Marine Decapod Crustacea of Southern Australia a Guide to Identification

Marine Decapod Crustacea of Southern Australia A Guide to Identification

With chapter on Stomatopoda by Shane Ahyong

Gary C. B. Poore This page intentionally left blank Marine Decapod Crustacea of Southern Australia A Guide to Identification © Museum Victoria 2004 All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO PUBLISHING for all permission requests.

National Library of Australia Cataloguing-in-Publication entry Poore, Gary C. B. Marine decapod Crustacea: a guide to identification. Includes index. ISBN 0 643 06906 2 (hardback). ISBN 0 643 09212 9 (netLibrary eBook). 1. Decapoda (Crustacea) – Australia, Southern – Identification. I. Title 595.380994

Available from CSIRO PUBLISHING 150 Oxford Street (PO Box 1139) Collingwood VIC 3066 Australia

Telephone: +61 3 9662 7666 Local call: 1300 788 000 (Australia only) Fax: +61 3 9662 7555 Email: [email protected] Web site: www.publish.csiro.au

Cover images, clockwise from top: Stenopus hispidus (photographer: Rudie H. Kuiter); Enoplometopus occidentalis (photographer: Rudie H. Kuiter); Munida gregaria (photographer: Rudie H. Kuiter); Paguristes frontalis (photographer: Rudie H. Kuiter); Bellidilia undecimspinosa (photographer: Michael Marmach); Actaea peronii (photographer: Karen Gowlett-Holmes); Rhynchocinetes australis (photographer: Michael Marmach); Galathea australiensis (photographer: Michael Marmach); Gnathophyllum taylori (photographer: Rudie H. Kuiter).

Set in Minion 9.5 pt Cover and text design by James Kelly Typeset by J & M Typesetting Printed in Australia by BPA Print Group TABLE OF CONTENTS

Acknowledgements...... ix

1. Introduction ...... 1 What is a crustacean?...... 1 Scope of this work ...... 2 Crustaceans of special interest ...... 5 History and resources ...... 5 Environment, ecology and biogeography ...... 7 Structure of this book, and some advice!...... 9

2. Systematics ...... 13 Classification of the Decapoda ...... 16 Key to suborders and infraorders of Decapoda and similar Crustacea...... 16

3. Dendrobranchiata – prawns and midwater shrimps ...... 23 Penaeoidea ...... 24 Aristaeidae...... 24 Benthesicymidae...... 27 Penaeidae...... 31 Box: Australian prawns ...... 40 Box: Southern Australian prawn fisheries ...... 41 Sicyoniidae ...... 41 Solenoceridae ...... 43 Sergestoidea ...... 45 Luciferidae...... 45 Sergestidae...... 46

4. Caridea – shrimps ...... 53 Box: Simple key to common shallow-water caridean families ...... 58 Pasiphaeoidea ...... 58 Pasiphaeidae ...... 58 Oplophoroidea ...... 63 Oplophoridae ...... 63 Atyoidea ...... 69 Atyidae ...... 69 Bresilioidea ...... 70 Bresiliidae ...... 70 Nematocarcinoidea ...... 73 Nematocarcinidae ...... 73 Rhynchocinetidae ...... 75 Stylodactyloidea ...... 78 Stylodactylidae ...... 78

v Table of contents

Campylonotoidea...... 80 Campylonotidae ...... 80 Palaemonoidea ...... 81 Gnathophyllidae ...... 81 Palaemonidae ...... 84 Alpheoidea ...... 98 Alpheidae ...... 98 Hippolytidae ...... 118 Ogyrididae ...... 126 Processoidea ...... 128 Processidae ...... 128 Pandaloidea ...... 129 Pandalidae ...... 129 Physetocaridoidea ...... 135 Physetocarididae ...... 135 Crangonoidea...... 136 Crangonidae ...... 136 Glyphocrangonidae ...... 141

5. Stenopodidea – coral shrimps and Venus shrimps ...... 145 Spongicolidae ...... 146 Stenopodidae ...... 148

6. Polychelida – deep-sea lobsters ...... 151 Polychelidae ...... 151

7. Astacidea – scampi and crayfish ...... 159 Enoplometopidae ...... 161 Nephropidae...... 162

8. Thalassinidea – ghost shrimps and sponge shrimps...... 169 Box: Common southern Australian ghost shrimps and sponge shrimps ...... 172 Axioidea ...... 173 Axiidae...... 173 Micheleidae ...... 176 Strahlaxiidae ...... 178 Callianassoidea...... 180 Callianassidae ...... 180 Ctenochelidae ...... 186 Laomediidae ...... 187 Upogebiidae ...... 189

9. Achelata – rock lobsters and bugs ...... 197 Palinuridae ...... 198 Box: Three common southern lobsters ...... 206 Scyllaridae ...... 206

10. Anomura – hermit crabs, porcelain crabs and squat lobsters ...... 215 Box: Quick key to hermits...... 217

vi Table of contents

Box: Six-legged crabs ...... 218 Galatheoidea ...... 219 Box: Miniature lobsters with long chelae...... 220 Chirostylidae ...... 220 Galatheidae ...... 228 Lomisidae ...... 240 Porcellanidae...... 242 Hippoidea ...... 247 Albuneidae ...... 247 Hippidae ...... 249 Paguroidea ...... 250 Diogenidae ...... 251 Lithodidae ...... 266 Paguridae ...... 270 Parapaguridae ...... 279 Pylochelidae ...... 286

11. Brachyura – crabs ...... 289 Box: Twenty-five common crabs ...... 296 Section Podotremata ...... 298 Subsection Dromiacea ...... 298 Dromiidae ...... 298 Dynomenidae ...... 308 Homolodromiidae ...... 310 Subsection Homoloidea...... 311 Homolidae...... 311 Latreilliidae ...... 314 Subsection Archaeobrachyura...... 316 Cyclodorippidae ...... 316 Cymonomidae ...... 318 Raninidae...... 320 Section Eubrachyura ...... 324 Subsection Heterotremata ...... 324 Dorippidae ...... 324 Calappidae...... 326 Leucosiidae ...... 331 Matutidae ...... 344 Majidae ...... 347 Box: Spider crabs and decorator crabs common in shallow-water ...... 349 Box: Spider crabs and decorator crabs found mainly at depths > 80 m or on seamounts ...... 349 Hymenosomatidae ...... 390 Parthenopidae ...... 398 Atelecyclidae ...... 400 Cancridae...... 401 Corystidae ...... 404 Geryonidae ...... 406 Portunidae...... 407 Goneplacidae...... 433

vii Table of contents

Hexapodidae ...... 439 Eriphiidae ...... 441 Pilumnidae ...... 448 Xanthidae ...... 460 Trapeziidae ...... 479 Domecidae...... 481 Cryptochiridae ...... 482 Subsection Thoracotremata ...... 484 Pinnotheridae ...... 484 Mictyridae ...... 487 Ocypodidae ...... 490 Palicidae...... 498 Grapsidae...... 500 Box: Grapsid crabs with potential for introduction to Australia ...... 512 Plagusiidae...... 512

12. Stomatopoda – mantis shrimps...... 517 Bathysquilloidea ...... 520 Bathysquillidae ...... 520 Erythrosquilloidea ...... 520 Erythrosquillidae ...... 520 Gonodactyloidea ...... 522 Gonodactylidae...... 523 Hemisquillidae ...... 525 Pseudosquillidae ...... 526 Lysiosquilloidea ...... 526 Tetrasquillidae...... 527 Lysiosquillidae ...... 528 Nannosquillidae ...... 530 Parasquilloidea ...... 535 Parasquillidae ...... 535 Squilloidea ...... 535 Squillidae...... 535

Glossary ...... 549

Credits...... 557

Taxonomic index ...... 561

Table 1. Species of decapod and stomatopod Crustacea introduced to Australia ...... 4

Table 2. A classification of living members of the subphylum Crustacea ...... 21

viii ACKNOWLEDGEMENTS

My interest in taxonomy in general and decapods in particular was encouraged in the early 1970s by Des Griffin, then at the Australian Museum. The collaborative work we began eventually lead me to work with the late Michèle de Saint Laurent in Paris, and to mix with numerous other decapod luminaries – Shane Ahyong, Keiji Baba, Fiona Bird, Chris Boyko, Sandy Bruce, Peter Castro, Alain Crosnier, Bill Dall, Peter Davie, Jacques Forest, Bella Galil, Danièle Guinot, Lipke Holthuis, Diana Jones, Brian Kensley, Rafael Lemaitre, Colin McLay, Enrique Macpherson, Peter Ng, Nguyen Ngoc-Ho, Bertrand Richer de Forges, Marcos Tavares and Christopher Tudge. All of these, either directly or indirectly, helped with this work. This project started in about 1988 with encouragement and help from my colleague and friend Robin Wilson. He made a significant contribution before he moved on to concentrate on polychaete systematics. In the final stages of pulling the manuscript together, Sherri Lehmann prepared illustrations, scanned them and made the plates, wrapped up text and wrote several sections from my basic notes. To both, my most sincere appreciation. Two scientific illustrators made hundreds of drawings from museum specimens. I am especially indebted to Kate Nolan (née Thompson) whose priceless drawings are the core of much of this contribution. Graham Milledge prepared many accurate drawings for early sections of the project and these are equally appreciated. Michael Marmach dedicated hours of his retirement to photographing crustaceans for this and other Crustacea Department endeavours. His patience is for ever appreciated and, I hope, rewarded. This work has only been made possible with access to the diverse and well catalogued collection at Museum Victoria. Collection staff, Robin Wilson, Elycia Wallis and Joanne Taylor, kept it in order. Tom Gunn volunteered many hours in the 1980s identifying unsorted specimens, teaching himself as he went from original literature. I thank all of these for their labours. Others provided hospitality and help in other institutions: Diana Jones at the Western Australian Museum, Penny Berents and Shane Ahyong at the Australian Museum, and Alain Crosnier and Michèle de Saint Laurent at the Museum national d’Histoire naturelle in Paris. Funding for the project, especially for the illustrations, came from Museum Victoria over several years. A financial contribution from the Australian Biological Resources Study, and the enthusiastic support of ABRS’s Pam Beesley, enabled the project to be completed almost on schedule. Most illustrations are original but I thank the publishers of several journals for allowing me to copy figures directly: Australian Museum, Museum and Art Gallery of the Northern Territory, Queensland Museum, CSIRO PUBLISHING, Muséum national d’Histoire naturelle, Paris, E.J. Brill, and The Crustacean Society. Several photographers allowed me to search their collections for suitable colour images and allowed me to use them freely. Special thanks to photographers, Bill Boyle, Neville Coleman, Karen Gowlett-Holmes, Rudie H. Kuiter, Mark Norman, Roger Springthorpe and Shane Ahyong, and to these individuals and the Australian Museum, CSIRO and Museum Victoria for copyright release. I appreciate the confidence of Nick Alexander, CSIRO PUBLISHING, in this project and his effort to get it published. More than anyone, I thank my wife Lynsey for supporting me in this and many other research projects. Only someone with her love of biology could tolerate my many hours absorbed in a world occupied only by lobsters, shrimps, isopods and their relatives.

ix This page intentionally left blank 1. INTRODUCTION

“For, whoever will follow its pages, crayfish in hand, and will try to verify for himself the state- ments which it contains, will find himself brought face to face with all the great zoological questions which excite so lively an interest at the present day …” (T. H. Huxley, 1880)

Charles Darwin’s friend and advocate, Thomas Huxley, wrote a book about the crayfish in 1880 that he hoped would stimulate an interest in the zoological questions that were important then. Many of ‘the great zoological questions’ to which he referred remain unanswered 124 years later. This guidebook is a list of the players acting out these questions. Crab, shrimp, prawn, lobster, crayfish, yabby and bug are just some of the names used in Australian English to describe members of the Crustacea. To these can be added more culinary terms from other languages – langoustine, scampi and udang. Most Australians would recognise a crustacean on the sea shore or on the dining table. This book sets out to document the diversity of some crustaceans found along the southern coast of Australia and to provide a means to tell one from another. Crustaceans are a group of related animals, meaning they probably had a common ancestor some time in the past. The features that they share are described in this introductory chapter. This volume deals with only some crustaceans, mainly those called ‘decapods’ and only those found in southern Australia. There are reasons for having such a narrow scope – the main one being that one volume can cope with no more. Much has been written about crustaceans before and in Chapter 1 this is reviewed to place present-day knowledge in context. In this introduction, some crustaceans of special interest for one reason or another are discussed. This introductory chapter explains how the book has been planned and what to expect from each section. Chapter 2 deals with the higher systematics of Crustacea and explains how the Decapoda and Stomatopoda, the only groups dealt with, are related to the many other crustaceans. General morphology is dealt with and specialist terms introduced. Having communicable names for species is essential and the principles that govern these are briefly explained. Chapters 3 to 12 cover crustacean groups in turn. Chapter 3 deals with prawns and related animals, Chapter 4 with most shrimps, Chapter 5 with coral shrimps, Chapter 6 with deep-sea lobsters, Chapter 7 with crayfish and scampi, Chapter 8 with sponge shrimps and ghost shrimps, Chapter 9 with rock lobsters and bugs, Chapter 10 with hermit crabs and squat lobsters, Chapter 11 with crabs, and Chapter 12 with mantis shrimps. The last was written by Shane Ahyong. There follows a glossary, because we have no choice but to learn a new language to identify these animals, with diagrams to explain the special terms applicable to different groups of decapods.

What is a crustacean? Insects, spiders, scorpions and millipedes are just some of the Phylum Arthropoda, that large group of animals distinguished from all others by their hard skin, or exoskeleton, and a segmented body. Crustaceans belong to this phylum too and may be called the insects of the sea. While insects are diverse and common on land, crustaceans are diverse and common in the sea. Unlike insects, which are more or less excluded from marine environments, crustaceans do get on to land and

1 Marine Decapod Crustacea of Southern Australia

into fresh water. Crustaceans, then, are arthropods that usually have many pairs of legs, a pair per body segment, and above all have two pairs of antennae (special sensory legs), one pair on each of the first two segments of the head. Insects, in contrast, have relatively few pairs of legs and only one pair of antennae. Chapter 2 goes into more detail about the classification of crustaceans. These details are of little importance when it comes to recognising a crustacean. We know from childhood what is a crab, shrimp or lobster and have never counted legs or antennae. In a catch of animals from a marine habitat it is simple to separate crustaceans from fish, snails, worms and seastars. Crustaceans are obvious to the marine biologist in the way that ants and flies are to land-lubbers for the simple reason that they are so abundant and so diverse. Densities may be in thousands of individuals per square metre of sea floor or cubic metre of ocean water and commonly many different kinds live side by side. The number of species in the world for which we have names is more than 50 000 but this is just a small fraction of the true number, which could be a million for all the world’s oceans. Or perhaps ten million. The reason for this underestimate is that only a small percentage has been described and are known by formal names. (Luckily for us this is not the case for the species in this book.) But for crustaceans as a whole, especially in Australia and especially for lesser known groups, this is true. Of all the c. 650 species of Crustacea in Port Phillip Bay, a well-studied bay near Melbourne, only about 13% are named. For the Order Isopoda, a group of small relatives of slaters, in deep marine seas off eastern Australia, only 10% of 359 species discovered have names (Poore et al., 1994). An area of half a metre-square of sand at the bottom of Port Phillip Bay may be home to as many as 70 species of crustaceans; most of these are undescribed. The significance of crustaceans in the world’s ecology can not be ignored. Crustaceans range in size from one-tenth of a millimetre maximum diameter (a tantulocarid parasite) to four metres across (a Japanese king crab). The mass of Antarctic krill in the oceans may be greater than that of any other single species. One species of oceanic copepod may be the most common animal on earth. And crustaceans have been found almost as far back in the fossil record as any group of complex animals. This contribution deals with but a few species from a small area.

Scope of this work This guide to species identification deals with just decapods and stomatopods. This is simply because these are the only major groups which can be reliably identified to species in southern Australia. While peracarid crustaceans such as amphipods (Lowry & Stoddart, 2003) and isopods (Poore, 2002) and copepods may be more common, very few are named. The Decapoda are just one Order of the Superorder Eucarida in the Subclass Eumalacostraca of the Class Malacostraca of the Subphylum Crustacea of the Phylum Arthropoda. The Stomatopoda are more primitive crustaceans in the generally-accepted ranking of the subphylum. They too comprise an Order but in another Subclass, Hoplocarida, of the Malacostraca. The morphological differences between stomatopods and decapods are profound. Both are included in this guidebook because individuals are much larger than typical examples of all other groups. Chapter 2 has more on crustacean taxonomy and the features that distinguish these taxa. This book confines itself to southern Australia for an equally practical reason. The marine fauna of southern Australia has a significant endemic component, meaning many of its species do not occur elsewhere. In contrast, the fauna of northern Australia is part of a much larger fauna extending from Japan through the western Pacific Ocean into the Indian Ocean, the so-called Indo-West Pacific fauna. The reasons for this dichotomy are explained in the Environment, Ecology and Biogeography section. Our task is made easier by restricting the coverage to a fauna that is smaller and taxonomically less confused. Even so, many Indo-West Pacific and northern Australian species reach as far south as ‘southern Australia’.Northern limits to this study are set at 32.5°S on the east coast and 31°S on the west coast to include the cities of Sydney and Perth (Fig. 1). All estuarine

2 Introduction

Gulf of Ashmore Reef Carpentaria

Great Barrier

North West Shelf

Reef

Shark Bay TASMAN SEA INDIAN OCEAN Elizabeth Reef 31.0oS

Great Australian Bight o 32.5 S Lord Howe Island ulf t Taupo Seamount

incen Spencer G Gulf St V Gascoyne Seamount Bass Strait SOUTHERN OCEAN

Tasmanian Seamounts Cascade Plateau

10˚S

Cape York

Darwin

Cairns Broome 20˚S Townsville NORTHERN TERRITORY Whitsunday Is North West Cape QUEENSLAND Rockhampton

Shark Bay Carnarvon WESTERN AUSTRALIA Brisbane Moreton Bay Geraldton SOUTH Southport Abrolhos I. AUSTRALIA Bryon Bay 30˚S Dongarra Lancelin I. Trial Bay Solitary Is Eucla NEW SOUTH WALES Coffs Harbour Perth Yamba Rottnest I. Fremantle Cockburn Sound Mandurah Esperance Adelaide Busselton Cape Naturaliste Port Jackson Cape Leeuwin Albany Sydney Botany Bay Kangaroo I. VICTORIA ACT Woolongong Robe Melbourne Eden Twofold Bay Gippsland Cape Howe Warrnambool Lakes Mallacoota Cape Otway ay Wilsons Promontory Flinders I. t Phillip B Por WesternTASMANIA Port 40˚S Hobart

Fig. 1. Top: geophysical and bathymetric map of Australia and surrounding continental shelf and seas. Major oceanographic features referred to in the text are indicated. The northern limits of the region covered, 32.5°S on the east coast and 31°S on the west coast, are marked. Bottom: political map of Australia with place names mentioned in text. States are abbreviated throughout as follows – Northern Territory (NT), Queensland (Qld), New South Wales (NSW), Victoria (Vic.), Tasmania (Tas.), South Australia (SA), Western Australia (WA).

3 Marine Decapod Crustacea of Southern Australia Other Australian Australian Other Not reported since reported Not Lakes localities Vic., Entrance Lakes Mallacoota; SA, Adelaide eastern coasts;eastern Vic., Vic., Flinders, Gippsland SA, central NSW, since reported Not Vic., Phillip Bay Port Tas., east and north- WA, Sound Cockburn Vic., Phillip Bay Port power stations power NSW, Sydney SA, Adelaide Tas., Hobart Eastern Tas. Eastern Eastern Tas. Eastern First recordFirst reported First 1970s 1990 1890s 1890s 1900 1990 1930s o Australia o Species Palaemon macrodactylus Palaemon novaezelandiae Charybdis japonica Halicarcinus Halicarcinus innominatus Petrolisthes elongatus Carcinus maenas Pyromaia tuberculata oratoriaOratosquilla Metacarcinus Metacarcinus Hymenosomatidae Caridea: Palaemonidae Shrimp Japanese Brachyura:Japanese Portunidae New Zealand Zealand New Brachyura: New Zealand Zealand New crab shore European Anomura: Brachyura: Portunidae Table 1.Table Species of t decapod introduced and stomatopod Crustacea nameCommon Taxonomy sea spider crabporcelain Porcellanidae swimming crab Spider crab shrimpMantis Brachyura: Majidae Stomatopoda: Squillidae Pie-crust crab Brachyura: Cancridae

4 Introduction

and marine environments are covered, from the intertidal down through the subtidal, across the continental shelf, down the continental slope to the deep sea. Most decapods live on the sea floor but also included are species that swim, especially mesopelagic oceanic species. Species known from seamounts off south-eastern Australia are included (Fig. 1, Pl. 1a). Within these geographic and environmental ranges some places, like shallow environments near population centres, are better explored than others and the chances of discovering new species are low. It would be no surprise, however, to find new records or even new species on the shelf of the Great Australian Bight, or on the continental slope or seamounts. Some species appear in the keys here on the basis of just one or two records. These may be genuinely rare but others are occasional vagrants from warmer waters. The fauna of Rottnest Island off Perth includes several species at the southern end of their range. There is evidence for warming of waters along the south-eastern coast during the 1990s and more subtropical species may turn up in future. This is principally a guidebook for identification but ecological observations are referred too in the few cases where they have been made. Davie’s (2002a, b) catalogues were published while this work was in progress and provided a valuable source of information. The few discrepancies between his lists and the coverage here are explained by newly identified material in museums not seen by him and not published. Very few new taxonomic decisions have been made in this work.

Crustaceans of special interest The crustaceans covered in this text are principally those met with on the shore or deeper marine environments of southern Australia. Of these, six crabs, one shrimp and one mantis shrimp are not native inhabitants and appear along with their 800 relatives in appropriate chapters (see Table 1). Decapods have been introduced to other parts of the world from their native areas by shipping and might be expected to eventually arrive, unwanted in Australia. Port Phillip Bay has a notorious reputation as a bay with a high percentage of exotic species (Hewitt et al., 1999, 2004) and species such as two shore crabs, Hemigrapsus pencillatus and H. sanguineus, and the Chinese mitten crab, Eriochier sinensis, now found in North America and Europe, might arrive unwanted. Just in case, these species are noted. If you find one, alert the local museum or fisheries authority! Edible decapods are another group of special interest, most not native to southern coasts. Keys are given to Australian species of prawns, lobsters and crabs found for sale in southern cities.

History and resources Like any group of animals, crustaceans have long attracted the attention of biologists and artists. Beautiful images of lobsters and crabs can be found in mosaics of ancient Roman cities, eighteenth century European still life painting and in the paintings of Australian Aborigines. The interest of these artists was surely because crustaceans are edible. Linnaeus included crustaceans in his Systema Naturae (1758), mostly in the genus Cancer. Many of the great European biological exploring expeditions of the late eighteenth and early nineteenth centuries collected marine and freshwater crustaceans along with other exotic wildlife. Australia attracted considerable attention from early European biological collectors, but plants and terrestrial mammals and birds fascinated James Cook, Joseph Banks, Matthew Flinders, Robert Brown and Ferdinand Bauer more than did marine crustaceans. French scientists first sailed and sampled in Australia in 1788, the year that Port Jackson was established as a convict settlement. But it was the expedition sent by Napoléon Bonaparte aboard the Le Géographe and Le Naturaliste, under the command of Nicholas Baudin, that showed the first real interest in southern Australian crustaceans. The voyage arrived in the region in 1801 with the naturalist François Peron. The collections that were taken back to the Jardin des Plantes, the natural history museum in Paris, provided a significant insight into the fauna. Many species of crabs and shrimps were described by Henri Milne Edwards (in 1837), and

5 Marine Decapod Crustacea of Southern Australia

their type specimens remain to this day in Paris. In another volume three years later, he published descriptions of many species of smaller crustaceans. Other foreign voyages followed including more by France. The United States Exploring Expedition visited Australia in 1839 with the notable geologist-zoologist James Dana on board. He published descriptions of several species but tragi- cally most of his material was lost in a shipwreck. The round-the-world voyage of H.M.S. Challenger set out to investigate the deep oceans and sampled offshore Australia in 1873. The Reports of the expedition are a constant source of reference for crustacean scientists even today but the contribution to the Australian fauna was small. Locally, Australian biologists in museums and universities in the major cities have always shown an interest in marine animals. Notable was William A. Haswell whose work in the 1880s in particular added numerous species to the fauna. He has been followed by numerous other carcinologists whose taxonomic contributions have accelerated knowledge of the fauna (Davie, 2002a). Casual shore collecting, dredging from steamships like the Thetis off New South Wales in the 1890s or from the Franklin a century later, scuba diving, and environmental surveys in modern times have all contributed museum material for study. The first Australian book on Australian Crustacea was Haswell’s Catalogue (1882) – a useful list but with little information on how to recognise species. It relied on reference to numerous esoteric articles in learned journals. The first book to treat the crustacean fauna of southern Australia in a way that enabled most species to be identified was Herbert M. Hale’s Crustaceans of South Australia, published in two parts (1927, 1929). The first part dealt with decapods, about 170 species, but does not cover many species of the Sydney or Perth regions. Hale’s book was reprinted unchanged in 1974 in response to demand but has not been revised since the first edition and is thus badly out of date. A more popular account was the slim volume by Anthony Healy and John Yaldwyn, Australian Crustaceans in Colour (1970). This appealing book relied heavily on colour photographs but it too is long out of print and covered few species. Its place has been taken by A Field Guide to Crustaceans of Australian Waters by Diana S. Jones and Gary J. Morgan (2002). This modern treatment has many photographs, each species expertly described. Crustaceans feature too in Graham Edgar’s Australian Marine Life – The Plants and Animals of Temperate Waters (1997, revised 2000). Books of colour photographs serve to introduce the amateur naturalist to the diversity of life, crustaceans in intertidal and shallow reef environments in these cases. While you might be lucky and be able to identify a species from a colour photo (many have distinctive colour patterns or are so different from everything else they can not be mistaken), you could be mislead. Several groups of species are subtly different from each other and nothing but careful examination of characters hidden in photos is the only solution. The two recent volumes of the Zoological Catalogue of Australia: Eucarida by Peter Davie (2002a, b) listed all the species of Australian decapod and stomatopod crustaceans known. They provided valuable taxonomic detail but are not an identification guide. There are no keys and no diagnoses beyond families and subfamilies. The volumes provide a valuable check against which this identification guide has been written and already new southern records can be noted. General books on Crustacea are found only in speciality libraries. Old books have a fascina- tion of their own because they established the classifications and terminology still used today. The Reverend T.R.R. Stebbing’s A History of Crustacea. Recent Malacostraca (1893) and William T. Calman’s The Life of Crustacea (1911) are classics. Patsy A. McLaughlin’s Comparative Morphology of Recent Crustacea (1980) is a useful practicable guide to the definition of living taxa but is not a taxonomic guide and has no ecological information. Alfred Kaestner’s (1980) Crustacea is a useful translation of a systematic German text but is an academic treatment, now out of print. Frederick R. Schram’s more detailed volume, also entitled Crustacea (1986), is packed with information and includes all the fossil taxa but has a controversial and idiosyncratic classification. It addresses only the most specialist audience, other crustacean biologists.

6 Introduction

Those determined to have every general text of crustaceans might find these in antiquarians. A more general text is the lighter Crustaceans by Waldo L. Schmitt (1973). It is anecdotal and, even if obtainable, does not deal in detail with identification. Similar, is Shrimps, Lobsters and Crabs: Their Fascinating Life Story by Dorothy E. Bliss (1982) with an American bias. J. Green’s A Biology of Crustacea (1961) covers identification scarcely at all and is now very old. And of course many general invertebrate zoology textbooks have a chapter on crustaceans. Now, in the early twenty-first century the worldwide web is a huge bank of information on crustaceans, as on anything else. Martin & Davis (2001) appended to their catalogue of the families of crustaceans a commentary on classification and a long list of web-based resources. As valuable as this was at the time, and may still be, no such printed list can ever be up to the minute. Museum Victoria’s website, Crustaceans of southern Australia (http://www.museum.vic.gov.au/crust) contains colour photographs of more than 100 marine crustacean species from this region.

Environment, ecology and biogeography Southern Australia lies in the temperate zone, in contact with the tropics in the north and facing the Southern Ocean. Sea-surface temperatures ranging from 15.5 to 25°C in summer and 11.5 to 20°C in winter. The intertidal and immediate subtidal coast is rocky for the most part but interspersed are lengths of long sandy beaches and bays. Notable among the bays and inlets are Port Phillip Bay and Western Port in Victoria, and Gulf St Vincent and Spencer Gulf in South Australia. The western and southern coasts lack major rivers and consequently estuaries are few and small. The Gippsland Lakes in Victoria are exceptions. Even the largest river in Australia, the Murray which reaches the coast in South Australia, struggles to maintain an opening to the sea and, even when it does, flow is quite salty. The lack of terrigenous input means that carbonate sediments cover most of the continental shelves. In contrast, siliceous sediments dominate the eastern coast of Australia. Tides are generally small, between 1.0 and 2.5 m, and much of the south and west coast is fully exposed to storm waves from the Southern Ocean (Bunt, 1987). Nevertheless, southern Australia offers a wide range of habitats to the one-third of the species of animals in marine environments that are Crustacea. These inhabit beaches and rocky shores, under stones, in the sediment and nestled in amongst algae and many other microhabitats. They also inhabit the open ocean and the deep sea. Most species live in a limited range of habitats and the ecological notes that accompany each species are a guide that might help in identification. It is the nature of species’ abundances that many are rare – one should be cautious identifying a new individual as a rare species, especially out of its range. Shrimps whose ecology is said to be ‘mesopelagic’ are never found intertidally and rarely even in midwater samples above the continental shelf. Most species found intertidally live also in the shallow subtidal (down to perhaps 10 m depth) but the reverse is not true. Many species of shallow reefs or of continental shelf sediments are never seen intertidally, unless washed up dead after a storm. The shallow-water marine biota of southern Australia has generally been described as highly endemic, with estimates for various taxonomic groups ranging from 85 to 95% (Womersley, 1981; Wilson & Allen, 1987). The history of the fauna is complex and these estimates may not be generally applicable. A brief review of the region’s oceanographic setting and geological history is illuminating. Southward-flowing currents exist on both the eastern and western coasts of Australia (Wilson & Allen, 1987). The warm Leeuwin Current arises in Indonesia as a western extension of the Equatorial Current and in winter it flows south along the coast of Western Australia and then west into the Great Australian Bight. From here it is replaced by the warm saline Great Australian Bight Current (Rochford, 1986) and then by the weaker Zeehan Current that runs as far east as the western coast of Tasmania (Baines et al., 1983). The East Australian Current also originates in the tropics and flows from the northern Tasman Sea south along the coasts of Queensland and New South Wales before veering off near 33°S back into the Tasman Sea. A series of warm-water eddies

7 Marine Decapod Crustacea of Southern Australia

can be pushed further south into eastern Bass Strait and sometimes as far as the eastern coast of Tasmania (Nilsson & Cresswell, 1981). Both of these currents help to bring tropical crustaceans into southern Australia. The southern Australian coastline began forming in the late Cretaceous (96 million years ago) as western Australia started rifting away from Antarctica. The resulting marine intrusion was colonised by warm-water species from the west. The eastern coast of Australia supported a cool- water fauna that extended around Antarctica to what is now South America. Australia and Antarctica finally split around 35.5 million years ago south of Tasmania, allowing the western and eastern faunas to mix (McLoughlin, 2001). Since that time, any endemic species that might have existed in southern Australia were supplemented by further invasions of tropical species as Australia drifted northwards and collided with South-East Asia as early as 20 million years ago, and by a smaller number of Southern Ocean species arriving by the circumpolar subantarctic currents (Darragh, 1985; Edgar, 1986; Poore, 1994). The Pleistocene period has been marked by over 60 glacial-interglacial cycles. Glacial times were characterised by a sea-level 110 m or more below current sea-level and cooler temperatures. Bass and Torres straits were generally dry, the Subtropical Convergence moved north towards the southern Australian coastline, the Leeuwin Current ceased to flow around south-western Australia and the East Australian Current may have flowed further south. Several interglacials were warmer than at present and during the last interglacial (125 000 years ago) corals were found as far south as Newcastle in New South Wales and Albany in south-western Western Australia indicating a water temperature 1–2°C warmer than present (see references in O’Hara & Poore, 2000). The closure of Bass Strait during glacial periods has been thought to drive speciation in south-eastern Australia and some examples of sister taxa do support this (Dartnall, 1974). Early research into the distributions of Australian marine fauna and flora focused on delin- eating distinct regions or biogeographical provinces (e.g. Knox, 1963; Womersley, 1981). Typically such schemes included a Flindersian Province that extended from Bass Strait to Geraldton, WA, a Peronian Province that extended from southern Queensland to Bass Strait, and sometimes a Maugean Province that included Bass Strait and Tasmania. Wilson & Gillett (1971) proposed a simplified scheme, recognising a broad temperate zone in southern Australia and a tropical zone in the north, overlapping on the east and west coasts. O’Hara & Poore (2000) found that, for 738 species of echinoderms and decapods at least, the southern zone was not as uniform as might be expected for a coast almost at a single latitude. Species richness is relatively constant from east to west but there are more species (in a given area) in the warm-temperate seas around Perth and Sydney than in cold-temperate southern Tasmania. Further, species composition turned over along the coast, especially at significant geographic features like bays and promontories. Only 20% of the species found at one end of the coast were present at the other. Species richness is not related to the area of continental shelf or average species range-size. There is some evidence that the southern distribution limit of some species is related to minimum sea-surface temperature. Historical and ecological factors contribute to explanations of the gradient of marine species richness in southern Australia. Tropical species have continued to invade as Australia has split from Gondwana and drifted northward. Some Gondwanan cool-temperate species at the limits of their range have gone extinct, as evidenced in the fossil record. Because of isolation from other cool- temperate continents, immigration has been slow. The high percentages of endemic species of southern molluscs (over 95%) and fish (85%) reported (Wilson & Allen, 1987) do not apply for decapods and stomatopods. Of the 800 species mentioned in this volume, 25 per cent (204) are endemic. Endemism is defined for this purpose as occurring somewhere on the southern coast and having a range not extending beyond southern Queensland or mid-Western Australia, i.e., as far as Wilson & Allen’s (1987) overlap zones. The total fauna on which this calculation is based includes species with different ecologies and

8 Introduction

biogeographies. Mesopelagic and bathypelagic shrimps and prawns tend to be widespread in the Indo-West Pacific or wider in the world’s oceans, at least on the basis of current taxonomy. Species of mesopelagic and bathypelagic Dendrobranchiata (especially Benthesicymidae and Sergestidae) and Caridea (Oplophoridae, Pandalidae, Nematocarcinidae and Pasiphaeidae) have been reported. Similarly, deep-water benthic crabs, lobsters and shrimps, restricted to 200 m depth or more (continental slope) are widespread (or known from few or unique specimens). These comprise Caridea (especially Crangonidae and Hippolytidae), lobsters (Nephropidae, Polychelidae), Anomura (Chirostylidae, Galatheidae, Parapaguridae) and Brachyura (Majidae and several primitive families). Deep-water species contribute 230 species. Eight species are introduced to southern Australia from other places (Table 1). Thirty-one species are essentially tropical species recorded rarely and only as a southern species at Rottnest Island or the nearby mainland WA. Most of these are xanthoid, majid or leucosiid crabs. Twenty-four species representing several diverse families are confined to the east coast of Australia, between Qld and southern NSW. If all these groups of species are excluded, the fraction of endemic species (those recorded from intertidal and shelf depths on the southern coast – not only on the southern parts of the eastern and western coasts) is 40 per cent. These ‘truly endemic’ species are represented across the taxa by many families having one or two species. In only a few families are much more than half the species endemic. All shallow-water species of two caridean shrimp families, Hippolytidae (six species) and Rhynchocinetidae (five species), are endemic while only 20 per cent of Alpheidae, the most species-rich family, are endemic. Most thalassinideans are shallow-water and endemic, the only exceptions being two species of Axiidae and four species of Upogebiidae. Deep-water chirostylid and galatheid anomurans are a mixture of endemic and widespread species but limited sampling could give misleading data. Anomuran hermit crabs are mostly endemic: Diogenidae (15 of 31 species) and Paguridae (most shallow-water species). More than half of the species of few shallow-water crab families are endemic: Dromiidae (12 of 15), Grapsidae (7 of 15) and Hymenosomatidae (8 of 14). The largest crab families are dominated by widespread species (Majidae, Portunidae, Xanthidae, Leucosiidae and Pilumnidae) as is the case for Stomatopoda. Genera that can be said to be rich in endemic species on the southern coast are few and are represented by few species, four to nine at most: Philocheras, Leontocaris, Tozeuma, Rhynchocinetes (Caridea), Austrodromidia, Ebalia, Leptomithrax, Micippa, Naxia, Halicarcinus, Pilumnus, (Brachyura), Agononida, Uroptychus, Galathea, Munida, Munidopsis, Paguristes (Anomura). This evidence suggests that evolution of local species has been widespread throughout the decapod and stomatopod taxa and that few genera have radiated more in southern Australia than elsewhere.

Structure of this book, and some advice! This volume follows what is becoming accepted as a new taxonomy of the decapods and stomatopods, each chapter devoted to a major group (Schram, 2001; Dixon et al., 2003). This classification differs in some details of the higher groupings from that used by Martin & Davis (2001). They provided a complete listing of families and discussion of taxonomic controversies summarised and updated for Decapoda in the next chapter. Sections within each chapter are hierarchical, species within genera, within families (with often subfamilies as well). For each taxon, information about diversity in Australia and elsewhere is summarised along with helpful characters for identification. Useful literature and key ecological or historical works are cited but the worldwide web now enables more up-to-date searches. Diagnoses of families are either original or simplified from more detailed diagnoses in Davie (2002a, b) or other sources. The families are arranged in alphabetical order within superfamilies or sections. This may aid identification if you have to resort to flicking through pictures to find something similar to the animal in your hand. Even the best taxonomists do this!

9 Marine Decapod Crustacea of Southern Australia

Literature cited within each family section is listed at the end of that section. It is an important source for further detail but does not necessarily include papers in which species and other taxa were originally described. See Davie (2002a, b) for that. Generic diagnoses are adapted from the literature but some, particularly well established and familiar ones, have never been diagnosed recently. This gap has been filled as well as possible and in general the diagnoses will serve to differentiate genera within this region if not across the globe. The information given for each species includes: genus and species names, author and year of description, common name if known, reference to a figure, diagnosis, size, geographical distribution, and ecological and depth distribution. Remarks may follow. If the author and year are in brackets, this is meaningful – the species has been moved from its original genus to another. The size given is usually the greatest dimension – sometimes carapace length (cl.), carapace width (cw.), or total length (tl.) are specified. Not all species have been recently diagnosed. Here, diagnoses are as short as possible, at least sufficient to tell one southern Australian species from another, but are often applicable on a wider scale. If in doubt, reference will have to be made to the cited literature and more elaborate descriptions and illustrations. Virtually all species are illustrated. One or more species of each family are figured in toto, a habitus drawing, and others by just key characters. Many are original drawings from museum specimens and others are copied from published illustrations. The sources of these figures are given in the credits. Some illustrations are electronic scans of published figures, acknowledged in captions. Identification is achieved through the use of dichotomous keys adapted from many originally published in the primary literature, or developed from scratch. Applicability of the keys varies but is explained in each heading. Some keys are to all taxa but most are to southern Australian taxa only. The temptation when using keys is to assume when you get an answer that your species is identified. For common species this will be so. But be warned! Cryptic species do exist and new discoveries are likely – species that are similar to those named but are subtly different. It is also highly probable on the northern margins of the region that species known further north will appear from time to time. This is true, for example, of midwater prawns and shrimps being carried southwards past Sydney in the summer by the Eastern Australian Current, or for species associated with coral on Rottnest Island. Reference to the relevant literature cited is the only solution when in doubt. Having said this, except on seamounts and in the deep sea, few new species have been reported over the last two decades. Even the most recently described species, common shore shrimps (Ahyong, 2003; Walker & Poore, 2003), were known as undescribed species to taxonomists in the 1970s. Nevertheless, as-yet-unrecognised cryptic species are probable even in the most obvious taxa. Parallels with the recent separation of the common intertidal biscuit star, Patiriella gunnii, into four species (O’Loughlin et al., 2003) could be predicted in the Decapoda. No new species are described in this work but some as-yet-unnamed are mentioned and figured. Some species are placed in different genera or higher taxa from those that other taxonomists have chosen. These are discussed on a case-by-case basis. Complete synonymies are inappro- priate but important revisions and alternative old names are given. The information in this volume derives from over 200 years of collecting by expert taxonomists in most accessible environments and published in thousands of papers in numerous journals in several languages. More than 800 of these papers and books are cited here. The same information was expertly summarised by Davie (2002a, b) for his catalogue of the entire Australian decapod and stomatopod fauna. The few differences from Davie’s lists result from my appraisal of collections of specimens now held in Museum Victoria, the Australian Museum, Western Australian Museum and South Australian Museum. Catalogues of these collections

10 Introduction

were reviewed, old names reviewed and enigmatic specimens examined. Some taxa not in the catalogues were revealed and extensions to published distributions revealed. Keys written by expert taxonomists for other taxonomists use arcane language. Terminology differs from one taxon to another. Every effort has been made here to simplify and explain terminology – but often one word of jargon conveys a complex idea and can not be avoided. The glossary should be helpful and is illustrated with special terms applicable to only some groups.

References Ahyong, S.T. 2003. Gnathophyllum taylori, a new species of caridean shrimp from southeastern Australia (Crustacea: Decapoda: Gnathophyllidae). Memoirs of Museum Victoria 60: 237–242. Baines, P.G., Edwards, R.J., & Fandry, C.B. 1983. Observations of a new baroclinic current along the western continental slope of Bass Strait. Australian Journal of Marine and Freshwater Research 34: 155–157. Bliss, D.E. 1982. Shrimps, Lobsters and Crabs: Their Fascinating Life Story. New Century Publishers: Piscatawey. 242 pp. Bunt, J.S. 1987. The Australian marine environment. Pp. 17–42 in: Dyne, G.W. (ed.) Fauna of Australia. Australian Government Publishing Service: Canberra. Calman, W.T. 1911. The Life of Crustacea. Methuen & Co.: London. 289 pp. Darragh, T.A. 1985. Molluscan biogeography and biostratigraphy of the Tertiary of southeastern Australia. Alcheringa 9: 83–116. Dartnall, A.J. 1974. Littoral biogeography. Pp. 171–194 in: Williams, W.D. (ed.) Biogeography and Ecology in Tasmania. Junk: The Hague. Davie, P.J.F. 2002a. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Davie, P.J.F. 2002b. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K.(eds), Zoological Catalogue of Australia. Vol.19.3B CSIRO Publishing: Melbourne. xiv, 641 pp. Dixon, C.J., Ahyong, S., & Schram, F.R. 2003. A new hypothesis of decapod phylogeny. Crustaceana 76: 935–975. Edgar, G.J. 1986. Biogeographic processes in the Southern Hemisphere marine environment. Medio Ambiente 1: 11–111. Edgar, G.J. 1997. Australian Marine Life – The Plants and Animals of Temperate Waters. Reed Books: Kew Victoria. 544 pp. Green, J. 1961. A Biology of Crustacea. Witherby: London. 180 pp. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Hale, H.M. 1929. The Crustaceans of South Australia. Part 2. South Australian Government Printer: Adelaide. 202–380 pp. Haswell, W.A. 1882. Catalogue of the Australian Stalk-and Sessile-eyed Crustacea. Australian Museum: Sydney. xxiv, 324 pp. Healy, A., & Yaldwyn, J.C. 1970. Australian Crustaceans in Colour. Reed: Sydney. 112 pp. Hewitt, C.L., Campbell, M.L., Thresher, R.E., & Martin, R.B. 1999. Marine biological invasions of Port Phillip Bay, Victoria. Centre for Research on Introduced Marine Pests, CSIRO Marine Research, Technical Report 20: 1–344. Hewitt, C.L., Campbell, M.L., Thresher, R.E., Martin, R.B., Boyd, S., Cohen, B.F., Currie, D.R., Gomon, M.F., Keough, M.J., Lewis, J.A., Lockett, M.M., Mays, N., McArthur, M.A., O’Hara, T.D., Poore, G.C.B., Ross, D.J., Storey, M.J., Watson, J.E., & Wilson, R.S. 2004. Introduced and cryptogenic species in Port Phillip Bay, Victoria, Australia. Marine Biology 144: 183–202.

11 Marine Decapod Crustacea of Southern Australia

Huxley, T.H. 1880. The Crayfish. An Introduction to the Study of Zoology (International Scientific Series Vol. 28). Paul & Co: London. 371 pp. Jones, D.S., & Morgan, G.J. 2002. A Field Guide to Crustaceans of Australian Waters. Reed New Holland: Sydney. 224 pp. 2nd edn. Kaestner, A. 1980. Invertebrate Zoology. Vol. 3 Crustacea Kreiger Publishing Co.: New York. 523 pp. Reprint edn. Knox, G.A. 1963. Littoral biogeography and ecology of the Australasian coasts. Oceanography and Marine Biology, Annual Review 1: 341–404. Lowry, J.K., & Stoddart, H.E. (eds) 2003. Crustacea: Malacostraca: Peracardia: Amphipoda, Cumacea, Mysidacea. In: Beesley, P.L.,& Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.2B. CSIRO Publishing: Melbourne. xii, 531 pp. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. McLaughlin, P.A. 1980. Comparative Morphology of Recent Crustacea. Freeman and Co.: San Francisco. 177 pp. McLoughlin, S. 2001. The breakup history of Gondwana and its impact on pre-Cenozoic floristic provincialism. Australian Journal of Botany 49: 271–300. Milne Edwards, H. 1837. Histoire Naturelle des Crustacés, comprenant l’Anatomie, la Physiologie et la Classification es ces Animaux. Vol. 2 Librairie Encyclopédique de Roret: Paris. 531 pp. Nilsson, C.S., & Cresswell, G.R. 1981. The formation and evolution of East Australian Current eddies. Progress in Oceanography 9: 133–183. O’Hara, T.D., & Poore, G.C.B. 2000. Patterns of distribution for southern Australian marine echinoderms and decapods. Journal of Biogeography 27: 1321–1335. O’Loughlin, M., Waters, J.M., & Roy, M.S. 2003. A molecular and morphological review of the asterinid, Patiriella gunnii (Gray) (Echinodermata: Asteroidea). Memoirs of Museum Victoria 60: 181–196. Poore, G.C.B. 1994. Marine biogeography of Australia. Pp. 189–213 in: Hammond, L.S., & Synnot, R. (eds), Marine Biology. Longman Cheshire: Melbourne. Poore, G.C.B. 2002. Crustacea: Malacostraca: Syncarida, Peracarida: Isopoda, Tanaidacea, Mictacea, Thermosbaenacea, Spelaeogriphacea. In: Houston, W.W.K. & Beesley, P.L. (eds) Zoological Catalogue of Australia. Vol. 19.2A. CSIRO Publishing: Melbourne. xii, 434 pp. Poore, G.C.B., Just, J., & Cohen, B.F.1994. Composition and diversity of Crustacea Isopoda of the south- eastern Australian continental slope. Deep-Sea Research 41: 677–693. Rochford, D.J. 1986. Seasonal changes in the distribution of Leeuwin Current Waters off Southern Australia. Australian Journal of Marine and Freshwater Research 37: 1–10. Schmitt, W.L. 1973. Crustaceans. David & Charles: Newton Abbot. 204 pp. Schram, F.R. 1986. Crustacea. Oxford University Press: New York. xii, 606 pp. Schram, F.R. 2001. Phylogeny of decapods: moving towards a consensus. Hydrobiologia 449: 1–20. Stebbing, T.R.R. 1893. A History of Crustacea. Recent Malacostraca. Kegan Paul, Trench, Trübner and C.: London. xvii, 466 pp. Walker, T.M., & Poore, G.C.B. 2003. Rediagnosis of Palaemon and differentiation of south-eastern Australian species (Crustacea: Decapoda: Palaemonidae). Memoirs of Museum Victoria 60: 243–256. Wilson, B.R., & Allen, G.R. 1987. Major components and distribution of marine fauna. Pp. 43–68 in: Dyne, G.W. (ed.) Fauna of Australia. Vol. 1A. General Articles. Australian Government Publishing Service: Canberra. Wilson, B.R., & Gillett, K. 1971. Australian Shells. Reed: Sydney. 168 pp. Womersley, H.B.S. 1981. Aspects of the distribution and biology of Australian marine macro-algae. Pp. 294–306 in: Pate and A.J. McComb, J.S. (ed.) The Biology of Australian Plants. University of Western Australia Press: Nedlands.

12 2. SYSTEMATICS

Besides being extraordinarily abundant in the sea, crustaceans are amongst the most morphologically diverse of any taxon. They are far more varied than insects or spiders. Most are not crabs, shrimps or lobsters but are small, unfamiliar and rarely have common names. Biologists know them as ostracods, copepods, amphipods, cumaceans, isopods, tanaids and a host of other names. Some are illustrated in Fig. 2 but unfortunately most will not be dealt with here. Some would argue that the Crustacea are not a monophyletic group but are a mixture of groups with independent ancestry that look similar. Schram & Koenemann (2004) observed that Cambrian fossil crustaceans are of two types and both may have given rise to Crustacea as we know them today. The Orsten fossils are minute short-bodied animals with few body segments while the Burgess Shale and similar deposits in China contain assemblages of diverse large multi- segmented crustacean-like arthropods. Others have used morphological and molecular evidence to support a polyphyletic ancestry (Spears & Abele, 1998) or monophyly (Edgecombe et al., 2000; Giribet et al., 2001). Others believe that other arthropod taxa have evolved from within the Crustacea – the view that insects are just flying crustaceans! For the practical purposes of identification to species, this debate does not matter. Ancestry does matter, however, at lower taxonomic levels. In theory at least, a species’ genus and family placement should tell you something about its ecology or evolutionary history. It is the attempts of taxonomists to reflect this history that induces them to move species from one genus or family to another. Table 2 is one hierarchy of all the major groups of Crustacea where it can be seen that the Decapoda and Stomatopoda are just small parts. This hierarchy of groups within groups is just one view of what a classification should be. The higher classification of Crustacea engrosses many taxonomists and there is not general agreement yet. This was discussed at length by Jody Martin and George Davis who sought the views of more than 90 specialists in their attempt to list and organise the 849 families of crustaceans (Martin & Davis, 2001). If one takes the view that a classification should reflect evolutionary history, the hierarchy becomes very fluid as new analyses of data from morphology and DNA intrude. The Subphylum Crustacea is divided into six classes: Branchiopoda – many branchiopods live in fresh water and only the Cladocera, popularly known as water fleas, have marine representatives. A useful guide to the identification of species in Australia is Williams (1980), a book sadly out of print. Hawking provided a useful lead into the more recent literature dealing with these and other taxa in Australia (Hawking, 2000). Remipedia – an unusual group of many-legged shrimps confined to fresh water in caves and aquifers (Schram et al., 1986; Koenemann et al., 2003). One species has been found in Western Australia (Yager & Humphreys, 1996). Cephalocarida – microscopic thin animals living between the sand grains on beaches and marine sediments (Sanders, 1963; Hessler, 1992; Hessler & Elofsson, 1992). Although species have been reported from New Zealand (Knox & Fenwick, 1977) and New Caledonia (Cals & Delamare- Deboutteville, 1970), none has yet been discovered in Australia. Maxillipoda – a diverse group of crustaceans, rich in small species from all environments. Copepods are a dominant example in the sea, a highly variable group of pelagic and benthic forms

13 Marine Decapod Crustacea of Southern Australia

(Huys & Boxshall, 1991; Boxshall & Halsey, 2004). Many copepods are parasitic or symbiotic (Kabata, 1970 and earlier publications). Some planktonic forms in Australia have been documented by various authors (Dakin & Colefax, 1940; McKinnon et al., 1992). Most benthic copepods belong to the Order Harpacticoida but few have been described from Australia (Hamond, 1973; Walker-Smith, 2001; Harris, 2002). Notable too in marine environments are barnacles. Parasitic barnacles are well studied generally but poorly known in Australia (Høeg, 1995; Høeg & Lützen, 1995). Shallow-water free-living barnacles in Australia are better understood (Jones, 1990, 1991, 1992). Ostracoda – another group rich in species in marine, freshwater and terrestrial environments, last reviewed for Australia many years ago (De Deckker & Jones, 1978). One major group comprising mostly small individuals enclosed in a calcareous carapace, Podocopida, includes a rich fossil fauna important in stratigraphy. The other group, Myodocopida, are larger and chitinous (Kornicker, 1994, 1995, 1996; Kornicker & Poore, 1996; Parker, 1998). Some are important benthic scavengers. Malacostraca – includes the most familiar crustaceans arranged in three subclasses, five superorders and 16 orders. A valuable handbook for the identification of freshwater malacostracan families was published by the Cooperatative Research Centre for Freshwater Ecology (Horwitz et al., 1995). One subclass includes only the Leptostraca. In Australia the group is rich in species but few are described (Walker-Smith, 1998, 2000; Walker-Smith & Poore, 2001). The second subclass, Hoplocarida, includes Stomatopoda, mantis shrimps, treated in Chapter 12. The third subclass is the Eumalacostraca. All eumalacostracans (and Hoplocarida) have 20 body segments (all except the first with paired limbs) plus a terminal telson. The body segments, or somites, are arranged into three sections, or tagmata, head or cephalon, thorax and abdomen. The head has eight pairs of limbs involved in sensation and feeding, the thorax eight pairs usually locomotory and variously involved with feeding, and the abdomen has six pairs of limbs involved with respiration, reproduction and swimming. Homologies between the segments in all Crustacea and between Crustacea and other arthropods has long fascinated zoologists; developmental genetics and the identification of genes responsible is illuminating the subject (Abzhanov & Kaufman, 2003; Schram & Koenemann, 2003). Figure 179 in the Glossary is a diagram explaining how the segmentation of a malacostracan crustacean appears in a decapod and stomatopod crustacean. There are three superorders within Malacostraca. The superorder Syncarida is important in Australia because of the diversity of Anaspidacea (Jarman & Elliott, 2000; Lake et al., 2002). Bathynellacea are cryptic cave and interstitial forms (Poore & Lew Ton, 2002). The superorder Peracarida includes nine orders of mostly abundant small benthic or epibenthic crustaceans. The numbers of species described from Australia are now thousands. The most abundant and diverse are amphipods (Gates et al., 2003; Lowry, 2003; Stoddart et al., 2003; Stoddart & Lowry, 2003b, c) and isopods (Poore, 2002). Less diverse but also less well studied are Cumacea (Stoddart & Lowry, 2003a) and Tanaidacea (Poore et al., 2002), and Mysidacea and Lophogastrida (Keable et al., 2003). Spelaeogriphacea (Poore & Humphreys, 2003) and Thermosbaenacea (Poore & Humphreys, 1992) are represented by only one or two species each. In aquifers in the Pilbara, WA. Mictacea is a rare deep-sea and anchialine order with one Australian species (Just & Poore, 1988). The superorder Eucarida is characterised by a carapace – the shield or shell encasing the segments of the head and thorax in such a way that the segmentation is concealed. In most other crustaceans, slaters for example, many of the segments are quite visible. All eucarids have eyes on moveable stalks. Eucarids comprise three orders, Euphausiacea, or krill (Mauchline, 1980; Baker et al., 1990), pelagic crustaceans famous for their role in the diet of whales, a single species of Amphionidacea (Davie, 2002), and Decapoda. It is only this order that features in this guide.

14 Systematics

e b d

a f

g h i

m j

k l n

q o p

t u

r s

Fig. 2. Representatives of some basic crustacean types. a, Copepoda. b, Ostracoda. c, Anostraca. d, Notostraca. e, Cladocera. f, Leptostraca. g, Anaspidacea. h, Cirripedia. i, Stomatopoda. j, Mysida. k, Tanaidacea. l, Isopoda. m, Amphipoda. n, Euphausiacea. o, Dendrobranchiata. p, Caridea. q, Astacidea. r, Anomura. s, Achelata. t, Thalassinidea. u, Brachyura. v, Polychelida.

15 Marine Decapod Crustacea of Southern Australia

Classification of the Decapoda Decapods differ from other eucarids in having the first three pairs of limbs attached to the segments of the thorax modified. The modified limbs are called maxillipeds and are feeding appendages. The consequence is that only the last five pairs of thoracic limbs are involved in loco- motion. These five pairs give the group its name, Decapoda, meaning ten-footed. The ten legs refer only to the claws and walking legs. The monophyly of the Decapoda is without doubt but the arrangement of taxonomic groups within the order has been debated over more than a century (Schram, 2001; Dixon et al., 2003). This book adopts the now widely held classification into two suborders: Dendrobranchiata and Pleocyemata. The latter has been traditionally divided into several infraorders corresponding to recognised body forms but this arrangement is already collapsing as a result of recent phylogenetic analyses. Relationships between Anomura and Brachyura are unclear (e.g. Spears et al., 1992; Tavares, 2003), between Anomura and Thalassinidea uncertain in some views (de Saint Laurent, 1979; Tudge, 1997), and between Astacidea and Palinura unresolved (Forest & de Saint Laurent, 1989). Doubts about the monophyly of the Palinura in particular were expressed by Scholtz & Richter (1995) and Schram & Ahyong (2002). Scholtz & Richter’s (1995) analysis of reptant decapod relationships was not based on unbiased character selection or dependent on parsimonious criteria but fuelled debate on the traditional classification of infraorders. Further rearrangement of relationships was suggested by cladistic analysis of morphological characters (Dixon et al., 2003). In their phylogeny, Palinura in particular are no longer recognised as such but comprise two clades not closely related: the Polychelida (family Polychelidae) and Achelata (Scyllaridae and Palinuridae). This guidebook adopts their recommendations and discusses their reasoning under relevant chapter headings. A key to differentiate the traditional suborders and infraorders of Decapoda may seem unnec- essary but is included as a quick critical guide. Several shrimp-like crustaceans look superficially alike and placing a shrimp in Thalassinidea, Caridea or Dendrobranchiata is not always obvious. Some non-decapod shrimp-like taxa are included here. Pereopods 1–5, the term used for legs in decapods, is equivalent to thoracic appendages 4–8 in other malacostracans. Exceptions could be found for most couplets, for example, secondarily chelate or simple limbs appear in most orders.

Key to suborders and infraorders of Decapoda and similar Crustacea 1. Carapace cylindrical, deep, enclosing thoracic appendages; 7 pairs of thoracic appendages in male, last absent in female; thoracic exopods rudimentary; female pleopod 1 extending forward as large brood-plate; rare pelagic shrimp ...... Amphionidacea … Amphionides reynaudii — These characters not combined ...... 2 2. Carapace short, exposing feathery thoracic gills; 8 thoracic appendages all similar, or last 1 or 2 lost, or first few modified; pelagic, shrimp-like ...... Euphausiacea — Carapace usually covering all gills (if short, gills absent); at least first pair of thoracic appendages modified as maxillipeds; pelagic or benthic ...... 3 3. Shrimp-like; with 6 or 7 pairs of biramous thoracic swimming appendages; females with thoracic oostegites (plates between legs and holding eggs) ...... 4 — Shrimp-, lobster- or crab-like; with 5 pairs of pereopods ...... Decapoda … 5 4. Carapace with clear longitudinal keels and/or spines, covering all thoracic somites; with dominant triangular rostrum; uropods without statocysts ...... Lophogastrida — Carapace usually unornamented, often not covering posterior thoracic somite; with very short flat rostrum; uropods with statocysts at base of endopods (swollen cavities containing granules) ...... Mysida

16 Systematics

5. Prawn-like; plate on side of abdominal somite 2 covering somites in front and behind; gills with branching fine filaments ...... Dendrobranchiata … p. 23 — Shrimp-, lobster- or crab-like; plates on sides of all abdominal somites overlapping somite behind, tile-like (or contiguous); gills with stacked plate-like branches . . . Pleocyemata … 6 6. Crabs; 1 (rarely more) pair of chelipeds and 4 pairs of walking legs; abdomen short and folded under carapace; uropods usually absent ...... Brachyura … p. 289 — Shrimps, lobsters, or secondarily crab-like; abdomen long and muscular or, if short, not engaging closely with underside of carapace; uropods usually present ...... 7 7. First 3 pairs of pereopods chelate, third pair longest ...... Stenopodidea … p. 145 — If chelate pereopods present, first or second pair longest ...... 8 8. Deep-sea lobsters with 4 or 5 pairs of chelate pereopods, first longest and thin; carapace flattened ...... Polychelida … p. 151 — Lobsters, shrimps or crabs with 3 or fewer pairs of chelate limbs (if other limbs chelate of different form) ...... 9 9. Lobsters (crayfish) with 3 pairs of chelate pereopods, first enlarged (fifth sometimes also chelate) ...... Astacidea … p. 159 — Fewer than 3 pairs of chelate pereopods ...... 10 10. Lobsters with no chelate pereopods ...... Achelata … p. 197 — 1 or 2 pairs of chelate pereopods ...... 11 11. Shrimps with 2 pairs of chelate pereopods ...... Caridea … p. 53 — 1 pair of chelipeds, much stronger than remaining walking legs ...... 12 12. Lobster-like; abdomen elongate and muscular; pereopod 2 with dense row of long setae on lower margin; pereopod 5 present, slightly reduced ...... Thalassinidea … p. 169 — Hermit-crab or crab-like; abdomen coiled or flattened; pereopod 2 without dense row of setae; pereopod 5 greatly reduced or absent ...... Anomura … p. 215

References Abzhanov, A., & Kaufman, T.C. 2003. Hox genes and tagmatization of the higher Crustacea (Malacostraca). In: Scholtz, G. (ed.), Evolutionary developmental biology of Crustacea. Crustacean Issues 15: 43–74. Baker, A.d.C., Boden, B.P.,& Brinton, E. 1990. A Practical Guide to the Euphausiids of the World. Natural History Museum Publications: London. 96 pp. Boxshall, G.A., & Halsey, S. 2004. An introduction to copepod diversity. Ray Society Publication 166: 2 volumes. Cals, P., & Delamare-Deboutteville, C. 1970. Une nouvelle èspece de Crustacé de l’hémisphère austral. Comptes Rendus de l’Académie des Sciences, Paris, Sciences de la vie 270: 2444–2447. Dakin, W.J., & Colefax, A.N. 1940. The plankton of the Australian coastal waters off New South Wales Part 1 with special reference to the seasonal distribution, the phyto-plankton, and the planktonic Crustacea, and in particular, the Copepoda and crustacean larvae, together with an account of the more frequent members of the groups Mysidacea, Euphausiacea, Amphipoda, Mollusca, Tunicata, Chaetognathia, and some reference to the fish eggs and fish larvae. Publications of the University of Sydney, Department of Zoology, Monograph 1: 1–215. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. De Deckker, P., & Jones, P.J. 1978. Check list of Ostracoda recorded from Australia and Papua New Guinea 1845–1973. Bureau of Mineral Resources, Geology and Geophysics, Report 195: 1–184.

17 Marine Decapod Crustacea of Southern Australia

Dixon, C.J., Ahyong, S., & Schram, F.R. 2003. A new hypothesis of decapod phylogeny. Crustaceana 76: 935–975. Edgecombe, G.D., Wilson, G.D.F., Colgan, D.J., & Gray, M.R. 2000. Arthorpod cladistics: a combined analysis of histone H3 and U2 sequences and morphology. Cladistics 16: 155–203. Forest, J., & de Saint Laurent, M. 1989. Nouvelle contribution à la connaissance de Neoglyphea inopinata Forest & de Saint Laurent, à propos de la description de la femelle adulte. In: Forest, J. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144: 75–92. Gates, J.E., Stoddart, H.E., & Lowry, J.K. 2003. Hyperiidea. Pp. 298–369 in: Beesley, PL., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.2B. CSIRO Publishing: Melbourne. Giribet, G., Edgecombe, G.D., & Wheeler, W.C. 2001. Arthropod phylogeny based on eight molecular loci and morphology. Nature 413: 157–161. Hamond, R. 1973. Four new copepods (Crustacea: Harpacticoida, Canuellidae) simultaneously occurring with Diogenes senex (Crustacea: Paguridea) near Sydney. Proceedings of the Linnean Society of New South Wales 97: 165–201. Harris, V.A. 2002. A new genus belonging to the family Porcellidiidae (Crustacea: Copepoda: Harpacticoida) with three new species from Australia. Records of the Australian Museum 54: 1–24. Hawking, J.H. 2000. Key to keys: A guide to keys and zoological information to identify invertebrates from Australian inland waters. Co-operative Research Centre for Freshwater Ecology Identification Guide 2: 1–71. Hessler, R.R. 1992. Reflections on the phylogenetic position of the Cephalocarida. Acta Zoologica, Stockholm 73: 315–316. Hessler, R.R., & Elofsson, F.W. 1992. Cephalocarida. Pp. 9–24 in: Harrison, F.W. (ed.) Microscopic Anatomy of Invertebrates. Wiley-Liss: New York. Høeg, J.T. 1995. The biology and life cycle of the Rhizocephala (Cirripedia). Journal of the Marine Biological Association of the United Kingdom 75: 517–550. Høeg, J.T., & Lützen, J. 1995. Life cycle and reproduction in the Cirripedia Rhizocephala. Oceanography and Marine Biology, Annual Review 33: 427–485. Horwitz, P.,Knott, B., & Williams, W.D. 1995. A preliminary key to the malacostracan families (Crustacea) found in Australian inland waters (Co-operative Research Centre for Freshwater Ecology Identification Guide 4). Cooperative Research Centre for Freshwater Ecology: Albury, NSW. 31 pp. Huys, R., & Boxshall, G.A. 1991. Copepod evolution. The Ray Society: London. 468 pp. Jarman, S.N., & Elliott, N.G. 2000. DNA evidence for morphological and cryptic Cenozoic speciations in the Anaspididae, ‘living fossils’ from the Triassic. Journal of Evolutionary Biology 13: 624–633. Jones, D.S. 1990. The shallow-water barnacles (Cirripedia: Lepadomorpha, Balanomorpha) of southern Western Australia. Pp. 333–437 in: Wells, F.E., Walker, D.I., Kirkman, H., & Lethbridge, R. (eds), Proceedings of the Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia. Western Australian Museum: Perth. Jones, D.S. 1991. A history of the discovery and description of Australian barnacles (Cirripedia: Thoracica), including a bibliography of reference works. Archives of Natural History 18: 149–178. Jones, D.S. 1992. A review of Australian fouling barnacles. Asian Marine Biology 9: 89–100. Just, J., & Poore, G.C.B. 1988. Second record of Hirsutiidae (Peracarida: Mictacea): Hirsutia sanderse- talia, new species, from southeastern Australia. Journal of Crustacean Biology 8: 483–488. Kabata, Z. 1970. Copepoda parasitic on Australian fishes X. families Educatylinidae and Pseudocycnidae. Journal of Natural History 4: 159–173. Keable, S.J., Fenton, G.E., & Lowry, J.K. 2003. Mysidacea. Pp. 419–471 in: Beesley, P.L., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol 19.2B. CSIRO Publishing: Melbourne. Knox, G.A., & Fenwick, G.D. 1977. Chiltoniella elongata n. gen. et sp. (Crustacea: Cephalocarida) from New Zealand. Journal of the Royal Society of New Zealand 7: 425–432. Koenemann, S., Iliffe, T.M., & Ham, J.v.d. 2003. Three new sympatric species of Remipedia (Crustacea) from Great Exuama Island, Bahamas Islands. Contributions to Zoology 72: 227–252.

18 Systematics

Kornicker, L.S. 1994. Ostracoda (Myodocopina) of the SE Australian continental slope, part 1. Smithsonian Contributions to Zoology 553: 1–200. Kornicker, L.S. 1995. Ostracoda (Myodocopina) of the SE Australian continental slope, part 2. Smithsonian Contributions to Zoology 562: 1–97. Kornicker, L.S. 1996. Ostracoda (Myodocopina) from shallow waters of the Northern Territory and Queensland, Australia. Smithsonian Contributions to Zoology 578: 1–97. Kornicker, L.S., & Poore, G.C.B. 1996. Ostracoda (Myodocopina) of the SE Australian continental slope, part 3. Smithsonian Contributions to Zoology 573: 1–186. Lake, P.S., Poore, G.C.B., & Lew Ton, H.M. 2002. Order: Anaspidacea Calman, 1904. Pp. 9–18 in: Houston, W.W.K., & Beesley, P.L. (eds), Zoological Catalogue of Australia. Vol 19.2A. CSIRO Publishing: Melbourne. Lowry, J.K. 2003. Amphipoda. Pp. 7–18 in: Beesley, P.L.,& Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol 19.2B. CSIRO Publishing: Melbourne. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. Mauchline, J. 1980. The biology of mysids and euphausiids. Advances in Marine Biology 18: 1–680. McKinnon, A.D., Kimmerer, W.J., & Benzie, J.A.H. 1992. Sympatric sibling species within the genus Acartia (Copepoda: Calanoida): a case study from Westernport and Port Phillip Bays, Australia. Journal of Crustacean Biology 12: 239–259. Parker, A.R. 1998. A new genus and two new species of Cypridinidae (Crustacea: Ostracoda: Myodocopina) from Australia. Records of the Australian Museum 50: 1–17. Poore, G.C.B. 2002. Crustacea: Malacostraca: Syncarida, Peracarida: Isopoda, Tanaidacea, Mictacea, Thermosbaenacea, Spelaeogriphacea. In: Houston, W.W.K., & Beesley, P.L. (eds), Zoological Catalogue of Australia. Vol. 19.2A. CSIRO Publishing: Melbourne. xii, 429 pp. Poore, G.C.B., & Humphreys, W.F. 1992. First record of Thermosbaenacea (Crustacea) from the Southern Hemisphere: a new species from a cave in tropical Western Australia. Invertebrate Taxonomy 6: 719–725. Poore, G.C.B., & Humphreys, W.F. 2003. Second species of Mangkurtu (Spelaeogriphacea) from north- western Australia. Records of the Western Australian Museum 22: 67–74. Poore, G.C.B., & Lew Ton, H.M. 2002. Order: Bathynellacea Chappuis, 1915. Pp. 19–23 in: Houston, W.W.K., & Beesley, P.L. (eds), Zoological Catalogue of Australia. Vol 19.2A. CSIRO Publishing: Melbourne. Poore, G.C.B., Lew Ton, H.M., & Sieg, J. 2002. Order: Tanaidacea Hansen, 1895. Pp. 346–374 in: Houston, W.W.K., & Beesley, P.L. (eds), Zoological Catalogue of Australia. Vol 19.2A. CSIRO Publishing: Melbourne. de Saint Laurent, M. 1979. Vers une nouvelle classification des Crustacés Décapodes Reptantia. Bulletin de l’Office Nationale de Pêche de Tunisie 3: 15–31. Sanders, H.L. 1963. Significance of the Cephalocarida. Museum of Comparative Zoology, Special Publication 13: 163–179. Scholtz, G., & Richter, S. 1995. Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113: 289–328. Schram, F.R. 2001. Phylogeny of decapods: moving towards a consensus. Hydrobiologia 449: 1–20. Schram, F.R., & Ahyong, S. 2002. The higher affinities of Neoglyphea inopinata in particular and the Glypheoidea (Decapoda, Reptantia) in general. Crustaceana 75: 629–636. Schram, F.R.,& Koenemann, S. 2003. Developmental genetics and arthropod evolution: on body regions of Crustacea. In: Scholtz, G. (ed.), Evolutionary developmental biology of Crustacea. Crustacean Issues 15: 75–92. Schram, F.R., & Koenemann, S. 2004. Shrimp cocktail: are the crustaceans monophyletic? In: Cracraft, J., & Donaghue, M.J. (eds), Assembling the tree of life, science, relevance, and challenges. American Museum of Natural History: New York. Schram, F.R.,Yager, J., & Emerson, M.J. 1986. Remipedia. Part I. Systematics. San Diego Society of Natural History, Memoir 15: 1–60.

19 Marine Decapod Crustacea of Southern Australia

Spears, T.,Abele, L.G., & Kim, W. 1992. The monophyly of brachyuran crabs: a phylogenetic study based on 18S rRNA. Systematic Biology 41: 446–461. Spears, T., & Abele, L.G. 1998. Crustacean phylogeny inferred from 18S rDNA. Pp. 169–188 in: Fortey, R.A., & Thomas, R.H. (eds), Arthropod Relationships. The Systematics Association Special Volume Series 55. Chapman and Hall: London. Stoddart, H.E., Keable, S.J., & Lowry, J.K. 2003. Gammaridea. Pp. 39–297 in: Beesley, P.L., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Crustacea: Malacostraca: Amphipoda, Cumacea, Mysidacea. CSIRO Publishing: Melbourne. Stoddart, H.E., & Lowry, J.K. 2003a. Cumacea. Pp. 373–418 in: Beesley, P.L., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol 19.2B. CSIRO Publishing: Melbourne. Stoddart, H.E., & Lowry, J.K. 2003b. Ingolfiellidea. Pp. 370–372 in: Beesley, P.L., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol 19.2B. CSIRO Publishing: Melbourne. Stoddart, H.E., & Lowry, J.K. 2003c. Caprellidea. Pp. 19–38 in: Beesley, P.L., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol 19.2B. CSIRO Publishing: Melbourne. Tavares, M.S. 2003. A new theoretical approach for the study of monophyly of the Brachyura (Crustacea: Decapoda) and its implication for the Anomura. Memoirs of Museum Victoria 60: 145–149. Tudge, C.C. 1997. Phylogeny of the Anomura (Decapoda, Crustacea): Spermatozoa and spermatophore morphological evidence. Contributions to Zoology 67: 125–141. Walker-Smith, G.K. 1998. A review of Nebaliella (Crustacea: Leptostraca) with description of a new species from the continental slope of southeastern Australia. Memoirs of the Museum of Victoria 57: 39–56. Walker-Smith, G.K. 2000. Levinebalia maria, a new genus and new species of Leptostraca (Crustacea) from Australia. Memoirs of the Museum of Victoria 58: 137–147. Walker-Smith, G.K. 2001. Porcellidium poorei, a new species of Porcelliidae (Copepoda: Harpacticoida) from seagrass in Port Phillip Bay, Victoria, Australia, and a review of the family. Journal of Crustacean Biology 21: 653–664. Walker-Smith, G.K., & Poore, G.C.B. 2001. A phylogeny of the Leptostraca (Crustacea) with keys to the families and genera. Memoirs of the Museum of Victoria 58: 383–410. Williams, W.D. 1980. Australian Freshwater Life. Macmillan: Melbourne. pp. xi, 321 pp. 2nd edn. Yager, J., & Humphreys, W.F. 1996. Lasionectes exleyi, sp. nov., the first remipede crustacean recorded from Australia and the Indian Ocean, with a key to the world species. Invertebrate Taxonomy 10: 171–187.

20 Systematics barnacles ing seed shrimps, ostracods goose barnaclesgoose water fleas water clam shrimpsclam copepods mantis shrimps y-larvae tantulocarids fish lice worms tongue mystacocarids fairy shrimps cephalocarids parasitic barnacles ascothoracican barnaclesascothoracican shield shrimps remipedes leptostracans, nebaliaceans acorn barnaclesacorn goose barnacles, wart barnacles, uborder Infraorder name Common Unipeltata Cladocera cea (Stomatopoda and Decapoda covered in this text shaded) in this text (Stomatopoda and Decapodacea covered Brachypoda Stomatopoda Sessilia Diplostraca 3 suborders Nectipoda 4 orders 10 orders Leptostraca Notostraca Thoracica Pedunculata 4 suborders Rhizocephala Anostraca Subclass/Infraclass Superorder Order S Hoplocarida Tantulocarida Branchiura Pentastomida Mystacocarida Copepoda Phyllopoda Thecostaca/Ascothoracica Thecostaca/Cirripedia bor Acrothoracica Malacostraca Phyllocarida Branchiopoda Sarcostraca Maxillipoda Thecostaca/Facetotecta Cephalocarida Remipedia Ostracoda 2 subclasses Table 2.Table A classification of living members of Crusta the subphylum Class

21 Marine Decapod Crustacea of Southern Australia ed) tanaidaceans lophogastrids spelaeogriphaceans amphionidaceans bathynellaceans shrimps squat lobsters thermosbaenaceans krill mountain shrimps opposum shrimps cumaceans prawns mictaceans isopods, slaters, woodlice amphipods, shrimps, skeleton whale lice StenopodideaPolychelida shrimps coral Astacidea deep-sea lobsters Thalassinidea lobsters, mud ghost and sponge lobsters, crayfish, yabbies AchelataAnomura lobsters, rock bugs Brachyura hermit crabs, crabs, porcelain crabs Pleocyemata Caridea shrimps Dendrobranchiata Suborder Infraorder name Common 10 suborders m Crustacea (Stomatopoda and Decapoda covered in this text shad in this text (Stomatopoda and Decapodam Crustacea covered Decapoda Anaspidacea Thermosbaenacea Lophogastrida Mysida Mictacea AmphipodaIsopoda 4 suborders TanaidaceaCumacea 3 suborders Amphionidacea Peracarida Spelaeogriphacea Eucarida Euphausiacea Eumalacostraca Syncarida Bathynellacea Table 2 (cont.).Table A classification of living members of the subphylu Class Subclass/Infraclass Superorder Order

22 3. DENDROBRANCHIATA – PRAWNS AND MIDWATER SHRIMPS

Prawns are the most familiar of all crustaceans to Australians. Solely because these feature on the menu. But prawns belong to just one of the seven families of Dendrobranchiata – the rest are rarely encountered. This chapter deals with one of the two major divisions of the Decapoda; all the others treat the other division, Pleocyemata. The Dendrobranchiata, therefore, are fundamentally different from all the others anatomically and ecologically. The essential anatomical character is found in the gills (branchia) under the sides of the carapace. In dendrobranchiates, the gills are ‘dendrobranchiate’ or composed or many branching filaments attached to a major stem. In all other pleocyemate decapods, crabs, shrimps, lobsters etc, the gills look like a stack of plates in a row along a main axis. The dendrobranchiate reproductive strategy also differs from the Pleocyemata. Eggs are released from the female and develop in the plankton until it is time to settle. Crabs and their relatives brood their eggs by attaching them to setae on the pleopods until they hatch as zoeal larvae. Prawns superficially look like caridean shrimps and in midwater samples examples of both often occur together. Carideans, besides having plate-like gills, differ from prawns in having the second abdominal pleuron (plate extending ventrally to shelter the pleopods) covering both the pleuron in front and behind. In dendrobranchiates, the abdominal pleura overlap like tiles, the first over the second, second over the third and so on. Also, dendrobranchiates have three pairs of claws (pereopods 1–3 chelate, although minutely so) while carideans usually have two pairs of chelate pereopods, the first often enlarged as claws. Identification of dendrobranchiates worldwide is facilitated by the well-illustrated and explained keys to families and genera of Pérez Farfante & Kensley (1997). This contribution draws heavily on their treatment. While their family treatment is agreed on by most modern authors, there is less sympathy for their genera of penaeids (see below). Other keys are available, especially for species of commercial importance (Holthuis, 1980; Burkenroad, 1983; Burukovskii, 1983; Dore & Frimodt, 1987). Burkenroad’s (1983) discussion of a phylogeny of the Dendrobranchiata is responsible for the current classification into superfamilies. His earlier work was responsible for the separation of dendrobranchiates from other decapods and the abandoning of divisions between natant (swimming) and reptant (walking) taxa. Because of their commercial significance, prawns have been well studied in Australia and elsewhere. The valuable guide by Grey et al. (1983) includes keys and colour photographs of commercially important Australian species.

Diagnosis. Abdominal pleuron 1 overlaps pleuron 2. Pereopods 1–3 chelate. Gills dendrobranchiate. Pleurobranchs appear latter in development than podobranchs and arthrobranchs. Appendices internae absent from pleopods, except on pleopods 1 and 2 of some males.

23 Marine Decapod Crustacea of Southern Australia

Key to superfamilies and families of Dendrobranchiata 1. At least 11 branchia (gills) on each side (some somites with at least 3 branchia on each side) ...... Penaeoidea … 2 — Never more than 8 branchia (gills) on each side (somites with at most 2 branchia on each side) ...... Sergestoidea … 6 2. Postorbital spine present ...... Solenoceridae … p. 43 — Postorbital spine absent ...... 3 3. Carapace rigid, with median dentate crest extending to posterior margin; pleopods with 1 ramus ...... Sicyoniidae … p. 41 — Carapace flexible, with median dentate crest extending only to mid-carapace; pleopods with 2 rami ...... 4 4. 1 or 2 (rarely 3) rostral and postrostral teeth ...... Benthesicymidae … p. 27 — More than 2 rostral and postrostral teeth ...... 5 5. Prosartema well developed (short branch on inner side of antennule, inside eyes) ...... Penaeidae … p. 31 — Prosartema reduced to setose boss ...... Aristeidae … p. 24 6. Branchia absent; body strongly compressed laterally ...... Luciferidae … p. 45 — Branchia present ...... Sergestidae … p. 46

References Burkenroad, M.D. 1983. Natural classification of Dendrobranchiata, with a key to Recent genera. In: Schram, F.R., Crustacean phylogeny. Crustacean Issues 6: 279–290. Burukovskii, R.N. 1983. Key to Shrimps and Lobsters. A.A. Balkema: Rotterdam. 174 pp. translation of 1974 Russian edn. Dore, I., & Frimodt, C. 1987. An Illustrated Guide to Shrimp of the World. Osprey Books and Scandinavian Fishing Year Book: Huntington and Hedehusene. Grey, D.L., Dall, W., & Baker, A. 1983. A Guide to the Australian Penaeid Prawns. Northern Territory Government Printing Office: Darwin. 140 pp. Holthuis, L.B. 1980. FAO species catalogue. Vol.1. Shrimps and prawns of the world. An annotated catalogue of species of interest to fisheries. FAO Fisheries Synopsis 1 (125): 1–261. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233.

Superfamily Penaeoidea Rafinesque-Schmaltz, 1815 Five families are included: Aristeidae, Benthesicymidae, Penaeidae, Sicyoniidae and Solenoceridae.

Aristeidae Wood-Mason, 1891 Pelagic shrimps of this family are deep-sea mesopelagic species found off the edge of the continental shelf at depths below 300 m. Some are of commercial interest because of their large size, up to 350 mm length, and high local densities. None occurs in shallow water. Some species may be confused superficially with oplophorid shrimps but can be separated on the dendrobranchiate gills and abdomen. A key character distinguishing the family from other

24 Dendrobranchiata – prawns and midwater shrimps

prawns is the disparity between the lengths of flagella of the antennule (first antenna). The taxonomy of the family in Australia was revealed by Dall (2001). He reported on 12 species representing all nine genera. Fewer have been reported from southern Australia. Pérez Farfante & Kensley (1997) diagnosed the family and genera in more detail than given here.

Diagnosis. Prosartema (short branch on inner side of antennule, inside eyes) reduced to a setose knob. Carapace with dentate crest not extending beyond middle of carapace. Upper flagellum of antennules markedly shorter than other and not attached together at tip of third article.

Key to species of Aristeidae from southern Australia 1. Rostrum with 5–9 dorsal teeth, hepatic spine present ...... Aristaeomorpha foliacea — Rostrum with usually 3 dorsal teeth, hepatic spine absent ...... 2 2. Pereopods with reduced exopods (Fig. 3e) ...... Austropenaeus nitidus — Pereopods without exopods ...... 3 3. Pereopods 1 and 2 without meral spines; scaphocerite in male elongate ...... Aristaeopsis edwardsiana — Pereopods 1 and 2 with subdistal meral spines; scaphocerite in male not elongate ...... 4 4. Cervical groove distinct and extending to top of carapace; postcervical groove present ...... Pseudaristeus sibogae — Cervical groove short, confined to hepatic region; postcervical groove absent . . . Aristeus … 5 5. Body and pereopods finely pubescent; pereopod 3 merus with mobile subdistal spine on lower margin ...... Aristeus virilis — Body and pereopods smooth; pereopod 3 merus without spine on lower margin ...... 6 6. Pereopod 5 merus as long as carpus; carpus with 4–8 photophores . . . . Aristeus mabahissae — Pereopod 5 merus shorter than carpus; carpus with 14–21 photophores ...... Aristeus semidentatus

Aristaeomorpha Wood-Mason, 1891 A single species of this genus occurs in southern Australia and is the most commonly recorded of the family. Diagnosis. Rostrum with 5–9 dorsal teeth. Hepatic spine present. Aristaeomorpha foliacea (Risso, 1827) giant prawn, red prawn, royal red prawn (Fig. 3a). Deep pink to wine-red (Grey et al., 1983). Females 255 mm, males 170 mm. Mediterranean, Indo- West Pacific, Atlantic, NSW, Vic., Tas., SA, WA; bathypelagic on continental slope, muddy bottoms, to 2000 m, usually 250–700 m depth. The species is commercially fished and its biology well studied (e.g. Cau et al., 2002; Kapiris et al., 2002)

Aristaeopsis Wood-Mason, 1891 There is only one species. Diagnosis. Rostrum with 3 dorsal teeth. Hepatic spine absent. Pereopods 1 and 2 without meral spines. Scaphocerite in male elongate. Pereopods without exopods. Aristaeopsis edwardsiana (Johnson, 1867) scarlet prawn (Fig. 3b). Females 350 mm, males 200 mm. Indo-West Pacific, Atlantic, Qld, NSW, Tas., WA; bathypelagic on continental slope, muddy bottoms, 200–1850 m, usually 450–600 m depth. The species was recorded as Plesiopenaeus edwardsianus by Kensley et al. (1987).

25 Marine Decapod Crustacea of Southern Australia

b c

d e

Fig. 3. Aristaeidae. a, Aristaeomorpha foliacea. Carapace: b, Aristaeopsis edwardsiana. c, Aristeus mabahissae. d, Austropenaeus nitidus. e, basis of pereopod with epipod and rudimentary exopod.

Aristeus Duvernoy, 1840 The weak cervical groove characterises the eight species, separated on subtle differences in proportions of articles of the pereopods (Dall, 2001). Only two species are certain in southern Australia. If legs are present and if preserved material is fresh photophores can help identification. They are visible in oblique light as round structures but invisible in old museum specimens. Diagnosis. Rostrum with 2 dorsal teeth plus 1 postrostral. Hepatic spine absent. Cervical groove short, confined to hepatic region; postcervical groove absent. Pereopods without exopods. Aristeus mabahissae Ramadan, 1938 shiny pink-striped prawn (Fig. 3c). cl. 53 mm. Indo-West Pacific, WA, NSW; mesopelagic, bathypelagic, continental slope, 366–1100 m, mainly 500–750 m depth. Aristeus semidentatus Bate, 1881. Australian records of this Indo-West Pacific species are probably attributable to A. mabahissae (Dall, 2001). Aristeus virilis (Bate, 1881) stout red shrimp. cl. 57 mm. Indo-West Pacific, WA, NT, Qld, NSW, Vic.; 188–1050 m, mainly 350–550 m depth.

26 Dendrobranchiata – prawns and midwater shrimps

Austropenaeus Pérez Farfante & Kensley, 1997 The featureless carapace and rudimentary exopods characterise the single species, one confined to more southern latitudes. Diagnosis. Rostrum with 2 dorsal teeth plus 1 postrostral. Hepatic spine absent. Cervical groove short, confined to hepatic region; postcervical groove absent; carinae weak. Pereopods with rudimentary exopods; pereopod 3 with rudimentary podobranch. Austropenaeus nitidus (Barnard, 1947) (Fig. 3d). cl. 38 mm. South Atlantic, Indian Ocean, NSW, Vic., SA, WA; abyssopelagic on continental slope, muddy bottoms, 457–1300 m, usually 1000–1300 m depth.

Pseudaristeus Crosnier, 1978 Six species are known but are difficult to distinguish. Diagnosis. Rostrum with 2 dorsal teeth plus 1 postrostral. Hepatic spine absent. Cervical groove distinct and reaching top of carapace; postcervical groove present. Pereopods without exopods. Pseudaristeus sibogae (de Man, 1911). Pereopods covered with minute setae. Eyestalk mesial margin 1.4 times width of distal margin. Orange. cl. 43 mm. Indian Ocean, SA, WA; bathypelagic, oceanic; 834–1305 m depth.

References Cau, M., Carbonell, A.C., Follesa, M.C., Mannini, A., Norrito, G., L., O.-R., Politou, C.-Y., Ragonese, S., & Rinelli, P.2002. MEDITS-based information on the deep-water red shrimps Aristaeomorpha foliacea and Aristeus antennatus (Crustacea: Decapoda: Aristeidae). Scientia Marina 66 (Suppl. 2): 103–124. Dall, W. 2001. Australian species of Aristeidae and Benthesicymidae (Penaeoidea: Decapoda). Memoirs of the Queensland Museum 46: 409–441. Grey, D.L., Dall, W., & Baker, A. 1983. A Guide to the Australian Penaeid Prawns. Northern Territory Government Printing Office: Darwin. 140 pp. Kapiris, K., Moraitou-Apostolopoulou, M., & Papaconstantinou, C. 2002. The growth of male secondary sexual characters in Aristaeomorpha foliacea and Aristeus antennatus (Decapoda, Aristeidae) in the Greek Ionian Sea (Eastern Mediterranean). Journal of Crustacean Biology 22: 784–789. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233.

Benthesicymidae Wood-Mason, 1891 Benthesicymid shrimps are, like aristeids, pelagic in deep offshore water. Populations in the Eastern Australian Current and the eddies that it generates were followed by Griffiths and cowork- ers (Griffiths & Brandt, 1983a, b; Griffiths & Wadley, 1986). Many species have been collected over great depth ranges, from the surface down to thousands of metres and it is assumed that they migrate vertically. Four genera are recognised, diagnosed with lists of species by Pérez Farfante & Kensley (1997). Dall (2001) provided keys to 15 species of Benthesicymus and nine species of Gennadas from the Indo-West Pacific. The key to species relies on differentiating males and females and is from Dall (2001). Males possess a petasma, a structure formed from enlarged and modified endopods of pleopods 1. The

27 Marine Decapod Crustacea of Southern Australia

pair of endopods hook together in the middle with clusters of hook-like cincinnuli. Couplets 3–9 differentiating males of Gennadas refer to lobes on the petasma (Fig. 4). Basically each half of the petasma has three lobes, an inner with the cincinnuli, a middle with an accessory lobe on its inner anterior margin, and an outer lobe but each lobe may be divided into two lobules. Females lack a petasma but have instead a thelycum, modified thoracic sternites between pereopods 4 and 5. Couplets 10–16 differentiating females refer to variously shaped plates on thoracic sternites 5 to 8, those between pereopods 2 to 5 (Fig. 4). A pair of seminal receptacles is often visible between sternites 6 and 7.

Diagnosis. Prosartema (short branch on inner side of antennule, inside eyes) absent. Carapace with dentate crest extending only to centre. Upper antennular flagellum as long as or longer than lower one.

Key to southern Australian genera and species of Benthesicymidae 1. Rostrum with 2 dorsal teeth; telson with 4 pairs of lateral moveable spines; podobranchs on maxilliped 2 – pereopod 3 ...... Benthesicymus investigatoris — Rostrum with 1 dorsal tooth; telson with 1 pair of lateral moveable spines; podobranch only on maxilliped 2 ...... Gennadas … 2 2. Males (with petasma, without thelycum) ...... 3 — Females (without petasma, with thelycum) ...... 10 3. Middle lobe undivided ...... 4 — Middle lobe divided ...... 6 4. Outer lobe divided by close blunt lobules ...... Gennadas tinayrei — Outer lobe undivided or with small medial process ...... 5 5. Accessory lobe bipartite ...... Gennadas capensis — Accessory lobe a single flap ...... Gennadas kempi 6. Lobules of outer lobe elongate, subequal, slender ...... Gennadas incertus — Lobules of outer lobe not elongate, unequal, not slender ...... 7 7. Lobules of middle lobe hooked ...... Gennadas bouvieri — Lobules of middle lobe not hooked ...... 8 8. Apex of inner lobe acute ...... Gennadas gilchristi — Apex of inner lobe rounded ...... 9 9. Inner lobule of middle lobe with acute apex ...... Gennadas propinquus — Inner lobule of middle lobe with truncate apex ...... Gennadas scutatus 10. Posteriorly directed tongue-like projection on sternite 5 ...... Gennadas tinayrei — Without tongue-like projection on sternite 5 ...... 11 11. Sternite 8 without plate ...... Gennadas capensis — Sternite 8 with well developed plate ...... 12 12. Plate on sternite 8 with anterior flap reaching sternite 6 ...... Gennadas scutatus — Plate on sternite 8 not reaching sternite 6 ...... 13 13. Plate on sternite 8 broad, anteriorly notched and overlapping sternite 7 . . Gennadas incertus — Plate on sternite 8 not anteriorly notched and not overlapping sternite 7 ...... 14 14. Plate on sternite 8 with 2 anteriorly-directed slender projections ...... Gennadas bouvieri — Plate on sternite 8 without 2 anteriorly-directed slender projections ...... 15

28 Dendrobranchiata – prawns and midwater shrimps

15. Plate on sternite 7 with 2 medial anterior projections ...... Gennadas gilchristi — Plate on sternite 7 without 2 medial anterior projections ...... 16 16. Plate on sternite 7 rectangular ...... Gennadas kempi — Plate on sternite 7 W-shaped ...... Gennadas propinquus

b c

d e

f g

Fig. 4. Benthesicymidae. a, Benthesicymus investigatoris. Petasma, upper right side, and thelycum: b, Gennadas bouvieri. c, Gennadas capensis. d, Gennadas gilchristi. h, Gennadas incertus. f, Gennadas kempi. g, Gennadas propinquus. h, Gennadas scutatus. i, Gennadas tinayrei.

29 Marine Decapod Crustacea of Southern Australia

Benthesicymus Bate, 1881 Most species of Benthesicymus live in the tropical Indo-West Pacific but only two have been discovered in Australia (Dall, 2001). Diagnosis. Telson with 4 pairs of lateral moveable spines. Podobranchs on maxilliped 2 to pereopod 3. Median carina present on abdominal somites 5–6, usually but weaker on somite 4. Benthesicymus investigatoris Alcock & Anderson, 1899 (Fig. 4a). Rostrum with 2 teeth. Branchiostegal spine behind margin of carapace, branchiostegal carina sharp, hepatic spine absent. Abdominal somite 5 with short posterodorsal spine; somite 6 with posterodorsal tip horizontal. Maxilliped 3 with dactylus subrectangular, having >1 spine. cl. 21 mm. Indo-West Pacific, WA, Qld, NSW; 580–1690 m depth.

Gennadas Bate, 1881 Gennadas is the most diverse genus of the Benthesicymidae with 18 species, half of which occur in the Indo-West Pacific (Pérez Farfante & Kensley, 1997; Dall, 2001). All except one of the Indo-West Pacific species occur off south-eastern Australia. Development passes through a mysis larval stage (Rivera & Guzman, 2002). Diagnosis. Rostrum not reaching end of eyestalk, with 1 dorsal tooth. Telson with 1 pair of lateral moveable spines. Podobranch only on maxilliped 2. Median carina only on abdominal somite 6. Gennadas bouvieri Kemp, 1909 (Fig. 4b). Female: plate on sternite 8 bearing 2 anteriorly directed projections, plate 7 attached to 6. Male: outer lobe divided in 2 broad lobules, middle lobe divided and hooked, inner lobe much shorter than middle. cl. 9 mm. Atlantic, Indo-Pacific, NSW, Vic., Tas., 17–39°S; meso- and bathypelagic, to 1988 m depth. Gennadas capensis Calman, 1925 (Fig. 4c). Female: plate on sternite 7 W-shaped, coxa 5 expanded and bilobed, coxa 4 with elongate process. Male: all lobes undivided, median lobe shorter than others, accessory lobe divided. cl. 13 mm. Atlantic, Indo-West Pacific, Qld, NSW, Vic., 22–39°S; meso- and bathypelagic, to 2000 m depth. Gennadas gilchristi Calman, 1925 (Fig. 4d). Female: plate on sternite 7 with pair of anterior projections, pair of red circular seminal receptacles in front; plate on sternite 8 apple-shaped., overlapping 7; coxa 3 with bilobed expansions. Male: outer lobe acute with smaller lobule at outer base, middle lobe with diverging slender lobules, inner lobe acute. cl. 7 mm. Southern Ocean, Indian Ocean, NSW,Vic., Tas., SA, S of 33°S; bathypelagic, to 3400 m, mostly 500–700 m depth. This is the commonest species of Gennadas in Australian waters (Griffiths & Brandt, 1983a; Kensley et al., 1987). Gennadas incertus (Balss, 1927) (Fig. 4e). Female: plate on sternite 8 indented anteriorly, overlapping sternite 7, plate on sternite 7 with acute anterolateral corners, somite 6 with large concave plate. Male: outer lobe divided into 2 long tapering lobules, middle lobe divided into 2 rounded lobules. cl. 8 mm. Atlantic, Indo-Pacific, NSW, 17–34°S; meso- and bathypelagic, 100–1406 m depth. Gennadas kempi Stebbing, 1914 (Fig. 4f). Female: plate on sternite 8 hexagonal, on sternite 7 subrectangular, with concave anterior margin, on sternite 6 subtriangular. Male: external, middle and inner lobes undivided, outer lobe acute, middle lobe broad and truncate, inner lobe truncate, accessory lobe a simple flap. cl. 10 mm. Southern Ocean, South Atlantic, Indian Ocean, Qld, NSW, Tas., SA, S of 32°S; meso- and bathypelagic, 250–3400 m, mostly 550– 640 m depth. Gennadas propinquus Rathbun, 1906 (Fig. 4g). Female: all 3 lobes divided apically, outer lobe with outer lobule acute, inner lobule rounded, middle lobe with prominent lobules, inner lobe with both lobules rounded, accessory lobe a simple flap. Male: ovate plate on sternite 8 excavate

30 Dendrobranchiata – prawns and midwater shrimps

anteriorly and posteriorly, plate on sternite 7 W-shaped, plate on sternite 6 with indented anterior apex. cl. 10 mm. Atlantic, Indo-West Pacific, NSW, Tas., S of 33°S; meso- and bathypelagic, to 1200 m depth. Griffiths & Brandt (1983a) recorded this species (as G. clavicarpus) associated with a warm-core eddy off eastern Australia. Gennadas scutatus Bouvier, 1906 (Fig. 4h). Female: plate on sternite 8 with wide rounded anterior flap extending to cover half sternite 6, plate on sternite 7 a wide flattened triangle, plate on sternite 6 triangular, with paired seminal receptacles at base, coxa 4 with inward-directed setae. Male: outer lobe divided into 2 rounded lobules, middle lobe with slender outer lobule, inner lobule with small accessory lobule, inner lobe divided into 2 rounded lobules, accessory lobe a simple flap. cl. 7 mm. Atlantic, Indo-West Pacific, NSW,Vic., 17–38°S; meso- and bathypelagic, surface to 3400 m, usually to 1200 m depth. Gennadas tinayrei Bouvier, 1906 (Fig. 4i). Female: plate on sternite 8 subrectangular, plate on sternite 7 a wide triangle with anterior processes, pair of plates on sternite 6, plate on sternite 5 projected posteriorly. Male: outer lobe slightly indented, edged with minute teeth and small lobule, middle lobe broad and convex, inner lobe with 2 rounded projections, accessory lobe rounded. cl. 7 mm. Atlantic, Indo-West Pacific, NSW, Tas., 33–42°S; mesopelagic, 92–950 m depth.

References Dall, W. 2001. Australian species of Aristeidae and Benthesicymidae (Penaeoidea: Decapoda). Memoirs of the Queensland Museum 46: 409–441. Griffiths, F.B., & Brandt, S.B. 1983a. Distribution of mesopelagic decapod Crustacea in and around a warm-core eddy in the Tasman Sea. Marine Ecology Progress Series 12: 175–184. Griffiths, F.B., & Brandt, S.B. 1983b. Mesopelagic Crustacea in and around a warm-core eddy in the Tasman Sea off eastern Australia. Australian Journal of Marine and Freshwater Research 34: 609–623. Griffiths, F.B., & Wadley, V.A. 1986. A synoptic comparison of fishes and crustaceans from a warm-core eddy, the East Australian Current, the Coral Sea and the Tasman Sea. Deep-Sea Research 33: 1907–1922. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233. Rivera, J., & Guzman, G. 2002. Description of mysis larvae of three mesopelagic shrimp species of the genus Gennadas (Decapoda: Aristeidae) from the southeastern Pacific waters. Investigaciones Marinas, Ciencias 30: 33–44.

Penaeidae Rafinesque-Schmaltz, 1815 It would make sense to call all members of the Penaeidae ‘prawns’ and members of the Caridea ‘shrimps’ and this is what most Australians do. King prawns and banana prawns are names understood in this continent for penaeids sold frozen or cooked in fish markets or caught in estuaries with trawl nets. The tiny shrimps bought in cans or frozen are imported carideans. The only confusion arises when we hear Americans refer to prawns as ‘shrimp’! Most of the edible prawns belong to Penaeidae. All have a strong serrated rostrum, a laterally flattened body and abdomen, three pairs of minutely chelate legs, and abdominal somites overlapping tile-like from the front. The last features distinguish prawns from the much smaller caridean shrimps who have at most only two pairs of chelate legs, and the second abdominal somite overlapping both in front and behind.

31 Marine Decapod Crustacea of Southern Australia

The prawns sold in markets are caught with trawl nets swimming above the bottom in the muddy waters of shallow bays and estuaries, mostly in the tropics. Sixty-six species have been recorded from Australia but the number that are commercially important is much smaller, about nine or ten. They go by such names in the Australian market as: bay prawn, greasy back prawn, school prawn, banana prawn, king prawn, tiger prawn, rough prawn and others. Many of these names refer to more than one species and some are prefixed by other adjectives, such as the blue- legged, western, eastern and redspot king prawns, or brown, blue, green and jumbo tiger prawns. While Australians have their own colourful names for the edible prawns found in this country, the FAO recommends its own ‘international’ common names (Holthuis, 1980). While confusion over what common name to use can be expected, because many names are only locally applicable, more certainty could be expected with taxonomic names. Unfortunately, this is not the case. Most commercial prawns have traditionally been placed in the genus Penaeus and only a few smaller genera recognised. Following a taxonomic revision based on morphological investigation by Pérez Farfante & Kensley (1997), Penaeus was divided into six smaller genera, largely by reviving names already proposed. Trachypenaeus was divided into three genera. Their proposals have not been universally accepted. Molecular evidence is ambivalent (Baldwin et al., 1998; Gusmao et al., 2000; Maggioni et al., 2001). Davie (2002) argued that the commercial importance of many species demanded that names be treated conservatively while uncertainty remains. This is what is done here – the generic names advocated by Pérez Farfante & Kensley are treated as subgenera. Nevertheless, few commercially important species are fished in southern Australia (see boxes: Australian Prawns and Southern Australian Prawn Fisheries). Except for one species in SA and WA, species of Penaeus are rarely found and then only as isolated individuals. Southern estuaries are home to species of Metapenaeus which are the basis of small commercial and amateur fisheries. The remainder are offshore, deep-water species encountered only by oceanographers. Pérez Farfante & Kensley (1997) recognised 26 genera of Penaeidae and provided a key. More conservative keys to the more common Australian species were published by Grey et al. (1983) who built on the much earlier review by Dall (1957). Because of their fishery interest in Australia and elsewhere, prawns are among the best understood of all Crustacea (Dall et al., 1990). The significance of penaeids to Australian fisheries was synthesised by Yearsley & Last (1999) who built on a large earlier literature.

Diagnosis. Prosartema (short branch on inner side of antennule, inside eyes) present. Flagella of antennules equal in length and attached together at tip of third article. Cervical groove never reaching more than two-thirds distance from hepatic spine to top of carapace. Pleopods with 2 rami.

Key to southern Australian species of Penaeidae 1. Rostrum with dorsal but without ventral teeth ...... 2 — Rostrum with dorsal and ventral teeth ...... Penaeus 17 2. Telson with 1 or more pairs of fixed subapical spines, with or without moveable spines . . 3 — Telson without fixed subapical spines (moveable spines if present) ...... 6 3. Telson with 1 pair of fixed subapical spines; mandibular incisor not elongate ...... 4 — Telson with 3 pairs of fixed subapical spines; mandibular incisor acutely elongate and transverse ...... Funchalia … 16 4. Maxilliped 3 and pereopod 2 without spine on inside of basis; petasma symmetrical . . . . . 5 — Maxilliped 3 and pereopod 2 with spine on inside of basis; petasma asymmetrical ...... Metapenaeopsis … 11

32 Dendrobranchiata – prawns and midwater shrimps

5. Carapace without longitudinal suture; telson with 2 or 3 small moveable spines ...... Penaeopsis eduardoi — Carapace with longitudinal suture (running back from near antennal spine); telson without moveable spines ...... Parapenaeus … 7 6. Pereopod 5 without exopod ...... Metapenaeus … 8 — Pereopod 5 with exopod ...... Trachypenaeus curvirostris 7. Rostrum as long as peduncle of antennule and scaphocerite . . . . . Parapenaeus australiensis — Rostrum little more half as long as peduncle of antennule and scaphocerite ...... Parapenaeus sextuberculatus 8. Telson with 4 pairs of conspicuous lateral moveable spines ...... Metapenaeus macleayi — Telson with numerous minute spines ...... 9 9. Pereopod 1 with well developed ischial spine (in addition to basal spines on pereopods 1–3); branchiocardiac ridge long ...... Metapenaeus ensis — Pereopod 1 without ischial spine (with basal spines on pereopods 1–3); branchiocardiac ridge long not prominent ...... 10 10. Abdominal somites with clearly developed areas of fine hairs; distal projections of petasma parallel and tubular; NSW and south-eastern Qld ...... Metapenaeus bennettae — Abdominal somites without areas of fine hairs; distal projections of petasma strongly divergent; WA and tropical Australia ...... Metapenaeus dalli 11. Posterior margin of carapace without stridulating organ ...... Metapenaeopsis velutina — Posterior margin of carapace with stridulating organ ...... 12 12. Abdominal somite 3 dorsal ridge without groove ...... 13 — Abdominal somite 3 dorsal ridge with groove ...... 14 13. Abdominal somite 3 dorsal ridge flat or weakly convex ...... Metapenaeopsis novaeguinae — Abdominal somite 3 dorsal ridge strongly hump-backed ...... Metapenaeopsis fusca 14. Stridulating organ raised on ridge above branchiostegite ...... Metapenaeopsis lindae — Stridulating organ not raised ...... 15 15. Fine stridulating organs on posterior carapace; abdominal somite 3 dorsal carina with narrow, deep well defined groove ...... Metapenaeopsis crassissima — Prominent stridulating organs on posterior carapace; abdominal somite 3 dorsal carina with wide, shallow groove ...... Metapenaeopsis palmensis 16. Rostrum with 4–6 dorsal teeth ...... Funchalia villosa — Rostrum with 10–13 dorsal teeth ...... Funchalia woodwardi 17. Adrostral ridge reaching most posterior of middorsal teeth; gastrofrontal ridge absent ...... Penaeus esculentus — Adrostral ridge almost reaching to posterior border of carapace; gastrofrontal ridge present 18 18. Gastrofrontal groove divided into 2 at posterior end; rostrum without accessory pair of ridges ...... Penaeus latisulcatus — Gastrofrontal groove divided into 3 at posterior end; rostrum with accessory pair of ridges ...... Penaeus plebejus

33 Marine Decapod Crustacea of Southern Australia

b d c

g h

i j k

Fig. 5. Penaeidae. a, Penaeus plebejus. Carapace: b, Funchalia villosa. c, Funchalia woodwardi. d, Metapenaeopsis crassissima. e, Metapenaeopsis lindae. f, Metapenaeopsis velutina. g, Parapenaeus australiensis. h, Parapenaeus sextuberculatus. i, Penaeopsis eduardoi. j, Penaeus latisulcatus. k, Trachypenaeus curvirostris.

Funchalia Johnson, 1867 Both Australian species of Funchalia are found offshore in southern Australian waters and are distributed through many of the world’s oceans besides. All five or six species of the genus are mesopelagic and are of no commercial interest. Australian species have been recorded from

34 Dendrobranchiata – prawns and midwater shrimps

large-scale eddies off the eastern coast of Australia (Griffiths & Brandt, 1983; Kensley et al., 1987) and in southern Australia (Hanamura, 1989). Diagnosis. Rostrum with dorsal teeth, usually without ventral teeth. Telson with 3 pairs of fixed subapical spines. Thoracic somite 8 (pereopod 5) with pleurobranch. Mandibular incisor elongate, scimitar-like, lying transversely behind labrum. Funchalia villosa (Bouvier, 1905) (Fig. 5b). Rostrum with 4–6 dorsal teeth, without ventral teeth; hepatic spine absent. Abdominal somites 4–6 carinate; lateral carinae on somites 5–6. cl. 19 mm. Atlantic, Indo-West Pacific, Antarctic, NSW; 50–666 m depth. Funchalia woodwardi Johnson, 1867 (Fig. 5c). Rostrum with 11–13 dorsal teeth, ventral margin unarmed; hepatic spine small, with marked ridge extending back to near posterior margin of carapace. Abdominal somites 4–6 carinate, except on posterior two-thirds of somite 4; lateral carinae on somites 4–6. cl. 38 mm. Atlantic, Indo-West Pacific, NSW, WA; 250–540 m depth.

Metapenaeopsis Bouvier, 1905 The Indo-West Pacific deep-water species were treated by Crosnier (1987, 1991) and shallow- water collections from Western Australia by Manning (1988) and Paulinose (1988). Some species are called velvet shrimps because of the texture of the exoskeleton. Diagnosis. Rostrum with dorsal teeth, without ventral teeth. Carapace without longitudinal or transverse sutures. Telson with 1 pair of fixed subapical spines followed by several pairs of large moveable spines. Thoracic somite 8 (pereopod 5) without pleurobranch. Maxilliped 3 with spine on inside of basis. Petasma asymmetrical. Fine stridulating organs (ridges) sometimes present on posterior carapace. Metapenaeopsis crassissima Racek & Dall, 1965 stout velvet shrimp (Fig. 5d). Fine stridulating organs present on posterior carapace. Dorsal carina of abdominal somite 3 with narrow, deep well-defined groove. Pterygostomial spine large and prominent. Rosy-pink with slightly darker mottling, without transverse bands on dorsal surface; pleopods and legs whitish. tl. 125 mm. WA, NT, SA (doubtful); to 30 m depth. The species is large but has shorter legs than others in the genus. Southern records may be M. lindae (Manning, 1988). Metapenaeopsis fusca Manning, 1988 (Fig. 6a). Stridulating organ narrow, slightly upcurved, not markedly raised above level of branchiostegite. Dorsal convex carina of abdominal somite 3 prominent, without groove. Pterygostomial spine short and sharp. Transparent with a choco- late-brown mottling. tl. 74 mm. Qld, (Torres Strait), and WA (Singleton to Cockburn Sound); on seagrass, 1–20 m depth (rare). Metapenaeopsis lindae Manning, 1988 (Fig. 6b, e). Stridulating organ raised on a ridge above the branchiostegite. Dorsal carina of abdominal somite 3 with well-defined groove at least half length of carina. Pterygostomial spine long, slender, sharp. Translucent white to light pink- brown with red-brown markings. tl. 62 mm. SA (E to Smoky Bay), WA (N to Rottnest I.); shallow water. Metapenaeopsis novaeguinae (Haswell, 1879). New Guinea prawn, northern velvet shrimp (Fig. 6c). Stridulating organ raised on a ridge above the branchiostegite. Dorsal carina of abdominal somite 3 flat on top. Pterygostomial spine long, slender, sharp. Mottled red-brown tl. 100 mm. East Indo-West Pacific, WA (S to Shark Bay), NT, Qld, NSW (S to Sydney), SA; 5–30 m depth. Metapenaeopsis palmensis (Haswell, 1879) southern velvet prawn. Prominent stridulating organs on posterior carapace. Dorsal carina of abdominal somite 3 with wide, shallow groove. Pterygostomial spine prominent. Pink to orange, lightly mottled with transverse bands on

35 Marine Decapod Crustacea of Southern Australia

dorsal surface; legs lightly banded. tl. 105 mm. Malaysia, Indonesia, New Guinea, WA (S to Shark Bay), NT, Qld, NSW (S to Sydney); 5–30 m depth. For information on biology see Watson & Keating (1989). Metapenaeopsis velutina (Dana, 1852). Stridulating organ absent. Dorsal carina of abdominal somite 3 high, narrow, without groove. Pterygostomial spine prominent. Translucent, marbled with white and orange-red. tl. 81 mm. Indo-West Pacific, WA,Qld, NSW (S to Broken Bay); 55–320 m depth. The species was redescribed by Crosnier (1991).

b c a

h d g e f

Fig. 6. Penaeidae. Abdominal somites 1–3. a, Metapenaeopsis fusca. b, Metapenaeopsis lindae. c, Metapenaeopsis novaeguinae. d, Metapenaeus bennettae (abdomen). e, Metapenaeopsis lindae (stridulating organ on posterior margin of carapace). Telson: f, Metapenaeus macleayi. g, Penaeopsis eduardoi. h, Penaeus plebejus (dorsal rostrum).

Metapenaeus Wood-Mason, 1891 Species of Metapenaeus are the only common prawns in estuaries in southern Australia, two endemic to south-eastern Australia and the others tropical but found in temperate Australia. They feature in commercial and amateur fisheries while larger representatives, the Endeavour prawns, contribute to Australia’s Northern Prawn Fishery. Miquel (1982) reviewed taxonomy, biology and fisheries worldwide for all 26 species. All species live in the Indo-West Pacific. He provided a key to distinguish them. Diagnosis. Rostrum with dorsal teeth, without ventral teeth. Telson without fixed subapical spines, with moveable spines. Pereopod 5 without exopod. Metapenaeus bennettae Racek & Dall, 1965 bay prawn, greentail prawn, greasyback prawn, river prawn (Fig. 6d). Telson with numerous minute spines. Pereopod 1 without obvious ischial spine. Branchiocardiac ridge not prominent. All abdominal somites with clearly developed areas of fine hairs. Distal projections of petasma parallel and tubular. Translucent body speckled with dark brown; tips of uropods, pleopods and antennal scales green; antennae red to brown. tl. 110 mm. Qld (N to Hervey Bay), NSW,Vic. (eastern estuaries only); estuarine, 0–30 m depth. Adult prawns are found in coastal rivers up to 15 km from the mouth, tolerating salinities between fully fresh and fully marine waters (Grey et al., 1983). The bay prawn is an important species in the amateur and commercial fisheries of NSW and Qld. Metapenaeus dalli Racek, 1957 western school prawn.As M. bennettae except abdominal somites without areas of fine hairs, distal projections of petasma strongly divergent. Semi-translucent,

36 Dendrobranchiata – prawns and midwater shrimps

pale green to brown with numerous dark brownish pigment spots; tips of uropods green to reddish. tl. 85 mm. Indonesia, Philippines, NT,WA (S to Mandurah); estuarine, 9–33 m depth. The Western school prawn is abundant in WA and forms the basis of an amateur fishery in the Swan River and other estuaries. Metapenaeus ensis (De Haan, 1844) offshore greasyback prawn. Telson with numerous minute spines. Pereopod 1 with long sharp ischial spine. Distinct long branchiocardiac ridge present. Pale brown to bright pink; bright red antennae; pereopods and uropods pink to red. tl. 160 mm. Indo-West Pacific, WA (S to Shark Bay), NT, Qld, NSW (S to 35°S); to 64 m depth. The species is of major commercial importance in Singapore, Malaysia, Indonesia, Japan and in northern Australia where it is caught with M. endeavouri. Metapenaeus macleayi (Haswell, 1879) eastern school prawn (Fig. 6f). Telson with 4 pairs of conspicuous lateral moveable spines. Pereopod 1 without obvious ischial spine. Abdominal somites without areas of fine hairs. Translucent with tinges of brown to green in irregular spots; antennae brown; tips of uropods blue in adults. tl. 160 mm. Qld (N to Moreton Bay), NSW,Vic. (W to Western Port); estuarine, to 60 m depth (endemic). The eastern school prawn is an important species exploited for the domestic market by commercial and amateur fisheries, especially in NSW where it is abundant.

Parapenaeus Smith, 1885 Species of Parapenaeus, which number ten, are deep-water prawns (Crosnier, 1985). Two species have been recorded in southern Australia. Another rare species is found elsewhere in Australia. In addition to the length of the rostrum which differentiates these species are important differences in the petasma. Diagnosis. Carapace with distinct longitudinal sutures; stridulating organs absent from posterior carapace. Rostrum with dorsal teeth, without ventral teeth. Telson with 1 pair of fixed subapical spines. Thoracic somite 8 (pereopod 5) without pleurobranch. Maxilliped 3 without spine on inside of basis. Petasma symmetrical. Parapenaeus australiensis Dall, 1957 red prawn. Rostrum reaching end of third article of antennule, as long as scaphocerite. Branchiostegal spine present, on anterior margin of carapace. Adrostral ridge ends in small oval depression. Thelycum without median tubercle between lateral flaps. tl. 160 mm. Western Pacific, Qld (N to Heron I.), NSW (S to Cape Howe); 100–250 m depth. Parapenaeus sextuberculatus Kubo, 1949. Rostrum reaching middle of second article of antennule, shorter than scaphocerite. Branchiostegal spine present, on anterior margin of carapace. Thelycum with median tubercle between lateral lobes. Indo-West Pacific, NSW; 140–350 m depth.

Penaeopsis Bate, 1881 A single species is found in southern Australia restricted to the east coast. Pérez Farfante (1980) rediagnosed the genus in which she recognised six species. Pérez Farfante (1978) found intersexes in one. The shrimps are benthic on the upper part of continental slopes, especially in the tropics of the Indo-West Pacific. Some species are sufficiently abundant to be commercially exploited. Diagnosis. Carapace without longitudinal or transverse sutures; orbital and branchiostegal spines absent; antennal, hepatic and pterygostomial spines well developed. Rostrum without ventral teeth. Telson with 1 pair of fixed subapical spines, 2 or 3 small moveable spines. Maxilliped 3 without spine on inside of basis. Petasma symmetrical, simple.

37 Marine Decapod Crustacea of Southern Australia

Penaeopsis eduardoi Pérez Farfante, 1977 red-flecked prawn (Fig. 6g). Telson with 2 pairs of moveable spines. Rostrum arched, longer than antennule peduncle. Thelycum with ventral costa produced as long spine. tl. 130 mm. East Indo-West Pacific, WA,NT, NSW; 250–570 m, mainly 250–400 m depth.

Penaeus Fabricius, 1798 The principal target species for all Australian commercial prawn fisheries belong to the genus Penaeus sensu lato. Members of the genus also form the most important of the world’s prawn fisheries but only one species is common enough in a few bays to be fished in southern Australia. Two subgenera, or genera according to Pérez Farfante & Kensley (1997), Melicertus and Penaeus,are represented in southern Australia. Diagnosis. Rostrum with dorsal and ventral teeth. Telson without spines or with 3 pairs of moveable spines. Mandibular incisor short, almost right-angled. Penaeus (Melicertus) latisulcatus Kishinouye, 1896 western king prawn (Pl. 7a). Adrostral ridge almost reaching to posterior border of carapace; postrostral groove present; gastrofrontal ridge present; gastrofrontal groove divided into 2 at posterior end. Rostrum without accessory pair of ridges. Body without cross-bands on carapace or abdomen, light yellow to brown; rostrum dark brown; legs light blue; pleopods yellow. tl. 200 mm. Indo-West Pacific, WA (S to Cape Naturaliste), NT, Qld, NSW, SA (Gulfs region); depths of 0–90 m. The only fisheries for large prawns in southern states rely on this species, in the Gulfs of SA and in WA. Penaeus (Melicertus) plebejus Hess, 1865 eastern king prawn (Fig. 6h, Pl. 7b). Adrostral ridge almost reaching to posterior border of carapace; postrostral groove present; gastrofrontal ridge present; gastrofrontal groove divided into 3 at posterior end. Rostrum with accessory pair of ridges. Body cream to yellow, uropods blue with red fringe. tl. 300 mm. Qld (N to 27°S), NSW, Vic. (W to eastern estuaries and Port Phillip Bay as isolated individuals); 2–220 m depth (Australian endemic). This species forms the basis of important fisheries in NSW and Qld. Penaeus (Penaeus) esculentus Haswell, 1879 brown tiger prawn. Adrostral ridge reaching most posterior of middorsal teeth; gastrofrontal ridge absent; hepatic ridge not reaching behind antennal ridge. Rostrum without accessory pair of ridges. Brown overall, carapace and abdomen with transverse stripes, 2 per segment; bases of pleopods yellow and blue. tl. 235 mm. WA (S to Shark Bay), NT, Qld, NSW (S to Port Jackson); estuaries to 200 m depth (Australian endemic). The species is fished commercially throughout its range with other species.

Trachypenaeus Alcock, 1901 A single species of this genus of thick-set pubescent prawns occurs in southern Australia. Others from tropical waters were figured by Grey et al. (1983). Trachysalambria is one of at least three genera into which Trachypenaeus was divided by Pérez Farfante & Kensley (1997). Diagnosis. Rostrum with dorsal teeth, without ventral teeth. Carapace suture short, weak. Telson without fixed subapical spines, with moveable lateral spines. Maxilliped 3 without spine on inside of basis. Pereopod 5 with exopod. Thoracic somites 4 and 5 (pereopods 1 and 2) without pleurobranch. Trachypenaeus (Trachysalambria) curvirostris (Stimpson, 1860) hardback prawn, southern rough prawn. Hepatic groove present. Telson with 3 or 4 pairs of lateral spines. Petasma with pair of large laterally-directed wing-like extensions. Pink to reddish-brown; legs whitish; pleopods and uropods red. tl. 100 mm. Indo-West Pacific, WA (S to Shark Bay), NT, Qld, NSW (S to Sydney); 13–150 m depth (rare).

38 Dendrobranchiata – prawns and midwater shrimps

References Baldwin, J.D., Bass, A.L., Bowen, B.W., & Clark, W.H. 1998. Molecular phylogeny and biogeography of the marine shrimp, Penaeus. Molecular Phylogenetics and Evolution 10: 399–407. Crosnier, A. 1985. Crustacés Décapodes: Penaeidae. Les espèces indo-ouest-pacifique du genre Parapeneaeus. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM I et II – Philippines (1976, 1980). Tome 2. Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. A (Zoologie) 133: 303–355. Crosnier, A. 1987. Les espèces indo-ouest-pacifiques d’eau profonde du genre Metapenaeopsis (Crustacea, Decapoda Penaeidae). Bulletin du Muséum National d’Histoire Naturelle, Paris 9: 409–453. Crosnier, A. 1991. Crustacea Decapoda: Les Metapenaeopsis indo-ouest-pacifiques sans appareil stridu- lent (Penaeidae). Deuxième partie. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM. Vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. A (Zoologie) 152: 155–297. Dall, W. 1957. A revision of the Australian species of Penaeinae (Crustacea Decapoda: Penaeidae). Australian Journal of Marine and Freshwater Research 8: 136–230. Dall, W., Hall, B.J., Rothlisberg, P.,& Staples, D.J. 1990. The biology of the Penaeidae. Advances in Marine Biology 27: 1–489. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Grey, D.L., Dall, W., & Baker, A. 1983. A Guide to the Australian Penaeid Prawns. Northern Territory Government Printing Office: Darwin. 140 pp. Griffiths, F.B., & Brandt, S.B. 1983. Distribution of mesopelagic decapod Crustacea in and around a warm-core eddy in the Tasman Sea. Marine Ecology Progress Series 12: 175–184. Gusmao, J., Laoski, C., & Sole-Cava, A.M. 2000. A new species of Penaeus (Crustacea: Penaeidae) revealed by allozyme and cytochrome oxidase I analysis. Marine Biology 137: 435–446. Hanamura, Y. 1989. Deep-sea shrimps (Crustacea: Decapoda) collected by the R.V.‘Soela’ from southern Australia. Bulletin of the Biogeographical Society of Japan 44: 51–69. Holthuis, L.B. 1980. FAO species catalogue. Vol.1. Shrimps and prawns of the world. An annotated catalogue of species of interest to fisheries. FAO Fisheries Synopsis 1 (125): 1–261. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Maggioni, R., Rogers, A.D., Maclean, N., & D’Incao, F. 2001. Molecular phylogeny of Western Atlantic Farfantepenaeus and Litopenaeus shrimp based on mitochondrial 16S partial sequences. Molecular Phylogenetics and Evolution 18: 66–73. Manning, R. 1988. Two new species of Metapenaeopsis (Crustacea: Decapoda: Penaeidae) from south Western Australia. Records of the Western Australian Museum 14: 91–103. Miquel, J.C. 1982. Le genre Metapenaeus (Crustacea, Penaeidae): Taxonomie, biologie et pêches mondi- ales. Zoologische Verhandelingen, Leiden 195: 1–137. Paulinose, V.T. 1988. Decapod Crustacea from the International Indian Ocean Expedition: larval and postlarval stages of three species of Metapenaeopsis Bouvier (Penaeidae: Penaeinae). Journal of Natural History 22: 1565–1577. Pérez Farfante, I. 1978. Intersex anomalies in shrimp of the genus Penaeopsis (Crustacea: Penaeidae). Fishery Bulletin 76: 687–691. Pérez Farfante, I. 1980. Revision of the penaeid shrimp genus Penaeopsis (Crustacea: Decapoda). Fishery Bulletin of the Fish and Wildlife Service, United States 77: 721–763. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233. Watson, R.A., & Keating, J.A. 1989. Velvet shrimps (Metapenaeopsis spp.) of Torres Strait, Queensland, Australia. Asian Fisheries Science 3: 45–56.

39 Marine Decapod Crustacea of Southern Australia

Yearsley, G.K., & Last, P.R. 1999. Crustaceans. Pp. 295–326 in: Yearsley, G.K., Last, P.R., & Ward, R.D. (eds), Australian Seafood Handbook. An Identification Guide to Domestic Species. CSIRO Marine Research: Hobart.

AUSTRALIAN PRAWNS The large prawns sold commercially throughout Australia and served in restaurants come mostly from tropical Queensland, Northern Territory and Western Australia. This key will separate only the dominant species. Commercially caught prawns are snap-frozen immediately they come on deck. Colour refers to live animals and is not necessarily retained on frozen individuals and certainly not on cooked ones! Maximum lengths are of females; males are smaller. 1. Rostrum with teeth underneath ...... Penaeus … 2 — Rostrum with teeth only on top ...... Metapenaeus … 8 2. Ridges on each side of rostrum reaching almost to posterior end of carapace ...... 3 — Ridges on each side of rostrum reaching to middle of carapace or shorter ...... 5 3. Red spot on side of abdomen; ridges on each side of rostrum not reaching to middle of carapace; first legs with spine on ischium ...... Penaeus longistylus (red spot king prawn, 180 mm) — No red spot on side of abdomen; first legs without spine on ischium ...... 4 4. Rostrum with intermediate pair of ridges between crest and major lateral ridges; end of groove running back from eye divided into 3; body yellowish with red tinges ...... Penaeus plebejus (eastern king prawn, 300 mm) — Rostrum without intermediate pair of ridges; end of groove running back from eye divided into 2; body yellow-brown, legs light blue ...... Penaeus latisulcatus (western king prawn, 200 mm) 5. With hepatic ridge, a short curved horizontal ridge behind lower spine at front of carapace; brownish with transverse darker bands at least on abdomen ...... 6 — Without hepatic ridge; body yellowish ...... 7 6. Ridges on side of rostrum extending past the most posterior dorsal tooth; dark green to brown with darker bands on abdomen ...... Penaeus semisulcatus (grooved tiger prawn, 250 mm) — Ridges on side of rostrum stopping at the most posterior dorsal tooth; dark red-brown with darker bands on carapace and 2 per segment on abdomen ...... Penaeus esculens (brown tiger prawn, 235 mm) 7. With gastro-orbital ridge, running obliquely down from behind notch at front of carapace; legs reddish-pink . . . . . Penaeus indicus (red-legged banana prawn, 230 mm) — Without gastro-orbital ridge; speckled with brown dots ...... Penaeus merguiensis (white banana prawn, 240 mm) 8. Telson with 3 pairs of moveable spines on sides; pale brown to pink, legs red, uropods and telson with bright blue tips ...... Metapenaeus endeavouri (blue endeavour prawn, 190 mm) — Telson with numerous, almost invisible minute spines on sides; pale brown to pink, legs, uropods and telson red . . . . . Metapenaeus ensis (red endeavour prawn, 160 mm)

40 Dendrobranchiata – prawns and midwater shrimps

SOUTHERN AUSTRALIAN PRAWN FISHERIES Three species of greasy-back prawns are commonly caught by amateurs in estuaries in NSW, eastern Victoria and Western Australia. Locality is a good guide to identification of the common species. Maximum lengths are of females; males are smaller. 1. WA, tropical Australia; telson without obvious lateral spines ...... Metapenaeus dalli (western school prawn, 85 mm) — NSW to southern Qld; telson with or without obvious lateral spines ...... 2 2. Telson with 4 pairs of moveable lateral spines; body with few hairs ...... Metapenaeus macleayi (New South Wales school prawn, 160 mm) — Telson without obvious lateral spines; sides of abdomen with areas of hairs ...... Metapenaeus bennettae (bay prawn, greentail prawn, 110 mm)

Sicyoniidae Ortmann, 1898 The rigid carapace is the most obvious feature distinguishing sicyoniids from all other dendrobranchiates. The legs are usually shorter and stronger than in other prawns. Besides this, they lack endopods on pleopods 3–5. Diagnosis. Rostrum with dorsal teeth, without ventral teeth. Carapace rigid, pubescent, without cervical groove, without postorbital pterygostomial spines, with hepatic spine and hepatic carina. Abdominal somites grooved or sculptured. Antennule with rudimentary prosartema and with short flagella. Exopod on maxilliped 1 only.

Sicyonia H. Milne Edwards, 1830 Sicyonia is the only genus but Crosnier (2003) recognised eight groups of species among 31 that he dealt with from the Indo-West Pacific. His work contains a key to species and colour photographs. Species number more than 40 worldwide (Pérez Farfante & Kensley, 1997). The genus is largely confined to the tropics.

Key to southern Australian species of Sicyonia 1. Carapace with 2 middorsal spines ...... Sicyonia australiensis — Carapace with 3 middorsal spines ...... Sicyonia metavitulus Diagnosis. As for family. Sicyonia australiensis Hanamura & Wadley, 1998 (Fig. 7a). Carapace with 2 middorsal spines. Rostrum reaching end of eyestalk, with straight lower margin, slightly tapering. Abdominal somites 1 and 2 each with sharp anterior dorsal spine, somite 5 with postventral angle. cl. 9 mm. Vic., Tas. (eastern Bass Strait); 123–230 m depth. Sicyonia metavitulans Crosnier, 2003 (Fig. 7b). Carapace with 3 middorsal spines. Rostrum reaching well beyond end of eyestalk, with straight lower margin, slightly tapering. Abdominal somite 1 only with sharp anterior dorsal spine, somite 5 with postventral tooth. cl. 14 mm. New Caledonia, NSW (Port Jackson); to 57 m depth.

41 Marine Decapod Crustacea of Southern Australia

References Crosnier, A. 2003. Sicyonia (Crustacea, Decapoda, Penaeoidea, Sicyoniidae) de l’Indo-ouest Pacifique. Zoosystema 25: 197–350. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233.

ac f

Fig. 7. Sicyoniidae. a, Sicyonia australiensis. b, Sicyonia metavitulans. Solenoceridae. c, Haliporoides cristatus. d, e, Haliporoides sibogae. f, Hymenopenaeus halli. g, Hymenopenaeus propinquus. h, Solenocera faxoni. Luciferidae. i, Lucifer hanseni (male). j, Lucifer typus (abdominal somite 6 of male).

42 Dendrobranchiata – prawns and midwater shrimps

Solenoceridae Wood-Mason, 1891 The solenocerid shrimps are rarely found over the continental shelf but prefer waters over the slope, at depths of hundreds of metres. Some are mesopelagic or bathypelagic while other are benthic. Members of the family are distinguished from other dendrobranchiates by the possession of a postorbital spine and usually a complete cervical groove (Pérez Farfante & Kensley, 1997). Dall’s (1999) review of the Australian species diagnosed the family, discussed anatomical features, and provided keys to all nine genera. He also reviewed the literature and gave keys to Indo-West Pacific species. Twenty-seven species were recorded by Dall from Australia but only five reach southern waters. Other genera and other species have been recorded as far south as northern NSW. Diagnosis. Rostrum with more than 3 dorsal teeth. Carapace with well developed cervical groove, usually reaching mid-dorsum, with postorbital and hepatic spines. Abdomen at least partially carinate. Antennule with prosartema (short branch on inner side of antennule, inside eyes) and with long flagella, often longer than carapace. Exopods on maxillipeds 1–3 and pereopods 1–4 (sometimes on 5). Petasma simple, tubular. Appendix masculina on pleopod 2 with spur on outer side of basal segment.

Key to southern Australian species of Solenoceridae 1. Dorsal and ventral antennular flagella flattened, forming respiratory tube; outer ramus of uropod without distolateral spine ...... Solenocera faxoni — Dorsal and ventral antennular flagellum subcylindrical; outer ramus of uropod with distolateral spine ...... 2 2. Postcervical (suprahepatic) spine present; 1 middorsal tooth in front of cervical groove (epigastric) separated from teeth on rostrum ...... Haliporoides … 3 — Postcervical (suprahepatic) spine absent; 2 middorsal teeth in front of cervical groove (first rostral and epigastric) separated from teeth on rostrum ...... Hymenopenaeus … 4 3 Rostrum high, arched behind first rostral tooth, up-turned, with 7–11 (usually 10) dorsal teeth plus epigastric, 3 ventral teeth ...... Haliporoides cristatus — Rostrum slender, almost straight, with 6–8 dorsal teeth plus epigastric, 1–2 ventral teeth ...... Haliporoides sibogae 4. Pereopod 1 basis and merus each with spine on lower margin; scaphocerite as long as peduncle ...... Hymenopenaeus halli — Pereopod 1 basis and merus without spine on lower margin; scaphocerite longer than peduncle ...... Hymenopenaeus propinquus

Haliporoides Stebbing, 1914 The suprahepatic spine that which defines the genus is minute and just posterior to the cervical groove above the hepatic spine. Although there are only four species, three additional subspecies have been described. Diagnosis. Dorsal and ventral antennular flagellum subcylindrical. Outer ramus of uropod with distolateral spine. Epigastric tooth separated from rostral teeth by long interval. Suprahepatic spine present. Haliporoides cristatus Kensley, Tranter & Griffin, 1987 (Fig. 7c). Rostrum high, arched behind first rostral tooth, up-turned, with 7–11 (usually 10) dorsal teeth plus epigastric, 3 ventral teeth. Yellowish with white stripe on upper surface of uropods. cl. 34 mm. Qld, NSW, to 35.5°S; 250–629 m depth.

43 Marine Decapod Crustacea of Southern Australia

Haliporoides sibogae de Man, 1907 (Fig. 7d, e). Rostrum slender, almost straight, with 6–8 dorsal teeth plus epigastric, 1–2 ventral teeth. Uniformly pink to red. cl. 50 mm. Indo-West Pacific, WA, NT, Qld, NSW, Vic., Tas.; 100–900 m, but all Australian records from deeper than 350 m depth. This species is more common than H. cristata and is fished commercially off eastern Australia (Davie, 2002). Dall (1999) was unconvinced of the taxonomic differences of Kensley et al.’s (1987) subspecies, H. sibogae australiensis.

Hymenopenaeus Smith, 1882 Hymenopenaeus is a genus of about a dozen small deepwater shrimps. Six occur in Australia. Diagnosis. Dorsal and ventral antennular flagellum cylindrical. Outer ramus of uropod with distolateral spine. Epigastric and first rostral teeth separated from remaining ones by long interval. Suprahepatic spine absent. Hymenopenaeus halli Bruce, 1966 (Fig. 7f). Pereopod 1 basis and merus each with spine on lower margin; scaphocerite as long as peduncle. White-yellowish. cl. 23 mm. Indo-West Pacific, Qld, NSW, to 34°S; 450–1000 m depth. Hymenopenaeus propinquus (de Man, 1907) (Fig. 7g). Pereopod 1 basis and merus without spine on lower margin; scaphocerite longer than peduncle. White-yellowish. cl. 27 mm. Indo-West Pacific, WA, NT, Qld, NSW, to 34°S; 450–1200 m depth.

Solenocera Lucas, 1849 Species of Solenocera are probably benthic on muddy sediments of the continental shelf and slope. The respiratory tube fashioned from the modified antennules suggest the shrimps bury themselves in the mud with the abdomen flexed upwards. Solenocera is the largest genus in the family, with more than 30 species of which 20 or so live in the Indo-West Pacific. Only one reaches southern Australia. Diagnosis. Dorsal and ventral antennular flagella flattened. Lateral ramus of uropod lacking distolateral spine. Solenocera faxoni de Man, 1907 (Fig. 7h). Pterygostomial spine absent. Rostrum reaching to middle of cornea. Postrostral carina not extending posterior to cervical groove. Hepatic carina ending in sharp elevated spine. Abdominal somite 3 without carina. cl. 27 mm. Indonesia to Japan, WA, NT, Qld, NSW; 200–400 m depth, uncommon.

References Dall, W. 1999. Australian species of Solenoceridae (Penaeoidea: Decapoda). Memoirs of the Queensland Museum 43: 553–587. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233.

44 Dendrobranchiata – prawns and midwater shrimps

Superfamily Sergestoidea Dana, 1852 There are two families: Luciferidae and Sergestidae.

Luciferidae De Haan, 1849 These tiny prawns appear in plankton and benthic samples from soft sediments, sometimes in large numbers. Their elongate shape, long neck, and dominant eyes make them immediately recognisable. Lucifer is the only genus, with only seven species worldwide. Three have been recorded from Australia, L. penicillifer in tropical regions in addition to the two below. Because they lack so many basic dendrobranchiate features, like gills, their relationships to other families and even their inclusion in Dendrobranchiata itself have been questioned (Burkenroad, 1983; Pérez Farfante & Kensley, 1997). The presence of a petasma in males is the only feature tying luciferids to dendrobranchiates but euphausiaceans possess this feature too. Loss of pereopods 4 and 5 is shared with some sergestids but is a dubious synapomorphy. Diagnosis. Carapace extremely laterally compressed, elongated anterior to pereopods and mandibles. Chelae on pereopods lacking or imperfect. Pereopods 4 and 5 absent, gills absent. Abdominal somite 6 of male with 2 ventral processes. Telson of male with strong ventral process.

Lucifer Thompson, 1829 Some authors have spelled the genus name, Leucifer, but the confusion was sorted out by Bowman & Holthuis (1968). Hansen (1919) and Petit (1973) illustrated and compared the species. Diagnosis. With the characters of the family.

Key to southern Australian species of Lucifer 1. Male abdominal somite 6 with blunt first ventral spine and larger hooked second ...... Lucifer typus — Male abdominal somite 6 with sharp first ventral spine and larger sharp second ...... Lucifer hanseni Lucifer hanseni Nobili, 1905 (Fig. 7i). First ventral process on abdominal somite 6 short, thick, closer to second larger triangular spine than to anterior margin of somite. 9 mm. Indo-West Pacific, NSW, Vic., Tas., SA; marine bays and shelf. Dakin & Colefax (1940) discussed the identity of specimens from NSW and noted some differences from typical specimens from the Red Sea. Hale’s (1927) record of L. affinis is probably a misidentification. Lucifer typus Milne Edwards, 1837 (Fig. 7j). First ventral process on abdominal somite 6 short, blunt, at one-third along somite; second process larger and hooked. Atlantic, Indo-Pacific, Qld, NSW; more offshore than previous species (Dakin & Colefax, 1940).

References Bowman, T.E., & Holthuis, L.B. 1968. Lucifer or Leucifer: which spelling is correct? Crustaceana 14: 216–217. Burkenroad, M.D. 1983. Natural classification of Dendrobranchiata, with a key to Recent genera. In: Schram, F.R. (ed.), Crustacean phylogeny. Crustacean Issues 6: 279–290. Dakin, W.J., & Colefax, A.N. 1940. The plankton of the Australian coastal waters off New South Wales Part 1 with special reference to the seasonal distribution, the phyto-plankton, and the planktonic

45 Marine Decapod Crustacea of Southern Australia

Crustacea, and in particular, the Copepoda and crustacean larvae, together with an account of the more frequent members of the groups Mysidacea, Euphausiacea, Amphipoda, Mollusca, Tunicata, Chaetognathia, and some reference to the fish eggs and fish larvae. Publications of the University of Sydney, Department of Zoology, Monograph 1: 1–215. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Hansen, H.J. 1919. The Sergestidae of the Siboga-Expedition. Siboga Expéditie Monographie 38: 1–65, pls 1–7. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233. Petit, D. 1973. Données sur la morphologie et la croissance chez le genre Lucifer (Décapodes Sergestidae): L. intermedius, L. penicillifer, L. hanseni, L. chacei, L. faxonii. Cahiers O.R.S.T.O.M., Série Océanographie 11: 207–227.

Sergestidae Dana, 1852 The combination of reduced last pereopods and a short rostrum serve to characterise sergestid shrimps. Sergestiids differ from other dendrobranchiates in having swimming legs fringed with long setae, although the number of legs may be fewer than the expected five pairs. Most are small species from offshore environments but at least one is common in estuaries on the Australian east coast. Species of the genera Sergia and Sergestes have the ability to produce light using either dermal photophores or the organs of Pesta. Dermal photophores are distinct phosphorescent organs, sometimes with external lenses that are associated with an internal pigment screen. Occurring on various parts of the body such as the carapace, abdomen and scaphocerite, photophores appear in preserved specimens as small or minute spots. Photophores bearing cuticular lenses appear to have a convex bulge when viewed side-on. Organs of Pesta are internal light-producing organs named after their discoverer. They are pigmented and modified areas of the gastrohepatic gland appearing as small, yellow, pea-like structures that can be seen through the carapace. Pérez Farfante & Kensley (1997) recognised six genera of which four can be found in southern Australia. In his detailed revision of Sergia, Vereshchaka (2000) tabulated differences between the genera, including Lucifer which has often been included.

Diagnosis. Carapace with thin integument, rostrum shorter than eyestalks, without antennal, branchiostegal and pterygostomial spines. Antenna with ventral flagellum modified as clasping organ in male. Pereopods 4 and 5 reduced or absent. Gills few (up to 8) or absent.

Key to southern Australian genera of Sergestidae 1. Pereopods 4 and 5 absent; bays and estuaries ...... Acetes — Pereopod 4 present; pereopod 5 present or absent; oceanic ...... 2 2. Gills absent or rudimentary above pereopod 4 ...... Petalidium — Gills present above pereopod 4 ...... 3 3. Dermal photophores absent; organs of Pesta present (luminous modifications of gastro-hepatic gland); pereopods 1 and 2 with ischium having well developed subterminal spine ...... Sergestes — Dermal photophores present; organs of Pesta absent; pereopods 1 and 2 with ischium lacking subterminal spine ...... Sergia

46 Dendrobranchiata – prawns and midwater shrimps

Acetes Milne Edwards, 1830 Many of the 14 species known live in estuaries or nearshore and are called paste shrimps by FAO. The complete absence of the last two pairs of legs distinguish them from all other shrimps and prawns. The genus is divided into groups, each differently distributed around the world (Chen & Chen, 2000). The Australian species is distinguished from other dendrobranchiates by the three dorsal teeth at the front of the carapace, scarcely produced into a rostrum. Diagnosis. Pereopods 1–3 minutely chelate, pereopods 4 and 5 absent. Arthrobranchs present on maxilliped 2 (rudimentary) to pereopod 4. Acetes sibogae Hansen, 1919 Australian paste shrimp (Fig. 9a). cl. 6 mm. Indo-West Pacific, WA (S to Swan R.), NT, Qld, NSW; estuaries and shallow marine bays and shelf. In summer the shrimp swarms near the surface in the Swan River. Acetes australis Colefax, 1940, described from Sydney, is a synonym of this widespread species (Hanamura, 1999). Intersexes are known (Hanamura & Ohtsuka, 2003).

Petalidium Bate, 1881 Three species of these mesopelagic or bathypelagic shrimps have been recorded, a cold-water one from southern Australia. Diagnosis. Rostrum without or with 1 small tooth. Pereopods 4 and 5 reduced, 5 more so. Arthrobranchs present on maxilliped 2–pereopod 3, reduced above pereopod 4. Petalidium foliaceum Bate, 1881 (Fig. 9b). cl. 18 mm. Southern circumpolar, Tas., Australian Antarctic Territory; bathypelagic, mesopelagic, oceanic (Hale, 1941).

Sergestes H. Milne Edwards, 1830 Preserved specimens of Sergestes and Sergia are difficult to place to genus because of the reliance on luminescent organs that are not obvious to the novice. Both have a short triangular rostrum. The organ of Pesta is a modification of the gastrohepatic gland inside the carapace. In Sergestes, red chromatophore are mostly near the front whereas in Sergia they are more evenly distributed. The absence of a spine on the ischium of the first legs is also helpful. There are numerous species worldwide and the works of Kensley (1972), Wasmer (1993) and Webber et al. (1990) are helpful for the Australian region. Griffiths & Brandt (1983) recorded several species of Sergestes associated with a warm-core eddy off eastern Australia. All are pelagic over and beyond the continental slope, rarely over the shelf. Diagnosis. Carapace with red chromatophores mainly concentrated in anterior part of body; rostrum a short triangular projection. Dermal photophores absent. Organs of Pesta present. Pereopods 1 and 2 with ischium having well developed subterminal spine; pereopods 4 and 5 without dactyli.

Key to southern Australian species of Sergestes 1. Maxilliped 3 as long as pereopod 3 ...... 2 — Maxilliped 3 longer than pereopod 3 ...... 5 2. Last 2 articles of pereopod 5 setose on both margins ...... 3 — Last 2 articles of pereopod 5 setose on 1 margin ...... 4 3. Anterodorsal surface of the body with orange stellate chromatophores . . . Sergestes seminudus — Anterodorsal surface of the body without orange stellate chromatophores . . Sergestes disjunctus

47 Marine Decapod Crustacea of Southern Australia

4. Antennular peduncle article 3 equal to or longer than 1; petasma lobes short, stumpy ...... Sergestes atlanticus — Antennular peduncle article 3 shorter than 1; petasma lobes elongate . . . . Sergestes arcticus 5. Last 2 articles of pereopod 5 setose on both margins ...... 6 — Last 2 articles of pereopod 5 setose on only 1 margin ...... 7 6. Dactyl and distal half of propodus of maxilliped 3 with numerous spines forming comb-like structure ...... Sergestes pectinatus — Dactyl and distal half of propodus of maxilliped 3 with spines not forming comb-like structure ...... Sergestes sargassi 7. Dactylus of maxilliped 3 of 4 articles ...... 8 — Dactylus of maxilliped 3 of 6 articles ...... Sergestes orientalis 8. Rostrum long, with an elongate tip ...... Sergestes armatus — Rostrum short and deep, with very short tip ...... Sergestes stimulator Sergestes arcticus Krøyer, 1855. Maxilliped 3 as long as pereopod 3. Last 2 articles of pereopod 5 setose on 1 margin. Petasma lobes elongate. Atlantic, Indo-Pacific, Mediterranean, Southern Ocean, NSW, SA, TAS, WA; bathypelagic, mesopelagic. Sergestes armatus Krøyer, 1855 (Fig. 8a). Maxilliped 3 longer than pereopod 3. Last 2 articles of pereopod 5 setose on 1 margin. Dactylus of maxilliped 3 of 4 articles. Rostrum long, with an elongate tip. Atlantic, Indo-Pacific, Mediterranean, NSW, Vic., Tas., WA; bathypelagic, mesopelagic; 280–2046 m depth during day, 130–172 m depth at night. Sergestes atlanticus Milne Edwards, 1830. Maxilliped 3 as long as pereopod 3. Last 2 articles of pereopod 5 setose on 1 margin. Petasma lobes short, stumpy. Atlantic, Indo-West Pacific, NSW, Tas.; bathypelagic, mesopelagic. Sergestes disjunctus Burkenroad, 1940. Maxilliped 3 as long as pereopod 3. Last 2 articles of pereopod 5 setose on both margins. Supraorbital spine present. All lobes, including ‘lobus armatus’ of petasma straight. cl. 16 mm. Atlantic, Indo-West Pacific, NSW, Vic., Tas., SA, WA; mesopelagic. Sergestes orientalis Hansen, 1919. Maxilliped 3 longer than pereopod 3. Last 2 articles of pereopod 5 setose on 1 margin. Dactylus of maxilliped 3 of 6 segments. Indo-West Pacific, NSW; 50–450 m depth. Sergestes pectinatus Sund, 1920. Maxilliped 3 longer than pereopod 3. Last 2 articles of pereopod 5 setose on both margins. Dactyl and distal half of propodus of maxilliped 3 with comb-like structure. Atlantic, Indo-West Pacific, NSW, WA; mesopelagic. Sergestes sargassi Ortmann, 1893. Maxilliped 3 longer than pereopod 3. Last 2 articles of pereopod 5 setose on both margins. Dactyl and distal half of propodus of maxilliped 3 armed with spines not forming comb-like structure. Atlantic, Indo-Pacific, Mediterranean, NSW,Vic., SA; mesopelagic. Sergestes seminudus Hansen, 1919. Maxilliped 3 as long as pereopod 3. Last 2 articles of pereopod 5 setose on both margins. Anterodorsal surface of the body with orange stellate chromatophores. tl. 55 mm. Indo-West Pacific, Qld, Vic.; slope, 250–350 m depth. Sergestes stimulator Burkenroad, 1940. Maxilliped 3 longer than pereopod 3. Last 2 articles of pereopod 5 setose on 1 margin. Rostrum short and deep, with a very short tip. cl. 16 mm. NSW; pelagic.

48 Dendrobranchiata – prawns and midwater shrimps

Fig. 8. Sergestidae. a, Sergestes armatus. b, Sergia prehensilis.

Sergia Stimpson, 1860 Sergia is difficult for the novice to differentiate from Sergestes (see above). Dermal photophores are regularly spaced, some in two curved rows on each side of the carapace, one above the branchiostegite and the other along the posterior half of the branchiostegite margin. These are often minute but are more easily visible on the scaphocerite, ventral margin of the abdominal somite 6,

49 Marine Decapod Crustacea of Southern Australia

and uropods. The photophores are visible only as opaque spots in preserved material except when they have domed cuticular lenses. They can number as many as 350. The ultrastructure of the photophores suggests that they are independently derived from other light-producing organs in other shrimps (Nowel et al., 2002). Vereshchaka (2000) reviewed the genus in a major study that included colour pictures. He also discussed the difficulties in recognising distinguishing characters, especially in the photophores. The key below is derived from his but should not be treated as exhaustive. Diagnoses are not given and new material should be confirmed using Vereshchaka (2000). Particular attention needs to be paid to the structure of the male petasma but this key avoids this feature. Because few surveys have been done in the region, chances of finding more species is high. Griffiths & Brandt (1983) recorded species of Sergia, some unidentified, associated with a warm-core eddy off eastern Australia. Diagnosis. Carapace with red cuticular pigment distributed over entire body and appendages; rostrum a short triangular projection. Dermal photophores usually present. Organs of Pesta absent. Pereopods 1 and 2 with ischium without well developed subterminal spine; pereopods 4 and 5 without dactyli.

Key to southern Australian species of Sergia 1. Hepatic tubercle absent; photophores absent ...... 2 — Hepatic tubercle or spine present; photophores present ...... 3 2. Cornea as wide as eyestalk; scaphocerite with inconspicuous distal tooth . . . Sergia japonica — Cornea wider than eyestalk; scaphocerite with clear but minute distal tooth . . . Sergia laminata 3. Ocular papilla present (inconspicuous); photophores without lenses; rostrum bidentate ...... Sergia potens — Ocular papilla absent; photophores with lenses; rostrum acute ...... 4 4. Lower branchiostegite row of 15–23 photophores; scaphocerite with 12–15 photophores; uropodal exopod with 8–14 photophores ...... Sergia prehensilis — Lower branchiostegite row of 8–14 photophores; scaphocerite with 7 photophores; uropodal exopod with 3 photophores ...... Sergia scintillans

a b

cd e

fg h i

Fig. 9. Sergestidae. Carapaces: a, Acetes sibogae. b, Petalidium foliaceum. c, Sergia laminata. d, Sergia scintillans. e, Sergia potens. Ends of uropodal exopods: f, Sergia japonica. g, Sergia laminata. h, Sergia potens. i, Sergia scintillans.

50 Dendrobranchiata – prawns and midwater shrimps

Sergia japonica (Bate, 1881) (Fig. 9f). Atlantic, Indian and Pacific oceans, Qld, NSW, Vic., Tas.; mesopelagic, to 1000 m depth. Sergia laminata (Burkenroad, 1940) (Fig. 9c, g). Atlantic, Indo-West to central Pacific oceans, NSW, WA; bathypelagic, to 2500 m depth. Sergia potens (Burkenroad, 1940) (Fig. 9e, h). Indian and Southern oceans, New Zealand, WA, Tas.; bathypelagic. Sergia prehensilis (Bate, 1881). Atlantic, Indo-West to central Pacific oceans, WA, NSW, Vic., Tas.; abyssopelagic to mesopelagic. Sergia scintillans (Burkenroad, 1940) (Fig. 9d, i). Indian, Pacific and Southern oceans, NSW, WA; bathypelagic, mesopelagic, oceanic province.

References Chen,Y.-H.,& Chen, I.-M. 2000. On the distribution of the epipelagic shrimp Acetes intermedius Omori, 1975 (Crustacea: Decapoda: Sergestidae) and its related environmental factors – a general review. Pp. 109–117 in: Hwang, J.-S., Wang, C.-H., & Chan, T.-Y. (eds), Proceedings of the International Symposium on Marine Biology in Taiwan – Crustacea and zooplankton taxonomy, ecology and living resources, 27–27 May, 1998, Taiwan. National Taiwan Museum: Taipei. Colefax, A.N. 1940. An Australian species of Acetes (Crustacea Macrura, Sergestidae) with remarks on the distribution and literature of the genus. Records of the Australian Museum 20: 341–352. Griffiths, F.B., & Brandt, S.B. 1983. Distribution of mesopelagic decapod Crustacea in and around a warm-core eddy in the Tasman Sea. Marine Ecology Progress Series 12: 175–184. Hale, H.M. 1941. Decapod Crustacea. British, Australian and New Zealand Antarctic Research Expedition, 1929–1931. Reports-Series B (Zoology and Botany) 4: 257–286, pl. 3. Hanamura, Y. 1999. Occurence of Acetes sibogae Hansen (Crustacea: Decapoda: Sergestidae) in Western Australia, with notes on the northern Australian population. Records of the Western Australian Museum 19: 465–468. Hanamura, Y., & Ohtsuka S. 2003. Occurence of intersex individuals in the sergestid shrimp, Acetes sibogae, in Darwin Harbour, Northern Territory, Australia. Crustaceana 76: 749–54. Kensley, B. 1972. Shrimps and Prawns of Southern Africa. South African Museum: Cape Town. 65 pp. Nowel, M.S., Shelton, P.M.J., Herring, P.J., & Gaten, E. 2002. Observations on the cuticular photophores of the sergestid shrimp Sergia grandis (Sund, 1920). Crustaceana 75: 551–566. Pérez Farfante, I., & Kensley, B. 1997. Penaeoid and sergestoid shrimps and prawns of the world. Keys and diagnoses for the families and genera. Mémoires du Muséum National d’Histoire Naturelle, Paris 175: 1–233. Vereshchaka, A.L. 2000. Revision of the genus Sergia (Decapoda: Dendrobranchiata: Sergestidae): taxonomy and distribution. Galathea Reports 18: 69–207, pls 2–5. Wasmer, R.A. 1993. Pelagic shrimps (Crustacea: Decapoda) from six USNS Eltanin cruises in the south- eastern Indian Ocean, Tasman Sea, and the southwestern Pacific Ocean to the Ross Sea. In: Biology of the Antarctic Seas 22. Antarctic Research Series 58: 49–91. Webber, W.R., Fenaughty, C.M., & Clark, M.R. 1990. A guide to some common offshore shrimp and prawn species of New Zealand. New Zealand Fisheries Occasional Publication 6: 1–42.

51 This page intentionally left blank 4. CARIDEA – SHRIMPS

In Australia members of the Caridea Dana, 1852 are best referred to as shrimps, a common name which conveniently distinguishes them from prawns, a word applied generally to the more edible members of the Penaeoidea, a superfamily of the Dendrobranchiata. In the USA shrimp is used in a culinary sense for penaeids, perhaps as well as for carideans. In southern Australia most small shrimp-like crustaceans from intertidal pools and subtidal benthic environments are carideans. Several other groups of crustaceans may be confused with caridean shrimps but if the second plate on the side of the abdomen curves forward and overlaps the one in front, it is a caridean. Mysids (opossum shrimps) have traditionally been regarded as members of the Superorder Peracarida (not covered in this book). In these transparent animals all the legs are similar and adapted for swimming rather than walking. There is an easily-seen spherical cavity containing a free particle (a statocyst) at the base of each uropod. Unlike benthic shrimps which rarely swim, mysids swarm just above the sediments surface in marine and estuarine habitats. Recent research based on molecular affinities rather than traditional morphological ones (Watling, 1999; Jarman et al., 2000) has shown that mysids may be decapods after all but the conservative view prevails in most classifications (Hessler & Watling, 1999; Martin & Davis, 2001). Confusion with other major crustacean taxa is more likely in pelagic habitats where caridean shrimps live with euphausiids (krill), lophogastrids (large mysid-like shrimps), dendrobranchiates (prawns and their relatives) and the rare amphionaceans. In none of these does the second abdominal plate overlap the first. The systematics of the Caridea has been well documented by two of the most thorough crustacean taxonomists of the twentieth century. Dr Lipke B. Holthuis was Curator of Crustacea at the National Museum of Natural History, Leiden, The Netherlands, until his retirement in 1992. His keys to families and genera first published in 1955 (Holthuis, 1955) was thoroughly revised almost 40 years later (Holthuis, 1993) and is an essential work for identification of any caridean to genus. The work is notable for the detailed synonymies but does not contain lists of species. Workers must go to local faunas for that information. Another key to families can be found in Chace (1992) and with the addition of a new family, another key to families in Vereshchaka (1997). Holthuis (1993) acknowledged that ‘the present form of [his] key to the caridean families is practically literally taken from his [Chace’s] recent paper on the subject’. Dr Fenner A. Chace, Emeritus Zoologist, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA, is another carcinologist who devoted his life to the taxonomy of decapods. Their arrangement of families within subfamilies is followed here. Another view of the higher superfamily classification of the Caridea is based on phylogenetic analysis (Christoffersen, 1990) but has almost no following among traditional workers who have seemed not to believe in the basis of the analysis. Holthuis (1993) admitted that his higher classification of the Caridea (in the Natantia, a taxon rarely recognised now) ‘will be generally considered old-fashioned’. A recent view (Dixon et al., 2003) places the Caridea as the most basal clade of the Pleocyemata, sister taxon to the Stenopodidea and Reptantia (all other non-dendrobranchiate Decapoda). These authors cited the ‘caridean lobe’ on maxilliped 1, chelate pereopods 1 and 2, and non-chelate pereopod 2 as defining characters.

53 Marine Decapod Crustacea of Southern Australia

h g e f

k l

n m

Fig. 10. Caridea. a, eye without ocellus. b, eye with ocellus. c, mandible with molar process and incisor. d, mandible with molar process. e, maxilliped 1 exopod abutting endite. f, maxilliped 1 far removed from endite. g, maxilliped 2 with last two articles side by side. h, maxilliped 2 with last two articles in sequence. i, arthrobranchs above pereopods 1–4. j, epipods (on maxilliped 3 and pereopods 1–4) without naked appendages. k, epipods ending in a naked appendages (Oplophoridae). l, detail of epipod ending in a naked appendage. m, pereopod 1 chela subchelate (Crangonidae). n, pereopod 1 chela prehensile.

54 Caridea – shrimps

The Caridea comprise some 325 genera worldwide of which only a small fraction are known to occur in southern Australia. Holthuis & Chace recognised 33 families but many are pelagic or demersal and oceanic and therefore rarely encountered in shallow coastal samples. All could be expected in tropical Australia. This key to families is an edited version of that of Holthuis and includes only those families so far reported from southern Australia. The even simpler version will do for most intertidal and shallow-water forms (see Box: Simple key to common shallow-water caridean families). Of the 130 species recorded, the number commonly seen in intertidal and shallow subtidal environments is few. The smallest shrimps are only a few millimetres long, the largest big enough to eat. Dimensions in this review are given as either total length (from the tip of the rostrum to the end of the telson) or as carapace length (from the eye to the back of the rostrum) depending on the conventions adopted in the literature for each family. In carideans, the legs are referred to as five pairs of pereopods, the first two pairs usually chelate. Maxilliped 3, the last of the mouthparts, is often as big as the first pereopods so it is often easier to count from the back to the front to get oriented. In many families, genera are distinguished by the presence, absence or position of carapace spines. A typical shrimp with these labelled appears in Fig. 183. Bauer’s (2004) book is a good overview of the natural history of caridean shrimps.

Diagnosis. Decapoda with well-developed abdomen; pleura of abdominal somite 2 overlapping those of somites 1 and 3. Pereopods 1 and 2 usually with chelae. Pereopod 3 without chelae. Gills phyllobranchiate. Female carrying developing eggs on pleopods.

Key to southern Australian superfamilies and families of Caridea Families marked * are mesopelagic or deep-sea and would not be encountered in shore or shelf benthic collections. 1. Pereopod 1 subchelate (palm transverse) (Fig. 10m) or prehensile (Fig. 10n) ...... Crangonoidea … 2 — Pereopod 1 truly chelate or rarely simple ...... 3 2. Pereopod 1 subchelate, dactylus closing against transverse palm which often ends in spine (Fig. 10m); carpus of pereopod 2 not subdivided; shallow water and deep-water ...... Crangonidae … p. 136 — Pereopod 1 prehensile, dactylus closing against inner surface of propodus (Fig. 10n); carpus of pereopod 2 subdivided; deep water ...... Glyphocrangonidae* … p. 141 3. Pereopods 1 and 2 similar, with long slender pectinate (comb-like) fingers (Fig. 11a) ...... Pasiphaeoidea: Pasiphaeidae … p. 158 — Pereopods 1 and 2 usually dissimilar, without pectinate fingers ...... 4 4. Carpus of pereopod 2 entire, not subdivided; pereopod 1 always with strong chela ...... 5 — Carpus of pereopod 2 usually subdivided into 2 or more short articles (Fig. 11b); if not, pereopod 1 not chelate ...... 13 5. Pereopods 1 and 2 similar, fingers extremely long, >5 times as long as short palm, fringed with long setae; maxilliped 2 with last 2 articles side by side (Fig. 10g) (deep-water only) ...... Stylodactyloidea: Stylodactylidae* … p. 78 — Pereopods 1 and 2 fingers not extremely long; maxilliped 2 with last 2 articles in sequence (Fig. 10h) ...... 6 6. Pereopods with epipods, ending in a naked appendix extending well into branchial cavity (Fig. 10k, l); pereopods 1 and 2 similar ...... Oplophoroidea: Oplophoridae* … p. 63 — Pereopodal epipods, if present, not ending in a naked appendix (Fig. 10j); pereopods 1 and 2 similar or dissimilar ...... 7

55 Marine Decapod Crustacea of Southern Australia

7. Pereopods 1 and 2 with similar chelae, fingers ending in dense brush of setae; estuarine ...... Atyoidea: Atyidae … p. 69 — Pereopods 1 and 2 with fingers not ending in dense brush of setae; marine ...... 8 8. Pereopods 1 stronger and heavier, though often shorter than pereopods 2 ...... 9 — Pereopods 1 usually more slender (rarely subequal) than pereopods 2 ...... 11 9. Pereopods without strap-like epipods; mandibular molar process conical, laminar or vestigial ...... Bresilioidea: Bresiliidae* … p. 70 — Pereopods 1–3 at least with strap-like epipods; mandibular molar process blunt, with grinding surface ...... Nematocarcinoidea … 10 10. Rostrum finely dentate; pereopods 1 and 2 slender, fingers without long spines; deep-sea ...... Nematocarcinidae* … p.73 — Rostrum grossly dentate, hinged and movable; pereopods 1 and 2 with long spines on fingers, forming a basket-like cage when closed ...... Rhynchocinetidae … p. 75 11. Pereopods 1–4 with arthrobranchs (Fig. 10i); antennule with dorsal flagellum simple; deep-water species ...... Campylonotoidea: Campylonotidae* … p. 80 — Pereopods without arthrobranchs; antennule with dorsal flagellum of 2 branches; shallow-water species ...... Palaemonoidea … 12 12. Maxilliped 3 slender, pereopod-like; mandible usually with incisor (Fig. 10c) ...... Palaemonidae … p. 84 — Maxilliped 3 with ischiomerus broad, sometime operculate; mandible without incisor (Fig. 10d) ...... Gnathophyllidae* … p. 81 13. Carapace merging anteriorly into an inflated, indiscrete rostrum; pereopod 2 with fixed finger curved around short dactylus ...... Physetocarididae* … p. 135 — Rostrum, if present, not as above; pereopod 2 with conventional chela ...... 14 14. Right pereopod 1 chelate, left usually simple; maxilliped 1 with exopod abutting endite, displacing palp (endopod) (Fig. 10e) ...... Processoidea: Processidae … p. 128 — Pereopods 1 both simple or chelate; maxilliped 1 with exopod far removed from endite (Fig. 10f) ...... 15 15. Pereopod 1 with chelae microscopic or simple ...... Pandaloidea: Pandalidae … p. 129 — Pereopod 1 distinctly chelate ...... 16 16. Eyestalks unusually elongate, as long as end of antennule peduncle; pereopods 1 and 2 equally strong ...... Ogyrididae … p. 126 — Eyestalks of normal length; pereopod 1 stronger than 2 ...... Alpheoidea … 17 17. Eyes usually covered partly or entirely by carapace; pereopods 1 unequal, larger one swollen ...... Alpheidae … p. 98 — Eyes not covered; pereopods usually equal, not swollen ...... Hippolytidae … p. 118

56 Caridea – shrimps

Fig. 11. Caridea. a, pereopod 1 with pectinate fingers (Pasiphaeidae). b, pereopod 2 with divided carpus (e.g. Alpheidae, Hippolytidae).

References Bauer, R.T. 2004 Remarkable Shrimps. Adaptations and Natrual History of the Caridean Shrimps. University of Oklahoma Press: Norman. 282 pp. Chace, F.A. 1992. On the classification of the Caridea (Decapoda). Crustaceana 63: 70–80. Christoffersen, M.L. 1990. A superfamily classification of the Caridea (Crustacea: Pleocyemata) based on phylogenetic pattern. Zeitschrift für Zoologische Systematik und Evolutionsforschung 28: 94–106. Dixon, C.J., Ahyong, S., & Schram, F.R. 2003. A new hypothesis of decapod phylogeny. Crustaceana 76: 935–975. Hessler, R.R., & Watling, L. 1999. Les péracarides : un groupe controversé. In: Forest, J. Traité de zoologie. Anatomie, systématique, biologie. Tome 7 Crustacés Fascicule 3A Péracarides. Mémoires de l’Institut Océanographique. Monaco 19: 1–10. Holthuis, L.B. 1955. The recent genera of the caridean and stenopodidean shrimps (Class Crustacea, Order Decapoda, Supersection Natantia) with keys for their determination. Zoologische Verhandelingen, Leiden 26: 1–153. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Jarman, S.N., Nicol, S., Elliott, N.G., & McMinn, A. 2000. 28rDNA evolution in the Eumalacostraca and the phylogenetic position of krill. Molecular Phylogenetics and Evolution 17: 26–36. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. Vereshchaka, A.L. 1997. New family and superfamily for a deep-sea caridean shrimp from the Galathea collections. Journal of Crustacean Biology 17: 361–373. Watling, L. 1999. Toward understanding the relationships of the peracaridan orders: the necessity of determining exact homologies. Pp. 73–89 in: Vaupel Klein, J.C.v. (ed.) Crustaceans and the biodiversity crisis. Proceedings of the Fourth International Crustacean Congress, Amsterdam, The Netherlands, July 20–24, 1998. Brill: Leiden.

57 Marine Decapod Crustacea of Southern Australia

SIMPLE KEY TO COMMON SHALLOW-WATER CARIDEAN FAMILIES This key can be used to identify most shrimps from intertidal and shallow subtidal environments to family. It eliminates common and distinctive families first. It is not useful for deep- water benthic or mesopelagic families. The genera and species listed are those commonly encountered in southern shelf or estuarine environments. Species of the last two families are possibly the most difficult to distinguish but unfortunately are the most common in the intertidal environment. First, ensure that you do not have a mysid, a shrimp-like crustacean with oval cavities inside the base of the uropods. 1. Eyes covered partly or entirely by carapace; pereopods 1 unequal, usually larger one swollen and with dactylus like a parrot’s beak (snapping shrimps) ...... Alpheidae — Not so ...... 2 2. Estuarine or freshwater near the coast; chelae with fingers ending in dense brush of setae ...... Atyidae (Paratya australiensis) — Marine; chelae without terminal brush of setae ...... 3 3. Carapace somewhat flattened; chelipeds with finger closing on a transverse palm ...... Crangonidae (Philoceras spp.) — Carapace more laterally flattened; chelipeds with typical pincer ...... 4 4. Eyes extremely elongate; shelf environment only ...... Ogyrididae (Ogyrides delli) — Not so ...... 5 5. Rostrum a short spine; pereopods 1 and 2 with long slender pectinate fingers ...... Pasiphaeidae (Leptochela sydniensis) — Rostrum usually elongate; pereopods without pectinate fingers ...... 6 6. Rostrum hinged, movable ...... Rhynchocinetidae (Rhynchocinetes spp.) — Rostrum fixed ...... 7 7. Pereopod 2 with subdivided carpus (3 or more short segments between 2 proximal long articles and distal palm and finger) ...... Hippolytidae — Pereopod 2 with simple carpus (single short article between 2 proximal long articles and distal palm and finger) ...... Palaemonidae

Superfamily Pasiphaeoidea Dana, 1852 Only one family is included.

Pasiphaeidae Dana, 1852 The Pasiphaeidae are a family of benthic or mesopelagic shrimps from shelf and offshore environments. Its members are immediately distinguished by the possession of the comb-like fingers on the first two pairs of pereopods. There are seven genera, six represented in Australia (Bruce, 1990; Holthuis, 1993; Hanamura & Evans, 1994; Hayashi, 1999). One species, Leptochela sydniensis,is common on the shelf of southern Australia while the only other representatives in this region are mesopelagic in deeper water (Kensley et al., 1987; Hanamura, 1989).

58 Caridea – shrimps

Diagnosis. Carapace smooth or with obscure lateral ridges; with or without antennal and branchiostegal spines. Rostrum absent, short or a deep blade. Maxilliped 2 without exopod. Pereopods 1 and 2 similar, with long slender pectinate fingers (bearing rows of fine closely spaced spines along cutting edge).

Key to southern Australian species of Pasiphaeidae 1. Mandibular palp absent; rostrum an erect spine posterior to front ...... Pasiphaea … 2 — Mandibular palp present; rostrum a normal projection of the carapace ...... 7 2. Carapace dorsally carinate. Posterior margin of telson deeply notched ...... 3 — Carapace dorsally rounded. Posterior margin of telson truncate or very slightly notched . 5 3. Merus of pereopod 1 with 0–3 spines ...... Pasiphaea barnardi — Merus of pereopod 1 with up to 8 spines ...... 4 4. Rostrum not reaching front of carapace, its anterior margin straight ...... Pasiphaea tarda — Rostrum extending well beyond front of carapace; its anterior margin convex ...... Pasiphaea rathbunae 5. Abdominal somites 1–5 dorsally rounded, somite 6 dorsally carinate . . . Pasiphaea longitaenia — All abdominal somites dorsally rounded ...... 6 6. Pereopod 2 merus with 6–11 ventral spines, suprabranchial ridge barely visible ...... Pasiphaea kapala — Pereopod 2 merus with 3–6 ventral spines, suprabranchial ridge well developed; without arthrobranchs ...... Pasiphaea australis 7. Pereopod 4 longer than pereopod 5, shorter than pereopod 3 ...... Leptochela sydniensis — Pereopod 4 distinctly shorter than either pereopods 3 or 5 ...... 8 8. Antennal and branchiostegal spines absent; dorsal margin of carapace not toothed ...... Parapasiphae sulcatifrons — Antennal and branchiostegal spines present; dorsal margin of carapace toothed ...... 9 9. Maxilliped 3 with 2 arthrobranchs ...... Eupasiphae gilesii — Maxilliped 3 with 1 arthrobranch ...... Glyphus marsupialis

Eupasiphae Wood-Mason & Alcock, 1893 Species of Eupasiphae are oceanic midwater shrimps recognisable because of the erect blade-like rostrum extending over the eyes. Two species occur in tropical Australia (Hanamura & Evans, 1994) but only one in southern waters. A similar pasiphaeid species, Glyphus marsupialis, has a smaller rostrum, abdominal somite 1 only obscurely ridged or rounded, and only one arthrobranch on maxilliped 3. Diagnosis. Rostrum an erect blade. Antennal and branchiostegal spines present. Dorsal margin of carapace toothed. Mandibular palp present. Pereopod 4 distinctly shorter than pereopod 5. Maxilliped 3 with 2 arthrobranchs. Eupasiphae gilesii (Wood-Mason & Alcock, 1893) (Fig. 12c). Rostrum not strongly compressed laterally and nearly triangular in adults. Carapace and abdomen dorsally carinate and serrate; abdominal somite 4 ending in medial spine. Branchiostegal spine immediately posterior to anterolateral margin of carapace. Telson dorsally sulcate, without spiniform setae. cl. 45 mm. Atlantic and Indian oceans, WA, NSW; mesopelagic, 340–770 m depth.

59 Marine Decapod Crustacea of Southern Australia

b e

h j

k f l g m

Fig. 12. Pasiphaeidae. a, Leptochela sydniensis. b, Pasiphaea barnardi. Carapace. c, Eupasiphae gilesii. d, Glyphus marsupialis. e, Parapasiphae sulcatifrons. f, g, h, Pasiphaea australis (proximal articles and epipod of pereopods 1, 2, chela of pereopod 2). i, Pasiphaea barnardi (pereopod 2). j, k, Pasiphaea kapala (meri of pereopods 1 and 2). l, m, Pasiphaea longitaenia (meri of pereopods 1 and 2).

Glyphus Filhol, 1884 A single species of this genus has been recorded in southern Australian waters. Diagnosis. Rostrum a normal anteriorly-directed prolongation of the carapace. Antennal and branchiostegal spines present. Mandibular palp present. Maxilliped 3 with 1 arthrobranch. Dorsal margin of carapace usually with teeth. Pereopod 4 distinctly shorter than pereopods 3 and 5. Glyphus marsupialis Filhol, 1884 (Fig. 12d). cl. 68 mm. Eastern Atlantic to eastern Pacific throughout the Indo-West Pacific; all Australia; mesopelagic, 1061–1080 m depth. This species was first recorded in the Pacific by Hanamura & Evans (1994) and only one subsequent record has been made from southern Australian waters (Museum Victoria collection).

60 Caridea – shrimps

Leptochela Stimpson, 1860 At least 14 species of Leptochela exist, six in northern Australia (Hanamura, 1987) and one in southern Australia. The genus was reviewed and all its species described in detail by Chace (1976). Diagnosis. Rostrum a short anteriorly-directed prolongation of the carapace, dorsally without teeth. Mandibular palp present. Pereopod 4 shorter than pereopod 3, longer than pereopod 5. Leptochela sydniensis Dakin & Colefax, 1940 (Fig. 12a). Ventral margin of orbit and suborbital angle unarmed. Antennal scale half as long as carapace. Pereopod 1 dactylus with 20–44 spines. Abdominal somite 5 with dorsal margin convex in lateral view, unarmed; abdominal somite 6 without dorsal lobe. Telson with 1 pair of dorsolateral spines set well posterior to pair of mesial spines. cl. 4.7 mm. Indo-West Pacific, WA, NT, NSW, Vic., Tas., SA; 16–400 m depth. This common shrimp is easily distinguished from all other shallow water carideans by its short pointed rostrum reaching only as far as the end of the eyestalk.

Parapasiphae Smith, 1884 Only one of species of this small genus has been recorded in southern Australian waters. Diagnosis. Rostrum a normal anteriorly-directed prolongation of the carapace. Mandibular palp of 2 articles. Antennal and branchiostegal spines absent. Dorsal margin of carapace usually without teeth. Fingers of chela of pereopod 2 not longer than palm. Pereopod 4 distinctly shorter than pereopods 3 and 5. Parapasiphae sulcatifrons Smith, 1884 (Fig. 12e). cl. 25 mm. Western Pacific, North and South Atlantic, Indian Ocean, off W coast of USA, NSW, Tas.; mesopelagic, 500–5340 m depth. Kensley et al. (1987) recorded the first specimen from the western Pacific.

Pasiphaea Savigny, 1816 This genus of perhaps a dozen species is well represented in midwater sampling in most oceans including Australia (Kensley et al., 1987; Hanamura, 1994; Hanamura & Evans, 1994). The species are superficially similar to sergestid prawns which often occur in the same mesopelagic samples. They can be separated by the reliable caridean character of abdominal somite 2 pleura. Three subantarctic species that may reach southern Australian oceanic waters are P. acutifrons Bate, 1888, P. balssi Burukovsky & Romensky, 1987 and P. scotiae (Stebbing, 1914) (Davie, 2002). Diagnosis. Rostrum formed by an erect postfrontal spine. Mandible without palp. Antennal spine absent. Branchiostegal spine present. Dorsal margin of carapace carinate, without teeth. Fingers of chela of pereopod 2 not longer than palm. Pereopod 4 distinctly shorter than pereopods 3 and 5. Pasiphaea australis Hanamura, 1989 (Fig. 12f, g, h). Carapace dorsally rounded. Suprabranchial ridge well developed. Merus of pereopod 1 unarmed. Pereopod 2 merus with 3–6 ventral spines. All abdominal somites dorsally rounded. Posterior margin of telson truncate or very slightly notched. cl. 16.5 mm. NSW, Vic., Tas., SA, WA; mesopelagic, 32–450 m depth. The species was referred to the genus Alainopasiphaea Hayashi, 1999 by Hayashi (2004). Pasiphaea barnardi Yaldwyn, 1971 (Fig. 12i). Carapace dorsally carinate. Merus of pereopod 1 with 0–3 spines, merus of pereopod 2 with 12–17 spines on posterior margin. Abdominal somites 2–6 sharply carinate dorsally. Telson deeply notched distally. cl. 42 mm. Southern Atlantic, Indo-West Pacific, WA, NSW, Vic.; mesopelagic, 1030–1140 m depth. Hanamura & Evans (1994) believed this species to be a senior synonym of P. berentsae Kensley et al., 1987. Pasiphaea kapala Kensley, Tranter & Griffin, 1987 (Fig. 12j, k). Carapace dorsally rounded. Merus of pereopod 1 unarmed; merus of pereopod 2 with 6–11 spines on posterior margin. All abdominal somites dorsally rounded. Posterior margin of telson truncate to very slightly notched. WA, NSW; mesopelagic, 446–900 m depth.

61 Marine Decapod Crustacea of Southern Australia

Pasiphaea longitaenia Kensley, Tranter & Griffin, 1987 (Fig. 12l, m). Carapace dorsally rounded. Merus of pereopod 1 armed with 9–14 spines; merus of pereopod 2 with 19–21 spines on posterior margin. Abdominal somites 1–5 dorsally rounded, somite 6 slightly compressed laterally along dorsal midline, but not sharply carinate, and ending in dorsomedial spine. Posterior margin of telson truncate or very slightly notched. cl. 22 mm. NSW; mesopelagic, 887–893 m depth. Pasiphaea tarda Krøyer, 1845. Carapace dorsally carinate almost entire length; rostrum not reaching frontal margin of carapace, its anterior margin vertical. Merus of pereopod 1 with 1–8 spines, merus of pereopod 2 with 11–22 spines. Abdominal somites 2–6 carinate dorsally. Posterior margin of telson deeply notched distally. cl. 63 mm. Atlantic, Indo-West Pacific, southern WA (Hanamura & Evans, 1994); mesopelagic, 880–960 m depth. Pasiphaea rathbunae (Stebbing, 1914). Carapace dorsally carinate almost entire length; rostrum overlapping frontal margin of carapace, its anterior margin convex. Merus of pereopod 1 with ?5–9 spines, merus of pereopod 2 with ?7–17 spines. Abdominal somites 2–6 carinate dorsally. Posterior margin of telson deeply notched distally. cl. 45 mm. Tas.; mesopelagic, 1710 m. A single specimen of this South African species was recorded by Hale (1941).

References Bruce, A.J. 1990. Two deep-sea shrimps new to the Australian fauna, Psathyrocaris hawaiiensis Rathbun (Pasiphaeidae) and Bresilia antipodarum, sp. nov. (Bresiliidae), with remarks on Encantada spinoc- ulata Wicksten (Bresiliidae). Invertebrate Taxonomy 4: 847–866. Chace, F.A. 1976. Shrimps of the pasiphaeid genus Leptochela with descriptions of three new species (Crustacea: Decapoda: Caridea). Smithsonian Contributions to Zoology 222: 1–51. Crosnier, A., & Forest, J. 1973. Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19: 1–409. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Hale, H.M. 1941. Decapod Crustacea. British, Australian and New Zealand Antarctic Research Expedition, 1929–1931. Reports-Series B (Zoology and Botany) 4: 257–286, pl. 3. Hanamura, Y. 1987. Caridean shrimps obtained by R.V.‘Soela’ from north-west Australia, with description of a new species of Leptochela (Crustacea: Decapoda: Pasiphaeidae). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 4: 15–33. Hanamura, Y. 1989. Deep-sea shrimps (Crustacea: Decapoda) collected by the R.V.‘Soela’ from southern Australia. Bulletin of the Biogeographical Society of Japan 44: 51–69. Hanamura, Y. 1994. A new species of Pasiphaea Savigny (Crustacea: Caridea: Pasiphaeidae) from north- western Australian waters. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11: 167–173. Hanamura, Y., & Evans, D.R. 1994. Deepwater caridean shrimps of the families Oplophoridae and Pasiphaeidae (Crustacea: Decapoda) from Western Australia, with an appendix on a lophogastri- dan mysid (Mysidacea). Crustacean Research 23: 46–60. Hayashi, K.-I. 1999. Crustacea Decapoda: Revision of Pasiphaea sivado (Risso, 1816) and related species, with descriptions of one new genus and five new species (Pasiphaeidae). In: Crosnier, A. (ed), Résultats des Campagnes MUSORSTOM, Vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180: 267–302. Hayashi, K.-I. 2004. Revision of Pasiphaea cristata Bate, 1888 species group of Pasiphaea Savigny, 1816, with descriptions of four new species and referral of P. australis to Alainopasiphaea Hayashi, 1999 (Crustacea: Decapoda: Pasiphaeidae). In: Marshall, B.A., and Richer de Forges, B. (eds), Tropical Deep-Sea Benthos, Vol 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191: 319–373. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331.

62 Caridea – shrimps

Superfamily Oplophoroidea Dana, 1852 Oplophoridae is the only family.

Oplophoridae Dana, 1852 The Oplophoridae are a diverse family of oceanic usually mesopelagic shrimps recognisable by the unusual pereopodal epipod which is L- or C-shaped (Fig. 14i). This feature distinguishes them from the nematocarcinids with which they are often taken in midwater nets. Of the ten genera recognised by Chace (1986) and Holthuis (1993) six have been reported from southern Australia. The only Australian records certainly do not accurately reflect true distributions. Several species found in Australia were described by Crosnier & Forest (1973). The shrimps are probably biolu- minescent. Diagnosis. Carapace sometimes with longitudinal ridges. Rostrum usually well developed and toothed, sometimes obsolete. Eyes visible. Pereopod 1 chelate, similar to pereopod 2. Pereopod 2 carpus not divided. Pereopods with epipods terminating in naked appendix extending vertically between branchia, and exopods.

Key to southern Australian species of Oplophoridae 1. Abdomen carinate middorsally on last 4 somites at least ...... 2 — Abdomen not carinate middorsally on somite 6 (obviously carinate on other somites) . . 10 2. Carapace with continuous lateral carina(e); posterior margin of hepatic groove marked by oblique carina ...... 3 — Carapace without continuous lateral carina; posterior margin of hepatic groove not marked by oblique carina ...... Acanthephyra … 6 3. Carapace with 1 lateral longitudinal carina; dorsal margin of carapace denticulate only anteriorly; abdominal somite 1 without mid-dorsal carina ...... Meningodora vesca — Carapace with more than 1 lateral longitudinal carina; dorsal margin of carapace denticulate over most of length; all abdominal somites with mid-dorsal carina ...... Notostomus … 4 4. Base of rostrum with 1 lateral carina on dorsal margin of orbit ...... Notostomus gibbosus — Base of rostrum with 2 lateral carinae ...... 5 5. Upper lateral rostral carina not extending posteriorly beyond orbit, anteriorly not as far as lower lateral rostral carina; abdominal somite 1 with median dorsal carina more than half its length ...... Notostomus elegans — Upper lateral rostral carina extending posteriorly beyond orbit, anteriorly beyond lower lateral rostral carina; abdominal somite 1 with median dorsal carina less than half its length ...... Notostomus auriculatus 6. Rostrum less than half length of carapace, with 1 ventral tooth ...... 7 — Rostrum more than half length of carapace, with several dorsal and ventral teeth ...... 8 7. Telson with 5–6 pairs of dorsolateral spiniform setae; abdominal somite 1 carinate ...... Acanthephyra acutifrons — Telson with 8–9 pairs of dorsolateral spiniform setae; abdominal somite 1 not carinate ...... Acanthephyra curtirostris 8. Telson with 7–11 pairs of dorsolateral spiniform setae ...... Acanthephyra sica — Telson with 3–4 pairs of dorsolateral spiniform setae ...... 9

63 Marine Decapod Crustacea of Southern Australia

9. Abdominal somite 1 with V-shaped notch in dorsal midline; telson with 3–4 pairs of dorsolateral spiniform setae ...... Acanthephyra smithi — Abdominal somite 1 with shallow sinuous dorsal midline margin; telson with 4 pairs of dorsolateral spiniform setae ...... Acanthephyra quadrispinosa 10. Pereopod 4 with epipod vestigial or absent; rostrum shorter than or just longer than eyestalk ...... Hymenodora …11 — Pereopod 4 with epipod well developed; rostrum much longer than eyestalk ...... 12 11. Carapace with hepatic region without a posterodorsal transverse groove between longitudinal grooves; rostrum just overreaching eyestalk ...... Hymenodora gracilis Carapace with hepatic region with a posterodorsal transverse groove between longitudinal grooves; rostrum falling short of eyestalk ...... Hymenodora glacialis 12. Abdominal somite 6 nearly twice as long as 5; telson with spinose end piece flanged by lateral spiniform setae ...... Systellaspis debilis — Abdominal somite 6 shorter than 1.5 times length of 5; telson with simple apex ...... Oplophorus … 13 13. Scaphocerite laterally spinose ...... Oplophorus spinosus — Scaphocerite laterally smooth ...... Oplophorus novaezeelandiae

Acanthephyra Milne Edwards, 1881 Acanthephyra is a genus of almost 30 species, most mesopelagic in oceanic waters (Chace, 1986). The narrow rostrum with regular rows of teeth and with a thickened base is characteristic. Kensley et al. (1987), Hanamura (1989) and Hanamura & Evans (1994) have published records of five species in Australia. Species occur together and with Systellaspis debilis which is superficially similar. Diagnosis. Rostrum narrow, with about as many dorsal as ventral teeth. Carapace not denticulate dorsally, usually without uninterrupted lateral carina, without hepatic spine. Scaphocerite without lateral teeth. Abdomen dorsally carinate on at least somites 3–6; somite 6 not more than 1.5 times somite 5. Telson blunt apically, without spinose end piece. Acanthephyra acutifrons Bate, 1888 (Fig. 14a). Rostrum shorter than half carapace, strongly tapered, with 1 ventral tooth; carapace with 11 teeth at base of rostrum. Abdominal somites all carinate, 3–6 with posteromedial tooth. Telson sulcate in midline, with 5–6 pairs of dorsolateral spiniform setae. Uniformly bright red. 160 mm. Indo-West Pacific, Atlantic, off central NSW; mesopelagic, 650–2400 m depth. Acanthephyra curtirostris Wood-Mason, 1891 (Fig. 14b). Rostrum shorter than half carapace, strongly tapered, with 1 ventral tooth; carapace with 11 teeth at base of rostrum. Abdominal somites 2–6 carinate, 3–6 with posteromedial tooth. Telson sulcate in midline, with 6–9 pairs of dorsolateral spiniform setae. Uniformly bright red. 60 mm. Indo-West Pacific, north-eastern Atlantic, Caribbean, off south to central NSW; mesopelagic, 630–4970 m depth. Acanthephyra quadrispinosa Kemp, 1939 (Figs 13a, 14l). Rostrum longer than carapace, with 8–9 dorsal and 5 ventral teeth. Abdominal somites 2–6 carinate, 3–6 with posteromedial tooth, 1 without medial V-shaped notch. Telson sulcate in midline, with 4 pairs of dorsolateral spiniform setae. Uniformly bright red. 160 mm. Indo-Pacific, south Atlantic, off west and east coasts S to Tas., western Vic.; mesopelagic, 250–1700 m depth. A. quadrispinosa is the commonest oplophorid in collections from eastern Australia, distinguishable from the less common A. sica which has more pairs of spiniform setae on the telson. It is an important component of the fauna of the warm-core eddies derived from the Eastern Australian Current (Griffiths & Brandt, 1983).

64 Caridea – shrimps

Fig. 13. Oplophoridae. a, Acanthephyra quadrispinosa. b, Notostomus auriculatus. c, Oplophorus novaezeelandiae.

65 Marine Decapod Crustacea of Southern Australia

Acanthephyra sica Bate, 1888 (Fig. 14m). Rostrum about as long as carapace, with 8–9 dorsal and 5–6 ventral teeth. Abdominal somites 2–6 carinate, 3–6 with posteromedial tooth, 1 with medial V-shaped notch. Telson sulcate in midline, with 7–11 pairs of dorsolateral spiniform setae. Uniformly bright red. 140 mm. Southern circumpolar, NSW, S to Tasman Rise; mesopelagic, 540–2166 m depth. The species has been reported as A. pelagica (Burukovsky & Romensky, 1982). Acanthephyra smithi Kemp, 1939. Rostrum more than half length of carapace, with 7–9 dorsal and 5 ventral teeth. Abdominal somites 2–6 carinate, 3–6 with posteromedial tooth, 1 with medial V-shaped notch. Telson sulcate in midline, with 3–4 pairs of dorsolateral spiniform setae. Uniformly bright red. 140 mm. Indo-Pacific, west and east coasts S to central NSW; mesopelagic, 216–800 m depth.

abc

def

h l m n

Fig. 14. Oplophoridae. Carapace: a, Acanthephyra acutifrons. b, Acanthephyra curtirostris. c, Hymenodora glacialis. d, Hymenodora gracilis. e, Meningodora vesca. f, Notostomus gibbosus. g, Notostomus elegans. h, Systellaspis debilis. i, Notostomus spp. (epipod and arthrobranch). Scaphocerite: j, Oplophorus novaezeelandiae. k, Oplophorus spinosus. Telson: l, Acanthephyra quadrispinosa. m, Acanthephyra sica. n, Systellaspis debilis.

Hymenodora Sars, 1877 Hymenodora is a genus of four species of mesopelagic and bathypelagic shrimps found in all oceans (Chace, 1986). Diagnosis. Rostrum narrow, with dorsal teeth only. Carapace not denticulate dorsally, without uninterrupted lateral carina, without hepatic spine. Abdomen without carina or posterior tooth

66 Caridea – shrimps

on somites 3–5; somite 6 longer than somite 5. Telson blunt, not tapering, rarely with spinose end piece. Scaphocerite without lateral teeth. Hymenodora glacialis (Buchholz, 1874) (Fig. 14c). Rostrum shorter than eyestalk. Carapace with hepatic region with a posterodorsal transverse groove between longitudinal grooves. Maxilliped 2 without a podobranch. Scaphocerite with subtruncate apex. Telson without spinose end piece. Atlantic, Indo-West Pacific, SA, WA; mesopelagic, to 5686 m depth. Hymenodora gracilis Smith, 1886 (Fig. 14d). Rostrum just longer than eyestalk, with 1–5 dorsal teeth. Carapace with hepatic region without a posterodorsal transverse groove between longitudinal grooves. Maxilliped 2 with podobranch. Scaphocerite with subtruncate apex. Telson without spinose end piece. cl. 15 mm. Indo-Pacific, Atlantic, Southern Ocean, off Tas.; mesopelagic, 500–5394 m depth.

Meningodora Smith, 1882 The short rostrum and single lateral carina distinguish this genus of five species from Notostomus (Chace, 1986). One species occurs rarely off southern NSW and another, M. mollis Smith, 1882, further north. Diagnosis. Rostrum tapering, with more dorsal than ventral teeth. Carapace not denticulate dorsally, with 1 lateral carina, without hepatic spine. Scaphocerite without lateral teeth. Abdomen dorsally carinate on at least somites 3–6, dentate on 4–6; somite 6 as long or longer than somite 5. Telson blunt apically, without spinose end piece. Meningodora vesca (Smith, 1886) (Fig. 14e). Rostrum one-third carapace length, with 9–12 dorsal and 1–2 ventral teeth. Abdominal somites 1–2 not carinate, somite 3 without posteromedial tooth, somites 4–6 carinate with tooth. 50 mm. Indo-West Pacific, Atlantic, off Qld, NSW; mesopelagic, to 950 m depth.

Notostomus Milne Edwards, 1881 The high crest on the carapace and tapering rostrum identify species of the mesopelagic genus Notostomus (Chace, 1986). Diagnosis. Rostrum tapering, with more dorsal than ventral teeth. Carapace with denticulate carina dorsally, with 2 or more lateral carinae, without hepatic spine. Scaphocerite without lateral teeth. Abdomen dorsally carinate on all somites; somite 6 longer than somite 5. Telson blunt apically, without spinose end piece. Notostomus auriculatus Barnard, 1950 (Fig. 13b). Base of rostrum with 2 lateral carinae, upper extending posteriorly beyond orbit, anteriorly further than lower lateral rostral carina; abdominal somite 1 with median dorsal carina less than half its length. 180 mm. South Atlantic, off NSW, Tas.; mesopelagic, to 900 m depth. Kensley et al. (1987) recorded a species as N. cf. crosnieri Macpherson, 1984 which seems very similar to this (Hanamura, 1989). Several specimens of this species (and a lophogastrid) were found in the stomach of a pygmy sperm whale (Museum Victoria record). Notostomus elegans Milne Edwards, 1881 (Fig. 14g). Base of rostrum with 2 lateral carinae, upper not extending posteriorly beyond orbit, nor anteriorly beyond lower lateral rostral carina; abdominal somite 1 with median dorsal carina more than half its length. 140 mm. Pacific and Atlantic, NSW; mesopelagic, 630–2780 m depth. Kensley et al. (1987) discussed variation in this species. Notostomus gibbosus Milne Edwards, 1881 (Fig. 14f). Base of rostrum with 1 lateral carina on dorsal margin of orbit, continuous with dorsal longitudinal lateral carina. Abdominal somite

67 Marine Decapod Crustacea of Southern Australia

1 with median dorsal carina more than half its length. 180 mm. Indo-West Pacific, Atlantic, WA, NSW; mesopelagic, 569–3932 m depth.

Oplophorus Milne Edwards, 1837 The elongate spines on the posterior margins of three abdominal somites characterise the four mesopelagic species of Oplophorus (Chace, 1986; Hanamura & Evans, 1994). Diagnosis. Rostrum narrow, with similar number or more dorsal than ventral teeth. Carapace not denticulate dorsally, without uninterrupted lateral carina, without hepatic spine. Scaphocerite with or without lateral teeth. Abdomen with strong posterior tooth on somites 3–5; somite 6 almost as long as somite 5. Telson acute posteriorly, without spinose end piece. Oplophorus novaezeelandiae de Man, 1931 (Fig. 14j). Carapace with posteroventral corner unarmed. Scaphocerite smooth along inner and outer margins. 50 mm. Indo-West Pacific, south-west Atlantic, throughout Australia; mesopelagic, 90–725 m depth. Oplophorus spinosus Brullé, 1839 (Fig. 14k). Carapace with posteroventral corner unarmed. Scaphocerite with obsolete spine on inner margin, denticulate on outer margin. 50 mm. Indo- West Pacific, Atlantic, throughout Australia; mesopelagic, 10–3923 m depth.

Systellaspis Bate, 1888 There are only five species of Systellaspis, most mesopelagic or benthic in tropical and temperate seas (Chace, 1986). Two occur in WA (Hanamura & Evans, 1994). Diagnosis. Rostrum narrow, with more dorsal than ventral teeth. Carapace not denticulate dorsally, without uninterrupted lateral carina, without hepatic spine. Scaphocerite without lateral teeth. Abdomen dorsally carinate on at least somites 3–5; somite 6 almost as long as somite 5. Telson with spinose end piece, flanked by pair of long spiniform setae. Systellaspis debilis Milne Edwards, 1881 (Fig. 14h, n). Rostrum little longer than carapace, with 12 dorsal teeth starting on a basal crest, 8–11 ventral teeth. Carapace without a lateral ridge. Abdominal somites 3 and 4 carinate, 3 with terminal spine; posterior margin of somite 4 dorsolateral denticulate. Telson with 4–8 pairs of dorsolateral spiniform setae. 70 mm. Indo- West Pacific, Atlantic, west and east coasts S to Tas.; mesopelagic, 150–1500 m depth. S. debilis can be confused with species of Acanthephyra but the lack of a posterior spine on abdominal somite 6 and the denticles along the margins of somite 4 are diagnostic.

References Burukovsky, R.N., & Romensky, L.L. 1982. New findings of several species of shrimps and description of Pasiphea natalensis sp. n. Zoologicheskii Zhurnal, Moscow 61: 1797–1801 (in Russian with English summary). Chace, F.A. 1986. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 4: families Oplophoridae and Nematocarcinidae. Smithsonian Contributions to Zoology 432: 1–82. Crosnier, A., & Forest, J. 1973. Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19: 1–409. Griffiths, F.B., & Brandt, S.B. 1983. Distribution of mesopelagic decapod Crustacea in and around a warm-core eddy in the Tasman Sea. Marine Ecology Progress Series 12: 175–184. Hanamura, Y. 1989. Deep-sea shrimps (Crustacea: Decapoda) collected by the R.V.‘Soela’ from southern Australia. Bulletin of the Biogeographical Society of Japan 44: 51–69. Hanamura, Y., & Evans, D.R. 1994. Deepwater caridean shrimps of the families Oplophoridae and

68 Caridea – shrimps

Pasiphaeidae (Crustacea: Decapoda) from Western Australia, with an appendix on a lophogastri- dan mysid (Mysidacea). Crustacean Research 23: 46–60. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331.

Superfamily Atyoidea De Haan, 1849 There is only one family.

Atyidae De Haan, 1849 Members of the Atyidae are primarily found only in fresh water and the only species included here could only just be called ‘marine’.It is found only at the freshest end of estuaries. The dense tufts of setae at the ends of the fingers of the first two pairs of legs immediately diagnose the family. The brushes of setae are probably used to scrape food from the substrate. Choy & Horwitz (1995) provided a preliminary key to 17 species in Australia but Davie (2002) recorded 22 species. Holthuis (1993) provided keys to all genera. Diagnosis. Eyes visible. Pereopods 1 and 2 similar, chelate, with dense tufts of setae at apices of fingers. Pereopod 2 carpus entire.

Paratya Miers, 1882 The genus is represented by only one species in Australia. Diagnosis. Supraorbital spine present, without pterygostomial spine. Pereopods 1–5 with exopods. Eyes well developed. Paratya australiensis Kemp, 1917 (Fig. 15). 30 mm. Qld to eastern SA, Tas.; streams, lakes and weak penetration of estuaries, usually associated with vegetation. This is the only species of shallow-water shrimp in Australia with a supraorbital spine. Detailed studies of morphology, taxonomy (Williams, 1977; Williams & Smith, 1979; Smith & Williams, 1980; Walsh &

a b

Fig. 15. Atyidae. Paratya australiensis. a, carapace with rostrum and antennae. b, cheliped.

69 Marine Decapod Crustacea of Southern Australia

Mitchell, 1995) make this species one of the most investigated of all Australian shrimps. The tufts of setae on the first two pereopods distinguish it from all others in the near-marine environment but it does not tolerate really estuarine environments.

References Choy, S., & Horwitz, P. 1995. Preliminary key to the species of Australian shrimps (Atyidae) found in inland waters. Pp. 51–59 in: Horwitz, P. (ed.) A Preliminary Key to the Species of Decapoda (Crustacea: Malacostraca) found in Australian Inland Waters (Identification Guide No. 5). Cooperative Research Centre for Freshwater Ecology: Albury. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Smith, M.J., & Williams, W.D. 1980. Infraspecific variation with the Atyidae: a study of morphological variation within a population of Paratya australiensis (Crustacea: Decapoda). Australian Journal of Marine and Freshwater Research 31: 397–407. Walsh, C.J., & Mitchell, B.D. 1995. The freshwater shrimp Paratya australiensis (Kemp, 1917) (Decapoda: Atyidae) in estuaries of south-western Victoria, Australia. Marine and Freshwater Research 46: 959–965. Williams, W.D. 1977. Some aspects of the ecology of Paratya australiensis (Crustacea: Decapoda: Atyidae). Australian Journal of Marine and Freshwater Research 28: 403–415. Williams, W.D., & Smith, M.J. 1979. A taxonomic revision of Australian species of Paratya (Crustacea: Atyidae). Australian Journal of Marine and Freshwater Research 30: 815–832.

Superfamily Bresilioidea Calman, 1896 Bresiliidae is the only family recorded from Australia although Martin & Davis (2001) recognised Disciadidae Rathbun, 1902. These authors included three further families not yet reported from Australia. The molar process is reduced, conical or laminar.

Bresiliidae Calman, 1896 The Bresiliidae are a family of small benthic shrimps which extend to the deep sea, some such as the blind species of Alvinocaris Williams & Chace, 1982 and Rimicaris Williams & Rona, 1986 from hydrothermal vents. Most records in southern Australia are in deep water, only one being found at shallow shelf depth. Prior to 1978 most genera were placed in the Disciadidae but in that year the family was synonymised with Bresiliidae (Chace & Brown, 1978), a family which now contains about 26 species in 11 genera (Holthuis, 1993). Bresiliids are rather undistinguished shrimps which on first appearance seem like the more common palaemonids or hippolytids. They differ from hippolytids in having an undivided carpus on pereopod 2 and from palaemonids in having the first pereopods stronger and heavier than the second. The Australian bresiliid fauna is little explored, with only seven species known. The unusual species from Queensland, Pseudocheles enigma Chace & Brown, 1978 can be recognised by having five pairs of chelate pereopods. The species from southern Australia are in two genera. More species could be expected in more remote environments.

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Diagnosis. Pereopod 1 chelate, stronger and shorter than second. Pereopod 2 chelate, carpus not subdivided. Mandible with molar process conical, laminar or vestigial. Maxilliped 2 with last article attached to penultimate article. Pereopods without strap-like epipods.

Key to southern Australian species of Bresiliidae 1. Abdominal somite 3 not hump-like; pereopods 1–5 with exopods; pereopod 1 ischium and merus fused; rostrum dorsoventrally flattened ...... Discias brownae — Abdominal somite 3 hump-like; pereopods 1–2 with exopods; pereopod 1 ischium and merus free; rostrum blade-like ...... Bresilia … 2 2. Pereopod 1 palm without robust setae on lower margin; posterodorsal process of abdominal somite 3 compressed; rostrum with 9 dorsal and 1 ventral teeth ...... Bresilia antipodarum — Pereopod 1 palm with robust setae on lower margin, dactylus dorsally carinate, dentate; posterodorsal process of abdominal somite 3 not strongly compressed; rostrum with 4 dorsal and no ventral teeth ...... Bresilia plumifera

f d

Fig. 16. Bresiliidae. a, Bresilia antipodarum. b, c, Discias brownae (habitus, pereopod 1 dactylus), d, Bresilia plumifera (anterior carapace). e, Bresilia plumifera (pereopod 1 chela), f, Bresilia antipodarum (pereopod 1 chela). (a, f from Bruce, 1990a; b from Kensley, 1983; e from Bruce, 1990b)

71 Marine Decapod Crustacea of Southern Australia

Bresilia Calman, 1896 Bresilia is a genus of four species, two in the North Atlantic and two in south-eastern Australia. Each one is known only from its holotype which indicates their rarity (Bruce, 1990a, b). Diagnosis. Carapace without supraorbital and branchiostegal spines. Abdominal somite 3 hump- like. Eyestalk without spine. Pereopod 1 with normal articulation between carpus and propodus; dactylus tapering. Pereopod 1 ischium and merus free. Pereopods 1–2 only with exopods. Pereopod 5 with reduced pleurobranch. Arthrobranchs present. Telson with 3 or more pairs of dorsolateral spiniform setae. Bresilia antipodarum Bruce, 1990 (Fig. 16a, f). Pereopod 1 with dactylus tapering. Posterodorsal process of abdominal somite 3 compressed. Rostrum with 9 dorsal and 1 ventral teeth. tl. 14.5 mm. Off Tas.; 800 m depth. Bresilia plumifera Bruce, 1990 (Fig. 16d, e). Pereopod 1 with dactylus dorsally carinate. Posterodorsal process of abdominal somite 3 not strongly compressed. Rostrum with 4 dorsal and 0 ventral teeth. tl. 6 mm. Taupo Seamount, off NSW; 133 m depth.

Discias Rathbun, 1902 Discias is a genus of seven species, one in south-eastern Australia (Kensley, 1983). The small rostrum and the unusual semicircular or disc-like dactylus on pereopod 1 are characteristic. The species are thought to be associates of sponges, usually in shallow water and the finely toothed margin of the dactylus and the slot in the fixed finger may be for scraping mucus from the host. The widespread D. atlanticus Gurney, 1939 and D. exul Kemp, 1920 occur on the Great Barrier Reef (Kensley, 1983). Diagnosis. Carapace without supraorbital and branchiostegal spines. Abdominal somite 3 not hump-like. Eyestalk without spine. Pereopod 1 with a lobe of propodus proximal to articulation with carpus; dactylus semicircular. Pereopod 1 ischium and merus fused. Pereopods 1–5 with exopods. Pereopod 5 with reduced pleurobranch. Arthrobranchs present. Telson with 3 or fewer pairs of dorsolateral spiniform setae. Discias brownae Kensley, 1983 (Fig. 16b, c). Uropod with outer ramus with entire lateral margin. Mandibular palp of 2 articles. Rostrum acute, with lateral margins entire. Telson with 3 apical spines. tl. 4 mm. NSW (Sydney region); 9–33 depth, from sponges. D. brownae has a small flattened rostrum and might be confused with the pasiphaeid Leptochela sydniensis,a common shelf shrimp with a short pointed rostrum.

References Bruce, A.J. 1990a. Two deep-sea shrimps new to the Australian fauna, Psathyrocaris hawaiiensis Rathbun (Pasiphaeidae) and Bresilia antipodarum, sp. nov. (Bresiliidae), with remarks on Encantada spinoc- ulata Wicksten (Bresiliidae). Invertebrate Taxonomy 4: 847–866. Bruce, A.J. 1990b. A second species of Bresilia, B. plumifera sp. nov., new to the Australian fauna (Crustacea: Decapoda: Bresiliidae). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 7: 1–8. Chace, F.A., & Brown, D.E. 1978. A new polychelate shrimp from the Great Barrier Reef of Australia and its bearing on the family Bresiliidae (Crustacea: Decapoda: Caridea). Proceedings of the Biological Society of Washington 91: 756–766. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Kensley, B. 1983. New records of bresiliid shrimp from Australia, South Africa, Caribbean, and Gulf of Mexico (Decapaoda: Natantia: Caridea). Smithsonian Contributions to Zoology 394: 1–31.

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Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124.

Superfamily Nematocarcinoidea Smith, 1884 Holthuis (1993) included four families, all with strap-like epipods on at least the first three pairs of pereopods and with a blunt molar process. Nematocarcinidae and Rhynchocinetidae occur in southern Australia. Xiphocarididae von Martens, 1872 are not known from Australia and Eugonatonotidae Chace, 1937 only from tropical seas.

Nematocarcinidae Smith, 1884 The Nematocarcinidae are a family of large deep-water demersal shrimps recognisable by the very fine rostrum and delicate long legs (Chace, 1986; Holthuis, 1993). About 25 species are known, many reportedly with wide distributions. The southern Australian species are poorly known and not all material in museum collections was able to be unambiguously identified. Diagnosis. Carapace without longitudinal ridges and spines. Rostrum thin, with fine teeth dorsally (often articulating) and sometimes ventrally. Eyes visible. Pereopod 1 chelate, stronger than pereopod 2, but both very delicate. Pereopod 2 carpus not divided. Pereopods 1–4 with simple epipods and exopods.

Key to southern Australian species of Nematocarcinidae 1. Pereopods 3–5 moderately long (1.5 times length of pereopod 1), carpus shorter than propodus; articulation between ischium and merus not thickened; rostrum upturned and with dorsal spines only basally ...... Lipkius holthuisi — Pereopods 3–5 extremely long (twice length of pereopod 1), carpus several times as long as propodus; articulation between ischium and merus thickened; rostrum straight and with dorsal spines to apex ...... Nematocarcinus … 2 2. Rostrum much longer than antennule peduncle ...... 4 — Rostrum shorter than antennule peduncle ...... 3 3. Rostrum with 12–22 evenly spaced dorsal teeth ...... Nematocarcinus gracilis — Rostrum with 7–14 dorsal teeth, more widely spaced anteriorly . . Nematocarcinus undulatipes 4. Rostrum with ventral teeth ...... Nematocarcinus sigmoideus — Rostrum without ventral teeth ...... Nematocarcinus sp.

Lipkius Yaldwyn, 1960 The genus is monotypic. Diagnosis. Rostrum upturned and with dorsal spines only basally. Pereopods 3–5 moderately long (1.5 times length of pereopod 1); carpus shorter than propodus; articulation between ischium and merus not thickened. Telson as long as uropods. Lipkius holthuisi Yaldwyn, 1960 (Fig. 17a). 200 mm. Partly red. New Zealand, NSW, Tas.; 400–1700 m depth. The upturned rostrum and the spine on the posterior margin of abdominal pleuron 5 distinguish the species from other deep-sea shrimps. The species is taken in fish trawls in New Zealand, often with species of Nematocarcinus (Webber et al., 1990).

73 Marine Decapod Crustacea of Southern Australia

c b

Fig. 17. Nematocarcinidae. a, Lipkius holthuisi. Anterior region of carapace, eye and basis of antenna: b, Nematocarcinus gracilis. c, Nematocarcinus undulatipes. d, Nematocarcinus sigmoideus. e, Nematocarcinus sp.

74 Caridea – shrimps

Nematocarcinus Milne Edwards, 1881 Several species have been described from deep waters north and north-west of Australia (Chace, 1986; Hanamura & Evans, 1996; Burukovsky, 2000) but the material from southern Australia reconciles with these only imperfectly. Kensley et al. (1987) reported two species off NSW and Hanamura (1989) another off Tas. All are about 170 mm long maximum and coloured partly red in life. Diagnosis. Rostrum straight and with dorsal spines to apex. Pereopods 3–5 extremely long (twice length of pereopod 1); carpus several times as long as propodus; articulation between ischium and merus thickened. Telson as long as uropods. Nematocarcinus gracilis Bate, 1888 (Fig. 17b). Rostrum shorter than antennal 1 peduncle, with 12–22 evenly spaced dorsal spines, and 1 distal ventral tooth. Indo-West Pacific, NSW; 165–1170 m depth. Nematocarcinus sigmoideus Macpherson, 1984 (Fig. 17c). Rostrum much longer than antennal 1 peduncle, with >26 dorsal spines, 5–8 postorbital, more spaced distally; 4–9 ventral spines. Southern Atlantic, southern Africa, NSW, Tas.; slope depths. Nematocarcinus undulatipes Bate, 1888 (Fig. 17d). Rostrum shorter than antennal 1 peduncle, with 7–14 dorsal teeth, more spaced anteriorly, and 1 distal ventral tooth. Indo-West Pacific, NSW; 366–1269 m depth. Nematocarcinus sp. (Fig. 17e). Rostrum much longer than antennal 1 peduncle, with 30 dorsal teeth, more obsolete distally; without ventral teeth. NSW, Tas.; slope depths.

References Burukovsky, R.N. 2000. Taxonomy of shrimps of the genus Nematocarcinus (Crustacea, Decapod, Nematocarcinidae). 7. Description of new species, N. hanamuri and N. evansi, from southwestern Australian waters. Zoologicheskii Zhurnal, Moscow 79: 1290–1293. Chace, F.A. 1986. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 4: families Oplophoridae and Nematocarcinidae. Smithsonian Contributions to Zoology 432: 1–82. Hanamura, Y. 1989. Deep-sea shrimps (Crustacea: Decapoda) collected by the R.V.‘Soela’ from southern Australia. Bulletin of the Biogeographical Society of Japan 44: 51–69. Hanamura, Y., & Evans, D.R. 1996. Deepwater caridean shrimps of the families Nematocarcinidae, Stylodactylidae, Pandalidae and Crangonidae (Crustacea: Decapoda) from Western Australia. Bulletin of Nansei National Fisheries Research Institute 29: 1–18. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Webber, W.R., Fenaughty, C.M., & Clark, M.R. 1990. A guide to some common offshore shrimp and prawn species of New Zealand. New Zealand Fisheries Occasional Publication 6: 1–42.

Rhynchocinetidae Ortmann, 1890 The family, the hinge-beaked shrimps, is easily distinguished from all others by the movable rostrum. All have fine curved striations on the carapace and abdomen, clearly visible only with oblique lighting. The family comprises only two genera totalling more than 20 species (Holthuis, 1995; Okuno, 1996, 1997; Chace, 1997; Okuno & Hoover, 1998).

75 Marine Decapod Crustacea of Southern Australia

Diagnosis. Carapace without longitudinal ridges and spines. Rostrum vertically hinged, prominent, with dorsal and ventral teeth. Eyes visible. Pereopod 1 chelate, stronger than pereopod 2. Pereopod 2 carpus not divided. Pereopods with some simple epipods, without exopods. Abdomen with bend at somite 3.

Rhynchocinetes Milne Edwards, 1837 Okuno (1994) provided a key to the Australian species and later (1997) added another. Five species occur in southern waters. The species are very similar and geography or colour can help in identification. The arthrobranchs (mentioned in the key) are small triangular gills between the lower ends of the larger oblique pleurobranchs. When present, the arthrobranch on pereopod 3 is between the third- and fourth-most posterior pleurobranchs. Diagnosis. Carapace covered with fine transverse striae, armed with 2 spines on median carina, anterior spine just behind rostral articulation. Supraorbital and antennal spine present. Abdominal somites 4 and 5 without a tooth above base of pleuron.

Key to southern Australian species of Rhynchocinetes 1. Arthrobranchs absent from maxillipeds and all pereopods ...... Rhynchocinetes enigma — Arthrobranchs present on at least maxillipeds and pereopod 1 ...... 2 2. Pereopod 2 without arthrobranch ...... Rhynchocinetes balssi — Pereopod 2 with arthrobranch ...... 3 3. Pereopod 3 with arthrobranch ...... Rhynchocinetes serratus — Pereopod 3 without arthrobranch ...... 4 4. Stylocerite reaches beyond distolateral angle of article 1 of antenna 1; endopod of male pleopod 1 without lobe on lateral margin ...... Rhynchocinetes australis — Stylocerite not reaching end of distolateral angle of article 1 of antenna 1; endopod of male pleopod 1 with lobe on lateral margin; ...... Rhynchocinetes kuiteri Rhynchocinetes australis Hale, 1941 (Fig. 18a, Pl. 7c). Rostrum with 2 spaced dorsal teeth, up to 4 apical dorsal teeth, and up to 12 curved ventral teeth. Stylocerite reaches beyond distolateral angle of article 1 of antenna 1. Arthrobranchs absent from pereopods 3–5. Endopod of male pleopod 1 without lobe on lateral margin. Carapace with irregular longitudinal reddish stripes and white dots; antennal bases bright yellow; abdominal somites 1 and 2 each with transverse red stripe; abdominal somites 3–6 with longitudinal lateral white and red stripes. 60 mm. Tas., Vic., SA, WA; subtidal to 245 m depth, reefs and algal assemblages. This species is far more common than R. kuiteri, being taken from crevices and in subtidal algal collections. It is one of the more brightly coloured shrimps. The life photographs published by Tiefenbacher (1983) and Edgar (1997) and labelled R. kuiteri show the same colours as specimens identified by me as R. australis. Rhynchocinetes balssi Gordon, 1936. Dorsal rostral margin armed with 2 proximal and 4 subterminal teeth, 11 ventral. Stylocerite reaches beyond distolateral angle of article 1 of antenna 1. Arthrobranchs absent from pereopods 2–5. Endopod of male pleopod 1 without lobe on lateral margin. Colour uncertain. Total length to 35 mm. Southern WA; 160–190 m depth. Rhynchocinetes enigma Okuno, 1997 (Fig. 18b). Dorsal rostral margin armed with 2 teeth in basal half and 3 subterminal teeth; 11 ventral teeth. Stylocerite reaches beyond distolateral angle of article 1 of antenna 1. Arthrobranchs absent from maxillipeds and all pereopods. Endopod of male pleopod 1 with lobe on lateral margin. Colour uncertain. cl. 9.0 mm. Western SA; 113–170 m (known from 3 specimens collected from the Great Australian Bight – Okuno, 1997).

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b c d

Fig. 18. Rhynchocinetidae. a, Rhynchocinetes australis. b, Rhynchocinetes enigma (carapace). c, Rhynchocinetes australis (pleopod 1 endopod). d, Rhynchocinetes serratus (pleopod 1 endopod).

Rhynchocinetes kuiteri Tiefenbacher, 1983. Rostrum with 2 spaced dorsal teeth, up to 6 apical dorsal teeth, and up to 14 curved ventral teeth. Stylocerite not reaching end of distolateral angle of article 1 of antenna 1. Arthrobranchs absent from pereopods 3–5. Endopod of male pleopod 1 with lobe on lateral margin. Colour uncertain. 37 mm. Vic.; subtidal, rocky reef. The species is known for certain only from four specimens taken off Portsea, Vic. The life photographs supposedly of this species in Tiefenbacher (1983) and Edgar (1997) are almost certainly of R. australis. Rhynchocinetes serratus (Milne Edwards, 1837) (Fig. 18d, Pl. 7d). Rostrum with 2 spaced dorsal teeth, up to 6 apical dorsal teeth, and up to 13 curved ventral teeth. Stylocerite reaches beyond distolateral angle of article 1 of antenna 1. Arthrobranchs absent from pereopods 4–5. Endopod of male pleopod 1 with lobe on lateral margin. Bluish; bright blue, yellow and red stripes and dots on a darker blue background. 60 mm. NSW (central and southern coasts), WA (south-western coast); intertidal and subtidal reefs. This species was known as R. rugulosus until its name was corrected by Holthuis (1995). Although widely reported from the western Pacific, the species may be more confined (Okuno, 1994).

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References Chace, F.A. 1997. The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae, Bathypalaemonellidae, Processidae, and Hippolytidae. Smithsonian Contributions to Zoology 587: i-v, 1–106. Edgar, G.J. 1997. Australian marine life – the plants and animals of temperate waters. Reed Books: Kew Victoria. 544 pp. Holthuis, L.B. 1995. Notes on Indo-West Pacific Crustacea Decapoda III to IX. Zoologische Mededelingen, Leiden 69: 139–151. Okuno, J. 1994. A new species of hinge-beak shrimp from the Western Pacific (Crustacea, Decapoda, Rhynchocinetidae). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11: 29–37. Okuno, J. 1996. Cinetorhynchus manningi, a new shrimp (Crustacea: Decapoda: Caridea: Rhynchocinetidae) from the western Atlantic. Proceedings of the Biological Society of Washington 109: 725–730. Okuno, J. 1997. A new shrimp of the genus Rhynchocinetes from the Great Australian Bight (Crustacea: Decapoda: Rhynchocinetidae). Records of the South Australian Museum 30: 13–18. Okuno, J., & Hoover, J.P. 1998. Cinetorhynchus hawaiiensis, a new shrimp forming a cryptic species pair with C. reticulatus Okuno, 1997, and new records of three congeneric species (Crustacea: Decapoda: Rhynchocinetidae). Natural History Research 5: 31–42. Tiefenbacher, L. 1983. A new species of Rhynchocinetes from south-Australia (Crustacea, Decapoda, Rhynchocinetidae). Revue Française d’Aquariologie 9: 121–124.

Superfamily Stylodactyloidea Bate, 1888 Stylodactylidae is the only family.

Stylodactylidae Bate, 1888 The Stylodactylidae are shrimps of the deep shelf or continental slope recognisable by their elongate similar setose first two pairs of pereopods. The family comprises about 24 species in five genera (Holthuis, 1993; Cleva, 1994). All genera are represented in tropical Australian waters, by nine species (Cleva, 1994; Hanamura & Evans, 1996). Only one has been reported from southern Australia but others occur as far south as northern NSW and could be expected in our region (Kensley et al., 1987). Diagnosis. Rostrum about as long or longer than carapace. Antennule flagella undivided. Maxilliped 2 with last 2 articles arising side by side from preceding article. Maxilliped 3 and pereopods 1 and 2 extremely long, without exopods, fringed with long setae. Pereopods 1 and 2 similar, with long slender fingers; carpus of 2 not subdivided.

Stylodactylus Milne Edwards, 1881 Chace (1983) and Cleva (1990) provided keys to species and diagnosed many in detail. Diagnosis. Mandibular palp of 2 articles. Stylocerite at least 4 times as long as wide; tip acute. Pereopods 3–5 without spine on distal lobe of carpus. Stylodactylus multidentatus Kubo, 1942 (Fig. 19). Abdominal somites without posterodorsal spines. Abdominal pleura without posteroventral spines. Rostrum with 14–23 ventral spines.

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Antennule peduncle article 2 shorter than article 1. Supraorbital spine well-developed, 1 or 2 secondary spines on carapace margin ventral to branchiostegal spine. Telson without posteromedial spine. 20 mm. West Pacific south to NSW; 152–412 m depth.

Fig. 19. Stylodactylidae. a, Stylodactylus multidentatus (without pereopods). b, c, Stylodactylus multidentatus (maxilliped 3, pereopod 1).

References Chace, F.A. 1983. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 1: family Stylodactylidae. Smithsonian Contributions to Zoology 381: 1–21. Cleva, R. 1990. Crustacea Decapoda: Les genres et les espèces indo-ouest pacifiques de Stylodactylidae. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145: 71–136. Cleva, R. 1994. Some Australian Stylodactylidae (Crustacea: Decapoda), with descriptions of two new species. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11: 53–64. Hanamura, Y., & Evans, D.R. 1996. Deepwater caridean shrimps of the families Nematocarcinidae, Stylodactylidae, Pandalidae and Crangonidae (Crustacea: Decapoda) from Western Australia. Bulletin of Nansei National Fisheries Research Institute 29: 1–18. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331.

79 Marine Decapod Crustacea of Southern Australia

Superfamily Campylonotoidea Sollaud, 1913 Campylonotidae is one of two families in the superfamily. The other, Bathypalaemonellidae de Saint Laurent, 1985 is represented in Australia by two tropical species (Bruce, 1986).

Campylonotidae Sollaud, 1913 The Campylonotidae comprise only a single genus of five species of midwater shrimps. The sculpture of the abdomen immediately identifies the only Australian species. Diagnosis. Pereopod 1 smaller than pereopod 2. Pereopods 2 subequal, carpus not subdivided. Pereopods 1–4 each with arthrobranch. Pereopods with epipods, without exopods. Maxilliped 2 with terminal article applied as a narrow strip to the mesial margin of the second last article. Dorsal antennular flagellum simple.

Campylonotus Bate, 1888 Diagnosis. As for family. Campylonotus rathbunae Schmitt, 1926 (Fig. 20). Rostrum narrow with 2 large basal teeth, ventral margin with 3–4 teeth. Carapace with 2 strong, lateral carinae. Abdomen armed with posterior dorsal spines on somites 3–5, and prominent tubercle on anterior part of third somite. 150 mm. New Zealand, NSW, SA, WA (Great Australian Bight); 270–810 m depth; mesopelagic. This species is distinguished by the lateral carinae on the carapace and the prominent tubercle on the third abdominal somite. Local records were summarised by Kensley et al. (1987) and Webber et al. (1990).

Fig. 20. Campylonotidae. Campylonotus rathbunae.

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References Bruce, A.J. 1986. Two new species of Bathypalaemonella Balss (Crustacea, Decapoda, Campylonotidae) from the Australian Northwest Shelf. Zoologica Scripta 15: 251–264. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Webber, W.R., Fenaughty, C.M., & Clark, M.R. 1990. A guide to some common offshore shrimp and prawn species of New Zealand. New Zealand Fisheries Occasional Publication 6: 1–42.

Superfamily Palaemonoidea Rafinesque, 1815 The Palaemonoidea is the largest of the caridean superfamilies with eight families according to Martin & Davis (2001). Only two occur in southern Australia. Kakaducarididae Bruce, 1993 is a tropical Australian freshwater family, while Anchistioididae Borradaile, 1915 (Bruce, 1981) and Hymenoceridae Ortmann, 1890 (Bruce, 1988b; Davie, 1992) are represented in northern Australian marine waters.

Gnathophyllidae Dana, 1852 The Gnathophyllidae are a small family of mainly tropical shrimps similar in many ways to pontoniine palaemonids. Bruce (1986) thought, partly on the similarity of larval stages, that the two groups should be synonymised but his view is not supported in recent classifications. About a dozen species are allocated to four genera (Chace, 1993; Holthuis, 1993). Only four species in two genera have been reported from Australia, the ranges of three just reaching southern coasts. Diagnosis. Antenna 1 with dorsal flagellum of 2 completely separate branches. Mandible without incisor (or incisor rudimentary). Maxilliped 2 with last 2 articles in sequence. Maxilliped 3 leaf- like, sometimes operculate, with ischiomerus slightly broader than and fused to basis. Pereopods without arthrobranchs. Pereopod 1 always with strong chela. Pereopods 1 usually more slender (rarely subequal) than pereopods 2. Carpus of pereopod 2 entire, not subdivided.

Key to southern Australian species of Gnathophyllidae 1. Pereopods 3–5 dactyls with bifid apex; with transverse colour bands . . Gnathophyllum … 2 — Pereopods 3–5 dactyls with simple apex; with longitudinal colour bands ...... Gnathophylloides … 3 2. Body dark with close dark bands; pereopods banded; 3–6 dorsal rostral teeth, most proximal arising behind anterior to orbital margin ...... Gnathophyllum americanum — Body pale dull yellow with narrow, widely spaced dark bands; pereopods without banding but with white speckling; 5 or 6 dorsal rostral teeth, most proximal arising behind posterior to orbital margin ...... Gnathophyllum taylori 3. Maxilliped 3 with antepenultimate article broad, widest centrally, exopod longer than endopod; rostrum concave, short, not overreaching eyes, with 4 dorsal teeth ...... Gnathophylloides mineri — Maxilliped 3 with antepenultimate article narrow, exopod shorter than endopod; rostrum straight, long, overreaching eyes, with 3 dorsal teeth ...... Gnathophylloides robustus

81 Marine Decapod Crustacea of Southern Australia

c f e

Fig. 21. Gnathophyllidae. a, Gnathophyllum americanum. b, Gnathophyllum taylori. Anterior carapace and rostrum: c, Gnathophylloides mineri. d, Gnathophylloides robustus. e, Gnathophyllum americanum. Maxilliped 3: f, Gnathophylloides mineri, g, Gnathophylloides robustus. h, Gnathophyllum americanum.

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Gnathophylloides Schmitt, 1933 Gnathophylloides is a genus of only two species, both in Australia, but its presence in southern Australia is doubtful. Both are associates of echinoids. Diagnosis. Pereopods 3–5 dactyls with simple apex, with rounded lobe on lower margin. Maxilliped 2 not reaching beyond 3. Maxilliped 3 about 4 times as long as wide, barely operculate. Gnathophylloides mineri Schmitt, 1933 (Fig. 21c). Rostrum depressed, concave dorsally, not exceeding eyes and with 4 dorsal teeth and small distal ventral tooth. Maxilliped 3 with antepenultimate article broad, widest centrally, exopod longer than endopod. Carapace length to 2.5 mm. Caribbean, tropical Pacific islands, Indian Ocean; Solitary Islands, NSW; subtidal associated with echinoids. There is only a single record of this tiny tropical shrimp in northern NSW (Bruce, 1988a) and it is unlikely to occur on truly southern coasts. The shrimp is coloured with a broad dorsal band of narrow white and pale brown stripes, a broader dorsolateral white band, lateral brown band along whole body, and a lower band of white. Its hosts in Australia are the echinoids Tripneustes gratilla and Pseudoboletia maculata. In the Caribbean, the shrimp lives in large groups, is cryptically coloured, and feeds on the epithe- lium of its host (Patton et al., 1985). Gnathophylloides robustus Bruce, 1973 (Fig. 21g). Rostrum straight, twice as long as the eyestalks, with 3 small acute dorsal teeth and unarmed ventrally. Maxilliped 3 with antepenultimate article narrow, exopod shorter than endopod. Carapace length to 6.2 mm. Off Point Moore, Geraldton, WA; 3 m depth (Bruce, 1973).

Gnathophyllum Latreille, 1819 Gnathophyllum is a genus of cryptic free-living or echinoid-associating shrimps. Nine species are known (Manning, 1963; Bruce, 1986; Chace, 1993; Ahyong, 2003). Diagnosis. Pereopods 3–5 dactyls with bifid apex, without rounded lobe on lower margin. Maxilliped 2 not reaching beyond 3. Maxilliped 3 about twice as long as wide, operculate. Gnathophyllum americanum Guérin-Méneville, 1856 (Fig. 21a, e, h, Pl. 7e). Rostrum, with 3–6 dorsal evenly-spaced teeth, the most proximal anterior to posterior orbital margin, and minute distal ventral tooth. Orbit with narrow posterodorsal sinus. Dark brown with numerous transverse white or cream bands, 6 on the carapace and 1–3 on each abdominal somite; gold bands on legs and near telson. cl. 4.4 mm. Tropical Atlantic, Caribbean, Indo-West Pacific, WA (S to Abrolhos Is.), NT, Qld, NSW (S to Sydney); free-living or associated with echinoids. The species was figured in colour by Healy & Yaldwyn (1970). Gnathophyllum taylori Ahyong, 2003 (Fig. 21b, Pl. 7f). Rostrum with 5 or 6 dorsal teeth and minute subdistal ventral denticle; proximal 1 or 2 teeth behind level of posterior orbital margin. Orbit without narrow posterodorsal sinus. Body pale dull yellow with narrow, widely spaced dark bands; pereopods without banding but with white speckling; sixth abdominal tergite and tail-fan transparent. cl. 5.7 mm. Elizabeth Reef, NSW (Central region S to Sydney); shallow subtidal associated with the echinoid, Centrostephanus rogersi. The species has been referred to as a colour variant of G. americanum under which name a photograph was published (Debelius, 1999).

References Ahyong, S.T. 2003. Gnathophyllum taylori, a new species of caridean shrimp from southeastern Australia (Crustacea: Decapoda: Gnathophyllidae). Memoirs of Museum Victoria 60: 237–242.

83 Marine Decapod Crustacea of Southern Australia

Bruce, A.J. 1973. Gnathophylloides robustus sp. nov., a new commensal gnathophyllid shrimp from Western Australia, with the designation of a new genus Levicaris (Decapoda, Caridea). Crustaceana 24: 17–32. Bruce, A.J. 1981. Pontoniine shrimps of Heron Island. Atoll Research Bulletin 245: 1–33. Bruce, A.J. 1986. Observations of the family Gnathophyllidae Dana, 1852 (Crustacea: Decapoda). Journal of Crustacean Biology 6: 463–470. Bruce, A.J. 1988a. A note on Gnathophylloides mineri Schmitt (Crustacea: Decapoda: Palaemonidae), including its first occurrence in Australian waters. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 5: 97–100. Bruce, A.J. 1988b. Further records of the hymenocerine shrimp, Phyllognathia simplex Fujino (Crustacea: Decapoda). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 5: 101–104. Chace, F.A. 1993. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 6: superfamily Palaemonoidea. Smithsonian Contributions to Zoology 543: 1–152. Davie, P.J.F. 1992. The rare shrimp, Phyllognathia ceratophthalma (Balss), discovered in Moreton Bay. Memoirs of the Queensland Museum 32: 98, colour fig. Debelius, H. 1999. Crustacea. Guide of the world. Ikan -Unterwasserarchiv: Frankfurt. 321 pp. Healy, A., & Yaldwyn, J.C. 1970. Australian Crustaceans in Colour. Reed: Sydney. 112 pp. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Manning, R.B. 1963. The east American species of Gnathophyllum (Decapoda, Caridea), with description of a new species. Crustaceana 5: 47–63. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. Patton, W.K., Patton, R.J., & Barnes, A. 1985. On the biology of Gnathophylloides mineri, a shrimp inhabiting the sea urchin Tripneustes ventricosus. Journal of Crustacean Biology 5: 616–626. Schmitt, W.L. 1933. Four new species of decapod crustaceans from Puerto Rico. American Museum Novitates 662: 1–9, figs. 1–4.

Palaemonidae Rafinesque, 1815 The Palaemonidae are a cosmopolitan family of many species and occur in most environments, including fresh water. The species range in size from among the smallest of shrimps, only a few millimetres long to examples like the edible tropical freshwater species, Macrobrachium rosenbergii, which attains 400 mm with its long chelipeds extended. The family is divided into four subfamilies of which two occur in Australia, Palaemoninae and Pontoniinae, but the distinction between them is not as clear as most keys make out (Chace, 1993; Bruce, 1994). Bruce (1994) pointed out that, although it has long been known that all palaemonids lack any pleurobranch associated with maxilliped 3, palaemonines alone are often wrongly reported to possess one. The number of posterior paired robust setae on the telson is the only reliable character separating the subfamilies, two pairs in Palaemoninae and three pairs in Pontoniinae. These robust setae have traditionally been referred to as ‘spines’ in earlier literature, a term I restrict to sharp nonarticulating cuticular projections. A third subfamily was suggested but immediately elevated to family rank; the Kakaducarididae include a single freshwater species from the Northern Territory (Bruce, 1993). In temperate waters of southern Australia palaemonine genera dominate and are among the most commonly seen shrimps in shallow water and rock pools. Pontoniines are diverse only in the tropics and few representatives are seen in southern Australia. The key to pontoniines below should not be relied on to identify species; all those so far discovered are rare and more could be anticipated at places like Rottnest I.

84 Caridea – shrimps

In most recent literature, definition of the palaemonine genera relies upon differentiating between a branchiostegal spine and a hepatic spine, both of which lie further down the front part of the carapace than the antennal spine which arises near the margin between the eyestalk and the antenna. Developmental studies have shown that the two are homologous and that the spine migrates as the shrimp grows. The spine is clearly on the hepatic region in Macrobrachium but branchiostegal or close to the branchiostegite in Palaemon. The branchiostegal groove, if present, passes to the front of or over the spine but is difficult to see until the light is angled correctly to show a slight depression.

Diagnosis. Antennule with dorsal flagellum of 2 completely separate branches. Maxilliped 2 with last 2 articles in sequence. Pereopods without arthrobranchs. Pereopod 1 always with strong chela. Pereopods 1 usually more slender (rarely subequal) than pereopods 2. Carpus of pereopod 2 entire, not subdivided.

Key to species of Palaemonidae from southern Australia 1. Telson with 2 pairs of posterior robust setae; 1 or 2 gills present on maxilliped 3 ...... Palaemoninae … 2 — Telson with 3 or more pairs of posterior robust setae; gills absent from maxilliped 3 ...... Pontoniinae … 10 2. Carapace spine (hepatic) well removed from anterior edge of carapace by a distance greater than its length; branchiostegal groove running from carapace edge to base of spine; pereopods 2 spinulose, larger in male; male larger than female ...... Macrobrachium novaehollandiae — Carapace spine (branchiostegal) on or overlapping carapace edge or set back; branchiostegal groove running back from carapace edge in a shallow arc up to or over spine; pereopods 2 smooth, same size in both sexes; female larger than male ...... 3 3. Carapace spine set back from carapace edge by less than its length and situated between branchiostegite and ridge of antennal spine; branchiostegal groove running up to dorsoposterior edge of spine but not past it; rostrum with 7–10 dorsal teeth, usually 8, 2–3 postorbital, proximal 4 incompletely articulated; 4–6, usually 5, ventral teeth ...... Palaemon intermedius — Carapace spine set below branchiostegal groove which may deviate up and over spine; spine on or set back from edge of carapace; rostrum not as above ...... 4 4. Carapace spine set back from the carapace edge, overlapping it at most only with tip . . . . 5 — Carapace spine at or very near carapace edge just under start of branchiostegal groove; groove running back from carapace edge in a shallow arc, not upwards sharply and down again ...... 6 5. Carapace spine almost or just reaching carapace edge with tip only; set above level of starting point of branchiostegal groove which runs upwards to front of spine, loops sharply over it and sweeps down again posteriorly; pereopod 2 carpus 1–1.5 times as long as merus; mandibular palp with 3 articles ...... Palaemon dolospina — Carapace spine set well back from the carapace edge which it never overlaps; branchiostegal groove deviating only slightly as it passes over spine; pereopod 2 carpus 1.3–1.5 times as long as merus; mandibular palp absent ...... Palaemonetes australis

85 Marine Decapod Crustacea of Southern Australia

6. Antennule inner flagellum fused for greater than half of its length, usually over about two thirds, 16–20 articles fused in adults; mandibular palp absent ...... Palaemonetes atrinubes — Antennule inner flagellum fused for half or usually less of its length, less than 15 articles fused in adults ...... 7 7. Pereopods slender (carpus of pereopod 2 longer than chela, longer than merus) ...... Palaemon cf. debilis — Pereopods not unusually slender (carpus of pereopod 2 shorter than chela, about as long as merus ...... 8 8. Pereopod 2 stout but short, 0.4–0.5 times body length ...... Palaemon litoreus — Pereopod 2 stout but long, 0.7–0.9 times body length ...... 9 9. Rostrum with 9–15 dorsal teeth (usually 10–12), all incompletely articulated; teeth directed upwards; only one-sixth of the antennule inner flagellum fused (4–6 articles) ...... Palaemon macrodactylus — Rostrum with 6–9 dorsal teeth, proximal 3 incompletely articulated; about one-third of antennule inner flagellum fused (about 12 articles in adults) ...... Palaemon serenus 10. Mandibular palp present, of 2 articles ...... 11 — Mandibular palp absent ...... 12 11. Pereopod dactyli ventrally smooth and concave; propodal spines about half propodus width ...... Palaemonella rotumana — Pereopod dactyli ventrally biconcave; propodal spines less than half propodus width ...... Palaemonella foresti 12. Hepatic spine absent; rostrum without dorsal teeth ...... 13 — Hepatic spine present; rostrum with dorsal teeth ...... Periclimenes … 15 13. Rostrum laterally compressed in distal half ...... Anchistus custos — Rostrum depressed and with a ventral keel ...... 14 14. Rostrum not dorsally concave; pereopod 1 with compressed non-spatulate fingers ...... Pseudopontonia minuta — Rostrum dorsally concave; pereopod 1 with elongate spatulate fingers ...... Notopontonia platycheles 15. Abdominal somite 3 with a strong dorsal lobe produced posteriorly; rostrum with 9–11 dorsal teeth, 3–4 on carapace proper; with 2–4 ventral teeth confined to distal quarter ...... Periclimenes aesopius — Abdominal somite 3 without a dorsal lobe; rostrum with 8–11 dorsal teeth, 2 on carapace proper; with 2–4 ventral teeth in distal half ...... 16 16. Rostrum a deep blade, 4 times as long as deep; eye with ocellus . . . . . Periclimenes yaldwyni — Rostrum tapering, 6 times as long as deep; eye without ocellus ...... 17 17. Rostrum slightly depressed, dorsally straight; slope depths, possibly with gorgonians ...... Periclimenes laccadivensis — Rostrum straight, dorsally slightly convex; subtidal, with crinoids . . Periclimenes carinidactylus

Palaemoninae Rafinesque, 1815 Some of the most common shallow water shrimps of southern Australia belong to the Palaemoninae and are characterised by a strong saw-toothed rostrum and long slender first chelipeds. The species are free-living usually preferring algal or seagrass habitats. Palaemonids

86 Caridea – shrimps

from these habitats are probably most easily confused with hippolytids but lack the divided carpus on pereopod 2 seen in the latter family. Chace (1993) provided useful keys to species of several of the 17 genera and listed all the species of others. Five genera occur in the southern Australian fauna. A sixth, Leandrites Holthuis, is represented in Australia only by L. celebensis de Man in the Northern Territory (Bruce, 1987).

Diagnosis. Maxilliped 3 usually with 2 arthrobranchs. Telson with 2 pairs of dorsal robust setae, 2 (rarely 3) pairs of posterior robust setae and with 1 or more pairs of hairs.

Fig. 22. Palaemonidae Palaemoninae. a, Palaemon serenus. b, Palaemon dolospina.

87 Marine Decapod Crustacea of Southern Australia

Leander Desmarest, 1849 Only three species are known (Chace, 1993). Hale (1927) illustrated three palaemonid species under the generic name Leander, from which they have all been now removed to other genera. Diagnosis. Hepatic spine absent. Branchiostegal spine present, submarginal. Branchiostegal groove absent. Mandibular palp 2-articled. Pereopods 3–5 with dactylus simple, shorter than propodus. Pereopod 5 propodus without transverse rows of setae in distal part of the posterior margin. Male pleopod 1 with appendix interna. Leander tenuicornis (Say, 1818) (Fig. 23a, b). Rostrum sexually dimorphic, with 2–9 dorsal and 6–7 ventral teeth; vertically deeper in female. Antennule with basal article straight or concave lateral to second article; stylocerite reaching almost to second article. Abdominal pleura 4 and 5 dentate posteroventrally. cl. 8 mm. Temperate and tropical Atlantic, Indian and western Pacific oceans, New Zealand, possibly throughout Australia; often associated with free float- ing Sargassum. Bruce & Cropp (1984) recorded this species from Australia but their source is uncertain.

a b c

d ef

gh i

Fig. 23. Palaemonidae Palaemoninae. a, b, Leander tenuicornis (female and male). c, Palaemon intermedius.d, Macrobrachium novaehollandiae.e, Palaemon cf. debilis.f, Palaemon litoreus.g, Palaemon macrodactylus.h, Palaemon serenus. i, Palaemon dolospina.j, Palaemonetes atrinubes.k, Palaemonetes australis.

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Macrobrachium Bate, 1868 Of the more than 175 described species and subspecies worldwide (Chace, 1993), few occur in Australia (Fincham, 1987). Most are from fresh water and from tropical regions and can be identified using the paper by Horwitz (1995). Macrobrachium danae (Heller, 1865), reported from Sydney, is a name not now used and the species to which it refers may be M. australe (Guérin- Méneville, 1838) but this is uncertain. One common southern Australian species from intertidal and very shallow shelf environments has been frequently referred to as Macrobrachium intermedium but is treated below as a species of Palaemon. Diagnosis. Hepatic spine set well back from carapace edge on or just below ridge of the antennal spine. Branchiostegal groove short, running from carapace edge to base of spine. Pereopods 2 markedly sexually dimorphic, larger in males, spinulose. Males larger than females. Mandibular palp 3-articled. Pereopods 3–5 with dactylus simple, shorter than propodus. Pereopod 5 propodus with transverse rows of setae in distal part of the posterior margin. Male pleopod 1 without appendix interna. Macrobrachium novaehollandiae (de Man, 1908) (Fig. 23d). Rostrum with 10 dorsal teeth, 3 posterior to orbit of the eye; 5 ventral teeth. Pereopod 2 carpus almost twice as long as the merus. Anteroventral corner of carapace slightly rounded. tl. 118 mm, cl. 46 mm. Qld, NSW, WA; estuarine.

Palaemon Weber, 1795 About 34 species comprise the genus Palaemon (Chace, 1993). Species are found worldwide in tropical and temperate seas or brackish and fresh water. Most are littoral inhabitants of algal communities. The species can be difficult to distinguish from each other but the proportions of the carpus and propodus of pereopod 2, length of the rostrum, and degree of fusion of the antennal flagella are critical. The New Zealand species, Palaemon affinis Milne-Edwards, has been reported from Australia but Yaldwyn (1954) dismissed these records. Diagnosis. Carapace smooth, with distinct branchiostegal spine on carapace edge (sometimes set back) and groove usually running back from carapace edge in a shallow arc, or over spine, or, into and over but not past (P. intermedius). Mandibular palp usually of 2 or 3 articles, but may be absent. Propodus of fifth pereopod with several transverse rows of setae on distal part of the posterior margin. First pleopod of male without or with rudimentary appendix interna on endopod. Palaemon cf. debilis Dana, 1852 (Fig. 23e). Rostrum exceeding tip of scaphocerite by less than quarter of its length, strongly upturned anteriorly; with 2–8 dorsal teeth (1 post-orbit) and 1 subterminal dorsal tooth, 3–4 ventral teeth. Antennule with free part of shorter dorsal flagellum as long a fused part. Carpus of pereopod 2 much longer than palm (propodus proximal to finger), reaching tip of scaphocerite. cl. 10 mm. Palaemon debilis is widespread in Indo- West Pacific and Qld. Populations in NSW and eastern Vic. have a shorter rostrum relative to the scaphocerite and fewer ventral rostral teeth (P. debilis has 5–7) and are doubtfully the same species. Palaemon dolospina Walker & Poore, 2003 (Figs 22b, 23i, Pl. 8a). Rostrum with 6–9 unevenly spaced dorsal teeth, 2 posterior to orbit of the eye; 3–6 ventral teeth, usually 4. Branchiostegal spine set back from and its tip barely reaching anterior carapace edge, high on branchiostegite. Branchiostegal groove looping over spine. Carpus of male pereopod 2 longer than palm (propodus proximal to finger), reaching to tip of scaphocerite. Distinct transverse red and/or occasionally olive stripes across all abdominal somites; less distinct longitudinal and oblique red stripes on carapace; antennule inner flagellum pink with few white flecks; often row of white chromatophores along lateral carina of the rostrum. cl. 15 mm. Tas.,Vic., SA, SA; 0–3 m

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depth. This common species is easily confused with P. intermedius. The two may co-occur on muddy sediments and in seagrasses. This species is common in bays where there is estuarine influence. Palaemon intermedius (Stimpson, 1860) (Fig. 23c). Rostrum with 7–10 dorsal teeth, usually 8, 2–3 posterior to orbit of the eye; 4–6 ventral teeth, usually 5. Branchiostegal spine falling short of carapace margin. Branchiostegal groove not running back along the carapace from hepatic spine. Pereopod 2 carpus longer than merus; fixed finger with cutting edge bearing 2 proximal teeth; 1 tooth between these on dactylus. Anteroventral corner of carapace square or slightly rounded. Distinct oblique and transverse red lines on carapace; few scattered red, olive and black chromatophores forming indistinct transverse lines on abdominal somites; antennule inner flagellum with definite red and white bars, persisting as purple bars in ethanol- preserved specimens. cl. 15 mm. Southern Qld, NSW, Vic., Tas., SA, southern WA; seagrass and algae, 0–22 m depth. This species is one of the most common shallow water carideans found in southern Australia. It is found on muddy soft sediments especially in the shelter of seagrasses. When alive the shrimp is translucent, marked with fine purple-red lines running along the carapace and more sparsely on the abdomen. This species, described as a species of Leander, was placed in Macrobrachium by Holthuis (1952) and has been commonly referred to as Macrobrachium intermedium in the ecological literature. Its misplacement has been long realised, on the basis of morphological examination (Walker, 1979), of allozymes (Boulton & Knott, 1984), and more recently following phylogenetic studies using 16S rRNA (Murphy & Austin, 2002; 2003). Walker & Poore (2003) redescribed the species in detail and argued for its placement in Palaemon; they recognised more slender forms in deeper water in Tasmania. Palaemon litoreus (McCulloch, 1909) (Fig. 23f). Rostrum reaching slightly beyond, but not far exceeding tip of scaphocerite, broad and straight; with 7–8 dorsal teeth, 2–3 ventral teeth. Flagella of antennule fused for 6–7 articles (one-fifth of length). Carpus of pereopod 2 shorter than palm (propodus proximal to finger), not reaching tip of scaphocerite. Saddle stripes of red across back and sides of whole body; less distinct on carapace; palm of pereopod 2 with narrow red band; fingers with similar band half way along. cl. 12 mm. NSW,Vic., SA, WA; 0–3 m on rocky habitats. Palaemon macrodactylus Rathbun, 1902 (Fig. 23g). Rostrum reaching to tip of scaphocerite, broad and straight; with 10–12 dorsal teeth and 1 subterminal tooth, 3–4 ventral teeth. Flagella of antennule fused for 5–9 articles (one-quarter of length). Carpus of male pereopod 2 longer than palm (propodus proximal to finger), reaching beyond tip of scaphocerite. Grey or olive- green; distinct wide, grey band across palm of pereopod 2; diffuse longitudinal and oblique rows on carapace; diffuse transverse lines on posterior edges of abdominal somites. cl. 20 mm. Japan, Korea, northern China, introduced to San Francisco Bay and to NSW, SA; estuarine and shallow marine waters. P. macrodactylus is a shrimp with long second pereopods in the male and with the branchiostegal spine only just overlapping the anterior margin of the carapace. The species was described in detail by Newman (1963) who recognised it as a species introduced to the USA about 1957. The species was first recognised as introduced to Australia in the late 1970s, in warm cooling ponds near power stations in SA and central NSW (Buckworth, 1979) and has not been reported since or in other places. The species lives in shallow marine and estuarine bays and lakes. Palaemon serenus (Heller, 1862) (Figs 22a, 23h, Pl. 8c). Rostrum reaching slightly beyond, but not far exceeding tip of scaphocerite, tapering and slightly upturned; with 6–9 dorsal and 1 subterminal teeth, 3–5 ventral teeth. Flagella of antennule fused for 12 articles (one-third of length). Carpus of male pereopod 2 longer than palm (propodus proximal to finger), reaching to or exceeding tip of scaphocerite. Broad, red band across most of palm of pereopod 2;

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fingers white; abdomen with longitudinal rows of few large spots of red, blue and black chromatophores; distinct longitudinal stripes on carapace. cl. 25 mm. Qld, NSW, Vic., Tas., SA, WA; 0–9 m depth on weedy rocky reefs or in rock pools. The species is the most common shrimp in rock pools along much of the coast appearing as a translucent shape with red bands and white tips to the second chelipeds.

Palaemonetes Heller, 1869 Holthuis (1950, 1993) regarded the number of articles of the mandibular palp as important for distinguishing Palaemonetes from Palaemon. Walker (1979) and Bray (1976) noted that this character is unreliable for distinguishing genera as some species of Palaemon have no mandibular palp and some Palaemonetes have a reduced palp. Murphy & Austin (2002, 2003) found on the basis of molecular analysis that species of Palaemonetes and Palaemon in Australia could not easily be untangled. While most individuals of both Australian species of Palaemonetes lack mandibular palps others have one or two articles. Diagnosis. Hepatic spine absent. Branchiostegal spine present. Branchiostegal groove present, usually extending posterior to branchiostegal spine. Mandibular palp absent or of 1–2 articles. Pereopods 3–5 with dactylus simple, shorter than propodus. Pereopod 5 propodus without transverse rows of setae in distal part of the posterior margin. Male pleopod 1 without appendix interna. Palaemonetes atrinubes Bray, 1976 (Fig. 23j). branchiostegal spine on anterior margin of carapace. Rostrum unarmed dorsally for one-third its length in distal region; 4–7 teeth dorsally, 2–4 ventrally. Antennule with 16 fused articles, twice length of 6 free articles. Distinct black spot at posterolateral end of abdominal somite 6; 3 distinct red and blue lines on carapace; 1 transverse line across posterior edge of abdominal somite 3. tl. 40 mm. NT, Qld, Vic., WA; shallow marine and estuarine habitats. The species is most easily distinguished from other species of Palaemonetes and species of Palaemon by the anteriorly unarmed rostrum. The biology and environment of this species is much like that of the following species. Bray (1976) found that unlike P. australis which hid during the day, P. atrinubes often swam over the substrate by day. Palaemonetes australis Dakin, 1915 (Fig. 23k). Branchiostegal spine distant by about its own length from anterior margin of carapace. Rostrum with regularly spaced teeth along entire length; 6 dorsally, 3 ventrally. Antennule with 10 fused articles, 0.6 length of 3 free articles. Overall olive-green or brown due to red, white, yellow and blue chromatophores; carapace with diffuse dorsal and anterior bands and indistinct mottling posteriorly; antennule inner flagellum pale red. tl. to 30 mm. WA; freshwater and estuarine habitats. The placement of the branchiostegal spine distinguishes this species from P. atrinubes. This is one of the few Australian shrimps about whose biology something is known. Bray (1976) found that it toler- ated salinities from 0 to >25 ppt in sheltered areas of rivers and estuaries of south-western WA, carried batches of up 70 eggs from September to April, and was a predator on smaller crustaceans, diatoms and forams.

Pontoniinae Kingsley, 1878 Pontoniine species are typically commensals or associates of sessile or larger mobile invertebrates such as sponges, hard and soft corals of all sorts, gastropods and bivalves, echinoderms and ascidians. Bruce (1983; 1990) listed all the Australian species then known and their associates. Most are tropical and the fauna is diverse on coral reefs. There is also a rich deep-sea fauna. Members of this subfamily are less common in southern Australian waters than the Palaemoninae. Of the 71 genera worldwide, only four have representatives in southern Australia.

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The key includes only the genera recorded so far. If a specimen does not seem to fit the descriptions, consult the papers of Holthuis (1993) and Bruce (1994) who gave keys to all genera. Species that might occur as far south as our region on the east coast are: Coralliocaris graminea (Dana, 1852), C. superba (Dana, 1852), Harpiliopsis depressa (Stimpson, 1860), Periclimenes holthuisi Bruce, 1969, P. madreporae Bruce, 1969, and P. soror Nobili, 1904.

Diagnosis. Maxilliped 3 without or with rudimentary arthrobranch. Telson usually with 2 pairs of dorsal robust setae, 3 pairs of posterior robust setae and with 1 or more pair of hairs.

Anchistus Borradaile, 1898 Only one of the seven known species (Chace, 1993) is recorded from southern Australia; others appear on tropical reefs. Diagnosis. Subcylindrical shrimps with smooth carapace and abdomen. Rostrum over-reaching eyes, dentate, laterally compressed in distal half. Antennal spine absent or weak, hepatic spine absent. Mandibular palp absent. Maxilliped 3 with exopod. Uropodal exopod with small mobile distolateral seta. Anchistus custos (Forskål, 1775). Carapace without antennal spine. Rostrum unarmed. Pereopod 1 chela lower edge produced into a downwardly bent flap thus forming an open tube. cl. 9 mm. Indo-West Pacific (E to Fiji), WA, Qld, SA; intertidal to shallow depths, living in bivalve molluscs. This species, like others in its genus, occurs as heterosexual pairs in association with bivalves of the families Tridactinidae, Pinnidae and Pectinidae (giant clams, razor mussels and scallops). It is one of the most common of all pontoniines.

Notopontonia Bruce, 1991 This genus contains one species whose flattened chelipeds are characteristic (Bruce, 1991a; Berggren, 1999). Diagnosis. Strongly depressed shrimps with smooth carapace and abdomen. Rostrum over-reaching eyes, unarmed, dorsally concave. Antennal and hepatic spines absent. Mandibular palp absent. Maxilliped 3 with exopod. Pereopod 1 with elongate spatulate fingers. Chelae of pereopod 2 strongly compressed and carried in the lateral plane. Uropodal exopod with small distolateral tooth and larger mobile seta medially. Notopontonia platycheles Bruce, 1991 (Fig. 24a). Males mostly pale pink/semitransparent, females red. tl. 36 mm. WA (N to Shark Bay), SA; commensal in branchial chamber of solitary tunicate Herdmania momus (Savigny, 1816), 7–64 m depth. The colour of this shrimp matches that of its host. It lives usually solitarily, infecting about three-quarters of host individuals (Berggren, 1999).

Palaemonella Dana, 1852 Of the 14 species worldwide only two species have been recorded from southern Australia (Chace, 1993). Bruce (2002) provided a key to Indo-West Pacific species of Palaemonella. Diagnosis. Rostrum overreaching eyestalks, armed dorsally and ventrally, not laterally expanded. Carapace with antennal and hepatic spines, without branchiostegal groove. Mandibular palp present, 2 articles. Maxilliped 3 bearing exopod. Abdominal somite 5 with sharp posterior corner on pleura.

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Fig. 24. Palaemonidae Pontoniinae. a, Notopontonia platycheles (from Bruce, 1991a). b, Periclimenes aesopius (from Bruce, 1977). c, Periclimenes carinidactylus (from Bruce, 1980).

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Palaemonella foresti Bruce, 2002 (Fig. 25c). Rostrum well exceeding scaphocerite, slender, with 7 dorsal teeth, 2 on carapace proper, and 4 ventral teeth Carapace without supraorbital ridge, spines or tubercles, postantennal ridge feebly present Mandible with 2-articled palp. Pereopod 2 carpus without preterminal tooth, merus with distoventral tooth, ischium unarmed, ambulatory dactyls not elongate, less than 0.2 of propodus length, ventrally biconcave, propodus with moderately long distoventral spines, numerous ventral spines. cl. 6.1 mm. WA (Cockburn Sound); harbour piles. Palaemonella rotumana (Borradaile, 1898) (Fig. 25d). Rostrum with 6–8 dorsal teeth, 2 on carapace proper, and 1–3 ventral teeth. Carapace with supraorbital spine (or tubercle). Pereopod 2 carpus armed distally with 1–2 marginal teeth; merus armed with sharp distal tooth on flexor margin. Pereopod 3 with flexor margin of dactylus concave. cl. 4.3 mm. Mediterranean, Red Sea, Indo-West Pacific, WA, NT, Qld, NSW; associated with coral on muddy bottom, to 128 m depth. This is the most widely distributed species of Palaemonella (Bruce, 2002).

c e d

Fig. 25. Palaemonidae Pontoniinae. a, Periclimenes yaldwyni (from Bruce & Cropp, 1984). b, Pseudopontonia minuta (from Bruce, 2003) c, Palaemonella foresti. d, Palaemonella rotumana. e, Periclimenes laccadivensis.

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Periclimenes Costa, 1844 Of more than 160 species described in this largely tropical genus (Chace, 1993) more than 60 have been recorded from Australia (Bruce, 1983, 1990). Four may be found in southern waters but none is common. The brief diagnoses below should be used only to differentiate these four. Diagnosis. Rostrum overreaching eyestalks, armed dorsally and/or ventrally, not laterally expanded. Carapace with antennal and hepatic spines, without branchiostegal groove. Scaphocerite well developed. Mandibular palp absent. Maxilliped 3 with exopod. Pereopods 3–5 with ischium and merus not fused. Abdominal somites with rounded pleura. Uropodal exopod with at least 1 movable lateral spine. Periclimenes aesopius (Bate, 1863) (Fig. 24b, Pl. 8b). Abdominal somite 3 strongly lobed posterodorsally. Rostrum slightly downturned, 6 times as long as deep, with 9–11 dorsal teeth, 3–4 on carapace proper; 2–4 ventral teeth in distal quarter. Eyestalk 2.3 times as long as wide, with ocellus. cl. 5.6 mm. Red spotted on rostrum, antennae and pereopods. SA, WA; possibly an associate of sponges. This rare species was redescribed in detail by Bruce (1977); it has not previously been reported from WA. Periclimenes carinidactylus Bruce, 1969 (Fig. 24c). Abdominal somite 3 not lobed posterodorsally. Rostrum slightly convex dorsally, 6 times as long as deep, with 10 dorsal teeth, 2 on carapace proper; 2 ventral teeth in distal half. Eyestalk twice as long as wide, without ocellus. cl. 6.6 mm, tl. 15.8 mm. Translucent, with yellowish-brown rostrum, eyestalks and pereopods 2, white dorsal band, reddish ventrally. NSW (Port Jackson), SA (Kangaroo I.); 6–9 m depth on crinoid, Comanthus trichoptera. The species was described in detail from only the second specimen by Bruce (1980). Periclimenes laccadivensis (Alcock & Anderson, 1894) (Fig. 25e). Abdominal somite 3 not lobed posterodorsally. Rostrum slightly upcurved, 6 times as long as deep, with 8–10 dorsal teeth, 2 on carapace proper; 2–4 ventral teeth in distal half. Eyestalk 1.5 times as long as wide, without ocellus. cl. 4.3 mm. Red spotted on rostrum, antennae and pereopods. Indo-West Pacific, Tas.; 274–1285 m depth; possibly an associate of gorgonians. The only Australian record is from very deep water (Bruce, 1991b, 1992a). Periclimenes yaldwyni Holthuis, 1959 (Fig. 25a). Abdominal somite 3 not lobed posterodorsally. Rostrum straight, 4 times as long as deep, with 8–9 dorsal teeth, 2 on carapace proper; 4–5 ventral teeth in distal quarter. Eyestalk 1.3 times as long as wide, with ocellus. Transparent with red spots and bars on carapace, rostrum, antennae and pereopods. cl. 5.3 mm. New Zealand, Tas.; 1–30 m, with red macroalgae. Bruce & Cropp (1984) redescribed P. yaldwyni, one of few Australian caridean species shared with New Zealand.

Pseudopontonia Bruce, 1992 A single species has been recorded from southern Australia (Bruce, 1992b, 2003). Diagnosis. Rostrum overreaching eyestalks, flattened dorsally, usually unarmed, laterally expanded into eave. Carapace without antennal and hepatic spines, without branchiostegal groove. Scaphocerite well developed. Mandibular palp absent. Maxilliped 3 with exopod. Pereopods 3–5 with ischium and merus not fused. Abdominal somites with rounded pleura. Uropodal exopod with 1 movable lateral robust seta. Pseudopontonia minuta (Baker, 1907) (Fig. 25b). Antennal spine absent. Pereopod 1 fixed finger with row of long setae on lower margin. Pereopod 3–5 dactylus tapering, with an accessory spine slightly more robust and shorter than unguis. Telson with rounded apex bearing 6 pairs of setae. cl. 4.6 mm. NSW, SA, WA; subtidal, in branchial sac of ascidian Polycarpa flava.The species was described in detail as a species of Pontonia by Bruce (1972). The male was

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described later (Bruce, 2003). The species has unique abbreviated larval development (Bruce, 1993).

References Berggren, M. 1999. Notopontonia platycheles, a little-known shallow-water shrimp from Western Australia (Decapoda: Pontoniinae). Journal of Crustacean Biology 19: 180–187. Boulton, A.J., & Knott, B. 1984. Morphological and electrophoretic studies of the Palaemonidae (Crustacea) of the Perth region, Western Australia. Australian Journal of Marine and Freshwater Research 35: 769–783. Bray, D.M. 1976. A review of two Western Australian shrimps of the genus Palaemonetes, P. australis Dakin 1915 and P. atrinubes sp. nov. (Decapoda, Palaemonidae). Records of the Western Australian Museum 4: 65–84. Bruce, A.J. 1972. Notes on some Indo-Pacific Pontoniinae, XVIII. A re-description of Pontonia minuta Baker, 1907, and the occurrence of abbreviated development in the Pontoniinae (Decapoda Natantia, Palaemonidae). Crustaceana 23: 65–75. Bruce, A.J. 1977. A redescription of Periclimemes aesopius (Bate, 1863) (Crustacea: Decapoda) with remarks on related species. Australian Zoologist 19: 217–229. Bruce, A.J. 1980. Periclemes carinidactylus Bruce, a crinoid-associated pontoniine shrimp, from South Australia. Crustaceana 38: 295–305. Bruce, A.J. 1983. The pontiniine shrimp fauna of Australia. Memoirs of the Australian Museum 18: 195–218. Bruce, A.J. 1987. Records of three palaemonid shrimps new the Australian fauna. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 4: 57–60. Bruce, A.J. 1990. Recent additions to the pontinine shrimp fauna of Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 7: 9–20. Bruce, A.J. 1991a. Notopontonia platycheles, new genus, new species (Decapoda: Pontoniinae) from South Australia, with remarks on Pontonia pinnophylax (Otto), the type species of Pontonia Latrielle. Journal of Crustacean Biology 11: 607–628. Bruce, A.J. 1991b. Crustacea Decapoda: further deep-sea palaemonoid shrimps from New Caledonian waters. In: A. Crosnier (ed.), Résultats des Campagnes MUSORSTOM,Vol.9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152: 299–411. Bruce, A.J. 1992a. Two new species of Periclimenes (Crustacea: Decapoda: Palaemonidae) from Lizard Island, Queensland, with notes on some related taxa. Records of the Australian Museum 44: 45–84. Bruce, A.J. 1992b. Designation of two Pontoniine shrimp genera (Decapoda: Palaemonidae). Journal of Natural History 26: 1273–1282. Bruce, A.J. 1993. Kakaducaris glabra gen. nov., sp. nov., a new freshwater shrimp from the Kakadu National Park, Northern Territory, Australia, Crustacea: Decapoda: Palaemonidae with designation of a new subfamily Kakaducaridinae. Hydrobiologia 268: 27–44. Bruce, A.J. 1994. A synopsis of the Indo-West Pacific genera of the Pontoniinae (Crustacea: Decapoda: Palaemonidae). Koeltz Scientific Books: Königstein. 172 pp. Bruce, A.J. 2002. Notes on some Indo-Pacific Pontoniinae, XLVI. Palaemonella foresti sp. nov., a new pontoniine shrimp from Western Australia (Decapoda, Palaemonidae), with a review of the Indo- West Pacific species of the genus Palaemonella Dana, 1852. Crustaceana 75: 277–298. Bruce, A.J. 2003. Further information on two pontoniine shrimps from ascidian hosts, Dasella brucei Berggren, 1990 and Pseudopontonia minuta (Baker, 1907) (Crustacea: Decapoda: Palaemonidae). Memoirs of the Queensland Museum 49: 111–114. Bruce, A.J., & Cropp, D.A. 1984. A redescription of Periclimenes yaldwyni Holthuis (Brachycarpus audouini Bate, 1888, Crustacea, Decapoda, Palaemonidae) and its occurrence in Australian waters. Pacific Science 38: 189–198.

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Buckworth, R.C. 1979. ‘Aspects of the population dynamics of Palaemon macrodactylus (Decapoda, Palaemonidae) in Lake Mannering, N.S.W., and in the laboratory’. M.Sc. thesis. University of New South Wales, Kensington Chace, F.A. 1972. Palaemon debilis from Hawaii and the status of the genus Palaemonetes (Decapoda, Palaemonidae). Crustaceana 23: 12–19. Chace, F.A. 1993. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 6: superfamily Palaemonoidea. Smithsonian Contributions to Zoology 543: 1–152. De Man, J.G. 1908. Description of a species of Palaemon from near Sydney, probably either a new species or he adult form of Palaemon (Eupalaemon) danae. Annals and Magazine of Natural History 1: 363–370, pl. 16. Fincham, A.A. 1987. A new species of Macrobrachium (Decapoda, Caridea, Palaemonidae) from Northern Territory, Australia and a key to Australian species of the genus. Zoologica Scripta 16: 351–354. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Holthuis, L.B. 1950. The Decapoda of the Siboga Expedition. Part X. The Palaemonidae collected by the Siboga and Snellius Expeditions with remarks on other species. I. Subfamily Palaemoninae. Siboga Expéditie Monographie 39a9: 1–268. Holthuis, L.B. 1952. On some Indo-westpacific Palaemoninae (Crustacea Decapoda Caridea). Zoologische Mededelingen, Leiden 31: 201–211. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Horwitz, P. 1995. A Preliminary Key to the Species of Decapoda (Crustacea: Malacostraca) found in Australian Inland Waters (Co-operative Research Centre for Freshwater Ecology Identification Guide 5). Co-operative Research Centre for Freshwater Ecology: Albury, NSW. 69 pp. Murphy, N.P., & Austin, C.M. 2002. A preliminary study of 16S rRNA sequence variation in Australian Macrobrachium shrimps (Palaemonidae: Decapoda) reveals inconsistencies in their classification. Invertebrate Systematics 16: 697–701. Murphy, N.P., & Austin, C. 2003. Molecular taxonomy and phylogenetics of some species of Australian palaemonid shrimps. Journal of Crustacean Biology 23: 169–177. Newman, W.A. 1963. On the introduction of an edible oriental shrimp (Caridea, Palaemonidae) to San Francisco Bay. Crustaceana 5: 119–132. Walker, T.M. 1979. ‘A study of sympatry in two species of Palaemoninae. PhD thesis. University of Tasmania: Hobart. 234 pp, plus appendices. Walker, T.M., & Poore, G.C.B. 2003. Rediagnosis of Palaemon and differentiation of south-eastern Australian species (Crustacea: Decapoda: Palaemonidae). Memoirs of Museum Victoria 60: 243–256. Yaldwyn, J.C. 1954. Studies on Palaemon affinis M.-Edw. 1837, (Crustacea, Decapoda, Natantia.) Part 1. Synonymy and external morphology. Transactions of the Royal Society of New Zealand 82: 169–187.

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Superfamily Alpheoidea Rafinesque, 1815 The three families included by Holthuis (1993) are represented in southern Australia. Martin & Davis (2001) added a fourth, Barbouriidae Christofferson, 1987, one of numerous families suggested for members of the Hippolytidae (Christoffersen, 1987).

Alpheidae Rafinesque, 1815 The family Alpheidae includes the snapping shrimps renowned for the sharp clicking noise made with the larger chela but not all alpheids have this capability. Nevertheless, the commonly collected ones in southern Australia do and comprise the most diverse of the caridean families. Individuals are frequently seen under rocks, in burrows and in sponges near low water level and subtidally. Alpheids are probably the most abundant family of caridean shrimps subtidally and their clicking noise can frequently be heard by divers or even through the hull of boats. The click generates a shock wave which stuns prey (other crustaceans) before being captured. There are almost 500 known species of Alpheidae and Holthuis (1993) provided a key to 31 genera but more have been described since. The family is especially diverse in coral rubble and reefs in the tropics but even in southern Australia almost 50 described species have been recorded. The late husband and wife team, Dora and Albert Banner, spent many years working on Australian alpheids in the 1960s and 1970s and contributed four papers which covered the fauna then known (Banner & Banner, 1973, 1975, 1982a, b). Although their keys, detailed drawings and descriptions are essential, it has not always been possible to identify tropical species from their works and many more species exist than the 140 dealt with by them. Keys here are derived from their papers and, for the genera, from Holthuis (1993). Of the 13 Australian genera, eight occur on southern coasts and shelf. Many currently accepted alpheid species are widespread throughout the Indo-West Pacific region and are morphologically variable, especially in the shape of the rostrum, development of spines and other taxonomically useful characters. This may indicate the presence of undescribed cryptic species. Many of the keys to species of alpheids rely on the shape of the larger chela (propodus and dactylus of the cheliped) but in material from dredge samples this is often lost. The dorsal view of the eyes, rostrum, ocular processes and associated grooves can be a reliable guide to identification. Many species have sexually dimorphic chelipeds, much larger in males. This could make identification of a single specimen difficult but often cryptic specimens from sponges and encrusting habitats occur in male-female pairs. Lengths of specimens are total lengths, rostrum to end of telson.

Diagnosis. Carapace smooth, without antennal and branchiostegal spines. Rostrum reduced. Eyes usually covered partially or entirely by anterior margin of carapace. Pereopods 1 dissimilar, larger one swollen. Pereopod 2 carpus of 3–5 articles.

Key to southern Australian genera of Alpheidae 1. Pereopods without epipods; anteroventral corner of carapace (pterygostomial angle) produced as a rounded or acute lobe ...... Synalpheus … p. 113 — At least pereopods 1 and 2 with epipods (short posteriorly directed curved appendages at the base of legs and covered by carapace); anteroventral corner of carapace rounded, rarely produced ...... 2 2. Pleuron of abdominal somite 6 without an articulating posteroventral triangular plate . . . 3 — Pleuron of abdominal somite 6 with an articulating posteroventral triangular plate ...... 4

98 Caridea – shrimps

3. Antepenultimate article of maxilliped 3 broad and flat, suboperculiform . . Metalpheus … p. 113 — Maxilliped 3 pediform, antepenultimate article not operculiform ...... Alpheus … p. 99 4. Rostrum absent or indistinct ...... Betaeus … p. 113 — Rostrum present ...... 5 5. Rostrum laterally flattened, its tip rounded ...... Athanopsis … p. 112 — Rostrum slender, pointed in lateral view ...... 6 6. Eyes entirely or partially visible in dorsal view and concealed in lateral view only by anterior tooth of carapace ...... 7 — Eyes concealed in dorsal view and partially to completely concealed in lateral view ...... Alpheopsis … p. 99 7. Pereopod 2 carpus with 5 articles ...... Athanas … p. 111 — Pereopod 2 carpus with 4 articles ...... Arete … p. 111

Alpheopsis Coutière, 1897 Banner & Banner (1973) recorded four species from Australia but only one occurs in the south. Diagnosis. Pereopods 1–3 at least with epipods. Pleuron of abdominal somite 6 without an articulating posterior triangular plate. Rostrum present, slender. Carapace covering eyes, at least dorsally. Chelipeds asymmetrical; large chela carried forward, fingers without plunger and socket. Alpheopsis trispinosa (Stimpson, 1861) (Figs 26a, b, 29a, m'). Carapace with rostrum and orbital teeth. Chelae of similar size and both with dorsal longitudinal and transverse grooves. 19 mm. Southern NSW, Tas.,Vic., SA; 11–135 m depth (common in Bass Strait).

Alpheus Fabricius, 1798 Alpheus is the largest genus of caridean shrimp in southern Australia with at least 27 described species known. It is distinguished from Synalpheus by the absence of a pterygostomial angle (anteroventral corner of the carapace) and possession of small epipods at the base of the first four pereopods. Davie (2002) listed 84 Australian species but repeated the view of A. Anker that several widespread species were probably complexes of species (bidens, collumianus, digitalis, facetus, lobidens, pacificus, sulcatus). Several species have subspecies and complex synonymies (Banner & Banner, 1982a; Bruce, 1994) indicating considerable variability over sometimes wide geographic areas. The proportions of the rostrum and development of the sculpture and spines may vary with size and differ from our figures. Identification of species can be difficult or impossible if the large cheliped is missing (often the case when specimens are not individually carefully collected) or, in some cases, without small chelipeds of both sexes. The shape of the rostrum and orbital hoods may be useful in these cases. The shape of the groove alongside the base of the rostrum, the orbitorostral groove, is specific to several taxa. Some species of Alpheus possess what is called a ‘balaeniceps’ small chela on the male or both sexes. This term derives from the similarity of the regular row of thick setae around the margin of the flattened dactylus to the baleen of whales. In the southern Australian species this remarkable structure is relatively easy to distinguish from the rows of more well spaced simple setae on the fingers of other species. Juveniles may not have this setation. Hale (1927) used the older rejected name Crangon for some species of Alpheus but this is no longer applicable to snapping shrimps. The key is adapted from Banner & Banner (1982a).

Diagnosis. Pereopods 1–4 with epipods. Pleuron of abdominal somite 6 without an articulating posterior triangular plate. Rostrum well developed and continued posteriorly as carina. Carapace

99 Marine Decapod Crustacea of Southern Australia

covering eyes completely in dorsal and lateral views, often with orbital processes; usually grooves between orbital hoods and rostral carina. Orbitorostral process usually present. Pterygostomial angle not produced. Chelipeds markedly asymmetrical, larger one swollen, carried extended, dactylus with plunger fitting into socket at base of fixed finger. Smaller cheliped with fingers simple.

Key to southern Australian species of Alpheus The large cheliped is usually essential for correct identification but if it is missing it may be possible to identify a common species from the shape of the anterior part of the carapace. Alpheus bunburius, known only from a single female, appears twice in the key. 1. Large chela with at most a depressed area on lower margin, never a shoulder; never a U- shaped transverse groove on propodus ...... 2 — Large chela with a definite shoulder on lower margin proximal to fixed finger; upper margin of propodus with U-shaped groove proximal to dactylar articulation usually expanding into depressed areas laterally ...... 18 2. Orbital hoods with acute teeth or teeth between hood and rostrum ...... 3 — Orbital hoods without acute teeth; no teeth between hood and rostrum ...... 12 3. Large chela with strong acute lateral and medial teeth near dactylar articulation; propodus with heavy longitudinal ridge ...... 4 — Large chela without acute teeth near dactylar articulation; propodus with only weak longitudinal sculpture ...... 7 4. Orbitorostral grooves short and slight ...... 5 — Orbitorostral grooves definite and extending behind eyes ...... 6 5. Large chela propodus upper margin longitudinal ridge terminating at tooth . . Alpheus astrinx — Large chela propodus upper margin longitudinal ridge terminating abruptly proximal to tooth ...... Alpheus hailstonei 6. Inner side of both chelae with setae so heavy as to almost obscure outlines ...... Alpheus deuteropus — Inner side of both chelae with only scattered setae ...... Alpheus collumianus 7. Rostral base between eyes gradually curving into orbitorostral grooves ...... 8 — Rostral base between eyes abruptly set off from orbitorostral grooves, sometimes overhanging ...... 10 8. Large chela with upper surface papillose; anterior carapace and chela hirsute . Alpheus villosus — Large chela with upper surface smooth; anterior carapace and chela with only scattered setae ...... 9 9. Large chela with slight transverse depression on upper margin, no longitudinal crest; frontal margin between orbital hoods and rostrum (rostrum-hood) recessed . . . . . Alpheus gracilis — Large chela without transverse depression on upper margin, with slight longitudinal crest; frontal margin between orbital hoods and rostrum (rostrum-hood) produced into convex prominence ...... Alpheus facetus 10. Large chela not sculptured; pereopod 3 dactylus apex rounded ...... Alpheus lottini — Large chela sculptured; pereopod 3 dactylus simple or bifid, acute ...... 11 11. Maxilliped 3 with penultimate article expanded distally into definite lobe . . Alpheus socialis — Maxilliped 3 with penultimate article expanded only slightly distally . . Alpheus parasocialis

100 Caridea – shrimps

12. Large chela rounded, not sculptured ...... Alpheus spongiarum — Large chela laterally compressed and/or with sculpture ...... 13 13. Rostrum flat, or if carinate not higher than orbital hoods; large chela oval to compressed in section, ridged but not pustulate ...... 14 — Rostrum carinate and elevated; large chela quadrangular in section (with angular upper, lateral and lower ridges) and pustulate (sandpaper-like) ...... 17 14. Large chela with transverse slit on upper surface near distal margin of propodus; rostrum not strongly overhanging orbitorostral groove laterally ...... 15 — Large chela without transverse slit on upper surface of propodus; rostrum strongly overhanging orbitorostral groove laterally ...... 16 15. Conspicuous flattened teeth mesial to base of orbital hood ...... Alpheus bidens — No teeth mesial of base to orbital hood ...... Alpheus novaezealandiae 16. Rostrum short, free section shorter than orbitorostral grooves, with stiff marginal setae; pereopod 2 with first carpal segment slightly longer than second ...... Alpheus sulcatus — Rostrum long, free section as long as orbitorostral grooves, without marginal setae; pereopod 2 with first carpal segment twice length of second ...... Alpheus australosulcatus 17. Carapace pustulate, like chela, especially anteriorly ...... Alpheus stephensoni — Carapace smooth ...... Alpheus digitalis 18. Small chela with only weak setation on fingers, not of balaeniceps form ...... 19 — Small chela of balaeniceps form, with row of stiff thick marginal setae on dactylus of male or both sexes ...... 22 19. Large chela with shoulder proximal to upper saddle, overhanging ...... 20 — Large chela with shoulder proximal to upper saddle, not overhanging ...... 21 20. With acute terminal or subterminal tooth on lower mesial margin of merus of large cheliped ...... Alpheus macrodactylus — With lower mesial margin of merus of large cheliped unarmed ...... Alpheus pacificus 21. Large chela with lower shoulder and another proximal to upper saddle low and rounded in profile ...... Alpheus bunburius — Large chela with both shoulders abrupt in profile ...... Alpheus tasmanicus 22. Large chela with shoulder proximal to upper saddle rounded to abrupt, not overhanging . 23 — Large chela with shoulder proximal to upper saddle overhanging ...... 24 23. Pereopod 3 dactylus flattened to subspatulate ...... 25 — Pereopod 3 dactylus simple and conical ...... 26 24. Small chela of balaeniceps form in both sexes; rostrum as long as wide at base ...... Alpheus australiensis — Small chela of balaeniceps form in male only; rostrum longer than wide at base ...... Alpheus lobidens 25. Pereopod 3 propodus setose laterally; large chela with plunger heavy and at a marked angle to distal margin ...... Alpheus richardsoni — Pereopod 3 propodus setose only marginally; large chela with plunger low and confluent with distal margin ...... Alpheus bunburius 26. Large chela with lower shoulder of outer face with numerous small papillae ...... Alpheus papillosus — Large chela with lower shoulder of outer face smooth ...... 27

101 Marine Decapod Crustacea of Southern Australia

27. Small chela of balaeniceps form in both sexes; rostrum twice as long as wide at base ...... Alpheus strenuus — Small chela of balaeniceps form in male only; rostrum 1.4 times as long as wide at base ...... Alpheus edwardsii Alpheus astrinx Banner & Banner, 1982 (Figs 26c, d, 29b, Pl. 8d). Rostrum a little longer than broad at base, separated from orbital hoods by broad grooves; orbital hoods with broad teeth, one-third as long as rostrum; rostrum-hood margins continuous. Large chela laterally compressed; lower margin barely concave; upper margin almost straight; with strong acute lateral and mesial teeth near dactylar articulation; propodus with weak longitudinal ridges, lateral one terminating at tooth; with only scattered setae. Small chela not of balaeniceps form. Bold bright red longitudinal stripes. 43 mm. WA (Perth), Vic. (Port Phillip Bay); subtidal in reef. Alpheus astrinx is a rare species and the single specimen from Victoria is the first record away from the type locality. Alpheus australiensis Banner & Banner, 1982 (Figs 26e, 29c, d). Rostrum conical, as long as wide at base; carina rounded; orbitorostral grooves rounded; orbital hoods without teeth; rostrum- hood margins convex. Large chela laterally compressed; lower margin with rounded shoulder next to broad notch more distal than upper saddle; upper margin with saddle defined proximally by rounded-abrupt shoulder and distally by lower elevation; without lateral and medial teeth near dactylar articulation; with sparse setae. Small chela of balaeniceps form in both sexes. Pereopod 3 dactylus simple and conical. Dark green. 35 mm. Qld (Whitsunday Group), NSW (S to Sydney); intertidal, estuarine muddy sediments. The small chela is of the balaeniceps form in both sexes but of this form only in the male in A. lobidens. This is one of several species burrowing in mud and associated with gobies, small fish which reside at the mouth of the shrimps’ burrows. Alpheus australosulcatus Banner & Banner, 1982 (Figs 26f, g, 29e, f). Rostrum flat, depressed apically, 1.5 times as long as wide at base, not bearing stiff setae laterally, free section as long as orbitorostral grooves which are overhung by base of rostrum and delimited from orbital hoods; orbital hoods without teeth but produced convexly; rostrum-hood margins deeply concave at base of rostrum. Large chela laterally compressed; lower and upper margins slightly concave; with obscure lateral longitudinal grooves; without lateral and mesial teeth near dactylar articulation; with numerous scattered setae on small bosses. Small chela not of balaeniceps form. Pereopod 2 with first carpal segment twice length of second. 65 mm. Central Vic.; intertidal and subtidal. Recent collections have been made in shallow water using the fish poison, rotenone. Although the species is rare in museum collections this may be only because it is very cryptic and lives deep in rocky environments. Alpheus bidens (Olivier, 1811) (Fig. 26h, i, Pl. 9a). Rostrum narrow, with carina leading back to second middorsal carina; orbitorostral grooves shallow and overhung posteriorly by pair of broadly triangular flat teeth; orbital hoods without teeth, overhanging anterior margin; rostrum-hood margin convex. Large chela cylindrical; lower margin slightly concave; upper margin with transverse slit near articulation with dactylus; without lateral and mesial teeth near dactylar articulation; with scattered setae medially. Small chela of balaeniceps form in male only, dactylus broadened. 77 mm. Eastern Pacific, Indonesia, Qld, Vic., SA, Tas., WA (Perth); intertidal to 83 m depth. The pair of flat dorsal teeth on the carapace immediately identify this species. Alpheus bunburius Banner & Banner, 1982 (Figs 26j, k, 29g, h). Rostrum triangular, slightly longer than broad, slightly carinate; orbitorostral grooves shallow; orbital hoods without teeth; rostrum-hood margins weakly concave. Large chela laterally compressed; lower margin with rounded shoulder; upper margin with shallow poorly defined saddle; without lateral and medial teeth near dactylar articulation; with sparse setae; dactylus half length of propodus,

102 Caridea – shrimps

d a bc

f g e

k i j h

m l n o

p q rs

t w u v

Fig. 26. Alpheidae. Dorsal and lateral views of anterior carapace: a, b, Alpheopsis trispinosa. c, d, Alpheus astrinx. e, Alpheus australiensis (from Banner & Banner, 1982a). f, g, Alpheus australosulcatus. h, i, Alpheus bidens. j, k, Alpheus bunburius (from Banner & Banner, 1982a). l, m, Alpheus edwardsii. n, o, Alpheus hailstonei. p, q, Alpheus lobidens. r, s, Alpheus lottini. t, u, Alpheus novaezealandiae. v, w, Alpheus pacificus.

103 Marine Decapod Crustacea of Southern Australia

with low plunger. Small chela of female not of balaeniceps form (male unknown). 38 mm. WA (Bunbury, known from only one individual). Alpheus collumianus Stimpson, 1860. Rostrum acute, short, carinate back to behind orbits; orbitorostral grooves deep but rounded; orbital hoods inflated, rounded, varying from armed with strong marginal tooth to unarmed; rostrum-hood margins convex. Carapace smooth. Large chela laterally compressed; margins nearly straight, upper margin arising proximally from narrow transverse saddle; with lateral tooth near dactylar articulation, without mesial tooth; with sparse setae on inner side. Small chela not of balaeniceps form; with only scattered setae. Indo-West Pacific (Madagascar to Japan and Society Is), Qld, WA; reef, to 76 m depth. Alpheus deuteropus Hilgendorf, 1879. Rostrum acute, curved abruptly upwards toward tip, carinate continuing to base of eye; orbitorostral grooves deep; orbital hoods inflated with acute teeth about as long as rostrum. Large chela strongly compressed, ovate in shape; upper margin interrupted by a deep transverse groove proximal to dactylar articulation; with lateral tooth near dactylar articulation, without medial tooth. Inner side of both chelae with setae so heavy as to almost obscure outlines. Small chela not of balaeniceps form. Indo-West Pacific (South Africa to Japan and Hawaii) Qld, WA; reef, to 35 m. Alpheus digitalis De Haan, 1844. Rostrum acute, carinate back to behind orbits; orbitorostral grooves deep; orbital hoods close together, without teeth; rostrum-hood margins not concave. Carapace smooth. Large chela laterally compressed; lower and upper margins straight; without lateral and medial teeth near dactylar articulation; with setae along ridges on lower and upper margins, and pustulate (like sandpaper) all over. Small chela not of balaeniceps form. 80 mm. Western Pacific, WA (S to Perth), NT, Qld, NSW (S to Botany Bay); on muddy sediments often in estuaries. Alpheus digitalis is a large species like A. stephensoni and is similar in possession of a sandpaper-like surface on the chelae. Unlike A. stephensoni, the carapace is smooth. Alpheus edwardsii (Audouin, 1827) (Figs 26l, m, 29i). Rostrum 1.4 times as long as wide at base; scarcely carinate; orbitorostral grooves deeply rounded; orbital hoods without teeth; rostrum- hood margins concave at base of rostrum. Large chela laterally compressed; lower margin with prominent shoulder next to broad notch running up laterally and opposite upper notch; upper margin with groove defined proximally by overhanging shoulder and distally by prominent elevation; without lateral and mesial teeth near dactylar articulation; with sparse setae. Small chela of balaeniceps form in male only. 40 mm. Indo-West Pacific, all coasts of Australia; intertidal to 25 m depth, muddy sediments under rocks. Alpheus facetus de Man, 1908. Rostrum acute, twice as long as broad at base, bluntly carinate, separated from orbital hoods by broad grooves; orbital hoods with converging acute teeth, one-third length of rostrum; rostrum-hood margins convex. Large chela laterally compressed; lower margin straight; upper margin with 2 longitudinal ridges separated by shallow groove; without lateral and medial teeth near dactylar articulation; with only scattered setae. Small chela not of balaeniceps form. 28 mm. Western Pacific, WA (S to Perth), NT, Qld, NSW (S to Sydney); intertidal to 30 m, in corals. Alpheus gracilis Heller, 1861. Rostrum as long as broad at base, separated from orbital hoods by broad grooves; orbital hoods with acute teeth, reaching two-thirds length of rostrum; rostrum-hood margins deeply incised. Large chela laterally compressed; lower margin barely concave; upper margin with shallow depression, without longitudinal ridges; without lateral and mesial teeth near dactylar articulation; propodus with weak longitudinal lateral depression; with only scattered setae. Small chela not of balaeniceps form. 35 mm; Indo-West Pacific, Qld, WA (Rottnest I.); intertidal and subtidal, rare in Australia. Alpheus hailstonei Coutière, 1905 (Figs 26n, o, 29j). Rostrum 1.5–2 times as long as broad at base, separated from orbital hoods by broad grooves reaching behind eyes; orbital hoods with

104 Caridea – shrimps

triangular teeth, two-thirds as long as rostrum; rostrum-hood margins continuous. Large chela laterally compressed; lower margin barely concave; upper margin almost straight; with strong acute lateral and medial teeth near dactylar articulation; propodus with weak longitudinal ridges, lateral one terminating before distal tooth; with only scattered setae. Small chela not of balaeniceps form. 30 mm. Eastern Indian Ocean, Indonesia, Japan, Hawaii, WA (S to Cape Naturaliste), Qld, NSW, Vic.; 25–536 m depth. Alpheus lobidens De Haan, 1844 (Figs 26p, q, 29k, l). Rostrum acute, narrow, carinate; orbitorostral grooves rounded; orbital hoods without teeth; rostrum-hood margins barely concave near rostral base. Large chela laterally compressed; lower margin with rounded shoulder next to broad notch more distal than upper saddle; upper margin with saddle defined proximally by rounded-abrupt shoulder and distally by lower elevation; without lateral and medial teeth near dactylar articulation; with sparse setae. Small chela of balaeniceps form in male only. Pereopod 3 dactylus simple and conical. 55 mm. Indo-West Pacific, WA, (S to Houtman Abrolhos Is), NT, Qld, NSW (S to Sydney); intertidal to 25 m depth, muddy sediments. Alpheus lottini Guérin-Méneville, 1829 (Figs 26r, s, 29m, n). Rostrum acute, twice as long as broad at base, flattened proximally, separated from orbital hoods by deep and narrow grooves; orbital hoods with converging acute teeth, one-third length of rostrum; rostrum-hood margins short and straight. Large chela laterally compressed; lower and upper margins straight and rounded; without ridges or grooves; without lateral and medial teeth near dactylar articulation; with only few scattered setae. Small chela not of balaeniceps form. Pereopod 3 dactylus with rounded apex. 38 mm. Indo-West Pacific, WA (S to Perth), NT, Qld, NSW (S to Sydney); subtidal in corals. Alpheus lottini is uniquely recognised by the rounded tips of the walking legs and the generally smooth orange-red carapace but again may be a complex of species. It is one of the most widespread members of the genus. It is an obligate associate of pocilloporid corals, living between their branches. Alpheus macrodactylus Ortmann, 1890. Rostrum slender, longer than broad at base, with slightly rounded carina; orbitorostral grooves shallow and disappearing at base of eyes; orbital hoods without teeth; rostrum-hood margins recessed at base of rostrum. Large chela 2.3 times as long as broad; with a definite shoulder on lower margin proximal to finger; upper margin with shoulder proximal to upper saddle and overhanging; without lateral and mesial teeth near dactylar articulation. Small chela not of balaeniceps form. Indo-West Pacific, Qld, NSW; subtidal. Alpheus novaezealandiae Miers, 1876 (Figs 26t, u, 29o). Rostrum narrow, acute, flattened between orbitorostral grooves which are overhung by base of rostrum and by medial margin of orbital hoods; orbital hoods with crest but without teeth; rostrum-hood margins concave. Large chela laterally compressed; lower margin slightly concave; upper margin with transverse slit near base of dactylus; without lateral and mesial teeth near dactylar articulation; with few scattered setae. Small chela of balaeniceps form in male only, dactylus not broadened. Dark colour. 68 mm. New Zealand, Lord Howe I., widespread in Australia; intertidal to 25 m depth, under rocks. This is a large species with deep well-defined orbitorostral grooves and narrow transverse slit on the large chela. Alpheus pacificus Dana, 1852 (Figs 26v, w, 29p, q). Rostrum acute, almost twice as long as broad, barely carinate; orbitorostral grooves shallow; orbital hoods without teeth; rostrum-hood margins concave near rostral base. Large chela laterally compressed; lower margin with strong rounded shoulder around an oblique lateral groove; upper margin with narrow groove defined proximally by overlapping shoulder; without lateral and medial teeth near dactylar articulation; with sparse setae. Small chela not of balaeniceps form but with lateral rows of crossed long setae. 40 mm. Indo-West Pacific, WA (S to Perth), NT, Qld, NSW (S to Sydney); intertidal to 20 m depth, under rocks.

105 Marine Decapod Crustacea of Southern Australia

Alpheus papillosus Banner & Banner, 1982. Rostrum conical, twice as long as wide at base, with lateral setae; carina rounded; orbitorostral grooves rounded; orbital hoods without teeth; rostrum-hood margins concave at base of rostrum. Large chela laterally compressed; lower margin with prominent square papillose shoulder next to broad notch running up laterally and more distal than upper groove; upper margin with groove defined proximally by overhanging shoulder and distally by prominent elevation; without lateral and medial teeth near dactylar articulation; with long setae near lower margin. Small chela of balaeniceps form in male only. 40 mm. WA (S to Perth), NT, Qld, NSW (S to Sydney), SA (Kangaroo I.); intertidal to 18 m depth. Alpheus parasocialis Banner & Banner, 1982 (Figs 27a, b, 29r, Pl. 8e). Rostrum acute, 3 times as long as broad at base, separated from orbital hoods by deep short grooves overhung by base of rostrum; orbital hoods with converging acute teeth, reaching half length of rostrum; rostrum- hood margins short and convex. Large chela laterally compressed; lower margin slightly concave, irregular proximally; upper margin with longitudinal groove; without lateral and medial teeth near dactylar articulation; with only few scattered setae. Small chela not of balaeniceps form. Maxilliped 3 with penultimate article not lobed on the lower margin. Dull orange and with orange-red chelipeds. 35 mm. Qld (N to Moreton Bay), NSW, Vic., Tas., WA (N to Shark Bay); intertidal to 29 m depth. Alpheus richardsoni Yaldwyn, 1971 (Figs 27c, d, 28l, 29s, t, Pl. 9b). Rostrum short and broad, smooth dorsally; orbitorostral grooves shallow or non-existent; orbital hoods without teeth; rostrum-hood margins barely concave near rostral base. Large chela laterally compressed; lower margin with rounded shoulder around an oblique lateral depression; upper margin with saddle defined proximally by rounded shoulder and distally by lower step; without lateral and medial teeth near dactylar articulation; with sparse setae; plunger a prominent cylinder. Small chela of balaeniceps form in male only, without saddle on upper margin. Deep green, banded with brown on the abdomen. 65 mm. Northern NZ, Qld (N to Moreton Bay), NSW, Vic., Tas., to north-western WA; intertidal to 24 m (one record at 412 m off NSW coast), muddy and estuarine environments. The species is commonly encountered in the muddy sediments of mangrove forests and deep muddy bays. Cohabiting pairs make elaborate burrows down to about 30 cm deep, often under a surface rock. The species was often treated as a subspecies of, or referred to as the Indo-West Pacific species A. euphrosyne. Alpheus socialis Heller, 1865 (Figs 27e, f, 29u, v, w). Rostrum acute, free section shorter than proximal part, separated from orbital hoods by deep short grooves overhung by base of rostrum; orbital hoods with converging acute teeth, almost length of rostrum; rostrum-hood margins broad and convex. Large chela laterally compressed; lower margin slightly concave, irregular proximally; upper margin with longitudinal groove; without lateral and medial teeth near dactylar articulation; with only few scattered setae. Small chela not of balaeniceps form. Maxilliped 3 with penultimate article lobed on the lower margin. Green. 35 mm. New Zealand, Lord Howe I., NSW (Sydney region), Vic., SA; intertidal to 108 m depth. Alpheus spongiarum Coutière, 1897. Rostrum broad, with carina reaching to base of eyes; orbitorostral grooves shallow; orbital hoods without teeth; rostrum-hood margins transverse. Large chela cylindrical, smooth; without lateral and medial teeth near dactylar articulation; with very few setae. Small chela not of balaeniceps form. Reddish. 18 mm. Indo-West Pacific, WA (S to Perth), NT, Qld (S to Heron I.); intertidal to 42 m depth, in sponges.

106 Caridea – shrimps

ab c d

e f g h

j k i l

m n o

q r s p

w tu v

Fig. 27. Alpheidae. Dorsal and lateral views of anterior carapace: a, b, Alpheus parasocialis. c, d, Alpheus richardsoni. e, f, Alpheus socialis. g, h, Alpheus stephensoni. i, j, Alpheus villosus. k, l, Arete dorsalis. m, Athanas dimorphus. n, Athanas granti. o, Athanas ornithorhynchus. p, q, Athanopsis australis. r, s, Betaeus australis. t, u, Metalpheus paragracilis. v, w, Synalpheus comatularum. (k–o from Banner & Banner, 1973; t, u from Banner & Banner, 1982a)

107 Marine Decapod Crustacea of Southern Australia

e f

h g k

Fig. 28. Alpheidae. Dorsal and lateral views of anterior carapace: a, b, Synalpheus fossor. c, d, Synalpheus harpagatrus. e, f, Synalpheus stimpsonii. g, h, Synalpheus streptodactylus. i, j, Synalpheus tumidomanus. Habitus: k, Synalpheus stimpsonii. l, Alpheus richardsoni.

108 Caridea – shrimps

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j l k i

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y x u v

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j’

f’ l’ h’ m’

Fig. 29. Alpheidae. Chelipeds, lateral views unless otherwise indicated (and other limbs): a, Alpheopsis trispinosa (major cheliped). b, Alpheus astrinx (major cheliped, upper). c, d, Alpheus australiensis (major and minor chelipeds). e, f, Alpheus australosulcatus (major cheliped and pereopod 2). g, h, Alpheus bunburius (major and minor chelipeds). i, Alpheus edwardsii (major cheliped). j, Alpheus hailstonei (major cheliped). k, l, Alpheus lobidens (major and minor chelipeds). m, n, Alpheus lottini (major cheliped, pereopod 3 propodus and dactylus). o, Alpheus novaezealandiae (major cheliped). p, q, Alpheus pacificus (major and minor chelipeds). r, Alpheus parasocialis (maxilliped 3). s, t, Alpheus richardsoni (major cheliped, pereopod 3 propodus and dactylus). u, v, w, Alpheus socialis (major cheliped, pereopod 3 dactylus, maxilliped 3). x, Alpheus stephensoni (major cheliped). y, Alpheus villosus (major cheliped). z, Arete dorsalis (pereopod 3). a’, Arete indicus (pereopod 3). b’, Metalpheus paragracilis (major cheliped). c’, Metalpheus paragracilis (major cheliped plunger and socket). d’, Synalpheus comatularum (minor cheliped). e, Synalpheus fossor (pereopod 3 dactylus). f’, Synalpheus harpagatrus (pereopod 3). g’, h’, i’, Synalpheus stimpsonii (major cheliped, fingers open and closed, minor cheliped). j’, Synalpheus streptodactylus (pereopod 3). k’, l’, Synalpheus tumidomanus (pereopod 3 and dactylus). m’, Alpheopsis trispinosa (pleonite 6 with articulating posterolateral plate). (c, d, g, h from Banner & Banner, 1982a; z–c’ from Banner & Banner, 1973)

109 Marine Decapod Crustacea of Southern Australia

Alpheus stephensoni Banner & Smalley, 1969 (Figs 27g, h, 29x). Rostrum acute, carinate back to almost half length of carapace; orbitorostral grooves moderate; orbital hoods close together, without teeth; rostrum-hood margins not concave. Carapace minutely pustulate, especially anteriorly. Large chela laterally compressed; lower and upper margins barely concave; without lateral and medial teeth near dactylar articulation; with setae along lower and upper margins, and pustulate (like sandpaper) all over. Small chela not of balaeniceps form but in male with fringes of opposing setae on fingers. At least 75 mm. Southern Qld, NSW (S to Sydney); intertidal to 18 m depth, muddy sediments. The rough carapace distinguishes this species from all others in southern Australia. Alpheus strenuus Dana, 1852. Rostrum acute, more than twice as long as broad, smooth dorsally; orbitorostral grooves shallow (in A. s. cremnus with abrupt margins to rostrum and hoods); orbital hoods without teeth; rostrum-hood margins concave near rostral base. Large chela laterally compressed; lower margin with strong rounded shoulder around an oblique lateral depression; upper margin with groove defined proximally by overlapping shoulder and distally by step; without lateral and mesial teeth near dactylar articulation; with sparse setae. Small chela of balaeniceps form in both sexes. 60 mm (95 mm reported). A. s. strenuus:Indo- West Pacific, WA (S to Perth), NT, Qld, NSW (S to Sydney); lower intertidal on sandy beaches. A. s. cremnus Banner & Banner, 1982: NSW, rare in WA, Qld and NT; muddy and estuarine environments. Banner & Banner (1982a) described a southern Australian subspecies on the basis of more defined orbitorostral grooves. The two subspecies overlap geographically but apparently not ecologically. Alpheus sulcatus Kingsley, 1878. Rostrum 1–2 times as long as wide at base, bearing stiff setae laterally, free section shorter than orbitorostral grooves which are overhung by base of rostrum and sharply delimited from orbital hoods; orbital hoods without teeth; rostrum- hood margins transverse. Large chela laterally compressed; lower margin slightly concave, irregular proximally; upper margin with longitudinal ridge and tubercles; without lateral and mesial teeth near dactylar articulation; with numerous scattered setae. Small chela not of balaeniceps form. Pereopod 2 with first carpal segment slightly longer than second. Green. 58 mm. Indo-West Pacific, eastern Pacific, eastern Atlantic, scattered throughout Australia except Tas.; intertidal to 24 m depth. Alpheus tasmanicus Banner & Banner, 1982. Rostrum acute, almost twice as long as broad, slightly carinate; orbitorostral grooves shallow; orbital hoods without teeth; rostrum-hood margins weakly concave. Large chela laterally compressed; lower margin with strong rounded shoulder; upper margin with saddle defined proximally by steep shoulder; without lateral and medial teeth near dactylar articulation; with sparse setae. Small chela not of balaeniceps form. 22 mm. Tas. (known only from the holotype). Alpheus villosus (Olivier, 1811) (Fig 27i, j, 29y, Pl. 9c, d). Entire body covered with short stiff setae interspersed with longer ones. Rostrum about twice as long as broad at base, carinate and with mid-dorsal tooth behind eyes, separated from orbital hoods by broad grooves; orbital hoods with small triangular teeth overhanging margin, one-quarter as long as rostrum; rostrum-hood margins with intermediate minute tooth. Large chela covered with long stiff setae, especially distally and medially; round in section; lower margin not concave; upper margin convex; without strong acute lateral and medial teeth near dactylar articulation; propodus with weak longitudinal ridges plus small tubercles. Small chela not of balaeniceps form. Pale orange, spotted with red, and sometimes bright orange on the chela, eyes are never strongly pigmented. 67 mm. South Africa, Philippines, Australia (except east coast); intertidal to 43 m depth; reported from corals. Alpheus villosus is the only very hairy species of shrimp in southern Australia and is therefore immediately recognisable. Banner & Banner (1982a) reported two morphological forms, one widespread, and the other along the southern Australian coast (Twofold Bay, NSW, to Perth, WA).

110 Caridea – shrimps

Arete Stimpson, 1860 This genus comprises only of two species which both occur in southern Australia. Diagnosis. Pereopods 1–2 or more with epipods. Pleuron of abdominal somite 6 with an articulating posterior triangular plate. Rostrum slender, pointed in lateral view. Eyes entirely or partially visible in dorsal view and concealed in lateral view only by anterior tooth of carapace. Anteroventral corner of carapace rounded, rarely produced. Pereopod 2 carpus with 4 articles.

Key to southern Australian species of Arete 1. Broad rostrum (1–2 times as long as broad, almost twice as long as eyestalks), pereopod 3 merus 3–4 times as long as wide, its lower distal tooth weak...... Arete dorsalis — Long rostrum (2.0–2.5 times as long as broad, more than twice as long as eyestalks), pereopod 3 merus narrow (4–5 times as long as wide) and lower distal tooth on merus developed ...... Arete indicus Arete dorsalis (Stimpson, 1861) (Figs 27k, l, 29z). Rostrum broad (1–2 times as long as broad), almost twice as long as eyestalks. Anterior margin of carapace without supraorbital and infraorbital teeth, extraorbital tooth reaching at least to middle of eye. Chelipeds slightly asymmetrical, sexually dimorphic, carried forward; propodus stout, fingers variable. Pereopod 3 merus 3–4 times as long as wide, its lower distal tooth weak. Orange-red, greenish-brown or black. Indo-West Pacific, Lord Howe I., Norfolk I., Qld (Great Barrier Reef), WA (only Rottnest I.); intertidal reef flat to 3 m depth. This species of variable colour is often associated with echinoids, typically between the spines of the sea urchins Heterocentrotus mammillatus and Anthocidaris crassispina and ophiuroids. The species may be a complex of similar species (Davie, 2002). Arete indicus Coutière, 1903 (Fig. 29a'). Rostrum triangular, lateral margins slightly depressed, tip reaching beyond distal end of article 2 of antennule peduncle. Anterior margin of carapace without supraorbital spine and infraorbital teeth, extraorbital tooth reaching to end of eye. Chelipeds slightly asymmetrical, carried forward. Pereopod 3 merus 4–5 times as long as wide, developed lower distal tooth. Colour variable. 15 mm. Indo-West Pacific, Qld, WA (S to Cape Naturaliste). Like the morphologically similar A. dorsalis, this species can be found on coral reefs associated with echinoderms, typically between the spines or underneath sea urchins Echinometra mathaei.

Athanas Leach, 1814 Banner & Banner (1973) recognised 11 species throughout Australia but only six or seven occur on the southern coasts. Diagnosis. Pereopods 1–2 or more with epipods. Pleuron of abdominal somite 6 with an articulating posterior triangular plate. Rostrum well developed, usually with 3 pairs of orbital processes. Carapace not covering eyes. Chelipeds asymmetrical and sexually dimorphic, chelate and swollen in male, simple and slender in female; large chela of male carried extended or flexed against expanded merus, fingers without plunger and socket. Pereopod 2 carpus with 5 articles.

Key to southern Australian species of Athanas 1. Pereopod 3 dactylus simple ...... 2 — Pereopod 3 dactylus bifid ...... 3 2. Carpus of female cheliped 1–2 times as long as propodus ...... Athanas dimorphus — Carpus of female cheliped shorter than propodus ...... Athanas hasswelli

111 Marine Decapod Crustacea of Southern Australia

3. Chelipeds carried forward ...... Athanas granti — Chelipeds carried flexed ...... 4 4. Large cheliped propodus with a pronounced lobe on lower margin; small cheliped with fingers dorsoventrally flattened ...... Athanas ornithorhynchus — Large cheliped propodus with simple lower margin; small cheliped with fingers not dorsoventrally flattened ...... Athanas parvus Athanas dimorphus Ortmann, 1894 (Fig. 27m). Rostrum reaching to or beyond second article of antennule. Carapace without supraorbital tooth, extraorbital tooth acute, infraorbital tooth slight, pterygostomial angle subacute. Chelipeds sexually dimorphic, symmetrical and flexed at merus-carpus in both sexes. Cheliped in adult male robust and inflated; merus excavated to accommodate chela when flexed; propodus circular in section, thickest at midpoint, 3 times as long as greatest width; dactylus 0.4 length of propodus. Chelipeds small and slender in female. Red or tan and white striped. 15 mm. Indo-West Pacific, WA (S to Perth), NT, Qld, NSW, (S to Sydney); intertidal pools under rocks or coral. Athanas granti Coutière, 1908 (Fig. 27n). Rostrum reaching past middle of third article of antennule. Carapace without supraorbital tooth, extraorbital tooth acute and extending beyond middle of eyestalk, infraorbital tooth small and rounded. Chelipeds not sexually dimorphic, almost symmetrical, and carried extended but capable of flexion; propodus with constriction proximally, fixed finger with convex cutting edge. Deep red with a pale dorsal stripe. Central NSW, Vic., Tas., SA, WA (N to Perth); intertidal with coral or an echinoid. Athanas hasswelli Coutière, 1908. This species, reported only from the type at Sydney and from SA by Hale (1927) is a mystery. It is differentiated from A. dimorphus on only one character which could be unreliable [see key and Banner & Banner (1973)]. Coutière clearly wanted to honour William Haswell with this species but misspelled his name! Athanas ornithorhynchus Banner & Banner, 1973 (Fig. 27o). Rostrum narrowly triangular and acute, reaching well beyond peduncle of antennule, with dorsal carina. Carapace without supraorbital tooth, extraorbital tooth large, acute and well above eyestalk, infraorbital tooth obsolete. Chelipeds asymmetrical, larger as long as carapace; merus excavated to accommodate propodus; propodus with lobe on lower margin; fingers rounded and crossing. Smaller cheliped with dorsoventrally flattened fingers (hence the specific name, the genus of the platy- pus). 11 mm. NT, WA (S to Cockburn Sound); to 18 m depth. Athanas parvus de Man, 1910. Rostrum reaching end of second article of antennule, with strong carina. Carapace without supraorbital tooth, extraorbital tooth and infraorbital tooth large, extending beyond eyestalk. Chelipeds almost symmetrical, sexually dimorphic. Male cheliped stout, propodus 3.5 times as long as broad, folding back on to expanded and excavated merus; dactylus crescent-shaped. Female chelipeds feeble. Pale blue with red spots and a dorsal pale stripe. 13 mm. Indo-West Pacific, WA (S to Rottnest I.), NT, Qld (to southern Great Barrier Reef); dead coral and sponges, intertidal under rocks on sandy beaches, to 130 m depth. Australian records of this species have been as Athanas sibogae de Man, 1910 (Davie, 2002)

Athanopsis Coutière, 1897 The genus is distinguished by the unusual rostrum, ever so subtly rounded in lateral view at the apex. The Australian species is only the second described (Banner & Banner, 1982a). Diagnosis. Pereopods 1–2 or more with epipods. Pleuron of abdominal somite 6 with an articulating posterior triangular plate. Rostrum well developed, with a broad vertical lamella ventrally, its tip broadly rounded, with supraorbital processes only. Carapace barely covering eyes. Chelipeds asymmetrical and sexually dimorphic, chelate and swollen in male, (unknown in female); large chela of male flexed against expanded merus, fingers without plunger and socket.

112 Caridea – shrimps

Athanopsis australis Banner & Banner, 1982 (Fig. 27p, q, Pl. 9e). Rostrum with tip developed as a laterally compressed lamella, a continuation of an obsolete dorsal carina, tip rounded in lateral view. Chela of male folded back against laterally expanded merus. Colour in life is translucent with red digestive gland apparent. 16 mm. Vic.; reef and sandy sediments, 5–8 m depth. The species is known from only four specimens. The species was listed as ‘vulnerable’ under IUCN criteria by O’Hara (2002).

Betaeus Dana, 1852 There is only one species in Australia. Diagnosis. Rostrum absent, front depressed. Carapace covering eyes, at least dorsally. Pleuron of abdominal somite 6 with an articulating posterior triangular plate. Chelipeds asymmetrical; large chela carried extended, fingers without plunger and socket. Pereopods with epipods. Betaeus australis Stimpson, 1860 (Fig. 27r, s). Anterior margin of carapace broadly rounded and covering eyes in dorsal view, without rostrum or orbital teeth. Dorsally purple or red, paler elsewhere, chela reddish-purple. 35 mm. Southern NSW, Vic., SA.; intertidally under rocks.

Metalpheus Coutière, 1897 Diagnosis. Body not usually compressed from side to side. Rostrum distinct, acute laterally. Carapace without high carina throughout length of dorsal midline. Eyes concealed from dorsal and all but anteroventral views. Mandible with palp and molar process, incisor process usually expanded. Abdominal somite 6 without articulating posterior triangular plate, telson not terminating in triangular tooth. Pereopods 1 dissimilar and unequal, carried forward with movable finger dorsal or lateral, not ventral. Pereopod 2 carpus with 5 articles. Pereopods 1–3 with epipods. Metalpheus paragracilis (Coutière, 1897) (Figs 27t, u, 29b', c'). Rostrum acute, reaching to distal end of article 1 of antennule peduncle; orbitorostral grooves shallow, extending only to base of eyes; orbital hoods subacute; rostrum-hood margins deeply indented. Large chela laterally compressed; lower margin with shoulder abrupt; upper margin with slight longitudinal ridge; without lateral and medial teeth near dactylar articulation; fingers with plunger and socket. Indo-West Pacific, Qld, WA; coral reef to 20 m depth.

Synalpheus Bate, 1888 Synalpheus and Alpheus are similar and best separated on the pterygostomial angle (the lower anterior corner of the carapace which wraps around the front under the antennae). It is produced and usually acute in Synalpheus while the whole anterior margin of the carapace is evenly rounded in Alpheus. The presence of pereopodal epipods, which can be seen at the base of the legs under the carapace, is a certain indicator of Alpheus. If in doubt, species without orbital processes or with a sculptured larger cheliped are Alpheus rather than Synalpheus. The converse is not true. Many species of Synalpheus are symbionts with crinoids, corals or sponges. It is suspected that the shrimp may feed on the mucus stream generated by its host but little is certain. Some species occur at high densities inside the cavities of sponges and may be trapped there. Some species are suspected of displaying a form of sociality (Duffy & Macdonald, 1999). Specimens are often colourful but as colour may mimic that of the host it can be an unreliable specific character. Banner & Banner (1975) listed, keyed and grouped more than 30 species of Australian Synalpheus. The key refers to a ‘orbitorostral process’ which can be difficult to see without lifting the rostrum and viewing from the front. It is a strong lobe protruding ventrally from the base of the rostrum between and in front of the eyes.

113 Marine Decapod Crustacea of Southern Australia

Diagnosis. Pereopods without epipods. Pleuron of abdominal somite 6 without an articulating posterior triangular plate. Rostrum well developed. Carapace covering eyes completely in dorsal and lateral views, usually with orbital processes; pterygostomial angle produced. Chelipeds markedly asymmetrical, larger one swollen, carried extended, dactylus with plunger fitting into socket at base of fixed finger. Smaller cheliped with fingers simple.

Key to southern Australian species of Synalpheus 1. Orbitorostral process absent (base of rostrum smooth ventrally) ...... 2 — Orbitorostral process present (a ventral lobe at the base of the rostrum) ...... 3 2. Small chela with dactylus sickle-shaped, longer than fixed finger, fingers crossing; pereopod 3 dactylus with lower claw a rudimentary angle ...... Synalpheus comatularum — Small chela with dactylus almost straight, fingers not crossing; pereopod 3 dactylus with lower claw a strong spine ...... Synalpheus stimpsonii 3. Small chela with dactylus broadened at least at base, subspatulate, fingers sometimes with teeth ...... 4 — Small chela with dactylus conical, fingers terminating in single tooth ...... 7 4. Pereopod 3 merus having a conspicuous row of spiniform setae along distal third; small cheliped fingers with apical teeth ...... 5 — Pereopod 3 merus unarmed; small cheliped fingers with single apical tooth ...... 6 5. Large chela propodus upper margin ending in acute tooth ...... Synalpheus harpagatrus — Large chela propodus upper margin ending in 2 conical, obtuse tubercules ...... Synalpheus bituberculatus 6. Telson with elongate dorsolateral spiniform setae; antennal articles with ventrolateral tooth much longer than stylocerite ...... Synalpheus theano — Telson with stout dorsolateral spiniform setae; antennal articles with ventrolateral tooth not far over-reaching stylocerite ...... Synalpheus neptunus 7. Pereopod 3 dactylus with 3 claws, upper one smallest ...... 8 — Pereopod 3 dactylus with 2 claws ...... 9 8. Pereopod 3 merus with a row of spiniform setae ...... Synalpheus nilandensis — Pereopod 3 merus unarmed ...... Synalpheus fossor 9. Small chela dactylus with definite longitudinal row of setae on crest or inner face; fingers may bear other setae but not in patterned row ...... Synalpheus lophodactylus — Small chela dactylus without definite rows or patches of setae ...... 10 10. Pereopod 3 merus with 2 or more spiniform setae on lower margin ...... 11 — Pereopod 3 merus without spiniform setae ...... 13 11. Pereopod 3 dactylus having upper claw half size of lower unguis . . . . . Synalpheus neomeris — Pereopod 3 dactylus having upper claw little smaller than lower claw ...... 12 12. Pereopod 3 having claws one-third to one-quarter length of dactylus ...... Synalpheus streptodactylus — Pereopod 3 having claws less than one-sixth length of dactylus ...... Synalpheus iocosta 13. Maxilliped 3 with numerous elongate spiniform setae on inner face of terminal article ...... Synalpheus echinus — Maxilliped 3 with only setae on inner face of terminal article and circlet of short apical spiniform setae ...... Synalpheus tumidomanus

114 Caridea – shrimps

Synalpheus bituberculatus de Man, 1910. Rostrum with tip upturned, rounded to acute, sides parallel to triangular, tip not reaching distal end of article 1 of antennule peduncle. Orbitorostral process present. Large chela propodus upper margin ending in 2 conical, obtuse tubercules. Small chela 2.8 times as long as broad, without spiniform setae on extensor margin of movable finger, ending in 2 acute teeth. Pereopod 3 merus inferior margin armed with 6–9 spiniform setae; dactylus with 2 claws. Telson with acute dorsolateral spiniform setae. Indo- West Pacific, NT, Qld, NSW, WA; coral reef, sponges commonly in pairs, to 36 m depth. Synalpheus comatularum (Haswell, 1882) (Figs 27v, w, 29d'). Rostrum 2–3 times as long as, and wider than, orbital processes, depressed, with sharp ridge running on to carapace; deep grooves between. Orbitorostral process absent. Large chela propodus upper margin ending in sharp tooth. Small chela with dactylus sickle-shaped, longer than fixed finger, fingers crossing. Pereopod 3 merus with acute distal tooth; dactylus with lower claw a rudimentary angle. Telson with strong dorsolateral spiniform setae. WA (Broome to Busselton), Qld, NSW; associated with crinoids. The strong carinae, laterally and running back from the rostrum, and the sickle-shaped dactylus on the smaller cheliped are characteristic of this species (but see S. stimpsonii). Synalpheus echinus Banner & Banner, 1975. Rostrum clearly longer and narrower than orbital processes, not depressed, with obtuse ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus upper margin ending in 2 rounded projections. Small chela fingers simple, each with simple apex. Pereopod 3 merus without spiniform setae on lower margin, no distal tooth; dactylus with 2 distal claws (about 0.2 length of dactylus). Telson with stout dorsolateral spiniform setae. Maxilliped 3 with numerous elongate spiniform setae on inner face of terminal article. 28 mm. WA (Perth, Rottnest I.); 0–3 m depth. Synalpheus fossor (Paul’son, 1875) (Figs 28a, b, 29e'). Rostrum longer and narrower than orbital processes, both with short apical setae, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus moderately swollen, upper margin ending in obtuse tooth. Small chela fingers each with simple tooth at tip. Pereopod 3 merus without spiniform setae on lower margin, no distal tooth; dactylus with 3 claws, upper one obtuse and broad, separate from next by U, middle and lower claws finer, separate by V. Telson with short dorsolateral spiniform setae. 25 mm. Indo-West Pacific, all coasts of Australia; to 50 m depth, in coral and sponges. This species was called S. bakeri by Hale (1927). Synalpheus harpagatrus Banner & Banner, 1975 (Fig. 28c, d, 29f'). Rostrum 1.5 times as long as, and narrower than, orbital processes, not depressed, without ridge running on to carapace; shallow grooves between. Orbitorostral process present. Large chela propodus upper margin ending in small acute tooth. Small chela with fingers broadened, weakly toothed at tips and opposing. Pereopod 3 merus with spiniform setae on lower margin, no distal tooth; dactylus with 2 claws, upper one broader but as long as lower one. Telson with moderate dorsolateral spiniform setae. 38 mm. SA, WA (Geraldton to Fremantle), NT; 0–17 m depth, with sponges. Synalpheus iocosta de Man, 1909. Rostrum scarcely longer and narrower than orbital processes, latter with concave inner and straight outer margins, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus upper margin ending in sharp tooth. Small chela fingers simple, each with simple apex. Pereopod 3 merus with spiniform setae on lower distal margin, no distal tooth; dactylus with 2 minute claws (<0.15 length of dactylus), upper one thinner than lower one. Telson with stout dorsolateral spiniform setae, all in distal half. 12 mm. Indonesia, southern WA, Qld (Gulf of Carpentaria); to 130 m depth, in sponges and bryozoans.

115 Marine Decapod Crustacea of Southern Australia

Synalpheus lophodactylus Coutière, 1908. Rostrum scarcely longer and much narrower than orbital processes, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus moderately swollen, upper margin ending in upturned tooth. Small chela fingers each with simple tooth at tip; dactylus setose on upper- inner surface. Pereopod 3 merus without spiniform setae on lower margin, no distal tooth; dactylus with 2 claws, scarcely curved. Telson with stout dorsolateral spiniform setae. 22 mm. Indian Ocean, central Pacific, WA (S to Perth), Qld; 1–2 m depth, in sponges. Synalpheus neomeris (de Man, 1897). Rostrum scarcely longer and narrower than orbital processes, all with apical setae, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus moderately swollen, upper margin ending in sharp tooth. Small chela fingers simple, each with simple apex. Pereopod 3 merus with spiniform setae on lower margin, no distal tooth; dactylus with 2 claws, upper one much smaller than lower one. Telson with stout dorsolateral spiniform setae, close together and distal. 25 mm. Indo-West Pacific, WA,NT, Qld, NSW; to 250 m depth, often with alcyonarians and sometimes sponges. Synalpheus neptunus Dana, 1852. Rostrum as long as, and narrower than, orbital processes, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus swollen, upper margin ending in rounded prominence. Small chela with dactylus broadened laterally; fingers each with slender tooth at tip. Pereopod 3 merus without spiniform setae on lower margin, no distal tooth; dactylus with 2 claws, upper one broader at base. Telson with stout dorsolateral spiniform setae. Two subspecies: S. n. neptunus. 26 mm. Indo-West Pacific, WA (S to Perth), NT, Qld, NSW (S to Sydney); to 70 m depth, in corals and sponges. S. n. germanus Banner & Banner, 1975. A subspecies based entirely on immature males and differing only slightly from the nominate subspecies. 15 mm. WA (Perth and Rottnest I.); sponges. Synalpheus nilandensis Coutière, 1908. Rostrum not overreaching distal end of article 1 of antennule peduncle. Orbitorostral process present. Large chela propodus upper margin ending in acute tooth. Small chela 3.5 times as long as broad, without spiniform setae on extensor margin of movable finger; fingers terminating in single distal tooth. Pereopod 3 merus inferior margin armed with 4 strong spiniform setae; dactylus with 3 claws. Telson with short, acute dorsolateral spiniform setae. Indo-West Pacific, WA; 18–135 m depth. Synalpheus stimpsonii (De Man, 1888) (Figs 28e, f, k, 29g', h', i'). Rostrum 2–3 times as long as, and wider than, orbital processes, depressed, with rounded ridge running on to carapace; grooves between. Orbitorostral process absent. Large chela propodus upper margin ending in tooth. Small chela with dactylus as long as fixed finger, fingers not crossing. Pereopod 3 merus with acute distal tooth; dactylus with lower claw acute. Telson with weak dorsolateral spiniform setae. 35 mm. Indo-West Pacific, WA (S to Perth), NT, Qld, NSW (S to Sydney); intertidal to 155 m depth, associated with crinoids and occasionally alcyonarians and dead coral. This species, or species complex (Davie, 2002), and S. comatularum differ from all others in the absence of the orbitorostral process and strong ridges and grooves on the anterior carapace. The rostrum is broader and shorter than in S. comatularum and the small cheliped lacks a sickle-shaped dactylus. Synalpheus streptodactylus Coutière, 1905 (Figs 28g, h, 29j'). Rostrum scarcely longer and narrower than orbital processes, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus upper margin ending in sharp tooth. Small chela fingers simple, each with simple apex. Pereopod 3 merus with spiniform setae on lower distal margin, no distal tooth; dactylus with 2 claws, upper one thinner than lower one. Telson with stout dorsolateral spiniform setae, all in distal half. 20 mm. Indo-West Pacific, all coasts of Australia; to 128 m depth, in sponges.

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Synalpheus theano de Man, 1910. Rostrum as long as, and narrower than, orbital processes, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela propodus swollen, upper margin ending in obtuse tooth. Small chela with dactylus broadened laterally; fingers each with slender tooth at tip. Pereopod 3 merus without spiniform setae on lower margin, no distal tooth; dactylus with 2 very similar claws. Telson with elongate dorsolateral spiniform setae. 22 mm. Indonesia, (Singapore), WA (S to Perth), NT, Qld, NSW (S to Sydney); to 62 m depth, in dead coral and sponges. Synalpheus tumidomanus (Paul’son, 1875) (Figs 28i, j, 29k', l', Pl. 9f). Rostrum clearly longer and slightly narrower than orbital processes, not depressed, without ridge running on to carapace; very shallow grooves between. Orbitorostral process present. Large chela swollen, propodus upper margin ending in variable projection. Small chela fingers simple, each with simple apex. Pereopod 3 merus without spiniform setae on lower margin, no distal tooth; dactylus with 2 distal claws, upper one longer than lower. Telson with stout dorsolateral spiniform setae. Maxilliped 3 with circlet of short apical spiniform setae and setae on inner face of terminal article. 25 mm. Eastern Mediterranean, Indo-West Pacific, all coasts of Australia; 0–148 m depth, in macroalgae, coral and sponges. Hale (1927) referred to this species as S. maccullochi and it has several other junior synonyms which may one day be revived as the species is probably several. It is one of the more commonly collected species on subtidal reefs in debris at the base of macroalgae. The shining pale-green cheliped, round in cross-section is diagnostic.

References Banner, D.M., & Banner, A.H. 1973. The alpheid shrimp of Australia. Part I: the lower genera. Records of the Australian Museum 28: 291–382. Banner, D.M., & Banner, A.H. 1975. The alpheid shrimp of Australia. Part 2: the genus Synalpheus. Records of the Australian Museum 29: 267–389. Banner, D.M., & Banner, A.H. 1982a. The alpheid shrimp of Australia. Part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum 34: 1–358. Banner, D.M., & Banner, A.H. 1982b. The alpheid shrimp of Australia. Supplement I. Records of the Australian Museum 34: 359–362. Bruce, A.J. 1994. Alpheus fenneri sp. nov. and A. williamsi sp. nov., two new Indo-West Pacific alpheid shrimps of the brevirostris group. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11: 15–28. Christoffersen, M.L. 1987. Phylogenetic relationships of hippolytid genera, with an assignment of new families for the Crangonoidea and Alpheoidea (Crustacea, Decapoda, Caridea). Cladistics 3: 348–362. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Duffy, J.E., & Macdonald, K.S. 1999. Colony structure of the social snapping shrimp Synalpheus filidigi- tatus in Belize. Journal of Crustacean Biology 19: 283–292. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. O’Hara, T. 2002. Endemism, rarity and vulnerability of marine species along a temperate coastline. Invertebrate Systematics 16: 671–684.

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Hippolytidae Bate, 1888 The Hippolytidae are one of the richest families with about 280 species in 41 genera. The rostrum is usually well developed and the carpus of pereopod 2 subdivided into a few or many articles. Most species in southern Australia inhabit algal environments on rocky shores or seagrasses. As usual there are many more species in tropical Australia than along southern shores. An early catalogue of world species (Holthuis, 1947) revised the names of some Australian species. Kensley et al. (1987) described some deep-water Australian species. Christoffersen (1987) proposed a phylogenetic analysis of hippolytid genera, concluding that the family as presently conceived is polyphyletic. He introduced new family names for some groups of genera but these have not been adopted by crustacean taxonomists. The most comprehensive treatment of the family as a whole (Chace, 1997) provided a key to genera and lists of species. The species in southern Australia are relatively easy to distinguish on the basis of the rostrum alone but keying through the genera and species to be certain of generic placement is more difficult. The key, based on that in Holthuis (1993), relies on two characters difficult to see. The arthrobranchs are gills at the bases of the legs and when present are small triangular gills directed dorsally between the pleurobranchs, larger gills which point down and are always present. The mandible is difficult to dissect out from among all the other mouthparts and it is often necessary to remove it completely to see whether the palp and incisor process are present; the incisor process is very thin and both are smaller than the molar process. Unidentified species of Eualus and Lebbeus have been identified using Holthuis’s (1993) key from collections from the Tasmanian Seamounts.

Diagnosis. Carapace with or without spines. Rostrum various, usually prominent. Eyes visible. Pereopods 1 and 2 chelate, first stouter but shorter than second. Pereopod 2 carpus divided.

Key to southern Australian genera and species of Hippolytidae 1. Pereopods 1–4 with arthrobranchs ...... 2 — Pereopods 1–4 without arthrobranchs ...... 4 2. Abdominal segment 6 without posterolateral articulated plate; deep-water ...... Merhippolyte chacei — Abdominal segment 6 with posterolateral articulated plate; coastal ...... 3 3. Mandible with incisor process ...... Saron marmoratus — Mandible without incisor process ...... Nauticaris marionis 4. Mandible with palp ...... 5 — Mandible without palp ...... 10 5. Carapace without supraorbital spine ...... 6 — Carapace with supraorbital spine ...... 9 6. Maxilliped 3 without exopod ...... Leontocaris ... 7 — Maxilliped 3 with exopod ...... Eualus sp. 7. Mandibular palp 2-articulate ...... Leontocaris bulga — Mandibular palp 1-articulate ...... 8 8. Exopod of uropod distolaterally serrate ...... Leontocaris yarramundi — Exopod of uropod distolaterally entire ...... Leontocaris amplectipes 9. Mandibular palp of 3 articles ...... Alope orientalis — Mandibular palp of 2 articles ...... Lebbeus spp.

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10. Mandible with incisor process ...... 11 — Mandible without incisor process ...... 13 11. Pereopod 2 carpus of 6–7 articles ...... Thor paschalis — Pereopod 2 carpus of 3 articles ...... 12 12. Rostrum narrow, with 3 dorsal spines and 1–2 ventral spines ...... Hippolyte caradina — Rostrum broad basally, unarmed dorsally and with 2–3 notches ventrally ...... Hippolyte australiensis 13. Body stout; maxilliped 3 with exopod; rostrum a deep blade of various shape separating the antennae ...... Latreutes compressus — Body slender; maxilliped 3 without exopod; rostrum a thin dagger as long or longer than carapace ...... 14 14. Body and limbs hairy ...... Tozeuma tomentosum — Body and limbs smooth ...... 15 15. Rostrum about as long as carapace, with 4–7 ventral teeth ...... Tozeuma pavoninum — Rostrum longer than carapace, with numerous ventral teeth ...... 16 16. Abdomen strongly curved but not geniculate; pereopod 2 with carpus having first article as long as third ...... Tozeuma elongatum — Abdomen geniculate; pereopod 2 with carpus having first article shorter than third ...... Tozeuma kimberi

Alope White, 1847 There are only two species of this genus, one widespread throughout the Indo-West Pacific and Australia, and the other in New Zealand. Both have a relatively weak rostrum. Diagnosis. Rostrum a short spike. Supraorbital spine present. Pereopod 2 carpus with 7 articles. Mandible with 3-articled palp, without incisor process. Maxilliped 3 with exopod. Pereopods without arthrobranchs. Abdominal somite 6 without articulating posterolateral corner. Alope orientalis (de Man, 1890) (Figs 30a, 32b, Pl. 9g). Rostrum little longer than eyestalk, with 4–7 teeth on upper margin; with supraorbital spine. 27 mm. Indo-West Pacific; throughout Australia; 0–46 m depth. This species, immediately recognisable by the arrangement of rostrum and supraorbital spine, was figured as A. australis by Hale (1927).

Eualus Thallwitz, 1892 Eualus comprises 32 species, with a large number recorded from the North Pacific Ocean. Only one species of Eualus, as yet undescribed, has been recorded from Australian waters. Diagnosis. Rostrum usually well-developed. Carapace smooth without supraorbital spine. Antennal and pterygostomial spine present. Pereopod 2 carpus with 7–8 articles. Mandible with 2-articled palp, with an incisor process. Maxilliped 3 with exopod. Pereopods without arthrobranchs. Abdominal 6 without articulating posterolateral corner. Eualus sp. Rostrum with 4 dorsal and 3–4 ventral teeth. Carapace not carinate with 2 postorbital teeth. Abdominal somites 1–3 with pleura rounded; somites 4–6 acute posteroventrally with spines. Pereopods 1–3 with an epipod. Southern Tasmanian Seamounts; 1300–1310 m depth.

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Fig. 30. Hippolytidae. a, Alope orientalis, b, Hippolyte australiensis, c, Latreutes compressus (female).

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Hippolyte Leach, 1814 Hippolyte was revised as part of a study of the species of the East Atlantic Ocean and the Mediterranean Sea (d’Udekem d’Acoz, 1996). This author listed 30 described species worldwide. Hippolyte ignobilis Kinahan, 1856 was described from Port Phillip Bay, Vic., but remains doubtful; the name has not been used since. Diagnosis. Rostrum variable. Supraorbital spine present. Pereopod 2 carpus with 3 articles. Mandible without palp, with incisor process. Maxilliped 3 with exopod. Pereopods without arthrobranchs. Abdominal somite 6 without articulating posterolateral corner. Hippolyte australiensis (Stimpson, 1860) (Fig. 30b, Pl. 10a, b). Rostrum prominent, broad at base, without dorsal teeth and with 2–6 ventral teeth. 25 mm. Southern NSW,Vic., Tas., SA, WA (N to central coast); 0–124 m depth. This is a very common species in shallow water algae in southern Australia whose colour varies with the algae it inhabits. Holthuis (1947) referred to it as H. ventricosa Milne Edwards, 1837, an Indo-West Pacific species found on drifting algae but the latter usually has dorsal rostral teeth. D’Udekem d’Acoz (1996) returned Stimpson’s name to specific status, rediagnosed it, and later redescribed it in detail (d’Udekem d’Acoz, 2001). He suggested that Hale’s (1927) figure of H. australiensis represents an undescribed species from southern Australia. Hippolyte caradina Holthuis, 1947 (Fig. 32c). Rostrum very slender, with 2–3 dorsal teeth, with 1–2 ventral teeth. 12 mm. Southern NSW, Vic., Tas., SA, WA (N to central coast); 0–82 m depth. This species from algal habitats was figured as H. tenuirostris by Hale (1927).

Latreutes Stimpson, 1860 The distinctive deep rostrum, often elevated with the carapace well above the eyes, is distinctive for this genus of almost 20 species. Diagnosis. Rostrum deep, variable and sexually dimorphic. Supraorbital spine absent. Mandible without palp, without incisor process. Pereopod 2 carpus with 3 articles. Maxilliped 3 with exopod. Pereopods without arthrobranchs. Abdominal somite 6 without articulating posterolateral corner. Latreutes compressus (Stimpson, 1860). (Figs 30c, 32d, Pl. 10c, d). Rostrum variable, with middorsal spine at base, extending well above eye and as a deep blade between eyes anteriorly, sometimes toothed distally; not so deep in male. 20 mm. Southern NSW, Tas., Vic., SA, to central WA; 0–56 m depth. The shape of the distinctive rostrum is variable, much deeper in females than males. The colour, green or yellowish, matches that of the algae or seagrass in which it is found. This species was figured as L. truncifrons by Hale (1927).

Lebbeus White, 1847 Lebbeus comprises over 40 species but only one of at least three in southern Australia is described. Two undescribed species were recorded from the Tasmanian Seamounts in 1100–1300 m depth by Poore et al. (1998). Diagnosis. Rostrum variable, usually a toothed blade. Carapace smooth with single supraorbital spine. Mandible with 2-articled palp, with incisor process. Pereopod 2 carpus with 7 articles. Maxilliped 3 without exopod. Pereopods without arthrobranchs. Abdominal somite 6 without articulating posterolateral corner. Lebbeus yaldwyni Kensley, Tranter & Griffin, 1987 (Fig. 32e). Carapace with a high dentate crest running on to short, variable rostrum. Pereopods 1–2 with epipod. 12 mm. NSW; 550–690 m depth.

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Fig. 31. Hippolytidae. a, Leontocaris amplectipes (from Bruce, 1990). b, Leontocaris bulga (anterior region). c, Leontocaris yarramundi. (b, c from Taylor & Poore, 1998)

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Leontocaris Stebbing, 1905 Leontocaris is a genus of seven deep-water slope and seamount species. All are characterised by the asymmetrical and folded first two pairs of pereopods. All species of Leontocaris possess a tympanum or thinning of the exoskeleton on the inner surface of the propodus of pereopod 2. The size of this varies between species and may play a part in extension of the folded limb (Taylor & Poore, 1998). A phylogenetic analysis of the world species included only the first named of the Australian species (Fransen, 2001). Diagnosis. Rostrum prominent, toothed. Mandible with incisor and 1- or 2-articled palp. Maxilliped 3 without exopod or epipod. Epipods present on maxilliped 1 and 2. Supraorbital spine absent. Pereopod 1 slender, carpus elongate. Pereopod 2 strongly asymmetrical, folded and extendable, carpus 4-articled. Major pereopod 2 with elongate and robust propodus and enlarged, chopper-shaped dactylus. Pereopods without arthrobranchs. Maxillipeds 2–3 and pereopods 1–4 with pleurobranchs. Leontocaris amplectipes Bruce, 1990 (Fig. 31a). Rostrum 0.58–0.85 of carapace length, shorter than antennular peduncle. Rostrum and carapace with 12–18 dorsal teeth (including 2–3 in epigastric region), and 2–4 ventral teeth. Mandibular palp of 1 article. Pereopod 3 with dactylus about 0.5 of propodus. Abdominal somite 5 pleuron rounded, unarmed. cl. 5.0–9.6 mm. Eastern Vic., Tas. Seamounts; 750–1450 m depth. Leontocaris bulga Taylor & Poore, 1998 (Fig. 31b). Rostrum subequal to carapace length. Rostrum and carapace with >19 dorsal teeth (including 1 in epigastric region), and >11 ventral teeth. Mandibular palp of 2 articles. Pereopod 3 with dactylus about 0.13 of propodus length. cl. 12.7 mm. Tas. Seamounts; 1400–1650 m depth. Leontocaris yarramundi Taylor & Poore, 1998 (Fig. 31c). Rostrum 1.05–1.46 times carapace length, exceeding antennule peduncle. Rostrum and carapace with 9 dorsal teeth (including 4 in epigastric region) and 15–18 ventral teeth. Mandibular palp of 1 article. Pereopod 3 with dactylus about 0.22 of propodus length. Abdominal somite 5 pleuron with 3 posterior teeth. cl. 6.2–11.2 mm. Tas. Seamounts; 1083–1448 m depth.

Merhippolyte Bate, 1888 This genus was reviewed by Crosnier & Forest (1973). They tabulated five species prior to the description of a new southern Australian species by Kensley et al. (1987). The species are distinguished mainly on the armature of the rostrum. Diagnosis. Rostrum prominent, toothed. Supraorbital spine absent. Mandible with 3-articled palp, with incisor process. Pereopod 2 carpus with about 14 articles. Maxilliped 3 with exopod. Pereopods with arthrobranchs. Abdominal somite 6 without articulating posterolateral corner. Merhippolyte chacei Kensley, Tranter & Griffin, 1987. (Fig. 32f). Rostrum upturned, 1.5 times length of carapace, with 4–5 dorsal teeth and 7 ventral teeth. 25 mm. NSW, Tas. Seamounts; 549–1000 m depth.

Nauticaris Bate, 1888 All of the four species of Nauticaris are from the subantarctic islands, only the one below being recorded from a continental coast. Diagnosis. Rostrum prominent, deep, toothed. Supraorbital spine absent. Mandible with 3-articled palp, without incisor process. Pereopod 2 carpus with about 15 articles. Maxilliped 3 without exopod. Pereopods with arthrobranchs. Abdominal somite 6 with articulating posterolateral corner.

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Nauticaris marionis Bate, 1888 (Fig. 32a, Pl. 9h). Rostrum with 7–10 dorsal teeth and 2–4 ventral teeth. 60 mm. Vic., Tas., WA; 0–230 m. This is a common species, especially from shelf sediments but also from algal communities, and was reported from Australia by Bruce & Cropp (1984). To confirm the identification of the Australian material, specimens were compared with material of N. marionis from both New Zealand and the Marion Islands. Although the Australian specimens differed from the borrowed material by consistently having only two ventral teeth (compared to up to 3–4 for other material), there were no other differences and thus the Australian specimens were considered to be N. marionis. The deep rostrum, with about 9 dorsal teeth running back on to the carapace and 2 ventral teeth, is diagnostic.

c d b

f g e

h i

Fig. 32. Hippolytidae. a, Nauticaris marionis. Anterior region of carapace. b, Alope orientalis. c, Hippolyte caradina, d, Latreutes compressus (male). e, Lebbeus yaldwyni. f, Merhippolyte chacei. g, Saron marmoratus. h, Tozeuma elongatum, i, Tozeuma tomentosum.

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Saron Thallwitz, 1891 Only two species are known, both in the Indo-West Pacific. Diagnosis. Rostrum prominent, deep, toothed. Supraorbital spine absent. Mandible with 3-articled palp, with incisor process. Pereopod 2 carpus with many articles. Maxilliped 3 with exopod. Pereopods with arthrobranchs. Abdominal somite 6 with articulating posterolateral corner. Saron marmoratus (Olivier, 1811) (Fig. 32g, Pl. 10e). Mid-dorsal ridge of carapace with 3–4 teeth; rostrum deep, distally upturned, 1 dorsal tooth and 7 strong ventral teeth. Indo-West Pacific, WA (S to Rottnest I.), NT, Qld; shallow water.

Thor Kingsley, 1878 Thor is a tropical genus of five species inhabiting coral reefs (Holthuis, 1947). Diagnosis. Rostrum short, toothed. Supraorbital spine present or absent. Mandible without palp, with incisor process. Pereopods without epipods. Pereopod 2 carpus with 6 articles. Maxilliped 3 without exopod. Pereopods without arthrobranchs. Abdominal somite 6 without articulating posterolateral corner. Thor paschalis (Heller, 1862). Rostrum as long as eyestalk, with 6 dorsoapical teeth. Supraorbital spine absent. Stylocerite without basal marginal spinule. 9 mm. Indo-West Pacific, WA (S to Rottnest I.), NT, Qld, NSW (S to south coast); coastal. This species, widespread and frequently reported from the tropics, is found only in the warmer parts of Australia.

Tozeuma Stimpson, 1860 The long thin rostrum and slender body characterise this genus of ten species. Except for rare occurrences in Victoria all four Australian species are from the SA Gulfs and coast (Baker, 1904). Diagnosis. Rostrum prominent, slender, sometimes longer than slender body. Supraorbital spine absent. Mandible without palp, without incisor process. Pereopod 2 carpus with 3 articles. Maxilliped 3 without exopod. Pereopods without arthrobranchs. Abdominal somite 6 without articulating posterolateral corner. Tozeuma elongatum (Baker, 1904) (Fig. 32h, Pl. 10f). Body smooth. Rostrum 1.5 times carapace, with numerous fine ventral teeth. Abdomen curved. Red with black dot fringed in yellow on abdominal somite 5. 50 mm. Vic. (W of Wilsons Promontory), SA; 0–27 m depth. Tozeuma kimberi (Baker, 1904). Body smooth. Rostrum as long as carapace, with numerous fine ventral teeth. Abdomen geniculate. Carapace and most of abdomen green with blue and red dots; rest red with dark spot on abdominal somite 5 (Edgar, 1997: colour photo, p. 193). 45 mm. SA (Gulf St Vincent); 7 m depth. Tozeuma pavoninum (Bate, 1863). Body smooth. Rostrum little shorter than rostrum, with 4–7 ventral teeth. Abdomen geniculate. Green, with a crimson and yellow rostrum, yellow striped carapace, and yellow, blue, red abdomen with ‘eyespots’.64 mm. SA (Gulf St Vincent); to 24 m depth. The specific name is from that of the peacock and alludes to the colour of the ‘eyespots’ on the side of the abdomen, said to ‘rival in brilliancy those of the peacock’s tail’. Tozeuma tomentosum (Baker, 1904) (Fig. 32i). Carapace and abdomen finely setose. Rostrum as long as carapace, with 5 ventral teeth. Abdomen not geniculate. 40 mm. Japan (doubtful?), SA; 36–50 m depth.

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References Baker, W.H. 1904. Notes on South Australian decapod Crustacea. Part I. Transactions of the Royal Society of South Australia 28: 146–161. Bruce, A.J. 1990. Leontocaris amplectipes sp. nov. (Hippolytidae), a new deep-water shrimp from southern Australia. Memoirs of the Museum of Victoria 51: 121–130. Bruce, A.J., & Cropp, D.A. 1984. A redescription of Periclimenes yaldwyni Holthuis (Brachycarpus audouini Bate, 1888, Crustacea, Decapoda, Palaemonidae) and its occurrence in Australian waters. Pacific Science 38: 189–198. Chace, F.A. 1997. The caridean shrimps (Crustacea: Decapoda) of the Albatross Philippine Expedition, 1907–1910, Part 7: Families Atyidae, Eugonatonotidae, Rhynchocinetidae, Bathypalaemonellidae, Processidae, and Hippolytidae. Smithsonian Contributions to Zoology 587: i-v, 1–106. Christoffersen, M.L. 1987. Phylogenetic relationships of hippolytid genera, with an assignment of new families for the Crangonoidea and Alpheoidea (Crustacea, Decapoda, Caridea). Cladistics 3: 348–362. Crosnier, A., & Forest, J. 1973. Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19: 1–409. d’Udekem d’Acoz, C. 1996. The genus Hippolyte Leach, 1814 (Crustacea: Decapoda: Caridea: Hipplytidae) in the East Atlantic Ocean and the Mediterranean Sea, with a checklist of all species in the genus. Zoologische Verhandelingen, Leiden 303: 1–133. d’Udekem d’Acoz, C. 2001. Redescription of Hippolyte australiensis (Stimpson, 1860) (Crustacea, Decapoda, Caridea). Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 71: 37–44. Edgar, G.J. 1997. Australian Marine Life – The Plants and Animals of Temperate Waters. Reed Books: Kew Victoria. 544 pp. Fransen, C.H.J.M. 2001. Leontocaris vanderlandi, a new species of hippolytid shrimp (Crustacea: Decapoda: Caridea) from the Seychelles, with an analysis of phylogenetic relations within the genus. Zoologische Verhandelingen, Leiden 334: 57–76. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Holthuis, L.B. 1947. The Decapoda of the Siboga Expedition. Part IX. The Hippolytidae and Rhynchocinetidae collected by the Siboga and Snellius Expeditions with remarks on other species. Siboga Expéditie Monographie 39a8: 1–100. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Poore, G.C.B., Hart, S., Taylor, J., & Tudge, C. 1998. Decapod crustaceans from Tasmanian seamounts. Pp. 65–78 in: Koslow, J.A., & Gowlett-Holmes, K. (eds), The seamount fauna of southern Tasmania: benthic communities, their conservation and impacts of trawling. Final report to Environment Australia and The Fisheries Research Development Corporation. CSIRO Marine Research: Hobart. Taylor, J., & Poore, G.C.B. 1998. A review of the genus Leontocaris (Crustacea: Caridea: Hippolytidae) with descriptions of three species from southeastern Australian seamounts and slope. Memoirs of the Museum of Victoria 57: 57–69.

Ogyrididae Holthuis, 1955 Ogyridid shrimps are immediately recognisable by their elongate eyestalks. There is a single genus. Diagnosis. Carapace without antennal and pterygostomial spines. Rostrum very small. Eyestalks very elongate, reaching nearly to distal end of antennal peduncle. Pereopods 1 and 2 similarly robust. Pereopod 2 carpus subdivided. Pereopods 3 and 4 distally setose.

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Ogyrides Stebbing, 1914 The genus has 12 species worldwide, only one in southern Australia and another in northern waters. Ogyrides delli Yaldwyn, 1971 (Fig. 33a) 4–8 movable spines on midline of anterior carapace; carpus of pereopod 2 with 4 (obscurely 5) articles. New Zealand, southern Qld, NSW, Vic., Tas.; burrowing in sand down to 60 m depth. The elongate eyestalks distinguish this species from all other shrimps in southern Australia. The species from south-eastern Australia was referred by Banner & Banner (1982) to a species originally described from New Zealand (Yaldwyn, 1971).

Fig. 33. Ogyrididae. a, Ogyrides delli. Processidae. b, Processa australiensis. Physetocarididae. c, Physetocaris microphthalma.

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References Banner, D.M., & Banner, A.H. 1982. The alpheid shrimp of Australia. Part III: the remaining alpheids, principally the genus Alpheus, and the family Ogyrididae. Records of the Australian Museum 34: 1–358. Yaldwyn, J.C. 1971. Preliminary descriptions of a new genus and twelve new species of natant decapod Crustacea from New Zealand. Records of the Dominion Museum 7: 85–94.

Superfamily Processoidea Ortmann, 1896 Processidae are the only family.

Processidae Ortmann, 1896 The family Processidae, of more than 60 species in five genera, are small nocturnal shrimps found in seagrass environments. In southern Australia, all species belong to Processa and can be reliably recognised by the asymmetry of the first pereopods, chelate on the right and simple on the left. A second genus with this characteristic, Nikoides, found in northern Australia differs in having an exopod on pereopod 1. The only genus with symmetrical chelate first limbs, Ambidexter, occurs in Florida. Chace (1997), Hayashi (1975) and Noel (1986) reviewed the family. Diagnosis. Rostrum short, with bifid or rarely simple apex. Carapace smooth, with antennal spines at most. Mandible without incisor and palp. Maxilliped 1 with exopod abutting endite, displacing palp. Pereopod 1 chelate on right, simple on left (both chelate in one genus). Pereopods 2 chelate, long carpus subdivided into numerous articles, merus and ischium sometimes subdivided also. Pereopods 1–4 or 2–4 each with arthrobranch.

Processa Leach, 1815 The numerous species of this immediately identifiable genus occur worldwide in temperate and tropical shallow waters. Both southern Australian species were figured by Hale (1927). Diagnosis. Pereopod 1 chelate on right, simple on left (rarely vice versa). Pereopod 1 without exopod.

Key to southern Australian species of Processa 1. Right chelate pereopod 1 with merus about as long as carpus and propodus together ...... Processa australiensis – Right chelate pereopod 1 with merus longer than carpus and propodus together ...... Processa gracilis Processa australiensis Baker, 1907 (Fig. 33b). Right chelate pereopod 1 with merus about as long as carpus and propodus together. Carapace about twice as long as deep. cl. 6 mm. Philippines, Indonesia, Saudi Arabia, NSW, SA Gulfs; to 32 m depth. Processa gracilis Baker, 1907. Right chelate pereopod 1 with merus longer than carpus and propodus together. Carapace about 3 times as long as deep. cl. 6 mm. SA Gulfs.

128 Caridea – shrimps

Superfamily Pandaloidea Haworth, 1825 Pandalidae are one of two families. The other, Thalassocarididae Bate, 1888, is pelagic in tropical Australia (Davie, 2002).

Pandalidae Haworth, 1825 In pandalids, pereopod 1 is simple or minutely chelate. The Pandalidae, comprising more than 140 species, range from the littoral zone to more than 3000 m depth. Benthic algal inhabitants and midwater species are included. Chace (1985), Kensley et al. (1987), Hanamura & Takeda (1987), Hanamura (1987) and Hanamura & Evans (1996) described numerous shelf and deep-water Australian species. The species in southern Australia are relatively easy to distinguish on the basis of the rostrum alone but keying through the genera and species to be certain of generic placement is more difficult. The key, based on those in Chace (1985) and Holthuis (1993), relies on strap-like epipods, a character difficult to see. Epipods can be seen as short posteriorly-directed curved appendages at the base of the legs and covered by the carapace. The key also relies upon the presence of pereopod 2, sometimes lost in netted specimens. If the pereopods are lost, the shape of the rostrum and the presence or absence of an exopod on maxilliped 3 should enable identification to at least genus level from the descriptions below. The papers of Hanamura & Takeda (1987) and Hanamura & Evans (1996) are useful for tropical Australia. Most species in the key are large species from midwater oceanic environments. Two, in the genus Chlorotocella, are from shallow water algal habitats (marked *) are distinguished from other shallow water shrimps by the minutely chelate pereopod 1 and 3-segmented carpus on pereopod 2.

Diagnosis. Rostrum various, well-developed and laterally compressed. Maxilliped 1 with exopod far removed from endite. Pereopods 1 and 2 usually dissimilar, without pectinate fingers. Pereopod 1 with chelae microscopic or simple. Pereopod 2 with conventional chela; carpus usually subdivided into 2 or more short segments.

Key to southern Australian species of Pandalidae 1. Lateral surface of carapace with longitudinal carinae ...... Heterocarpus sibogae — Lateral surface of carapace without longitudinal carinae ...... 2 2. Carpus of pereopod 2 of 2 or 3 articles ...... 3 — Carpus of pereopod 2 of more than 3 articles ...... 5

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3. Arthrobranchs and epipods present at bases of pereopods 1–4; maxilliped 3 with exopod; carpus of pereopod 2 2-articled ...... Chlorotocus novaezealandiae — Pereopods without arthrobranchs and epipods; maxilliped 3 without an exopod; carpus of pereopod 2 3-articled ...... Chlorotocella … 4 4. Abdominal somite 3 normal, not swollen dorsally ...... Chlorotocella spinicaudus* — Abdominal somite 3 swollen dorsally ...... Chlorotocella gibber* 5. Last article of maxilliped 2 longer than broad, not applied as strip to the penultimate article but with its narrow base ...... Stylopandalus richardi — Last article of maxilliped 2 broader than long, attached strip-like to the penultimate article with its longer side ...... Plesionika .. 6 6. Without strap-like epipods on coxae of any pereopods ...... 7 — With strap-like epipods on coxae of pereopods 1–4 (reduced and obscure in P. edwardsii) .8 7. Pleura of abdominal somites 4 and 5 each with posteroventral tooth; maxilliped 3 without epipod ...... Plesionika spinipes — Pleura of abdominal somites 4 rounded, pleuron 5 terminating in sharp posteroventral tooth; maxilliped 3 with epipod ...... Plesionika laurentae 8. Abdomen with acute posteromedial tooth on somite 3 ...... Plesionika reflexa — Abdomen with tergum of somite 3 evenly rounded and unarmed posteriorly ...... 9 9. Dorsal margin of rostrum and carapace with more than 20 teeth closely set ...... Plesionika edwardsii — Dorsal margin of rostrum and carapace with fewer than 20 teeth less closely set ...... 10 10. Telson bearing 5 pairs of dorsolateral small spiniform setae, including pair adjacent to lateral pair of posterior spiniform setae ...... Plesionika bifurca — Telson bearing 4 pairs of dorsolateral small spiniform setae, including pair adjacent to lateral pair of posterior spiniform setae ...... 11 11. Rostrum usually armed ventrally with more than 35 closely set teeth, serrate ...... Plesionika martia — Rostrum armed ventrally with fewer than 25 rather widely spaced teeth ...... 12 12. Pereopods 2 subequal, chelate, carpus consisting of 26–32 segments; abdominal somite 4 pleura without posteroventral tooth; eyes well developed, lacking ocellus (spot on dorsal side of eye stalk) ...... Plesionika alcocki — Pereopods 2 unequal: left long and slender, carpus of about 114 segments; right with 25 segments; abdominal somite 4 pleura with posteroventral tooth. Eye somewhat compressed; ocellus large subcircular, narrowly joined to cornea ...... Plesionika grahami

Chlorotocella Balss, 1914 The slender rostrum with comparatively few and feeble teeth and presence of a supraorbital spine are distinctive for this genus. Unlike other members of the family in southern Australia, its species are benthic on the shelf or coastal algal dwellers. Diagnosis. Rostrum long and slender mostly unarmed or with few, feeble teeth on dorsal and ventral margins. Carapace with supraorbital spine. Maxilliped 2 with terminal article nearly as long as wide; maxilliped 3 without exopod. Pereopods lacking epipods and arthrobranchs. Pereopods 2 subequal and similar, carpus with 3 articles. Abdomen with posterodorsal tooth on somite 6.

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Chlorotocella gibber (Hale, 1927). Abdominal somite 3 swollen dorsally in front of strong curva- ture. 31 mm. SA. According to Hale (1927) this species is found only in Gulf St Vincent, SA. He described it as a subspecies, Parapandalus leptorhynchus gibber. Chlorotocella spinicaudus (Milne Edwards, 1837) (Fig. 34a, Pl. 10g). Abdominal somite 3 strongly curved so remainder of abdomen directed downwards, not swollen dorsally. 25 mm. Vic., Tas., SA, WA; 3–115 m depth. This common delicate, semitransparent species was figured as Parapandalus leptorhynchus by Hale (1927). Its name was corrected by Holthuis (1995).

Fig. 34. Pandalidae. a, Chlorotocella spinicaudus. b, Heterocarpus sibogae. c, Plesionika martia.

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Chlorotocus Milne Edwards, 1882 There are only three species (Chace, 1985) and only one found in southern mesopelagic Australian waters. Diagnosis. Rostrum armed with teeth on dorsal and ventral margins; carapace without supraorbital spine or longitudinal lateral carinae. Maxilliped 2 with terminal article distinctly wider than long. Maxilliped 3 with exopod. Pereopods 2 subequal and similar; carpus of 2 articles. Epipods present at bases of pereopods 1–4. Abdomen without posterodorsal spine on somite 6. Chlorotocus novaezealandiae (Borradaile, 1916) (Fig. 35a). Rostrum just falling short of, just reaching, or barely over-reaching the distal antennular peduncle article, with 10–13 dorsal and 3–5 ventral teeth. Anterior rostral teeth evenly spaced. Chela on pereopod 2 only. Transparent with red/pink to yellow/orange markings. 65 mm. New Zealand, NSW; surface to about 200 m depth in oceanic waters. This species was first recorded outside New Zealand waters by Kensley et al. (1987).

a b

c d

i h

Fig. 35. Pandalidae. Carapace and rostrum: a, Chlorotocus novaezealandiae. b, Plesionika alcocki. c, Plesionika edwardsii. d, Plesionika grahami. e, Plesionika laurentae. f, Plesionika spinipes. g, Stylopandalus richardi. h, Plesionika laurentae (pleonites 1–4). i, Plesionika reflexa (pleonites 1–4). j, Stylopandalus richardi (pleonite 3). k, Plesionika bifurca (telson).

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Heterocarpus Milne Edwards, 1881 Of the 20 species of the mesopelagic genus Heterocarpus found worldwide (Chace, 1985), eight occur in northern WA (Hanamura & Takeda, 1987; Hanamura & Evans, 1996) but only one occurs in southern Australia. Diagnosis. Rostrum armed on dorsal and ventral margins. Carapace without supraorbital spine, dorsally carinate nearly to posterior margin, and with 1 or more longitudinal lateral carinae. Maxilliped 2 with terminal article wider than long. Epipods present at the bases of pereopods 1–4. Pereopods 2 distinctly unequal and dissimilar: carpus of left with 7–12 articles; right with 18–25 articles. Heterocarpus sibogae de Man, 1917 (Fig. 34b). Longitudinal postantennal carinae on the lateral surface of the carapace reaching posterior margin of carapace. Abdomen with strong dorsomedial carinae on somites 1 and 2. cl. 37 mm. Andaman Sea, Indonesia, southern Philippines, Japan, New Caledonia, Vanuatu, Fiji, NSW, Vic., WA; mesopelagic, 247–828 m depth. This species is distinguished from other southern Australian pandalids by the longitudinal carinae on the lateral surface of the carapace. Heterocarpus hiyashii Crosnier, 1988 is similar but the dorsal spination does not reach the midlength of the carapace as it does in this species. The broadly based high dorsal ridge on the carapace is also characteristic. Chace (1985) discussed variation of the species in the Philippines.

Plesionika Bate, 1888 Of the 65 described species of the mesopelagic genus Plesionika found worldwide, 15 occur in north-western Australia (Hanamura, 1987; Hanamura & Takeda, 1987; Hanamura & Evans, 1996) and at least eight in southern Australia. Diagnosis. Rostrum armed with at least some fixed teeth on dorsal and ventral margins. Carapace without supraorbital spine or lateral carinae, dorsally carinate anteriorly, rounded posteriorly. Maxilliped 2 with terminal article wider than long. Maxilliped 3 with exopod. Epipods present at bases of pereopods 1–4 (except P. spinipes and P. laurentae). Carpus of pereopod 2 divided into more 3 three articles. Abdominal somite 3 unarmed or bearing fixed posteromedial tooth. Plesionika alcocki (Anderson, 1896) (Fig. 35b). Rostrum dorsally unarmed anterior to crest, ventral margin with fewer than 20 fairly widely spaced teeth; rostral crest of 5 sharp teeth, posterior 3 small and moveable. Eyes well developed, lacking ocellus. Pereopods 2 subequal, carpus with 26–32 articles. With strap-like epipods on pereopodal coxae. cl. 30 mm. Indo- West Pacific, Qld, to southern NSW; mesopelagic, 500–1412 m depth. Plesionika bifurca Alcock & Anderson, 1894 (Fig. 35k). Rostral crest of 4 teeth behind orbital margin and 2–3 teeth anterior on dorsal margin; ventral margin with 3–6 teeth and small subapical tooth. Eyes well developed, lacking ocellus. Pereopods 2 unequal, carpus with 10–15 and 15–24 articles. With strap-like epipods on pereopodal coxae. Telson bearing 5 pairs of dorsolateral spiniform setae, including pair adjacent to lateral pair of posterior spiniform setae. cl. 15 mm. Indo-West Pacific to South China Sea and Japan, Qld, to southern NSW; mesopelagic, 245–1412 m depth. This is the only southern Australian species of Plesionika with 5 pairs of dorsolateral small spiniform setae on the telson. Plesionika edwardsii (Brandt, 1851) (Fig. 35c). Rostrum with about 4 sharp teeth basally and more than 20 teeth closely set along dorsal margin; ventral margin with more than 30 teeth. Eyes well developed, ocellus present. Pereopods 2 subequal, carpus with 21–25 articles. With reduced and obscure strap-like epipods on coxae of pereopods 1–4. cl. 20 mm. Eastern and western Atlantic, Mediterranean, Gulf of Mexico, Indo-West Pacific, Qld, to southern NSW;

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mesopelagic, 50–689 m depth. The strap-like epipods on coxae of pereopods 1–4 are reduced and obscure compared to those of other species of the genus. Plesionika grahami Kensley, Tranter & Griffin, 1987 (Fig. 35d). Rostrum armed dorsally with 6 or 7 teeth on rostral crest, 3 or 4 posterior to orbital margin, 3 posterior-most teeth articulated; 3 subapical teeth present; armed ventrally with 18–22 teeth. Eye somewhat compressed, ocellus large, subcircular, narrowly joined to cornea. Pereopods 2 unequal, carpus with c. 25 and c. 114 articles. With strap-like epipods on pereopodal coxae. NSW; mesopelagic, 400–500 m depth. Plesionika laurentae Chan & Crosnier, 1991 (Fig. 35e, h). Rostrum with 46–55 closely set dorsal teeth, teeth on the basal crest largest; 28–36 ventral teeth, posterior teeth well spaced. Eyes well developed, ocellus present. Maxilliped 3 with well-developed epipod. Without strap-like epipods on pereopodal coxae. Pereopods 2 subequal, carpus with 21–31 articles. Pleura of abdominal somites 4 rounded, pleuron 5 terminating in sharp posteroventral tooth. cl. 19.5 mm. NSW; mesopelagic, 150–210 m depth. Plesionika martia (Milne Edwards, 1883) (Fig. 34c). Rostral crest of 7–9 sharp teeth, dorsal margin anterior to crest smooth, ventral margin with 31–54 teeth. Eyes well developed, ocellus present. Pereopods 2 subequal, carpus with 18–36 articles. With strap-like epipods on pereopodal coxae. cl. 30 mm. Western and eastern Atlantic, Indo-West Pacific, Qld, NSW; mesopelagic, 190–1215 m depth. This species is distinguished by the 7–9 teeth on the dorsal margin of the rostrum. It is a widespread and common midwater shrimp found in southern Australian waters. The species varies and local populations can be distinguished, e.g., that from southern Australia differs from those from other oceans (Hanamura, 1989). Plesionika reflexa Chace, 1985 (Fig. 35i). Rostrum with 5 or 6 sharp teeth proximally, remainder of dorsal margin smooth except for subapical spine; ventral margin with more than 30 teeth. Eyes well developed. Pereopods 2 subequal, carpus with >30 articles. With strap-like epipods on pereopodal coxae. Abdomen with acute posteromedial tooth on somite 3. cl. 20 mm. South China Sea, Indonesia, Philippines, WA,Qld, NSW; mesopelagic, 191–629 m depth. The species is distinguished by the acute posteromedial tooth on abdominal somite 3. Plesionika spinipes Bate, 1888 (Fig. 35f). Rostrum with 39–57 closely set dorsal teeth and 22–36 more widely spaced ventral teeth. Eyes well developed, ocellus present. Maxilliped 3 without epipod. Pereopods 2 subequal, carpus with 21–28 articles. Without strap-like epipods on pereopodal coxae. Pleura of abdominal somites 4 and 5 each with posteroventral tooth. cl. 20 mm. Indo-West Pacific, Qld, NSW; mesopelagic, 216–360 m depth.

Stylopandalus Coutière, 1905 The genus contains a single species. The elongate last article of maxilliped 2 is characteristic (Chace, 1985). Diagnosis. Rostrum armed with mostly fixed teeth on dorsal and ventral margins. Carapace without supraorbital spine or lateral carinae, dorsally carinate anteriorly, rounded posteriorly. Terminal article of maxilliped 2 longer than broad. Maxilliped 3 with well developed exopod. Pereopods lacking epipods. Pereopods 2 subequal, carpus divided into more than 3 articles. Abdominal somite 3 bearing slender, basally articulated posteromedial tooth or minute stout setae (sometimes lost). Stylopandalus richardi (Coutière, 1905) (Fig. 35g, j). Rostrum dorsally armed with 15–21 teeth, ventrally with 16–27 small teeth. Pereopod 2 carpus subequal, of 7–13 articles. cl. 10 mm. Cosmopolitan in tropical and temperate seas, NSW; mesopelagic, surface to 3600 m depth. This species can be distinguished by the minute (sometimes lost), posteromedial tooth on abdominal somite 3, and the elongate terminal article of maxilliped 2.

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References Chace, F.A. 1985. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 3: families Thalassocarididae and Pandalidae. Smithsonian Contributions to Zoology 411: 1–143. Chan, T.-Y., & Crosnier, A. 1991. Crustacea Decapoda: Studies of the Plesionika narval (Fabricius, 1787) group (Pandalidae) with descriptions of six new species. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152: 413–461. Crosnier, A., & Forest, J. 1973. Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19: 1–409. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Hanamura, Y. 1987. Caridean shrimps obtained by R.V.‘Soela’ from north-west Australia, with description of a new species of Leptochela (Crustacea: Decapoda: Pasiphaeidae). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 4: 15–33. Hanamura, Y. 1989. Deep-sea shrimps (Crustacea: Decapoda) collected by the R.V.‘Soela’ from southern Australia. Bulletin of the Biogeographical Society of Japan 44: 51–69. Hanamura, Y., & Evans, D.R. 1996. Deepwater caridean shrimps of the families Nematocarcinidae, Stylodactylidae, Pandalidae and Crangonidae (Crustacea: Decapoda) from Western Australia. Bulletin of Nansei National Fisheries Research Institute 29: 1–18. Hanamura, Y., & Takeda, Y. 1987. Family Pandalidae (Crustacea, Decapoda, Caridea) collected by the RV ‘Soela’ from the northwest Australian shelf. Bulletin of the National Science Museum, Tokyo 13: 103–121. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Holthuis, L.B. 1995. Notes on Indo-West Pacific Crustacea Decapoda III to IX. Zoologische Mededelingen, Leiden 69: 139–151. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331.

Superfamily Physetocaridoidea Chace, 1940 The superfamily contains but one species.

Physetocarididae Chace, 1940 The family Physetocarididae comprises only a single species of oceanic shrimp, immediately recognisable from the rostrum which dominates the carapace. Diagnosis. Rostrum a broadly triangular inflated prolongation of the carapace. Carapace inflated and with 2 lateral carinae, integument thin. Maxilliped 3 and pereopods without exopods. Pereopod 1 simple. Pereopods 2 chelate, carpus subdivided into 4 articles. Pereopods 1–4 each with arthrobranch.

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Physetocaris Chace, 1940 Diagnosis. As for family. Physetocaris microphthalma Chace, 1940 (Fig. 33c). cl. 16 mm. Atlantic, north-west Pacific, NSW; mesopelagic, surface to 1600 m depth. This species is distinguished from all other mesopelagic shrimps by the unusual inflated carapace with two lateral carinae, abdomen lacking dorsal carina or spines, and minute eyes. Only a single specimen has been taken in Australia (Kensley et al., 1987).

References Crosnier, A., & Forest, J. 1973. Les crevettes profondes de l’Atlantique oriental tropical. Faune Tropicale 19: 1–409. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331.

Superfamily Crangonoidea Haworth, 1825 Two families with characteristic subchelate or prehensile first pereopods are both represented in southern Australia.

Crangonidae Haworth, 1825 Crangonids are small, often somewhat flattened shrimps with characteristic subchelate first pereopods. The prehensile relationship between the dactylus and palm is seen elsewhere only in the Glyphocrangonidae, a family known only from deep water and with a more sculptured carapace. Holthuis (1993) recognised 21 genera of which at least seven occur in Australia and surrounding deep sea. Worldwide more than 160 species are described. They can be reliably distinguished on the basis of sculpture of the carapace and abdomen. At least two species are common in soft sediments of the shelf and other subtidal environments in southern Australia. Others are more rare and some of these have been attributed to species originally described from the Mediterranean or tropical Indo-West Pacific.

Diagnosis. Carapace with longitudinal ridges and spines. Rostrum short, sometimes with lateral basal teeth. Eyes visible. Pereopod 1 prehensile, dactylus closing against oblique distal margin propodus; palm defined by articulating spine. Pereopod 2 carpus not divided. Abdomen sometimes sculptured.

Key to southern Australian genera and species of Crangonidae 1. Pereopod 2 absent ...... Paracrangon australis — Pereopod 2 present ...... 2 2. Pereopod 2 simple, not chelate ...... 3 — Pereopod 2 chelate ...... 4 3. Pereopod 2 rudimentary, thin, short, failing to reach end of pereopod 1 merus ...... Lissosabinea tridentata — Pereopod 2 well-developed, reaching beyond pereopod 1 merus; carapace and abdomen sculptured ...... Vercoia gibbosa

136 Caridea – shrimps

4. Pereopod 2 about as long as other pereopods ...... Metacrangon spp. — Pereopod 2 much shorter than other pereopods ...... 5 5. Gills with ventral tips clearly directed posteriorly; rostrum with simple apex ...... 6 — Gills with ventral tips directed anteriorly; rostrum with bifid apex ...... 10 6. Pereopod 1 with exopod ...... Parapontophilus gracilis — Pereopod 1 without exopod ...... Philocheras ...7 7. Middorsal line on carapace divided into 2 ridges, each defined anteriorly by tooth; submedian oblique tooth lateral to anterior median tooth; dorsolateral ridge unarmed ...... Philocheras obliquus — Middorsal ridge with 1 or more sharp teeth; no submedian spine between middorsal ridge and dorsolateral ridge; dorsolateral ridge with or without spines ...... 8 8. Dorsolateral ridge on carapace unarmed (obscure); middorsal ridge with 1 anterior tooth ...... Philocheras flindersi — Dorsolateral ridge on carapace with row of spines; middorsal ridge with 1–3 teeth ...... 9 9. Middorsal ridge on carapace with 3 similar and evenly spaced teeth ...... Philocheras victoriensis — Middorsal ridge on carapace with 1 large tooth anteriorly, sometimes a smaller one behind ...... Philocheras intermedius 10. Middorsal ridge on carapace with 7–9 teeth ...... Pontocaris propensalata — Middorsal ridge on carapace with 4 or 5 teeth ...... Aegaeon … 11 11. Middorsal ridge on carapace with 5 teeth ...... Aegaeon rathbunae — Middorsal ridge on carapace with 4 teeth ...... Aegaeon lacazei

Aegaeon Agassiz, 1846 One species recorded from NSW and previously referred to Pontocaris is placed in Aegaeon as defined by Holthuis (1993). Diagnosis. Carapace about half length of abdomen. Rostrum apically bifid, with minute basal teeth. Carapace middorsal ridge with 4–5 teeth, without submedian spine, with dorsolateral ridge having 6 or more spines, hepatic spine present, lateral ridge spinose. Gills (pleurobranchs) ventrally directed anteriorly. Pereopod 1 with exopod. Pereopod 2 chelate, much shorter than other pereopods. Aegaeon lacazei (Gourret, 1887) (Fig. 36e). Carapace middorsal ridge with 4 teeth. Dorsolateral ridge on carapace with 6–7 spines. Lateral ridge with 9 spines, first 2 separated from rest by groove. Abdominal somites 3–4 with median and pair of oblique ridges, 1, 5 and 6 with paired submedian ridges. 45 mm. Mediterranean, eastern Atlantic, Indo-Pacific, NSW, Vic.; 30–759 m depth. Australian specimens differ from those from the Mediterranean in having fewer spines in the lateral row (9 vs 12–20). Aegaeon rathbunae de Man, 1918. Carapace about 1.5 times as long as wide. Carapace middorsal ridge with 5 teeth. Lateral ridge with 4–5 spines. Abdominal somite 4 with 3 pairs of diverging dorsolateral carinae. 28 mm. Indonesia, Hawaii, NSW; 11–1600 m depth. Kensley et al. (1987) recorded the species as member of Pontocaris.

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efghi

Fig. 36. Crangonidae. a, Lissosabinea tridentata (from Dardeau & Heard, 1983). b, Paracrangon australis (from Hanamura et al., 1999). c, Philocheras victoriensis. d, Vercoia gibbosa (from Duris, 1992). Dorsal view of carapace: e, Aegaeon lacazei. f, Metacrangon sp. g, Philocheras flindersi. h, Philocheras intermedius. i, Philocheras obliquus.

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Lissosabinea Christoffersen, 1988 The genus with its characteristic forked rostrum is known from a single species and this new Australian record appears indistinguishable. Diagnosis. Carapace without submedian spine, dorsolateral ridge with 1–2 spines, hepatic spine present. Eyes well developed; cornea present, large, well pigmented. Pereopod 1 rudimentary, thin, short, failing to reach end of pereopod 2 merus. Pereopod 2 simple, not chelate. Abdominal somites 1, 2, 4 and 5 without dorsal carina. Lissosabinea tridentata Pequegnat, 1970 (Fig. 36a). Rostrum acute, armed laterally with pair of sharp spines. Carapace middorsal ridge with 3 teeth. Submedian spine absent. Dorsolateral ridge with 1 spine. Hepatic spine present. Lateral ridge without spines. Pereopod 1 rudimentary, thin, short, failing to reach end of pereopod 2 merus. South-eastern Gulf of Mexico, 391 m depth (Dardeau & Heard, 1983); Vic. (new record); 260–265 m depth.

Metacrangon Zarenkov, 1965 Unidentified species of Metacrangon (Fig. 36f) occur on the south-eastern shelf and around 500–600 m depth off eastern Tasmania. Diagnosis. Carapace much shorter than abdomen. Rostrum simple. Carapace middorsal ridge with 2 teeth, submedian spine present , dorsolateral ridge unarmed, hepatic spine present, lateral ridge unarmed. Gills (pleurobranchs) ventrally directed posteriorly. Pereopod 1 without exopod. Pereopod 2 chelate, subequal in length to other pereopods.

Paracrangon Dana, 1852 This genus of six species, mostly in the Northern and Eastern Pacific was reviewed by Hanamura et al. (1999) while describing a new Australian species. Diagnosis. Carapace much shorter than abdomen. Rostrum long, narrow, oblique and with ventral teeth. Carapace middorsal ridge with 2–5 teeth, submedian spines present, dorsolateral ridge with spines, hepatic spine present, lateral ridge with spines. Gills (pleurobranchs) ventrally directed posteriorly. Pereopod 1 without exopod. Pereopod 2 absent. Paracrangon australis Hanamura, Wadley & Taylor, 1999 (Fig. 36b, Pl. 10h). Carapace with distinct irregular reticulate structure; rostrum shorter than carapace; obtuse ventral projections of abdominal pleura 1 and 2 shorter than depth of carapace; pereopods 4 and 5 with dactylus about one-fifth length of propodus. 70 mm. South-east Tas., Macquarie I.; 363–1422 m depth.

Parapontophilus Christoffersen, 1988 Diagnosis. Carapace much shorter than abdomen. Hepatic spine present. Gills (pleurobranchs) ventrally directed posteriorly. Pereopod 1 with rudimentary exopod. Pereopod 2 chelate, much shorter than other pereopods. Parapontophilus gracilis (Smith, 1882). Carapace much shorter than abdomen. Carapace middorsal ridge with 2 teeth and small denticle anteriorly. Submedian spine absent. Dorsolateral ridge with 1 tooth (rarely 2). Hepatic spine present. Lateral ridge without spines. Pereopod 2 well-developed, reaching beyond pereopod 2 merus. Atlantic and Indo-West Pacific; abyssal, benthic (Davie, 2002). Three subspecies have been recorded in Australia, P. gracilis profundus (Bate, 1888) by the Challenger Expedition off Sydney, NSW, at abyssal depths.

139 Marine Decapod Crustacea of Southern Australia

Philocheras Stebbing, 1900 Since Fulton & Grant’s (1902) publication, species of Philocheras in Australia were referred to Pontophilus Leach, 1817. Chace’s (1984) revision differentiated the two genera. Now, virtually all shallow and shelf specimens can be placed in Philocheras on account of the simple rounded rostrum and absence of an exopod on pereopod 1. An undescribed species from slope depths in south-eastern Australia exists in museum collections. Diagnosis. Carapace about half length of abdomen. Rostrum simple, basally depressed. Carapace middorsal ridge with at least 1 tooth, with dorsolateral ridge spinose or not, hepatic spine present, lateral ridge with 1 or more spines. Gills (pleurobranchs) ventrally directed posteriorly. Pereopod 1 without exopod. Pereopod 2 chelate, much shorter than other pereopods. Philocheras flindersi (Fulton & Grant, 1902) (Fig. 36g). Middorsal ridge with 1 tooth anteriorly. Submedian spine absent. Dorsolateral ridge unarmed, curving along front margin. Hepatic spine present. Lateral ridge with 2 spines. 35 mm. NSW, Tas.,Vic., SA; shelf sediments, 18–127 m depth. P. flindersi is the dominant crangonid in Bass Strait sediments. Philocheras intermedius (Bate, 1863) (Fig. 36h, Pl. 10i). Middorsal ridge with 1 anterior tooth, sometimes smaller second. Submedian spine absent. Dorsolateral ridge with 4 spines. Hepatic spine present. Lateral ridge with 1–3 spines. 45 mm. SA (Gulf St Vincent); shelf sediments. P. intermedius differs from P. victoriensis only in the possession of 1 rather than 3 medial teeth. Hale (1927) described the colour as ‘exquisite’,mottled pale grey on top, like the sediments in which it lives, with white eyes, brilliant red posterior somites, three velvety black marks on the fifth abdominal somite, and sides and limbs purple, reddish or yellow. Philocheras obliquus (Fulton & Grant, 1902) (Fig. 36i). Middorsal ridge with anterior and posterior halves, each defined anteriorly by tooth. Submedian spine strong and oblique. Dorsolateral ridge unarmed. Hepatic spine present. Lateral ridge without spines. 23 mm. Tas., Vic., SA; subtidal and shelf sediments, 0–18 m depth, rare. Philocheras victoriensis (Fulton & Grant, 1902) (Fig. 36c). Middorsal ridge with 3 similar and evenly spaced teeth. Submedian spine absent. Dorsolateral ridge with 4–5 spines. Hepatic spine present. Lateral ridge with 1–4 spines. 24 mm. Tas.,Vic., SA, southern WA; subtidal and bay sediments, 0–42 m depth. This is the most common crangonid species encountered in sheltered environments such as Port Phillip Bay and Western Port.

Pontocaris Bate, 1888 Pontocaris differs from Aegaeon in possessing more than five middorsal teeth on the carapace (Chan, 1996). Diagnosis. Carapace about half length of abdomen. Rostrum apically bifid, with minute basal teeth. Carapace middorsal ridge with more than 5 teeth, without submedian spine, with dorsolateral ridge having more than 6 spines, hepatic spine present, lateral ridge spinose. Gills (pleurobranchs) ventrally directed anteriorly. Pereopod 1 with exopod. Pereopod 2 chelate, much shorter than other pereopods. Pontocaris propensalata Bate, 1888. Carapace about 1.2 times as long as wide. Carapace middorsal ridge with 7–9 teeth. Lateral ridge obscurely tuberculate. Abdominal somite 4 with 3 pairs of diverging dorsolateral carinae. 28 mm. Philippines, Indonesia, NSW; 49–349 m depth.

140 Caridea – shrimps

Vercoia Baker, 1904 Vercoia is known from three species (Duris, 1992; Komai, 1995), only one Australian. Diagnosis. Carapace little shorter than abdomen. Rostrum broader than long, with apical projection. Carapace middorsal ridge apparent only posteriorly, dorsolateral ridge apparent anteriorly, hepatic spine absent. Gills (pleurobranchs) ventrally directed anteriorly. Pereopod 1 without exopod. Pereopod 2 simple, much shorter than other pereopods. Vercoia gibbosa Baker, 1904. (Fig. 36d). Merus of pereopod 2 not especially enlarged. 17 mm. Qld (Lizard I.), SA; shelf sediments, 3–54 m depth. The compact form and strong sculpture of the carapace and abdomen immediately distinguish V. gibbosa from species of Philocheras.

References Chace, F.A. 1984. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 2: families Glyphocrangonidae and Crangonidae. Smithsonian Contributions to Zoology 397: 1–63. Chan, T.-Y. 1996. Crustacea Decapoda Crangonidae: revision of the three closely related genera Aegaeon Agassiz, 1846, Pontocaris Bate, 1888 and Parapontocaris Alcock, 1901. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 15. Mémoires du Muséum National d’Histoire Naturelle, Paris 168: 269–336. Dardeau, M.R., & Heard, R.W. 1983. Crangonid shrimps (Crustacea: Caridea), with a description of a new species of Pontocaris. Memoirs of the Hourglass Cruises 6: 1–39. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Duris, Z. 1992. Revision of Vercoia Baker (Crustacea: Decapoda: Crangonidae). Invertebrate Taxonomy 6: 1437–1457. Fulton, S.W., & Grant, F.E. 1902. Some little known Victorian decapod Crustacea with descriptions of new species. No. II. Proceedings of the Royal Society of Victoria 15: 59–68, pls 8–10. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Hanamura, Y., Wadley, V., & Taylor, J. 1999. Description of a new species of the Pacific shrimp genus Paracrangon (Crustacea: Decapoda: Crangonidae) from southern Australia with a key to the genus. Memoirs of the Museum of Victoria 57: 311–317. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Komai, T. 1995. Vercoia japonica, a new species of crangonid shrimp (Crustacea: Decapoda: Caridea) from Japan. Natural History Research 3: 123–132.

Glyphocrangonidae Smith, 1884 The family comprises only one genus of more than 73 species (Komai 2004). All are large highly sculptured shrimps from upper slope to abyssal depths. Lateral carinae usually end anteriorly in strong antennal and branchiostegal spines. Chace (1984) provided a key to the species then known and Komai (2004) treated Indo-West Pacific species. Diagnosis. Carapace rigid, with prominent sculpture of longitudinal ridges and spines. Rostrum prominent, with lateral teeth. Eyes visible. Pereopod 1 prehensile, dactylus closing against truncate end of propodus. Pereopod 2 carpus divided. Abdomen sculptured.

141 Marine Decapod Crustacea of Southern Australia

Glyphocrangon Milne Edwards, 1881 Kensley et al. (1987) recorded four species from eastern Australia but some of the many known from Indonesian waters might be expected in northern Australia. Diagnosis. As family.

Key to southern Australian species of Glyphocrangon 1. Lateral anterior (fourth) carina not expanded anterolaterally, not reaching margin of orbit ...... Glyphocrangon kapala — Lateral anterior (fourth) carina expanded anterolaterally, overlapping or almost reaching margin of orbit ...... Glyphocrangon novacastellum

b c

Fig. 37. Glyphocrangonidae. a, Glyphocrangon kapala. b, c, Glyphocrangon novacastellum (carapace lateral and dorsal).

Glyphocrangon kapala Komai, 2004 (Fig. 37a). Dorsolateral posterior (second) carina tuberculate. Antennal posterior (third) carina without anterior sharp tooth. Lateral anterior (fourth) carina not reaching margin of orbit, not expanded anterolaterally. 110 mm. S Qld, NSW; 460–810 m depth. The species was referred to G. assimilis de Man, 1918 by Kensley et al. (1987).

142 Caridea – shrimps

Glyphocrangon novacastellum Kensley, Tranter & Griffin, 1987 (Fig. 37b, c). Dorsolateral posterior (second) carina tuberculate. Antennal posterior (third) carina with anterior sharp tooth. Lateral anterior (fourth) carina overlapping margin of orbit, expanded anterolaterally. 120 mm. Southern NSW; 576–823 m depth. This is one of three very similar species described by Kensley et al. (1987) from the eastern Australian coast but the only one recorded as far south as Sydney.

References Chace, F.A. 1984. The caridean shrimps (Crustacea; Decapoda) of the Albatross Philippine Expedition, 1907–1910, part 2: families Glyphocrangonidae and Crangonidae. Smithsonian Contributions to Zoology 397: 1–63. Kensley, B., Tranter, H.A., & Griffin, D.J.G. 1987. Deepwater decapod Crustacea from eastern Australia (Penaeidea and Caridea). Records of the Australian Museum 39: 263–331. Komai, T. 2004. A review of the Indo-West Pacific species of the genus Glyphocrangon A. Milne- Edwards, 1881 (excluding the G. caeca species group) (Crustacea: Decapoda: Caridea: Glyphocrangonidae). In: Marshall, B.A., and Richer de Forges, B. (eds), Tropical Deep-Sea Benthos, Vol. 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191: 375–610.

143 This page intentionally left blank 5. STENOPODIDEA – CORAL SHRIMPS AND VENUS SHRIMPS

Most shrimps belong to one of several families of the Infraorder Caridea (Chapter 4). However, coral shrimps and Venus shrimps are so different from the rest that a separate infraorder is warranted. They belong to another evolutionary branch, the Infraorder Stenopodidea Bate, 1888. The picturesque coral shrimps are the most familiar members of the group because they are much photographed animals characteristic of tropical reefs. Nevertheless, they represent only one of two stenopodidean families – the other family, called Venus shrimps, includes rarely-seen deep-water species. Stenopodideans differ from carideans in having the pleura of the first abdominal somite overlapping the second, as in dendrobranchiate prawns and lobsters, instead of the second overlapping both the first and third. The third pereopod is chelate and larger than the second, unlike carideans where pereopod 2 is usually the largest and pereopod 2 is not chelate. Dendrobranchiates have chelate pereopods 3 too. Males do not have a petasma (modified pleopods 1) and gills are trichobranchiate, with thin filaments. Females carry eggs on the pleopods as do carideans. According to Dixon et al. (2003), stenopodideans are sister taxon to all other reptant decapods. Key diagnostic features that separate stenopodideans from carideans and dendrobranchiates have been extracted from Davie’s (2002) longer description. Holthuis’s (1993) book has keys to the two families and all genera. Diagnosis. Pereopods 1–3 chelate, third usually much longer than others; without exopods. Abdominal somites with pleura not expanded, overlapping in sequence posteriorly. Pleopods uniramous in both sexes; male without petasma, appendix interna or appendix masculina. Gills trichobranchiate, central axis with irregular rows of simple filaments. Females carry eggs attached to pleopods.

Key to families of Stenopodidea 1. Body compressed; telson ending in 2 strong spines; uropodal endopod with 2 longitudinal ridges ...... Stenopodidae — Body depressed; telson ending in 3–5 strong spines; uropodal endopod with single longitudinal ridge ...... Spongicolidae

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Dixon, C.J., Ahyong, S., & Schram, F.R. 2003. A new hypothesis of decapod phylogeny. Crustaceana 76: 935–975. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn.

145 Marine Decapod Crustacea of Southern Australia

Spongicolidae Schram, 1986 The family of small, spiny, shrimps comprises five genera worldwide (Holthuis, 1946, 1993). Four genera have been recorded from depths of hundreds or thousands of metres, individual shrimps often associated with sponges, as the type genus name Spongicola and family name suggest. Microprosthema, however, is confined to shallow reefs on the shelf. This genus is the most similar to stenopodids and the others are more derived. Symbiosis with hexactinellid sponges has resulted in loss of gills, exopods, and spination (Saito & Takeda, 2003). Some deep-water species have direct development enabling fully developed juveniles hatching from large eggs to reinvade their sponge hosts (Saito & Konishi, 1999; Saito et al., 2002). Species of Microprosthema have typical planktonic development. Two species representing these different clades and very different habitats occur in southern Australian waters but others could be expected.

Diagnosis. Body depressed. Maxilliped 2 propodus with unarmed lower margin; maxilliped 3 exopod absent or rudimentary. Telson rounded, broad, ending in 3–5 subequal spines. Uropodal endopod usually with a single longitudinal dorsal ridge. Chelae with chitinous ridges distally on cutting edges, rarely with few small sharp teeth; chela of pereopod 3 propodus broad or robust. Gills number 11–19 branchial exites. (adapted from Davie, 2002)

Key to southern Australian genera of Spongicolidae 1. Carapace highly spinose; maxilliped 3 with long exopod; pereopod 1 with setiferous organ under carpus-propodus ...... Microprosthema — Carapace with few microscopic spines only; maxilliped 3 without exopod; pereopod 1 without setiferous organ ...... Spongicoloides

Microprosthema Stimpson, 1860 Four species are distributed on reefs and shallow shelves of the Indo-West Pacific (Holthuis, 1946). All are spinose shrimps with large strong chelate third pereopods. Diagnosis. Carapace highly spinose. Telson with 1 lateral tooth. Maxilliped 3 with long exopod. Pereopod 1 with setiferous organ under carpus-propodus. Microprosthema validum Stimpson, 1860 (Fig. 38a, Pl. 11a). Pereopod 3 propodus with distinct crest on upper margin. Abdominal somite 3 with short dorsal carina on posterior half. Scaphocerite with 2 or 3 strong outer teeth. White with golden-brown patterns (Debelius, 1999). tl. 16 mm. Indo-Pacific, NT, Qld, NSW (S to Port Jackson); silty intertidal pools, and subtidal to 70 m depth.

Spongicoloides Hansen, 1908 The association between species of Spongicoloides and hexactinellid sponges, popularly known as Venus flower baskets, gives shrimps of this genus the name of Venus shrimps. Both the sponge and the shrimp, which lives in a cavity in the sponge, are restricted to deep water. Four species are known from the North Atlantic and one from New Zealand (Baba, 1979). The last is now recorded from Australia.

Diagnosis. Carapace with few microscopic spines only. Uropodal exopod with single median longitudinal ridge. Maxilliped 3 without exopod. Pereopod 1 without setiferous organ. Spongicoloides novaezelandiae Baba, 1979 (Fig. 38b, c). Pereopods without epipods; maxilliped 2–pereopod 2 with 2 arthrobranchs. Rostrum reaching to end of first article of antennule,

146 Stenopodidea – coral shrimps and venus shrimps

Fig. 38. Spongicolidae. a, Microprosthema validum. b, c, Spongicoloides novaezelandiae (carapace dorsal and lateral). Stenopodidae. Rostrum: d, Stenopus hispidus. e, Stenopus tenuirostris.

with 11 small dorsal teeth, with 2 small distoventral teeth. Carapace with 3 small spines on pterygostomial margin, patches of spinules on branchiostegite. 20 mm. New Zealand, Tasmanian Seamounts, 990–1450 m depth.

References Baba, K. 1979. A new stenopodidean shrimp (Decapoda, Natantia) from the Chatham Rise, New Zealand. Pacific Science 33: 311–314. Borradaile, L.A. 1910. Penaeidae, Stenopidea and Reptantia from the Western Indian Ocean. Transactions of the Linnean Society of London 13: 257–264, 16 pls. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp.

147 Marine Decapod Crustacea of Southern Australia

Debelius, H. 1999. Crustacea. Guide of the World. Ikan-Unterwasserarchiv: Frankfurt. 321 pp. Holthuis, L.B. 1946. Biological results of the Snellius Expedition XIV. The Decapoda Macrura of the Snellius Expedition I. The Stenopodidae, Nephropsidae, Scyllaridae and Palinuridae. Temminckia 7: 1–178,11pls. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Saito, T., & Konishi, K. 1999. Direct development in the sponge-associated deep-sea shrimp Spongicola japonica (Decapoda : Spongicolidae). Journal of Crustacean Biology 19: 46–52. Saito, T., & Takeda, M. 2003. Phylogeny of the family Spongicolidae (Crustacea : Stenopodidea): evolutionary trend from shallow-water free-living to deep-water sponge-associated habitat. Journal of the Marine Biological Association of the United Kingdom 83: 119–131. Saito, T., Uchida, I., & Takeda, M. 2002. Skeletal growth of the deep-sea hexactinellid sponge Euplectella oweni, and host selection by the symbiotic shrimp Spongicola japonica (Crustacea : Decapoda : Spongicolidae). Journal of Zoology, London 258: 521–529.

Stenopodidae Claus, 1872 Coral shrimps, sometimes called boxer shrimps, of the family Stenopodidae are spiny benthic shrimps, typically of coral reefs. They are invariably found in pairs and are a favourite of under- water photographers and more recently of aquarists (Calado et al., 2003). All species are brightly coloured and patterned (see photos of nine species in Debelius, 1999). The banded coral shrimp, Stenopus hispidus, is the best known and features in many colour articles and books about coral reef life. Species of Stenopus are known as ‘fish cleaners’, establishing stations that fish visit for removal of external parasites, notably gnathiid isopods, or dead tissue (Jonason, 1987). Pairs of shrimps display elaborate mating behaviour (Zhang et al., 1998). Holthuis (1993) provided a key to four genera of which two occur in Australia. Species of Spongicolidae were previously treated in this family. Diagnosis. Body compressed. Maxilliped 2 propodus with cristate lower margin, often with 1 or more acute hooked proximal spines; maxilliped 3 with distinct exopod. Telson long, narrow, lanceolate, subtriangular, ending in 2 strong spines, sometimes with median spinule. Uropodal endopod generally with 2 longitudinal dorsal ridges. Chelae cutting edges with series of peg-like teeth separated by chitinous lamellae distally; chela of pereopod 3 long and slender. Gills number 19 branchial exites. (adapted from Davie, 2002)

Stenopus Latreille, 1819 Species of Stenopus are colourful and popular inhabitants of coral reefs but reach can be found as far south as Port Jackson, NSW. Coral shrimps are always found in pairs, usually at the entrance to small caves. Males are smaller than females. Goy (1992) described species from Australia and presented a key to eight Indo-West Pacific species. Diagnosis. Carapace and abdomen densely and uniformly covered with strong spines, which are sometimes in rows, erect and curved. Pereopods 4 and 5 with biungulate dactyli.

Key to southern Australian species of Stenopus 1. Rostrum without spine on lower margin, reaching middle of antennular peduncle; abdominal somite 3 without bare shield-shaped area ...... Stenopus hispidus — Rostrum with 3–9 spines on lower margin; reaching beyond end of antennular peduncle; abdominal somite 3 with bare shield-shaped area ...... Stenopus tenuirostris

148 Stenopodidea – coral shrimps and venus shrimps

Stenopus hispidus (Olivier, 1811) banded coral shrimp, banded cleaner shrimp (Fig. 38d, Pl. 11b). Banded red and white on carapace, abdomen, and on large pereopod 3 where tips are white; antennae and other pereopods white. tl. 60 mm. Indo-Pacific, New Zealand, West Atlantic , NT, Qld, NSW (S to Sydney); subtidal to 210 m depth. Stenopus tenuirostris de Man, 1888 (Fig. 38e, Pl. 11c). Carapace purplish; abdomen yellow with white patches, red bands on somites 3 and 5; large pereopod 3 with purple base and banded red, white and yellow, tips red; other pereopods purple-white. tl. 28 mm. Indo-West Pacific , NSW; coral and other reef.

References Calado, R., Lin, J.D., Rhyne, A.L., Araújo, R., & Narcisco, L. 2003. Marine ornamental decapods - popular, pricey, and poorly studied. Journal of Crustacean Biology 23: 963–973. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A CSIRO Publishing: Melbourne. xii, 551 pp. Debelius, H. 1999. Crustacea. Guide of the world. Ikan-Unterwasserarchiv: Frankfurt. 321 pp. Goy, J.W. 1992. A new species of Stenopus from Australia, with a redescription of Stenopus canoscelis (Crustacea: Decapodsa: Stenpodidea). Journal of Natural History 26: 79–102. Holthuis, L.B. 1946. Biological results of the Snellius Expedition XIV. The Decapoda Macrura of the Snellius Expedition I. The Stenopodidae, Nephropsidae, Scyllaridae and Palinuridae. Temminckia 7: 1–178, 11pls. Holthuis, L.B. 1993. The Recent Genera of the Caridean and Stenopodidean Shrimps (Crustacea, Decapoda) with an Appendix on the Order Amphionidacea. Nationaal Natuuristorisch Museum: Leiden. 328 pp. revised edn. Jonason, M. 1987. Fish cleaning behavior of shrimp. Journal of Zoology, London 213: 117–131. Zhang, D., Lin, J., & Creswell, R.L. 1998. Mating behavior and spawning of the banded coral shrimp Stenopus hisidus in the laboratory. Journal of Crustacean Biology 18: 511–518.

149 This page intentionally left blank 6. POLYCHELIDA – DEEP-SEA LOBSTERS

The family Polychelidae has usually been included with the rock or spiny lobsters and slipper lobsters in the infraorder Palinura. Evidence is increasing that this is entirely artificial and that polychelids are only remotely related to spiny lobsters. Stefan Richter and Gerhard Scholtz (Scholtz & Richter, 1995; Richter & Scholtz, 2001) argued that the flattened deep-sea lobsters belong to a monophyletic clade with a unique morphology. They revived the name Polychelida De Haan, 1841. Polychelids are also different in possessing a eryoneicus larva while the other ‘palinuran’ lobsters have a phyllosoma larva. Their conclusions were supported by a rigorous phylogenetic analysis of morphological characters by Dixon et al. (2003) who, however, disputed some of the defining characters. There is only one living family.

Diagnosis. Carapace dorsally flattened, with a sharp lateral margin, with 1 or 2 small rostral spines. Eyestalk present, without cornea. Pereopods 1–4, sometimes 5, chelate; pereopod 1 more robust than others, elongate and fine. Antennules and antennae long, filamentous. Antennal gland opening dorsally. Mandible without without molar process, with curved palp. Maxilliped 3 without exopod. Pereopodal basis-ischium-merus fused.

References Dixon, C.J., Ahyong, S., & Schram, F.R. 2003. A new hypothesis of decapod phylogeny. Crustaceana 76: 935–975. Richter, S., & Scholtz, G. 2001. Phylogenetic analysis of the Malacostraca (Crustacea). Journal of Zoological, Systematic and Evolutionary Research 39: 113–136. Scholtz, G., & Richter, S. 1995. Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113: 289–328.

Polychelidae Wood-Mason, 1875 Polychelids are unusual deep-water lobsters of no commercial interest. They are best recognised by the possession of extremely long and chelate first pereopods. They reach a size of about 150 mm long. In life, they are predominantly red or orange over most of the carapace. Little is known of their biology in Australia or elsewhere (Wicksten, 1980). George (1983) recorded two species from the North West Shelf and Griffin & Stoddart (1995) ten from eastern Australia. Since those times the family has been revised on worldwide basis (Galil, 2000) and additional Australian species added (Ahyong & Brown, 2002). There are five genera with 35–40 species worldwide (Gore, 1984; Holthuis, 1991; Galil, 2000; Ahyong & Brown, 2002). A fourth genus name, Eryoneicus Bate, seen in the older literature refers to the late larval stage of polychelids. The family is the only extant one in the Polychelida but similar fossils assigned to other families are known. Traditionally, Polychelidae have been assigned to the Palinura with spiny lobsters and slipper lobsters, an infraorder not recognised here.

Diagnosis. As for Polychelida.

151 Marine Decapod Crustacea of Southern Australia

Key to southern Australian species of Polychelidae 1. Eyestalks below and parallel to anterior margin of carapace; finger of pereopod 1 with subapical tooth ...... Willemoesia pacifica — Eyestalks in deep incisions on anterior margin of carapace; finger of pereopod 1 without tooth ...... 2 2. Maxilliped 3 epipod longer than ischium ...... Pentacheles . . . 3 — Maxilliped 3 epipod shorter than ischium ...... Polycheles . . . 4 3. Abdominal somites 1–3 with distinct middorsal anterior spine ...... Pentacheles laevis — Abdominal somites 1–3 without middorsal anterior spine ...... Pentacheles validus 4. Frontal margin of carapace with at least single spine on internal angle of orbital sinus . . . . 5 — Internal angle of orbital sinus produced, not spinose ...... Polycheles baccatus 5. Frontal margin of carapace with several spinules on each side of rostral spine/s ...... 6 — Frontal margin of carapace with single spine on internal angle of orbital sinus ...... 8 6. Median carina on abdominal somites 2–5 notched or crenulate; somites 2–5 with distinct oblique grooves; dorsal surface of carapace strongly granulate ...... Polycheles martini — Median carina on abdominal somites 2–5 without notch; somites 2–5 smooth, without distinct oblique grooves; dorsal surface of carapace smooth or spinose ...... 7 7. Branchial carinae indicated at most by low granules; gastric region with 1 or 2 spines of similar size to spines of median carina; posterior margin of cervical groove without spine between median and branchial carinae ...... Polycheles enthrix — Branchial carinae indicated by row of 4–6 spines; gastric region with numerous spines of similar size to spines of median carina; posterior margin of cervical groove with spine between median and branchial carinae ...... Polycheles kermadecensis 8. Median carina on abdominal somites 2–5 bicuspidate ...... Polycheles suhmi — Median carina on abdominal somites 2–5 not bicuspidate ...... 9 9. Frontal margin of carapace with 2 rostral spines ...... 10 — Frontal margin of carapace with 1 rostral spine ...... Polycheles typhlops 10. Branchial groove with 1 or more anterior spines ...... Polycheles nanus — Branchial groove unarmed ...... 11 11. Posterior margin of cervical groove with 3–4 spines between median and branchial carinae ...... Polycheles surdus — Posterior margin of cervical groove with 1 spine between median and branchial carinae . . . 12 12. Abdominal pleuron 2 with anterior spine; median submarginal tooth prominent ...... Polycheles aculeatus — Abdominal pleuron 2 without anterior spine; median submarginal tooth small ...... Polycheles sculptus

Pentacheles Bate, 1878 Members of the genus are distinguished from those of the superficially similar genus Polycheles by the possession of a long maxilliped 3 epipod. Diagnosis. Carapace with postorbital carina obsolescent, postcervical carina well defined, neither swollen. Frontal margin with orbital sinus. Antenna with basal article quadrate and lamellar, mesially granular. Maxilliped 3 epipod longer than ischium. Pereopod 5 chelate in female, subchelate in male.

152 Polychelida – deep-sea lobsters

a b

Fig. 39. Polychelidae. a, Pentacheles laevis. b, Polycheles nanus.

153 Marine Decapod Crustacea of Southern Australia

Pentacheles laevis Bate, 1878 (Figs 39a, 41a). Abdominal somites 1–3 with distinct middorsal anterior spine. Lateral margin with 14–15 spines posterior to postcervical groove. cl. 52.7 mm. Atlantic, Indo-Pacific, NSW, Vic., Tas., southern WA; 347–2505 m depth. The WA record is new for Australia (Museum Victoria collection). Pentacheles validus Milne Edwards, 1880 (Fig. 41b). Abdominal somites 1–3 without middorsal anterior spine. cl. 75.1 mm. Atlantic, Indo-Pacific, NSW, Vic., Tas.; 914–3365 m depth

Polycheles Heller, 1862 There are at least 24 species worldwide and four were recorded from southern Australian by Griffin & Stoddart (1995) as species of Stereomastis. Stereomastis was considered a synonym by Galil (2000). All are from slope depths. Diagnosis. Carapace with postorbital carina obsolescent, postcervical carina well defined, neither swollen. Frontal margin with orbital sinus. Antennule with basal article proximally rounded, anteromesially produced. Maxilliped 3 epipod rudimentary. Pereopod 5 chelate in female, simple or subchelate in male. Polycheles aculeatus Galil, 2000 (Figs 40a, 41d). Frontal margin of carapace with 2 rostral spines; median submarginal tooth prominent; internal angle of orbital sinus spinose; anterior margin of carapace smooth beyond outer angle of orbital notch; lateral margin with 8–10 spines posterior to postcervical groove; branchial groove unarmed; posterior margin of cervical groove with 1 spine between median and branchial carinae; posterior margin of carapace with 2 prominent submedian spines. Abdominal pleuron 2 with anterior spine. cl. 60 mm. WA, Qld, NSW, Tas.; 144–1053 m depth.

a bcd

i efg Fig. 40. Polychelidae. Dorsal carapace: a, Polycheles aculeatus. b, Polycheles baccatus. c, Polycheles enthrix. d, Polycheles kermadecensis. e, Polycheles sculptus. f, Polycheles suhmi. g, Polycheles surdus. h, Polycheles martini. i, Willemoesia pacifica.

154 Polychelida – deep-sea lobsters

Polycheles baccatus Bate, 1878 (Fig. 40b). Frontal margin of carapace with 2 rostral spines; median submarginal tooth prominent; internal angle of orbital sinus produced, unarmed; anterior margin of carapace spinose beyond outer angle of orbital notch; lateral margin with 21–25 spines posterior to postcervical groove; posterior margin of cervical groove granulate between median and branchial carinae; posterior margin of carapace granular, submedian pair largest. cl. 36 mm. Indo-West Pacific, northern Australia to NSW; 350–916 m depth.

c f

e h

Fig. 41. Polychelidae. Lateral abdomen: a, Pentacheles laevis. b, Pentacheles validus. c, Polycheles nanus. d, Polycheles aculeatus. e, Polycheles sculptus. f, Polycheles suhmi. g, Polycheles surdus. h, Polycheles typhlops. i, Willemoesia pacifica.

155 Marine Decapod Crustacea of Southern Australia

Polycheles enthrix (Bate, 1878) (Fig. 40c). Frontal margin of carapace with 2 rostral spines; median submarginal tooth short, inconspicuous; internal angle of orbital sinus spinose; anterior margin of carapace smooth beyond outer angle of orbital notch; lateral margin with 13–16 spines posterior to postcervical groove; posterior margin of cervical groove without spine between median and dorsal carinae; posterior margin of carapace with 3 pairs of submedian spines. cl. 73 mm. Western Pacific, Qld, NSW; 229–1152 m depth. Polycheles kermadecensis (Sund, 1920) (Fig. 40d). Frontal margin of carapace with 2 rostral spines; median submarginal tooth short, inconspicuous; internal angle of orbital sinus spinose; anterior margin of carapace spinose beyond outer angle of orbital notch; lateral margin with 12–17 spines posterior to postcervical groove; posterior margin of cervical groove with spine between median and dorsal carinae; posterior margin of carapace with 2 pairs of submedian spines. cl. 57 mm. Kermadec I., New Zealand, Qld (N to central coast), NSW (S to Newcastle); 549–951 m depth. Polycheles martini (Ahyong & Brown, 2002) (Fig. 40h). Frontal margin of carapace with 2 rostral spines; median submarginal tooth short, inconspicuous; internal angle of orbital sinus with blunt spinules; anterior margin of carapace spinose beyond outer angle of orbital notch; lateral margin with 15–18 spines posterior to postcervical groove; posterior margin of carapace strongly granulate. cl. 47 mm. NSW (Yamba to Sugarloaf Point); 256–477 m depth. Polycheles nanus (Smith, 1884) (Fig. 41c). Frontal margin of carapace with 2 rostral spines; internal angle of orbital sinus spinose; anterior margin of carapace smooth beyond outer angle of orbital notch; lateral margin with 6–7 spines posterior to postcervical groove; middorsal ridge in gastric region with 2 single spines in front of pair of spines; posterior margin of carapace spinose, with submedian spines largest. Median carinae of abdominal somites 4 and 5 posteriorly smooth. cl. 43 mm. Atlantic, Indo-Pacific, Qld, NSW, Tas.; 300–4000 m depth. The species has been reported as Stereomastis andamanensis. Polycheles sculptus Smith, 1880 (Figs 40e, 41e). Frontal margin of carapace with 2 rostral spines; internal angle of orbital sinus spinose; anterior margin of carapace smooth beyond outer angle of orbital notch; lateral margin with 6–7 spines posterior to postcervical groove; middorsal ridge in gastric region with 1 single spine in front of pair of spines; posterior margin of cervical groove with 1 spine between median and branchial carinae; posterior margin of carapace with 2 prominent submedian spines. Median carinae of abdominal somites 1–5 each with anterior spine. cl. 49 mm. Atlantic, Indo-Pacific, NSW, western Vic.; 200–4000 m depth. Polycheles suhmi (Bate, 1878) (Figs 40f, 41f). Frontal margin of carapace with 2 rostral spines; internal angle of orbital sinus spinose; anterior margin of carapace smooth beyond outer angle of orbital notch; lateral margin with 7–9 spines posterior to postcervical groove; middorsal ridge in gastric region with 2 single spines in front of 2 pairs of spines; posterior margin of carapace with 2 prominent submedian spines. Median carina on abdominal tergites 2–5 bicuspidate. cl. 37 mm. South Atlantic, South Africa, South Pacific, NSW, western Vic, Tas.; 223–2195 m depth. Polycheles surdus Galil, 2000 (Figs 40g, 41g). Frontal margin of carapace with 2 rostral spines; internal angle of orbital sinus spinose; anterior margin of carapace smooth beyond outer angle of orbital notch; lateral margin with 10–14 spines posterior to postcervical groove; posterior margin of cervical groove with 3–4 spines between median and branchial carinae; posterior margin of carapace with 2 prominent submedian spines. Abdominal pleuron 2 without anterior spine. cl. 49 mm. WA, Qld, NSW, western Vic.; 144–1053 m depth. Polycheles typhlops Heller, 1862 (Fig. 41h). Frontal margin of carapace with 1 rostral spine; internal angle of orbital sinus spinose; anterior margin of carapace weakly spinose beyond outer angle of orbital notch; lateral margin with 24–26 spines posterior to postcervical groove;

156 Polychelida – deep-sea lobsters

posterior margin of carapace with 3–4 pairs of spines. cl. 57 mm. Atlantic, Mediterranean, Caribbean, Indian Ocean, WA, Qld, NSW; 183–2195 m depth. Polycheles typhlops has been reported from 300–500 metres on the North West Shelf (de Man, 1916; George, 1983). Griffin & Stoddart (1995) considered the eastern Australian specimens to be the subspecies P. typhlops typhlops.

Willemoesia Grote, 1873 The genus is abyssal, rare and little known; there were eight species (distinguished in a table by Gore, 1984) of which only four are now considered valid (Galil, 2000). One has been reported from southern Australia. It differs from other polychelids in having the last leg chelate in both sexes and lacking orbits. Diagnosis. Carapace with postorbital and postcervical carinae well defined, not swollen. Frontal margin without orbital sinus, eyestalks below and parallel to anterior margin of carapace. Antennule with basal article with interior lamellar process. Maxilliped 3 epipod longer than ischium. Pereopod 5 chelate in both sexes. Willemoesia pacifica Sund, 1920 (Figs 40i, 41i). Abdominal somite 6 smooth. cl. 50 mm. South- eastern Atlantic, South Pacific, SA; 28–2520 m depth. The species is known in Australia from a single specimen reported as W. leptodactyla by Bage (1938) and as W. bonaspei Kensley, 1968 by Griffin & Stoddart (1995).

References Ahyong, S., & Brown, D.E. 2002. New species and new records of Polychelidae from Australia (Crustacea: Decapoda). Raffles Bulletin of Zoology 50: 53–79. Bage, F. 1938. Crustacea Decapoda (in part). Australasian Antarctic Expedition, 1911–14. Scientific Reports. Series C. -Zoology and Botany 2: 1–13,pl.4. Galil, B. 2000. Crustacea Decapoda: review of the genera and species of the family Polychelidae Wood- Mason, 1874. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 21. Mémoires du Muséum National d’Histoire Naturelle, Paris 184: 285–387. George, R.W. 1983. New finds of deepwater ‘lobsters’ on the Northwest Shelf. FINS (Fisheries News), Perth 16: 16–20. Gore, R.H. 1984. Abyssal lobsters, genus Willemoesia (Palinura, Polychelidae), from the Venezuela Basin, Caribbean Sea. Proceedings of the Academy of Natural Science of Philadelphia 136: 1–11. Griffin, D.J.G., & Stoddart, H.E. 1995. Deep-water decapod Crustacea from eastern Australia: lobsters of the families Nephropidae, Palinuridae, Polychelidae and Scyllaridae. Records of the Australian Museum 47: 231–263. Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125: 1–292. de Man, J.G. 1916. Eryonidae, Palinuridae, Scyllaridae and Nephropsidae. Siboga Expéditie Monographie 23: 1–122. Wicksten, M.K. 1980. New records of Stereomastis sculpta pacifica (Faxon) (Decapoda: Polychelidae) in the eastern Pacific Ocean. Proceedings of the Biological Society of Washington 93: 914–919.

157 This page intentionally left blank 7. ASTACIDEA – SCAMPI AND CRAYFISH

The Astacidea Latreille, 1802 are the true freshwater and marine crayfish, scampi and lobsters. They are differentiated from most other decapods (except stenopodid and other specialised shrimps) usually by the possession of three pairs of chelate limbs of which the first is a strong claw. The southern Australian marine ‘rock lobsters’, otherwise called ‘crayfish’, are not astacideans but belong to Achelata and do not have chelate limbs or a rostrum. Much of our detailed knowledge of the anatomy of Crustacea was gained in early research on astacideans and T.H. Huxley’s (1880) classic work, The Crayfish: An Introduction to the Study of Zoology, is based on the common European freshwater crayfish, Astacus astacus Linnaeus. Scholtz & Richter (1995) undertook a phylogenetic analysis of the Decapoda and found that the Astacidea are not, in their view, a monophyletic group. They found that the Astacidae, Cambaridae and Parastacidae (essentially the freshwater crayfish) belong to a monophyletic taxon that they called Fractosternalia, sister taxon to Anomura, Thalassinidea and Brachyura. Schram (2001) and Schram & Ahyong (2002) confirmed their results and found that the Glypheoidea belong in Fractosternalia also. The taxon is characterised primarily in the articulation between thoracic sternites 7 and 8. Dixon et al. (2003) came to a different conclusion on the basis of a cladistic analysis of morphological characters. They united all the following taxa in a group for which they revived the name Astacura Borradaile, 1907. They retained ‘Astacida Latreille, 1802’ for a clade of the same taxa except Glypheoidea. Most Astacura possess telson spines and all have an unusual spine articulating between the basis and coxa of pereopod 1. Martin & Davis (2001) divided Astacidea into five superfamilies. The superfamily Glypheoidea Zittel [excluded from Astacidea by Dixon et al. (2003)] were a group of fossil species until the recent discovery of a living species in the Glypheidae Zittel, 1885. Neoglyphea inopinata Forest & de Saint Laurent, 1975 (Forest et al., 1976; Forest & de Saint Laurent, 1989) was first recorded from the Philippines but does occur in northern Australia (Forest & de Saint Laurent, 1989) along with a fossil species (Feldmann & de Saint Laurent, 2002). Their strong subchelate first limb and simple pereopods 2–5 make them similar to Palinura where they have been traditionally placed (Holthuis, 1991) but Forest & de Saint Laurent (1989) argued for their affinities with Astacidea. The living species is rare, living in the deep waters (186–300 m) of the Timor Sea near Australia. The superfamily Astacoidea Latreille which comprise two families (Astacidae Latreille, 1802 and Cambaridae Hobbs, 1942) of Northern Hemisphere freshwater crayfishes do not concern us here (see Hobbs, 1974, for a review of species). The superfamily Parastacoidea Huxley, of which there is a single family Parastacidae Huxley, 1879, are the Southern Hemisphere freshwater crayfishes, the most common called yabbies in Australia. The fauna is diverse in Australia and because many species are edible and some culti- vated commercially are of considerable interest. The family is the subject of several recent systematic reviews (Morgan, 1986, 1988; Horwitz, 1990; Morgan, 1997). Horwitz & Austin (1995) provided a key to all species. The superfamily Enoplometopoidea are the reef lobsters now placed in their own superfamily comprising one family and genus, Enoplometopus Milne Edwards, with nine species. De Saint

159 Marine Decapod Crustacea of Southern Australia

Laurent (1988) and Manning & Camp (1989) are critical reviews. One species occurs in southern Australia. The fifth superfamily contains marine species present in southern Australia. The Nephropoidea Dana, 1852 comprises two families. The Thaumastochelidae Bate, 1888 include two genera and four species of deepwater ‘pincer lobsters’ (Holthuis, 1974). The only species from Australian seas are tropical. Thaumastochelopsis wardi Bruce, 1988 has been recorded from 425 m on the Maria Plateau off Queensland (Bruce, 1988). Another, Thaumastocheles dochmiodon,is known from the Timor Sea (Chan & de Saint Laurent, 1999). The family is recognised by being blind and having one large chela with spinose fingers. Dixon et al. (2003) believed this family to be quite different from nephropids and enoplometopids. The other family, Nephropidae Dana, 1852 is the major one of Astacidea relevant to this book. Holthuis (1974; 1991) provided detailed and up-to-date treatments of the family.

Diagnosis. Lobster-like decapods with body divided into a cephalothorax and strong abdomen ending in broad tail fan. Uropodal exopod with suture. Carapace well developed, not fused to epistome, usually with prominent rostrum and divided by a cervical groove. Pereopods 1–3 chelate, first a strong claw, 4 and 5 with simple dactylus (or pereopod 1 a strong subchela, pereopods 2–5 simple in Glypheoidea). Pereopod 1 ischium with long proximal process articulating with coxa. Eggs hatching as mysis stage.

Key to southern Australian families of Astacidea 1. Body setose; pereopods 1 stout, chelate; pereopods 2–5 slender, subchelate (less so on 5); abdominal pleura rounded ...... Enoplometopidae — Body smooth; pereopods 1 slender, chelate; pereopods 2 and 3 slender, chelate, 4 and 5 simple; abdominal pleura acutely toothed ...... Nephropidae

References Bruce, A.J. 1988. Thaumastochelopsis wardi, gen. et sp. nov., a new blind deep-sea lobster from the Coral Sea (Crustacea: Decapoda: Nephropidea). Invertebrate Taxonomy 2: 903–914. Chan, T.-Y., & de Saint Laurent, M. 1999. The rare lobster genus Thaumastocheles (Decapoda: Thaumastochelidae) from the Indo-Pacific, with description of a new species. Journal of Crustacean Biology 19: 891–901. Dixon, C.J., Ahyong, S., & Schram, F.R. 2003. A new hypothesis of decapod phylogeny. Crustaceana 76: 935–975. Feldmann, R.M., & de Saint Laurent, M. 2002. Glyphea foresti n. sp. (Decapoda) from the Cenomanian of Northern Territory, Australia. Crustaceana 75: 359–374. Forest, J., & de Saint Laurent, M. 1989. Nouvelle contribution à la connaissance de Neoglyphea inopinata Forest & de Saint Laurent, à propos de la description de la femelle adulte. In: Forest, J. (ed.), Résultats des Campagnes MUSORSTOM, vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144: 75–92. Forest, J., de Saint Laurent, M., & Chace, F.A. 1976. Neoglyphea inopinata: a crustacean ‘living fossil’. Science 192: 884. Hobbs, H.H. 1974. Synopsis of the families and genera of crayfishes (Crustacea: Decapoda). Smithsonian Contributions to Zoology 164: 1–32. Holthuis, L.B. 1974. Biological results of the University of Miami Deep-Sea Expeditions. 106. The lobsters of the superfamily Nephropidea of the Atlantic Ocean (Crustacea: Decapoda). Bulletin of Marine Science 24: 723–884. Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125: 1–292.

160 Astacidea – scampi and crayfish

Horwitz, P. 1990. A taxonomic revision of species in the freshwater crayfish genus Engaeus Erichson (Decapoda: Parastacidae). Invertebrate Taxonomy 4: 427–614. Horwitz, P., & Austin, C.M. 1995. Preliminary key to the species of Australian freshwater crayfish (Parastacidae). Pp. 51–59 in: Horwitz, P. (ed.) A preliminary key to the species of Decapoda (Crustacea: Malacostraca) found in Australian inland waters (Identification Guide No. 5). Cooperative Research Centre for Freshwater Ecology: Albury. Huxley, T.H. 1880. The Crayfish. An Introduction to the Study of Zoology (International Scientific Series Vol. 28). Paul & Co.: London. 371 pp. Manning, R.E.B., & Camp, D.K. 1989. Additional records for an Atlantic reef lobster, Enoplometopus antillensis Lütken, 1865 (Crustacea, Decapoda, Enoplometopidae). Proceedings of the Biological Society of Washington 102: 411–417. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. Morgan, G.J. 1986. Freshwater crayfish of the genus Euastacus (Decapoda: Parastacidae) from Victoria. Memoirs of the Museum of Victoria 47: 1–57. Morgan, G.J. 1988. Freshwater crayfish of the genus Euastacus Clark (Decapoda: Parastacidae) from Queensland. Memoirs of the Museum of Victoria 49: 1–49. Morgan, G.J. 1997. Freshwater crayfish of the genus Euastacus Clark (Decapoda: Parastacidae) from New South Wales, with a key to all the species of the genus. Records of the Australian Museum, Supplement 23: 1–110. de Saint Laurent, M. 1988. Enoplometopoidea, nouvelle superfamille de Crustacés Décapodes Astacidea. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 307: 59–62. Scholtz, G., & Richter, S. 1995. Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113: 289–328. Schram, F.R. 2001. Phylogeny of decapods: moving towards a consensus. Hydrobiologia 449: 1–20. Schram, F.R., & Ahyong, S. 2002. The higher affinities of Neoglyphea inopinata in particular and the Glypheoidea (Decapoda, Reptantia) in general. Crustaceana 75: 629–636.

Enoplometopidae de Saint Laurent, 1988 On first impression, reef lobsters look like deep-water scampi, lobsters of the family Nephropidae. But their taxonomic affininities have long been enigmatic. They were placed in this family until Holthuis (1974) removed them to the thalassinidean family Axiidae, which reef lobsters also resemble. De Saint Laurent (1988) returned them to the Astacidea and erected a new superfamily to contain the sole genus Enoplometopus. Two species occur in Australia. This is the first record of a species as far south as Sydney. Diagnosis. Body cylindrical, with tufts of long stiff setae; cervical groove weak. Carapace with well developed rostrum with lateral spines. Eyes pigmented. Antennae long; scaphocerite with inner margin unarmed, curved. Chelipeds equal, robust. Walking legs subchelate, less so on last. Abdominal pleura rounded, sometimes with spines. Tailfan entirely calcified; telson with moveable spines, posterior margin convex. Male pleopod 2 with appendix masculina.

Enoplometopus Milne Edwards, 1862 Enoplometopus is the only genus but two subgenera or genera have been suggested (Holthuis, 1983) and rejected (Poupin, 2003). Chan & Yu (1998) and Poupin (2003) provided keys to eleven species. All are brightly coloured, usually patterns over a background of red (see photos in Holthuis, 1983; Chan & Yu, 1998) and have become popular as aquarium pets. They generally inhabit coral reefs from shallow water to shelf depths. Diagnosis. As for family.

161 Marine Decapod Crustacea of Southern Australia

Enoplometopus occidentalis (Randall, 1840) reef lobster (Fig. 43g, Pl. 11d). Carapace with 1 large postcervical tooth; 2 intermediate teeth (between supraocular and median teeth), 4 median teeth. Abdominal pleura broadly convex. Telson with 1 pair of lateral spines. Orange-red with conspicuous white spots on abdomen, fewer on carapace; posterior pereopods with alter- natinng white and orange bands (colour photo in Healy & Yaldwyn, 1970). cl. 60 mm. Indo- West Pacific, Qld, NSW (off Sydney); coral reefs, subtidal rocky reef to 100 m depth.

References Chan, T.-Y., & Yu, H.-P. 1998. A new reef lobster of the genus Enoplometopus A. Milne Edwards, 1862 (Decapoda, Nephropoidea) from the western and southern Pacific. Zoosystema 20: 183–192. Healy, A., & Yaldwyn, J.C. 1970. Australian Crustaceans in Colour. Reed: Sydney. 112 pp. Holthuis, L.B. 1946. Biological results of the Snellius Expedition XIV. The Decapoda Macrura of the Snellius Expedition I. The Stenopodidae, Nephropsidae, Scyllaridae and Palinuridae. Temminckia 7: 1–178,11pls. Holthuis, L.B. 1974. Biological results of the University of Miami Deep-Sea Expeditions. 106. The lobsters of the superfamily Nephropidea of the Atlantic Ocean (Crustacea: Decapoda). Bulletin of Marine Science 24: 723–884. Holthuis, L.B. 1983. Notes on the genus Enoplometopus, with descriptions of a new subgenus and two new species (Crustacea, Decapoda, Axiidae). Zoologische Mededelingen, Leiden 56: 281–298. Poupin J. 2003. Reef lobsters Enoplometopus A. Milne Edwards, 1862 from French Polynesia, with a brief revision of the genus (Crustacea, Decapoda, Enoplometopidae). Zoosystema 25: 643–64. de Saint Laurent, M. 1988. Enoplometopoidea, nouvelle superfamille de Crustacés Décapodes Astacidea. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 307: 59–62.

Nephropidae Dana, 1852 The best known nephropids are the scampi, Nephrops norvegicus (Linnaeus), which is the basis of a fishery in Europe, and the Maine lobster, Homarus americanus Milne Edwards, which is also fished. However, most species are distributed in the tropical Indo-West Pacific. The Nephropidae include 44 known species in 10 genera and three subfamilies. Holthuis (1991) and Macpherson (1993) provided valuable keys. Three genera are known from Australia with greatest representa- tion of species on the tropical North West Shelf. In that area there is a significant fishery for species of Metanephrops. The key includes these potentially commercial species from northern Australia and New Zealand that are unlikely to be found on southern coasts: Acanthacaris tenuimana Bate, 1888 (Indo-West Pacific, Qld; 600–2161 m); Metanephrops arafurensis (de Man, 1905) (Arafura Sea; 560 m); M. australiensis (Bruce, 1966) (north-western Australia; 418–500 m); M. challengeri (Balss, 1914) (New Zealand, shelf depths); M. neptunus (Bruce, 1965) (north-western Australia; 238–940 m); M. sibogae (de Man, 1916) (Indonesia, northern Australia; 260–480 m); Nephropsis holthuisi Macpherson, 1993 (Indonesia, north-western Australia, NSW; 900–1105 m); N. stewarti Wood-Mason, 1873 (Indo-West Pacific, northern WA; 170–1060 m); and N. sulcata Macpherson, 1990 (Indo-West Pacific, Qld, NSW; 415–1115 m). Tshudy & Babcock (1997) related the extant and fossil genera of nephropids to another fossil family. Rode & Babcock (2003) found that both families were basal to a clade containing freshwater crayfishes.

Diagnosis. Body cylindrical, without setae or with tomentum; cervical groove deep. Carapace with well developed rostrum, usually with lateral spines. Eyes pigmented, sometimes feeble. Antennae long; scaphocerite with inner margin unarmed, curved. Chelipeds equal, robust. Walking legs 1–3 chelate, 4 simple. Abdominal pleura acute. Tailfan entirely calcified; telson with fixed spines, posterior margin convex. Male pleopod 2 without appendix masculina.

162 Astacidea – scampi and crayfish

Key to species of Nephropidae from waters near Australia and New Zealand * species from southern Australia 1. Rostrum laterally compressed for most of length, without dorsal and ventral teeth; carapace with branchiostegal spine; body uniformly spinulose; telson with about 8 lateral spines ...... Acanthacaris tenuimana — Rostrum dorsoventrally depressed, with lateral teeth; carapace without branchiostegal spine; body ornamented but not uniformly spinulose; telson with at most 3 lateral spines ...... 2 2. Scaphocerite absent; carapace without postorbital spine; eyes small, non-pigmented ...... Nephropsis … 3 — Scaphocerite present; eyes large, pigmented ...... Metanephrops … 9 3 Uropodal exopod without suture ...... Nephropsis suhmi* — Uropodal exopod with suture ...... 4 4. Telson with an erect basal spine at midpoint ...... Nephropsis acanthura* — Telson without a basal spine ...... 5 5. Rostrum with 2 pairs of lateral spines ...... Nephropsis sulcata — Rostrum with 1 pair of lateral spines ...... 6 6. Abdominal somites with middorsal carina ...... 7 — Abdominal somites without middorsal carina ...... 8 7. Pereopod 1 merus reaching as far as end of rostrum; subdorsal ridge granular ...... Nephropsis holthuisi* — Pereopod 1 merus reaching as far as lateral spine on rostrum; subdorsal ridge spinose ...... Nephropsis macphersoni* 8. Subdorsal carinae granulate ...... Nephropsis stewarti — Subdorsal carinae with at least 3 well-developed spines ...... Nephropsis serrata* 9. Carapace smooth between ridges and large spines ...... 10 — Carapace uniformly spinulose ...... 13 10. Chelae heavily ridged and spinulose ...... Metanephrops velutinus* — Chelae weakly ridged and finely granular ...... 11 11. Without longitudinal cardiac ridge ...... Metanephrops challengeri — Spinulose longitudinal cardiac ridge present ...... 12 12. Pereopod 1 merus with distinct spine in middle of inner margin . . . Metanephrops boschmai* — Pereopod 1 merus without such a spine ...... Metanephrops sibogae (Pl. 11f) 13. Region between postrostral carinae heavily spinulose ...... Metanephrops neptunus — Region between postrostral carinae smooth ...... 14 14. Lateral longitudinal furrows on abdominal somites ...... Metanephrops arafurensis — No lateral longitudinal furrows on abdominal somites . . . . Metanephrops australiensis (Pl. 11e)

Metanephrops Jenkins, 1972 Metanephrops was described for 13 living species and some fossils most of which had been previously assigned to Nephrops. The latter is now confined to the European scampi, N. norvegicus. Chan (1997) provided a key to species and discussed relationships and ecology. In Australia there is a trawl fishery for scampi on the North West Shelf. Catches are small and the lobsters are sold as an expensive luxury food. Commercial quantities have only been recently

163 Marine Decapod Crustacea of Southern Australia

discovered and biological data relevant to the fishery has been gathered by CSIRO Division of Fisheries. The most abundant species in the area were M. australiensis (Pl. 11e) and M. velutinus but their distribution and that of the other species is patchy. In general scampi are found on the upper slope, between about 300 and 500 m depth, on soft muddy sediments. All species are reddish or pinkish with white. Females carry about 300 blue eggs each. A species similar to M. velutinus, M. andamanicus, was studied by Berry (1969) in South Africa where a small fishery also exists. Only two species have been recorded from southern Australia. The New Zealand species, M. challengeri (Balss, 1914), is included in the key because it is imported for sale in Australian fish markets.

Diagnosis. Rostrum elongate with 1 pair of lateral spines; lateral carina extending on to carapace bearing 3–5 prominent spines. Antenna spine lamellate; scaphocerite broad. Eyes pigmented. 7 longitudinal ridges on carapace posterior to cervical groove. Chelipeds almost symmetrical. (rewritten from Jenkins, 1972)

Fig. 42. Nephropidae. a, Nephropsis serrata. b, Metanephrops boschmai.

164 Astacidea – scampi and crayfish

Metanephrops boschmai (Holthuis, 1964) Bight lobster (FAO), Bight scampi, Boschma’s scampi (Fig. 42a, Pl. 11f). Sides of carapace smooth; longitudinal spinulose cardiac ridge present. Abdominal somites without transverse grooves. Chelae weakly ridged and finely granular, merus with distinct spine in middle of inner margin. 180 mm. WA, NT, SA (Great Australian Bight); 300–500 m depth. There is a small fishery incorporating this species (Wallner & Phillips, 1988). Metanephrops velutinus Chan & Yu, 1991 velvet lobster (FAO), velvet scampi (Fig. 43a). Sides of carapace smooth, furry; postrostral carinae with at most 3 teeth. Abdominal somites weakly sculptured, fifth without spines on lateral carina. Chelae ridged and spinulose but without large spines. 180 mm. Patterned in yellowish-pink, pink and white. Philippines, Indonesia, WA, NT, Qld, SA (Great Australian Bight); 238–702 m depth. This species contributes most to the scampi fishery of the North West Shelf (Wallner & Phillips, 1988). It was previously confused with M. andamanensis.

f g

bc d e

Fig. 43. Nephropidae. a, Metanephrops velutinus (carapace, lateral). b, Nephropsis holthuisi (carapace and left cheliped, dorsal). c, Nephropsis macphersoni (carapace and left cheliped, dorsal). d, Nephropsis acanthura (carapace, dorsal). e, Nephropsis suhmi (carapace, dorsal). f, Metanephrops velutinus (cheliped). Enoplometopidae. g, Enoplometopus occidentalis

165 Marine Decapod Crustacea of Southern Australia

Nephropsis Wood-Mason, 1873 A deep-water genus of 14 species worldwide (Macpherson, 1990, 1993; Holthuis, 1991) is represented in southern Australia by four species. Unlike Metanephrops, none is of commercial interest. Diagnosis. Eyes small, not pigmented. Antenna without scaphocerite. Rostrum without ventral teeth. Abdominal pleura not overlapping, second acutely pointed. Nephropsis acanthura Macpherson, 1990 spinetail lobsterette (FAO) (Fig. 43b). Rostrum with 1 pair of lateral spines. Dorsal surface of telson with an erect basal spine at midpoint; uropodal exopod with suture. cl. 60 mm. Indo-West Pacific, WA, Qld, NSW, SA (Great Australian Bight); 750–1305 m depth, rare. Nephropsis holthuisi Macpherson, 1993 (Fig. 43c). Rostrum with 1 pair of lateral spines; subdorsal ridge granular. Pereopod 1 merus reaching as far as end of rostrum. Abdominal somites with middorsal carina; telson without a basal spine on dorsal surface; uropodal exopod with suture. cl. 60 mm. Indonesia, NSW, WA (off Ashmore Reef); 900–1105 m depth. Nephropsis macphersoni Watabe & Iizuka, 1999 (Fig. 43d). Rostrum with 1 pair of lateral spines; subdorsal ridge spinose. Pereopod 1 merus reaching as far as spine on rostrum. Abdominal somites with middorsal carina; telson without a basal spine on dorsal surface; uropodal exopod with suture. cl. 40 mm. NSW; 900–1135 m depth, rare. The species was previously recorded as N. holthuisi (Griffin & Stoddart, 1995; Watabe & Iizuka, 1999). Nephropsis serrata Macpherson, 1993 (Fig. 42b). Rostrum with 1 pair of lateral spines; subdorsal carinae with 2–5 teeth, commonly 3. Abdominal somites without median carinae; telson without a basal spine on dorsal surface; uropodal exopod with suture. cl. 49 mm. Indonesia, SA (Great Australian Bight), WA; 390–1430 m depth. Nephropsis suhmi Bate, 1888 (Fig. 43e). Rostrum with 3 pairs of lateral spines. Uropodal exopod without suture. Abdominal somite 2 with tooth on anterior margin of pleura. cl. 59 mm. Arabian Sea, Indonesia, Qld, NSW, SA, WA (Great Australian Bight); 786–2029 m depth.

References Berry, P.F. 1969. The biology of Nephrops andamanicus Wood-Mason (Decapoda, Reptantia). South African Association for Marine Biological Research, Investigational Report 22: 1–55. Chan, T.-Y. 1997. Crustacea Decapoda: Palinuridae, Scyllaridae and Nephropidae collected in Indonesia by the KARUBAR cruise with an identification key for the species of Metanephrops In: Crosnier, A and Bouchet, P. (eds), Résultats des Campagnes MUSORSTOM, vol. 16. Mémoires du Muséum National d’Histoire Naturelle, Paris 172: 409–431. Chan, T.-Y., & Yu, H.-P. 1991. Studies on the Metanephrops japonicus group (Decapoda, Nephropidae), with descriptions of two new species. Crustaceana 60: 18–51. Griffin, D.J.G., & Stoddart, H.E. 1995. Deep-water decapod Crustacea from eastern Australia: lobsters of the families Nephropidae, Palinuridae, Polychelidae and Scyllaridae. Records of the Australian Museum 47: 231–263. Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125: 1–292. Jenkins, R.J.F. 1972. Metanephrops, a new genus of late Pliocene to Recent lobsters, (Decapoda, Nephropidae). Crustaceana 22: 161–77, pls 1, 2. Macpherson, E. 1990. Crustacea Decapoda: on a collection of Nephropidae from the Indian Ocean and Western Pacific. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145: 289–328. Macpherson, E. 1993. New records for the genus Nephropsis Wood-Mason (Crustacea, Decapoda,

166 Astacidea – scampi and crayfish

Nephropidae) from northern Australia, with the description of two new species. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 10: 55–66. Rode, A.L., & Babcock, L.E. 2003. Phylogeny of fossil and extant freshwater crayfish and some closely related nephropid lobsters. Journal of Crustacean Biology 23: 418–435. Tshudy, D., & Babcock, L.E. 1997. Morphology-based phylogenetic analysis of the clawed lobsters (family Nephropidae and the new family Chilenohphoberidae). Journal of Crustacean Biology 17: 253–263. Wallner, B., & Phillips, B. 1988. From scampi to deepwater prawns: developments in the Northwest Shelf deepwater trawl fishery. Australian Fisheries 47: 34–38. Watabe, H., & Iizuka, E. 1999. A new species of the bathyal lobster genus Nephropsis (Crustacea: Decapoda: Nephropidae) from Australian waters, with a redescription of N. holthuisi. Species Diversity 4: 371–380.

167 This page intentionally left blank 8. THALASSINIDEA – GHOST SHRIMPS AND SPONGE SHRIMPS

The Thalassinidea Latreille, 1831 are a loose assemblage of shrimp-like reptant decapods united by the possession of a dense row of long setae on the lower margin of pereopod 2. The group has traditionally held together by the possession of a linea thalassinica (Fig. 44a), one on each side of the cephalothorax, a longitudinal hinge line between the dorsal regions and the branchiostegal region or side-flap covering the branchial chamber. Although it is well expressed in the Callianassidae where it runs from the base of the antennae straight back to the post margin of cephalothorax, in other families it is variously reduced and or absent. Dixon et al.’s (2003) analysis of morphological characters in Decapoda recognised ‘Thalassinida’, well defined by a curved articulation between ischium and merus of pereopod 1, the long setae on pereopod 2, and enlarged thoracic sternite 7. On the other hand, Morrison et al. (2002) and Tudge & Cunningham (2002) found equivocal support for the monophyly of Thalassinidea on the basis of molecular data. Tudge & Cunningham (2002) found weak support for the Upogebiidae, Axianassidae, Thalassinidae and Laomediidae being more closely related to some Palinura and Anomura than to other thalassinideans. They were more confident that these families belonged to a single clade distinct from a second clade comprising Strahlaxiidae and Callianassidae. This dichotomy differs from that proposed by Poore (1994) on the basis of morphology. He found axiids, strahlaxiids and calocaridids to be the most isolated and not close to callianassids. Dixon et al. (2003) raised new doubts about the relationships of families. Most of the other characters traditionally used to define the Thalassinidea are valid for other reptant groups but there are useful give-aways for distinguishing them from caridean shrimps and other shrimp-like reptants. In most thalassinideans the legs vary in form. Pereopod 1 is a larger cheliped, chelate or subchelate, sometimes asymmetrical. Pereopods 2–4 may be flattened, especially distally. Pereopod 5 is smaller than the others, is often minutely chelate or subchelate, and its coxae are well separated from those of more anterior limbs which form a compact ventral mass. The rostrum is often short and if prominent is flattened dorsoventrally. The abdominal somites are weakly articulating and have weak pleura. The familial composition has been fluid but Poore’s (1994) phylogenetic analysis recognised three supefamilies containing eleven families into which 73 genera are placed. He provided keys to all families and genera. Seven genera have been added since that date. Not all families are represented in southern Australia. The Thalassinidae (only family of Thalassinoidea) are represented in tropical mangrove forests of Australia by the mud shrimp, Thalassina squamifera de Man, 1915, Calocarididae by some deep-water species in northern Australia, and Thomassiniidae by species of Thomassinia in sediments of the Great Barrier Reef (Poore & Griffin, 1979; Poore, 1997). The monotypic Callianideidae do not occur in Australia. Most thalassinideans are burrowers in soft sediments or live under rocks or in crevices (Murphy & Kremer, 1992; Nickell & Atkinson, 1995; Astall et al., 1997).

169 Marine Decapod Crustacea of Southern Australia

a c

f e

h i

l j k

Fig. 44. Thalassinidea. a, Linea thalassinica present. b, pleuron of abdominal somite 1 produced (Paraxiopsis brocki, Axiidae). c, pleuron of abdominal somite 1 not produced (Michelea microphylla, Micheleidae from Poore, 1997). d, abdominal somite 1 with anterolateral lobe (Laomediidae). e, pereopod 3 propodus with heel (Callianassidae). f, mandibular incisor not toothed, asymmetrical (Axiidae). g, maxilla 2 with several long setae on posterior lobe (Laomedia healyi). h, pleopods 2–5 laterally lobed (Micheleidae). i, pleopod 3 rami lanceolate (Ctenochelidae). j, uropodal exopod with secondary lobe (Eucalliax aequimanus, Callianassidae). k, uropodal endopod ovate (Axiopsis werribee, Axiidae). l, uropodal endopod triangular (Strahlaxius waroona, Strahlaxiidae).

Diagnosis. Reptant decapods without a thelycum in the female. Maxilliped 1 without arthrobranch. Pereopods 1–5 with basis and ischium fused. Pereopod 1 chelate and carpus-propodus articulation slight. Pereopod 2 chelate or simple, always with dense row of long setae on lower margin. Pereopod 3 simple. Pereopod 4 simple or subchelate. Pereopod 5 chelate or subchelate.

170 Thalassinidea – ghost shrimps and sponge shrimps

Key to southern Australian superfamilies and families of Thalassinidea 1. Linea thalassinica absent; antennule with article 3 about as long as article 2 . . Axioidea ...2 — Linea thalassinica present (Fig. 44a) at least anteriorly; antennule with article 3 usually longer than 2 ...... Callianassoidea . ..4 2. Abdominal somite 2 less than twice as long as 1; pleuron of abdominal somite 1 produced (Fig. 44b); coxa of pereopod 4 more or less cylindrical; propodus of pereopods 3 and 4 more or less linear; uropodal endopod ovate (Fig. 44k); pereopods 2–4 and abdominal somites without setal-rows ...... Axiidae … p. 173 — Abdominal somite 2 twice as long as 1; pleuron of abdominal somite 1 not produced (Fig. 44c); coxa of pereopod 4 flattened; propodus of pereopods 3 and 4 more or less flattened; uropodal endopod triangular (Fig. 44l) or ovate; some of pereopods 2–4 and abdominal somites with setal-rows ...... 3 3. Rostrum spinose, apically bifid; exopods of pleopods 2–5 not laterally lobed; longitudinal and vertical setal-rows on carapace absent; antennule, article 1 as long as 2; mandibular incisor toothed and symmetrical; epipods and podobranchs reduced posteriorly ...... Strahlaxiidae ...p.178 — Rostrum not spinose; exopods of pleopods 2–5 laterally lobed (Fig. 44h); longitudinal and vertical setal-rows on carapace present; antennule, article 1 longer than 2; mandibular incisor not toothed and asymmetrical (Fig. 44f); epipods and podobranchs rarely reduced posteriorly ...... Micheleidae ...p.176 4. Posterior margin of carapace with lateral lobes; abdominal somite 1 with anterolateral lobe (Fig. 44d); maxilla 2 scaphognathite with several long setae on posterior lobe (Fig. 44g) ...... Laomediidae ...p.187 — Posterior margin of carapace evenly curved; abdominal somite 1 without anterolateral lobes; maxilla 2 scaphognathite with 1 or without long seta on posterior lobe ...... 5 5. Pereopods 1 equal; pereopod 2 simple; rostrum usually broad and setose; eyestalks cylindrical; coxa of pereopod 4 cylindrical ...... Upogebiidae ...p.189 — Pereopods 1 unequal, rarely equal; pereopod 2 chelate; rostrum obsolete; eyestalks flattened; coxa of pereopod 4 flattened ...... 6 6. Pleopod 2 similar to pleopods 3–5, rami lanceolate (Fig. 44i); pereopod 3 propodus linear or weakly ovate; uropodal exopod simply ovate ...... Ctenochelidae ...p.186 — Pleopod 2 reduced, sexually modified; pleopods 3–5 with broad interacting rami; pereopod 3 propodus with proximal heel on lower margin (Fig. 44e); uropodal exopod with secondary setose lobe (Fig. 44j) ...... Callianassidae ...p.180

171 Marine Decapod Crustacea of Southern Australia

Murphy, R.C., & Kremer, J.N. 1992. Benthic community metabolism and the role of deposit-feeding callianassid shrimp. Journal of Marine Research 50: 321–340. Nickell, L.A., & Atkinson, R.J.A. 1995. Functional morphology of burrows and trophic modes of three thalassinidean shrimp species, and a new approach to the classification of thalassinidean burrow morphology. Marine Ecology Progress Series 128: 181–197. Poore, G.C.B., & Griffin, D.J.G. 1979. The Thalassinidea (Crustacea: Decapoda) of Australia. Records of the Australian Museum 32: 217–321. Poore, G.C.B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54: 79–120. Poore, G.C.B. 1997. A review of the thalassinidean families Callianideidae Kossmann, Micheleidae Sakai, and Thomassiniidae de Saint Laurent (Crustacea: Decapoda) with descriptions of fifteen new species. Zoosystema 19: 345–420. Tudge, C.C., & Cunningham, C.W. 2002. Molecular phylogeny of the mud lobsters and mud shrimps (Crustacea: Decapoda: Thalassinidea) using nuclear 18s rDNA and mitochondrial 16s rDNA. Invertebrate Systematics 16: 839–847.

COMMON SOUTHERN AUSTRALIAN GHOST SHRIMPS AND SPONGE SHRIMPS Shrimp-like animals belong to a range of quite different higher taxa. Thalassinidean shrimps: • have either one claw stronger than the other or two similar strong claws, sometimes one or both flattened (never thin claws) • live in burrows or under rocks, never freely walking • have a fringe of long setae on the lower margin of the second leg Although there are numerous species of thalassinideans in several families, few are common. Only the intertidal ghost shrimp Trypaea australiensis would be familiar to most shore collectors. The others are rarely seen. A green alpheid shrimp, Alpheus richardsoni, is sometimes found with the ghost shrimp in seagrass meadows but it snapping fingers give it away. This simple key separates the families most likely to be encountered intertidally in southern Australia. Only resorting to the more technical keys will differentiate the species. 1. Soft-bodied, elongate; usually with one claw much bigger than the other; pink or translucent; in burrows on intertidal mudflats ...... Callianassidae (ghost shrimps) — Compact shrimps; claws more or less similar in size ...... 2 2. Both uropodal rami with sutures; in burrows among mangroves ...... Laomedia healyi (mangrove shrimps) — One or neither uropodal ramus with suture ...... 3 3. One uropodal ramus with suture ...... Axiidae (sponge shrimps) — Neither uropodal ramus with suture ...... 4 4. Flat rostrum, marginally dentate ...... Strahlaxiidae (sponge shrimps) — Very setose rostrum ...... Upogebiidae (sponge shrimps)

172 Thalassinidea – ghost shrimps and sponge shrimps

Superfamily Axioidea Huxley, 1879 Three of four families are found in Australia. Only Calocarididae Ortmann, 1891 is absent. Axioids are more compact shrimps than the flexible, elongate callianassoids. The linea thalassinica is absent and article 3 of the antennule is about as long as article 2.

Axiidae Huxley, 1879 The Axiidae are represented by more than 100 species in 22 genera and have been extensively reviewed by Sakai & de Saint Laurent (1989). This paper contains many errors and should be used with caution. Poore (1994) provided a key to all the genera but three have been added since that date. The family is most diverse in tropical seas where many undescribed species are to be found. Little is known of axiid biology but they are generally weak burrowers or live under stones or in cavities in coral rubble. One species, Axius serratus, build burrows up to 3 m deep. Sakai (1994) provided a key for the identification of 16 named Australian species. He included the two species of Strahlaxius (Strahlaxiidae) in Axiidae. The review of Poore & Griffin (1979) has been largely superseded with the discovery of new species and a new generic arrangement (Kensley, 1989; Sakai & de Saint Laurent, 1989; Poore, 1994). Four species in three genera occur in southern Australia but none is commonly found intertidally. The two species assigned to Axiopsis should be placed in another genus.

Diagnosis. Rostrum usually spinose, apically acute; eye usually pigmented; pleuron of abdominal somite 1 produced; abdominal somite 2 less than twice as long as 1; propodus of pereopods 3 and 4 linear or broadened; coxa of pereopod 4 more or less cylindrical; male pleopod 2 endopod foliaceous with small appendix masculina attached mesially; exopods of pleopods 2–5 not laterally lobed; uropodal endopod ovate; pereopods 2–4 without setal-rows, abdominal somite 6 with longitudinal setal-row.

Key to southern Australian species of Axiidae 1. Gastric region elevated and separate from rostrum ...... Dorphinaxius kermadecensis — Gastric region convex and continuously curved down to rostrum ...... 2 2. Antenna acicle short, bifurcate; pleopods 2–5 without appendix interna . . . Paraxiopsis brocki — Antenna acicle long, simple; pleopods 2–5 with appendix interna ...... Axiopsis ...3 3. Submedian carinae finely tuberculate in gastric region; middorsal carina extending on to rostrum as a tuberculate ridge ...... Axiopsis australiensis — Submedian carinae unarmed in gastric region; middorsal carina ending anteriorly as 1–3 teeth ...... Axiopsis werribee

Axiopsis Borradaile, 1903 The two species from southern Australia were listed by Poore & Griffin (1979) in Axiopsis and erroneously listed in two other different genera by Sakai & de Saint Laurent (1989), Sakai (1994), and Davie (2002). Probably they deserve their own genus and the diagnosis given is true only of these two species. Diagnosis. Rostrum triangular, longer than eyestalks. Antenna acicle long, simple. Gastric region convex, with middorsal spines and carina and submedian carinae. Pleurobranchs 5–7 present. Pereopods 1 subequal. Pereopods 3 and 4 propodi with transverse rows of spiniform setae. Telson

173 Marine Decapod Crustacea of Southern Australia

longer than wide, rounded. Pleopods 1 of male present; pleopods 2 of male with appendix interna and appendix masculina. Uropodal exopod with transverse suture. Axiopsis australiensis de Man, 1925 (Figs 45a, b, 46a). Submedian carinae finely tuberculate in gastric region; middorsal carina extending on to rostrum as a tuberculate ridge. NSW, Vic.; intertidal. Axiopsis werribee Poore & Griffin, 1979 (Figs 45c, d, 46b, Pl. 11h). Submedian carinae unarmed in gastric region; middorsal carina ending anteriorly as 1–3 teeth. Vic. (Port Phillip Bay), Tas.; 10–25 m depth.

b a

c d

h g

Fig. 45. Axiidae. Dorsal and lateral views of carapace: a, b, Axiopsis australiensis. c, d, Axiopsis werribee. e, f, Dorphinaxius kermadecensis. g, h, Paraxiopsis brocki.

174 Thalassinidea – ghost shrimps and sponge shrimps

b a

c d

Fig. 46. Axiidae. Telson and left uropod: a, Axiopsis australiensis. b, Axiopsis werribee. c, Dorphinaxius kermadecensis. d, Paraxiopsis brocki.

Dorphinaxius Sakai & de Saint Laurent, 1989 The steeply sloping frontal profile of Dorphinaxius kermadecensis, the only species in the genus, is characteristic. A similar genus, Scytoleptus Gerstaecker, is represented in northern Australia by an undescribed species which has frequently been confused with S. serripes Gerstaecker, 1856 (see Poore & Griffin, 1979). Diagnosis. Rostrum short, trilobed, little longer than eyestalks. Antenna acicle comma-shaped. Gastric region conspicuously elevated from level of rostrum, with middorsal and submedian carinae. Pleurobranchs 5–7 small. Pereopods 1 strongly unequal. Pereopods 3 and 4 propodi with transverse rows of spiniform setae. Telson longer than wide, rounded. Pleopods 1 of male absent; pleopods 2–5 with appendix interna, sexually dimorphic, those of female more slender; pleopod 2 of male with appendix masculina. Uropodal exopod with transverse suture. Dorphinaxius kermadecensis (Chilton, 1911) (Figs 45e, f, 46c). Poore & Griffin (1979) described this as D. appendiculis. Kermadec Is, New Zealand, central NSW; intertidal–8 m depth.

Paraxiopsis de Man, 1905 Nine species are known, most from the Atlantic (Kensley, 1996). Only one reaches the southern coast of Australia. Diagnosis. Rostrum distinctly set at lower level than anterior carapace, apically acute, marginally armed, longer than eyestalks. Antenna acicle short, bifurcate. Gastric region convex, with middorsal carina entire, submedian and dorsolateral carinae usually dentate. Pleurobranchs absent. Pereopods 1 subequal. Pereopods 3 and 4 propodi with transverse rows of spiniform setae. Telson about as long as wide, with pair of oblique spinose carinae, with posteromedian spine. Pleopods 1 of male absent; pleopods 2–5 without appendix interna; pleopod 2 of male with appendix masculina. Uropodal exopod with transverse suture.

175 Marine Decapod Crustacea of Southern Australia

Paraxiopsis brocki (de Man, 1888) (Figs 45g, h, 46d). Indonesia, NT, WA; intertidal to shallow reef. The species can be recognised in southern Australia by the strong spines on the rostrum and anterodorsal margin of the cephalothorax and the oblique rows of spines on the telson. The species has been previously placed in the genera Axius, Axiopsis and Eutrichocheles.

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells,A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A. CSIRO Publishing: Melbourne. xii, 551 pp. Kensley, B. 1989. New genera in the thalassinidean families Calocarididae and Axiidae (Crustacea: Decapoda). Proceedings of the Biological Society of Washington 102: 960–967. Kensley, B. 1996. The genus Paraxiopsis De Man, with descriptions of new species from the Western Atlantic (Crustacea: Decapoda: Axiidae). Bulletin of Marine Science 58: 709–729. Poore, G.C.B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54: 79–120. Poore, G.C.B., & Griffin, D.J.G. 1979. The Thalassinidea (Crustacea: Decapoda) of Australia. Records of the Australian Museum 32: 217–321. Sakai, K. 1994. Eleven species of Australian Axiidae (Crustacea: Decapoda: Thalassinidea) with descriptions of one new genus and five new species. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11: 175–202. Sakai, K., & de Saint Laurent, M. 1989. A check list of Axiidae (Decapoda, Crustacea, Thalassinidea, Anomula), with remarks and in addition descriptions of one new subfamily, eleven new genera and two new species. Naturalists, Publications of Tokushima Biological Laboratory, Shikoku University 3: 1–104.

Micheleidae Sakai, 1992 The Micheleidae are essentially a tropical family of four genera and about 24 species (Poore, 1994, 1997). Many species are associated with coral environments where they are probably weak burrowers. The combination of setal-rows and the form of the equal first pereopods are diagnostic. Only two species have been reported from southern Australia, each represented by a single specimen. The Victorian species, Michelea microphylla, was listed as ‘vulnerable’ under IUCN criteria by O’Hara (2002). Two species, Michelea leura (Poore & Griffin) and M. paraleura Poore are known from the Great Barrier Reef and another, Meticonaxius mindoro Poore, from the North West Shelf. Diagnosis. Rostrum flat or obsolete. Eye usually pigmented. Article 1 of antennule peduncle longer than article 2. Propodus of pereopods 3 and 4 more or less broadened. Pereopod 4 coxa flattened. Pleuron of abdominal somite 1 obsolete. Abdominal somite 2 twice as long as 1. Male pleopod 2 endopod foliaceous with small appendix masculina attached mesially. Pleopods 2–5 exopods laterally lobed. Uropodal endopod ovate. Carapace, pereopods 2–4 and abdominal somites with some setal-rows.

Michelea Kensley & Heard, 1991 Michelea is a very distinctive genus, more callianassid-like than other micheleids which tend to look more like axiids. Species of Michelea are easily distinguished from callianassids with which it might be first confused by the similarity of pleopod 2 to those following, by the pleopodal lamellae on both rami, and the elongate first article of the antennule. The flat filaments around the margins of the pleopods are unique. There are ten species described, mostly from reef environments but none is very common. Both southern species are known from a single individual each.

176 Thalassinidea – ghost shrimps and sponge shrimps

Diagnosis. Rostrum obsolete or obscurely dentate, not carinate. Anterolateral cephalothorax with 1 longitudinal, 1 marginal and 1 vertical setal-rows. Article 1 of antennule peduncle elongate. Pereopods 3 and 4 with lateral spiniform setae on propodus and dactylus. Abdominal somites 1–5 each with 1 lateral setal-row, sixth with 3 pairs of setal-rows. Pleopods 2–5 with marginal lamellae on at least endopod and usually both rami. Telson broader than long, distally rounded.

a d b c

Fig. 47. Micheleidae. a, Michelea microphylla. b, c, Michelea hortus (pleopods 3, 4). d, Michelea microphylla (pleopod 2). (from Poore, 1997)

Key to southern Australian species of Michelea 1. Gills minute; pleopods 2–5 rami each with about 20 lateral lamellae . . . . Michelea microphylla — Gills well developed; only pleopods 3–5 endopods with 4–5 lateral lamellae . . . Michelea hortus Michelea hortus Poore, 1997 (Fig. 47b, c). Pleopods 3–5 with 4–5 marginal lamellae on endopods only. With epipods and branchia. cl. 3.6 mm. WA (Garden I.); shelf depths. Michelea microphylla Poore, 1997 (Fig. 47a, d). Pleopods 2–5 with marginal lamellae on both rami. With few rudimentary epipods and branchia. Juvenile cl. 4.0 mm. Vic. (Western Port); sand-gravel, 19 m depth.

References O’Hara, T. 2002. Endemism, rarity and vulnerability of marine species along a temperate coastline. Invertebrate Systematics 16: 671–684. Poore, G.C.B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54: 79–120. Poore, G.C.B. 1997. A review of the thalassinidean families Callianideidae Kossmann, Micheleidae Sakai, and Thomassiniidae de Saint Laurent (Crustacea: Decapoda) with descriptions of fifteen new species. Zoosystema 19: 345–420.

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Strahlaxiidae Poore, 1994 Strahlaxiids are distinguished from axiids by several characters of which the triangular rather than ovate uropodal endopod is the most obvious. The family has three genera, two occurring in Australia (Poore & Griffin, 1979; Poore, 1994). Neaxius is represented in Australia by two tropical species and Strahlaxius by two. Diagnosis. Rostrum usually spinose, apically bifid. Eye usually pigmented. Article 1 of antenna 1 peduncle as long as article 2. Propodus of pereopods 3 and 4 broadened. Pereopod 4 coxa more or less flattened. Pleuron of abdominal somite 1 not produced. Abdominal somite 2 twice as long as 1. Pereopods 2–4 and abdominal somites with some setal-rows. Male pleopod 2 endopod foliaceous with small appendix masculina attached mesially. Exopods of pleopods 2–5 laterally lobed. Uropodal endopod triangular.

Strahlaxius Sakai & de Saint Laurent, 1989 The genus is confined to the two Australian species. Diagnosis. Antenna 2 acicle simple. Pleopods 1 of male with 3 articles. Pleurobranchs 5–7 absent. Telson as wide or wider than long, dorsally smooth. Uropodal exopod without transverse suture.

Key to Australian species of Strahlaxius 1. Rostrum strongly spinose, spines of medial lobe distinct from those of lateral lobes; uropodal exopod U-shaped, widest at midpoint; cephalothorax up to 23 mm long ...... Strahlaxius plectrorhynchus — Rostrum obsoletely spinose, spines of medial lobe not distinct from those of lateral lobes; uropodal exopod subtriangular, widest distally; cephalothorax up to 13 mm long ...... Strahlaxius waroona Strahlaxius plectrorhynchus (Strahl, 1861) (Fig. 48a, b, c). Rostrum spinose, spines of medial lobe distinct from one at end of lateral lobe. Uropodal exopod U-shaped, widest at midpoint. cl. 23 mm. Indonesia, Qld, SA, WA; reef and coral, intertidal to 13 m depth. This species is more commonly encountered intertidally in southern Australia than any axiid shrimp but all are cryptic. Strahlaxius waroona (Poore & Griffin, 1979) (Fig. 48d, e, f, Pl. 12e). Rostrum obsoletely spinose, spines of medial lobe not distinct from one at end of lateral lobe. Uropodal exopod subtriangular, widest distally. cl. 13 mm. Qld, Vic., Tas., SA, WA; under rocks, intertidal.

References Poore, G.C.B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54: 79–120. Poore, G.C.B., & Griffin, D.J.G. 1979. The Thalassinidea (Crustacea: Decapoda) of Australia. Records of the Australian Museum 32: 217–321.

178 Thalassinidea – ghost shrimps and sponge shrimps

d b

Fig. 48. Strahlaxiidae. a, b, c, Strahlaxius plectrorhynchus (habitus, dorsal carapace and rostrum, telson and left uropod). d, e, f, Strahlaxius waroona (dorsal carapace and rostrum, lateral carapace, telson and left uropod).

179 Marine Decapod Crustacea of Southern Australia

Superfamily Callianassoidea Dana, 1852 Tropical Thomassiniidae de Saint Laurent, 1979 are the only one of five callianassoid families not so far reported from Australia. All callianassoids possess at least a partially developed linea thalassinica, a hinge between the dorsal carapace and branchiostegite, supposedly to facilitate respiration in a burrow.

Callianassidae Dana, 1852 Callianassids are elongate, very soft-bodied shrimps, white or pale pink, with an abdomen longer than the carapace. They are popularly called ghost shrimps and there are more than 200 species, mostly in the tropics where they build permanent burrows in soft sediments of the shelf or reef lagoons. Burrow structure and feeding behaviour is well known (de Vaugelas & de Saint Laurent, 1984; Witbaard & Duineveld, 1989; Nickell & Atkinson, 1995; Berkenbusch & Rowden, 1998; Dworschak, 1998; Tamaki & Ueno, 1998; Bird et al., 1999; Bird & Poore, 1999; Stapleton et al., 2002). Some are filter feeders but others pull seagrass down into the burrows to harvest as food (Stamhuis et al., 1998). There have been several studies of their larval development (eg. Nates et al., 1997). The largest callianassids are up to 15 cm long and are sometimes used as food. More commonly, ghost shrimps are captured for fish bait using a ‘yabby pump’ or ‘bait pump’. There seems little consensus on the generic arrangement but at least 20 genera are known (Manning & Felder, 1991; Sakai, 1999b; Tudge et al., 2000). Poore & Griffin (1979) listed 15 species of callianassids from Australia and covered all but one of the southern species now known. Sakai (1988) and Ngoc-Ho (1994) described more from the tropics and Sakai (1999a, 2000) others from Tasmania and South Australia, together bringing the total to 26 from Australia.

Diagnosis. Rostrum obsolete or a spike. Carapace posterior margin evenly curved, rarely with cardiac prominence; linea thalassinica present, lateral to antennae. Eyestalks flattened, rarely cylindrical. Maxilla 2 scaphognathite without long seta on posterior lobe. Pereopods 1 unequal or equal; merus straight or toothed lower margin. Pereopod 2 chelate. Pereopod 3 propodus without distal spiniform seta on lower margin, with proximal heel on lower margin. Pereopod 4 coxa flattened, mobile. Thoracic sternite 7 narrow. Abdominal somite 1 without anterolateral lobes, weakly chitinised. Abdominal somite 6 with setal-rows. Pleopod 2 different from pleopods 3–5, sexually modified. Pleopods 3–5 with broad interacting rami. Uropodal exopod with thickened anterodorsal setose margin.

Key to southern Australian species of Callianassidae 1. Antennule peduncle heavier and longer than antenna peduncle, extremely setose on lower margin; maxilliped 3 merus distally expanded ...... Trypaea australiensis — Antennule and antenna peduncles of similar lengths, not extremely setose; maxilliped 3 merus not expanded distally ...... 2 2. Maxilliped 3 with long exopod; merus of large cheliped with a strong unarmed lobe along entire lower edge, almost as broad as carpus ...... Paraglypturus tooradin — Maxilliped 3 without exopod; merus of large cheliped with hook or complex dentition on lower margin, narrower than carpus ...... 3 3. Rostrum with medial and 2 lateral spines ...... Corallianassa collaroy — Rostrum without spines ...... 4 4. Chelipeds subequal; telson much shorter than uropods ...... Eucalliax aequimanus — Chelipeds unequal; telson as long or little shorter than uropods ...... 5

180 Thalassinidea – ghost shrimps and sponge shrimps

5. Merus of large cheliped with dentate lower margin ...... Neocallichirus angelikae — Merus of large cheliped with strong hook or dentate lobe on lower margin . . . Biffarius … 6 6. Propodus of maxilliped 3 almost as long as broad, 3 times as wide as dactylus ...... Biffarius limosus — Propodus of maxilliped 3 much longer than broad, about twice as wide as dactylus ...... 7 7. Telson tapering slightly distally, as long as uropodal endopod; male cheliped merus with strongly dentate ridge, carpus shorter than long; adults to 80 mm ...... Biffarius ceramicus — Telson subquadrate or tapering, shorter than uropodal endopod; male cheliped merus with denticulate hook, carpus longer than wide; adults less than 43 mm ...... 8 8. Larger cheliped with notch in gape; dactylus toothed ...... Biffarius arenosus — Larger cheliped without notch in gape, dactylus simple ...... Biffarius poorei

Biffarius Manning & Felder, 1991 Four of the following species were described as species of the larger genus Callianassa, and one as Neocallichirus, but all were included in Biffarius in Tudge et al.’s (2000) phylogeny. This may not be their final generic placement. Diagnosis. Carapace with dorsal oval, rostrum and anterolateral projections obsolete. Eyes flattened. Antennules and antennae peduncles similar in dimensions. Maxilliped 3 operculate, ischium not produced, dactylus linear. Chelipeds equal or unequal; major cheliped with meral hook; minor cheliped with denticulate ischium. Pleopods 3–5 with stubby appendix masculina. Biffarius arenosus (Poore, 1975) (Fig. 49b, d, Pl. 12a). Larger cheliped merus with hook on lower margin; propodus excavate in gape in male only; dactylus with 2 strong teeth in male, simple in female. Telson as wide as long, shorter than uropods. 43 mm. Southern Qld, NSW, Vic., Tas.; intertidal to shallow subtidal sandflats, often estuarine. This is one of the few Australian species studied. The paper in which the species was described detailed its density in Port Phillip Bay (Poore, 1975) and comparable data is available for Western Port (Coleman & Poore, 1980). The species is an active burrower important in sedimentary nutrient dynamics (Bird et al., 1999; Bird & Poore, 1999). It makes a complex burrow and feeds on the sediment it excavates (Pl. 12d; Stapleton et al., 2002). Biffarius ceramicus (Fulton & Grant, 1906) (Fig. 49d, e). Larger cheliped merus with serrated tooth and ridge on lower margin; carpus in male shorter than broad; propodus excavate and with minor tooth in gape; fixed finger straight; dactylus irregularly dentate, more so in male. Telson longer than wide, as long as uropods. 80 mm. Tas., Vic., SA, southern WA.; intertidal and shallow subtidal mudflats and sandy beaches. Biffarius limosus (Poore, 1975) (Fig. 49f, g, Pl. 12b). Chelipeds unequal and sexually dimorphic; larger cheliped merus with proximal hook on lower margin; propodus smooth; fixed finger simple; dactylus irregularly toothed in male. Telson tapering to convex posterior margin, as long as uropods. Cornea fully formed. 30 mm. NSW, Vic., Tas.; subtidal muddy sediments to 100 m depth. This species is extremely common on muddy sediments in Port Phillip Bay (Poore, 1975). Biffarius poorei (Sakai, 1999) (Fig. 50a). Larger cheliped merus with triangular serrated tooth on lower margin; carpus in male longer than broad; propodus scarcely excavate and without minor tooth in gape; fixed finger straight; dactylus smooth, more so in male. Telson longer than wide, shorter than uropods. 22 mm. Tas., Vic.; shallow subtidal mudflats beaches. This smaller subtidal species was confused with the much larger B. ceramicus by Poore (1975) and later publications.

181 Marine Decapod Crustacea of Southern Australia

e d

f g

Fig. 49. Callianassidae. a, Trypaea australiensis. Large cheliped, uropod and left uropod. b, c, Biffarius arenosus. d, e, Biffarius ceramicus. f, g, Biffarius limosus.

182 Thalassinidea – ghost shrimps and sponge shrimps

a b

d c

g h

Fig. 50. Callianassidae. a, Biffarius poorei (large cheliped). b, Corallianassa collaroy (carapace and eyes). Large cheliped, telson and left uropod: c, d, Corallianassa collaroy (from Poore & Griffin, 1979). e, f, Eucalliax aequimanus. g, h, Neocallichirus angelikae (from Sakai, 2000). i, j, Paraglypterus tooradin. k, l, Trypaea australiensis (small cheliped, telson and left uropod).

183 Marine Decapod Crustacea of Southern Australia

Corallianassa Manning, 1987 Sakai (1999b) treated most species of Corallianassa as members of a much larger Neocallichirus. Diagnosis. Carapace with dorsal oval, front trispinose. Eyes globular. Antennules and antennae peduncles similar in dimensions. Maxilliped 3 operculate, ischium not produced, propodus broad, dactylus linear. Chelipeds unequal. Pleopods 3–5 with stubby appendix masculina. Corallianassa collaroy (Poore & Griffin, 1979) (Fig. 50b, c, d). Rostrum trispinose. Chelipeds unequal and similar in both sexes; larger cheliped merus spinose on lower margin; fixed finger and dactylus stout, unarmed. Telson with concave posterior margin, much shorter than uropods. 123 mm. NSW; intertidal.

Eucalliax Manning & Felder, 1991 Sakai (1999b) did not differentiate Eucalliax from Calliax de Saint Laurent, 1973. Diagnosis. Carapace without dorsal oval, rostrum obsolete. Maxilliped 3 dactylus ovate. Chelipeds equal. Eucalliax aequimanus (Baker, 1907) (Fig. 50e, f). Rostrum short. Chelipeds equal and similar in both sexes; merus unarmed, carinate on upper margin; propodus carinate on upper and lower margins; fixed finger with proximal tooth; dactylus usually smooth. Telson with concave posterior margin, much shorter than uropods. 84 mm. Southern Qld to WA, not Tas.; intertidal mudflats–9 m depth, often estuarine.

Neocallichirus Sakai, 1988 Diagnosis. Carapace with dorsal oval, rostrum obsolete. Eyes minute. Antennules and antennae peduncles similar in dimensions. Maxilliped 3 operculate, ischium not produced, propodus broad, dactylus linear. Chelipeds unequal. Pleopods 3–5 with stubby appendix masculina. Neocallichirus angelikae Sakai, 2000 (Fig. 50g). Chelipeds unequal (only female known); larger cheliped merus with dentate lower margin; propodus with rugose surfaces; fixed finger toothed near base; dactylus notched in female. Telson tapering to posterior margin with small median spine, shorter than uropods. Cornea reduced. 47 mm. Western SA; intertidal.

Paraglypturus Türkay & Sakai, 1995 Diagnosis. Carapace without dorsal oval, rostrum distinct or obsolete. Maxilliped 3 with exopod, dactylus ovate. Chelipeds unequal, major without meral hook. Paraglypturus tooradin (Poore & Griffin, 1979) (Fig. 50h, i). Rostrum short. Maxilliped 3 with long exopod. Chelipeds unequal (male not known); merus strongly lobed on lower margin; propodus tapering; fixed finger and dactylus simple. Telson much wider than long, much shorter than uropods. 24 mm. Western Port, Vic.; subtidal to 2.5 m depth. The species is very rare but is distinguished from all other Australian callianassids by the possession of an exopod on maxilliped 3. The species (as Eucalliax tooradin) was listed as ‘vulnerable’ under IUCN criteria by O’Hara (2002).

Trypaea Dana, 1852 The Australian species is the only representative of the genus, sunk into Callianassa by Sakai (1999b). Diagnosis. Antennule peduncle much heavier and longer than antenna peduncle, article 3 about twice as long as article 2. Maxilliped 3 ischium-merus operculiform, merus projecting beyond articulation with carpus; propodus and dactylus digitiform. Male pleopod 2 absent.

184 Thalassinidea – ghost shrimps and sponge shrimps

Trypaea australiensis Dana, 1852 Bass yabby, ghost nipper (Figs 49a, 50i, k, Pl. 12c). 63 mm. Qld, NSW, eastern Vic.; intertidal sand and mudflats, often estuarine. The species is most easily recognised by the dense setation of the antennules, antennae and maxilliped 3. It has until recently been referred to the genus Callianassa along with most other species of the family. This species is the most common of the Australian callianassids and is popularly called ‘yabbies’ or ‘Bass yabbies’ to differentiate them from freshwater yabbies (Parastacidae), or ‘ghost nippers’.They are frequently taken with a yabby pump by fishermen for bait. In suitable environments densities may reach hundreds of individuals per square metre (Hailstone & Stephenson, 1961; Coleman & Poore, 1980). Hailstone & Stephenson (1961) reported on growth, morphometrics and reproduction in this species in Queensland. Dakin & Colefax (1940) described the planktonic larval stages. The shrimp builds a Y-shaped burrow and feeds on the sediment it excavates and resuspends (Stapleton et al., 2002).

References Baker, W.H. 1907. Notes on South Australian decapod Crustacea. Part V. Transactions of the Royal Society of South Australia 31: 173–190, pls 23–25. Berkenbusch, K., & Rowden, A.A. 1998. Population dynamics of the burrowing ghost shrimp Callianassa filholi on an intertidal sandflat in New Zealand (Decapoda: Thalassinidea). Ophelia 49: 55–69. Bird, F.L., Ford, P.W., & Hancock, G.J. 1999. Effect of burrowing macrobenthos on the flux of dissolved substances across the water-sediment interface. Marine and Freshwater Research 50: 523–532. Bird, F.L., & Poore, G.C.B. 1999. Functional burrow morphology of Biffarius arenosus (Decapoda: Callianassidae) from southern Australia. Marine Biology 134: 77–87. Coleman, N., & Poore, G.C.B. 1980. The distribution of Callianassa species (Crustacea, Decapoda) in Western Port, Victoria. Proceedings of the Royal Society of Victoria 91: 73–78. Dakin, W.J., & Colefax, A.N. 1940. The plankton of the Australian coastal waters off New South Wales Part 1 with special reference to the seasonal distribution, the phyto-plankton, and the planktonic Crustacea, and in particular, the Copepoda and crustacean larvae, together with an account of the more frequent members of the groups Mysidacea, Euphausiacea, Amphipoda, Mollusca, Tunicata, Chaetognathia, and some reference to the fish eggs and fish larvae. Publications of the University of Sydney, Department of Zoology, Monograph 1: 1–215. Dworschak, P.C. 1998. The role of tegumental glands in burrow construction by two Mediterranean callianassid shrimps. Senckenbergiana Maritima 28: 143–149. Hailstone, T.S., & Stephenson, W. 1961. The biology of Callianassa (Trypaea) australiensis Dana 1852 (Crustacea, Thalassinidea). University of Queensland, Department of Zoology, Papers 1: 259–282, pls 1–3. Manning, R.B., & Felder, D.L. 1991. Revision of the American Callianassidae (Crustacea: Decapoda: Thalassinidea). Proceedings of the Biological Society of Washington 104: 764–792. Nates, S.F., Felder, D.L., & Lemaitre, R. 1997. Comparative larval development in two species of the burrowing ghost shrimp genus Lepidophthalmus (Decapoda: Callianassidae). Journal of Crustacean Biology 17: 497–519. Ngoc-Ho, N. 1994. Some Callianassidae and Upogebidae from Australia with description of four new species (Crustacea: Decapoda: Thalassinidea). Memoirs of the Museum of Victoria 54: 51–78. Nickell, L.A., & Atkinson, R.J.A. 1995. Functional morphology of burrows and trophic modes of three thalassinidean shrimp species, and a new approach to the classification of thalassinidean burrow morphology. Marine Ecology Progress Series 128: 181–197. O’Hara, T. 2002. Endemism, rarity and vulnerability of marine species along a temperate coastline. Invertebrate Systematics 16: 671–684. Poore, G.C.B. 1975. Systematics and distribution of Callianassa (Crustacea, Decapoda, Macrura) from Port Phillip Bay, Australia, with descriptions of two new species. Pacific Science 29: 197–209.

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Poore, G.C.B., & Griffin, D.J.G. 1979. The Thalassinidea (Crustacea: Decapoda) of Australia. Records of the Australian Museum 32: 217–321. Sakai, K. 1988. A new genus and five new species of Callianassidae (Crustacea: Decapoda: Thalassinidea) from northern Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 5: 51–69. Sakai, K. 1999a. A new species, Callianassa poorei, sp. nov. (Decapoda: Crustacea: Callianassidae) from Tasmania. Journal of the Marine Biological Association of the United Kingdom 79: 373–374. Sakai, K. 1999b. Synopsis of the family Callianassidae, with keys to subfamilies, genera and species, and the description of new taxa (Crustacea: Decapoda: Thalassinidea). Zoologische Verhandelingen, Leiden 326: 1–152. Sakai, K. 2000. A new species of Neocallichirus, N. angelikae from South Australia (Decapoda: Callianassidae). Mitteilungen aus dem Hamburgischen Zoologischen Institut und Museum 97: 91–98. Stamhuis, E.J., Dauwe, B., & Videler, J.J. 1998. How to bite the dust: morphology, motion pattern and function of the feeding appendages of the deposit-feeding thalassinid shrimp Callianassa subterranea. Marine Biology 132: 43–58. Stapleton, K.L., Long, M., & Bird, F.L. 2002. Comparative feeding ecology of two spatially coexisting species of ghost shrimp; Biffarius arenosus and Trypaea australiensis (Decapoda: Callianassidae). Ophelia 55: 141–150. Tamaki, A., & Ueno, H. 1998. Burrow morphology of two callianassid shrimps, Callianassa japonica Ortmann, 1891 and Callianassa sp. (=Callianassa japonica: De Man, 1928)(Decapoda: Thalassinidea). Crustacean Research 27: 28–39. Tudge, C.C., Poore, G.C.B., & Lemaitre, R. 2000. Preliminary phylogenetic analysis of generic relationships within the Callianassidae and Ctenochelidae (Decapoda: Thalassinidea: Callianassoidea). Journal of Crustacean Biology 20 (Special Issue 2): 129–149. de Vaugelas, J., & de Saint Laurent, M. 1984. Premières données sur l’écologie de Callichirus laurae de Saint Laurent sp. nov. (Crustacé Décapode Callianassidae): son action bioturbatrice sur les forma- tions sédimentaires du golfe d’Aqaba (Mer Rouge). Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 6: 147–152. Witbaard, R., & Duineveld, G.C.A. 1989. Some aspects of the biology and ecology of the burrowing shrimp Callianassa subterranea (Montagu) (Thalassinidea) from the southern North Sea. Sarsia 74: 209–219.

Ctenochelidae Manning and Felder, 1991 There are five genera, mostly tropical (Poore, 1994); only one species occurs in southern Australia and that is rarely found probably because it is a deep subtidal burrower. Diagnosis. Rostrum obsolete or a spike. Carapace posterior margin evenly curved, with cardiac prominence; linea thalassinica present, lateral to antennae. Eyestalks flattened. Maxilla 2 scaphognathite without long seta on posterior lobe. Pereopods 1 unequal; merus with straight or toothed lower margin. Pereopod 2 chelate. Pereopod 3 propodus without distal spiniform seta on lower margin. Pereopod 3 propodus linear or weakly ovate. Pereopod 4 coxa flattened, mobile. Thoracic sternite 7 narrow. Abdominal somite 1 without anterolateral lobes, weakly chitinised. Abdominal somite 6 sometimes with setal-rows. Pleopod 2 similar to pleopods 3–5, rami lanceolate. Uropodal exopod simply ovate.

Ctenocheles Kishinouye, 1926 Ctenocheles is a widespread genus of four species, some from deep water (de Saint Laurent & Le Loeuff, 1979; Sakai, 1999). There is only one species in Australia but it has been taken infrequently. Diagnosis. Large cheliped with fingers elongate and pectinate.

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Ctenocheles collini Ward, 1945 (Fig. 51). Moreton Bay, Qld, and eastern Bass Strait. The species is immediately recognisable by its elongate pectinate fingers. Specimens from Moreton Bay are large (120 mm overall) but those collected so far from offshore NSW and Bass Strait, possibly juveniles, are much smaller (15 mm).

b a

Fig. 51. Ctenochelidae. Ctenocheles collini. a, carapace. b, c, large and small chelipeds.

References Poore, G.C.B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54: 79–120. de Saint Laurent, M., & Le Loeuff, P. 1979. Campagnes de la Calypso au large des côtes Atlantiques Africaines (1956 et 1959) (suite) 22. Crustacés Décapodes Thalassinidea. I. Upogebiidae et Callianassidae. In: Forest, J. (ed.), Résultats Scientifiques des Campagnes de la Calypso. Fasc. 11 (22). Annales de l’Institut Océanographique, Monaco et Paris 55 suppl.: 29–101. Sakai, K. 1999. Redescription of Ctenocheles balssi Kishinouye, 1926, with comments on its systematic position and establishment of a new subfamily Gourretiinae (Decapoda, Callianassidae). Crustaceana 72: 85–97.

Laomediidae Borradaile, 1903 The diagnosis is based on Kensley & Heard (1990), Poore (1994) and Ngoc-Ho (1997) who included the Axianassidae in the Laomediidae. The last author treated the two families as subfamilies of Laomediidae on the basis of a cladistic analysis of adult and larval characters. She tabulated the characters of the five genera. There are about 15 species worldwide but others are known only from larvae (Wear & Yaldwyn, 1966; Goy & Provenzano, 1979). Only one genus is known from Australia. Diagnosis. Compact shrimps, cephalothorax with posterior margin separate from thickened margins of posterolateral extensions of branchial region, with rostrum at least as long as eyestalks. Linea thalassinica present, cervical groove distinct. Antenna with acicle. Maxilla 2 scaphognathite with several long setae extending into branchial chamber. Maxilliped 3 pediform, epipod with serrate margin. Pleopods 1 of male absent, of female uniramous; pleopods 2–5 with broad rami. Epipods 3–7 present.

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Laomedia De Haan, 1841 Ngoc-Ho (1997) provided a key to the only four species, one each from Japan, Vietnam, southern Australia, and northern Queensland (Yaldwyn & Wear, 1972; Ngoc-Ho, 1997; Ngoc-Ho & Yaldwyn, 1997). Diagnosis. Rostrum armed. Antennule peduncle with short article 3. Antenna peduncle with reduced acicle and short article 4. Maxilliped 3 with exopod; pereopods 2–5 with reduced exopods. Pereopods 1 chelate, dissimilar. Pereopods 2 and 5 subchelate. Uropodal rami both with sutures. Laomedia healyi Yaldwyn & Wear, 1970 (Fig. 52). 28 mm. Qld, NSW, Vic.; intertidal, often in mangroves. This species may be confused with upogebiids but the combination of rostrum, chelipeds and uropods is unique. When alive it is brick-red all over. The species was redescribed and compared with others by Ngoc-Ho (1997).

b c

Fig. 52. Laomediidae. a, b, c, Laomedia healyi (habitus, dorsal carapace, telson and left uropod).

References Goy, J.W., & Provenzano, A.J. 1979. Juvenile morphology of the rare burrowing mud shrimp Naushonia crangonoides Kingsley, with a review of the genus Naushonia (Decapoda: Thalassinidea: Laomediidae). Proceedings of the Biological Society of Washington 92: 339–359. Kensley, B., & Heard, R. 1990. The genus Axianassa (Crustacea: Decapoda: Thalassinidea) in the Americas. Proceedings of the Biological Society of Washington 103: 558–572. Ngoc-Ho, N. 1997. The genus Laomedia De haan, 1841 with description of a new species from Vietnam (Crustacea, Thalassinidae, Laomediidae). Zoosystema 19: 729–747.

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Ngoc-Ho, N., & Yaldwyn, J.C. 1997. A new species of Laomedia (Crustacea, Thalassinidea, Laomediidae) from Australia with notes on its ecology. Zoosystema 19: 337–343. Poore, G.C.B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54: 79–120. Wear, R.G., & Yaldwyn, J.C. 1966. Studies on thalassinid Crustacea (Decapoda, Macrura Reptantia) with a description of a new Jaxea from New Zealand and an account of its larval development. Zoology Publication from the Victoria University of Wellington 41: 1–27. Yaldwyn, J.C., & Wear, R.G. 1972. The eastern Australia, burrowing mud-shrimp Laomedia healyi (Crustacea, Macrura Reptantia, Laomediidae) with notes on the larvae of the genus Laomedia. Australian Zoologist 17: 126–141, pls 6, 7.

Upogebiidae Borradaile, 1903 Upogebiids, sometimes called mud shrimps or sponge shrimps, are rarely encountered but are found under rocks, inside sponges or burrowing in sediments. Most are instantly recognised by their furry flat rostrum. Nothing is known of their biology in Australia but the physiology and ecology of burrowing species has been widely studied elsewhere. For example, the eastern Pacific species, Upogebia pugettensis, forms male-female pairs living in permanent U-shaped burrows up to one metre deep in soft muds of tidal flats. There, they pump water through the burrow to feed on plankton and suspended detritus (Williams, 1986; Atkinson & Taylor, 1988; Nickell & Atkinson, 1995; Astall et al., 1997). Most species (about 100 worldwide) are members of the genus Upogebia and are very similar in general appearance. Until recently, most species were referred to this genus. There are now seven other genera and a key to all except one was provided by Poore (1994); the exception was added by Ngoc-Ho (2001). Twenty-three upogebiid species are known from Australia (Poore & Griffin, 1979; Sakai, 1982, 1993; Ngoc-Ho, 1994).

Diagnosis. Eyestalks cylindrical, cornea large and terminal. Cephalothorax with cervical groove well-marked; linea thalassinica obvious on anterior region but variable posteriorly; gastric region continuous with flat rostrum, usually extensive and ornamented anteriorly with spines or tubercles and thick pile of plumose setae. Epistome visible laterally at base of antennae. Abdomen reflexed, depressed, pleura weak. Epipods 4–8 absent. Pereopods 1 (chelipeds) equal. Pereopods 2 not chelate. Pereopods 3 and 4 with dactyli more or less spatulate. Pereopod 5 subchelate. Pleopod 1 absent in male, simple in female. Pleopods 2–5 foliaceous, without appendix interna, without appendix masculina in male. Uropodal rami never with suture.

Key to southern Australian species of Upogebiidae 1. Medial and lateral lobes of rostrum ventrally spinose, dorsally smooth ...... Austinogebia spinifrons — Lobes of rostrum not ventrally spinose, with marginal dorsal spines ...... 2 2. Lateral and medial lobes of rostrum separated by a broad U; telson apically broadly rounded ...... Acutigebia simsoni — Lateral and medial lobes of rostrum separated by a narrow slit; telson apically truncate ...... Upogebia ...3 3. Pereopod 1 subchelate, fixed finger shorter than dactylus ...... 4 — Pereopod 1 chelate, fixed finger and dactylus equal ...... 5 4. Lateral lobes of rostrum obsolete or at most one-eighth as long as medial lobe; telson with 2 transverse carinae ...... Upogebia dromana

189 Marine Decapod Crustacea of Southern Australia

— Lateral lobes of rostrum about half as long as median lobe and separated from it by a deep U; telson smooth or with 1 transverse carina ...... Upogebia neglecta 5. Eyestalks exceeding rostrum; medial and lateral lobes of rostrum separated by a broad concavity ...... Upogebia tractabilis — Eyestalks not exceeding rostrum; rostrum trilobed or with strong tubercles laterally . . . . . 6 6. Medial lobe of rostrum with about 12 short equal closely-spaced spines; distal medial edge of propodus of pereopod 1 with 2 strong spines; posterior margin of telson concave ...... Upogebia bowerbankii — Medial lobe of rostrum unevenly spinose; distal mesial edge of pereopod 1 with minute denticles; posterior margin of telson convex ...... 7 7. Medial lobe of rostrum with 2 or 3 spines each side; lateral lobes reaching less than halfway towards most posterior spine of medial lobe; merus of pereopod 1 with denticles on lower edge ...... Upogebia darwinii — Medial lobe of rostrum with about 6 tubercles each side; lateral lobes reaching to most posterior tubercle of medial lobe; merus of pereopod 1 smooth on lower edge ...... Upogebia australiensis

Acutigebia Sakai, 1982 The trilobed rostrum distinguishes members of this small genus from other upogebiids in Australia (Sakai, 1993). Diagnosis. Rostrum without inferior spines, medial lobe separated from prominent lateral lobes by a broad U. Maxilliped 3 ischium with row of spines (crista dentata) on mesial surface. Acutigebia simsoni (Thomson, 1893) (Figs 53a, b, c, 54d, 55a, Pl. 12f). Rostrum trilobed, medial lobe with up to 10 tubercles each side; lateral lobes half as long as medial lobe, separated by broad U. Pereopod 1 merus spinose on lower edge; distal margin of carpus with strong lower spine, shorter upper spines; propodus with denticles on lower margin and small upper spine distally; fixed finger one-fifth length of dactylus. Telson wider than long, smooth, margin concave. cl. 40 mm. NT, Qld, NSW, Vic., Tas., SA; intertidal to high subtidal.

Austinogebia Ngoc-Ho, 2001 Austinogebia is one of two genera with spines under the rostrum (Ngoc-Ho, 2001). The other, Gebiacantha Ngoc-Ho, 1989 is represented in tropical Australia (Ngoc-Ho, 1994). Diagnosis. Rostrum with 2–4 ventral spines on median lobe; setose, without dorsal median furrow; lateral lobes strongly projecting and separated from medial lobe, setose, each with 1–3 spines underneath. Uropodal endopod with prominent knob on lateral margin, not longer than telson; tail fan not operculiform. Austinogebia spinifrons (Haswell, 1881) (Figs 53a, b, c, 54e, 55b). Rostrum strongly trilobed, medial lobe with 4 spines ventrally; lateral lobes half as long as medial lobe, with 1 ventral spine, separated by narrow U. Pereopod 1 merus spinose on lower edge; distal margin of carpus with strong lower spine, 6–8 mesial and dorsal spines; propodus with up to 10 strong spines on upper margin, spinose distally; fixed finger less than half length of dactylus. Telson wider than long, with groove in midline. cl. 25 mm. Qld, NSW; 12–25 m depth. Austinogebia spinifrons is the only species in southern Australia with spines on the ventral surface of the rostral lobes. Sakai (1984) recorded this species from Japan; his figures suggest a slightly different species.

190 Thalassinidea – ghost shrimps and sponge shrimps

c b

d e

g f

Fig. 53. Upogebiidae. a, b, c, Austinogebia spinifrons (habitus, dorsal and lateral carapace). Dorsal carapace: d, Acutigebia simsoni. e, Upogebia australiensis. f, Upogebia bowerbankii. g, Upogebia darwinii. (e from Poore & Griffin, 1979)

191 Marine Decapod Crustacea of Southern Australia

b a

f e

Fig. 54. Upogebiidae. Dorsal carapace: a, Upogebia dromana. b, Upogebia neglecta. c, Upogebia tractabilis. Pereopod 1: d, Acutigebia simsoni. e, Austinogebia spinifrons. f, Upogebia darwinii.

Upogebia Leach, 1814 Upogebia is by far the largest genus in the family from with others have been hived off. It is distributed worldwide, except in polar waters, with numerous species in the tropics. Diagnosis. Rostrum usually without ventral spines; with dorsal median furrow; lateral lobes only weakly separated from medial lobe. Uropodal exopod broad, not longer than telson; tail fan not operculiform. Maxilliped 3 with proximal hook on ischium. Upogebia australiensis de Man, 1927 (Figs 53e, 55c). Medial lobe of rostrum triangular, with about 6 tubercles each side; lateral lobes reaching to most posterior tubercle of medial lobe and separated by broad V. Pereopod 1 merus smooth on lower edge; distal margin of carpus with dorsal denticles, a mesial and ventral spine; fixed finger of propodus almost as long as dactylus. Telson as wide as long, with strong transverse ridge and pair of longitudinal carinae, spinulose. cl. 15 mm. Qld, NSW, WA; often from sponges, intertidal to 60 m depth. This species is not a synonym of U. bowerbankii (Sakai, 1982; Davie, 2002). Upogebia bowerbankii (Miers, 1884) (Figs 53f, 55d). Medial lobe of rostrum broadly rounded, with about 6 spines each side; lateral lobes reaching to most posterior spine of medial lobe and separated by deep U. Pereopod 1 merus smooth on lower edge; carpus with strong mesiodistal spine and 3 or 4 minute mesial spines; propodus with minute spines on upper edge and 4 strong spines at base of dactylus; fixed finger and dactylus subequal. Telson as wide

192 Thalassinidea – ghost shrimps and sponge shrimps

a b

c d

e f

h g

Fig. 55. Upogebiidae. Telson and left uropod (setae not shown on all): a, Acutigebia simsoni. b, Austinogebia spinifrons. c, Upogebia australiensis. d, Upogebia bowerbankii. e, Upogebia darwinii. f, Upogebia dromana. g, Upogebia neglecta. h, Upogebia tractabilis. (c, d, g, h from Poore & Griffin, 1979)

as long, with spinulose transverse ridge and pair of longitudinal carinae, margin concave. cl. 17 mm. Madagascar, SA, southern WA; often from sponges, 6–200 m depth. Upogebia darwinii (Miers, 1884) (Figs 53g, 54f, 55e). Medial lobe of rostrum rounded-truncate, with 2 or 3 tubercles each side; lateral lobes quarter length of medial lobe and separated by shallow V. Pereopod 1 merus with denticles on lower edge; distal margin of carpus with 4 or 5

193 Marine Decapod Crustacea of Southern Australia

short mesial denticles and short dorsal spines; propodus with scattered denticles; fixed finger of propodus as long as dactylus. Telson wider than long, with weak transverse ridge and pair of longitudinal carinae, spinulose, margin convex. cl. 15 mm. Indonesia, NT, Qld, WA; often from sponges, intertidal to 20 m depth. This was incorrectly called Upogebia hexaceras (Ortmannn) by Poore & Griffin (1979). The adult and its larval development were described by Ngoc-Ho (1977). Upogebia dromana Poore & Griffin, 1979 (Figs 54a, 55f). Rostrum not obviously trilobed, with 4–7 tubercles each side of medial lobe; lateral lobes barely distinguishable by short hiatus in row of tubercles from medial rostrum. Pereopod 1 merus spinose on lower edge; carpus with strong mesiodorsal spine and smaller mesial spine; propodus without spines; fixed finger one-fifth length of dactylus. Telson about as wide as long, with 2 transverse ridges and median longitudinal carina, margin convex. cl. 9 mm. NSW, Vic., Tas.; bays. benthic sediments, 10–20 m depth. This species is yellowish-orange with widely-spaced red chromatophores. In Port Phillip Bay quantitative samples have indicated a density of up to 16 specimens per square metre. Upogebia neglecta de Man, 1927 (Figs 54b, 55g). Rostrum trilobed, tuberculate; lateral lobes separated by narrow V from medial lobe. Pereopod 1 merus spinose on lower edge; carpus with strong curved lower spine, shorter upper spines; propodus with denticles on lower margin and spines distally; fixed finger one-fifth length of dactylus. Telson wider than long, smooth, margin convex. cl. 15 mm. NSW (known only from the holotype from Port Stephens). Upogebia tractabilis Hale, 1941 (Figs 54c, 55h, Pl. 12g). Rostrum trilobed, medial lobe much wider than long, shorter than eyestalks, with about 6 small spines each side; lateral lobes one-fifth as long as medial lobe, separated by shallow U. Pereopod 1 merus spinose on lower edge; distal margin of carpus with lower spines; propodus with 2 spines on lower margin; fixed finger and dactylus subequal. Telson wider than long, smooth, margin convex. cl. 8 mm. SA, WA; often with sponges, to 75 m depth. This species is the only one in southern Australia in which the eyestalks are longer than the rostrum.

References Astall, C.M., Taylor, A.C., & Atkinson, R.J.A. 1997. Behavioural and physiological implications of a burrow-dwelling lifestyle for two species of a upogebiid mud-shrimp (Crustacea: Thalassinidea). Estuarine, Coastal and Shelf Science 44: 155–168. Atkinson, R.J.A., & Taylor, A.C. 1988. Physiological ecology of burrowing decapods. Symposia of the Zoological Society of London 59: 201–226. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A. CSIRO Publishing: Melbourne. xii, 551 pp. Ngoc-Ho, N. 1977. The larval development of Upogebia darwini (Crustacea, Thalassinidae) reared in the laboratory, with a redescription of the adult. Journal of Zoology, London 181: 439–464. Ngoc-Ho, N. 1994. Some Callianassidae and Upogebidae from Australia with description of four new species (Crustacea: Decapoda: Thalassinidea). Memoirs of the Museum of Victoria 54: 51–78. Ngoc-Ho, N. 2001. Austinogebia, a new genus in the Upogebiidae and rediagnosis of its close relative, Gebiacantha Ngoc-Ho, 1989 (Crustacea: Decapoda: Thalassinidea). Hydrobiologia 449: 47–58. Nickell, L.A., & Atkinson, R.J.A. 1995. Functional morphology of burrows and trophic modes of three thalassinidean shrimp species, and a new approach to the classification of thalassinidean burrow morphology. Marine Ecology Progress Series 128: 181–197. Poore, G.C.B. 1994. A phylogeny of the families of Thalassinidea (Crustacea: Decapoda) with keys to the families and genera. Memoirs of the Museum of Victoria 54: 79–120.

194 Thalassinidea – ghost shrimps and sponge shrimps

Poore, G.C.B., & Griffin, D.J.G. 1979. The Thalassinidea (Crustacea: Decapoda) of Australia. Records of the Australian Museum 32: 217–321. Sakai, K. 1982. Revision of Upogebiidae (Decapoda, Thalassinidea) in the Indo-West Pacific region. Researches on Crustacea, Special Number 1: 1–106. Sakai, K. 1984. A new record of Upogebia spinifrons (Haswell, 1882) (Decapoda, Thalassinidea) from Naruto, Japan, showing possible hermaphroditism. Crustaceana 47: 209–214. Sakai, K. 1993. On a collection of Upogebiidae (Crustacea, Thalassinidea) from the Northern Territory Museum, Australia, with the description of two new species. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 10: 87–114. Williams, A.B. 1986. Mudshrimps, Upogebia, from the Eastern Pacific (Thalassinidea: Upogebiidae). San Diego Society of Natural History, Memoir 14: 1–60.

195 This page intentionally left blank 9. ACHELATA – ROCK LOBSTERS AND BUGS

The Achelata Scholtz & Richter, 1995 are entirely marine and include a variety of lobster-like crustaceans. Diagnostic features of the adults are the possession of a thickened, strong cylindrical or flattened carapace covering all of the head and thorax, and absence of a prominent rostrum. All the pereopods are simple (which distinguishes them from the freshwater yabbies which have an enlarged claw). In most forms the abdomen is strong and ends as a broad tail fan formed from the telson and uropodal rami. This enables a quick reverse escape response by swimming. Scholtz & Richter (1995), supported later by Dixon et al. (2003), believed on the basis of a phylogenetic analysis of morphology that Palinura Latreille, 1802, as traditionally considered, are polyphyletic. Palinura is the term used for these lobsters plus polychelid lobsters. These authors all found Polychelidae to be not at all related to the others but much more basal in the decapod clade, members of a group called Polychelida. The Achelata are unique in passing their early larval stages as a pelagic phyllosoma larva (Fig. 56). The phyllosoma is long-lived, almost two years in some palinurids, and is known to travel widely in oceanic currents from the shore environments where the adults live. Other features linking the Achelata are enlarged antennae, basal antennal articles fused with the epistome, pereopods without chelae, and a unique knob attaching the carapace to the last thoracic somite. Baisre (1994) derived a phylogeny of genera of the Achelata (what he called the Palinuroidea) based on larval characters. He found Palinuridae and Scyllaridae sister taxa with Synaxidae related to these two. His findings did not differ from earlier or subsquent workers whose conclusions were based on adult features (e.g. Dixon et al., 2003). The two families found in Australia are easy to separate on the basis of general shape. The Palinuridae are cylindrical, large rock lobsters or crayfish, and the Scyllaridae are flattened shovel- nosed lobsters, Balmain bugs and the like in Australia. Only one other family is recognised, Synaxidae Bate, 1881, comprising three species (Holthuis, 1991).

Diagnosis. Abdomen well-developed and muscular. Uropodal exopod without a suture. Antenna without a spine on peduncle. Posteroventral sides of carapace held in place by a projection of the last thoracic pleuron which engages the inner side of the carapace. Pereopod 1 simple. Bases of antennae, epistome and anterolateral borders of carapace fused. Egg hatching as phyllosoma larva.

Key to Australian families of Achelata 1. Carapace subcylindrical; antennae long and filamentous ...... Palinuridae ...p.197 — Carapace flattened; antennae short and flattened ...... Scyllaridae ...p.206

References Atkinson, J.M., & Boustead, N.C. 1982. The complete larval development of the scyllarid lobster Ibacus alticrenatus Bate, 1888 in New Zealand waters. Crustaceana 42: 275–287. Baisre, J.A. 1994. Phyllosoma larvae and the phylogeny of Palinuroidea (Crustacea: Decapoda): a review. Australian Journal of Marine and Freshwater Research 45: 925–944.

197 Marine Decapod Crustacea of Southern Australia

Dixon, C.J., Ahyong, S., & Schram, F.R. 2003. A new hypothesis of decapod phylogeny. Crustaceana 76: 935–975. Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125: 1–292. Scholtz, G., & Richter, S. 1995. Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113: 289–328.

Fig. 56. Achelata. Phyllosoma stage of Ibacus alticrenatus.

Palinuridae Latreille, 1802 The Palinuridae are rightly called the rock lobsters or spiny lobsters but are often called crayfish. They should not be confused with the freshwater crayfish or yabbies which belong to a very different family of crustaceans. The lobsters of North America and Europe also differ and, with their large claws, are immediately recognisable. The abdomen of palinurids is large, muscular and edible. In Australia lucrative fisheries exist for the commonest species. Fishing is usually by the use of baited pots set from small boats close to shore. While several species do occur in southern Australia and others from the tropics may be seen in fish markets, only three species are commonly seen in the region (See Box: Three common southern lobsters). Because they are so commercially important much is known about the biology and fisheries potential of rock lobsters (see extensive review in books by Cobb & Phillips, 1980; Phillips et al., 1980; Williams, 1988; Holthuis, 1991). Conferences on lobster biology and management are held every four years (See Marine and Freshwater Research 52(8) 2002).They inhabit shallow nearshore

198 Achelata – rock lobsters and bugs

rocky or reef environments, rarely to 1000 metres depth. They are cryptic, hiding in rock crevices during the day and coming out at night to feed. Analyses of stomach contents tell us that they usually feed on small invertebrates and occasionally on algae. The translucent flat phyllosoma larva lives in the plankton for up to 22 months feeding on gelatinous zooplankton (Johnston & Ritar, 2002). In the Western Australian species, Palinurus cygnus, the phyllosoma passes through nine instars lasting 9–11 months and is transported by currents well into the Indian Ocean before moulting to the puerulus stage looking much like a translucent miniature adult (Phillips et al., 1978). The puerulus swims across the continental shelf to settle in shallow inshore reef areas. George & Main (1967) recognised two lines of evolution in the Palinuridae based on the presence (in the Stridentes) or absence (in the Silentes) of a sound-producing organ. This is a stridulating apparatus formed by an expansion of the antenna which is ventrally ribbed and rubs across a ridge on the antennule. George (1997) revisited the evolution of the group, especially of Jasus and Panulirus, in the light of a more modern understanding of plate tectonics. He concluded that the genus Jasus is at least 20 million years old, speciating as Gondwanan continents and islands became separated and larval exchange was limited. Panulirus is more recent, not more than nine million years old, and diverged as more habitats became available with the collision of southern land masses with Asia. George (1966) presented colour photographs of nine species of rock lobsters in Australia and later (George, 1983) figured and keyed some other deepwater species from the North West Shelf which then were becoming increasingly interesting as a commercial resource. There are eight genera and 46 species of Palinuridae worldwide (Phillips et al., 1980; Holthuis, 1991; Chan, 1998). Another family, Synaxidae for which keys and good descriptions were provided by Holthuis (1991), was synonymised with Palinuridae by Davie (1990). Five genera and 13 named species occur in Australia and their systematics can be expected to remain relatively stable. The keys are to Australian genera and include tropical species because of their commercial importance and because some occasionally occur in south-western Australia.

Diagnosis. Carapace subcylindrical, calcified, often spinose, with two horns (supraocular spines) over eyes, sometimes fused. Pereopods simple, first short and stout. Antennules and antennae long and filamentous. Eyes stalked.

Key to southern Australian genera and subgenera of Palinuridae 1. Carapace prismatic, i.e., with strong longitudinal ridges; female pleopods 2 with stylamblys (second endopodal article) as long as in remaining pleopods ...... 2 — Carapace cylindrical, without strong longitudinal ridges but with ridges of spines; female pleopods 2 with stylamblys absent or minute ...... 3 2. Supraorbital spines fused in midline as a broad swollen rostrum ...... Linuparus — Supraorbital spines separated ...... Puerulus 3. Antennule with 2 very long flagella, much longer than peduncle ...... Panulirus — Antennule with 2 short flagella, shorter than half length of peduncle ...... 4 4. Dorsum of carapace with 2 submedian longitudinal rows of spines ...... Projasus — Dorsum of carapace densely and irregularly spinose ...... Jasus … 5 5. Scalloped sculpture on upper surface of abdominal somites ...... Jasus (Jasus) — Upper surface of abdominal somites smooth ...... Jasus (Sagmariasus)

199 Marine Decapod Crustacea of Southern Australia

Jasus (Jasus) Parker, 1883 The six species of Jasus each has a limited distribution on the continents and islands around the Southern Ocean and each is the basis of a commercial fishery. Much is known of the biology of the species in Australia, New Zealand and South Africa (Cobb & Phillips, 1980; Frusher et al., 1999). Two species live and are fished in Australia and frozen rock lobsters are imported from New Zealand for sale in Australia. Diagnosis. Carapace cylindrical. Supraorbital spines almost vertical. Eyes elevated. Without stridulating apparatus on antennae. Antennule with short flagella. Scalloped sculpture of upper surface of abdominal somites. Female pleopod 2 endopod broad, with reduced second article (stylamblys). Jasus (Jasus) edwardsii (Hutton, 1875) Australian subspecies: J. e. novaehollandiae Holthuis, 1963 southern rock lobster (Pls 2, 13a). Abdomen with dorsal squamiform sculpture; rostrum smaller than supraorbital spines; red. Maximum body length about 50 cm but individuals this long are rarely seen today. NSW (N to central coast), Vic., Tas., SA, WA (N to Dongara); crevices in immediate subtidal to about 90 m, sometimes to 200 m depth on rocky reefs. This is the commonest species of rock lobster in south-eastern Australia where it is the usual species in fish-shops and restaurants. Numbers are now reduced through commercial and recreational fishing since the mid-twentieth century but the commercial fishery is still valuable. Egg-laying occurs from May to July and the females carry the eggs on the pleopods and are said to be ‘in berry’. The larvae hatch from July to December. Phyllosoma larvae may actively migrate by vertical swimming. They remain in the water for 16–24 months passing through 11 moults and growing from 2 to 50 mm long. The rock lobster settles as a transparent puerulus preferring hard-walled rough shelters (Booth, 2002). Providing they escape capture, lobsters may live more than 20 years (Frusher et al., 1999). Because of its commercial importance there has been considerable interest in the biology and fishery potential of this species in New Zealand and in Australia (Mckoy, 1983: movements; Kailola et al., 1993: summary of resource use; Tudge et al., 1998: spermatozoal ultrastructure; Kelly et al., 1999: aggregations; Phillips & Melville-Smith, 1999: biology in Australia; Yearsley et al., 1999: identification as seafood). Southern rock lobsters from New Zealand and Australia are difficult to distinguish (George & Kensler, 1970) and are now regarded as conspecific on the basis of adult and larval morphology (Booth et al., 1990; Phillips & McWilliam, 1986), cross-breeding (Kittaka et al., 1988) and molecular studies that found that Australian and New Zealand populations have overlapping mitochondrial DNA haplotypes (Ovenden et al., 1992; Ovenden & Brasher 1994). In New Zealand lobsters the plates on abdominal somite 2 are said to be in two transverse rows and the antennular flagellum lacks banding; the plates are not in two rows and the flagellum has broad pale bands in Australian lobsters but these characters are unreliable. Phyllosomas from this complex of species are widespread through the Tasman Sea and the degree of genetic exchange is probably high with those from New Zealand settling in Australia. Holthuis (1963) discriminated adults of the two populations and gave the Australian one specific rank, J. novaehollandiae. George (1997) compromised and regarded them as emerging subspecies.

Jasus (Sagmariasus) Holthuis, 1991. This subgenus includes a single species (Holthuis, 1991). Diagnosis. Antennae slender. Rostrum as long as supraorbital spines. Dorsal surface of abdominal somites smooth. Jasus (Sagmariasus) verreauxi (Milne Edwards, 1851) eastern rock lobster (Pl. 3). Abdomen dorsally smooth; rostrum about as long as supraorbital spines; green. Maximum length about 60 cm. Southern Qld, NSW, Vic., eastern SA, northern Tas.; usually 20–100 m depth.

200 Achelata – rock lobsters and bugs

This species is rare over most of its range but common only around the Sydney region where it forms the basis of a commercial fishery (Montgomery, 1999). It is more common than the southern rock lobster only in these warmer waters. Holthuis (1991) erected the subgenus, Sagmariasus, for this species to distinguish it from the other six species of Jasus and Booth & Webber (2001) treated it as a full genus. What has long been regarded as the same species in northern New Zealand (packhorse lobster) is almost certainly a separate species. Brasher et al. (1992) and Ovenden & Brasher (1994) discovered significant genetic differences in mitochondrial DNA between the two populations. If so, either J. huegelii (Heller, 1862) or J. tumidus (Kirk, 1880) are applicable to the New Zealand species (George, 1997). Fisheries biology has been studied by numerous authors in Australia and New Zealand (Kensler, 1966: bibliography; 1967: fecundity, and other papers by this author). Feeding in the phyllosoma was the subject of a study by Nelson et al. (2002). Linuparus White, 1847 There are three species in the tropical Indo-West Pacific; two occur in Australia (Holthuis, 1991; Griffin & Stoddart, 1995). Diagnosis. Carapace prismatic. Supraorbital spines flattened and fused. Female pleopod 2 endopod narrowly lobed, with long second article (stylamblys). Eyes horizontal. With stridulating apparatus on antennae.

Key to southern Australian species of Linuparus 1. Paired ventral ridges (epistomal ridges) in front of mouth without sharp anterior tooth ...... Linuparus sordidus — Paired ventral ridges (epistomal ridges) in front of mouth with sharp anterior tooth ...... Linuparus trigonus

a b c

e f g h

Fig. 57. Palinuridae. Dorsal carapace: a, Linuparus sordidus. b, Projasus parkeri. c, Puerulus angulatus. d, Linuparus sordidus (epistome, maxillipeds 3). e, Linuparus trigonus (epistome). Abdominal somites 2 and 3, lateral: f, Panulirus cygnus. g, Panulirus homarus. h, Panulirus penicillatus (oblique view or anterior carapace).

201 Marine Decapod Crustacea of Southern Australia

Linuparus sordidus Bruce, 1965 Oriental spear lobster, sometimes box lobster or white spear lobster (Fig. 57a, d). Paired ventral ridges (epistomal ridges) in front of mouth without sharp anterior tooth, coarsely granulate. Length including antennae about 40 cm. Taiwan, South China Sea, NSW, WA (North West Shelf); rocky bottoms, 200–414 m depth, rare. Linuparus trigonus (von Siebold, 1824) Japanese spear lobster, sometimes barking lobster or champagne lobster in Australia (Fig. 57e). Paired ventral ridges (epistomal ridges) in front of mouth with sharp anterior tooth, feebly granulate; reddish-pink with cream bands and markings. Length including antennae about 50 cm. Japan, Korea, China, Taiwan, Philippines, WA (north-west), Qld (off Townsville), NSW; 30–414 m depth. The species occurs in sufficient densities in small areas around 200–250 m in tropical Australia to be fished occasionally and is sometimes for sale in southern states (Haddy et al., 2004).

Panulirus White, 1847 Of the 19 species that occur in the tropics and subtropics seven or eight occur in Australia (Holthuis, 1991; Chan & Ng, 2001). The key is to Australian species but only one, Panulirus cygnus, is common in the south. Keys to species of the genus have also been given by George & Holthuis (1965), George (1968) and Holthuis (1991). Other species, e.g. P. longipes and P. ornatus,may occasionally be seen on the south-eastern or south-western coasts. Differentiation of species is supported by morphological differences but colour patterns are also reliable (Jones & Morgan, 1994). Ptacek et al. (2002), on the basis of mitochondrial genes, found two major lineages of species within Panulirus.

Diagnosis. Carapace cylindrical. Supraorbital spines almost vertical. Female pleopod 2 endopod broad, without second article (stylamblys). Eyes elevated. Without stridulating apparatus on antennae. Antennule with long flagella.

Key to Australian species of Panulirus 1. Abdominal somites each with a transverse groove ...... 2 — Abdominal somites each without a transverse groove ...... 5 2. Anterior margin of abdominal transverse grooves crenulate; legs and abdomen spotted ...... Panulirus homarus (Linnaeus, 1758) Indo-West Pacific, including northern Australia — Anterior margin of abdominal transverse grooves not crenulate ...... 3 3. Antennular plate with 4 strong spines fused at base; legs striped, abdominal pleura spotted . Panulirus penicillatus (Olivier, 1791) Indo-West Pacific, including north-western Australia — Antennular plate with 2 equal principal spines ...... 4 4. Posterior margin of thoracic sternum of adult female with 2 distinct teeth; posterior half of abdominal somite 2 without pubescent area; carapace brightly spotted ...... Panulirus longipes Milne Edwards, 1868 Indian Ocean, north-eastern Australia to southern Qld; P. longipes bispinosus Borradaile, 1899 is the subspecies occuring in Australia — Posterior margin of thoracic sternum of adult female without 2 distinct teeth; posterior half of abdominal somite 2 with pubescent area behind transverse groove; carapace plain ...... Panulirus cygnus George, 1962 (see below) 5. Maxilliped 2 exopod with multiarticulate flagellum; merus of pereopods blotched, abdomen with narrow transverse pale bands ...... Panulirus polyphagus (Herbst, 1793) India, South-East Asia, northern Australia — Maxilliped 2 exopod flagellum small or absent ...... 6

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6. Abdomen uniformly coloured, with spots on sides, without narrow transverse pale bands, legs banded ...... Panulirus ornatus (Fabricius, 1798) Indo-West Pacific, N Australia to southern WA and NSW — Abdomen with conspicuous narrow transverse pale bands, carapace decorated ...... Panulirus versicolor (Latreille, 1804) Indo-West Pacific, including northern Australia Panulirus cygnus George, 1962 western rock lobster (Fig. 57f). WA (between North West Cape and Cape Leeuwin); on limestone reefs, 30–90 m and rarely to 200 m depth. This is the only palinurid regularly seen in south-western Australia and is the basis of an important fishery. Considerable research on the biology of the species and management of the fishery has been done by Western Australian Department of Fisheries and CSIRO Division of Fisheries until it is now one of the best understood marine animals in Australia. There is an excellent book written about it presenting detailed information on its biology to a lay audience (Gray, 1991) and a summary in Phillips & Melville-Smith (1999). The adult lobsters live under ledges in the extensive limestone reefs within 60 km of the south part of the western coast of WA where they feed on small invertebrates and encrusting seaweeds. Adults are largely confined to the outer shelf and mating occurs in July and August. Egg-laying on to the pleopods of the female occurs between August and December. A single full-grown female may hold as many as one million eggs at one time and many produce two broods a year. Incubation time varies from about 20 days in warmer climates to almost 70 further south. Hatching is generally complete at the end of February or March and the 1.5 mm long phyllosoma larva begins its planktonic life. In their 9–11 months at sea the phyllosomas may travel as far as 1500 km from the coast and grow to 35 mm long. The final moult to the puerulus stage takes place near the edge of the continental shelf, most frequently in the spring when the Leeuwin Current is strongest (Caputi et al., 2002), and this miniature lobster swims to the shore to settle. With luck, the lobster will live to 10 or 15 years, reaching maturity at about 6–7 years. Animals as big as 450 mm total length have been recorded but strong commercial fishing pressure ensures that none survives to reach that size today. The discreteness of this species from P. longipes was confirmed on the basis of allozyme variation by Thompson (1996).

Projasus George & Grindley, 1964 The genus comprises two species, one in Chile and one in Australia, New Zealand and South Africa. Both are rarely taken. Diagnosis. Dorsum of carapace with 2 submedian longitudinal rows of spines. Supraorbital spines almost vertical. Female pleopod 2 endopod broad, with reduced second article (stylamblys). Eyes elevated. Without stridulating apparatus on antennae. Antennule with short flagella. Projasus parkeri (Stebbing, 1902) Parker’s crayfish (Fig. 57b). Merus and ischium of pereopods without spinules on posterior margin. Southern NSW, eastern Vic., Tas., south-western WA; 370–880 m depth. A single individual puerulus stage of Projasus from off NSW was described by Webber & Booth (1988) but not assigned to species. Adults of P. parkeri were first reported by Griffin & Stoddard (1995).

Puerulus Ortmann, 1897 Four species are described, all from the Indo-West Pacific (Holthuis, 1991; Griffin & Stoddart, 1995). Puerulus is both a genus name as here and the name given to the first-settled stage of palinurids in general.

203 Marine Decapod Crustacea of Southern Australia

Diagnosis. Carapace prismatic. Supraorbital spines separate, small, flattened. Female pleopod 2 endopod narrowly lobed, with long second article (stylamblys). Eyes horizontal. With stridulating apparatus on antennae. Puerulus angulatus (Bate, 1888) banded whip lobster (Fig. 57c). Post orbital spine absent; median keel of carapace with 3 post cervical and 2 intestinal teeth; 3 teeth between frontal horn and cervical groove. Light orange overall, pereopods white, bands on abdomen. Indo-West Pacific, WA (north), Qld, NSW; 192–536 m depth. The species is too rare to be fished. Griffin & Stoddart (1995) reported the species from eastern Australia.

References Booth, J.D. 2002. Habitat preferences and behaviour of newly settled Jasus edwardsii (Palinuridae). Marine and Freshwater Research 52: 1055–1065. Booth, J.D., Street, R.J., & Smith, P.J. 1990. Systematic status of the rock lobsters Jasus edwardsi from New Zealand and Jasus novaehollandiae from Australia. New Zealand Journal of Marine and Freshwater Research 24: 239–249. Booth, J.D., & Webber, R. 2001. All the pretty lobsters – part 1. Seafood New Zealand 9 (11): 20–23. Brasher, D.J., Ovenden, J.R., Booth, J.D., & White, R.W.G. 1992. Genetic subdivison of Australian and New Zealand populations of Jasus verreauxi (Decapoda: Palinuridae) preliminary evidence from the mitochondrial genome. New Zealand Journal of Marine and Freshwater Research 26: 53–58. Caputi, N., Chubb, C., & Pearce, A. 2002. Environmental effects on recruitment of the western rock lobster, Panulirus cygnus. Marine and Freshwater Research 52: 1167–1174. Chan, T.Y. 1998. Lobsters. Pp. 973–1043 in: Carpenter, K.E., & Niem, V.H. (eds), The Living Resources of the Western Central Pacific. FAO: Rome. Chan, T.-Y., & Ng, P.K.L. 2001. On the nomenclature of the commercially important spiny lobsters Panulirus longipes femoristriga (Von Martens, 1872), P. bispinosus Borridaile, 1899, and P. albiflagel- lum Chan & Chu, 1996 (Decapoda, Palinuridae). Crustaceana 74: 123–127. Cobb, J.S., & Phillips, B.F. 1980. The Biology and Management of Lobsters. 1 Academic Press: New York. 463 pp. Davie, P.J.F. 1990. A new genus and species of marine crayfish, Palibythus magnificus, and new records of Palinurellus (Decapoda: Palinuridae) from the Pacific Ocean. Invertebrate Taxonomy 4: 685–695. Frusher, S., Prescott, J., & Edmunds, M. 1999. Southern rock lobsters. Pp. 106–113 in: Andrew, N. (ed.) Under Southern Seas. The Ecology of Australia’s Rocky Reefs. UNSW Press: Sydney. George, R.W. 1966. Marine crayfish or spiny lobsters of Australia. Australian Fisheries Newsletter. George, R.W. 1968. Tropical spiny lobsters, Palinurus spp., of Western Australia (and the Indo-West Pacific). Journal of the Royal Society of Western Australia 51: 33–38. George, R.W. 1983. New finds of deepwater ‘lobsters’ on the Northwest Shelf. FINS (Fisheries News), Perth 16: 16–20. George, R.W. 1997. Tectonic plate movements and the evolution of Jasus and Panulirus spiny lobsters (Palinuridae). Marine and Freshwater Research 48: 1121–1130. George, R.W., & Holthuis, L.B. 1965. A revision of the Indo-West Pacific spiny lobsters of the Panulirus japonicus group. Zoologische Verhandelingen, Leiden 72: 1–36,pls1–5. George, R.W., & Kensler, C.B. 1970. Recognition of marine spiny lobsters of the Jasus lalandii group (Crustacea: Decapoda: Palinuridae). New Zealand Journal of Marine and Freshwater Research 4: 292–311. George, R.W., & Main, A.R. 1967. The evolution of the spiny lobsters (Palinuridae): a study of evolution in the marine environment. Evolution 21: 803–820. Gray, H. 1991. The Western Rock Lobster Palinurus cygnus. Book 1: A Natural History. Westralian Books: Geraldton. 112 pp. Griffin, D.J.G., & Stoddart, H.E. 1995. Deep-water decapod Crustacea from eastern Australia: lobsters of

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the families Nephropidae, Palinuridae, Polychelidae and Scyllaridae. Records of the Australian Museum 47: 231–263. Haddy, J.A., Roy, D.P., & Courtney, A.J. 2004. The fishery and reproductive biology of barking crayfish, Linuparus trigonus (Von Siebold, 1824) along Queenslands east coast. Crustaceana 76: 1189–1200. Holthuis, L.B. 1963. Preliminary descriptions of some new species of Palinuridea (Crustacea Decapoda, Macrura Reptantia). Proceedings Koninklijke Nederlandse Akademie van Wetenschappen 66: 54–60. Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125: 1–292. Johnston, D.J., & Ritar, A. 2002. Mouthpart and foregut ontogeny in phyllosoma larvae of the spiny lobster Jasus edwardsii (Decapoda: Palinuridae). Marine and Freshwater Research 52: 1375–1386. Jones, D.S., & Morgan, G.J. 1994. A Field Guide to Crustaceans of Australian Waters. Reed: Sydney. 216 pp. Kailola, P.J., Williams, M.J., Stewart, P.C., Reichelt, R.E., McNee, A., & Grieve, C. 1993. Australian Fisheries Resources. Bureau of Resource Sciences, Department of Primary Industries and Energy, and Fisheries Research and Development Corporation: Canberra. 422 pp. Kelly, S., MacDiarmid, A.B., & Babcock, R.C. 1999. Characteristics of spiny lobster, Jasus edwardsii, aggregations in exposed reef and sandy areas. Marine and Freshwater Research 50: 409–416. Kensler, C.B. 1966. An annotated bibliography of the marine spiny lobster Jasus verreauxi (H. Milne Edwards) (Crustacea, Decapoda, Palinuridae). Transactions of the Royal Society of New Zealand (Zoology) 19: 207–210. Kensler, C.B. 1967. Fecundity in the marine spiny lobster Jasus verreauxi (H. Milne Edward) (Crustacea: Decapoda: Palinuridae). New Zealand Journal of Marine and Freshwater Research 2: 143–155. Kittaka, J., Iwai, M., & Yoshimura, M. 1998. Culture of a hybrid of spiny lobster genus Jasus from egg stage to peurulus. Nippon Suisan Gakkaishi 54: 413–417. McCoy, F. 1885–1890. Prodromus of the Zoology of Victoria: or Figures and Descriptions of the Living Species of All Classes of the Victorian Indigenous Animals. Vol 2. Government Printer: Melbourne. Mckoy, J.L. 1983. Movements of rock lobsters, Jasus edwardsii (Decapoda: palinuridae), tagged near Stewart Island, New Zealand. New Zealand Journal of Marine and Freshwater Research 17: 357–366. Montgomery, S. 1999. Eastern rock lobsters. Pp. 114–117 in: Andrew, N. (ed.) Under Southern Seas. The Ecology of Australia’s Rocky Reefs. UNSW Press: Sydney. Nelson, M.M., Cox, S.L., & Ritz, D.A. 2002. Function of mouthparts in feeding behaviour of phyllosoma larvae of the packhorse lobster, Jasus verreauxi (Decapoda: Palinuridae). Journal of Crustacean Biology 22: 595–600. Ovenden, J.R., & Brasher, D.J. 1994. Stock identity of the red (Jasus edwardsii) and green (Jasus verreauxi) rock lobsters inferred from mitochondrial DNA analysis. Pp. 230–249 in: Phillips, B.F., Cobb, J.S., & Kittaka, J. (eds), Spiny Lobster Management. Blackwell Scientific Publications: Oxford. Ovenden, J.R., Brasher, D.J., & White, R.W.G. 1992. Mitochondrial DNA analyses of the red rock lobster Jasus edwardsii supports an apparent absence of population subdivision throughout Australia. Marine Biology 112: 319–326. Phillips, B.F., Cobb, J.S., & George, R.W. 1980. General biology. Pp. 2–89 in: Cobb, J.S., & Phillips, B.F. (eds), The Biology and Management of Lobsters. Academic Press: New York. Phillips, B.F., & Melville-Smith, R. 1999. Chapter 14 Western Rock Lobsters. Pp. 118–125 in: Andrew, N. (ed.) Under Southern Seas. The Ecology of Australia’s Rocky Reefs. UNSW Press: Sydney. Phillips, B.F.,& McWilliam, P.S.1986. The pelagic phase of spiny lobster development. Canadian Journal of Fisheries and Aquatic Sciences 43: 2153–2163. Phillips, B.F., Rimmer, D.W., & Reid, D.D. 1978. Ecological investigations of the late stage phyllosoma and puerulus larvae of the western rock lobster Panulirus longipes cygnus. Marine Biology 45: 347–357. Ptacek, M.B., Sarver, S.K., Childress, M.J., & Herrnkind, W.F. 2002. Molecular phylogeny of the spiny lobster genus Panulirus (Decapoda: Palinuridae). Marine and Freshwater Research 52: 1037–1047.

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Thompson, A.P.1996. A genetic perspective on the specific status of the Western Rock Lobster along the coast of Western Australia – Panulirus cygnus George 1962 or P. longipes A Milne Edwards 1868? Journal of the Royal Society of Western Australia 79: 207–210. Tudge, C.C., Scheltinga, D.M., & Jamieson, B.G.M. 1998. Spermatozoal ultrastructure in the spiny lobster Jasus novaehollandiae Holthuis, 1963 (Palinuridae, Palinura,Decapoda). Journal of Morphology 236: 117–126. Webber, W.R., & Booth, J.D. 1988. Projasus parkeri (Stebbing, 1902) (Crustacea, Decapoda, Palinuridae) in New Zealand and description of a Projasus puerulus from Australia. National Museum of New Zealand Records 3: 81–92. Williams, A.B. 1988. Lobsters of the World -an Illustrated Guide. Osprey Books: New York. 186 pp. Yearsley, G.K., Last, P.R., & Ward, R.D. 1999. Australian Seafood Handbook. An Identification Guide to Domestic Species. CSIRO Marine Research: Hobart. 461 pp.

THREE COMMON SOUTHERN LOBSTERS Only two of the 13 species of Australian rock lobsters are commonly caught in southern Australia. A third occurs in NSW. 1. Antennules (smaller of the 2 pairs) with long flagella; red carapace, spotted abdomen (WA only) ...... western rock lobster Panulirus cygnus — Antennules (smaller of the 2 pairs) with very short flagella; red or green all over . . . . 2 2. Red; abdomen with plate-like sculpture; common commercially caught species ...... southern rock lobster Jasus (Jasus) edwardsii — Green; abdomen smooth; uncommon, south-eastern coast only ...... eastern rock lobster Jasus (Sagmariasus) verreauxi Other lobster-like animals are scyllarids, of which only the Balmain bug (Ibacus peronii) is common, and the deep-water, long-handed polychelids.

Scyllaridae Latreille, 1825 Scyllarids are easily recognised by their flattened antennae which give them the common name of shovel-nosed lobsters. They are also known as slipper lobsters but in Australia the two most common edible species are the Balmain bug (Ibacus peronii) and the Moreton Bay bug (Thenus sp.). A popular account of the biology of the Australian species was given by George & Griffin (1972) but the definitive works on their systematics are by Holthuis (1985; 2002). He also treated the family from a fisheries perspective (Holthuis, 1991) and then from an historical angle (Holthuis, 1996). Unlike the palinurid lobsters which are associated with reefs, scyllarids inhabit soft substrates and burrow into the sand or mud. Scyllarids have phyllosoma larvae like palinurids that have often been recorded associated with medusae (jelly fishes) in the open ocean. The larval development of the species now known as Petrarctus demani (Holthuis, 1946) from northern Australia is known (Ito & Lucas, 1990). Twenty genera are recognised with more than 50 named species. Almost 30 species in 15 genera occur in Australia; to those catalogued by Davie (2002) must be added new records of the subfamily Scyllarinae (Holthuis, 2002). Several species from the New Caledonian region described by Holthuis (2002) might be expected to be encountered in deep water in southern Australia in the future. This last revision excluded the well-known genus Scyllarus Fabricius, 1775 from the Indo- West Pacific and placed all species from the region in new genera. For identification of species outside this region refer to Williams (1988).

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The Moreton Bay bug, as it is known in Australia, is a complex of species not occurring in southern Australia but seen in fish markets. Balmain bugs are more frequent in Melbourne shops, commonly mislabelled as Moreton Bay bugs. Moreton Bay bugs have been frequently referred to Thenus orientalis (Lund, 1793), widespread throughout the Indo-West Pacific, and sometimes to T. indicus Leach 1816, but recent molecular, spermatological and morphological research has discovered that neither of these species occurs in Australia (Jones, 1990, 1993; Davie, 2002). The species of Thenus are immediately identifiable by being flat and having the eyes on the extreme anterolateral corners of the carapace. The species are caught in prawn trawls on muddy and sandy bottoms along the northern coast of Australia between Moreton Bay, Qld, and Exmouth Gulf, WA. In Moreton Bay itself there is a striped-legged species on muddy bottoms with clear to moderately turbid inshore waters, while a spotted-legged species lives on sandy or shelly bottoms around reefs in cleaner water (Davie, 1998). Mikami & Greenwood (1997) described the complete larval development of these two new species from Moreton Bay. Johnston with others detailed the diet and functional morphology of the feeding apparatus in species of Thenus (Johnston, 1994; Johnston & Yellowlees, 1998; Johnston & Alexander, 1999).

Diagnosis. Antenna peduncle wide and flat, flagellum transformed to a single wide flat unseg- mented plate. Carapace more or less flattened.

Key to southern Australian genera of Scyllaridae (plus Thenus) 1. Exopods of all maxillipeds with multiarticulate flagellum ...... 2 — Maxillipeds 1 and 3 exopods without flagellum; maxilliped 2 exopod flagellum transformed to a single laminate segment; maxilliped 3 exopod rudimentary ...... 4 2. Carapace strongly depressed, with a deep lateral cervical incision ...... Ibacus — Carapace highly vaulted, with shallow or no lateral cervical incision ...... 3 3. Abdominal somite 1 with dorsal transverse groove between anterior and posterior sections; abdominal somites distinctly sculptured laterally; carapace with postorbital spine . . . Arctides — Abdominal somite 1 without dorsal transverse groove; abdominal somites with only median ridge; carapace without postorbital spine ...... Scyllarides 4. Orbits on anterolateral angles of carapace; carapace strongly depressed . . . Thenus Leach, 1816 — Orbits on anterior margin of carapace; carapace highly vaulted ...... 5 5. Posterior section of dorsal surface of abdominal somites 2–4 with wide transverse groove over middle, sometimes with tubercles on either side, but without arborescent pattern of narrow grooves; abdominal somites with distinct median longitudinal carina sharply set off from rest of dorsal surface ...... 6 — Posterior section of dorsal surface of abdominal somites 2–4 with arborescent pattern of a narrow central transverse groove with side grooves, that often are branched ...... 7 6. Anterior margin of thoracic sternum truncate or convex, sometimes with median tubercle, without median incision; this margin on about the same level as the anterolateral teeth of rostrum; abdominal somite 3 without elevated medial ridge; pereopod 1–4 propodi often with setae along lower margin ...... Bathyarctus — Anterior margin of thoracic sternum U- or V-shaped, not sunken, with median incision; pereopods 1, 2 and 4 without setae on lower margin ...... Antarctus 7. Pleura of abdominal somites 2–4 ending in sharp posteriorly directed point . . . .Galearctus — Pleura of abdominal somites 2–4 with the apex blunt or rectangularly rounded . . . Crenarctus

207 Marine Decapod Crustacea of Southern Australia

ab c

Fig. 58. Scyllaridae. a, Antarctus mawsoni. b, Arctides antipodarum. c, Crenarctus crenatus. d, Ibacus alticrenatus. e, Ibacus peronii.

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Antarctus Holthuis, 2002 The Australian species is the only member of the genus. Diagnosis. Carapace with rostral, pregastric, gastric and cardiac teeth in midline. Posterior section of dorsal surface of abdominal somites 2–4 each with transverse groove and median carina; without arborescent arrangement of narrow grooves. Antenna article 4 with 2 oblique dorsal carinae. Maxillipeds 1 exopod without flagellum. Maxilliped 2 exopod flagellum a single laminate segment. Maxilliped 3 exopod rudimentary. Pereopods without fringes of setae. Thoracic sternum with anterior margin having deep median V-shaped notch between 2 teeth. Antarctus mawsoni (Bage, 1938) (Fig. 58a). Anterior margin of antenna article 4 with 5–8 acute triangular spines; with 2 similar strong acute middorsal carinae. tl. 90 mm. Southern NSW, Vic., Tas., SA; 80–540 m depth.

Arctides Holthuis, 1960 There are three species of which one occurs in southern Australia. Diagnosis. Carapace highly vaulted, with shallow or no lateral cervical incision, with postorbital spine. Exopods of all maxillipeds with multiarticulate flagellum. Abdominal somite 1 with dorsal transverse groove between anterior and posterior sections. Abdominal somites distinctly sculptured laterally. Arctides antipodarum Holthuis, 1960 rough Spanish lobster (Fig. 58b). With weak abdominal sculpture. 300 mm, usually 100 mm. Northern New Zealand, central NSW; rocky habitats, 5–146 m, uncommon. Known in some older works as Scyllarus sculptus.

Bathyarctus Holthuis, 2002 Six species are known, all in deep water; only one has been recorded from Australia and another from New Caledonia. Diagnosis. Carapace with pregastric, gastric and cardiac teeth in midline, without sharp rostral tooth. Antenna article 4 with 2 oblique dorsal carinae. Maxilliped 1 exopod without flagellum. Maxilliped 2 exopod flagellum a single laminate segment. Maxilliped 3 exopod rudimentary. Abdominal somites with prominent median carinae, fourth higher than on third; without arborescent arrangement of narrow grooves. Thoracic sternum with anterior margin not incised. Bathyarctus rubens (Alcock & Anderson, 1894) (Fig. 59h). Thoracic sternum with anterior margin slightly concave. Abdominal somites 1–4 with posterior section smooth; median carina 4 conspicuously higher than 3. Pale brown. cl. 26 mm. Indian Ocean, Philippines, Fiji, NSW; 183–732 m depth.

Crenarctus Holthuis, 2002 Only one species occurs in southern Australia. Diagnosis. Carapace highly vaulted, rostral and pregastric teeth distinct, gastric and cardiac teeth each 2 tubercles; orbits on anterior margin. Antenna, article 4 with single oblique carina over its full length. Maxillipeds 1 exopod without flagellum. Maxilliped 2 exopod flagellum a single laminate segment. Maxilliped 3 exopod rudimentary. Anterior margin of thoracic sternum with U-shaped incision with 2 small tubercles flanking median incision. Posterior section of dorsal surface of abdominal somites 2–4 with arborescent pattern of a narrow central transverse groove with side grooves; pleura with apex blunt or rectangularly rounded.

209 Marine Decapod Crustacea of Southern Australia

b cd

ef g

hih

Fig. 59. Scyllaridae. Lateral carapace and anterior abdomen: a, Antarctus mawsoni. b, Crenarctus crenatus. Carapace and antennae: c, Ibacus brucei. d, Ibacus chacei. e, Galearctus umbilicatus (thoracic sternum). f, Ibacus alticrenatus (maxillipeds 3 and epistome). g, Ibacus peronii (maxillipeds 3 and epistome). h, Bathyarctus rubens (lateral abdomen). i, Galearctus umbilicatus (abdominal somite 2). j, Scyllarides haanii (abdomen).

Crenarctus crenatus (Whitelegge, 1900) (Figs 58c, 59b, Pl. 13b). Submedian carina ending in strong tooth. Abdominal somite 2 with median area elevated. Anterior margin of antenna with 5 rounded lobes. cl. 22 mm. Qld, NSW, Vic., Tas., WA (Rottnest I.); intertidal (rare) to 250 m depth. Jones & Morgan’s (1993) record of Scyllarus bicuspidatus (de Man, 1905) from Rottnest I., WA, is of this species.

Galearctus Holthuis, 2002 Of five Indo-West Pacific species one has been recorded from Australia and others from the New Caledonian region. Diagnosis. Carapace highly vaulted, with rostral, gastric and cardiac teeth, without pregastric tooth; orbits on anterior margin. Antenna, article 4 with single oblique carina over its full length.

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Maxillipeds 1 exopod without flagellum. Maxilliped 2 exopod flagellum a single laminate segment. Maxilliped 3 exopod rudimentary. Anterior part of thoracic sternum tongue-like, produced forward, anterior margin a triangle with a minute median incision. Posterior section of dorsal surface of abdominal somites 2–4 with arborescent pattern of a narrow central transverse groove with side grooves; pleura ending in sharp posteriorly-directed point. Galearctus umbilicatus (Holthuis, 1977) (Fig. 59e). Carapace with high gastric tooth. Anterior margin of antenna with acute triangular spines. Pereopod 2 propodus narrower than that of pereopod 3. Abdominal somites, anterior section with transverse grooves lined with setae; somites 2–5 with 2 crenulated grooves. cl. 24 mm. Southern Qld, NSW; 55–232 m depth.

Ibacus Leach, 1815 Eight species are known, all from the Indo-West Pacific (Holthuis, 1985; Brown & Holthuis, 1998). Brown & Holthuis (1998) provided a key and colour pictures. In south-eastern Australia four species exist, all known commonly as Balmain Bugs, but others inhabit northern Australia. They are a popular local delicacy. Diagnosis. Carapace strongly depressed, with a deep lateral cervical incision, dorsal surface smooth, with distinct longitudinal medial and branchial carinae; distance between eyes about equal to distance from eye to margin. Exopods of all maxillipeds with multiarticulate flagellum.

Key to southern Australian species of Ibacus 1. Maxilliped 3 merus raised, not coloured differently from rest, with 6–8 transverse septa; posterior epistomal tooth sharp ...... 2 — Maxilliped 3 merus flat, yellowish, without transverse septa; posterior epistomal tooth rounded or truncate ...... 3 2. Posterior branchial ridges of carapace convex; maxilliped 3 merus evenly swollen; carapace evenly granulate and with short setae ...... Ibacus peronii — Posterior branchial ridges of carapace almost straight; maxilliped 3 merus with distal spherical knob; carapace evenly punctate and naked ...... Ibacus chacei 3. Carapace pitted, sparsely pubescent; article 4 of antenna widening laterad before narrowing to apex, with distinct toothed lateral margin ...... Ibacus brucei — Carapace evenly pubescent; article 4 of antenna slender, without lateral teeth ...... Ibacus alticrenatus Ibacus alticrenatus Bate, 1888 velvet fan lobster (Fig. 58d). Maxilliped 3 merus flat, without transverse septa. Branchial ridges straight. Carapace evenly pubescent. tl. 200 mm. Northern New Zealand, Qld, NSW,Vic., northern Tas.,SA; soft sediments, 20–686 m depth. This species which appears in Sydney fish markets is best separated from the more common I. peronii by having straight branchial ridges. The larval stages were described by Atkinson & Boustead (1982). Ibacus brucei Holthuis, 1977 glabrous fan lobster (Fig. 59c). Maxilliped 3 merus flat without transverse septa. Branchial ridges straight and diverging. Carapace minutely pitted, almost naked. tl. 130 mm. Kermadec Is., New Zealand, Southern Qld, central NSW; 83–559 m depth (uncommon). Ibacus chacei Brown & Holthuis, 1998 (Fig. 59d). Maxilliped 3 merus swollen, rounded, produced as distal knob, with transverse septa. Branchial ridges almost straight. Carapace evenly punctate and naked. tl. 23 cm. Central Qld, NSW; 22–330 m depth. The species contributes to the catch of Balmain bugs in Qld and NSW. The species rarely breeds as far south as central NSW (Stewart et al., 1997).

211 Marine Decapod Crustacea of Southern Australia

Ibacus peronii Leach, 1815 Balmain bug, or butterfly fan lobster in FAO Catalogue (Figs 58e, 59g, Pl. 6). Maxilliped 3 merus raised with transverse septa. Branchial ridges markedly curved. Carapace evenly granulate, with cover of short setae. Antennule article 4 laterally toothed. Dull reddish. tl. 230 mm. Qld (N to Southport), NSW, Vic., northern Tas., SA, WA (N to Geraldton); soft sand and mud, 4–250 m depth. This is the most common species encountered in south-eastern Australia where it lives in bays and on the shelf. It is excellent eating but not as popular as rock lobster, said by some to taste and smell strongly of garlic. The species was figured as I. incisus by Hale (1927) who called it the ‘squagga’ or ‘prawn killer’,names not in common use today. The phyllosoma stages have been described by Ritz & Thomas (1973). Stewart with others described the life history (Stewart & Kennelly, 1997; Stewart et al., 1997).

Scyllarides Gill, 1898 The 13 species were keyed and illustrated by Holthuis (1991). Some are large and edible. These are essentially tropical species occasionally taken by divers and in lobster pots on coral reefs. Diagnosis. Carapace highly vaulted, with shallow or no lateral cervical incision; carapace without postorbital spine. Abdominal somite 1 without dorsal transverse groove; abdominal somites with only median ridge. Exopods of all maxillipeds with multiarticulate flagellum.

Key to southern Australian species of Scyllarides 1. Abdominal somite 4 with strong median hump, larger than more anterior ones; abdominal somite 2 with blunt tooth posteriorly ...... Scyllarides haanii — Abdominal somites with median carinae of similar size; abdominal somite 2 evenly curved posteriorly ...... Scyllarides squammosus Scyllarides haanii (De Haan, 1841) Aesop slipper lobster (Fig. 59j). Abdominal somite 4 with strong median hump, larger than more anterior ones. Abdominal somite 2 with blunt tooth posteriorly. tl. 500 mm. Indo-West Pacific, WA (S to Rottnest I.), Qld, Lord Howe I.; 10–135 m depth. Scyllarides squammosus (Milne Edwards, 1837) blunt slipper lobster. Abdominal somites with median carinae of similar size. Abdominal somite 2 evenly curved posteriorly. tl. 400 mm. Indo-West Pacific, WA (S to central coast), NSW; 20–80 m depth, rare in southern waters.

References Atkinson, J.M., & Boustead, N.C. 1982. The complete larval development of the scyllarid lobster Ibacus alticrenatus Bate, 1888 in New Zealand waters. Crustaceana 42: 275–287. Brown, D.E., & Holthuis, L.B. 1998. The Australian species of the genus Ibacus (Crustacea: Decapoda: Scyllaridae), with description of a new species and addition of new records. Zoologische Mededelingen, Leiden 72: 113–141. Davie, P.D.F. 1998. Wild Guide to Moreton Bay: Wildlife and Habitats of a Beautiful Australian Coast –Noosa to the Tweed. Queensland Museum: Brisbane. 408 pp. Davie, P.J.F. 2002. Crustacea: Malacostraca: Phyllocarida, Hoplocarida, Eucarida (Part 1). In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3A. CSIRO Publishing: Melbourne. xii, 551 pp. George, R.W., & Griffin, D.J.G. 1972. The shovel nosed lobsters of Australia. Australian Natural History: 227–231. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Holthuis, L.B. 1985. A revision of the family Scyllaridae (Crustacea: Decapoda: Macrura). I. Subfamily Ibacinae. Zoologische Verhandelingen, Leiden 218: 1–130.

212 Achelata – rock lobsters and bugs

Holthuis, L.B. 1991. FAO species catalogue. Vol. 13. Marine lobsters of the world. An annotated and illustrated catalogue of species of interest to fisheries known to date. FAO Fisheries Synopsis 125: 1–292. Holthuis, L.B. 1996. The scyllarid lobsters (Crustacea: Decapoda: Palinuridea) collected by F. Péron and C. A. Lesueur during the 1800–1804 expedition to Australia. Zoologische Mededelingen, Leiden 70: 261–270. Holthuis, L.B. 2002. The Indo-Pacific scyllarine lobsters (Crustacea, Decapoda, Scyllaridae). Zoosystema 24: 499–683. Ito, M., & Lucas, J.S. 1990. The complete larval development of the scyllarid lobster, Scyllarus demani Holthuis, 1946 (Decapoda, Scyllaridae), in the laboratory. Crustaceana 58: 144–167. Johnston, D.J. 1994. Functional morphology of the membraneous lobe within the preoral cavity of Thenus orientalis (Crustacea: Scyllaridae). Journal of the Marine Biological Association of the United Kingdom 74: 787–800. Johnston, D.J., & Yellowlees, D. 1998. Relationships between dietary preferences and digestive enzyme complement of the slipper lobster Thenus orientalis (Decapoda: Scyllaridae). Journal of Crustacean Biology 18: 656–665. Johnston, D.J., & Alexander, C.G. 1999. Functional morphology of the mouthparts and alimentary tract of the slipper lobster Thenus orientalis (Decapoda: Scyllaridae). Marine and Freshwater Research 50: 213–223. Jones, C.M. 1990. Morphological characteristics of bay lobsters, Thenus Leach species (Decapoda, Scyllaridae), form north-eastern Australia. Crustaceana 59: 265–275. Jones, C.M. 1993. Population structure of Thenus orientalis and T. indicus (Decapoda: Scyllaridae) in northeastern Australia. Marine Ecology Progress Series 97: 143–155. Jones, D.S., & Morgan, G.J. 1993. An annotated checklist of the Crustacea of Rottnest Island, Western Australia. Pp. 135–162 in: Wells, F.E., Walker, D.I., Kirkman, H., & Lethbridge, R. (eds), Proceedings of the Fifth International Marine Biological Workshop: The Marine Flora and Fauna of Rottnest Island, Western Australia. Western Australian Museum: Perth. McCoy, F. 1885–1890. Prodromus of the Zoology of Victoria: or Figures and Descriptions of the Living Species of all Classes of the Victorian Indigenous Animals. Vol. 2. Government Printer: Melbourne. Mikami, S., & Greenwood, J.G. 1997. Complete development and comparative morphology of larval Thenus orientalis and Thenus sp. (Decapoda: Scyllaridae) reared in the laboratory. Journal of Crustacean Biology 17: 289–308. Ritz, D.A., & Thomas, L.R. 1973. The larval and postlarval stages of Ibacus peronii Leach (Decapoda, Reptantia, Scyllaridae). Crustaceana 24: 5–16, 1 pl. Stewart, J., & Kennelly, S.J. 1997. Fecundity and egg-size of the Balmain bug Ibacus peronii (Leach, 1815) (Decapoda, Scyllaridae) off the east coast of Australia. Crustaceana 70: 191–197. Stewart, J., Kennelly, S.J., & Høegh-Guldberg, O. 1997. Size at sexual maturity and the reproductive biology of two species of scyllarid lobster from New South Wales and Victoria, Australia. Crustaceana 70: 344–367. Williams, A.B. 1988. Lobsters of the World – An Illustrated Guide. Osprey Books: New York. 186 pp.

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b c

d e Plate 1. a, South-eastern Australia viewed from the South (computer-generated image prepared from swath mapping data for the National Oceans Office by Geoscience Australia. Provided courtesy of the National Oceans Office). The continental Australian and Tasmanian land masses and other islands are red with higher mountains in white; continenal shelf orange; continental slope and rise yellow to green; and abyssal depths blue. Depths are considerably exaggerated. Typical marine habitats of south-eastern Australia: b, estuary in eastern Victoria. c, intertidal rock platforms of western Victoria. d, epifauna trawled from shelf sediments of Bass Strait. e, subtidal epifauna on wharf pile, Port Victoria, South Australia. Plate 2. Achelata Palinuridae. Jasus edwardsii novaehollandiae. Reproduced from McCoy (1887: pl. 149) Prodromus of the Zoology of Victoria. Plate 3. Achelata Palinuridae. Jasus verreauxi. Reproduced from McCoy (1888: pl. 159) Prodromus of the Zoology of Victoria. Plate 4. Brachyura Eriphiidae. Pseudocarcinus gigas (juvenile). Reproduced from McCoy (1889: pl. 179) Prodromus of the Zoology of Victoria. Plate 5. Brachyura Eriphiidae. Pseudocarcinus gigas (adult male). Reproduced from McCoy (1889: pl. 180) Prodromus of the Zoologyof Victoria. Plate 6. Achelata Scyllaridae. Ibacus peronii. Reproduced from McCoy (1890: pl. 199) Prodromus of the Zoology of Victoria. a

e f Plate 7. Dendrobranchiata Penaeidae. a, Penaeus latisulcatus. b, Penaeus plebejus. Caridea Rhynchocinetidae. c, Rhynchocinetes australis. d, Rhynchocinetes serratus. Gnathophyllidae. e, Gnathophyllum americanum. f, Gnathophyllum taylori. a b

d e Plate 8. Caridea Palaemonidae. a, Palaemon dolospina. b, Periclimenes aesopius. c, Palaemon serenus. Alpheidae. d, Alpheus astrinx. e, Alpheus parasocialis. a b

c d

e f

g h Plate 9. Caridea Alpheidae (cont.). a, Alpheus bidens. b, Alpheus richardsoni. c, d, Alpheus villosus (in situ and in vitro). e, Athanopsis australis. f, Synalpheus tumidomanus. Hippolytidae. g, Alope orientalis. h, Nauticaris marionis. a b

c d

e g

h i Plate 10. Caridea Hippolytidae (cont.). a, b, Hippolyte australiensis (two colour morphs on red and green algae). c, d, Latreutes compressus (two colour morphs). e, Saron marmoratus. f, Tozeuma elongatum. Pandalidae. g, Chlorotocella spinicaudus. Crangonidae. h, Paracrangon australis. i, Philocheras intermedius. a b

c d

f g

h Plate 11. Stenopodidea Spongicolidae. a, Microprosthema validum. Stenopodidae. b, Stenopus hispidus. c, Stenopus tenuirostris. Astacidea Enoplometopidae. d, Enoplometopus occidentalis. Nephropidae. e, Metanephrops australiensis. f, Metanephrops boschmai. g, Metanephrops sibogae. Thalassinidea Axiidae. h, Axiopsis werribee. a b

c d

e f g Plate 12. Thalassinidea Callianassidae. a, Biffarius arenosus. b, Biffarius limosus. c, Trypaea australiensis. d, resin cast of burrow of Biffarius arenosus showing larger inhalent siphon and smaller exhalent siphon reaching to sediment surface and turning chambers. Strahlaxiidae. e, Strahlaxius waroona. Upogebiidae. f, Acutigebia simsoni. g, Upogebia tractabilis (pair from sponge). a

b c

e f Plate 13. Palinura Palinuridae.a, Jasus edwardsii novaehollandiae (in typical reef shelter with Rhynchocinetes australis). Scyllaridae. b, Crenarctus crenatus. Anomura Galatheidae. c, Munida gregaria. d, Galathea australiensis. e, Galathea magnifica. f, Munida gregaria. g, Allogalathea elegans. a b

c d Plate 14. Anomura Porcellanidae. a, Petrolisthes elongatus. b, Pisidia dispar. c, Polyonyx transversus. Lomisidae. d, Lomis hirta. b

c e

f g Plate 15. Anomura Diogenidae. a, Cancellus typus (chelipeds protecting hermit crab in rock). b, Clibanarius taeniatus. c, Dardanus arrosor. d, Dardanus pedunculatus. e, Paguristes frontalis. f, g, Paguristes brevirostris (individual and group around feeding welk). a b

c d

e f

g h Plate 16. Anomura Diogenidae. a, Strigopagurus elongatus. b, Strigopagurus strigimanus. Lithodidae. c, Lithodes longispina. Paguridae. d, Lophopagurus nanus. e, Micropagurus acantholepis. f, Pagurixus handrecki. g, Pagurus sinuatus. h, Porcellanopagurus tridentatus. a b

c d

e f

g h Plate 17. Anomura Parapaguridae. a, Paragiopagurus diogenes. b, Parapagurus bouvieri. c, Sympagurus dimorphus. Pylochelidae. d, Trizocheles spinosus. Brachyura Dromiidae. e, Austrodromidia octodentata. f, Dromia wilsoni. g, Dromidiopsis australiensis. h, Haledromia bicavernosa. b

a c

f g Plate 18. Brachyura Dromiidae (cont.). a, b, Lamarckdromia globosa (with large ascidian and sponge). Homolidae. c, Dagnaudus petterdi. Latreilliidae. d, Eplumula australiensis. Raninidae. e, Lyreidus tridentatus. Calappidae. f, Calappa philargius. g, Mursia curtispina. a b

c d

e f Plate 19. Brachyura Leucosiidae. a, Bellidilia laevis. b, Bellidilia undecimspinosa. c, Ebalia dentifrons. d, Ebalia intermedia. e, Merocryptus lambriformis. Matutidae. f, Ashtoret lunaris. a b

c d

e Plate 20. Brachyura Majidae. a, Huenia australis. b, Dumea latipes. c, Pyromaia tuberculata. d, e, Leptomithrax gaimardii (individual and en masse). a b

c d

e f

g h Plate 21. Brachyura Majidae (cont.). a, Naxia aurita. b, Naxia aries. c, Notomithrax minor. d, Notomithrax ursus. e, Schizophrys rufescens. Hymenosomatidae. f, Amarinus laevis. g, Halicarcinus ovatus. h, Trigonoplax longirostris. a b

c d

g f

Plate 22. Brachyura Atelecyclidae. a, Trichopeltarion sp. Cancridae. b, Metacarcinus novaezelandiae. Portunidae. c, Lissocarcinus laevis. d, Carcinus maenas. e, Nectocarcinus tuberculosus. f, g, Liocarcinus corrugatus (individual and mating group). b

c e

f g

h i Plate 23. Brachyura Portunidae (cont.). a, b, Ovalipes australiensis (in vitro and burrowing). c, Ovalipes molleri. d, e, Portunus pelagicus (in defensive posture and burrowing). f, Portunus rubromarginatus. g, Portunus rugosus. h, Portunus sanguinolentus. i, Charybdis natator. a b

c d

f g Plate 24. Brachyura Goneplacidae. a, Carcinoplax meridionalis. b, Carcinoplax victoriensis. c, Litocheira bispinosa. Hexapodidae. d, Hexapinus granuliferus. Pilumnidae. e, Zebrida brevicarinata (on echinoid, Heliocidaris erythrogramma). f, Actumnus setifer. g, Pilumnopeus serratifrons. a b

c d

e f Plate 25. Brachyura Pilumnidae (cont.). a, Pilumnus acer. b, Pilumnus etheridgei. c, Pilumnus monilifer. d, Pilumnus tomentosus. e, Heteropilumnus fimbriatus. f, Ceratoplax luteus. a b

c d

e f

g h Plate 26. Brachyura Xanthidae. a, Lybia tessellata (not a southern Australian species). b, Actaea calculosa. c, d, Actaea peronii (two colour morphs). e, Megametope carinata. f, Megametope rotundifrons. g, Xanthias sp. Pinnotheridae. h, Ostracotheres subglobosus. a

b Plate 27. Brachyura Mictyridae. a, b, Mictyris longicarpus (en masse and on mangrove shore). a b

d e

f g Plate 28. Brachyura Mictyridae (cont.). a, b, Mictyris platycheles (individual and crab with pelletised sediment from burrows). Ocypodidae. c, Ocypode ceratophthalma. d, Heloecius cordiformis. Grapsidae. e, Cyclograpsus audouinii. f, Cyclograpsus granulosus. g, Helograpsus haswellianus. a b

c d

e f

g Plate 29. Brachyura Grapsidae (cont.). a, Paragrapsus gaimardii. b, Paragrapsus laevis. c, Paragrapsus quadridentatus. d, Leptograpsodes octodentatus. e, Parasesarma erythodactyla. f, Brachynotus spinosus. Plagusiidae. g, Plagusia chabrus. a b c

d e f Plate 30. Stomatopoda Bathysquillidae. a, Bathysquilla microps. Hemisquillidae. b, Hemisquilla australiensis. Lysiosquillidae. c, Lysiosquilla colemani. Nannosquillidae. d, Hadrosquilla perpasta. Squillidae. e, Anchisquilloides mcneilli. f, Belosquilla laevis. a b

c d Plate 31. Stomatopoda Squillidae. a, Erugosquilla grahami. b, Harpiosquilla harpax. c, Harpiosquilla melanoura. d, Harpiosquilla sinensis. a b

c d Plate 32. Stomatopoda Squillidae. a, Kempina mikado. b, Oratosquilla oratoria. c, Oratosquillina berentsae. d, Quollastria kapala. 10. ANOMURA – HERMIT CRABS, PORCELAIN CRABS AND SQUAT LOBSTERS

The Anomura McLeay, 1838 are a group of decapods which on first appearance seem to have little in common; some look like crabs, others are hermit crabs, others resemble small lobsters, and others not easily placed in one or other of these well-known crustacean types. The only easy character unit- ing them is the small fifth thoracic leg. To further complicate the issue, animals of any one of these forms cannot easily be placed in one or other of the three superfamilies into which the infraorder is classified. Crab-like forms occur in the Paguroidea, Galatheoidea and Hippoidea. None of the anomuran lobster-like crustaceans have strong abdomens like lobsters but some galatheoids are often called squat lobsters. And shell-carrying hermit crabs, easily recognisable as such, belong in four families. Two boxes are useful for the anomuran groups. One is for six-legged crabs and the other, a quick key to hermits. The key to superfamilies should be used to be sure of an identification. New evidence based on molecular evidence goes some way to explaining the uncertainty surrounding the relationships of the established anomuran families and superfamilies. Pérez- Losada et al. (2002) reviewed the views based on morphology dating from the middle of the nineteenth century and provided a new interpretation based on 18S ribosomal DNA. They confirmed the accepted composition of Galatheoidea (Porcellanidae, Chirostylidae, Galatheidae and Aeglidae) and differed only in the relative position of the families from earlier studies (Martin & Abele, 1986; Tudge, 1997). Pérez-Losada et al. (2002) found that Galatheoidea and Paguroidea more related to each other than to Hippoidea, a view not shared by Morrison et al. (2002) who placed Hippoidea closest to Paguroidea. Clearly, more work will be completed before there is general agreement on higher relationships. It has long been recognised that the larvae of the crab-like lithodids resemble those of the pagurid hermit crabs. Cunningham et al.’s (1992) cladogram based on DNA sequences from the gene encoding mitochondrial large-unit ribosomal RNA placed two genera of Lithodidae within the genus Pagurus, a finding which at least reinforces the polyphyly of this large genus and of the Paguridae. It seems certain that carcinisation, return to a crab-like habitus, has evolved several times in the Anomura. This phenomenon was discussed in detail by McLaughlin & Lemaitre (1997) and McLaughlin et al. (2004) but they disagreed with Cunningham et al.’s conclusions. The name of this group has been debated and sometimes termed Anomala and has variously included several groups of anomalous decapods. McLaughlin & Holthuis (1985) agreed on the widely used name Anomura which is now confined to the three superfamilies, Paguroidea for hermit crabs, Galatheoidea and Hippoidea. They fixed on the authorship and date of the name. From time to time other superfamilies have been included or the families differently arranged. Thalassinoidea, once thought to be anomuran, are here placed in a separate suborder Thalassinidea. The classification used here is based on McLaughlin’s (1983a) phylogenetic reappraisal supported by Martin & Abele’s (1986) analysis. It was followed by Martin & Davis (2001) and McLaughlin (2003) but not by Davie (2002) who recognised two superfamilies for hermit crabs, Paguroidea (including Lithodidae) and Coenobitoidea (including Lomisidae). His classification was supported by Forest (1987 citing unpublished work by M. de Saint Laurent) and McLaughlin & Lemaitre (2001) who all relied on undocumented evidence from the skeleton.

215 Marine Decapod Crustacea of Southern Australia

The enigmatic monotypic family Lomisidae has often been treated as a superfamily (McLaughlin, 1983b; Martin & Davis, 2001) but is almost certainly a highly derived hermit crab, possibly related to Lithodidae on the basis of larval morphology. Its place in the key to superfamilies is quite artificial. Given the present-day confusion, the superfamily Coenobitoidea is not recognised here; its members are placed with the Paguroidea. The key and diagnoses of the superfamilies were derived using characters from McLaughlin (1983a, 1983b, 2003). Davie (2002) has given more extended diagnoses of all the families.

Diagnosis. Epistome protected by sides of carapace. Pereopod 5 reduced and its somite (thoracic somite 8) loosely connected to the preceding one. Uropodal exopod without suture. Innervation of abdominal somite 1 from ganglion attached to thoracic ganglionic mass. Key to superfamilies of Anomura (and Lomis) 1. Abdomen asymmetrical (symmetrical in most Pylochelidae); pereopod 4 reduced, smaller than 2 and 3 (except in Lithodidae); ocular scales present (reduced in Lithodidae) ...... Paguroidea … p. 250 — Abdomen symmetrical; pereopod 4 ambulatory, like 2 and 3; ocular scales absent ...... 2 2. Uropods absent (or invisible in natural view and pleopod-like only in female) ...... Lomis hirta … p. 240 — Uropods present as part of tail-fan and visible in natural position ...... 3 3. Pereopods 1 elongate chelipeds; pereopods 2–4 with claw-like dactyli; telson and uropods forming tail-fan ...... Galatheoidea … p. 219 — Pereopods 1 short, chelate or subchelate; pereopods 2–4 with flattened dactyli; telson longer than wide, not forming tail-fan with uropods ...... Hippoidea … p. 247

References Cunningham, C.W., Blackstone, N.W., & Buss, L.W. 1992. Evolution of king crabs from hermit crab ancestors. Nature 355: 539–542. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda - Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Forest, J. 1987. Les Pylochelidae ou “Pagures symétriques” (Crustacea Coenobitoidea). Mémoires du Muséum National d’Histoire Naturelle, Paris 137: 1–254, 9 pls. Martin, J.W., & Abele, L.G. 1986. Phylogenetic relationships of the genus Aegla (Decapoda: Anomura: Aeglidae), with comments on anomuran phylogeny. Journal of Crustacean Biology 6: 576–616. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. McLaughlin, P.A. 1983a. Hermit crabs - are they really polyphyletic? Journal of Crustacean Biology 3: 608–621. McLaughlin, P.A. 1983b. A review of the phylogenetic position of the Lomidae (Crustacea: Decapoda: Anomala). Journal of Crustacean Biology 3: 431-447. McLaughlin, P.A.2003. Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60: 111–144. McLaughlin, P.A., & Holthuis, L.B. 1985. Anomura versus Anomola. Crustaceana 49: 204–229. McLaughlin, P.A.,& Lemaitre, R. 1997. Carcinization in the Anomura – fact or fiction? I. Evidence from adult morphology. Contributions to Zoology 67: 79–123. McLaughlin, P.A., & Lemaitre, R. 2001. A new family for a new genus and new species of hermit crab of the superfamily Paguroidea (Decapoda: Anomura) and its phylogenetic implications. Journal of Crustacean Biology 21: 1062–1076.

216 Anomura – hermit crabs, porcelain crabs and squat lobsters

McLaughlin, P.A., Lemaitre, R., & Tudge, C.C. 2004. Carcinization in the Anomura – fact or fiction? II. Evidence from larval, megalopal and early juvenile morphology. Contributions to Zoology 73: 165–205. Morrison, C.L., Harvey, A.W., Lavery, S., Tieu, K., Huang, Y., & Cunningham, C.W. 2002. Mitochondrial gene rearrangements confirm the parallel evolution of the crab-like form. Proceedings of the Royal Society of London B 269: 345–350. Pérez-Losada, M., Jara, C.G., Bond-Buckup, G., Porter, M.L., & Crandall, K.A. 2002. Phylogenetic position of the freshwater anomuran family Aeglidae. Journal of Crustacean Biology 22: 661–669. Tudge, C.C. 1997. Phylogeny of the Anomura (Decapoda, Crustacea): Spermatozoa and spermatophore morphological evidence. Contributions to Zoology 67: 125–141.

QUICK KEY TO HERMITS Hermit crabs (Paguroidea) are not the only crustaceans which use shells as homes. Some tanaidaceans do too and look like hermits crabs until they are removed from the shell. Of course, tanaidaceans are tiny, in shells never more than about 5 mm long. They are not decapods so are not covered in this book. Some amphipods carry fragments of bivalve shells as temporary homes and others live commensally in the deeper cavities of gastropods with hermit crabs. Some crabs carry bivalves. A first step is to check that your animal is a hermit crab and then to determine which of the two common families it belongs to. Deep-water species, those from depths greater than 200 m, may belong to other families not in these keys. 1. Thorax segmented dorsally, without carapace; with chelipeds plus 6 pairs of walking legs ...... not a hermit crab – Tanaidacea: Pagurapseudidae — Thorax covered dorsally by carapace shield; with chelipeds, 3 pairs of walking legs, fifth legs reduced ...... Paguroidea … 2 2. Chelipeds of distinctly different size, right always the larger; maxillipeds 3 widely spaced at base ...... Paguridae — Chelipeds equal or subequal in size, or left distinctly larger; maxillipeds adjacent at base ...... Diogenidae PAGURIDAE (RIGHT-HANDED HERMIT CRABS) Large (5 cm overall) right-handed hermit crabs from shallow water usually belong to the genus Pagurus but it appears that most species in southern Australia are undescribed. Small right-handed hermit crabs are more tricky and a microscope is needed. 1. Ocular scale multispinose and with a pair of small lobes between ...... Micropagurus acantholepis — Ocular scale simple, without lobes between ...... 2 2. Small left chela with dorsal ridge; subtidal; south-east only . . . . . Lophopagurus nanus — Small left chela smoothly rounded dorsally; some species common intertidally ...... Pagurixus (several species, some geographically restricted) DIOGENIDAE (LEFT- OR EQUAL-HANDED HERMIT CRABS) The technical key can be difficult without a microscope and some knowledge of hermit crab anatomy. Here is a rough key which eliminates unusual species first. It is necessary to check the species in the main text. 1. Living in a stone ...... Cancellus typus (and another rare species in WA) — Living in a shell ...... 2

217 Marine Decapod Crustacea of Southern Australia

2. Insides of chelae with wash-board-like ridges (presumably to rub together for sound) ...... Trizopagurus strigimanus (large, common shelf species; one other rarer species) — Chelae variously ornamented but never with regular ridges ...... 3 3. Tips of fingers white, calcareous ...... Calcinus spp. — Tips of fingers black, corneous ...... 4 4. Small; female with dorsal brood-pouch; sometimes intertidal species ...... Paguristes spp. (P. frontalis and P. squamosus the only intertidal species) — Usually large, subtidal species ...... 5 5. Rostrum hinged; drab ...... Diogenes spp. (in NSW, D. senex, intertidal; others in WA, subtidal) — Rostrum fixed; colourful ...... 6 6. Shield with a Y-shaped groove; chelipeds unequal; subtidal ...... Dardanus spp. — Shield without Y-shaped groove; chelipeds equal ...... Clibanarius spp.

SIX-LEGGED CRABS Most, but not all, crabs with only three pairs of walking legs are anomurans, or not true crabs. Some deep-water crabs appear to have only three pairs of legs. Lithodids, for example, are anomuran crabs. But members of brachyurans families, Homolidae and the like, and the rare Dynomenidae which may occur in shallow water have the fifth legs reduced or held erect and only slightly smaller than the others. On the shore and subtidally, small crab-like forms are commonly encountered in which the fifth leg is invisible or tiny. They could belong to one of several families separated as follows. 1. Abdomen tightly held under carapace, especially in male; chela roughly cylindrical ...... Brachyura: Hexapodidae … 2 — Abdomen loosely held under carapace; chela flattened ...... Anomura … 3 2. Sternum in male without grooves at end of abdomen ...... Hexapinus granuliferus — Sternum in male with broad deep oblique grooves running between bases of maxillipeds 3 and end of abdomen ...... Hexapus sexpes 3. Digging in sand ...... 4 — Under rocks or subtidal ...... 5 4. Carapace flat, square ...... Austrolepidopa trigonops (Albuneidae) — Carapace oval ...... Hippa spp. (Hippidae) 5. Hairy; limbs compact; with bright blue antennae ...... Lomis hirta (Lomisidae) — Smooth: limbs, especially chelae elongate; antennae with long filtering setae ...... Porcellanidae … 6 6. Subtidal, muddy sediments ...... Polyonyx transversus — Intertidal, rocky shores and subtidal ...... 7 7. Chelae spiky ...... Petrocheles australiensis — Chelae smooth ...... Petrolisthes elongatus (Tas. only) or Pidisia dispar (WA only)

218 Anomura – hermit crabs, porcelain crabs and squat lobsters

Superfamily Galatheoidea Samouelle, 1819 The Galatheoidea comprise four families of unusual decapods, some crab-like and others more lobster-like. They share with all anomurans only four pairs of well-developed legs of which the first is a stronger cheliped. The fifth leg is weak and often hidden under the carapace. Galatheoids are symmetrical (unlike hermit crabs) and the uropods and telson form a broad tailfan at the end of the depressed abdomen which folds up against itself or the thorax. There are four families. One, Aeglidae Dana, 1852, is confined to freshwater environments in South America. The other three are marine and occur in Australia. Martin & Abele’s (1986) study of this family confirmed its placement in this superfamily. The Galatheidae (squat lobsters) and Porcellanidae (false crabs or porcelain crabs) are common in southern Australia and the Chirostylidae (also squat lobsters) are most common in deep water. A guide to the most common shallow shelf taxa can be found in the box ‘Miniature lobsters with long chelae’.

Diagnosis. Body symmetrical. Uropods spatulate, forming tail-fan. Pleopods 3–5 paired. Male gonopods often present. Female pleopods 1 absent. Ocular scales absent. 4–5 antennal peduncle articles. Epistome unarmed. 11–14 pairs of gills. Pereopods 2–5 dactyli with claw, 4 ambulatory.

Key to Australian families of Galatheoidea 1. Abdomen folded up against thorax; body crab-like; chelipeds moderately elongate, stout or flattened ...... Porcellanidae ...p.242 — Abdomen folded up against itself (not against thorax); body shrimp-like; chelipeds greatly elongate, slender ...... 2 2. Telson with a transverse suture (sometimes difficult to see) and folded beneath the preceding abdominal somites with the tailfan; without sternal plate on last thoracic somite (no plate between the last legs); usually with antennal scale ...... Chirostylidae ...p.220 — Telson without a transverse suture and not folded beneath the preceding abdominal somites with the tailfan; with well developed sternal plate on last thoracic somite between last legs; without antennal scale ...... Galatheidae ...p.228

Reference Martin, J.W., & Abele, L.G. 1986. Phylogenetic relationships of the genus Aegla (Decapoda: Anomura: Aeglidae), with comments on anomuran phylogeny. Journal of Crustacean Biology 6: 576–616.

219 Marine Decapod Crustacea of Southern Australia

MINIATURE LOBSTERS WITH LONG CHELAE No members of the Galatheidae are found intertidally in Australia but galatheids are among the most common of all decapods on shelf sediments. Large lobster-like animals with long chelae are found only in the deep sea. These are members of the Polychelidae (Polychelida), quite a different group from those in shallower water, usually squat lobsters (Superfamily Galatheoidea). Galatheoids are easy to recognise but the species can be difficult to distinguish, especially in the tropics. This is a rough guide to the commonest species of the shallow shelf. 1. Rostrum of 3 spines ...... Munida spp. (M. haswelli is the most common species) — Rostrum triangular ...... 2 2. Rostrum extremely elongate ...... Allogalathea elegans (WA only) — Rostrum only a bit elongate ...... 3 3. Rostrum with lateral teeth ...... Galathea spp. (G. australiensis is the most common) — Rostrum leaf-like ...... Phylladiorhynchus pusillus The family Chirostylidae and the galatheid genus Munidopsis occur only at slope depths and beyond.

Chirostylidae Ortmann, 1892 Chirostylids look superficially much like galatheids but generally occur in deeper shelf waters and beyond. These squat lobsters are often collected associated with soft corals, antipatharians and gorgonians. The fifth legs attaching free, and without a sternal plate between, distinguishes them. The definitive work on the systematics of the Chirostylidae was by Baba (1988) who recognised five genera and more than 100 species worldwide. There are many described species particularly in the Indo-West Pacific region which is the centre of richest diversity. The fauna of southern Australian slopes and seamounts is better known than that of tropical seas although some of the species reported by Baba (1991 and in paper in press in Galathea Reports) might be expected also. Numerous species of four genera have been reported (Poore et al., 1998; Ahyong & Poore, 2004). The key to these genera and to a fifth from Hawaii is from Baba (1988). Diagnosis. Body shrimp-like. Telson with a transverse suture (sometimes difficult to see) and folded beneath the preceding abdominal somites with the tail-fan. Chelipeds elongate, slender. Without sternal plate on last thoracic somite. Usually with antennal scale.

Key to genera of Chirostylidae 1. 1 or 2 pairs of supraocular spines on margin of carapace; mandible smooth or feebly dentate on incisor margin ...... 2 — No supraocular spines (may be spines behind eyes on carapace); mandible strongly dentate on incisor margin ...... 3 2. 2 pairs of supraocular spines; 3 pairs of hepatic spines in oblique line ...... Eumunida — 1 pair of supraocular spines; hepatic spines absent ...... Pseudomunida (Hawaii only) 3. Pereopods usually spinose and slender ...... 4 — Pereopods not extremely spinose, short or of moderate length ...... Uroptychus 4. Rostrum absent ...... Chirostylus (not in Australia) — Rostrum present ...... Gastroptychus

220 Anomura – hermit crabs, porcelain crabs and squat lobsters

Eumunida Smith, 1883 The genus of mostly deep-water squat lobsters is recognised by the combination of five rostral and supraocular spines and oblique rows of three spines just posterior to these. Twenty-four species in the Indo-West Pacific are known (de Saint Laurent & Macpherson, 1990a, b; de Saint Laurent & Poupin, 1996), including at least two from tropical Australia. Diagnosis. Rostrum present. 2 pairs of supraocular spines. 3 pairs of hepatic spines in oblique rows. Mandible smooth or weakly dentate on incisor margin. Pereopods with spinose merus and carpus. Eumunida australis de Saint Laurent & Macpherson, 1990 (Fig. 60a). Carapace with 2 pairs of anterolateral spines, 4 pairs of lateral spines, first hepatic spine stronger than following 2; cheliped merus with 2 longitudinal rows of spines. 25 mm. Qld (off Southport), NSW (to 35°S); continental slope, 300–685 m depth.

Gastroptychus Caullery, 1896 Baba (1988, 1991) stated that there are 17 described species and later described the first from southern Australia (Baba, 2000). Ahyong & Poore’s (2004) key to species of the Indo-West Pacific included 13 species. Most species have a spinose carapace, the Australian species especially so. Diagnosis. Rostrum present. No supraocular spines. Mandible strongly dentate on incisor margin. Pereopods spinose, slender.

Key to southern Australian species of Gastroptychus 1. Abdominal somites 2–5 without transverse rows of spines . . . . Gastroptychus sternoornatus — Abdominal somites 2–5 with transverse rows of spines ...... 2 2. Abdominal somite 3 without spines except on pleura; maxilliped 3 without spines on propodus and carpus ...... Gastroptychus hendersoni — Abdominal somite 3 with spines all over; maxilliped 3 with spines on propodus and carpus ...... Gastroptychus rogeri Gastroptychus hendersoni (Alcock & Anderson, 1899). Carapace with spines over whole surface, especially prominent medially; abdominal somite 3 without spines, except small spines on pleura. 37 mm. Indo-West Pacific, Tas.; 787–1469 m depth. Gastroptychus rogeri Baba, 2000 (Fig. 60b). Carapace with spines over whole surface, especially prominent medially; all abdominal somites spinose. 28 mm. NSW, Tas.; 476–1000 m depth. Gastroptychus sternoornatus (Van Dam, 1933). Carapace with spines over whole surface, especially prominent behind eyes, medially on gastric region and near posterior margin; all abdominal somites unarmed. Indo-West Pacific, Tas.; 787–1469 m depth.

Uroptychus Henderson, 1888 This is by far the most common genus of Chirostylidae in southern Australian collections from deep water, Twenty new species were described by Ahyong & Poore (2004) from south-eastern and southern Australia of which 14, plus three others, occur in our region. Carapace length does not include the rostrum. Diagnosis. Rostrum present. No supraocular spines. Mandible strongly dentate on incisor margin. Pereopods not extremely spinose, short or of moderate length.

221 Marine Decapod Crustacea of Southern Australia

Fig. 60. Chirostylidae. a, Eumunida australis. b, Gastroptychus rogeri. c, Uroptychus australis. d, Uroptychus babai. e, Uroptychus calcar. f, Uroptychus litosus.

Key to southern Australian species of Uroptychus 1. Carapace with lateral margins smooth or crenulate ...... 2 — Carapace with lateral margins dentate or spinose ...... 11 2. Carapace with lateral margins smooth or crenulate ...... 3 — Carapace (excluding rostrum) length equal to or greater than width ...... 8 3. Walking legs with dactyli having 2 distal spines ...... Uroptychus pilosus — Walking legs with dactyli having more than 8 spines or denticles along lower margin . . . . 4 4. Antennal peduncle having last and second-last articles each with distal spine ...... 5 — Antennal peduncle having last and second-last articles each without distal spine ...... 6

222 Anomura – hermit crabs, porcelain crabs and squat lobsters

5. Antennal peduncle having basal article with outer spine ...... Uroptychus latus — Antennal peduncle having basal article without outer spine ...... Uroptychus laperousazi 6. Antennal scale overreaching apex of distal peduncle article; basal antennal artcles with small outer spine; outer orbital angle acute ...... U. babai — Antennal scale reaching to about midlength of distal peduncle article; basal antennal article without outer spine; outer orbital angle rounded ...... 7 7. Cheliped about 3 times carapace length; dorsal margin of propodal palm about 2.5 times as long as dactylus; teeth lining flexor margins of dactyli of walking legs slender and longer than wide ...... Uroptychus longvae — Cheliped about twice carapace length; dorsal margin of propodal palm about 2.5 times as long as dactylus; teeth lining flexor margin of dactyli of walking legs short, about as long as wide ...... Uroptychus patulus 8. Walking legs dactyli having spines along inner margin oriented parallel to margin ...... Uroptychus australis — Walking legs dactyli having spines along inner margin oriented oblique to margin ...... 9 9. Chelipeds robust, about 3 times carapace length; carapace margins divergent ...... Uroptychus litosus — Chelipeds slender, about 3.5–4 times carapace length; carapace margins subparallel . . . . 10 10. Walking leg 3 merus about half length of merus of walking leg 2; sternite 4 with field of granules ...... Uroptychus empheres — Walking leg 3 merus about two-thirds length of merus of walking leg 2; sternite 4 smooth ...... Uroptychus gracilimanus 11. Carapace lateral margin with single strong spine above base of walking leg 1 ...... Uroptychus raymondi — Carapace lateral margin with more than 1spine or tooth ...... 12 12. Walking leg 1 markedly more slender than walking leg 2 ...... Uroptychus nowra — Walking leg 1 similar to walking leg 2 ...... 13 13. Antennal peduncle with second last article unarmed distally, about quarter length of last ...... Uroptychus flindersi — Antennal peduncle with second last article with distal spine, half or more as long as last .14 14. Antennal scale reaching to midlength of last peduncle article; second last article with distal spine; last article unarmed ...... Uroptychus calcar — Antennal scale reaching to midlength of last peduncle article; second last article with distal spine; last article unarmed ...... 15 15. Carapace with lateral margins denticulate or with small spines ...... 16 — Carapace with lateral margins with strong spines ...... 17 16. Outer orbital angle rounded; carapace setose ...... Uroptychus hesperius — Outer orbital angle spiniform; carapace not setose ...... Uroptychus subsolanus 17. Walking legs dactyli with 8–10 strong spines on inner margin; carapace smooth except for epigastric spines ...... Uroptychus zeidleri — Walking legs dactyli with 16–20 small spines on inner margin; carapace rugose, setose ...... Uroptychus cardus

223 Marine Decapod Crustacea of Southern Australia

b c

e d

g f h

Fig. 61. Chirostylidae. a, Uroptychus cardus. b, Uroptychus empheres. c, Uroptychus flindersi. d, Uroptychus gracilimanus. e, Uroptychus hesperius. f, Uroptychus laperousazi. g, Uroptychus latus. h, Uroptychus nowra.

Uroptychus australis (Henderson, 1885) (Figs 60c, 62f). Carapace longer than broad; lateral margins unarmed. Antennal basal article with distinct outer spine; ultimate and penultimate articles unarmed. Chelipeds about 3.5–4 times carapace length; merus usually with cluster of tubercles on inner proximal margin. Pereopods 2–4 dactyli with 7–9 small spines on inner margin, oriented parallel to dactylar margin. Northern New Zealand, Indonesia, NSW, Vic., Tas., Tasmanian Seamounts; 458–1800 m depth, the most common species of the genus in south-eastern Australia.

224 Anomura – hermit crabs, porcelain crabs and squat lobsters

Uroptychus babai Ahyong & Poore, 2004 (Figs 60d, 62i). Carapace slightly broader than long; lateral margins irregular, crenulate, distinctly convex, broadest posterior to midlength; with distinct, anteriorly directed anterolateral spine. Basal antennal article with distinct outer spine; ultimate and penultimate articles unarmed; scale longer than peduncle. NSW; 880–920 m depth. Uroptychus calcar Ahyong & Poore, 2004 (Fig. 60e). Carapace longer than broad; lateral margins with anterolateral spine and 7–12 lateral spines; dorsum smooth, with cluster of 3–4 small epigastric spines on each side. Antennal basal article with outer spine; penultimate article with distal spine. Chelipeds about 3 times carapace length; all articles glabrous. NSW, Vic.; 326–461 m depth. Uroptychus cardus Ahyong & Poore, 2004 (Fig. 61a). Carapace longer than broad; lateral margins with anterolateral spine and 7–8 lateral spines; dorsum rugose. Antennal basal article with outer spine; ultimate and penultimate articles with distal spine. Chelipeds about 3.5 times carapace length; all articles acutely rugose and sparsely setose; inner margin of merus 2 rows of stout spines. Tasmanian Seamounts; 987–1200 m depth. Uroptychus empheres Ahyong & Poore, 2004 (Fig. 61b). Carapace longer than broad; margins slightly divergent, irregularly tuberculate behind base of cervical groove; without posterolateral ridge; anterolateral spine small. Outer orbital spine extending to level of anterolateral spine. Basal antennal article with lateral spine; ultimate and penultimate articles unarmed; ultimate article slightly exceeding twice length of penultimate article. Cheliped 3.5–4 times carapace length; distal 3 articles glabrous on upper outer margin; inner and ventral margin of propodal palm, carpus and merus granular. Tasmanian Seamounts; 800 m depth. Uroptychus flindersi Ahyong & Poore, 2004 (Fig. 61c). Carapace distinctly longer than broad; lateral margins with anterolateral spine and distinct, bifid spine at base of cervical groove followed by 7–8 small teeth or serrations. Antennal basal article with acute tooth on outer margin; ultimate peduncle article with small distal spine; penultimate article unarmed. Chelipeds about 3.5 times carapace length; articles rugose ventrally, with distinct scales. Tas., WA; 520–714 m depth. Uroptychus gracilimanus (Henderson, 1885) (Fig. 61d). Carapace longer than broad; lateral margins subparallel or slightly divergent, unarmed; with small anterolateral spine; posterior quarter with low but distinct ridge; outer orbital angle produced to small spine. Antennal basal article with distinct outer spine; ultimate and penultimate articles unarmed. Chelipeds 3.5–4 times carapace length; merus with tubercles on inner proximal margin. Indo-West Pacific, 421–1668 m; NSW (off Port Jackson); 750 m depth. Uroptychus hesperius Ahyong & Poore, 2004 (Fig. 61e). Carapace broader than long; lateral margins serrated, convex, divergent, broadest posterior to midlength; with strong anteriorly directed anterolateral spine; outer orbital angle rounded; dorsum finely setose. Ultimate and penultimate articles of antennal peduncle with distal spine. Cheliped slender, about twice carapace length; all articles rugose and sparsely setose; propodal palm about 3 times as long as fixed finger. Southern WA; 1011–1020 m depth. Uroptychus laperousazi Ahyong & Poore, 2004 (Figs 61f, 62k). Carapace distinctly broader than long; lateral margins slightly crenulate, distinctly convex, broadest posterior to midlength; with short, anteriorly directed anterolateral spine; posterior fifth with low, indistinct ridge. Outer orbital angle produced to small tooth, not extending anteriorly beyond anterolateral spine. Basal antennal article without outer spine; ultimate and penultimate articles armed. Cheliped about 2 times carapace length. SA; 984–1110 m depth. Uroptychus latus Ahyong & Poore, 2004 (Figs 61g, 62j). Carapace distinctly broader than long; lateral margins smooth or slightly irregular, distinctly convex, broadest posterior to midlength; with short, anteriorly directed anterolateral spine; posterior fifth with low, indistinct ridge; outer orbital angle produced to small tooth, not extending anteriorly beyond

225 Marine Decapod Crustacea of Southern Australia

anterolateral spine; dorsum minutely punctate, finely but sparsely-setose, unarmed. Basal antennal article with outer spine; ultimate and penultimate articles armed; antennal scale extending almost to apex of ultimate peduncle article. Chelipeds about 3 times carapace length. Vic; 1073 m depth. Uroptychus litosus Ahyong & Poore, 2004 (Figs 60f, 62j). Carapace as long as broad; lateral margins distinctly convex, with irregular tubercles along margin; with anterolateral spine; outer orbital angle produced to small spine. Basal antennal article with distinct outer spine; ultimate and penultimate articles unarmed; ultimate article about twice length of penultimate article. Chelipeds robust, slightly compressed, about 3 times carapace length; merus and ischium with distinct tubercles on inner proximal margin. Tasmanian Seamounts; 800–1120 m depth. Uroptychus longvae Ahyong & Poore, 2004. Carapace as broad as long (similar to U. patulus); lateral margins smooth, distinctly convex, broadest posterior to midlength; with triangular anterolateral spine; outer orbital angle sharp; dorsum finely but sparsely-setose, unarmed. Basal antennal article without outer spine; ultimate and penultimate articles unarmed. Antennal scale extending about to third of ultimate peduncle article. Chelipeds about 3 times carapace length. Teeth on dactyli of walking legs slender and longer than wide. SA (Great Australian Bight); 805–816 m depth. Uroptychus nowra Ahyong & Poore, 2004 (Fig. 61h). Carapace broader than long; lateral margins convex, divergent; with strong anteriorly directed anterolateral spine and 9 lateral spines; anterior 2 lateral spines small, third spine large, stout, at base of indistinct cervical groove, followed by 2 small spines and 4 large spines. Outer orbital angle produced to a small spine. Basal antennal article with inner and outer spine; ultimate article of antennal peduncle about twice as long as penultimate article, both with distal spine. Cheliped slender, all articles rugose and sparsely setose. NSW; 1100 m depth. Uroptychus patulus Ahyong & Poore, 2004 (Fig. 62a). Carapace distinctly broader than long; lateral margins smooth, distinctly convex, broadest posterior to midlength; with distinct, slightly incurved anterolateral spine; posterior fifth with low, indistinct ridge; outer orbital angle rounded, unarmed; dorsum minutely punctate, finely but sparsely-setose, unarmed. Basal antennal article without outer spine; ultimate and penultimate articles unarmed. Antennal scale extending about to midlength of ultimate peduncle article. Chelipeds about 2.5 times carapace length. Teeth on dactyli of walking legs short, about as long as wide. Vic., Tas.; 955–1190 m depth. Uroptychus pilosus Baba, 1981 (Fig. 62b). Body entirely covered with fine setae. Carapace slightly wider than long, widest near midlength; with small anterolateral spinules; dorsum and lateral margins unarmed, pilose. Basal antennal article with outer spine; antennal scale as long as or slightly longer than penultimate peduncle article. Chelipeds slender, subcylindrical; 4–5 times carapace length; setose. Japan, NSW; 987–1160 m depth. Uroptychus raymondi Baba, 2000. (Fig. 62c). Carapace slightly broader than long; lateral margins convex, irregularly tuberculate, with strong midlateral spine; anterolateral spine stout; outer orbital angle triangular; dorsum setose and sparsely tuberculate. Basal antennal article with lateral tooth; ultimate and penultimate articles with distal tubercle. Antennal scale extending beyond midlength of ultimate peduncle article. Chelipeds setose, not more than 5 times carapace length, carpus and merus distally dentate. Vic., Tas.; 644–650 m depth. Uroptychus subsolanus Ahyong & Poore, 2004 (Fig. 62d). Carapace broader than long; lateral margins serrated, convex, divergent, broadest posterior to midlength; with strong anteriorly directed anterolateral spine and distinct spine at base of cervical groove; outer orbital angle produced to a small spine not reaching beyond midlength of anterolateral spine; dorsum minutely punctate, without setae; lateral margins sparsely setose. Ultimate and penultimate article of antennal peduncle with distal spine. Cheliped slender, about twice carapace length;

226 Anomura – hermit crabs, porcelain crabs and squat lobsters

propodal palm about 2 times as long as fixed finger. Pereopods with propodus broadened distally, bearing 6–8 moveable spines on lower distal margin, distalmost paired; dactylus with 7–8 triangular, corneous, obliquely directed spines on inner margin. Vic., SA; 999–1110 m depth. Uroptychus zeidleri Ahyong & Poore, 2004 (Fig. 62e). Carapace broader than long; lateral margins divergent; with anterolateral spine and 11–15 lateral spines; anterior 2 lateral spines small, third spine large, stout, at base of indistinct cervical groove, remainder stout, closely spaced, decreasing in size posteriorly; outer orbital angle produced to small acute spine. Basal antennal article with 1 or 2 small outer spines; ultimate and penultimate articles with distal spine. Chelipeds about 3 times carapace length; propodus with scales; carpus and merus with setose scales or small spinules, distal margin spinose. Tas.; 520 m depth.

c i a b

j f

d e g k

Fig. 62. Chirostylidae. a, Uroptychus patulus. b, Uroptychus pilosus. c, Uroptychus raymondi. d, Uroptychus subsolanus. e, Uroptychus zeidleri. Distal articles of walking leg: f, Uroptychus australis. g, Uroptychus litosus. h, Uroptychus subsolanus. Antennal scale and peduncle: i, Uroptychus baba. j, Uroptychus laperousazi. k, Uroptychus latus.

References Ahyong, S.T., & Poore, G.C.B. 2004. The Chirostylidae of southern Australia (Crustacea: Decapoda: Anomura). Zootaxa 436: 1–88. Baba, K. 1988. Chirostylid and galatheid crustaceans (Decapoda: Anomura) of the ‘Albatross’ Philippine Expedition, 1907–1910. Researches on Crustacea, Special Number 2: 1–203.

227 Marine Decapod Crustacea of Southern Australia

Baba, K. 1991. Crustacea Decapoda: Chirostylus Ortmann, 1892, and Gastroptychus Caullery, 1896 (Chirostylidae) from New Caledonia. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152: 463–477. Baba, K. 2000. Two new species of chirostylids (Decapoda: Anomura: Chirostylidae) from Tasmania. Journal of Crustacean Biology 20 (Special Number 2): 246–252. Poore, G.C.B., Hart, S., Taylor, J., & Tudge, C. 1998. Decapod crustaceans from Tasmanian seamounts. Pp. 65–78 in: Koslow, J.A., & Gowlett-Holmes, K. (eds), The seamount fauna of southern Tasmania: benthic communities, their conservation and impacts of trawling. Final report to Environment Australia and The Fisheries Research Development Corporation. CSIRO Marine Research: Hobart. de Saint Laurent, M., & Macpherson, E. 1990a. Crustacea Decapoda: Le genre Eumunida Smith, 1883 (Chirostylidae) dans les eaux néo-calédoniennes. In: Crosnier, A. (ed.), Résultats des Campagnes Musorstom, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145: 227–288. de Saint Laurent, M., & Macpherson, E. 1990b. Les espéces atlantiques du genre Eumunida Smith, 1883 (Crustacea: Decapoda: Chirostylidae). Journal of Natural History 24: 647–666. de Saint Laurent, M., & Poupin, J. 1996. Crustacea, Anomura: les espèces indo-ouest pacifiques du genre Eumunida Smith, 1880 (Chirostylidae) Description de six espèces nouvelles. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 15. Mémoires du Muséum National d’Histoire Naturelle, Paris 168: 337–385.

Galatheidae Samouelle, 1819 Galatheids are commonly called squat lobsters although the names, plated lobsters, lobster krill and craylets, have been used. All the species are benthic on rough sedimentary bottoms at shelf and greater depths. The family is one of the most diverse of all decapods and is one of the most obvious in photographs taken of hydrothermal vents at abyssal depths. In southern New Zealand, as in other places, some species of galatheids swarm in astronomical numbers forming ‘red tides’ near the coast. This phenomenon has not been reported in Australia but several species are very common in dredge samples from the shelf. Baba (1988) reviewed the systematics of the Galatheidae and provided a key to the 16 genera known. Nine genera have been added since then by Baba (1991; 1993; 1996) and Macpherson & Machordom (2000). Five occur in both the Atlantic and Indo-West Pacific and the remainder are confined to the Indo-West Pacific. Hundreds of species are known, including many undescribed from the tropical South Pacific (see Macpherson, 2004). At least ten genera are present in Australia, some represented by undescribed species. The Australian fauna is richest in northern waters (Haig, 1974; Baba, 1988) but numerous species have been recorded from the southern coast. Six species were recorded from the Tasmanian Seamounts (Poore et al., 1998) and recently described (Ahyong & Poore, 2004). Genera and species are distinguished by the shape of the rostrum and the arrangement of spines on the carapace and abdominal somites. Note that abdominal somite 1 is easily overlooked when the animal is viewed from dorsal view. The spines on the end of both the antennule and antenna are sometimes important.

Diagnosis. Body shrimp-like. Telson without a transverse suture and not folded beneath the preceding abdominal somites with the tail-fan. Chelipeds elongate, slender.

Key to southern Australian genera of Galatheidae 1. Eyes usually reduced, not pigmented; maxilliped 1 exopod without flagellum ...... Munidopsis … p. 235 — Eyes usually well-developed, pigmented; maxilliped 1 exopod with flagellum ...... 2 2. Rostrum spiniform, distinct from supraorbital spines (if triangular not tapering from orbit) 5

228 Anomura – hermit crabs, porcelain crabs and squat lobsters

— Rostrum triangular, evenly tapering from orbit and including supraorbital spines ...... 3 3. Rostrum extremely elongate, with 5–9 lateral teeth ...... Allogalathea ...p.231 — Rostrum moderately elongate, with 2–5 lateral teeth ...... 4 4. Rostrum leaflet-like with tiny distolateral and distinct basilateral teeth ...... Phylladiorhynchus ...p.238 — Rostrum with 4 distinct lateral teeth ...... Galathea ...p.231 5. Pereopods 1–3 with epipod; frontal margin of carapace with spine mesiad to anterolateral spine ...... Raymunida … p. 239 — Pereopods 1–3 without epipod. Frontal margin of carapace without spine mesiad to anterolateral spine ...... 6 6. Carapace with row of spines across epigastric region, sometimes pair more developed; sometimes with pair of postcervical spines; male gonopod 1 present . . . . Munida ...p.232 — Carapace with pair of epigastric spines behind supraocular spines and with pair of postcervical spines; male gonopod absent ...... 7 7. Carapace with clear transverse ridges; dactyli of walking legs compressed laterally, without lateral ridge ...... Agononida ...p.229 — Carapace without clear transverse ridges, with scales or short ridges; dactyli of walking legs depressed vertically, with lateral and mesial ridges ...... Paramunida ...p.238

Agononida Baba & de Saint Laurent, 1996 Species of Agononida belong to a group of deep-water genera without a second male gonopod. They can be separated from the more abundant Munida by fewer carapace spines, dominated by the pairs of epigastric and postcervical spines sometimes other pairs in between. Ahyong & Poore (2004) discussed variability in Australian species. Macpherson (2004) listed many species from Tonga and Fiji. Diagnosis. Eyes well-developed. Rostrum spiniform, well-developed, remote from supraocular spines. Carapace with distinct transverse ridges, with pair of epigastric spines behind supraocular spines, and with pair of postcervical spines. Pereopods 1–3 without epipod; dactyli of walking legs compressed laterally, without lateral ridge. Frontal margin of carapace without spine mesiad to anterolateral spine. Male gonopod 1 absent.

Key to southern Australian species of Agononida 1. Antenna with basal article produced as long slender spine beyond eyes ...... 2 — Antenna with basal article produced as short triangular spine, not beyond eyes ...... 4 2. Carapace without cardiac spines ...... Agononida incerta — Carapace with cardiac spines ...... 3 3. Carapace with median mesogastric spines ...... Agononida eminens — Carapace without median mesogastric spines ...... Agononida marini 4. Carapace with pair of protogastric spines behind pair of epigastric spines; with 2–4 cardiac spines ...... Agononida procera — Carapace without protogastric spines; with single median gastric spine . Agononida squamosa Agononida eminens (Baba, 1988) (Fig. 63a). Carapace with pair of epigastric spines; without hepatic spine; with 1 or 2 medial cardiac spines; with pair of postcervical spines followed by line of 2 branchiocardiac spines each side; posterior margin with 2–4 antrorse spines; lateral

229 Marine Decapod Crustacea of Southern Australia

margin of carapace anterior to cervical groove with 2 spines (including anterolateral) and 4 spines posterior to cervical groove. 29 mm. Philippines, Indonesia, New Caledonia, Qld, NSW; 564–1051 m depth. Agononida incerta (Henderson, 1888). Carapace with pair of epigastric spines; without protogastric, hepatic and cardiac spines; with pair of postcervical spines followed by line of 2 branchiocardiac spines each side; posterior margin without spines; lateral margin of carapace anterior to cervical groove with 2 spines (including anterolateral) and 4 spines posterior to cervical groove. 34 mm. Indo-West Pacific, WA, Qld, NSW; 17–720 m depth. Two forms of this species differ in colour and spination (Ahyong & Poore, 2004).

a b c d

h g

Fig. 63. Galatheidae. a, Agononida eminens. b, Agononida marini. c, Agononida procera. d, Agononida squamosa. e, Allogalathea elegans. f, Galathea australiensis. g, Galathea magnifica. h, Galathea whiteleggii.

230 Anomura – hermit crabs, porcelain crabs and squat lobsters

Agononida marini Macpherson, 1994 (Fig. 63b). Carapace with pair of epigastric spines, without protogastric and hepatic spines; with 3 median cardiac spines; with pair of postcervical spines followed by 2 branchiocardiac spines each side; posterior margin with 1 antrorse spines; lateral margin of carapace anterior to cervical groove with 4 spines (including long anterolateral) and 1 spine posterior to cervical groove. 27 mm. New Caledonia, Qld, NSW; 457–548 m depth. Agononida procera Ahyong & Poore, 2004 (Fig. 63c). Carapace with pairs of epigastric and protogastric spines; small hepatic spine; with 1 or 2 pairs of cardiac spines; with pair of postcervical spines with line of 2–3 branchiocardiac spines each side; posterior margin with 4–13 antrorse spines; lateral margin of carapace anterior to cervical groove with 2 spines (including anterolateral) and 4 spines posterior to cervical groove. 23 mm. Qld, NSW; 675–824 m depth. Agononida squamosa (Henderson, 1885) (Fig. 63d). Carapace with pair of epigastric spines; without protogastric and hepatic spines; with 1 median cardiac spine; with pair of postcervical spines; posterior margin with 2 antrorse spines; lateral margin of carapace anterior to cervical groove with 2 spines (including anterolateral) and 3 or 4 spines posterior to cervical groove. 30 mm. South-West Pacific, NSW; 156–752 m depth.

Allogalathea Baba, 1969 The genus is monotypic. Diagnosis. Eyes well-developed. Maxilliped 1 exopod with flagellum. Rostrum triangular, elongate, with 5–9 lateral teeth. Allogalathea elegans (Adams & White, 1848) (Fig. 63e, Pl. 13f). The elongate rostrum and longitudinal dark stripes immediately characterise this species. It is believed to associate with comatulid crinoids (Baba, 1979). There are various colour forms but the specimen in Jones & Morgan’s (2002) photograph has a yellow median stripe and yellow legs to match its host. 22 mm. Red Sea, E coast of Africa to Japan and Fiji, Qld, NT, WA (rarely S to Cockburn Sound); intertidal to 146 m. Galathea Fabricius, 1793 Haig (1972) listed eight species of Galathea from north-western Australia and later (1973) provided a key to all nine from eastern Australia; recent collecting suggests the fauna is considerably larger in tropical Australia. Only four species have been recorded from southern Australia but there is a good chance that more of the WA species occur in the southern part of this state. Haig (1973) provided more detailed descriptions and synonymies for some species. Diagnosis. Eyes well-developed. Maxilliped 1 exopod with flagellum. Rostrum triangular, with 4 (rarely 3–5) distinct lateral teeth.

Key to southern Australian species of Galathea 1. Anterior half of carapace with short, curved scales only ...... 2 — Anterior half of carapace with uninterrupted transverse striae ...... 3 2. Fingers of cheliped more than half as long as propodus; maxilliped 3 merus with spines on outer margin; pereopods 1–3 with epipod ...... Galathea subsquamata — Fingers of cheliped half as long as propodus; maxilliped 3 merus unarmed on outer margin; only cheliped with epipod ...... Galathea magnifica 3. Rostrum as long as broad at base, with pair of gastric spines near base ...... Galathea australiensis — Rostrum longer than broad at base, without pair of gastric spines near base ...... Galathea whiteleggii

231 Marine Decapod Crustacea of Southern Australia

Galathea australiensis Stimpson, 1858 (Fig. 63f, Pl. 13d). Rostrum as long as broad at base. Carapace with transverse striae and pair of anterior gastric spines. 7 mm. Central NSW, Vic., Tas., SA, WA (N to Shark Bay); common at shelf depths. Galathea magnifica Haswell, 1882 (Fig. 63g, Pl. 13e). Carapace with short curved scales. Pair of anterior gastric spines. Fingers of cheliped half as long as propodus. Dark brown with broad stripe down back; white bands on legs. 13 mm. Qld, NSW,Vic., SA, WA (S coast); shelf to 110 m depth. Galathea subsquamata Stimpson, 1858. Carapace with squamiform striae in gastric region; body and limbs with more stiff hairs than other species. Pair of anterior gastric spines. Fingers of cheliped more than half as long as propodus. 13 mm. Japan, eastern Pacific, NT, Qld, WA (S to Rottnest I.); shore to 238 m. This species was known as G. aculeata Haswell, 1882 in earlier literature. Galathea whiteleggii Grant & McCulloch, 1906 (Fig. 63h). Rostrum longer than width at base. Carapace with transverse striae and with only weak pair of anterior gastric spines if present. 5 mm. Qld, NSW; shallow shelf depths.

Munida Leach, 1820 Munida is a genus of more than 120 species recognisable by its spiniform rostrum and distinct transverse ridges on the carapace. More than half the species live in the Indo-West Pacific at shelf depths and beyond. None are found in less than 100 m depth in Australia although M. gregaria can be exceptionally abundant in shallow water in southern New Zealand and Chile. So abundant that the sea and shore can be red with millions of individuals. Such ‘red tides’ have not been reported in Australia. Keys to the identification of Indo-West Pacific species were provided by Baba (1988) and Macpherson & Baba (1993), Macpherson & de Saint Laurent (2002) for the western and southern Indian Ocean, and Ahyong & Poore (2004) for Australian species. Macpherson (2004) described more from Tonga and Fiji. Seven new species from eastern Australia were described by Ahyong & Poore (2004) and some of those might yet be discovered further south. Identification depends on careful attention to spines on the carapace and antennules. Diagnosis. Eyes well-developed. Maxilliped 1 exopod with flagellum. Rostrum spiniform, well- developed. Carapace with distinct transverse ridges. Pereopods 1–3 without epipod. Frontal margin of carapace without spine mesiad to anterolateral spine. Male gonopod 1 present.

Key to southern Australian species of Munida 1. Carapace with lateral margin having 3 spines behind cervical groove ...... Munida kapala — Carapace with lateral margin having 4 spines behind cervical groove ...... Munida rogeri — Carapace with lateral margin having 5 spines behind cervical groove ...... 2 2. Abdominal somite 3 with spines on anterior margin ...... 3 — Abdominal somite 3 without spines on anterior margin ...... 5 3. Abdominal somite 4 with pair of spines on anterior margin ...... Munida gregaria — Abdominal somite 4 without pair of spines on anterior margin ...... 4 4. Antennular basal article with inner distal spine longer than outer ...... Munida chydaea — Antennular basal article with inner distal spine shorter than outer . . . . . Munida asprosoma 5. Antennular basal article with inner distal spine as long as outer; rostrum laterally compressed ...... Munida rubridigitalis — Antennular basal article with inner distal spine unequal to outer; rostrum spiniform . . . . 6 6. Antennular basal article with inner distal spine longer than outer ...... Munida haswelli — Antennular basal article with inner distal spine shorter than outer ...... 7

232 Anomura – hermit crabs, porcelain crabs and squat lobsters

7. Sternite 7 with patch of granules laterally ...... Munida spinicruris — Sternite 7 without patch of granules laterally ...... 8 8. Cornea as wide or little wider than eyestalk ...... Munida endeavourae — Cornea much wider than eyestalk ...... Munida isos

a b b

d e

f g

Fig. 64. Galatheidae. a, Munida asprosoma. b, Munida chydaea. c, Munida endeavourae. d, Munida isos. e, Munida kapala. f, Munida rogeri. g, Munida spinicruris.

Munida asprosoma Ahyong & Poore, 2004 (Fig. 64a). Carapace with 4 or 5 pairs of epigastric spines in addition to smaller scattered spinules; with paired parahepatic, paired anterior branchial and paired postcervical spines; lateral margins of carapace with 5 spines posterior to cervical groove. Eyes much wider than eyestalk. Antennular basal article with 2 distal spines, outer longer. Abdominal somite 2 with 10 spines; somite 3 with 2 spines on anterior border. 29 mm. Qld, NSW; 823–1053 m depth.

233 Marine Decapod Crustacea of Southern Australia

Munida chydaea Ahyong & Poore, 2004 (Fig. 64b). Carapace with pairs of epigastric and protogastric spines and numerous spinules on anterior quarter of carapace; with paired parahepatic, paired anterior branchial and paired postcervical spines; lateral margins of carapace with 5 spines posterior to cervical groove. Eyes much wider than eyestalk. Antennular basal article with distomesial spine longer than distolateral spine. Abdominal somite 2 with 8–10 anterior spines; somite 2 with 2 spines. 44 mm. NSW (N to Sydney), Vic., Tas., SA; 146–700 m depth. This is the most common species of Munida in south-eastern Australia, frequently misidentified as M. haswelli. Munida endeavourae Ahyong & Poore, 2004 (Fig. 64c). Carapace with transverse row of 4–6 epigastric spines; with 0–3 minute spinules behind epigastric spines; with 0–1 minute parahepatic spinules, 1–3 pairs of anterior branchial and 1–2 pairs of postcervical spines; margins of carapace with 5 spines posterior to cervical groove. Eyes scarcely wider than eyestalk. Antennular basal article with 2 distal spines, mesial shorter. Abdominal somite 2 with 6–8 anterior spines. 26 mm. NSW (N to Green Cape), Vic., Tas.; 860–1169 m depth. The species has been previously reported as M. microps. Munida gregaria (Fabricius, 1793) (Pl. 13c). Carapace dorsally rugose and spinulose; 1 spine plus 1–2 spinules on each side of epigastric area, spinule on end of first stria; 1 protogastric, 1 anterior branchial and 2 postcervical spines on each side; anterolateral spine strong. Supraocular spines about third length of spiniform rostrum. Abdominal somite 2 with 4 spines, somite 3 with 2–4 spines, somite 4 with 2 spines. 25 mm. New Zealand, southern South America to 1100 m depth; southern NSW, eastern Vic., eastern Tas.; 7–429 m depth. Munida haswelli Henderson, 1885. Carapace dorsally rugose and spinulose, striae numerous; 1 epigastric spine, 1 protogastric, 1 or more anterior branchial and 1 postcervical spines on each side. Anterolateral spine stronger than other laterals. Abdominal somite 2 with 6–10 spines, somite 3 without spines. 22 mm. NSW (N to Sydney), Vic., Tas., SA, WA (to central Great Australian Bight); 121–329 m depth. This species superficially resembles M. chydaea but lacks spines on the abdominal margin of the third abdominal somite. Its maximum size is half that of M. chydaea. Munida isos Ahyong & Poore, 2004 (Fig. 64d). Carapace with long, scattered setae; with transverse row of 6–7 (usually 6) epigastric spines; with numerous granules on anterolateral region; with 1–2 anterior branchial spines; with or without postcervical spine; margins of carapace with 5 spines posterior to cervical groove. Eye much wider than eyestalk. Antennular basal article with 2 distal spines, mesial shorter. Abdominal somite 2 with 6 anterior spines. 21 mm. NSW (N to Broken Bay), Tas.; 732–1281 m depth. Munida kapala Ahyong & Poore, 2004 (Fig. 64e). Carapace with 4 pairs of epigastric spines; margins of carapace anterior to cervical groove with 2 spines (including anterolateral); with 3 spines posterior to cervical groove. Abdominal somite 2 with 8 spines on anterior border; other abdominal somites unarmed. Eyes much wider than eyestalk. Antennular basal article with 2 distal spines, mesial shorter; with 2 lateral spines. 11 mm. Qld, NSW, Taupo Seamount; 244–549 m depth. Munida rogeri Macpherson, 1994 (Fig. 64f). Carapace dorsally rugose; 10 epigastric spines on first stria, 1 anterior branchial and 1 postcervical spines on each side. Supraocular spines subparallel, about half or less length of spiniform rostrum. Abdominal somite 2 with 8 anterior spines. 13 mm. New Caledonia, Chesterfield Is, WA (S to Perth); 146–390 m depth. Previous records of Munida japonica from Australia are referable to M. rogeri. Munida rubridigitalis Baba, 1994. Carapace dorsally rugose, striae numerous and closely spaced; 6 epigastric spines. Rostrum laterally compressed, not spiniform. Antennular basal article with inner distal spine as long as outer. Abdominal somite 2 with 8 spines, somite 3 without spines. Chela stout, distinctly shorter than body length. 18 mm. Coral Sea, NSW (S to Long Reef); 156–549 m depth.

234 Anomura – hermit crabs, porcelain crabs and squat lobsters

Munida spinicruris Ahyong & Poore, 2004 (Fig. 64g). Carapace with 5–6 pairs of epigastric spines; with several spinules on lateral anterior quarter of carapace in addition to paired parahepatic, paired anterior branchial and paired postcervical spines; margins of carapace with 5 spines posterior to cervical groove. Last sternite with coarse granules laterally. Abdominal somite 2 with 6–8 spines on anterior border. Eyes wider than eyestalks. Antennular basal article with 2 distal spines, mesial longer. 13 mm. Taupo Seamount; 143 m depth.

Munidopsis Whiteaves, 1874 There are more than 150 described species of Munidopsis (Baba, 1988) of which 17 are now reported from Australia (Baba, 1994; Baba & Poore, 2002; Ahyong & Poore, 2004). All Australian species are from deep water on the continental slope or seamounts. The key is an artificial one and the older literature needs to be consulted for accurate identifications. General carapace shape is the most useful means of recognising the species. Diagnosis. Eyes small, eyestalks often produced as spines. Maxilliped 1 exopod without flagellum. Rostrum variable, often flattened.

Key to southern Australian species of Munidopsis 1. Rostrum with prominent lateral oblique spines, its apex upturned; carapace with 2 strong median spines and otherwise dorsally microscopically tuberculate . . . . Munidopsis rostrata — Rostrum simply triangular or with subapical notches; carapace without strong spines . . . . 2 2. Rostrum with subapical notch separating apex from triangular tooth ...... 3 — Rostrum simple, triangular ...... 4 3. Carapace with 4 lateral spines, including anterolateral ...... Munidopsis serricornis — Carapace with 3 lateral spines, including anterolateral ...... Munidopsis treis 4. Eyestalks barely visible under rostrum which is expanded subproximally; carapace granular ...... Munidopsis marginata — Eyestalks clearly visible alongside narrow rostrum; carapace smooth or hirsute ...... 5 5. Carapace with pair of prominent spines at base of rostrum ...... 6 — Carapace with pair of weak tubercles or no spines at base of rostrum ...... 9 6. Spines at base of rostrum sharp ...... 7 — Spines at base of rostrum blunt ...... 8 7. Rostrum quarter length of rest of carapace ; all limbs without epipods . . Munidopsis verrilli — Rostrum half length of rest of carapace; cheliped with epipod . . . . Munidopsis subsquamosa 8. Cheliped palm about 4 times as long as high; antennal process of carapace sharp ...... Munidopsis proales — Cheliped palm about twice as long as high; antennal process of carapace broadly blunt ...... Munidopsis tasmaniae 9. Without spine at end of narrow eyestalk ...... Munidopsis kensleyi — With spine at end of short, broad eyestalk ...... 10 10. Carapace without dense coat of fine setae, with squamae only . . . . Munidopsis bispinoculata — Carapace densely covered with fine setae ...... 11 11. Anterolateral margin of carapace convex, denticulate, separated by notch from posterolateral margin; eyestalk with prominent spine, cornea lateral ...... Munidopsis edwardsii — Anterolateral and posterolateral margins of carapace largely continuous, together weakly convex; eyestalk with major and minor terminal spines ...... Munidopsis victoriae

235 Marine Decapod Crustacea of Southern Australia

c b a

g ef

k j l

Fig. 65. Galatheidae. a, Munidopsis bispinoculata. b, Munidopsis edwardsii. c, Munidopsis kensleyi. d, Munidopsis marginata. e, Munidopsis proales. f, Munidopsis rostrata. g, Munidopsis serricornis. h, Munidopsis subsquamosa. i, Munidopsis tasmaniae. j, Munidopsis treis. k, Munidopsis verrilli. l, Munidopsis victoriae.

236 Anomura – hermit crabs, porcelain crabs and squat lobsters

Munidopsis bispinoculata Baba, 1988 (Fig. 65a). Carapace smooth, with interrupted ridges posteriorly, lateral margins with anterolateral and smaller spines, anterior margin with supraocular and larger antennal spine. Rostrum carinate. Abdominal somites 2 and 3 each with 2 transverse ridges. Eyestalks immoveable, with 2 distal spines, inner one ventral and smaller. Pereopods without epipods. 15 mm. Indonesia, NSW slope; 933–2363 m depth. Munidopsis edwardsii (Wood-Mason, 1891) (Fig. 65b). Carapace covered with fine setae, with ripples posteriorly, lateral margins subparallel, with anterolateral spine anteriorly, anterior margin without supraocular or antennal spine. Rostrum carinate. Abdominal somites 2 and 3 each with 2 transverse ridges. Eyestalks slightly moveable, with dominating distal spine, cornea lateral. 28 mm. Indian Ocean, NSW; 1896–2610 m depth. Munidopsis kensleyi Ahyong & Poore, 2004 (Fig. 65c). Carapace longer than wide, covered with plumose setae, smooth except for 2–5 spines on posterior margin, anterior margin oblique, anterolateral and posterolateral margins parallel, barely distinct; with anterolateral spine anteriorly. Rostrum carinate, unarmed. Abdominal somites 2 and 3 each with 2 transverse ridges. Eyestalks slender, without distal spines. Chelipeds with epipods. 25 mm. NSW; 214–1939 m depth (reported as M. dasypus by Baba & Poore, 2002). Munidopsis marginata (Henderson, 1885) (Fig. 65d). Carapace granular, lateral margins flanged behind prominent triangular lobe, anterior margin without supraocular or antennal spines. Rostrum broad subproximally, flattened and carinate. Abdominal somites 2 and 3 each with 2 transverse ridges. Eyes fused to side of rostrum, with small distal spine. Pereopods without epipods. 15 mm. New Zealand, NSW slope; about 2000 m depth. Munidopsis proales Ahyong & Poore, 2004 (Fig. 65e). Carapace excluding rostrum longer than wide, with strongly rugose squamae and tubercles; cervical groove distinct; regions well- defined; with pair of blunt epigastric processes; frontal margin with blunt antennal process; lateral margins with 4 large, anterolaterally directed teeth; posterior margin unarmed. Rostrum unknown. Abdominal somites unarmed. Eyestalk moveable, with short dorsal processes. Pereopods 1–3 with epipod. 9 mm. WA (S coast); 513–540 m depth. Munidopsis rostrata (Milne Edwards, 1880) (Fig. 65f). Carapace with tubercles, armed with 2 small epigastric, 1 strong mesogastric and 1 cardiac spines; lateral margin with 2 prominent spines anteriorly. Rostrum with lateral spines, upturned apically. Abdominal somites 2–4 each with median spine. Pereopods 1–3 with epipods. 38 mm. Widespread in Atlantic, Indo-Pacific and Southern Ocean; Qld, NSW and Vic. slope; 1650–3294 m. Munidopsis serricornis (Lovén, 1852) (Fig. 65g). Carapace, with obsolete interrupted transverse ridges posteriorly, lateral margins subparallel, with 3–4 spines, anterior margin with antennal spine. Rostrum carinate, distally trifid. Abdomen unarmed. Eyestalks moveable, without distal spine, cornea distal. Pereopods without epipods. 15 mm. Atlantic Ocean, Indo-Pacific, Vic., Tas. Seamounts, Cascade Plateau; 100 m in Norway, to 2165 m depth. Munidopsis subsquamosa Henderson, 1885 (Fig. 65h). Carapace with pair of epigastric spines, interrupted ridges all over; anterior margin with broad antennal spine; lateral margin with anterolateral spine, 3 spines plus fourth and others behind cervical constriction; posterior margin unarmed. Rostrum carinate, upturned, unarmed. Abdominal somites 2 and 3 each with 2 transverse ridges. Eyestalks with 2 distal spines, inner one larger. Cheliped with epipod. 40 mm. East and west Pacific, NSW; 2690–3430 m depth. Munidopsis tasmaniae Ahyong & Poore, 2004 (Fig. 65i). Carapace with sparsely setose squamae and tubercles; with pair of broad, blunt, flattened epigastric processes; frontal margin with blunt antennal process; lateral margins with 4 blunt teeth; posterior margin unarmed. Rostrum triangular, about one-third remaining carapace length, broad basally, margins convex and serrate, carinate, sparsely tuberculate. Abdominal somites unarmed. Eyestalk moveable, with short, papillate, tubercular process mesiodorsally. Pereopods 1–3 with epipod. 16 mm. Tas.; 1100–1135 m depth.

237 Marine Decapod Crustacea of Southern Australia

Munidopsis treis Ahyong & Poore, 2004 (Fig. 65j). Carapace without dorsal spines; frontal margins oblique, with slender antennal spine, strong anterolateral spine, 2 strong spines on hepatic margin. Rostrum broad, trifid distally. Abdominal somites unarmed. Eyestalk unarmed. Pereopods without epipod. 16 mm. Tas., Tasmanian Seamounts, SA; 366–820 m depth (reported as M. serricornis by Baba & Poore, 2002). Munidopsis verrilli Benedict, 1902 (Fig. 65k). Carapace with stiff setae, with 2 epigastric spines, interrupted transverse ridges posteriorly, lateral margins convex, with 5 prominent spines, anterior margin with antennal spine. Rostrum carinate, distally upturned. Abdomen unarmed. Eyestalks moveable, with 2 distal spines, mesial one larger, cornea between spines. Pereopods without epipods. 29 mm. North-eastern Pacific Ocean (off USA), Tas.; 1500–2010 m depth. Munidopsis victoriae Baba & Poore, 2002 (Fig. 65l). Carapace with short pilosity, smooth, lateral margin with prominent anterolateral spine, anterior margin with supraocular and smaller antennal spine. Rostrum carinate. Abdominal somites 2–4 each with transverse ridge and groove. Eyestalks immoveable, with 2 distal spines, inner one ventral and smaller. Pereopods without epipods. 24 mm. Western Vic.; 990 m depth.

Paramunida Baba, 1988 Ahyong & Poore (2004) recorded three species of Paramunida from Australia and Macpherson (2004) several from Tonga and Fiji. All are from deep water. The genus differs from Munida in having a plate-like base to the rostrum and supraocular spines. Diagnosis. Eyes well-developed. Rostrum spiniform, short, fused basally with supraocular spines. Carapace without clear transverse ridges, with scales or short ridges, with pair of epigastric spines behind supraocular spines, and with pair of postcervical spines. Pereopods 1–3 without epipod; dactyli of walking legs compressed dactyli of walking legs depressed vertically, with lateral and mesial ridges. Frontal margin of carapace without spine mesiad to anterolateral spine. Male gonopod 1 absent. Paramunida antipodes Ahyong & Poore, 2004 (Fig. 66a). Carapace with pair of epigastric spines, protogastric spines, large mesogastric spine; cardiac region with longitudinal row of 2–3 spines; posterior margin with median spine; branchiocardiac region with longitudinal row of 3 spines on each side. Antennular basal article lateral margin with 4–5 small spines. Antennal article 1 with long, sinuous mesial spine; article 2 with sharp apex on mesial spine. 13 mm. NSW; 420–548 m depth.

Phylladiorhynchus Baba, 1969 Of the three Indo-Pacific species only one occurs in Australia (Baba, 1969, 1991, 1993). Diagnosis. Eyes well-developed; maxilliped 1 exopod with flagellum; rostrum leaf-like, with tiny distolateral and distinct basilateral teeth; carapace with distinct transverse striae anteriorly. Phylladiorhynchus pusillus (Henderson, 1885) (Fig. 66b). Rostrum with marginal spine near apex. 1–2 pairs protogastric spines. Abdominal somites 2 and 3 with median transverse groove. Pereopods without epipods. 7 mm. Eastern Indian Ocean, Japan, New Caledonia, New Zealand, southern Qld, NSW, Vic., Tas., SA, WA (N to Shark Bay); intertidal to 310 m depth. This small species is distinguishable from all other galatheids in southern Australia by the form of the rostrum. See Haig (1973) for synonymy, and Baba (1991).

238 Anomura – hermit crabs, porcelain crabs and squat lobsters

Raymunida Macpherson & Machordom, 2000 Eight species of the Raymunida are presently recognised (Macpherson & Machordom, 2000, 2001 Lin et al., 2004). A phylogenetic analysis based on morphology and molecular data confirmed the importance of colour as a taxonomic character (Macpherson & Machordom, 2001). The small spine in the orbit is diagnostic of the genus. Dorsal ornamentation of the two southern Australian species is very similar.

Diagnosis. Eyes well-developed. Maxilliped 1 exopod with flagellum. Rostrum spiniform, well- developed. Carapace with distinct transverse ridges. Pereopods 1–3 with epipod. Frontal margin of carapace with spine mesiad to anterolateral spine.

Key to southern Australian species of Raymunida 1. Antennal peduncle first article with mesial spine overreaching basal article of antennular peduncle ...... Raymunida formosanus — Antennal peduncle first article with mesial spine not reaching basal article of antennular peduncle ...... Raymunida elegantissima Raymunida elegantissima (De Man, 1902). Carapace dorsally rugose; 10 epigastric spines; 1 lateral protogastric, 2 anterior branchial and 1 postcervical spines on each side. Antennal peduncle first article with mesial spine not reaching basal article of antennular peduncle. 14 mm. Indian Ocean, Indonesia, Japan, WA (S to Perth), NT, Qld (S to Cape Moreton); 20–200 m depth. Raymunida formosanus Lin, Chan & Chu, 2004 (Fig. 66c, d). Carapace dorsally striate; 12 epigastric spines; 1 lateral protogastric, 2 anterior branchial and 1 postcervical spines on each side. Walking legs with distinct scale-like striae. Antennal peduncle first article with mesial spine overreaching basal article of antennular peduncle. 20 mm. Taiwan, NSW; 104 m depth.

b c

Fig. 66. Galatheidae. a, Paramunida antipodes. b, Phylladiorhynchus pusillus. c, d, Raymunida formosanus (with right antennule and antenna).

References Ahyong, S.T., & Poore, G.C.B. 2004. Deep-water Galatheidae (Crustacea: Decapoda: Anomura) from southern and eastern Australia. Zootaxa 472: 1–76. Baba, K. 1969. Four new genera with their representatives and six new species of the Galatheidae in the collection of the Zoological Laboratory, Kyushu University, with redefinition of the genus Galathea. Ohmu 2: 4–32.

239 Marine Decapod Crustacea of Southern Australia

Baba, K. 1979. Expédition Rumphius II (1975) Crustacés parasites, commensaux, etc. (Th. Monod et R. Sèrene, éd.) VII. Galatheid crustaceans (Decapoda, Anomura). Bulletin du Muséum National d’Histoire Naturelle, Paris A, 1: 643–657. Baba, K. 1988. Chirostylid and galatheid crustaceans (Decapoda: Anomura) of the ‘Albatross’ Philippine Expedition, 1907–1910. Researches on Crustacea, Special Number 2: 1–203. Baba, K. 1991. Crustacea Decapoda: Alainius gen. nov., Leiogalathea Baba, 1969, and Phylladiorhynchus Baba, 1969 (Galatheidae) from New Caledonia. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152: 479–491. Baba, K. 1993. Anomoeomunida, a new genus proposed for Phylladiorhynchus caribensis Mayo, 1972 (Crustacea: Decapoda: Galatheidae). Proceedings of the Biological Society of Washington 106: 102–105. Baba, K. 1994. Deep-sea galatheid crustaceans (Anomura: Galatheidae) collected by the ‘Cidaris I’ Expedition off central Queensland, Australia. Memoirs of the Queensland Museum 35: 1–21. Baba, K. 1996. Crustacea Decapoda: revision of the genus Bathymunida Balss, 1914, and description of six new related genera (Galatheidae). In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 15. Mémoires du Muséum National d’Histoire Naturelle, Paris 168: 433–502. Baba, K., & Poore, G.C.B. 2002. Munidopsis (Decapoda, Anomura) from south-eastern Australia. Crustaceana 75: 231–252. Haig, J. 1972. The anomuran crabs of Western Australia: their distribution in the Indian Ocean and adjacent seas. Journal of the Marine Biological Association of India 14: 443–451. Haig, J. 1973. Galatheidea (Crustacea, Decapoda, Anomura) collected by the F.I.S. Endeavour. Records of the Australian Museum 28: 269–289. Haig, J. 1974. The anomuran crabs of Western Australia: their distribution in the Indian Ocean and adjacent seas. Journal of the Marine Biological Association of India 14: 443–451. Jones, D.S., & Morgan, G.J. 2002. A Field Guide to Crustaceans of Australian Waters. Reed New Holland: Sydney. 224 pp. 2nd edn. Lin, C.-W., Chan, T.-Y., & Chu, K.H. 2004. A new squat lobster of the genus Raymunida (Decapoda: Galatheidae) from Taiwan. Journal of Crustacean Biology 24: 149–156. Macpherson, E. 2004. Species of the genus Munida Leach, 1820 and related genera from Fiji and Tonga (Crustacea: Decapoda: Galatheidae). In: Marshall, B.A., & Richer de Forges, B. (eds), Tropical Deep-Sea Benthos. Vol. 23. Mémoires du Muséum National d’Histoire Naturelle, Paris 191: 231–292. Macpherson, E., & Baba, K. 1993. Crustacea Decapoda: Munida japonica Stimpson, 1858, and related species (Galatheidae). In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156: 381–420. Macpherson, E., & Machordom, A. 2000. Raymunida, new genus (Decapoda: Anomura: Galatheidae) from the Indian and Pacific Oceans. Journal of Crustacean Biology 20 (Special Number 2): 253–258. Macpherson, E., & Machordom, A. 2001. Phylogenetic relationships of species of Raymunida (Decapoda: Galatheidae) based on morphology and mitochondrial cytochrome oxidase sequences, with the recognition of four new species. Journal of Crustacean Biology 21: 696–714. Macpherson, E., & de Saint Laurent, M. 2002. On the genus Munida Leach, 1820 (Decapoda, Galatheidae) from the western and southern Indian Ocean, with the description of four new species. Crustaceana 75: 465–484. Poore, G.C.B., Hart, S., Taylor, J., & Tudge, C. 1998. Decapod crustaceans from Tasmanian seamounts. Pp. 65–78 in: Koslow, J.A., & Gowlett-Holmes, K. (eds), The seamount fauna of southern Tasmania: benthic communities, their conservation and impacts of trawling. Final report to Environment Australia and The Fisheries Research Development Corporation. CSIRO Marine Research: Hobart.

Lomisidae Bouvier, 1895 The family is monotypic. The family name has alternatively been spelt Lomidae (occasionally since 1969) but in the opinion of Dr L. Holthuis, considered the world expert on crustacean names, the spelling Lomisidae is to be preferred. The reason for this is that Lomis is not a Greek or

240 Anomura – hermit crabs, porcelain crabs and squat lobsters

Latin word so has no stem (like Lom-) to which the -idae ending can be added. In fact, the original author of Lomis coined the French common name ‘Lomisinés’ for these Australian crabs. Diagnosis. Body symmetrical. Ocular scales absent. Antennal peduncle with 6 articles. Epistome with spine. 14 pairs of branchial gills. Pleopods 3–5 paired. Female pleopods 1 present. Uropods non-spatulate, vestigial in male. Male gonopods present. Pereopods 2–5 dactyli with claw; last pereopod vestigial. Lomis hirta (Lamarck, 1818) hairy stone crab (Fig. 67, Pl. 14d). 16 mm. Vic., Tas., SA, WA (N to Shark Bay). The hairy stone crab is a very unusual false crab common under rocks in the intertidal zone of exposed rocky shores of southern Australia. In Victoria it is recorded only from open coast west of Wilsons Promontory and in the far east of the state (Phillips et al., 1984). It is best recognised by its flattened form, grey hairy and muddy appearance, with legs and chelipeds so well fitting together that it clings closely to its rocky surface. The antennae are brilliant blue. It differs from true brachyuran crabs in possessing vestigial fifth pereopods. It resembles porcellanid crabs more closely but differs in the reduced uropods, presence of female first pleopods, and eyes contiguous and flattened. The species’ systematic position has long been enigmatic and after a history of being shifted from one anomuran group to another was sometimes thought to be so different from all others that it deserves its own superfamily. Pilgrim (1965) presented detailed information on morphology and discussed its systematic position. McLaughlin (1983) and Tudge (1997) discussed the matter further and decided on the classification adopted here. The first stage zoea larva was described by Cormie (1993) who suspected it may have an abbreviated larval life.

Fig. 67. Lomisidae. Lomis hirta.

241 Marine Decapod Crustacea of Southern Australia

References Cormie, A.K. 1993. The morphology of the first zoeal stage of Lomis hirta (Lamarck, 1818) (Decapoda, Lomisidae). Crustaceana 64: 249–255. McLaughlin, P.A. 1983. A review of the phylogenetic position of the Lomidae (Crustacea: Decapoda: Anomala). Journal of Crustacean Biology 3: 431–447. Phillips, D.A.B., Handreck, C.P., Bock, P.E., Burn, R., Smith, B.J., & Staples, D.A. 1984. Coastal invertebrates of Victoria. An atlas of selected species. Marine Research Group of Victoria in association with the Museum of Victoria: Melbourne. 168 pp. 1st edn. Pilgrim, R.L.C. 1965. Some features in the morphology of Lomis hirta (Lamarck) (Crustacea: Decapoda) and a discussion of its systematic position and phylogeny. Australian Journal of Zoology 13: 545–557. Tudge, C.C. 1997. Spermatological evidence supports the taxonomic placement of the Australian endemic hairy stone crab, Lomis hirta (Decapoda: Anomura: Lomidae). Memoirs of the Museum of Victoria 56: 235–244.

Porcellanidae Haworth, 1825 The family Porcellanidae, porcelain crabs or false crabs, is so easily recognised that it has not been diagnosed in recent literature. The species can be easily told from true brachyuran crabs in having the fifth pereopod not immediately obvious (held up over the carapace), in having the antenna with an elongate flagellum, and in the abdomen not fitting closely to the underside of the cephalothorax. They are always very flattened which is the easiest way to distinguish them from the larger deepwater lithodids (also with reduced fifth legs). Most often porcelain crabs are found clinging to the undersides of intertidal rocks. The translation of Petrolisthes, one of the genera, as ‘stone slider’ is a good description of their habits. As Hale (1927) reported ‘when alarmed some of the species exhibit a ferocious aspect which, consid- ering their diminutive size, is ludicrous; they often part with their large claws when captured.’They have long third maxillipeds fringed with long plumose setae which are the main food gathering organs. Although there are more than 200 species worldwide in more than a dozen genera (Haig, 1978 and numerous other papers by this author since 1960) only five occur on the southern coast of Australia. Tropical Australia is home to more than 30 species in at least nine genera (Haig, 1965, 1972, 1981, 1988; Jones, 1990). Colour patterns differ between species as shown in Jones & Morgan’s (2002) photographs of three tropical species. Haig’s (1965) key and diagnoses of the Western Australian species are still useful; our key and diagnoses are derived from this work.

Diagnosis. Abdomen folded up against thorax; body crab-like; chelipeds moderately elongate, stout or flattened.

Key to southern Australian genera and species of Porcellanidae 1. Antenna peduncle with moveable articles freely accessible to orbit ...... 2 — Antenna peduncle with moveable articles excluded from orbit by projection of basal article meeting anterior margin of carapace ...... 4 2. Front straight; posterior part of lateral carapace walls separated from rest by membranes ...... Pachycheles granti — Front with triangular rostrum; lateral walls of carapace entire ...... 3 3 Front a strongly produced triangular rostrum, armed with row of strong spines; telson with 5 plates ...... Petrocheles australiensis — Front broadly rounded; telson with 7 plates ...... Petrolisthes elongatus

242 Anomura – hermit crabs, porcelain crabs and squat lobsters

4. Carapace much broader than long ...... Polyonyx transversus — Carapace as long as or longer than broad ...... 5 5. Fingers of one or both chelipeds twisted, especially in males; carapace with 2 hepatic spines; front appearing transverse in dorsal view, median lobe sharply deflexed . . . . . Pisidia dispar — Fingers twisted only in smaller cheliped (especially of large males); carapace smooth; front obtusely triangular in dorsal view ...... Ancylocheles gravelei

a b

Fig. 68. Porcellanidae. a, Ancylocheles gravelei. b, Pachycheles granti. c, Pisidia dispar. d, Polyonyx transversus.

243 Marine Decapod Crustacea of Southern Australia

Ancylocheles Haig, 1978 The Australian species is the type species and the only one. Diagnosis. Antenna peduncle with moveable articles excluded from orbit by projection of basal article meeting anterior margin of carapace. Carapace about as long as broad, strongly convex front to back, smooth or somewhat roughened, regions clearly marked; lateral margins ridged, unarmed posterior to epibranchial angle. Front horizontal or deflexed, broadly triangular or rounded in dorsal view; median lobe deflexed. Chelipeds unequal, especially in large males, flattened; dactylus of smaller cheliped twisted. Walking legs with 1 well-developed claw and moveable spinules ventrally. Telson with 7 plates. Ancylocheles gravelei Sankolli, 1963 (Fig. 68a). 7 mm. India, NSW, Qld, NT, WA (S to Esperance); intertidal to 17 m. The genus, and therefore this species, is very similar to Pisidia dispar but differs in lacking a strongly tridentate front and less pronounced sexual dimorphism and twisting of the chelipeds. The species has been recorded as Porcellana gravelei by Haig (1965) and Jones (1990).

Pachycheles Stimpson, 1858 There are at least four species in Australia (Haig, 1965). Diagnosis. Antenna peduncle with moveable articles freely accessible to orbit, basal article not produced. Carapace rounded or subquadrate, front rounded or transverse, without rostrum; lateral margins usually without strong spines; lateral walls incomplete, posterior part divided by membrane from rest. Chelipeds unequal; robust; carpus with strong spines on margins and dorsally. Telson with 5 or 7 plates. Pachycheles granti Haig, 1965 (Fig. 68b). Carapace smooth. Cheliped carpus and chela with 5 longitudinal rows of tubercles, covered with short hairs; carpus with 3 broad teeth on inner margin. 8 mm. Qld, NSW (S to Sydney); upper subtidal.

Petrocheles Miers, 1876 The Australian species and another in New Zealand are the only members of the genus. Diagnosis. Antenna peduncle with moveable articles freely accessible to orbit, basal article not produced. Carapace cordate, wider posteriorly, with strongly produced triangular rostrum; lateral margins with strong spines. Chelipeds subequal; chela long, slender and flattened; carpus with strong spines on margins and dorsally. Telson with 5 plates. Petrocheles australiensis (Miers, 1876) spiny porcelain crab (Fig. 69a, b). Protogastric region with pair of spines. Rostrum with 3 (usually 4) spines each side. Chelipedal carpus with 6–7 spines on inner margin. Reddish-brown. 23 mm. Western Vic., Tas., SA, eastern WA (Phillips et al., 1984); intertidal on open coasts.

Petrolisthes Miers, 1876 Petrolisthes is a large genus with 35 species in the Indo-West Pacific of which at least ten occur in northern Australia (Haig, 1965, 1972). Only one occurs in southern Australia. Diagnosis. Antenna peduncle with moveable articles freely accessible to orbit, basal article not produced. Carapace rounded or subquadrate, about as broad as long; front triangular or trilobate, often prominent, not armed. Chelipeds subequal; chela flattened. Telson with 7 plates. Petrolisthes elongatus (Milne Edwards, 1837) (Fig. 69c, d, Pl. 14a). Carapace finely granulate, without dorsal spines, without supraocular spine; front broadly rounded-triangular. Cheliped

244 Anomura – hermit crabs, porcelain crabs and squat lobsters

carpus inner margin with obsolete ridge proximally, outer margin with 1–3 weak spines and sharp distal angle; propodus unarmed, weakly granulate. Merus of walking legs unarmed. 21 mm long. Steely blue, sometimes greenish. New Zealand, eastern Tas.; intertidal rocky shores. It has been confirmed that this species was introduced to Tasmania from New Zealand, probably with ballast rocks at the end of the 1800s (King, 1997).

b d

Fig. 69. Porcellanidae. a, b, Petrocheles australiensis (with telson). c, d, Petrolisthes elongatus (with telson).

Pisidia Leach, 1820 There are 13 species of Pisidia worldwide, of which seven occur in the Indo-West Pacific (Haig, 1978). Only one occurs in Australia, others reported for this genus having been transferred to other genera. Diagnosis. Antenna peduncle with moveable articles excluded from orbit by projection of basal article meeting anterior margin of carapace. Carapace usually rounded, with lateral spine; front prominent, trilobate. Chelipeds unequal, fingers twisted especially in smaller cheliped and more so in males. Carapace and chelipeds more spinose in females and juveniles. Dactylus of walking leg with single claw, lower margin with row of moveable spinules, the most distal stout. Telson of 7 plates. Pisidia dispar (Stimpson, 1858) polished porcelain crab (Fig. 68c, Pl. 14b). Front depressed, median lobe may only be visible in anterior view, acute, longer than lateral lobes. Carapace with hepatic spine; lateral margin with 1–3 small postcervical spines. Cheliped merus with

245 Marine Decapod Crustacea of Southern Australia

strong mesial lobe; carpus with 3 mesial lobes or teeth and 2–4 lateral spines; propodus with lateral row of spinules. Merus of walking leg unarmed. 6 mm. New Caledonia, Molucca Is, Japan, SA, WA, NT, Qld, NSW; littoral to 55 m depth. Hale (1927) figured this widespread species as a species of Porcellana.

Polyonyx Stimpson, 1858 Polyonyx is a moderately large genus with at least 14 species in the Indo-West Pacific of which at least seven occur in Western Australia. There is only one species on southern coasts. Diagnosis. Antenna peduncle with moveable articles excluded from orbit by projection of basal article meeting anterior margin of carapace. Carapace broader than long, convex; front deflexed, trilobate or transverse. Chelipeds large, unequal. Walking legs with at least 2 well-developed claws and often with spinules on lower margin. Telson with 7 plates. Pleopods usually present in males. Polyonyx transversus (Haswell, 1882) (Fig. 68d, Pl. 14c). Carapace about 1.4 times as broad as long, with fringe of hairs along frontal margin; lateral margins unarmed. Cheliped merus with vestigial lobe; carpus hairy on inner margin, developed into a prominent, convex lobe, broadest distally and unarmed; chela dorsally hairy. Propodus of walking legs with more than 12 spinules on lower margin; dactylus with lower claw larger than upper, with 2 stout spines on lower margin. 11 mm. Southern Qld, NSW,Vic., SA, WA (N to Perth, WA); intertidal to 20 m depth. Polyonyx transversus is an inhabitant of muddy environments and although the basal colour is orange animals appear very muddy. They are often taken in the paper tubes of the large polychaete Chaetopterus variopedatus and may be commensal with this species. This is certainly the case in Port Phillip Bay and Haig (1965) repeated anecdotal evidence about their occurrence in pairs in tubes 35 cm below the surface of intertidal mud.

References Haig, J. 1965. The Porcellanidae (Crustacea, Anomura) of Western Australia), with descriptions of four new Australian species. Journal of the Royal Society of Western Australia 48: 97:118. Haig, J. 1972. The anomuran crabs of Western Australia: their distribution in the Indian Ocean and adjacent seas. Journal of the Marine Biological Association of India 14: 443–451. Haig, J. 1978. Contribution toward a revision of the porcellanid genus Porcellana (Crustacea: Decapoda: Anomura). Proceedings of the Biological Society of Washington 91: 706–714. Haig, J. 1981. Porcellanid crabs from the Indo-west Pacific, Part II. Steenstrupia 7: 269–291. Haig, J. 1988. Two new mangrove-dwelling porcellanid crabs, of genus Petrolisthes Stimpson, from tropical Australia (Crustacea: Decapoda: Anomura). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 5: 71–76. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Jones, D.S. 1990. Annotated checklist of marine decapod Crustacea from Shark Bay, Western Australia. Pp. 169–208 in: Berry, P.B., Bradshaw, S.D., & Wilson, B.R. (eds), Research in Shark Bay: Report of the France-Australe Bicentenary Expedition Committee. Western Australian Museum: Perth. Jones, D.S., & Morgan, G.J. 2002. A Field Guide to Crustaceans of Australian Waters. Reed New Holland: Sydney. 224 pp. 2nd edn. King, R. 1997. Systematic relationships between Tasmanian and New Zealand populations of Petrolisthes elongatus (Crustacea: Anomura: Porcellanidae). B.Sc.(Hons) thesis. University of Melbourne: Melbourne 37 pp. Phillips, D.A.B., Handreck, C.P., Bock, P.E., Burn, R., Smith, B.J., & Staples, D.A. 1984. Coastal Invertebrates of Victoria. An Atlas of Selected Species. Marine Research Group of Victoria in association with the Museum of Victoria: Melbourne. 168 pp. 1st edn.

246 Anomura – hermit crabs, porcelain crabs and squat lobsters

Superfamily Hippoidea Latreille, 1825 Hippoids, the mole and sand crabs, are not true crabs at all. They live buried in sandy sediments from the low intertidal to offshore. There are three families, Albuneidae, Hippidae and Blepharipodidae, separated on the basis of the form of carapace, gills and first leg (Boyko, 2002). The first two are represented in the Indo-West Pacific and Australia by 37 species (Boyko & Harvey, 1999) but neither family is common on the southern coast. Blepharopodids do not occur in the south Pacific. Diagnosis. Body symmetrical. Uropodal rami not forming a tail-fan, without rasps. Ocular scales absent. Pereopods 2–4 dactyli similar.

Key to families of Hippoidea 1. Carapace subquadrangular, flattened, without lateral expansions covering pereopods; pereopod 1 subchelate ...... Albuneidae … p. 247 — Carapace oval, strongly convex, with lateral expansions covering most pereopods; pereopod 1 simple ...... Hippidae … p. 249

Albuneidae Stimpson, 1858 Albuneids, sand crabs, burrow deeply in sand and are therefore rarely taken. It has been suggested that species of the genus Albunea are filter feeders, collecting suspended food particles with the long first antennae but Boyko (2002) discredited this and proposed that they are scavengers or predators. The family contains six genera of which four occur in Australia (Efford & Haig, 1968; Haig, 1972; Davie, 2002). Boyko & Harvey (1999) reported on several undescribed species in Australia masquerading as Albunea steintizi and provided a key to tropical Indo-West Pacific species. There is a key to genera in Efford & Haig’s (1968) paper which describes new genera from Australia and another to two subfamilies and eight genera in Boyko (2002). Only one species occurs in southern Australia but the family is represented by eight other species in more tropical Australia (Davie, 2002). Diagnosis. Carapace subquadrangular, flattened, without lateral expansions covering pereopods. Telson ovoid. Mandible well-developed. Maxilliped 3 with exopod. Pereopod 1 subchelate. Pereopods 2–4 dactyli sickle-shaped. Gills phyllobranchiate.

Austrolepidopa Efford & Haig, 1968 Two of the three species are Australian. Only the western one reaches far enough south to be included in this book. The authors of the genus and Boyko (2002) gave more extensive diagnoses and discussed relationships. Diagnosis. Carapace with 1 lateral spine, above linea anomurica; anterior margin with median truncate lobe. Eyes narrow, lamelliform, separated by distance equal to eye-width. Maxilliped 3 ischium-merus unarmed.

247 Marine Decapod Crustacea of Southern Australia

Austrolepidopa trigonops Efford & Haig, 1969 (Fig. 70a). Median lobe on anterior margin of carapace almost as long as lateral lobes; anterior transverse depression interrupted only in midline, if at all; setal field across anterior part of carapace not extending more anterior than base of lateral carapace spine. Eyestalk apex acute. Maxilliped 3 exopod of 2 articles. Telson as wide as long. 17 mm. WA (Beagle Bay to Perth); 11–36 m. The diagnosis is sufficient to distinguish this species from A. schmitti from southern Queensland (and perhaps NSW) and A. caledonica from WA and New Caledonia.

c b

Fig. 70. Albuneidae, a, Austrolepidopa trigonops. Hippidae. b, Hippa australis. c, Hippa pacifica.

References Boyko, C.B. 2002. A worldwide revision of the Recent and fossil sand crabs of the Albuneidae Stimpsonand Blepharipodidae, new family (Crustacea: Decapoda: Anomura: Hippoidea). Bulletin of the American Museum of Natural History 272: 1–396. Boyko, C.B., & Harvey, A.W. 1999. Crustacea Decapoda: Albuneidae and Hippidae of the tropical Indo- West Pacific region. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180: 379–406. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Efford, I.E., & Haig, J. 1968. Two new genera and three new species of crabs (Decapoda: Anomura: Albuneidae) from Australia. Australian Journal of Zoology 16: 897–914. Haig, J. 1972. The anomuran crabs of Western Australia: their distribution in the Indian Ocean and adjacent seas. Journal of the Marine Biological Association of India 14: 443–451.

248 Anomura – hermit crabs, porcelain crabs and squat lobsters

Hippidae Latreille, 1825 Hippids are commonly called mole crabs. They comprise a small family of sand-burrowing crustaceans found subtidally worldwide. No work has been done on the biology of the local species but research on others suggests they are scavengers or predators, some feeding on the Portuguese- man-o-war Physalia (Haig, 1974). The Australian Hippidae were reviewed by Haig (1974) who diagnosed and keyed all the species. An extensive taxonomic study undertaken on the fauna of the Indo-West Pacific region (Boyko & Harvey, 1999) has overtaken Haig’s work. Of the three genera only two are known to occur in Australia and are represented by three species. The southern Australian fauna comprises only two species in the genus Hippa. Diagnosis. Carapace oval, strongly convex, with lateral extensions covering all except pereopod 1. Mandible reduced. Maxilliped 3 without exopod. Pereopod 1 simple. Pereopods 2 and 3 dactyli curved and flattened, 4 less so. Telson elongate, lanceolate.

Hippa Fabricius, 1787 The genus is represented by at least 15 species worldwide for which a list and a key to some were given by Haig et al. (1986). The tropical Indo-West Pacific species were listed and keyed by Boyko & Harvey (1999). Of the three (possibly four) Australian species only two occur on the southern coast. They can be distinguished on the shape of the frontal margin and other characters. Diagnosis. Antennal flagella shorter than carapace. Pereopod 1 dactylus less than quarter length of carapace, non-articulated.

Key to southern Australian species of Hippa 1. Front with low broad obtusely triangular rostrum; orbits shallowly concave, outer orbital lobes rounded ...... Hippa australis — Front with 4 rounded lobes, outer pair more prominent than inner which are separated by shallow concavity ...... Hippa pacifica Hippa australis Hale, 1927 (Fig. 70b). Carapace covered with shallow setiferous pits, nearly absent anteriorly, elsewhere in more or less even transverse rows, anterolaterally close-set oblique lines near lateral margins. Front with low broad obtusely triangular rostrum; orbits shallowly concave, outer orbital lobes rounded. Antennule longer flagellum of 32–40 articles, shorter flagellum with 5–7 articles in males, 7–12 in females. Pereopods 2 and 3 dactyli with concave margin evenly curved. Males to 10 mm, females to 17 mm. SA (Spencer Gulf), southern WA. Hippa pacifica (Dana, 1852) (Fig. 70c). Carapace covered with wavy transverse lines; row of 30–40 shallow setiferous elongate pits near each lateral margin. Front with 4 rounded lobes, outer pair more prominent than inner which are separated by shallow concavity. Pereopods 2 and 3 dactyli with concave margin obtusely angled. Males to 18 mm, females to 28 mm. Indo-West Pacific, Africa to western central America including islands, WA (S to Rottnest I.), NT, Qld, NSW (S to Coffs Harbour); intertidal. This species has often been referred to as Remipes testudinarius. In Central America, it feeds on animal matter washed ashore (Wenner, 1977).

References Boyko, C.B., & Harvey, A.W. 1999. Crustacea Decapoda: Albuneidae and Hippidae of the tropical Indo- West Pacific region. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180: 379–406. Haig, J. 1974. A review of the Australian crabs of family Hippidae (Crustacea, Decapoda, Anomura). Memoirs of the Queensland Museum 17: 175–189.

249 Marine Decapod Crustacea of Southern Australia

Haig, J., Murugan, T., & Nair, N.B. 1986. Hippa indica, a new species of mole crab (Decapoda, Anomura, Hippidae) from the south west coast of India. Crustaceana 51: 286–292. Wenner, A.M. 1977. Food supply, feeding habits and egg production in Pacific mole crabs (Hippa pacifica Dana). Pacific Science 31: 39–47.

Superfamily Paguroidea Latreille, 1802 The Paguroidea are the hermit crabs, coconut crab, and stone crabs, a heterogeneous assemblage of over 800 species united most obviously by the asymmetry of the abdomen. In the rare case of symmetry (Pylochelidae) a reduced fourth pereopod ties the family with other paguroids. Hermit crabs range from polar to tropical latitudes and from semiterrestrial to abyssal habitats and are often encountered on intertidal rocky shores of southern Australia. Stone crabs are less commonly met, being confined to slope depths. Because hermit crabs are peculiar in inhabiting the shells of dead gastropod molluscs this behaviour has attracted considerable attention. The fauna of the tropics is more diverse than of southern Australia and includes several colourful species, some photographed in a guide book for the Great Barrier Reef by Tudge (1995). Some of the species covered by Tudge occur in southern Australia also. Definition of the Paguroidea is drawn from three important papers by McLaughlin (McLaughlin, 1983a, b, 2003) in which the superfamily Coenobitoidea is no longer recognised. The implication of her findings is that the six families of hermit crabs and one of stone crabs comprise a monophyletic group. In many ways this makes more common sense than earlier classifications which had the families of hermit crabs in two superfamilies but this is not the view of one of the world’s leading hermit crab specialists, Jacques Forest, who believed the superfamilies to have very different skeletal structure (reported in McLaughlin & Lemaitre, 2001). The family of terrestrial hermit crabs, Coenobitidae, does not occur in southern Australia. It includes the tree-climbing coconut crab, Birgus latro Linnaeus, of the islands of the Indian and Pacific oceans, the largest terrestrial invertebrate known. This species does not inhabit a shell. ‘crazy crabs’, Coenobita variabilis McCulloch, 1909, captured from the shores of tropical states are sold as pets in southern Australia. The monotypic family Pylojacquesidae is known from only one specimen collected off Brisbane, Qld, at 139 m depth (McLaughlin & Lemaitre, 2001). Two keys are presented.

Diagnosis. Body asymmetrical (rarely symmetrical). Uropodal rami not forming a tail-fan, with rasps, or absent. Pleopods 3–5 paired, unpaired or absent. Ocular scales well-developed or secondarily reduced. 6 antennal articles. 11 pairs of gills. Pereopod 1 a cheliped; pereopods 2 and 3 ambulatory, dactyli with claw; pereopod 4 ambulatory or reduced, simple, chelate or subchelate; pereopod 5 reduced.

Key 1 to families of southern Australian hermit crabs from intertidal and shelf environments 1. Left cheliped larger than right or chelipeds subequal; abdominal somite 1 distinct from last thoracic somite; usually maxillipeds 3 adjacent at base ...... Diogenidae … p. 251 — Right cheliped larger than left; abdominal somite 1 fused to last thoracic somite; usually maxillipeds 3 widely separate at base ...... Paguridae … p. 270

Key 2 to all families of hermit crabs and allies (adapted from McLaughlin & Lemaitre, 2001) 1. Pereopods 4 developed as normal walking legs; body crab-like; abdomen recurved and carried under cephalothorax ...... Lithodidae (king crabs) … p. 266

250 Anomura – hermit crabs, porcelain crabs and squat lobsters

— Pereopods 4 not developed as normal walking legs; body not crab-like; abdomen usually not recurved and carried under cephalothorax ...... 2 2. Mandible with incisor process mostly corneous, armed with prominent, acute teeth; sternites 11 and 12 distinctly separated by membranous area ...... Pylojacquesidae — Mandible with incisor process calcareous or at most with only mesial edge corneous, lacking acute teeth; sternites 11 and 12 fused ...... 3 3. Maxillipeds 3 generally widely separated basally; chelipeds unequal or less frequently subequal, right largest ...... 4 — Maxillipeds 3 generally close together basally; chelipeds equal, subequal or unequal, left frequently largest ...... 5 4. Maxilliped 1 exopod with flagellum ...... Paguridae … p. 270 — Maxilliped 1 exopod without flagellum ...... Parapaguridae … p. 279 5. Antenna with upper flagella stick-like; semiterrestrial hermit crabs ...... Coenobitidae — Antenna with upper flagella not stick-like ...... 6 6. Paired pleopods on abdominal somites 2–5; deep-sea; body straight . Pylochelidae … p. 286 — No paired pleopods on abdominal somites 4 and 5; typical hermit crab ...... Diogenidae … p. 251

References McLaughlin, P.A. 1983a. Hermit crabs – are they really polyphyletic? Journal of Crustacean Biology 3: 608–621. McLaughlin, P.A. 1983b. A review of the phylogenetic position of the Lomidae (Crustacea: Decapoda: Anomala). Journal of Crustacean Biology 3: 431–447. McLaughlin, P.A.2003. Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60: 111–144. McLaughlin, P.A., & Lemaitre, R. 2001. A new family for a new genus and new species of hermit crab of the superfamily Paguroidea (Decapoda: Anomura) and its phylogenetic implications. Journal of Crustacean Biology 21: 1062–1076. Tudge, C.C. 1995. Hermit Crabs of the Great Barrier Reef and Coastal Queensland. School of Marine Science, The University of Queensland & Backhuys Publishers: Brisbane. 40 pp.

Diogenidae Ortmann, 1892 Diogenid hermit crabs are most easily distinguished from pagurids by having the left cheliped larger than the right, or equal chelipeds. The northern Australian, deep-water genus, Tisea Morgan & Forest, 1991, which includes only one species, probably the largest of all marine hermit crabs. The Australian fauna as a whole includes 89 species in ten genera, three times as many as Paguridae (Davie, 2002). Curiously, Paguridae are more diverse on a world scale. Literature on the fauna of southern Australia is scattered but recent research by Morgan (1987b; 1989; 1991), Morgan & Jones (1991), McLaughlin & Gunn (1992), Gunn & Morgan (1992) has begun to pull this together. Morgan (1987c; 1987d; 1990) and Morgan & Forest (1991c) did the same thing for the northern fauna. Jones & Morgan’s (2002) book, and Tudge’s (1995) guide have colour photos of several species. More general taxonomic reviews of the world’s fauna are referred to under the appropriate headings. Dimensions, where given, are of the length of the shield, the anterior harder part of the carapace.

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About half of the 19 genera occur in southern Australia. Keys to genera have been offered by the renowned French carcinologist Jacques Forest whose prolific writings on anomurans have contributed so much to knowledge of this difficult group. His key (Forest, 1984) was later updated (Forest, 1995) and another key has appeared more recently (McLaughlin, 2003). Morgan (1989) provided a key to the species of south-western Australia. The first two couplets of the key are difficult and involve careful microscopic investigation under the carapace. All members of the family have ten arthrobranchs, two on the soft cuticle of each of the five thoracic somites 3–7 (carrying maxilliped 3 to pereopod 4). They also all have a pleurobranch on the hard exoskeleton above the limb on thoracic somites 5–7 (carrying pereopods 2–4) and may have one on the last thoracic somite. Put more simply, in the 14-gilled genera there is a gill above the last leg and in the 13-gilled genera there is not. Unless your hermit crab is a species of Cancellus (living in a stone), Strigopagurus (a large shelf species with stridulating medial surfaces on the ends of the chelipeds), or Dardanus (with a forked groove on the posterior part of the shield) you can safely start at couplet 4. Pereopod 1 is the cheliped, pereopods 2 and 3 the two pairs of walking legs, and pereopods 4 and 5 reduced.

Diagnosis. Abdomen well-developed, asymmetrical, terga weakly calcified, integument membranous. Chelipeds unequal or sometimes equal; if unequal left larger (except Petrochirus – not in Australia). Pleopods 2–5 unpaired, on left side only, sometimes absent in male, or pleopods 1 (or 1 and 2) paired. Abdominal somite 1 distinct from last thoracic somite. Maxillipeds 3 usually adjacent at base. Antennule flagellum elongate.

Key to southern Australian genera of Diogenidae 1. 1 pleurobranch above pereopod 5 (14 branchia on each side) ...... 2 — No pleurobranch above pereopod 5 (13 branchia on each side) ...... 4 2. Maxilla 1 with lateral flagellum on endopod; propodus and carpus of chelipeds armed with ridges of horny stridulating teeth ...... Strigopagurus — Maxilla 1 without lateral flagellum on endopod; chelipeds without stridulating apparatus . 3 3. Chelipeds equal, forming an operculum with pereopods 2 and 3; mesogastric region not marked by forked grooves; uropods equal or subequal; pleopods absent in male, 4 biramous pleopods on left in female ...... Cancellus — Chelipeds equal or unequal, not forming an operculum with pereopods 2 and 3; mesogastric region marked posteriorly by forked grooves; left uropod generally much stronger than right; 4 biramous pleopods on left, in female pleopods 2–4 endopods bifurcating ...... Dardanus 4. Pleopods 1 and 2 paired in male and forming copulatory apparatus, only 1 pair of pleopods in female, single pleopods on left; chelipeds generally equal, sometimes unequal . . . . . Paguristes — Pleopods 1 and 2 not paired; chelipeds equal or left stronger than right ...... 5 5. Front rounded in middle, not as prominent as strong laterofrontal points; median plate (sometimes reduced) between ocular scales; antenna flagellum with long setae; left cheliped stronger than right ...... Diogenes — Front more or less produced, small, but always defined, acute and exceeding weak laterofrontal points; no plate between ocular scales; antenna flagellum with short setae . . . 6 6. Chelipeds identical, exceptionally left stronger than right, surface of propodus rugose, granulose, tuberculate or spinose, claws corneous, carpus-propodus axis of articulation almost parallel to sagittal plane (vertical) ...... Clibanarius — Left cheliped generally much stronger than right, surface of left propodus often partly smooth and polished, claws calcareous, carpus-propodus axis of articulation very oblique with respect to sagittal plane ...... Calcinus

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Calcinus Dana, 1853 The calcareous, rather than corneous, black tips to the fingers of the large left cheliped immediately identify members of the genus. The genus Calcinus was reviewed and defined by Morgan (1991) who provided a list of 33 world species and key to 17 Australian species. Poupin & McLaughlin (1998) described more species and gave a key to Indo-West Pacific species. Tudge (1995) described with colour photos some species from the Great Barrier Reef. Most species are intertidal or from the shallow shelf and are tropical or subtropical but three occur in southern Australia.

Diagnosis. Rostrum small. No plate between slender ocular scales. Left cheliped much stronger than right, surface of left propodus often partly smooth and polished, claws calcareous, carpus- propodus axis of articulation very oblique, left carpus with prominent upper-lateral tubercle. Pereopod 4 chelate. Pleopods not paired. Without pleurobranch on thoracic somite 8. Maxilliped 3 with well-developed crista dentata, without accessory tooth.

Key to southern Australian species of Calcinus 1. Pereopodal 3 dactylus and distal part of propodus with brush of long setae on lower margin, much more setose than pereopod 2; left cheliped dactylus with prominent proximal lateral and mesial red spot ...... Calcinus imperialis — Pereopodal 3 dactylus and distal part of propodus without brush of long setae, only slightly more setose than, or similar to, pereopod 2; left cheliped dactylus without prominent spot 2 2. Major chela smooth laterally; pereopods 2 and 3 banded in white, red and brown, with small orange spots; telson with left and right posterior lobes each with 1 spine directed ventrally from posterior margin ...... Calcinus dapsiles — Major chela with lateral tubercles, especially on fixed finger; pereopods 2 and 3 with several short dark red-purple stripes proximally on dactylus; telson with left posterior lobe with several marginal spines, right lobe with 1 spine (rarely 0 or 2) directed ventrally from posterior margin ...... Calcinus latens Calcinus dapsiles Morgan, 1989 (Fig. 71a). Ocular scales simple. Left cheliped propodus with small lateral tubercles. Right cheliped carpus and propodus with pronounced spines on upper margin. Pereopod 3 without brush of long setae. Telson with left and right posterior lobes each with 1 spine directed ventrally from posterior margin. Shield brown or salmon-brown; pereopods 2 and 3 banded in white, red and brown, with small orange spots at setal pores. WA (150 km N of Carnarvon to E of Albany); intertidal to 97 m, rocky and coral reefs, sand, seagrass. This species is extremely common in south-western Australia. It uses a variety of gastropod shells as well as serpulid worm tubes (Morgan, 1991). Calcinus imperialis Whitelegge, 1901 (Fig. 71b). Ocular scales simple. Left cheliped propodus with 2 or 3 broad lateral depressions. Right cheliped carpus and propodus with spines on upper margin. Pereopod 3 with brush of long setae on lower margin of distal part of propodus and dactylus. Telson with left and right posterior lobes each with 1 spine directed ventrally from posterior margin. Left cheliped dactylus with proximal red patch on mesial and lateral face; pereopods 2 and 3 banded in white, red-brown and olive. Southern Great Barrier Reef, Qld, NSW (S to Sydney), islands and reefs of eastern Tasman Sea; intertidal to 2 m. Calcinus latens (Randall, 1840) (Fig. 71c). Ocular scales simple (rarely additional spinule). Left cheliped propodus smooth laterally. Right cheliped carpus and propodus carinate and spinose on upper margin. Pereopod 3 with brush of long setae only on dactylus. Telson with left posterior lobe with several marginal spines, right lobe with 1 spine (rarely 0 or 2) directed ventrally from posterior margin; pereopods 2 and 3 with dark red-purple stripe proximally

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on dactylus. Indo-West Pacific, WA (S to Rottnest I.), NT, Qld, NSW (S to Sydney); intertidal to 5 m, rocky and coral reefs, rubble and sand. The species is one of the most widespread hermit crabs in the Indo-West Pacific and has two Australian synonyms, C. terraereginae Haswell and C. abrolhensis Morgan.

Cancellus Milne Edwards, 1836 Species of Cancellus are curious hermit crabs. Unlike most others, the chelipeds, abdomen and uropods are perfectly symmetrical. This is explained by the manner in which they inhabit cavities in stones rather than shells and use the chelipeds and second pereopods as an effective operculum. Mayo (1973) reviewed the genus and diagnosed it and its nine species then known; the second Australian species is a tenth (Morgan & Forest, 1991a). Diagnosis. Chelipeds equal, forming an operculum with pereopods 2 (and rarely 3). Abdomen symmetrical. Uropods equal. Pleopods absent in male, 4 biramous pleopods on left in female. Pleurobranch on thoracic somite 8.

Key to Australian species of Cancellus 1. Pereopod 3 propodus lobate on upper margin; coxa 5 of male more or less triangular, pair barely touching in midline ...... Cancellus typus — Pereopod 3 propodus not lobate on upper margin; coxa 5 of male more or less rectangular, pair contiguous in midline ...... Cancellus quadraticoxa Cancellus quadraticoxa Morgan & Forest, 1991 miner hermit crab (Fig. 75f). Operculum formed of pereopods 1 and 2; shield with inflated frontal rim divided into 3 parts, anterior part of shield weakly tuberculate; cheliped propodus striate on lower surface; ocular scale with 2–5 terminal spines; pereopod 3 propodus with tubercles on upper margin; coxa 5 of male more or less rectangular, pair contiguous in midline. 5 mm. WA (W of Rottnest I. and Garden I.); 137–219 m. Cancellus typus Milne Edwards, 1836 miner hermit crab (Fig. 75d, e, Pl. 15a). Operculum formed of pereopods 1 and 2; shield with inflated frontal rim divided into 3 parts, anterior part of shield tuberculate; cheliped propodus striate on lower surface; ocular scale with 1 terminal spine; pereopod 3 propodus lobate on upper margin; coxa 5 of male more or less triangular, pair barely touching in midline. 10 mm. Central NSW, Vic., Tas., SA, WA (N to Cervantes); low intertidal to 88 m. The curious miner hermit crab is immediately recognised in Australia by living in a tubular cavity in a stone. Although the stone can be fist-sized the hermit is able to move it about, only at night in specimens kept in aquaria. Although it has been alleged that the hermits build or expand its burrows this has not been established and it is probable that they inhabit tubes made by boring molluscs. Hale (1941) described the young stages brooded by the adult female in its home. This species is much more widespread than C. quadraticoxa.

Clibanarius Dana, 1852 Species of this large genus are recognised by the weak rostrum, ocular scales close together, and absence of paired pleopods. The genus is widespread throughout the warmer seas of the Indo- West Pacific and Mediterranean (Miyake, 1978). Both southern Australian species feed on organic detritus, macroalgae and the associated small invertebrates (Kunze & Anderson, 1979). There are colour photos of both species in Tudge (1995). Diagnosis. Rostrum small, but always defined, acute and exceeding weak laterofrontal points. Ocular scales approximate. No transverse groove on anterior part of cardiac region. Chelipeds identical, exceptionally left stronger than right, surface of propodus ornamented or spinose, claws

254 Anomura – hermit crabs, porcelain crabs and squat lobsters

a e

f b

Fig. 71. Diogenidae. a, Calcinus dapsiles (left cheliped). b, Calcinus imperialis (left cheliped). c, Calcinus latens (left cheliped). d, Cancellus typus. Pair of coxae 5: e, Cancellus typus. f, Cancellus quadraticoxa. g, Clibanarius taeniatus.h, Clibanarius virescens (left cheliped). i, j, Dardanus arrosor (front of carapace, left cheliped). k, Dardanus australis (left cheliped).

255 Marine Decapod Crustacea of Southern Australia

corneous, carpus-propodus axis of articulation almost parallel to sagittal plane (vertical). Pereopod 4 subchelate. Pleopods 2–5 not paired in both sexes. No pleurobranch on thoracic somite 8.

Key to southern Australian species of Clibanarius 1. Shield and pereopods 2 and 3 with longitudinal stripes; cheliped carpus with spines on upper crest and laterally ...... Clibanarius taeniatus — Shield and pereopods 2 and 3 uniformly coloured; cheliped carpus with spines only on upper crest ...... Clibanarius virescens Clibanarius taeniatus (Milne Edwards, 1848) (Fig. 1g, Pl. 15b). Shield with 2 broad submedian and 2 narrower lateral blue-green bands on cream background, each band bordered in darker colour; chelipeds and pereopods blue-green, irregularly longitudinally banded. WA (S to Shark Bay), NT, Qld, NSW (S to Port Hacking); intertidal platforms and pools. The striking colouration of this species immediately distinguishes it from any other hermit crab in our region (Morgan, 1987d). The chelipeds are more spinose than in the following species. Clibanarius virescens (Krauss, 1843) (Fig. 71h). Carapace brownish; chelipeds dark with while- yellow spines and fingers; pereopods 2 and 3 blue; eyestalk with white ring around eye. Indo- West Pacific, including eastern Africa and Fiji, WA (S to Geraldton), NT, Qld, NSW (S to Sydney); intertidal and shallow subtidal. Colour may vary considerably in this species but the ring around the eye is a quick clue to its identity.

Dardanus Paul’son, 1875 Dardanus is a large genus with many large colourful species in the tropics of the Indo-West Pacific. Miyake’s (1978) key is applicable for some northern Australian species and can be supplemented with Cook’s (1989) paper. In most species the eyestalks are short and stout and the eyes well-developed. The forked groove on the posterior half of the shield immediately identifies the genus. Diagnosis. Rostrum absent. Ocular scales short with oblique and denticulate distal border. Mesogastric region with forked grooves; transverse cardiac line sometimes present. Both chelipeds with strong corneous claws, left generally stronger than right, variously ornamented. Left uropod much stronger than right. Both sexes with 4 biramous pleopods on left, in female pleopods 2–4 endopods bifurcate (whole apparently trifurcate). Pleurobranch on thoracic somite 8. Maxilla 1 without lateral flagellum on endopod.

Key to southern Australian species of Dardanus 1. Chelipeds and pereopods 2 and 3 with outer faces ornamented with transverse striae fringed with short setae ...... Dardanus arrosor — Chelipeds and pereopods 2 and 3 with outer faces not so ornamented ...... 2 2. Left pereopod 3 only (second walking leg) propodus and dactylus flattened and with tessellated margins ...... 3 — Left and right pereopods 3 similar, propodus and dactylus not flattened ...... 4 3. Carapace and pereopods mottled red and white, eyestalks and antennae uniformly pink-orange ...... Dardanus setifer — Carapace and legs reddish brown with light purple lateral edges; few scattered purple spots; 2 prominent longitudinal stripes; distinct purple patch on upper surface of carpus of chelipeds and walking legs ...... Dardanus crassimanus

256 Anomura – hermit crabs, porcelain crabs and squat lobsters

4. Chelipeds subequal ...... Dardanus australis — Left cheliped clearly larger than right ...... 5 5. Carapace and limbs orange-red with many white spots ringed with black; eyestalks reddish with small white spots ...... Dardanus megistos — Carapace mottled tan and cream, shield with median red-brown spot near anterior margin; eyestalks white with red bands ...... Dardanus pedunculatus — Carapace mottled browny-green and white with branching patterns on branchial region; eyestalks white with dark brown transverse band proximally and lighter brown band distally, cornea green ...... Dardanus deformis Dardanus arrosor (Herbst, 1796) (Fig. 71i, j, Pl. 15c). Eyestalks shorter than antennule peduncle. Left cheliped much larger than right; chelipeds and pereopods 2 and 3 with outer faces ornamented with transverse striae fringed with short setae. Red and white rings around eyes, red and white stripes on carapace and legs; chelipeds with blue-purple stripes. Mediterranean and Red seas, Indo-West Pacific, including Japan and New Zealand, throughout Australia; 2–160 m depth. Dardanus australis Forest & Morgan, 1991 (Fig. 71k). Eyestalks shorter than antennule peduncle. Chelipeds subequal; propodus covered with strong spines, less well developed mesially. Chelipeds and pereopods 2 and 3 moderately setose. South-western WA; 10–188 m depth, most 130–150 m. This species from deeper water is, at a shield length up to 36 mm, one of the largest species in the genus. Dardanus crassimanus (Milne Edwards, 1836). Eyestalks shorter than antennule peduncle. Left cheliped much larger than right; propodus with spinose upper and outer surface, each spine surrounded by short and long setae, lower margin with 2 irregular rows of long spines. Left pereopod 3 propodus and dactylus flattened and tessellated. Carapace and legs reddish brown with light purple lateral edges; few scattered purple spots; 2 prominent longitudinal stripes; distinct purple patch on upper surface of carpus of chelipeds and walking legs. Indo-West Pacific, WA (S to Rottnest I.); 10–30 m depth. The species is easily confused with D. setifer and there may be a species complex in what are currently known as these two species (Morgan, 1989). It appears that in southern Australia this is the WA species and D. setifer the NSW species. For a colour photo see Tudge (1995). Dardanus deformis (Milne Edwards, 1836). Eyestalks shorter than antennule peduncle. Left cheliped larger than right and acting as an operculum when animal is retracted, upper margin with 2 or 3 rows of spines, lower margin with single row. 7 mm. Indo-West Pacific, WA, Qld, NSW; intertidal and subtidal; associated with anemones. For a colour photo see Tudge (1995). Dardanus megistos (Herbst, 1804). Eyestalks shorter than antennule peduncle. Left cheliped much larger than right; propodus with spinose upper and outer surface, each spine surrounded by short and long setae, lower margin without specially differentiated spines. Carapace and limbs orange-red with many white spots ringed with black; red setae in the middle of the spots. Eyestalks reddish with small white spots. Indo-West Pacific, Qld, NSW, WA, SA; sublittoral reef to 10 m depth. This is a large distinctively white-spotted species, sometimes reaching 30 cm long in deep water. For a colour photo see Tudge (1995) or Jones & Morgan (2002). Dardanus pedunculatus (Herbst, 1804) (Fig. 72b, Pl. 15d). Eyestalks shorter than antennule peduncle. Left cheliped much larger than right; propodus with 2–3 longitudinal rows of tubercles on upper part of surface, smooth on lower surface. Chelipeds and pereopods 2 and 3 not setose. Carapace mottled tan and cream, shield with median red-brown spot near anterior margin. Eyestalks white with red bands. Western Pacific, Indonesia, Qld, NSW (S to Port Jackson); 10–60 m depth. This species is symbiotically associated with anemones (Ross, 1975).

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Dardanus setifer (Milne Edwards, 1836) (Fig. 72a). Eyestalks longer than antennule peduncle. Left cheliped much larger than right; propodus of left cheliped spinose, each spine on outer surface surrounded by fans of short setae, lower margin with a palisade of usually 2 lateral and 1 mesial spine. Propodus and dactylus of left pereopod 3 broadened, with longitudinal grooves and crenulate sculpture. Carapace and pereopods mottled red and white, eyestalks and antennae uniformly pink-orange. Indo-West Pacific, NT, Qld, NSW (S to Sydney); subtidal. The fans of short setae around the spines on the outer face of the left chela and the rows of short spines along the lower margin distinguish this species (in south-eastern Australia) from D. crassimanus (in south-western Australia) although colour may be more reliable (Morgan, 1987d). Kunze & Anderson (1979) discovered that, like many hermit crabs, this species feeds on macroalgae and detritus and was rarely a predator.

Fig. 72. Diogenidae. a, Dardanus setifer (from Alcock, 1905). Left cheliped: b, Dardanus pedunculatus. c, Diogenes custos.d, Diogenes lophochir.e, Diogenes senex..

258 Anomura – hermit crabs, porcelain crabs and squat lobsters

Diogenes Dana, 1851 The hinged rostrum, often multispinose, immediately identifies species of Diogenes. Most species are found in the Indo-West Pacific from the intertidal down to shelf depths. Morgan & Forest (1991a) listed several species from northern Australia. Five described species have been recognised in southern Australia. Diagnosis. Rostrum developed as moveable median plate (sometimes reduced) between ocular scales. No transverse groove on anterior part of cardiac region. Chelipeds unequal, claws corneous, carpus-propodus axis of articulation oblique. Pereopod 4 chelate. Pleopods 2–5 not paired in both sexes. No pleurobranch on thoracic somite 8.

Key to southern Australian species of Diogenes 1. Eyestalks shorter than both antennal peduncles ...... 2 — Eyestalks longer than antennal 2 peduncles ...... 4 2. Outer face of left chela with distinct curved row of tubercles ...... Diogenes lophochir — Outer face of left chela more or less evenly covered with small tubercles ...... 3 3. Antennal acicle clearly bifurcate, inner branch third to half as long as outer; carpus of left cheliped with 2 similar rows of small spines on upper edge ...... Diogenes dubius — Antennal acicle obscurely bifurcate, inner branch represent by 1 or 2 spines; carpus of left cheliped with 2 rows of spines on upper edge, outer row stronger ...... Diogenes custos 4. Chelipeds propodus obscurely tuberculate on upper edge; rostrum without ventral spine ...... Diogenes senex — Chelipeds carpus and propodus with spines on upper edge stronger than others; rostrum with ventral spine obscured in dorsal view by bifid apex ...... Diogenes serenei Diogenes custos (Fabricius, 1798) (Fig. 72c). Eyestalks shorter than antennal peduncles. Rostrum simple. Outer surface of palm of left chela with many tubercles, weaker on lower area; carpus of left cheliped with 2 rows of spines on upper edge, outer row stronger. Antennal acicle obscurely bifurcate, inner branch represent by 1 or 2 spines. 6 mm. Indo-West Pacific, Qld, NSW (S to Port Jackson); sublittoral. Many literature citations of this species throughout the Indo-West Pacific refer to other species (McLaughlin & Holthuis, 2001). These authors rediagnosed and illustrated the species. Diogenes dubius (Herbst, 1804). Eyestalks shorter than antennal peduncles. Rostrum simple. Outer surface of palm of left chela with many tubercles, weaker on lower area; carpus of left cheliped with 2 similar rows of small spines on upper edge. Antennal acicle clearly bifurcate, inner branch third to half as long as outer. Indian Ocean, Qld, NSW; sublittoral. This species has been referred to as D. miles (by Whitelegge, 1900 in Australia and by others) or D. custos (McLaughlin & Holthuis, 2001). These authors rediagnosed the species. Diogenes lophochir Morgan, 1989 (Fig. 72d). Eyestalks shorter than antennal peduncles. Rostrum simple. Chelipeds not hairy, outer face of left chela with pronounced curved row of tubercles. 4 mm. WA (Rottnest I.); 34 m depth. Diogenes senex Heller, 1865 (Fig. 72e). Eyestalks longer than antennal peduncles. Rostrum apex bifid, without ventral spine. Left cheliped carpus and propodus with upper margin rounded, with 1 or more rows of tubercles, upper outer face of propodus with 1 longitudinal row of small spines. 5 mm. NT, Qld, NSW, eastern Vic.; intertidal, especially sand beaches. Diogenes serenei Forest, 1956. Eyestalks longer than antennal peduncles. Rostrum apex bifid, obscuring strong ventral spine. Left cheliped carpus and propodus with upper margin bearing row of strong spines (especially in females), outer faces with longitudinal row of sharp

259 Marine Decapod Crustacea of Southern Australia

spines. 5 mm. Basically cream, mottled with brown. West Indian Ocean, Vietnam, New Guinea, NT, Qld, WA; sublittoral, coral and estuary.

Paguristes Dana, 1851 The genus is distributed worldwide from intertidal to 200 m depth and is best recognised by the copulatory apparatus seen on the male abdomen and paired first pleopods in the female. These are close to pereopods 5 and are hidden by a lobe on the underside of the abdomen. Most species have a prominent rostrum. Morgan’s (1987b) key has been modified here. Diagnosis. Chelipeds usually equal, sometimes unequal. Pereopod 4 not chelate. Pleopods 1 and 2 paired in male and forming copulatory apparatus, in female only pleopods 1 paired. Female often with brood pouch on left side of abdomen enclosing eggs on the pleopods and covering dorsal side. No pleurobranch above pereopod 5.

Key to Australian species of Paguristes 1. Chelipeds subequal ...... 2 — Left cheliped much larger than right ...... 8 2. Chelipeds with very long setae, not obscuring spines; pereopods 2 and 3 with long scattered setae ...... Paguristes longisetosus — Chelipeds with setae obscuring spines along upper-lateral margin; pereopods 2 and 3 with dense fringes of setae ...... 3 3. Ocular scales simple ...... Paguristes aciculus — Ocular scales with 2 or more spines ...... 4 4. Ocular scales with 2 spines ...... 5 — Ocular scales with 3 or more spines ...... 7 5. Cheliped carpus with distal rounded boss on upper surface ...... Paguristes pugil — Cheliped carpus without boss on upper surface ...... 6 6. Peduncle of antennule longer than eyestalks ...... Paguristes laurentae — Peduncle of antennule about as long as eyestalks ...... Paguristes squamosus 7. Ocular scales with about 4 apical spines; chelipeds with dense setae obscuring red-tipped spines; rostrum narrow, exceeding lateral projections ...... Paguristes sulcatus — Ocular scales multispinose; chelipeds with large acute spines; rostrum broad, shorter than lateral projections ...... Paguristes brevirostris 8. Rostrum short, not exceeding lateral projections; left cheliped propodus grossly tuberculate ...... Paguristes tuberculatus — Rostrum long, exceeding lateral projections; left cheliped propodus finely tuberculate . . . 9 9. Right cheliped dactylus with different-sized tubercles and spines; pereopods salmon with scattered red spots; antennae 2 red-orange ...... Paguristes frontalis — Right cheliped dactylus with similarly-sized tubercles; pereopods cream-orange with dense red speckles; antennae 2 purple ...... Paguristes purpureantennatus Paguristes aciculus Grant, 1905 (Fig. 73a). Rostrum narrow, exceeding lateral projections. Ocular scales simple. Antennule peduncles longer than eyestalks. Chelipeds equal; carpus and propodus covered on upper surface with solid spines. Pereopods 2 and 3 with dactyli longer than propodi. Telson posterior lobes with small teeth. Colour unknown. 6 mm. NSW, Vic., Tas.; 360 m depth.

260 Anomura – hermit crabs, porcelain crabs and squat lobsters

a c b

f d

Fig. 73. Diogenidae. a, Paguristes aciculus.b, Paguristes brevirostris.c, d, Paguristes frontalis.e, Paguristes laurentae. f, Paguristes frontalis (coxae 5 and gonopods 1 and 2). Left cheliped: g, Paguristes pugil.h, Paguristes purpureantennatus.i, Paguristes squamosus.j, Paguristes sulcatus. k, Paguristes tuberculatus.

261 Marine Decapod Crustacea of Southern Australia

Paguristes brevirostris Baker, 1905 (Fig. 73b, Pl. 15f, g). Rostrum broad not exceeding lateral projections. Ocular scales with 7–8 apical spinules, separate. Antennule peduncles longer than eyestalks. Chelipeds subequal; propodi and dactyli covered with large acute spines on lateral and upper surfaces. Pereopods 2 and 3 with dactyli longer than propodi. Telson posterior lobes with about 6 small lateral spines. Chelipeds and legs pale orange; eyestalks red with blue cornea. 8 mm. SA Gulfs, western Tas.; to 29 m depth. Paguristes frontalis (Milne Edwards, 1836) (Fig. 73c, d, f, Pl. 15e). Rostrum narrow, exceeding lateral projections. Ocular scales simple, approximate distally. Antennule peduncles as long or shorter than eyestalks. Chelipeds very unequal; left propodi and dactyli and right dactylus covered with uneven spines and tubercles; propodi with weak lateral longitudinal ridge. Pereopods 2 and 3 with dactyli longer than propodi. Telson posterior lobes unarmed. Shield, pereopods and antennae salmon, only pereopods with few red spots at setal pores. 25 mm. Vic. (E to Wilsons Promontory), SA, WA (W to Cape Naturaliste); common in intertidal to shallow subtidal, rocks and seagrass. This species is large, brightly coloured and common. It is one of several in the genus in which the glaucothoe larval stage hatches from the egg and undergoes abbreviated development inside the gastropod shell of its parent (Morgan, 1987a). Paguristes laurentae Morgan & Forest, 1991 (Fig. 73e). Rostrum broad not exceeding lateral projections. Ocular scales with 2–3 apical spinules, separate. Antennule peduncles longer than eyestalks. Chelipeds subequal; propodi and dactyli covered with large acute spines on lateral and upper surfaces. Pereopods 2 and 3 with dactyli longer than propodi. Telson posterior lobes with 2–3 very large spines. Cheliped marbled orange-red, pereopods 2 and 3 with red stripes. 4.6 mm. WA (Point Cloates to Rottnest I.); 134–146 m. There is an undescribed south- eastern deep-water species which keys, but is not this species . Paguristes longisetosus Morgan, 1987. Rostrum narrow, exceeding lateral projections. Ocular scales multispinose. Antennule peduncles as long or shorter than eyestalks. Chelipeds subequal and spinose. Chelipeds, pereopods and tail-fan with long setae not obscuring spines. Telson posterior lobes with marginal spines. Brown with cream or white spines, scattered red patches. 7.3 mm. WA (Hopetoun to Albany); shallow subtidal, sand. Paguristes pugil McCulloch, 1913 (Fig. 73g). Rostrum narrow, exceeding lateral projections. Ocular scales with 2 apical spinules, separate. Antennule peduncles shorter than eyestalks. Chelipeds subequal; carpus with rounded distal boss on upper surface. Pereopods 2 and 3 fringed with dense setae. Telson posterior lobes with 4 short spines. 6 mm. NSW (Port Jackson), Tas.,Vic., Bass Strait; 6–69 m depth. Paguristes purpureantennatus Morgan, 1987 (Fig. 73h). Rostrum narrow, exceeding lateral projections. Ocular scales simple, approximate distally. Antennule peduncles as long or shorter than eyestalks. Chelipeds very unequal; left propodi and dactyli and right dactylus covered with similarly-sized tubercles; propodi with distinct lateral longitudinal ridge. Pereopods 2 and 3 with dactyli longer than propodi. Telson posterior lobes unarmed. Shield bright red, pereopods with red flecks, antennae purple. 22 mm. WA (Dongara to Albany); subtidal to 10 m, rocks and seagrass. This species is best separated from P. frontalis on the colour of the antennae; the two co-occur at Albany. Paguristes squamosus McCulloch, 1913 (Fig. 73i). Rostrum narrow, exceeding lateral projections. Ocular scales with 2 apical spinules. Antennule peduncles equal to eyestalks. Chelipeds subequal; setose laterally, squamous, carpus with 4 spines on upper surface. Pereopods 2 and 3 fringed with dense setae. Telson lobes with long hooked spines. 11 mm. Pink tinged with green and brown. NSW (Sydney region), eastern Vic. Tas.; intertidal rock platforms. Kunze & Anderson (1979) found that filamentous and coralline algae, and dead crustaceans and molluscs were collected by the chelipeds for feeding. The dense tomentum of setae on the chelipeds and pereopods may enable particulate filter feeding.

262 Anomura – hermit crabs, porcelain crabs and squat lobsters

Paguristes sulcatus Baker, 1905 (Fig. 73j). Rostrum narrow, exceeding lateral projections. Ocular scales with about 4 apical spinules. Antennule peduncles longer than eyestalks. Chelipeds subequal; dense setae obscuring red-tipped spines. Pereopods 2 and 3 fringed with dense setae. Telson lobes with small spines. Generally cream. 17 mm.Vic. (E to Wilson Promontory), Tas., SA, WA (N to Rottnest I.); subtidal to 25 m, rocks. Morgan & Forest (1991b) confirmed the name of this species. Densities of more than 10 m–2 have been recorded for South Australia (Mower & Shepherd, 1988). Paguristes tuberculatus Whitelegge, 1900 (Fig. 73k). Rostrum broad not exceeding lateral projections. Ocular scales simple acute spines. Antennule peduncles longer than eyestalks. Chelipeds unequal; left propodus with strong tuberculate projections on outer surface. Pereopods 2 and 3 with dactyli longer than propodi. Telson posterior lobes spinose. 5 mm. Colour unknown. NSW (N to Port Macquarie), Vic., SA, Tas., WA (N to Perth); shallow subtidal, sand and seagrass. Many specimens of this small species are associated with colonies of encrusting bryozoans which surround the mouth of the shell in which the animal lives.

Strigopagurus Forest, 1995 For many years one large abundant species was known from the shelf of the southern coast – Trizopagurus strigimanus. Forest erected a new genus for five species, one in northern Australia in addition to the two below, previously confused as T. strigimanus (Forest, 1995). The strong stridulating apparatus formed from rows of smooth plates on the facing surfaces of the chelipeds gives the genus away in Australia. An abbreviated key is given to the two similar species which appear to not overlap geographically. Diagnosis. Ocular scales not toothed. Chelipeds similar, left slightly larger than right, propodus and carpus armed with strong teeth, each with sharp horny point, facing surfaces of propodus and dactylus ornamented with rows of corneous striae forming a stridulating organ. Rostrum broad, weakly produced. Shield convex; mesogastric region usually marked by lateral grooves which meet posteriorly as sharp angle and continue as median groove; cardiac region smooth. Walking legs with dactyli longer than propodi. Usually a pair of pleopods (pleopods 2) and 3 single pleopods on left side in male; 4 unpaired pleopods in female, the last similar to the others. Pleurobranch present on thoracic somite 8. Maxilla 1 with lateral flagellum on endopod. Abdominal somites 2–5 with chitinous terga.

Key to southern Australian species of Strigopagurus 1. Eyestalks with red stripe between mesial and lateral white borders; cheliped and pereopods with red spots at ends ...... Strigopagurus elongatus — Eyestalk completely red except for white ring around eye; cheliped and pereopods without red spots ...... Strigopagurus strigimanus Strigopagurus elongatus Forest, 1995 (Fig. 74c, Pl. 16a). Eyestalks with red stripe between mesial and lateral white borders. Cheliped and pereopods with red spots at ends. Male with only left pleopod 2 present. 26 mm. Vic. (E to Cape Otway), SA, WA (N to Mandurah); ?85–220 m. Colour pattern of the eyes is the distinguishing feature of this species. Strigopagurus strigimanus (White, 1847) (Fig. 74a, b, Pl. 16b). Eyestalk completely red except for white ring around eye. Cheliped and pereopods without red spots. Male with pleopod 2 paired. 37 mm. NSW (N to Newcastle), Vic. (W to Melbourne), Tas.; 3–130 m. This large species is extremely common on the shelf of south-eastern Australia, much more common than T. elongatus. There are some very subtle morphological differences between the two species but colour and the male pleopods are the most reliable. This is a large hermit crab and has sometimes been offered for sale in fish markets in Melbourne.

263 Marine Decapod Crustacea of Southern Australia

b c

Fig. 74. Diogenidae. a, Strigopagurus strigimanus. Eyestalk: b, Strigopagurus strigimanus. c, Strigopagurus elongatus.

References Cook, S.D. 1989. Dardanus imbricatus (H. Milne Edwards) and descriptions of three new species of Dardanus (Decapoda, Anomura, Diogenidae). Memoirs of the Queensland Museum 27: 111–122. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda - Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Forest, J. 1984. Révision du genre Aniculus Decapoda Diogenidae. Crustaceana Supplement 8: 1–91. Forest, J. 1995. Crustacea Decapoda Anomura: révision du genre Trizopagurus Forest, 1952 (Diogenidae), avec l’établissement de deux genres nouveaux. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 13. Mémoires du Muséum National d’Histoire Naturelle, Paris 163: 9–149. Gunn, S.W., & Morgan, G.J. 1992. A new species of Pagurixus (Crustacea: Decapoda: Paguridae) from southern Australia. Memoirs of the Museum of Victoria 53: 31–41. Hale, H.M. 1941. Decapod Crustacea. British, Australian and New Zealand Antarctic Research Expedition, 1929-1931. Reports-Series B (Zoology and Botany) 4: 257–286, pl. 3. Jones, D.S., & Morgan, G.J. 2002. A Field Guide to Crustaceans of Australian Waters. Reed New Holland: Sydney. 224 pp. 2nd edn. Kunze, J., & Anderson, D.T. 1979. Functional morphology of the mouthparts and gastric mill in the hermit crabs Clibanarius taeniatus (Milne Edwards), Clibanarius virescens (Krauss), Paguristes squamosus (McCulloch) and Dardanus setifer (Milne Edwards) (Anomura: Paguridae). Australian Journal of Marine and Freshwater Research 30: 683–722. Mayo, B.S. 1973. A review of the genus Cancellus (Crustacea: Diogenidae) with the description of a new species from the Caribbean Sea. Smithsonian Contributions to Zoology 150: 1–63. McLaughlin, P.A.,& Gunn, S.W. 1992. Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species. Part IV. Lophopagurus McLaughlin and Australeremus McLaughlin. Memoirs of the Museum of Victoria 53: 43–99.

264 Anomura – hermit crabs, porcelain crabs and squat lobsters

McLaughlin, P.A.2003. Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60: 111–144. McLaughlin, P.A., & Holthuis, L.B. 2001. In pursuit of J.F.W. Herbst’s species of Diogenes (Anomura: Paguridea: Diogenidae). Journal of Crustacean Biology 21: 249-265. Miyake, S. 1978. The Crustacean Anomura of Sagami Bay collected by His Majesty the Emperor of Japan. Biological Laboratory, Imperial Household: Tokyo. 161 pp. Morgan, G.J. 1987a. Abbreviated development in Paguristes frontalis (Milne Edwards, 1836) (Anomura: Diogenidae) from southern Australia. Journal of Crustacean Biology 7: 536–540. Morgan, G.J. 1987b. Two new species of Paguristes (Decapoda: Anomura: Diogenidae) from southwestern Australia. Proceedings of the Biological Society of Washington 100: 726–734. Morgan, G.J. 1987c. A new and aberrant species of Paguristes (Anomura: Diogenidae) from northern Australia. Records of the Western Australian Museum 13: 379–386. Morgan, G.J. 1987d. Hermit crabs (Decapoda, Anomura: Coenobitidae, Diogenidae, Paguridae) of Darwin and Port Essington, Northern Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 4: 165–186. Morgan, G.J. 1989. The hermit crabs (Decapoda: Anomura: Diogenidae, Paguridae) of southwestern Australia, with descriptions of two new species. Records of the Western Australian Museum 14: 391–417. Morgan, G.J. 1990. A collection of Thalassinidea, Anomura and Brachyura (Crustacea: Decapoda) from the Kimberley region of northwestern Australia. Zoologische Verhandelingen, Leiden 265: 1–90. Morgan, G.J. 1991. A review of the hermit crab genus Calcinus Dana (Crustacea: Decapoda: Diogenidae) from Australia, with descriptions of two new species. Invertebrate Taxonomy 5: 869–913. Morgan, G.J., & Forest, J. 1991a. Seven new species of hermit crabs from Northern and Western Australia (Decapoda, Anomura, Diogenidae). Bulletin du Muséum National d’Histoire Naturelle, Paris A, 12: 649-689. Morgan, G.J., & Forest, J. 1991b. Recognition of the Paguristes species P. tomentosus (H. Milne Edwards, 1848) and P. sulcatus Baker, 1905 (Decapoda, Anomura, Diogenidae). Crustaceana 60: 108–111. Morgan, G.J., & Forest, J. 1991c. A new genus and species of hermit crab (Crustacea, Anomura, Diogenidae) from the Timor Sea, north Australia. Bulletin du Muséum National d’Histoire Naturelle, Paris 13: 189–202. Morgan, G.J., & Jones, D.S. 1991. Checklist of marine decapod Crustacea of southern Western Australia. Pp. 483–497 in: Wells, F.E., Walker, D.I., Kirkman, H., & Lethbridge, R. (eds), Proceedings of the Third International Marine Biological Workshop: The Marine Flora and Fauna of Albany, Western Australia. Western Australian Museum: Perth. Mower, A.G.J., & Shepherd, S.A. 1988. The crab fauna of West Island, South Australia: their abundance, diet and role as predators of abalone. Transactions of the Royal Society of South Australia 112: 83–86. Poupin, J., & McLaughlin, P.A. 1998. Additional records of Calcinus species (Decapoda: Anomura: Diogenidae) from French Polynesia with description of three new species and a key to Indo-West Pacific species of the genus. Crustacean Research 27: 9–27. Ross, D.M. 1975. The behavior of pagurids in symbiotic associations withh actinians in Japan. Publications of the Seto Marine Biological Laboratory 22: 157–170. Tudge, C.C. 1995. Hermit Crabs of the Great Barrier Reef and Coastal Queensland. School of Marine Science, The University of Queensland & Backhuys Publishers: Brisbane. 40 pp. Whitelegge, T. 1900. Scientific results of the trawling expedition of H.M.C.S. ‘Thetis’ off the coast of New South Wales in February and March, 1898. Crustacea. Part I. Memoirs of the Australian Museum 4: 135-199, pls 32–35.

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Lithodidae Samouelle, 1819 The name of the family can be translated to mean stone crabs and this is often what they are called. The name, king crab, which is applied to the large commercial species might be a better ‘common name’. However, apart from these northern hemisphere species such as the Alaskan King Crab, Paralithodes camtschatica, which reach Australia only frozen or in cans, none is ‘common’ here. Only one local species can be considered large enough to be called a ‘king crab’. Lithodids differ from all other paguroids in being crab-like rather than hermit crabs, not inhabiting gastropod shells and having a calcified flattened abdomen. The species differ from brachyuran crabs in having only three pairs of walking legs and an asymmetrical abdomen and pleopods. Many species, especially in the subfamily Lithodinae, are confined to deep water and this is true of all southern Australian species. Because of the high commercial value of a least one Northern Pacific species and the potential value of others the family has attracted a lot of attention. It has been the special interest of Elliot Dawson of Wellington, New Zealand, and his bibliography (Dawson, 1989) listed some 3850 papers, notes, newspaper articles and other printed material dealing with everything known at that time about lithodids. All 95 described species in 16 genera were listed by Dawson (1989: appendix 1) but about a dozen have been described since by Macpherson in several papers. The subfamily Haplogastrinae (five genera, nine species) is intertidal and immediate subtidal only in the Northern Pacific. Most species are in the Lithodinae and are from slope depths and beyond except in polar and subpolar seas. Macpherson’s (1988; 1990) studies of the faunas of the Atlantic and Pacific oceans are the most recent definitive works on the systematics of the family; the first includes a key to genera. An unpublished manuscript prepared by Dawson & Yaldwyn for the International King Crab Symposium held in Anchorage in 1985 also contains a key to genera, key to species of Lithodes, and a list of species (Dawson & Yaldwyn, 1985). Lithodes murrayi Henderson, 1888 is common off Macquarie Island and throughout the subantarctic and there are at least five other species in south-eastern Australia. The type localities of all the species names used are not in Australia and identification of Australian specimens is provisional. Macpherson (1990) described some species from seas near Australia and these might be expected to occur in northern Australian waters.

Diagnosis. Crab-like. Maxilliped 1 exopod with flagellum. Maxillipeds 3 widely separate at base. Epistomal spines absent. Chelipeds equal or subequal. Pereopod 4 ambulatory. Abdomen asymmetrical in females, symmetrical in males, calcified, abdominal terga with supplemental calcified plates lateral to paired plates; abdominal somite 1 fused to last thoracomere. Uropod absent. Pleopods 2–5 unpaired.

Key to southern Australian species of Lithodidae 1. Sternite between pereopods 2 (first walking legs) without deep longitudinal median groove, shallow depression instead (carapace generally evenly spinose) ...... Paralomis … 2 — Sternite between pereopods 2 (first walking legs) with deep longitudinal median groove (carapace with mixture of long and short spines) ...... 4 2. Carapace and legs sparsely tuberculate and spinose ...... Paralomis cf. birsteini — Carapace and legs densely covered with long sharp spines ...... 3 3. Median gastric spine stronger than others ...... Paralomis cf. phryxa — No median gastric spine stronger than others ...... Paralomis cf. histrix 4. Abdominal somite 2 of 3 plates (with long tapering rostrum bearing 2 apical and 2 dorsal spines) ...... Lithodes longispina

266 Anomura – hermit crabs, porcelain crabs and squat lobsters

Fig. 75. Lithodidae. Neolithodes brodiei, a, habitus (right limbs only). b, abdominal somite 2. c, abdominal somites 3–7.

267 Marine Decapod Crustacea of Southern Australia

— Abdominal somite 2 of 5 plates, 1 median, 1 lateral pair and 1 marginal pair; abdominal somites 3–5 comprising small spiniform nodules; males without well defined plates, females with well developed left lateral plates (rostrum short, conical and with pair of lateral basal tubercles, carapace otherwise with strong and weak spination) ...... Neolithodes brodiei

Lithodes Latreille, 1806 Macpherson (1988, 1990) gave a more complete diagnosis of the genus and listed the 16 species known. Six species occur in the western Pacific including Lithodes murrayi Henderson, 1888 near Macquarie Island. Diagnosis. Carapace armed with spines and granules, varying in size. Rostrum normally bifid, with 1 or 2 pairs of dorsal spines and strong curved basal spine. Sternal region with a deep median groove between first walking legs. Abdominal somite 2 of median plate fused with pair of lateral plates and pair of marginal plates. Abdominal somites 3–5 with calcified nodules in place of median plates. Lateral and marginal plates clearly differentiated in both sexes. Lateral plates in females more developed on left side than on right and fused with marginal plates. Lithodes longispina Sakai, 1971(Fig. 76a, Pl. 16c). Long bifid rostrum, about 0.7 carapace length, with pair of dorsal spines near its base; carapace with 5–7 pairs of long dorsal spines plus marginal spines; walking legs covered on upper surfaces by spinules. 15 cm carapace length. Japan, western Pacific, Qld, NSW, Vic., Tas. (including seamounts), SA; 900–1200 m depth. Lithodes longispina resembles but has longer carapace spines than L. richeri Macpherson, 1990 from New Caledonia (Macpherson, 1990), a species that might also occur in south-eastern Australia (Macpherson, 2001). Dorsal carapace spines and marginal spines can be highly variable in number and size.

Neolithodes Milne Edwards & Bouvier, 1894 Macpherson (1988) listed nine species scattered through all oceans. Diagnosis. Carapace armed with spines and granules, varying in size. Rostrum formed by a basal spine slanting slightly upwards and a pair of divergent upwardly directed dorsal spines. Sternal region with a deep median groove between first walking legs. Abdominal somite 2 of 5 plates, 1 median plate, pair of lateral plates and pair of marginal plates. Abdominal somites 3–5 with small spiniform nodules, Plates not well differentiated in males. A lateral plate in females more developed on left side than on right. Neolithodes brodiei Dawson & Yaldwyn, 1970 (Fig. 75). Carapace compact, gastric region promoted anteriorly, well delimited from cardiac and branchial regions; rostrum short, with pair of stout oblique basal spines and another pair of smaller basal spines; branchial region with 10 major dorsal and 15 major lateral spines on each side; walking legs elongate and flattened. cl. 130 mm. New Zealand, Vanuatu, Tas., SA; 800–1180 m depth. The species was first well described and illustrated by McLay (1988). The species was originally described from a single specimen taken at 832 m depth off southern New Zealand (Dawson & Yaldwyn, 1970). It is common between the Tasmanian Seamounts, eastern Tas., and the central Great Australian Bight where it is often taken by deep-water trawlers. Its known depth range reflects the activity of the trawlers seeking blue grenadier. It is recorded as far north as Vanuatu (Macpherson, 2001).

268 Anomura – hermit crabs, porcelain crabs and squat lobsters

Paralomis White, 1856 With 48 species throughout the world’s slopes (Macpherson, 1988, 1992; de Saint Laurent & Macpherson, 1997; Macpherson, 2001) this is the biggest and most variable lithodid genus but no described species has been reported in the literature from Australia (Davie, 2002). Nevertheless, at least three species have been found in deep water off south-eastern Australia but identification with known species is provisional. This key will differentiate them but is not diagnostic. See too Macpherson (2003). Diagnosis. Carapace armed with short spines and granules. Rostrum formed by a basal spine and at least 1 pair of divergent upwardly directed dorsal spines. Sternal region without a deep median groove between first walking legs. Abdominal somite 2 of a single plate. Abdominal somites 3–5 each with median plate and pairs of lateral and marginal plates clearly differentiated in both sexes. Paralomis cf. birsteini Macpherson, 1988 (Fig. 76b). Carapace and legs sparsely tuberculate and spinose; rostrum with strong ventral spine. cl. 80 mm. Ross Sea, Tasmanian Seamounts; 1100–2000 m depth. Paralomis cf. phrixa Macpherson, 1992 (Fig. 76c). Carapace and legs densely covered with long sharp spines, median gastric spine stronger than others; rostrum with basal ventral spinule. cl. 105 mm. Tasmanian Seamounts; 987 m depth. Paralomis cf. histrix (De Haan, 1844) (Fig. 76e). Carapace and legs densely covered with long sharp spines, median gastric spine not differentiated from others; rostrum with rudimentary basal ventral spinules. cl. 105 mm. Japan, New Zealand, Vic.; 640 m depth. This Japanese species appears in Sakai’s (1976) key to species. Yamaguchi & Baba (1993) confirmed the spelling and date of publication of this species.

b c a

d e

Fig. 76. Lithodidae. Dorsal carapace. a, Lithodes longispina. b, Paralomis cf. birsteini. c, Paralomis cf. phrixa. Lateral anterior carapace. d, Paralomis cf. birsteini. e, Paralomis cf. histrix.

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References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda - Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Dawson, E.W. 1989. King Crabs of the World (Crustacea: Lithodidae) and their fisheries: a comprehensive bibliography. New Zealand Oceanographic Institute Miscellaneous Publication 101: 1–338. Dawson, E.W., & Yaldwyn, J.C. 1970. Diagnosis of a new species of Neolithodes (Crustacea: Anomura: Lithodidae) from New Zealand (note). New Zealand Journal of Marine and Freshwater Research 4: 227–228. Dawson, E.W., & Yaldwyn, J.C. 1985. King Crabs of the world or the world of King Crabs: an overview of identity and distribution -with illustrated diagnostic keys to the genera of the Lithodidae and to the species of Lithodes. Pp. 69–106 in: Proceedings of the International King Crab Symposium, Anchorage, Alaska, 22–24 Jan 1985. Macpherson, E. 1988. Revision of the family Lithodidae Samouelle, 1819 (Crustacea, Decapoda, Anomura) in the Atlantic Ocean. Monografías de Zoología Marina 2: 9–153. Macpherson, E. 1990. Crustacea Decapoda: on some species of Lithodidae from the Western Pacific. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 6. Mémoires du Muséum National d’Histoire Naturelle, Paris 145: 217–226. Macpherson, E. 1992. Paralomis phrixa (Decapoda, Anomura, Lithodidae), a new species from northern Peru, and a key to the eastern Pacific species of the genus. Crustaceana 63: 313-317. Macpherson, E. 2001. New species and new records of lithodid crabs (Crustacea, Decapoda) from the southwestern and central Pacific Ocean. Zoosystema 23: 797-805. Macpherson, E. 2003. Some lithodid crabs (Crustacea: Decapoda: Lithodidae) from the Solomon Islands (SW Pacific Ocean), with the descriptions of a new species. Scientia Marina 67: 413–418. McLay, C.L. 1988. Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22: 1–403. de Saint Laurent, M., & Macpherson, E. 1997. Une nouvelle espèce du genre Paralomis White, 1856, des sources hydrothermales du Sud-ouest Pacifique (Crustacea, Decapoda, Lithodidae). Zoosystema 19: 721–727. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates. Yamaguchi, T., & Baba, K. 1993. Crustacean specimens collected in Japan by Ph. F. von Siebold an H. Bürger and held by the National Natuurhistorisch Museum in Leiden and other museums. Pp. 145–570 in: Yamaguchi, T. (ed.) Ph. F. von Siebold and Natural History of Japan, Crustacea. The Carcinological Society of Japan: Tokyo.

Paguridae Latreille, 1802 Hermit crabs are immediately recognisable from their use of gastropod shells for protection of the soft abdomen. Although many biologists are familiar with the use of the word ‘pagurid’ to describe hermit crabs, most of the commonly encountered species on the southern coast are not members of the family from which this name is derived – most are diogenids. The easiest way to differentiate the two shallow-water families is to compare the chelipeds. Species with the right hand larger than the left are members of the Paguridae, species with the left bigger than the right or of the same size are members of the Diogenidae. This rule does not apply to the deep-water species which may belong to Parapaguridae or Pomatochelidae. Most pagurid species recorded from southern Australia are from deep water. The curious relationship between hermit crabs and shells has been investigated many times (Hazlett, 1981). Hermit crabs adopt numerous feeding strategies, most species are not especially fussy. Schembri’s (1982) study of fifteen species in New Zealand found deposit-feeding, scaveng- ing, predation and filter-feeding to be options for many species. The Paguridae have the reputation of being one of the most difficult of all decapod families for the general taxonomist and consequently have attracted few specialists in recent years. The

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complex literature dealing with named genera and species, the enormous number of undescribed species especially in the tropical Indo-West Pacific and, until recently, the absence of a recent synthesis of almost 80 genera are all discouraging. Luckily, there are few representatives in shallow water of southern Australia but exploration of the richer unworked fauna of northern Australia and of the deep sea will require a good knowledge of the literature as many of the species described from South-East Asia or other deep oceans may occur here. Present-day systematics of the family builds on the works of Henderson (1888) and Alcock (1905). Gordan (1956) presented a checklist of species and comprehensive bibliography but the modern work of M. de Saint Laurent, J. Forest, P. McLaughlin, G. Morgan and R. Lemaitre are the most useful. Most of the papers by these authors are limited in scope. McLaughlin’s (1997; 2003a) keys and generic diagnoses have provided a way into the literature but more genera have appeared since. Keys to the Japanese genera (Miyake, 1978) may be useful for the fauna of northern Australia which has been little studied. Grant & McCulloch (1906), McCulloch (1913), Lewinsohn (1978), Morgan (1987; 1990; 1993), Haig & Ball (1988), McLaughlin & Haig (1989), Jones (1990) and McLaughlin (1994) between them listed only about a dozen northern species not found in southern Australia. De Saint Laurent (1966) divided the family into three groups based largely on the number of thoracic gills, a scheme that may be no longer appropriate. The 13-gilled Pylopaguropsis-group (with 2 arthrobranchs each on maxilliped 3–pereopod 4, pleurobranch on pereopods 2–4) comprises few genera of which the only named Australian species are members of Pylopaguropsis and Propagurus. The 10-gilled Ostraconotus-group (with 2 arthrobranchs each on maxilliped 3–pereopod 4, no pleurobranchs) was revised by de Saint Laurent in a series of six papers concluding in 1970 (de Saint Laurent, 1970). Undescribed Australian species of this group occur and they are common in the Indonesian archipelago. The 11-gilled Pagurus-group (with 2 arthrobranchs each on maxilliped 3–pereopod 4, pleurobranch on pereopod 4) comprises the majority of genera. A subgroup in which females possess paired first pleopods was reviewed in a series of papers beginning with McLaughlin (1981) and of which McLaughlin & Gunn (1992) is the most relevant to Australia. Dimensions are given as shield length (sl.). Pereopod 1 is the cheliped, pereopods 2 and 3 the two pairs of walking legs, and pereopods 4 and 5 reduced. Diagnosis. Abdomen well-developed, asymmetrical (rarely symmetrical), integument membranous. Chelipeds unequal, right larger than left. Usually 3 (male) or 4 (female) biramous pleopods on left side only, sometimes pleopods 1 paired in male or female, rarely pleopods 1 and 2 paired in male. Abdominal somite 1 distinct from last thoracic somite. Maxillipeds 3 separate at base. Antennule flagellum elongate.

Key to southern Australian genera of Paguridae 1. Abdomen reduced, symmetrical; living in bivalve shell ...... Porcellanopagurus — Abdomen typically large and twisted ...... 2 2. Pleurobranchs on pereopods 2–4 (13 pairs) ...... 3 — Pleurobranchs on pereopod 4 only (11 pairs) ...... 5 3. Chelipeds subequal, left sometimes longer but not stronger; pereopod 4 rasp with 1 row of scales ...... Bythiopagurus — Right cheliped stronger than left; pereopod 4 rasp with 1 or 2 or more rows of scales . . . . 4 4. Right cheliped massive, chela flat, almost circular and operculate ...... Bathypaguropsis — Right cheliped elongate, cylindrical, covered with rows of spikes ...... Propagurus

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5. Females with paired pleopods 1 modified as gonopods ...... Lophopagurus — Females without modified paired pleopods 1 ...... 6 6. Right chelipeds sexually dimorphic, larger and smooth in male; right male gonopore on coxa 5 obscured by medially directed tuft of long setae; rostrum prominent ...... Pagurixus — Right cheliped not sexually dimorphic; right male gonopore without tuft of setae; rostrum usually weak ...... 7 7. Right cheliped palm elongate, carpus with prominently produced denticulate lower margin ...... Goreopagurus — Right cheliped not elongate, lower carpal margin not produced ...... 8 8. Left cheliped somewhat flattened; ocular scale multispinose; left male gonopore on coxa 5 extended as a curved sexual tube ...... Micropagurus — Left cheliped not flattened; ocular scale triangular or multispinose; male gonopores simple apertures or with short sexual tube on coxa 5 ...... Pagurus

Bathypaguropsis McLaughlin, 1994 Bathypaguropsis is a deep-water genus immediately recognised by the dominating circular right chela. It may be confused with some parapagurids. Four species are known, two on the eastern coast of Australia. Diagnosis. Ocular scale simple. Carapace shield well-calcified, posterior part membranous. Maxilliped 3 ischium with 1 accessory tooth. Chelipeds grossly unequal, right larger, chela flattened, almost circular and operculate. Pereopod 4 propodus with rasp of 1 or more rows of scales. Male with simple paired gonopores. Male and female with unpaired left pleopods 2–5. Uropods asymmetrical. Telson with transverse suture, margin with 2 oblique lobes. 13 quadriserial gills on each side (2 arthrobranchs on maxilliped 3–pereopod 4; pleurobranch on thoracic somites 5–7, above pereopods 2–4). Bathypaguropsis yaldwyni McLaughlin, 1994 (Fig. 77a). Palm of right chela with 3 stout spaced spines and cluster of spines distally along upper margin (not a continuous row of spines or smooth). sl. 12 mm. New Zealand, Vic., Tas. including seamounts; 117–620 m depth.

Bythiopagurus McLaughlin, 2003 Bythiopagurus is known only from the species from Tasmanian Seamounts (McLaughlin, 2003b). Diagnosis. Ocular scale triangular. Carapace shield well-calcified. Maxilliped 3 ischium with 1 or 2 accessory teeth. Chelipeds subequal, left longer. Pereopod 4 propodus with rasp of 1 row of scales. Male with paired gonopores, each with papilla or short sexual tube. Male with paired pleopods 1 and 2, plus 3 unpaired left pleopods; female with 4 unpaired biramous pleopods. Uropods asymmetrical. Telson with transverse suture, margin with 2 lobes. 13 quadriserial gills on each side (2 arthrobranchs on maxilliped 3–pereopod 4; pleurobranch on thoracic somites 5–7, above pereopods 2–4). Bythiopagurus macrocolus McLaughlin, 2003 (Fig. 77b). sl. 6 mm. Tasmanian Seamounts; 1083–1300 m depth. The rows of regular spines along the upper and lower margins and on the face of the right cheliped characterise this deep-water species.

Goreopagurus McLaughlin, 1988 This deep-water genus of three species was recognised in Australia after specimens belonging to a new species were first placed in Michelopagurus (Lemaitre & McLaughlin, 2003a).

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g e d j h

m l

n o

Fig. 77. Paguridae. a, Bathypaguropsis yaldwyni (right cheliped). b, Bythiopagurus macrocolus (right cheliped). c, Goreopagurus poorei (cheliped). d, e, Lophopagurus nanus (left cheliped, and in cross- section). f, Lophopagurus triserratus (right cheliped). Micropagurus acantholepis (g, front; h, left view of male pereopods 4 and 5 showing sexual tube). Telson: i, Pagurixus amsa. j, Pagurixus granulimanus. k, Pagurixus jerviensis. Pagurixus jerviensis (l, front; m, male right cheliped; n, female right cheliped; o, coxa 5). p, Pagurixus handrecki (right cheliped). q, Pagurixus granulimanus (right cheliped).

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Diagnosis. Ocular scale triangular with submarginal spine. Maxilliped 3 ischium with accessory tooth. Chelipeds unequal. Right cheliped with elongate chela, carpus produced ventrally as blade. Pereopod 4 propodus with rasp of 1 row of spines; dactylus with or without preungual process. Male with simple gonopores or with prominent sexual tube on right gonopore, shorter or papilla on left. Male with 3 unpaired biramous pleopods, female with paired pleopods 1 and unpaired biramous pleopods 2–5. Uropods asymmetrical. Telson with transverse suture, margin with 2 lobes. 11 phyllobranch gills on each side. Goreopagurus poorei Lemaitre & McLaughlin, 2003 (Fig. 77c). Male gonopores without sexual tubes. Gills quadriserial. 5 mm. Tas. (slope and seamounts); 500–1300 m depth. The right cheliped has a broad carpus with strong toothed ridges on the upper and lower margins, and elongate palm with short fingers.

Lophopagurus McLaughlin, 1981 This genus of 13 small species from shelf depths in New Zealand and Australia now includes species previously put in Australoremus (McLaughlin & Gunn, 1992; de Saint Laurent & McLaughlin, 2000; Lemaitre & McLaughlin, 2003b). Females are characterised by a pair of gonopods, pleopods 1 modified as small cylindrical limbs that lie close to and between coxa 5. The gonopods are much smaller than the unpaired pleopods and difficult to see in such small animals. Diagnosis. Maxilliped 3 ischium with accessory tooth. Chelipeds unequal or subequal. Right cheliped with propodus-carpus articulation more or less rotated from horizontal plane. Pereopod 4 propodus with rasp of 1 row of spines. Male with simple paired gonopores. Male with 3 unpaired biramous pleopods, female with paired pleopods 1 modified as gonopods, and unpaired biramous pleopods 2–5 (endopod on 5 reduced). Uropods asymmetrical. Telson with transverse suture, margin with 2 lobes. 11 phyllobranch biserial gills on each side. Lophopagurus nanus (Henderson, 1888) (Fig. 77d, e, Pl. 16d). Left cheliped with strong convex lateral ridge on propodus, propodus triangular in cross-section and scarcely rotated with respect to carpus, carpus with strong lateral spines. 5 mm. Southern Qld, NSW, Vic., Tas., SA; 14–128 m depth. The species is especially common in Bass Strait. It belongs in the subgenus Lophopagurus. Lophopagurus triserratus (Ortmann, 1892) (Fig. 77f). Left cheliped with flat lateral surface on propodus, propodus strongly twisted with respect to carpus, carpus without lateral spines. 4 mm. West Pacific, New Zealand, Vic. (Bass Strait); often in serpulid polychaete tubes, 60–400 m depth. The species is placed in the subgenus Australoremus McLaughlin, 1981.

Micropagurus McLaughlin, 1986 The unusual sexual tube at the base of the left fifth leg distinguishes this genus which is confined to four Pacific species. There is only one species in southern Australia; it occurs in shallow water. It has multispinose ocular scales. Diagnosis. Ocular scale multispinose. Pair of lobes between ocular scales. Maxilliped 3 ischium with accessory tooth. Chelipeds unequal, right larger. Left cheliped somewhat flattened. Pereopod 4 propodus with rasp of 1–3 rows of spines; dactylus without preungual process. Male with sexual tube on left coxa 5, with or without gonopore on right. Male with 3 unpaired biramous pleopods, female with 4 unpaired biramous pleopods 2–5. Uropods asymmetrical. Telson without transverse suture, margin entire. 11 phyllobranch gills on each side. Micropagurus acantholepis (Stimpson, 1858) (Fig. 77g, h, Pl. 16e). With right male gonopore. Pereopods not dorsally armed. sl. 4 mm. Red-brown mottled on background of yellow.

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Southern NSW, Vic., Tas., SA; 0–43 m depth. The multispinose ocular scales with a pair of small lobes between distinguish this species from all other southern Australian pagurids; the sexual tube of the male is especially obvious (Tudge & Lemaitre, 2004).

Pagurixus Melin, 1939 McLaughlin & Haig (1984) provided a diagnosis and keyed all the species then known. Now there are 12 species in the genus which is confined to the Indo-West Pacific. Seven named species occur in Australia and undescribed ones exist in deeper water. It is the most common pagurid genus on southern shores. Three species may be found together at Rottnest I., WA (Morgan, 1993). The key is adapted from those in the works mentioned. Diagnosis. Ocular scale triangular with small submarginal spine. Maxilliped 3 ischium with accessory tooth. Chelipeds unequal, right larger. Right cheliped sexually dimorphic, dorsal surface of chela smooth and palm often swollen or elongate in adult male, smaller in female. Left cheliped subtriangular in cross-section, armed. Pereopod 4 propodus with rasp of 1 row of spines; dactylus without preungual process. Male with right gonopore obscured by tuft of long mesially-directed setae. Male with 3 unpaired biramous pleopods, female with 4 unpaired biramous pleopods. Uropods asymmetrical. Telson with transverse suture, margin with 2 symmetrical lobes. 11 phyllobranch gills on each side.

Key to southern Australian species of Pagurixus 1. Right male cheliped palm with mesial ridge running between articulation with carpus and base of dactylus for about two-thirds of length ...... Pagurixus handrecki — Right male cheliped palm with mesial ridge only proximal or absent ...... 2 2. Right male cheliped palm coarsely tuberculate, especially mesially . Pagurixus granulimanus — Right male cheliped palm minutely granular or smooth ...... 3 3. Pereopods 2 and 3 dactyli 4.0–4.5 times as long as broad, much shorter than propodi; telson with narrow shallow median cleft ...... Pagurixus jerviensis — Pereopods 2 and 3 dactyli 5.5–6.5 times as long as broad, as long as propodi; telson with broad deep median cleft ...... Pagurixus amsa Pagurixus amsa Morgan, 1993 (Fig. 77i). Right male cheliped palm minutely granular. Pereopods 2 and 3 dactyli 5.5–6.5 times as long as broad, as long as propodi. Telson with broad deep median cleft, lobes oblique. Shield cream with brown mottling; chelipeds salmon-pink; pereopods 2 and 3 banded in cream and salmon-brown, with longitudinal red-brown stripes. sl. 2.7 mm. Southern WA (Albany to Shark Bay); subtidal to 20 m; rocks. Pagurixus granulimanus Morgan, 1993 (Fig. 77j, q). Right male cheliped palm with acute granules. Pereopods 2 and 3 dactyli 6 times as long as broad, as long as propodus. Telson with strong median cleft, lobes oblique. Shield cream with brown mottling; chelipeds pale orange- pink; pereopods 2 and 3 cream, with longitudinal red stripes. sl. 3.4 mm. WA (Rottnest I.); subtidal to 9 m; rocks. Pagurixus handrecki Gunn & Morgan, 1992 (Fig. 77p, Pl. 16f). Right male cheliped palm with mesial ridge for about two-thirds of length. Pereopods 2 and 3 dactyli 6 times as long as broad, as long as propodi. Telson with strong median cleft, lobes almost straight. Shield cream, often with 2 red submedian spots; chelipeds cream-brown; pereopods 2 and 3 with longitudinal red-brown stripes. sl. 3.4 mm. Vic. (W of Lakes Entrance), Tas., SA, WA (N to Rottnest I.); intertidal to 42 m; rocks. Pagurixus jerviensis McLaughlin & Haig, 1984 (Fig. 77k, l, m, n). Right male cheliped palm with mesial ridge only proximally. Pereopods 2 and 3 dactyli 4 times as long as broad, shorter than

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propodi. Telson with narrow shallow median cleft, lobes straight. Shield orange tinted; chelipeds orange; pereopods 2 and 3 with longitudinal stripes and diffuse transverse bands. sl. 4.3 mm. Central NSW, Vic., SA, southern WA; intertidal to 22 m. This is one of the most common species of hermit crab on open coastal reefs in southern NSW and in some older books was referred to as Pagurus lacertosus. It can be distinguished from P. handrecki by its larger size, poorer development of the mesial ridge on the right male cheliped, and colour.

Pagurus Fabricius, 1775 The genus Pagurus is one of the most loosely defined, referred to as a ‘catch-all’ by McLaughlin (1997). She provided a modern diagnosis in 2003. About 170 species are known. The many North Atlantic species were reviewed by McLaughlin (1974) in a work which is a useful source for information on the anatomy and systematics of the family. She recognised eight species-groups, some of which now have separate generic status. In much of the older literature the name Eupagurus was used but this is no longer valid. Its species are often species of Pagurus but because there are so many recent genera this cannot be guaranteed. There are several described species known from northern Australia and several, possibly undescribed, from south-eastern slope depths. Two species supposedly recorded from the Sydney region, P. filholi (de Man, 1887) and P. minutus Hess, 1865, are unlikely to exist in this region (Sandberg & McLaughlin, 1993). Diagnosis. Ocular scale triangular or multifid. Carapace shield well-calcified, posterior part membranous. Maxilliped 3 ischium with accessory tooth. Chelipeds unequal, right much larger. Pereopod 4 propodus with well-developed rasp of spines. Male with simple paired gonopores. Male with 3 or 4 unpaired biramous pleopods, female with 4 unpaired biramous pleopods. Uropods asymmetrical. Telson with transverse suture, margin with 2 lobes. 11 phyllobranch gills on each side. Pagurus sinuatus (Stimpson, 1858) (Fig. 78a, Pl. 16g). Upper surface of chelipeds and pereopods 2 and 3 hairy, with plumose setae. Orange and red with red-violet patches; pereopods banded. 14 mm. Sydney and south coast (NSW), Vic., SA, Bremer Bay to Shark Bay (WA); intertidal to 20 m, exposed rocky localities. This is a large, very active species usually living in globose shells. The solid hairy chelipeds immediately identify it. The diagnosis is sufficient only to distinguish the species from P. hirtimanus Miers, 1880 from Queensland, New Guinea and elsewhere. What appears to be another species, Pagurus janitor Alcock, 1905 (Fig. 78b), originally described from the Maldives, has been taken from the shore at Margaret River, WA. It is only half the size and has a much more rounded propodus and dactylus on the larger cheliped than P. sinuatus.

Porcellanopagurus Filhol, 1885 This is an unusual genus of four species in the Pacific. The short squat abdomen enables the animal to carry one shell of a bivalve rather than inhabit a gastropod as do most hermit crabs (Miyake, 1978). The eggs are carried on top. There is one Australian species described in detail by its author (Whitelegge, 1900). Diagnosis. Chelipeds unequal, right larger, dactylus oblique. Male with simple paired gonopores. Male without pleopods, female with 3 unpaired biramous pleopods. Abdomen much reduced, heart-shaped; uropods and telson symmetrical. 11 phyllobranch gills on each side. Porcellanopagurus tridentatus Whitelegge, 1900 (Fig. 78d, Pl. 16h). New Caledonia, Kermadec Is, NSW (Sydney to Eden); 100–628 m depth. The broad carapace, about 7 mm long, shorter symmetrical abdomen, and weak cheliped make this species unmistakable.

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Fig. 78. Paguridae. Right cheliped: a, Pagurus sinuatus. b, Pagurus cf. janitor. c, Propagurus deprofundis. d, Porcellanopagurus tridentatus. Distal articles of walking leg 2: e, Propagurus deprofundis. f, Propagurus haigae.

Propagurus McLaughlin & de Saint Laurent, 1998 Propagurus (and Bythiopagurus) are recognised by the asymmetrical, ‘quadriserial’ gill structure. This gill structure is a modified phyllobranchiate gill in which each of the pairs of lamella bears a small outer lobe. McLaughlin & de Saint Laurent (1998) provided detailed descriptions and a key to the four species of which two occur in southern Australia. Diagnosis. Ocular scale simple. Carapace shield well-calcified, posterior part membranous. Maxilliped 3 ischium with 1 accessory tooth. Chelipeds unequal, right larger. Pereopod 4 propodus with well-developed rasp of 2 or more rows of scales. Male with simple paired gonopores. Male usually with 3 unpaired left pleopods; female with 4 unpaired biramous pleopods. Uropods asymmetrical. Telson with transverse suture, margin with 2 lobes. 13 quadriserial gills on each side (2 arthrobranchs on maxilliped 3–pereopod 4; pleurobranch on thoracic somites 5–7, above pereopods 2–4).

Key to southern Australian species of Propagurus 1. Pereopod 2 (first walking leg) with propodus with longitudinal keel on mesial face; lateral face of palm of chelipeds with rows of tubercles ...... Propagurus deprofundis — Pereopod 2 (first walking leg) with propodus without longitudinal keel on mesial face; lateral face of palm of chelipeds without rows of tubercles ...... Propagurus haigae Propagurus haigae (McLaughlin, 1997) (Fig. 78f). Pereopod 2 propodus without longitudinal keel on mesial face; lateral face of palm of chelipeds without rows of tubercles. sl. 9 mm. Indonesia, New Caledonia, Qld, NSW, Vic., Tas.; 265–660 m depth. Propagurus deprofundis (Stebbing, 1924) (Fig. 78c, e). Pereopod 2 propodus with longitudinal keel on mesial face; lateral face of palm of chelipeds with rows of tubercles. sl. 17 mm. South

277 Marine Decapod Crustacea of Southern Australia

Africa, Philippines, Hawaii, New Zealand, slope of NSW, Vic., Tas. (including seamounts); 200–1100 m depth.

References Alcock, A. 1905. Catalogue of the Indian Decapod Crustacea in the Collection of the Indian Museum. Part II. Anomura. Fasciculus I. Pagurides. Trustees of the Indian Museum: Calcutta. 193, 15 pls pp. Gordan, J. 1956. A bibliography of pagurid crabs, exclusive of Alcock, 1905. Bulletin of the American Museum of Natural History 108: 253–352. Grant, F.E., & McCulloch, A.R. 1906. On a collection of Crustacea from the Port Curtis district, Queensland. Proceedings of the Linnean Society of New South Wales 1906: 2–53, pls1-4. Haig, J., & Ball, E.E. 1988. Hermit crabs from north Australian and eastern Indonesian waters (Crustacea Decapoda: Anomura: Paguroidea) collected during the 1975 Alpha Helix Expedition. Records of the Australian Museum 40: 151–196. Hazlett, B.A. 1981. The behavioural ecology of hermit crabs. Annual Review of Ecology and Systematics 12: 1–22. Henderson, J.R. 1888. Report on the Anomura collected by H.M.S. Challenger during the years 1873–76. ‘Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76.’ Zoology 27: 1–221, 21 pls. Jones, D.S. 1990. Annotated checklist of marine decapod Crustacea from Shark Bay, Western Australia. Pp. 169-208 in: Berry, P.B., Bradshaw, S.D., & Wilson, B.R. (eds), Research in Shark Bay: Report of the France-Australe Bicentenary Expedition Committee. Western Australian Museum: Perth. Lemaitre, R., & McLaughlin, P.A.2003a. New species of Goreopagurus (Decapoda: Anomura: Paguridae) from Tasmania and reevaluation of sexual tubes in hermit crab systematics. Memoirs of Museum Victoria 60: 221–227. Lemaitre, R., & McLaughlin, P.A. 2003b. Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species. Addendum and taxonomic summary. Proceedings of the Biological Society of Washington 116: 464–486. Lewinsohn, C. 1978. Bemerkungen zur taxonomie von Paguritta harmsi (Gordon) (Crustacea Decaopoda, Anomura) und beschreibung einer neuen art der gleichen gattung aus Australien. Zoological Miscellany 53: 243–252. McCulloch, A.R. 1913. Studies in Australian Crustacea. No. 3. Records of the Australian Museum 9: 321–353. McLaughlin, P.A. 1974. The hermit crabs (Crustacea Decapoda, Paguridea) of northwestern North America. Zoologische Verhandelingen, Leiden 130: 1–396. McLaughlin, P.A. 1981. Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with the descriptions of new genera and species. Part I. Ten new genera of the Paguridae and a redescription of Tomopagurus A. Milne Edwards and Bouvier. Bulletin of Marine Science 31: 1–30. McLaughlin, P.A. 1994. A new genus and two new species of deep-water hermit crabs (Decapoda: Anomura: Paguridae) from the Southern Ocean. Proceedings of the Biological Society of Washington 107: 469–481. McLaughlin, P.A. 1997. Crustacea Decapoda: hermit crabs of the family Paguridae from the KARUBAR cruise in Indonesia. In: Crosnier, A and Bouchet, P.(eds), Résultats des Campagnes MUSORSTOM, vol. 16. Mémoires du Muséum National d’Histoire Naturelle, Paris 172: 433–572. McLaughlin, P.A. 2003a. Illustrated keys to families and genera of the superfamily Paguroidea (Crustacea: Decapoda: Anomura), with diagnoses of genera of Paguridae. Memoirs of Museum Victoria 60: 111–144. McLaughlin, P.A. 2003b. A new genus and species of hermit crab (Decapoda: Anomura: Paguridae) from seamounts off south-eastern Tasmania, Australia. Memoirs of Museum Victoria 60: 229–236. McLaughlin, P.A.,& Gunn, S.W. 1992. Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species. Part IV. Lophopagurus McLaughlin and Australeremus McLaughlin. Memoirs of the Museum of Victoria 53: 43–99.

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McLaughlin, P.A.,& Haig, J. 1984. A review of Pagurixus (Decapoda, Anomura, Paguridae) and descriptions of new species. Crustaceana 47: 121–148. McLaughlin, P.A., & Haig, J. 1989. On the status of Pylopaguropsis zebra (Henderson), P. magnimanus (Henderson), and Galapagurus teevanus Boone, with descriptions of seven new species of Pylopaguropsis (Crustacea: Anomura: Paguridae). Micronesica 22: 123–171. McLaughlin, P.A., & de Saint Laurent, M. 1998. A new genus for four species of hermit crabs formerly assigned to the genus Pagurus Fabricius (Decapoda: Anomura: Paguridae). Proceedings of the Biological Society of Washington 111: 158–187. Miyake, S. 1978. The Crustacean Anomura of Sagami Bay collected by His Majesty the Emperor of Japan. Biological Laboratory, Imperial Household: Tokyo. 161 pp. Morgan, G.J. 1987. Hermit crabs (Decapoda, Anomura: Coenobitidae, Diogenidae, Paguridae) of Darwin and Port Essington, Northern Australia. The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 4: 165–186. Morgan, G.J. 1990. A collection of Thalassinidea, Anomura and Brachyura (Crustacea: Decapoda) from the Kimberley region of northwestern Australia. Zoologische Verhandelingen, Leiden 265: 1–90. Morgan, G.J. 1993. Three new species of Pagurixus (Crustacea, Decapoda, Paguridae) from Western Australia, with notes on other Australian species. Pp. 163–181 in: Wells, F.E.,Walker, D.I., Kirkman, H., & Lethbridge, R. (eds), The marine flora and fauna of Rottnest Island, Western Australia. Western Australian Museum and Australian Marine Sciences Association: Perth. de Saint Laurent, M. 1970. Révision des genres Catapaguroides et Cestopagurus et description de quatre genres nouveau. V. Trichopagurus de Saint Laurent (Crustacés Décapodes Paguridae). VI. Conclusion. Bulletin du Muséum National d’Histoire Naturelle, Paris 42: 210–222. de Saint Laurent, M., & McLaughlin, P.A.2000. Superfamily Paguroidea, family Paguridae. In: Forest, J., de Saint Laurent, M., McLaughlin, P.A., and Lemaitre, R. The marine fauna of New Zealand: Paguridea (Decapoda: Anomura) exclusive of the Lithodidae. NIWA Biodiversity Memoir 114: 104–209. de Saint Laurent-Dechancé, M. 1966. Iridopagurus, genre nouveau de Paguridae (Crustacés Décapodes) des mers tropicales américaines. Bulletin du Muséum National d’Histoire Naturelle, Paris 38: 151–173. Sandberg, L., & McLaughlin, P.A. 1993. Reexamination of Pagurus minutus Hess, 1865, and Pagurus filholi (De Man, 1887) (Crustacea: Anomura: Paguridae). Zoologische Mededelingen, Leiden 67: 197–206. Schembri, P.J. 1982. Feeding behaviour of fifteen species of hermit crabs (Crustacea: Decapoda: Anomura) from the Otago region, southeastern New Zealand. Journal of Natural History 16: 859–878. Tudge, C.C., & Lemaitre, R. 2004. Studeis of male sexual tubes in hermit crabs (Crustacea, Decapoda, Anomura, Paguroidea). I. Morphology of the sexual tube in Micropagurus acantholepis (Stimpson, 1858), with comments on function and evolution. Journal of Morphology 259: 106–118. Whitelegge, T. 1900. Scientific results of the trawling expedition of H.M.C.S. ‘Thetis’ off the coast of New South Wales in February and March, 1898. Crustacea. Part I. Memoirs of the Australian Museum 4: 135–199, pls 32–35.

Parapaguridae Smith, 1882 Parapagurids are unusual hermit crabs from deep water, recorded from shelf to hadal depths (5000 m) where they may be extraordinarily common in trawl samples. They are especially conspicuous on continental slopes. The unusual association between parapagurids and anemones takes a step further than the oft-cited symbiosis between shore hermits which carry a solitary anemone on their shell homes. In parapagurids the home is often made only of a cavity in a flat gelatinous colony of a dozen or more anemones or zoanthids (Fautin, 1987). The protective corona of anthozoans may as the Reverend Stebbing said ‘contribute to the commissariat by throwing out its darts

279 Marine Decapod Crustacea of Southern Australia

as some swift gliding shrimp passes by, and thus reducing it to a condition in which it may be captured by the pagurid’. The systematics of the Parapaguridae was treated by de Saint Laurent (1972) and updated by Lemaitre (1996; 1999) who recognised and keyed ten genera for more than 40 species worldwide. Five genera and about half of the world’s species are known from the Australian region. Eleven species have been recognised in southern Australian waters but many species are widespread so more might be expected, including those reported from more northern Australia. Pereopod 1 is the cheliped, pereopods 2 and 3 the two pairs of walking legs, and pereopods 4 and 5 reduced.

Diagnosis. Pleurobranch only on pereopod 4 (rarely rudimentary on pereopod 5); branchial lamellae divided into two lobes, entire or subdivided. Abdomen well-developed, asymmetrical, integument membranous. Telson entire, without transverse suture. Chelipeds very unequal, right larger than left. Male sometimes with paired pleopods 1 or 2 or both; pleopods 3–5 on left only, exopod short. Female pleopods 2–4 or 2–5 on left only. Maxillipeds 3 separate at base. Epistome with 0–2 median spines.

Fig. 79. Parapaguridae. Parapagurus bouvieri in zoanthid shelter.

Key to southern Australian genera of Parapaguridae 1. Shield distinctly broader than long; dactyli of ambulatory legs straight or curved only at end; cornea strongly dilated; males with pleopod 2 having short exopod and strongly twisted distal article (Fig. 80f) ...... Strobopagurus — Shield about as broad as long; dactyli of ambulatory legs usually evenly curved; cornea moderately or weakly dilated; males with pleopod 2 lacking exopod and distal article not twisted (Fig. 80e) ...... 2

280 Anomura – hermit crabs, porcelain crabs and squat lobsters

2. Vestigial pleurobranch present on last thoracic somite (above pereopod 5) (Fig. 80d) ...... Sympagurus — Vestigial pleurobranch absent on last thoracic somite (above pereopod 5) ...... 3 3. Epistomal spine strongly curved upwards ...... Oncopagurus — Epistomal spine straight (Fig. 80b) or absent ...... 4 4. Bi- or quadriserial gills; antennule peduncle with article 4 armed with dorsodistal spine (Fig. 80c); eyestalks at least as half as long as shield ...... Paragiopagurus — Quadriserial gills; antennule peduncle with article 4 unarmed; eyestalks less than half as long as shield (except P. bouvieri) ...... Parapagurus

d e

a f

Fig. 80. Parapaguridae. a, Paragiopagurus diogenes (shield, eyes and ocular acicles). b, Paragiopagurus diogenes (illustrating straight epistome). c, Paragiopagurus diogenes (antenna 1 peduncles). d, Sympagurus sp. (vestigial pleurobranch on last thoracic somite). e, male pleopod 2, most taxa. f, Strobopagurus sibogae. (male pleopod 2).

Oncopagurus Lemaitre, 1996 Lemaitre (1996) recognised ten species all having the characteristic curved epistomal spine. Besides the three from southern Australia is another reaching tropical Western Australia. All are very small species. Diagnosis. 11 pairs of quadriserial gills only. Shield about as broad as long. Eyestalks weakly dilated. Antennule peduncle with article 4 armed. Epistomal spine strongly curved upwards. Dactyli of ambulatory legs evenly curved. Males with pleopod 2 with distal article not twisted.

Key to southern Australian species of Oncopagurus 1. Cornea cone-shaped ...... Oncopagurus minutus — Cornea not cone-shaped ...... 2 2. Male with paired first gonopods; cheliped of female with mesial face of right palm expanded distally and with row of spines delimited longitudinally and obliquely Oncopagurus indicus — Male without paired first gonopods; cheliped of female and male with mesial face of right palm not expanded distally and with row of spines delimited longitudinally ...... Oncopagurus cidaris

281 Marine Decapod Crustacea of Southern Australia

Oncopagurus cidaris Lemaitre, 1996 (Fig. 81a). Cornea round. Male without paired first gonopods. Cheliped of female and male with mesial face of right palm not expanded distally and with row of spines delimited longitudinally. 3 mm. Qld, NSW (S to 35°S); 439–1150 m depth. Inhabiting gastropod shells. Oncopagurus indicus (Alcock, 1905) (Fig. 81b, c). Cornea round. Male with paired first gonopods. Cheliped of female with mesial face of right palm expanded distally and with row of spines delimited longitudinally and obliquely. 4 mm. Indo-West Pacific, Hawaii, WA, Qld, NSW (S to Newcastle); 183–1480 m depth. Inhabiting gastropod shells, occasionally with anthozoans attached. Oncopagurus minutus (Henderson, 1896) (Fig. 81d). Cornea cone-shaped. Male with paired first gonopods. Cheliped mesial face of right palm rounded, with few scattered tubercles and with row of tubercles delimited longitudinally. 4 mm. Indo-West Pacific, NSW (off Newcastle); 800–2308 m depth. Probably inhabiting gastropod shells.

Paragiopagurus Lemaitre, 1996 Lemaitre (1996) transferred 16 species from Sympagurus to his new genus including six Australian species. Only one, the type species, is from southern Australia. Diagnosis. 11 pairs of bi- or quadriserial gills. Shield about as broad as long. Eyestalks weakly to moderately dilated. Antennule peduncle with article 4 armed with dorsodistal spine. Epistomal spine straight or absent. Dactyli of ambulatory legs evenly curved. Males with pleopod 2 with distal article not twisted. Paragiopagurus diogenes (Whitelegge, 1900) (Fig. 81e, Pl. 17a). Ocular acicles simple. Right chela without transverse furrows on ventral face of chela or carpus, with well-delimited upper- lateral margin. Female with strongly oblique (45°) palm. Male with paired pleopods 1.11 mm. Chelipeds white, iridescent, body generally orange or reddish, legs with darker red band on carpi. Inhabiting shells. Western Pacific, WA (S to south-west coast), Qld, NSW (to eastern Bass Strait); 40–695 m. This is the commonest and most shallowly recorded parapagurid in southern Australia. Its iridescent chelipeds are characteristic.

Parapagurus Smith, 1879 Lemaitre (1989, 1999) reviewed Parapagurus, a genus of 17 species found worldwide from midshelf depths down to 5000 m. Members of the genus are characterised by gills consisting of series of four filamentous or flattened branches arranged along the axis, reduced eyestalks and corneae, elongated antennule and antenna, and rounded mesial and lateral faces of the right chelipeds. Three species occur in southern Australia. The key is adapted from Lemaitre’s (1999) larger one for Indo-West Pacific species. Diagnosis. 11 pairs of quadriserial gills. Eyestalk less than half as long as shield (except P. bouvieri), cornea weakly dilated. Antennule peduncle with article 4 unarmed. Epistomal spine short if present. Dactyli of ambulatory legs evenly curved. Males with pleopod 2 with distal article slightly twisted.

Key to southern Australian species of Parapagurus 1. Walking legs with lateral and mesial faces of meri weakly calcified, indicated by brown patches; eyestalks longer than half shield length ...... Parapagurus bouvieri — Walking legs with lateral faces of meri well calcified medially; eyestalks half or less shield length ...... 2 2. Pereopod 4 propodal rasp of 1 row of ovate contiguous scales (sometimes more rows proximally) ...... Parapagurus richeri — Pereopod 4 propodal rasp of 2 rows of conical separate scales ...... Parapagurus latimanus

282 Anomura – hermit crabs, porcelain crabs and squat lobsters

b a

e f

k h

g l

Fig. 81. Parapaguridae. Right cheliped. a, Oncopagurus cidaris. b, c, Oncopagurus indicus (mesial and outer face). d, Oncopagurus minutus. e, Paragiopagurus diogenes. f, Parapagurus bouvieri. g, Parapagurus bouvieri (walking leg 1). Walking leg 4, propodus and dactylus with detail of spination. h, i, Parapagurus bouvieri. j, k, Parapagurus latimanus. l, Parapagurus richeri.

283 Marine Decapod Crustacea of Southern Australia

Parapagurus bouvieri Stebbing, 1910 (Figs 79, 81f, g, h, i, Pl. 17b). Walking legs with lateral and mesial faces of meri weakly calcified, with brown patches. Eyestalks longer than half shield length. Pereopod 4 propodal rasp of 2 or 3 rows of lanceolate scales. 15 mm. South-eastern Atlantic, Indo-West Pacific, Qld, NSW, SA; 247–990 m. Inhabiting zoanthid shelters, probably Epizoanthus sp. Parapagurus latimanus Henderson, 1888 (Fig. 81j k). Walking legs with lateral faces of meri well calcified medially. Eyestalks half shield length. Pereopod 4 propodal rasp of 2 often irregular rows of conical separate scales. 16 mm. Indo-West Pacific, New Zealand, NSW, Tas., SA; 400–2500 m. Inhabiting zoanthid shelters, probably Epizoanthus sp. Parapagurus richeri Lemaitre, 1999 (Fig. 81l). Walking legs with lateral faces of meri well calcified medially. Eyestalks half shield length. Pereopod 4 propodal rasp of 1 row of ovate contiguous scales (sometimes more rows proximally). 12 mm. Indo-West Pacific, Tasman Sea, Qld, NSW (S to 35°S); 311–4470 m. Inhabiting gastropod shells, often with anthozoan; occasionally scaphopod shells.

Strobopagurus Lemaitre, 1989 Strobopagurus was diagnosed by Lemaitre (1996). Of its three species one occurs in southern Australia. Diagnosis. 11 pairs of bi- or quadriserial gills. Shield distinctly broader than long. Eyestalks stout, cornea strongly dilated. Antennule peduncle with article 4 unarmed. Epistomal spine absent. Dactyli of ambulatory legs curved only at end. Males with pleopod 2 having short exopod and strongly twisted distal article. Strobopagurus sibogae (de Saint Laurent, 1972) (Fig. 82a). Right chela with strong acute marginal spines on propodus and dactylus. 8 mm. Western Pacific from Japan, China to Australia, Qld, NSW (S to Sydney); 40–550 m.

Sympagurus Smith, 1883 Sympagurus and several species were diagnosed by Lemaitre (1996, 2004); there are 17 species of which four occur in southern Australia. Diagnosis. 11 pairs of trichobranchiate gills plus 1 pair of vestigial gills (small conical lobes) on last thoracic somite. Shield as broad as long. Eyestalks weakly dilated. Antennule peduncle with article 4 unarmed or weakly so. Epistomal spine straight or absent. Dactyli of ambulatory legs evenly curved. Males with pleopod 2 with distal article not twisted.

Key to southern Australian species of Sympagurus 1. Pereopod 4 propodal rasp with 1 row of scales ...... Sympagurus soela — Pereopod 4 propodal rasp with 2 or more rows of scales ...... 2 2. Antennule article 4 armed with dorsodistal spine; chelipeds tuberculate ...... Sympagurus dimorphus — Antennule article 4 unarmed; chelipeds mostly smooth ...... 3 3. Ambulatory legs with numerous long stiff setae on upper margins of 4 last segments; upper margin of chelae with numerous stiff setae plus mat of short setae . . . . Sympagurus villosus — Ambulatory legs with few short setae on upper margins of 4 last segments; upper margin of chelae with only mat of short setae ...... Sympagurus burkenroadi Sympagurus burkenroadi Thompson, 1943 (Fig. 82c, f). Chelipeds dissimilar, right massive, smooth or barely tuberculate on most surfaces. Ambulatory legs with few short setae on upper

284 Anomura – hermit crabs, porcelain crabs and squat lobsters

a b

e c

Fig. 82. Parapaguridae. Right cheliped: a, Strobopagurus sibogae. b, Sympagurus dimorphus (female, male is narrower). c, Sympagurus burkenroadi. d, Sympagurus soela. Pereopod 4 propodus and dactylus: e, Sympagurus dimorphus. f, Sympagurus burkenroadi. g, Sympagurus soela.

margins of 4 last segments; upper margin of chelae with only mat of short setae. Pereopod 4 propodal rasp with 2–3 rows of scales. Antennule article 4 unarmed. 17 mm. Indo-West Pacific, NT, Qld, NSW (S to southern coast); 205–960 m depth. Inhabiting colonies of zoanthid, Epizoanthus sp. S. pappossus Lemaitre, 1996 is a junior synonym (Lemaitre, 2004). Sympagurus dimorphus (Studer, 1883) (Fig. 82b, e, Pl. 17c). Chelipeds massive in female, narrow in male, tuberculate on most surfaces. Pereopod 4 propodal rasp with 2–5 rows of scales. Antennule article 4 armed with dorsodistal spine. 16 mm. Southern Hemisphere, 25°S–57°S,

285 Marine Decapod Crustacea of Southern Australia

southern NSW, Tas.; 91–1995 m depth. Inhabiting gastropod shells or colonies of zoanthid, Epizoanthus sp. Lemaitre (1996) redescribed the species and Lemaitre & McLaughlin (1992) described the adults and larval stages. Sympagurus soela Lemaitre, 1996 (Fig. 82d, g). Chelipeds dissimilar, right elongate, setose and tuberculate on most surfaces. Pereopod 4 propodal rasp with 1 row of scales. Antennule article 4 usually unarmed. 6.5 mm. Qld, NSW (S to central coast); 274–704 m depth. Inhabiting gastropod shells. Sympagurus villosus Lemaitre, 1996. Chelipeds dissimilar, right massive, smooth or barely tuberculate on most surfaces. Ambulatory legs with numerous long stiff setae on upper margins of 4 last segments; upper margin of chelae with numerous stiff setae plus mat of short setae. Pereopod 4 propodal rasp with 2–3 rows of scales. Antennule article 4 unarmed. 14 mm. Qld, Tas.; 490–1100 m depth. Inhabiting colonies of zoanthid, Epizoanthus sp.

References Fautin, D.G. 1987. Stylobates loisetteae, a new species of shell-forming sea anemone (Ceolenterata: Aciniidae) from western Australia. Proceedings of the California Academy of Science 45:1–7. Lemaitre, R. 1989. Revision of the genus Parapagurus (Anomura: Paguroidea: Parapaguridae), including redescriptions of the Western Atlantic species. Zoologische Verhandelingen, Leiden 253: 1–106. Lemaitre, R. 1996. Hermit crabs of the family Parapaguridae (Crustacea: Decapoda: Anomura) from Australia: species of Strobopagurus Lemaitre, 1989, Sympagurus Smith, 1883 and two new genera. Records of the Australian Museum 48: 163–221. Lemaitre, R. 1999. Crustacea Decapoda: a review of the species of the genus Parapagurus Smith, 1879 (Parapaguridae) from the Pacific and Indian Oceans. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180: 303–378. Lemaitre, R. 2004. A worldwide review of hermit crab species of the genus Sympagurus Smith, 1883 (Crustacea: Decapoda: Parapaguridae). In: Marshall, B., and Richer de Forges, B. (eds), Tropical Deep-Sea Benthos, Vol. 23. Mémoires du Muséum National d'Histoire Naturelle, Paris 191: 85–149. Lemaitre, R., & McLaughlin, P.A. 1992. Descriptions of megalopa and juveniles of Sympagurus dimorphus (Studer, 1883), with an account of the Parapaguridae (Crustacea: Anomura: Paguroidea) from Antarctic and subantarctic waters. Journal of Natural History 26: 745–768. de Saint Laurent, M. 1972. Sur la famille des Parapaguridae Smith, 1882. Description de Typhlopagurus foresti gen. nov., sp. nov., et de quinze espèces ou sous-espèces nouvelles de Parapagurus Smith (Crustacea, Decapoda). Bijdragen tot de Dierkunde 42: 97–123.

Pylochelidae Bate, 1888 The pylochelids are referred to as the ‘symmetrical hermit crabs’ and most species differ from all other hermit crabs in having a calcified segmented symmetrical abdomen. The members of the family are mostly represented between 200 and 500 m depth, rarely reaching as shallow as 30 m or as deep as 1570 m. Instead of gastropod shells, the usual hermit crab home, pylochelids inhabit burrows in small pieces of wood, stone or sponge, or use tusk shells or bamboo as homes. One species of deep-water pagurid, Porcellanopagurus tridentatus, is also straight and symmetrical but uses a bivalve for protection – it should not be confused with pylochelids. Forest’s (1987a, 1987b) excellent revisions with keys and detailed discussion of systematics and biology are the last word on Pylochelidae, often erroneously called Pomatochelidae. He provided more detailed diagnoses of the family, genus and species than given here. He recognised 28 species, some with subspecies, in seven genera of which six inhabit the Indo-West Pacific. Only one has been reported from Australia but others known from the region of Indonesia might be

286 Anomura – hermit crabs, porcelain crabs and squat lobsters

expected in deep tropical waters. Diagnosis. Abdomen straight (asymmetrical only in Mixtopagurus Forest, 1987), somites calcified and distinct; chelipeds equal; pleopods 1–5 paired. Maxillipeds 3 adjacent at base.

Trizocheles Forest, 1987 There are 17 species each distributed locally in the Indo-West Pacific. Diagnosis. Shield completely separated from posterior part of carapace. Pereopod 4 propodus with numerous imbricating squamiform setae. Ocular scales present. Abdomen symmetrical. Telson longer than wide. Trizocheles spinosus (Henderson, 1888) (Fig. 83, Pl. 17d). Contacting faces of cheliped and pereopod 2 with stridulating apparatus. Pereopod 2 propodus with 4 or more dorsal teeth ranged in line. Pereopod 3 propodus with 4–5 dorsal teeth. Eyestalk two-thirds length of shield. 7 mm. Southern Qld, NSW, to southern Tas.; 137– 470 m (subspecies T. spinosus spinosus). New Caledonia, south-eastern New Zealand; 127–550 m (subspecies T. spinosus bathamae Forest & de Saint Laurent, 1987). Forest’s material was found in a sponge; the species also occurs in tusk shells at depths beyond those initially reported.

Fig. 83. Pylochelidae. Trizocheles spinosus.

References Forest, J. 1987a. Les Pylochelidae ou ‘Pagures symétriques’ (Crustacea Coenobitoidea). Mémoires du Muséum National d’Histoire Naturelle, Paris 137: 1-254, 9 pls. Forest, J. 1987b. Ethology and distribution of Pylochelidae (Crustacea Decapoda Coenobitoidea). Bulletin of Marine Science 41: 309-321.

287 This page intentionally left blank 11. BRACHYURA – CRABS

To imagine crustaceans is to imagine crabs, perhaps the best known group but, in fact, one of the most atypical of the Crustacea as a whole and even of Decapoda. The absence of a ‘tail’ or abdomen obvious when viewed from the top defines most Brachyura unambiguously. In fact, the abdomen is present but tucked under the rest of the body enclosed in the carapace. The abdomen tightly fits into a cavity in males of more advanced crab families but is not so restricted in females. In more primitive families, as in the anomuran families Lithodidae and Lomisidae, the abdomen is larger and clearly visible in dorsal view. Nevertheless, it is never large and muscular as in lobsters and shrimps. In males, there are almost always only two pairs of pleopods (but in homolodromiids and some dromiids there are more vestigial pleopods in males), both modified and interacting as gonopods in the transfer of sperm to the female. Crabs are exceptionally diverse in form although never deviating far from the basic body plan of a carapace enclosing most of body somites on top and the sides, one pair of claws (pereopods 1) and four pairs of walking legs. The walking legs have simple dactyli, except in primitive families where the last legs are also minutely clawed. Over 6500 species have been described, the vast majority from the Indo-West Pacific region surrounding Australia to the north and east. Of these, almost one thousand inhabit Australia itself (Davie, 2002). Worldwide the numbers of crab species declines from the tropics to the poles with none known, for example, from Antarctica and almost none from the subantarctic. This gradient has been especially clearly demonstrated for the Americas (Boschi, 2000) but applies in Australia also. Only one-third of species recorded in Australia are known from southern Australia, as defined here. Some of those are rare vagrants or tropical species at the southern limit of their ranges in places like Rottnest Island. The diversity of crab morphology correlates with a diversity of crab ecology. In general, all are benthic (although some species live in the open ocean, usually associated with drifting objects, like turtles or algae), none swims for more than a brief time, and most are cryptic under stones or other shelters. A few have specialised associations, living associated with corals or sponges, or symbiotic with molluscs, worms or echinoderms. Some burrow in sand on beaches, others prefer mud and some spend most of their adult life up trees. Most species and even some families have restricted depth ranges, intertidal of shallow subtidal, continental shelf sediments, or deep sea. These preferences are discussed for each family in turn and can help eliminate taxa during the process of identification. The number of estuarine species in southern Australia is few but are included with others in this volume. Not included in this book on marine crustaceans are the truly terrestrial and freshwater species. Australia has a poor fauna of non-marine crabs compared to Asia, Africa or the Americas. The only truly terrestrial crabs in Australia are members of Gecarcinidae, Cardisoma carnifex, among tropical mangroves on the mainland and on Lizard I. on the northern Great Barrier Reef (Quinn et al., 1991). Australia also claims the famously abundant Christmas Island red crab, Geocarcoides natalis, one of five land crabs occurring there (Hicks et al., 1984). Freshwater crabs number seven species of Parathelphusidae in Australia of which one, Austrothelphusa transversa, reaches the drier areas of southern Australia far from the coast. This one may be a complex of three or more species.

289 Marine Decapod Crustacea of Southern Australia

The taxonomy of the Brachyura has been long debated and countless family names have been proposed, many later synonymised and some revived. Sometimes, the splitting and lumping of families served only the whims of older taxonomists wanting to label branches within a larger bush. For example, Portunidae has 22 known synonyms and Majidae 50 known synonyms, some valid for subfamilies, but both families are probably monophyletic. On the other hand, Grapsidae with eight synonyms, may not be monophyletic and some of its family-level taxa may be more related to other families than to most grapsids. Family names used here follow Davie (2002); Martin & Davis (2001) elevated some of subfamily names to family level. For many years groupings of the families relied on divisions based on overall body shape and shape of the mouth-field but beginning in the late 1970s, new ideas were proposed by Guinot (1977) in Paris. Her groupings have been debated on the basis of evidence from adult morphology (Guinot, 1978; de Saint Laurent, 1980; Tavares, 2003), larvae (Rice, 1980; Pohle & Telford, 1981; Martin, 1988), sperm structure (Jamieson & Tudge, 1990; Guinot et al., 1994, 1998), fossils (Glaessner, 1969; Guinot, 1993; Guinot & Tavares, 2001), eye structure (Fincham, 1980; Gaten, 1998) and molecular data (Spears et al., 1992; Scholtz & Richter, 1995; Kitaura et al., 2002). All are trying to derive a classification that reflects perceived evolutionary relationships. The questions about relationships are not trivial because fundamental to them is the monophyly of the Brachyura (Tavares, 2003). Guinot (1977) proposed three groups based on the position of the female gonopore, on the sixth sternite (sternite of walking leg 2 = pereopod 3) in Heterotremata and Thoracotremata, and on the coxa of the same limb in Podotremata. The monophyly of the Heterotremata + Thoracotremata clade, termed Eubrachyura by de Saint Laurent (1980) is not in doubt. The position of the female gonopore and the presence of a sella turcica, a structure within the internal skeleton (Secretan, 1998) are synapomorphies defining the clade. Synapomorphies supporting Thoracotremata are the sternal positions of both male and female gonopores. This is certainly a monophyletic group but the families included are debated. Guinot & Richer de Forges (1997) took a more conservative view than the one adopted here and restricted the group to just four families. The Heterotremata, all of the Eubrachyura except thoracotremes, crabs with female gonopores on the sternum and male gonopores on the sternum or coxae, remain probably paraphyletic. Questions remain about who to include within the Brachyura and whether any or all of the so-called podotremes are more or less related to Anomura. Podotremes, unlike all other crabs, share with anomurans, which might be closely related, and with all other malacostracans, female gonopores on the coxae of the sixth thoracic limbs (Tavares, 2003). Podotreme crabs themselves share one synapomorphy, a double spermatheca (Tavares & Secretan, 1993) but this does not inform about whether they are advanced anomurans or primitive crabs. The current view is that three clades exist, each with probably synapomorphies, but that nothing unites them. This is the view that is used here to arrange the families in an order that keeps similar families together. The following list explains; the ranks above family attempt a reconciliation with Martin & Davis (2001). This arrangement differs from theirs especially for the Podotremata. They allocated all families to superfamilies and their ranking of eubrachyurans is followed. More than 20 families of marine crabs listed by them are not found in southern Australia. Another eight are terrestrial.

Section Podotremata Subsection Homoloidea Subsection Dromiacea 4. Homolidae 1. Dromiidae 5. Latreilliidae 2. Dynomenidae Subsection Archaeobrachyura 3. Homolodromiidae 6. Cyclodorippidae 7. Cymonomidae 8. Raninidae

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Section Eubrachyura 22. Hexapodidae Subsection Heterotremata 23. Eriphiidae 9. Dorippidae 24. Pilumnidae 10. Calappidae 25. Xanthidae 11. Leucosiidae 26. Trapeziidae 12. Matutidae 27. Domeciidae 13. Majidae 28. Cryptochiridae 14. Hymenosomatidae Subsection Thoracotremata 15. Parthenopidae 29. Pinnotheridae 16. Atelecyclidae 30. Mictyridae 17. Cancridae 31. Ocypodidae 18. Corystidae 32. Palicidae 19. Geryonidae 33. Grapsidae 20. Portunidae 34. Plagusiidae 21. Goneplacidae

The key to families is adapted from Ng (1998) by deleting the tropical and terrestrial families. It does not reflect relationships but tries to eliminate unusual crabs first. Ecological notes are added if this helps eliminate families.

Key to southern Australian families of Brachyura 1. 3 pairs of walking legs visible (Fig. 142) (but see box – Six-legged crabs) ...... Hexapodidae . ..p.439 — 4 pairs of walking legs visible (last pair or 2 pairs may be smaller than others) ...... 2 2. Eyestalk exceptionally thin, basal segment much longer than second segment; deep-water (Fig. 91) ...... Latreilliidae . . .p.314 — Eyestalk thick, of one segment or with short, thin stalk ...... 3 3. Last pair of walking legs subchelate or chelate (Figs 84a, 95) or reduced, held over carapace 4 — Last pair of legs similar to others, held laterally ...... 10 4. Maxilliped 3 with merus distinctly triangular; crab carries shells etc over carapace ...... 5 — Maxilliped 3 with merus quadrate or more or less rounded ...... 7 5. Maxilliped 3 not covering large anterior part of buccal cavern; rare . . Dorippidae ...p.324 — Maxilliped 3 covering most or all of buccal cavern; deep shelf and slope ...... 6 6. Carapace hexagonal to subovate; orbits distinct; maxilliped 3 exopod usually without flagellum ...... Cyclodorippidae ...p.316 — Carapace rectangular to squarish; orbits absent; maxilliped 3 exopod with flagellum ...... Cymonomidae . . . p. 318 7. Carapace pear-shaped; orbits incomplete (e.g. Fig. 113); carapace, chelipeds and legs often with hooked setae; oopore of female on thoracic sternum ...... Majidae (part) ...p.347 — Carapace not pear-shaped; orbits usually complete; without hooked setae; oopore of female on coxae of second walking leg ...... 8 8. Carapace longitudinally rectangular, dorsal surface smooth or with scattered stiff setae; walking legs 4 with propodus and dactylus subchelate-chelate; deep shelf and slope ...... Homolidae ...p.311 — Carapace longitudinally circular or hexagonal, dorsal surface usually with dense soft setae; walking legs 3 and 4 both with propodus and dactylus subchelate-chelate; crab usually carrying sponge, tunicate or bivalve shell ...... 9

291 Marine Decapod Crustacea of Southern Australia

9. Carapace more or less circular or wider than long; uropods visible externally as small external plates at base of telson; intertidal to deep-sea ...... Dromiidae ...p.298 — Carapace longer than wide; uropods visible only internally if at all; deep-sea only ...... Homolodromiidae ...p.310 10. Maxilliped 3 merus distinctly triangular ...... 11 — Maxilliped 3 merus quadrate-ovate ...... 14 11. Carapace longitudinally ovate; chela with finger transverse ...... Raninidae ...p.320 — Carapace not much longer than wide; chela normally crab-like ...... 12 12. Opening for respiratory current below orbits, with distinct channel alongside base of buccal cavern, visible when maxillipeds 3 pushed aside (Figs 98a, b, , 99f–h); sides of carapace never expanded; chela without special cutting tooth ...... Leucosiidae ...p.331 — Opening of respiratory current at base of chela, without a channel; sides of carapace expanded; chela with strong tooth ...... 13 13. Carapace with posterolateral expansion covering walking legs; walking legs with dactyli tapered and pointed ...... Calappidae ...p.326 — Carapace with spine at most lateral point; walking legs flattened, especially distally ...... Matutidae ...p.344 14. Abdomen with small lateral plates between abdominal somite 6 and telson; walking leg 4 reduced to short appendage ...... Dynomenidae ...p.308 — Abdomen without lateral plate between abdominal somite 6 and telson; walking leg 4 similar to others or thin, short but otherwise normal ...... 15 15. Walking leg 4 strongly reduced (Fig. 158); deep shelf and slope ...... Palicidae ...p.498 — Walking leg 4 similar in size to others or not very much smaller ...... 16 16. Sockets for antenna 1 square or longer than broad; antenna 1 folds longitudinally ...... 17 — Sockets for antenna 1 broader than long; antenna 1 folds transversely or obliquely . . . . . 21 17. Carapace pear-shaped, triangular or circular; orbits incomplete or absent ...... 18 — Carapace longitudinally or transversely ovate, hexagonal or circular; orbits complete . . . 19 18. Carapace well calcified, dorsal surface convex, usually covered with spines or granules, often with hooked setae; abdomen with 6 (rarely 5) somites and telson; male genital openings on coxa of last walking leg ...... Majidae (part) ...p.347 — Carapace poorly calcified, soft, flat, never with spines or granules, without hooked setae; abdomen with 5 somites and telson; male genital opening on sternum ...... Hymenosomatidae ...p.390 19. Antennal flagellum slightly setose or smooth; body wider than long, smooth ...... Cancridae ...p.401 — Antennal flagellum distinctly setose; body longer or as long as broad, more or less setose . . 20 20. Antenna as long or longer than body; body longer than wide; burrowing in sand; rare ...... Corystidae ...p.404 — Antenna much shorter than body; body circular; deep shelf and slope ...... Atelecyclidae ...p.400 21. Carapace triangular or hexagonal, highly sculptured; front triangular, forked or spiniform; chelipeds usually stout and elongate ...... Parthenopidae ...p.398 — Carapace not as above; front usually truncate or multidentate; chelipeds short or, if elongate, slender ...... 22

292 Brachyura – crabs

22. Maxilliped 3 merus much smaller than ischium or fused with it; male genital openings on sternum; carapace circular or oval, smooth; often found in association with molluscs, worms or echinoderms ...... 23 — Maxilliped 3 merus square, at least half size of ischium; male genital openings on coxa (rarely sternum); carapace various; free-living ...... 24 23. Carapace circular or oval, smooth; usually living inside bivalves, holothurians or other sessile invertebrates ...... Pinnotheridae ...p.484 — Carapace longer than wide, tuberculate; living in corals ...... Cryptochiridae ...p.482 24. Last walking leg with dactylus paddle-shaped or at least slightly flattened and blade-like; usually on sandy or muddy sediments, sometimes under intertidal rocks ...... Portunidae ...p.407 — Last walking leg with dactylus similar to others, acute ...... 25 25. Distinct rhomboid gap between maxillipeds 3; mandibles usually visible when mouthparts closed; includes intertidal rocky shore crabs ...... 26 — Maxillipeds 3 closed together and concealing mandibles ...... 27 26. Front with 3 lobes separated by antennular notches, such that antennules are visible in dorsal view ...... Plagusiidae ...p.512 — Front entire, without notches, concealing antennules from dorsal view ...... Grapsidae (part) ...p.500 27. Front simple, triangular, very narrow compared to carapace width ...... 28 — Front truncate, multilobed or multidentate, at least a third of carapace width ...... 29 28. Body globular; orbits absent; intertidal sandy beaches ...... Mictyridae ...p.487 — Body flat, typical crab-shape; orbits long and transverse ...... Ocypodidae ...p.490 29. Dactylus of walking legs T-shaped; large, deep-water ...... Geryonidae ...p.406 — Dactylus of walking legs usually quadrate or oval; shallow or deepwater crabs ...... 30 30. Carapace squarish to longitudinally rectangular, posterior margins subparallel, dorsal surface flat or gently convex; intertidal or estuarine or shallow-water crabs ...... Grapsidae (part) ...p.500 — Carapace not as above, dorsal surface usually convex, posterolateral margins gently to strongly converging; sublittoral to deep-water crabs ...... 31 31. Walking legs with special dactylus-propodus articulation (formed by rounded submedian extension of lateral margin of propodus, sliding under a projecting button at base of dactylus) (Fig. 149d, e) ...... 32 — Walking legs without such special articulation ...... 35 32. Chelipeds with fingers distinctly spoon-shaped; male abdominal somites 3–5 immoveable, fused although sutures may be partially visible; cryptic intertidal and shelf ...... Xanthidae (part) . . .p.460 — Chelipeds with fingers not spoon-shaped, acute; male abdominal somites 3–5 freely moveable or fused ...... 33 33. Carapace flat, hexagonal or pentagonal; male gonopod 1 very slender, S-shaped; male gonopod 2 short, less than quarter length of gonopod 1, comma-shaped; commensals of echinoderms ...... Pilumnidae (Eumedoninae) ...p.449 — Carapace square or round; male gonopod 1 stout, sinuous – straight; male gonopod 2 short, more than quarter length of gonopod 1, not comma-shaped ...... 34

293 Marine Decapod Crustacea of Southern Australia

34. Carapace flattened, with 4-lobed front; chelipeds smooth ...... Trapeziidae ...p.479 — Carapace convex, with bilobed front, chelipeds spinose or granular . . Domeciidae ...p.481 35. Male abdominal somites 3–5 distinct, moveable; carapace regions poorly defined ...... 36 — Male abdominal somites 3–5 fused, immoveable; carapace regions usually well defined . . 38 36. Male gonopod 1 stout, straight ; male gonopod 2 elongate, as long as or longer than gonopod 1 ...... Eriphiidae ...p.441 — Male gonopod 1 slender to stout, sinuous to straight ; male gonopod 2 shorter than gonopod 1 ...... 37 37. Male gonopod 1 slender, S-shaped to almost straight ; male gonopod 2 short, comma-shaped ...... Pilumnidae (part) ...p.448 — Male gonopod 1 stout, sinuous ; male gonopod 2 third to half length of gonopod 1 ...... Goneplacidae (part) . . . p. 433 38. Carapace usually transversely rectangular or ovate, or subcircular; male gonopod 2 usually third to as long as gonopod 1 ...... Goneplacidae (part) ...p.433 — Carapace usually hexagonal or transversely ovate; male gonopod 2 short, comma-shaped, no more than third gonopod 1, not sigmoid ...... Xanthidae (part) ...p.460

References Boschi, E.E. 2000. Biodiversity of marine decapod brachyurans of the Americas. Journal of Crustacean Biology 20 (Special Number 2): 337–342. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Fincham, A.A. 1980. Eyes and the classification of the malacostracan crustaceans. Nature 287: 729–731. Gaten, E. 1998. Optics and phylogeny: is there an insight? The evolution of superposition eyes in the Decapoda (Crustacea). Contributions to Zoology 67: 223–235. Glaessner, M.F. 1969. Decapoda. Pp. R399–R651 in: Moore, R.C. (ed.) Treatise on Invertebrate Paleontology, Part R, Arthropoda 4. Vol. 2. Geological Society of America and University of Kansas Press Guinot, D. 1977. Propositions pour une novelle classification des Crustacés, Décapodes, Brachyoures. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 285: 1049–1052. Guinot, D. 1978. Principes d’une classification évolutive des Crustacés Décapodes Brachyoures. Bulletin Biologique de la France et de la Belgique 112: 211–292. Guinot, D. 1993. Données nouvelles sur les crabes primitifs (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 316: 1225–1232. Guinot, D., Jamieson, B.G.M., & Richer de Forges, B. 1994. Relationship of Homolidae and Dromiidae: evidence from spermatozoal ultrastructure (Crustacea, Decapoda). Acta Zoologica, Stockholm 75: 255–267. Guinot, D., Jamieson, B.G.M., Richer de Forges, B., & Tudge, C.C. 1998. Comparative spermatozoal ultrastructure of the three dromiacean families exemplified by Homolodromia kai (Homolodromiidae), Spaerodromia lamellata (Dromiidae), and Dynomene tanensis (Dynomenidae (Podotremata: Brachyura). Journal of Crustacean Biology 18: 78–94. Guinot, D., & Richer de Forges, B. 1997. Affinités entre les Hymenosomatidae MacLeay, 1838 et les Inachoididae Dana, 1851 (Crustacea, Decapoda, Brachyura). Zoosystema 19: 453–502. Guinot, D., & Tavares, M. 2001. Une nouvelle famille de crabes du Crétacé, et la notion de Podotremata Guinot, 1977 (Crustacea, Decapoda, Brachyura). Zoosystema 23: 507–546.

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Hicks, J., Rumpff, H., & Yorskston, H. 1984. Christmas Crabs. Christmas Island Natural History Association: Christmas Island. Jamieson, B.G.M., & Tudge, C.C. 1990. Dorippids are Heterotremata: evidence from ultrastructure of the spermatozoa of Neodorippe astuta (Dorippidae) and Portunus pelagicus (Portunidae) Brachyura: Decapoda. Marine Biology 106: 347–354. Kitaura, J., Wada, K., & Nishida, M. 2002. Molecular phylogeny of grapsoid and ocypodoid crabs with special reference to the genera Metaplax and Macrophthalmus. Journal of Crustacean Biology 22: 682–693. Martin, J.W. 1988. Phylogenetic significance of the brachyuran megalopa: evidence from the Xanthidae. Symposia of the Zoological Society of London 59: 69–102. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. Ng, P.K. 1998. Crabs. Pp. 1045–1155 in: Carpenter, K.E., & Niem, V.H. (eds), The Living Resources of the Western Central Pacific. FAO: Rome. Pohle, G., & Telford, M. 1981. Morphology and classification of decapod crustacean larval setae: a scan- ning electron microscope study of Dissodactylus crinitichelis Moreira, 1901 (Brachyura: Pinnotheridae). Bulletin of Marine Science 31: 736–752. Quinn, D.J., Diele, K., & Meischner, U. 1991. Reproductive behaviour of Cardiosoma carnifex (Herbst, 1794) (Brachyura: Gecarcinidae) at Lizard Island, Great Barrier Reef. Memoirs of the Queensland Museum 31: 399. Rice, A.L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Transactions of the Zoological Society of London 35: 271–424. de Saint Laurent, M. 1980. Sur la classification et la phylogénie des Crustacés Décapodes Brachyoures. I. Podotremata Guinot, 1977 et Eubrachyura sect. nov. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 290: 1265–1268. Scholtz, G., & Richter, S. 1995. Phylogenetic systematics of the reptantian Decapoda (Crustacea, Malacostraca). Zoological Journal of the Linnean Society 113: 289–328. Secretan, S. 1998. The sella turcica and the endophragmal system of decapods. Journal of Natural History 32: 1753–1767. Spears, T.,Abele, L.G., & Kim, W. 1992. The monophyly of brachyuran crabs: a phylogenetic study based on 18S rRNA. Systematic Biology 41: 446–461. Tavares, M.S. 2003. A new theoretical approach for the study of monophyly of the Brachyura (Crustacea: Decapoda) and its implication for the Anomura. Memoirs of Museum Victoria 60: 145–149. Tavares, M., & Secretan, S. 1993. La notion de thelycum et de spermathèque chez les Crustacés Décapodes. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 316: 133–138.

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TWENTY-FIVE COMMON CRABS Few southern Australia crabs are common intertidally or on shallow reefs. This key is a short- cut to some common species but certainly not all. Once you have worked through the key, look in the main chapters to compare with similar species and confirm your identification. 1. Body globular, bright blue, living on sandy beaches ...... Mictyridae: Mictyris platycheles (soldier crabs) — Body more typically crab-like ...... 2 2. Carapace flattened, oval, with prominent anterolateral teeth ...... Portunidae (swimming crabs) … 3 — Carapace square, pear-shaped, triangular or, if flattened, not with prominent teeth . . 4 3. Last leg paddle-like, with 2 red spots on carapace ; sandy beaches ...... Ovalipes australiensis (sand crab) — Last leg with pointed, flattened last article; rocky shores, bays ...... Carcinus maenas (European shore crab) 4. Carapace oval, flattened, with H-shaped groove on back; less than 12 mm wide ...... Hymenosomatidae (flat-backed spider crabs) ...5 — Carapace various, not flat ...... 7 5. Carapace with single broad rostrum; in estuaries and freshwater . . . . . Amarinus laevis — Carapace with rostrum of 3 teeth ...... 6 6. Rostrum of 3 evenly-spaced and sized teeth ...... Halicarcinus ovatus — Rostrum of dominant median spine and 2 lateral teeth ...... Halicarcinus rostratus 7. Carapace highly convex, more or less pear-shaped, often with algae and sponges attached; with double rostrum and spines around eyes ...... Majidae (spider crabs and decorator crabs) … 8 — Carapace rectangular or oval, not tapering to acute rostrum; if strongly globular then carapace polished ...... 11 8. Large subtidal spider-crab, carapace to 165 mm long; eyes protected behind by cupped spine ...... Leptomithrax gaimardii (giant spider crab) — Moderate-size decorator crab, carapace to 80 mm long; eyes protected spines but none cupped ...... 8 9. Eave over eye broadly rounded anteriorly, without spine; walking legs cylindrica ...... Notomithrax minor — Eave over eye with spine on anterior corner; walking legs with propodi expanded and flattened ...... Naxia ... 10 10. Rostral spines slender, strongly curved outwards; eave without spines ...... Naxia aries (ramshorn crab) — Rostral spines stout, straight; eave with spine or sharp angles front and back ...... Naxia aurita (smooth seaweed-crab) 11. Carapace oval, covered with scattered or dense hairs ...... Pilumnidae ...12 — Carapace shape various, without hairs ...... 13

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12. Hairs scattered except on underside of carapace ...... Pilumnopeus serratifrons (smooth-handed crab) — Hair dense especially anteriorly on carapace . . . Pilumnus monilifer (beaded hairy crab) 13. Carapace globular, highly domed; cheliped much longer than body ...... Leucosiidae (pebble crabs) ...14 — Carapace flat, more or less rectangular; cheliped compact ...... 15 14. Carapace almost spherical, without prominent spines ...... Bellidilia laevis (smooth pebble crab) — Carapace with produced 4-lobed front and prominent posterior projection ...... Ebalia intermedia (smooth nut crab) 15. Eyestalks long; front between only quarter width of carapace . . . . . Ocypodidae ...16 — Eyestalks short; front between about half width of carapace ...... 17 16. Carapace almost twice as wide as long, narrow at front; chelipeds with fingers twisted; ...... Heloecius cordiformis (semaphore crab) — Carapace square or little wider, widest at front; chelipeds not twisted ...... Macrophthalmus latifrons (southern sentinel crab) 17. Carapace with 1 spine on each side and parallel-sided behind ...... Goneplacidae: Litocheira bispinosa — Carapace with convex sides, with or without notches but not spines . . . Grapsidae . . . 18 18. Carapace oval, with 1 prominent tooth immediately behind orbital angle and 2 or 3 small teeth more posterior; large supratidal crab ...... Leptograpsodes octodentatus (burrowing shore crab) — Carapace rectangular or slightly curved lateral margins; with evenly-spaced small teeth if any; crab of muddy or rocky intertidal ...... 19 19. Carapace widest just behind eyes; chelipeds bright orange . Parasesarma erythodactyla — Carapace with curved or toothed lateral margins; cheliped not orange ...... 20 20. Carapace without lateral teeth ...... Cylcograpsus granulosus — Carapace with lateral teeth ...... 21 21. Carapace with 3 lateral teeth on each side; fingers of cheliped with mat of hairs on inner and outer surfaces ...... Brachynotus spinosus (little shore crab) — Carapace with 1 or 2 lateral teeth on each side; fingers of cheliped without hairs . . . 22 22. Carapace with front curving evenly downwards ...... Helograpsus haswellianus (Haswell’s shore crab) — Carapace with front not curving strongly downwards Paragrapsus (shore crabs) ...23 23. Anterolateral margins with 1 tooth ...... Paragrapsus quadridentatus — Anterolateral margins with 2 teeth ...... 24 24. First walking legs with felt on anterior surface ...... Paragrapsus laevis — First walking legs with only inner edge felted ...... Paragrapsus gaimardii

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Section Podotremata Guinot, 1977 The intermediate position of podotreme crabs between eubrachyurans and anomurans has stim- ulated an unresolved debate over their classification. Here, the families are arranged in three groups, best called subsections: Dromiacea, Homoloidea and Archaeobrachyura.

Subsection Dromiacea De Haan, 1833

Dromiidae De Haan, 1833 The dromiids are the sponge crabs, so called because of their habit of disguise using a piece of sponge or other encrusting animal carried over the back. In many species the sponge appears to have been carefully trimmed to fit the crab and is sometimes much larger than its owner. Most species are cryptic and are rarely seen intertidally. The Dromiidae are often considered one of the most primitive families of Brachyura and are sometimes excluded from this order. McLay (1993) believed that the ancestral dromiid had four legs used for walking, and with spines on the propodus and dactylus. From this has evolved patterns of spination adapted for carrying camouflage, and later, replacement of this behaviour with reduction in size of posterior legs and more ornamented carapace. McLay (1993) has provided a systematic review of the family. He summarised earlier nomenclatural confusion while providing several new generic names for many common Australian species. While he concentrated on descriptions of New Caledonian and Philippine species his revision is worldwide at the generic level. He recognised 30 genera for about 100 species and provided keys to them all. His illustrations of key characters are good but the publication itself is difficult to obtain in Australia. Another three genera were added for species from the Northern Territory and Singapore (McLay, 1998, 2001, 2002). McLay’s was not the last word and Guinot & Tavares (2003) have provided a new subfamily arrangement (Dromiinae, Hypochonchinae, Sphaerodrominae) based largely on the female reproductive structures. Only the Dromiinae are found in southern Australia. They also rediagnosed some genera and erected eight new genera. The dromiid fauna of southern Australia comprises 15 described species. The most common species, based on the records of Museum Victoria are Stimdromia lateralis, S. lamellata, Lamarckdromia globosa, Dromia wilsoni and Austrodromia australis. Some species, Epipedodromia thomsoni, Haledromia bicavernosa, Dromia wilsoni, Lamarckdromia globosa and the two species of Fultodromia are readily recognisable from the drawings of the carapaces presented here but the rest are more difficult. The presence of an epipod attached to the coxa of the cheliped is a character of major taxonomic importance in distinguishing genera but it can be difficult to see. It is small and bifurcate, concealed inside the gill chamber by the edge of the carapace but is visible on the posterior edge of the coxa if the cheliped is moved forward or detached.

Diagnosis. Carapace typically convex and subglobular, ovoid, subcircular, or pentagonal. Lateral linea present. Orbit, antennular and antennal fossae defined. Maxilliped 3 operculiform. Last 2 pairs of legs smaller, similar in size, carried subdorsally; armed with a chela formed by a distal propodal spine and dactylus. Abdomen of both sexes of 7 somites (including telson) generally without pleurae; males rarely with vestigial pleopods on somites 3–5; somite 1 of females with pair of uniramous pleopods. Abdomen usually with a retaining mechanism, often involving dorsal uropods. Uropods always present (except in Ascidiophilus) in form of dorsal plates, sometimes intercalated laterally, rarely ventral. Penis forming a long penial tube. Spermathecal apertures small and rounded. (adapted from Davie, 2002)

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Fig. 84. Dromiidae. a, Austrodromidia australis. b, Dromia wilsoni.

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Key to southern Australian species of Dromiidae 1. Cheliped usually with epipod; last 2 abdominal somites fused or free ...... 2 — Cheliped usually without epipod; last 2 abdominal somites free ...... 9 2. Carapace longer than wide, surface sparsely tuberculate; anterolateral margin with irregular teeth; walking legs 1 and 2 with several conspicuous nodules ...... Fultodromia ... 3 — Carapace usually wider than long or as wide as long, surface not tuberculate, smooth, finely denticulate or with areas clearly defined; anterolateral margin smooth or with regular teeth; walking legs 1 and 2 with few or no conspicuous nodules ...... 4 3. Carapace with large tubercles extending on to anterolateral teeth; walking legs 3 and 4 with 3 spines on outer margin of propodus ...... Fultodromia nodipes — Carapace sparsely covered with small tubercles not extending on to anterolateral teeth; walking legs 3 and 4 with 1 or 2 spines on outer margin of propodus . . . . . Fultodromia spinifera 4. Carapace with pair of kidney-shaped cavities each side of rostrum; uropodal plates vestigial or absent, not visible externally ...... Haledromia bicavernosa — Carapace without pair of kidney-shaped cavities each side of rostrum; uropodal plates small but visible externally ...... 5 5. Walking leg 4 with spine on outer margin of dactylus ...... 6 — Walking leg 4 without spine on outer margin of dactylus ...... 7 6. Carapace longer than wide; last 2 abdominal somites fused . . . . . Dromidiopsis australiensis — Carapace about as long as wide; last 2 abdominal somites free . . . . . Lamarckdromia globosa 7. Chelipeds and walking legs 1 and 2 strongly tuberculate; carapace with dorsal mat of soft hairs ...... Dromia wilsoni — Chelipeds and walking legs 1 and 2 ridged; carapace finely granulate . . . . . Stimdromia ... 8 8. Subhepatic region (area under and between eye and anterolateral tooth) with a strong blunt tubercle; inner surface of male cheliped propodus entirely hairy, outer surface smooth; walking legs 1 and 2 with carpus and propodus weakly ridged ...... Stimdromia lateralis — Subhepatic region with a low straight ridge; inner surface of male cheliped propodus hairy only distally, outer surface granular; walking legs 1 and 2 with carpus and propodus sharply ridged ...... Stimdromia lamellata 9. Carapace covered with large granular tubercles ...... Epigodromia sculpta — Carapace smooth ...... 10 10. Uropodal plates small but visible externally ...... Cryptodromia tumida — Uropod plates concealed or absent ...... 11 11. Rostrum bidentate; with 4-lobed transverse ridge behind rostrum; walking legs 3 and 4 without spines on outer margin of propodus ...... Epipedodromia thomsoni — Rostrum tridentate; without transverse ridge behind rostrum; walking legs 3 and 4 with spines on outer margin of propodus; last leg much shorter than others ...... Austrodromidia ... 12 12. Without anterolateral teeth; carapace about as wide as long ...... 13 — Anterolateral teeth present; carapace wider than long ...... 14 13. Lateral rostral and supraorbital teeth acute ...... Austrodromidia incisa — Lateral rostral and supraorbital teeth blunt ...... Austrodromidia insignis 14. Uropodal plates small, visible ventrally between somite 6 and telson; 4 anterolateral teeth ...... Austrodromidia australis — Uropodal plates vestigial, appearing as scales on ventral surface of somite 6; 4–5 anterolateral teeth ...... Austrodromidia octodentata

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Fig. 85. Dromiidae. a, Fultodromia nodipes. b, Lamarckdromia globosa.

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Fig. 86. Dromiidae. a, Austrodromidia australis (with sponge cap). b, Epipedodromia thomsoni. c, Lamarckdromia globosa. d, Austrodromidia insignis. e, f, Dromidiopsis australiensis.

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c b

d e

Fig. 87. Dromiidae. a, Dromidiopsis australiensis. b, Stimdromia lamellata (ventral carapace). c, Stimdromia lateralis (ventral carapace). Inside view of posterior abdominal somites of male showing uropods: d, Austrodromidia australis. e, Austrodromidia octodentata.

Austrodromidia McLay, 1993 The four southern Australian species are the only members of the genus which is confined to this region. All are covered with a dense tomentum and carry a well fitting camouflage cap of a dense sponge or ascidian. Females carry unusually large eggs, 2 mm in diameter, and at least A. octodentata broods its young (Hale, 1925). Guinot & Tavares (2003) rediagnosed the genus. Diagnosis. Carapace as wide or wider than long; covered with dense felt, with longer setae on fringes. Rostrum tridendate, with supraorbital tooth and blunt or obtuse postorbital tooth. Female sternal grooves end together between walking legs 1. Cheliped without epipod. Walking legs 1 and 2 with carpus and propodus produced. Walking leg 3 with 1–2 spines on propodus opposing dactylus plus up to 3 on outer margin. Walking leg 4 shorter than walking leg 1; with 1 spine on propodus opposing dactylus plus up to 3 on outer margin. Male uropods ventral, concealed or obsolete. Last 2 somites of abdomen free. Telson rounded. Austrodromidia australis (Rathbun, 1923) southern sponge-crab (Figs 84a, 86a, 87d). Carapace wider than long, with blunt lateral rostral and supraorbital teeth, with anterolateral teeth. Uropodal plates small, visible ventrally between somite 6 and telson. 35 mm. NSW, Vic., Tas., SA, WA; intertidal to 68 m depth. This species is common but easily confused with A. octodentata. It is distinguished from the latter by the presence of small uropods seen inside between the last abdominal somite and the telson when the abdomen is lifted, and by the more prominent marginal teeth. The species was previously known as Dromidia australis (Hale, 1927). Austrodromidia incisa (Henderson, 1888) Carapace as wide as long, with acute lateral rostral and supraorbital teeth, without anterolateral teeth. 12 mm. NSW (Twofold Bay); 200 m, known only from type specimens covered with ascidian. Austrodromidia insignis (Rathbun, 1923) adorned sponge-crab (Fig. 86d). Carapace about as wide as long, with blunt lateral rostral and supraorbital teeth, without anterolateral teeth; lateral rostral teeth with row of clubbed setae. Uropodal plates small, visible ventrally between somite 6 and telson. 17 mm. NSW,Vic., Tas., SA; 128–183 m depth. A. insignis is a rarely taken shelf species, previously placed in the genus Dromidia.

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Austrodromidia octodentata (Haswell, 1882) bristled sponge-crab (Fig. 87e, Pl. 17e). Carapace wider than long, with blunt lateral rostral and supraorbital teeth, with anterolateral teeth. Uropodal plates vestigial, appearing as scales on ventral surface of somite 6. 35 mm. NSW, Vic., SA, WA; subtidal to 79 m depth. The species is easily confused with A. australis but its uropods are almost invisible. The species was previously known as Cryptodromia octodentata (Hale, 1927).

Cryptodromia Stimpson, 1858 Members of the genus are best recognised by the lobed or nodular walking legs and smooth carapace. All are relatively small, less than 20 mm. Its 13 species are distributed through the tropical Indo-West Pacific (McLay, 1993), four reaching northern Australia. Only one is found, rarely, in southern WA. Diagnosis. Carapace as wide or wider than long; surface smooth, convex. Rostrum tridendate, with small supraorbital tooth, blunt postorbital tooth, and well developed suborbital tooth. Female sternal grooves end apart on small tubercles between walking legs 1. Cheliped without, or rarely with, epipod. Walking legs 1 and 2 with carpus and propodus lobed or nodular. Walking legs 1 and 2 with up to 6 spines on dactyli. Walking legs 3 and 4 reduced; with 1 spine on propodus opposing dactylus plus up to 2 on outer margin. Uropods well developed and visible externally. Abdomen of 6 somites. Telson usually rounded. Cryptodromia tumida Stimpson, 1858. Carapace wider than long, smooth, lateral cardiac grooves not marked, with 3 equal anterolateral teeth; rostrum with lateral teeth more prominent than median. Cheliped with rows of granules. Walking legs 1 and 2 with lobed carpi and propodi. Abdominal somites smooth. 15 mm. Indo-West Pacific, southern WA; sublittoral.

Dromia Weber, 1795 The type genus of the family comprises ten species of large sponge crabs, most restricted to either the Atlantic Ocean or Indo-West Pacific region. McLay (1993) provided a key. A key character for their recognition is the externally visible uropods. The only species from southern Australia has a wide distribution in other seas. Diagnosis. Carapace wider than long, dorsally smooth or slightly sculptured. Rostrum tridentate, lateral teeth rounded. Cheliped with epipod. Walking legs smooth. Walking leg 3 with 1 spine on propodus opposing dactylus, without spines on outer margin. Walking leg 4 shorter than walking leg 1; with 1–2 spines on propodus opposing dactylus plus 1 spine on outer margin; without spine on outer margin of dactylus. Female sternal grooves end apart or together between or behind chelipeds. Uropods usually visible externally. Last 2 somites of abdomen free. Telson rounded. Dromia wilsoni (Fulton & Grant, 1902) (Fig. 84b, Pl. 17f) Anterolateral margin with 3 well developed teeth, 2 directed upward. Carapace not strongly sculptured, covered with dense tomentum. 70 mm. New Zealand, South Africa, Japan, St Helena, NSW, Vic., Tas., SA, WA; subtidal to 520 m depth. The carapace of this common species is dark grey, densely hairy and sculptured. A sponge or ascidian cap is usual. The species, originally described from Victoria, is one of the most widespread of dromiid species (McLay, 1988). The species has been mentioned by many authors under the name Petalomera wilsoni (e.g., Hale, 1927).

Dromidiopsis Borradaile, 1900 Dromidiopsis is an Indo-West Pacific genus of four species (Guinot & Tavares, 2003). Following Guinot & Tavares’ (2003) rediagnosis, some Australian species appearing under this genus name

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belong to other genera. McLay (1993) provided a key and a clear delineation from Dromia. Species of Dromidiopsis are smaller (carapace less than 40 mm wide) than of Dromia. Diagnosis. Carapace longer than wide, dorsally smooth. Rostrum tridentate, lateral teeth rounded, not prominent. Female sternal grooves end on medial tubercle, behind chelipeds. Cheliped with epipod. Walking legs smooth. Walking leg 3 with 1 spine on propodus opposing dactylus, with 2 spines on outer margin. Walking leg 4 about as long as walking leg 1; with 1–2 spines on propodus opposing dactylus plus 1 spine on outer margin; with 1 spine on outer margin of dactylus. Uropods visible externally, involved in holding abdomen. Male abdomen with somites 5–6 fused. Telson long, ovate, rounded. Dromidiopsis australiensis (Haswell, 1882) (Fig. 87a, Pl. 17g). Carapace front and orbital margin without teeth or a dense fringe of setae. Anterolateral margin with 1–3 teeth. 33 mm. NSW, WA; shallow water. Records outside Australia of this rarely taken species are of other species (Lewinsohn, 1984). Dromidiopsis abrolhensis Montgomery, 1931 is a junior synonym.

Epigodromia McLay, 1993 Epigodromia is a genus of nine Indo-West Pacific species, all small, with a projecting rostrum and a granulate carapace. Only one species occurs in southern Australia but another lives in Queensland and three in New Caledonia (McLay, 1993). Diagnosis. Carapace width less than, equal to or greater than length; surface granular. Rostrum tridentate, teeth blunt, divergent; anterolateral teeth usually broad granular lobes. Female sternal grooves end apart between walking legs 1. Cheliped without epipod (rarely present). Walking legs 1 and 2 tuberculate. Walking leg 3 with 1 spine on propodus opposing dactylus, without spine on outer margin. Walking leg 4 much shorter than walking leg 1; with 1 spine on propodus opposing dactylus, without spine on outer margin. Uropods visible externally. Last 2 somites of abdomen free; terminal somites with granules and transverse ridges. Telson truncate or bilobed. Epigodromia sculpta (Haswell, 1882). Carapace wider than long, not divided into areas, covered with large granulated tubercles; anterolateral margin with 3 granulate teeth. 11 mm. NSW, Tas.; subtidal.

Epipedodromia André, 1932 There is only one species (Guinot & Tavares, 2003). Diagnosis. Carapace wider than long, flattened, subpentagonal; margins with short stiff setae; surface minutely granular. Rostrum bidentate, eave-like, frontal margin strongly deflexed; prominent curved ridge behind front divided into 4 equal parts by grooves. Female sternal grooves end apart on sinuous ridge between chelipeds. Cheliped without epipod. Walking legs 1 and 2 fringed with setae. Walking legs 3 and 4 reduced, with 1 spine on propodus opposing dactylus, without spine on outer margin. Uropods absent. Last 2 somites of abdomen free. Telson rounded. Epipedodromia thomsoni (Fulton & Grant, 1902) flat sponge-crab (Fig. 86b). 12 mm. Vic. (E to Western Port), SA; to 60 m depth. The species is instantly recognised by its flat carapace with four lobes along the front. It has been known until recently as Platydromia thomsoni but was moved because this generic name applied to something else.

Fultodromia McLay, 1993 The only two species of this genus occur in southern Australia (Guinot & Tavares, 2003). They are distinguished by the granules on the carapace.

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Diagnosis. Carapace almost as wide as long; surface sparse tuberculate. Rostrum tridentate, teeth broad and blunt; anterolateral margin with well developed teeth. Female sternal grooves end apart between or behind chelipeds. Cheliped with epipod. Walking legs 1 and 2 strongly nodular. Walking leg 3 with 1–2 spines on propodus opposing dactylus, with up to 3 spines on outer margin. Walking leg 4 shorter than walking leg 1; with 1–2 spines on propodus opposing dactylus plus up to 3 spines on outer margin. Uropods visible externally. Last 2 somites of abdomen free. Telson rounded in female, bilobed in male. Fultodromia nodipes (Guérin-Méneville, 1832) warty sponge-crab (Fig. 85a). Carapace with large tubercles, extending on to anterolateral teeth. Propodus of walking legs 3 and 4 armed with 3 spines on outer margin. 30 mm. SA, WA. The species is variable particularly in the size and arrangement of anterolateral teeth and the tubercles. Hale (1925) and others have treated this species under the name Petalomera depressa (Baker, 1907). Fultodromia spinifera (Montgomery, 1931). Carapace with small tubercles, not extending on to anterolateral teeth. Propodus of walking legs 3 and 4 armed with 1 or 2 spines on outer margin. 14 mm. WA.

Haledromia McLay, 1993 There is only one species and it is confined to southern Australia. The carapace is distinctive, having deep, red kidney-shaped cavities on each side of the rostrum, and vestigial uropods. The cavities are not seen in any other Australian species. Diagnosis. Carapace much wider than long, dorsally smooth. Rostrum tridendate, anterolateral margin without teeth. Female sternal grooves end together between chelipeds. Cheliped with epipod. Walking legs smooth. Walking leg 3 with 1 spine on propodus opposing dactylus, without spines on outer margin. Walking leg 4 shorter than walking leg 1; with 1 spine on propodus opposing dactylus, without spine on outer margin; without spine on outer margin of dactylus. Uropods vestigial, concealed. Last 2 somites of abdomen free. Telson rounded. Haledromia bicavernosa (Zietz, 1887) eared sponge-crab (Pl. 17h). 93 mm. SA; intertidal or shallow water. The species has large eggs, 2.8 mm in diameter, and may have direct development.

Lamarckdromia Guinot & Tavares, 2003 The genus was erected for one species to distinguish it from Dromidiopsis on the basis of abdominal and uropodal structure (Guinot & Tavares, 2003). Diagnosis. Carapace about as long as wide, dorsally smooth. Front narrow, appearing tridentate, with strongly deflexed rostrum, lateral teeth rounded, not prominent. Female sternal grooves end together on medial tubercle, near bases of walking leg 1. Cheliped with epipod. Walking legs smooth. Walking leg 3 with 1 spine on propodus opposing dactylus, with 2 spines on outer margin. Walking leg 4 about as long as walking leg 1; with 1–2 spines on propodus opposing dactylus plus 1 spine on outer margin; with 1 spine on outer margin of dactylus. Uropods visible as ventral ovate plates, not visible externally, not involved in holding abdomen. Male abdomen with somites free. Telson broader than long, rounded. Lamarckdromia globosa (Lamarck, 1818) shaggy sponge-crab (Figs 85b, 86c, Pl. 18a, b). Carapace with dense fringe of setae across front; anterolateral margin with 1 tooth. 41 mm. Central NSW, Vic., Tas., SA, WA (N to Fremantle) (Griffin, 1972); rare intertidally, more common subtidally to 180 m depth. This endemic Australian species has been widely known as Dromidiopsis excavata. The fringe of setae across the front immediately identify this sponge crab. The species is unusual in having large red eggs, 2 mm in diameter, which develop directly and are brooded by the parent under the abdomen (Hale, 1941).

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Stimdromia McLay, 1993 Stimdromia was erected for five Indo-West Pacific species, all with few nodules on distal parts of anterior walking legs. Two occur in southern Australia. Both were previously included in the genus Petalomera (e.g., by Griffin, 1972). Diagnosis. Carapace as wide or wider than long; surface smooth or granular. Rostrum tridentate, teeth broad and blunt; anterolateral margin with well developed teeth. Female sternal grooves end apart between walking legs 1. Cheliped with epipod. Walking legs 1 and 2 strongly nodular or ridged, like chelipeds. Walking leg 3 with 1 spine on propodus opposing dactylus, with 1 spine on outer margin. Walking leg 4 shorter than walking leg 1; with 1 spine on propodus opposing dactylus plus 1 spine on outer margin. Uropods visible externally. Last 2 somites of abdomen free. Telson rounded or bilobed. Stimdromia lamellata (Ortmann, 1894) (Fig. 87b). Subhepatic region (area under and between eye and anterolateral tooth) with a low straight ridge. Inner surface of male cheliped propodus hairy only distally, outer surface granular. Walking legs 1 and 2 with carpus and propodus sharply ridged. 24 mm. Qld, NSW, Vic., Tas., SA, southern WA; intertidal to 120 m depth; numerous records from Port Phillip Bay and Western Port. Stimdromia lateralis (Gray, 1831) ridged sponge-crab (Fig. 87c). Subhepatic region (area under and between eye and anterolateral tooth) with a strong blunt tubercle. Inner surface of male cheliped propodus entirely hairy, outer surface smooth. Walking legs 1 and 2 with carpus and propodus weakly ridged. 20 mm. New Zealand, central Qld, NSW, Vic., Tas., SA, WA (N to North West Cape); intertidal to 220 m depth. Stimdromia lateralis is one of the most common dromiids in southern Australia. It has direct development (Hale, 1925).

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Griffin, D.J.G. 1972. Brachyura collected by Danish expeditions on south-eastern Australia (Crustacea, Decapoda). Steenstrupia 2: 49–90. Guinot, D., & Tavares, M. 2003. A new subfamilial arrangement for the Dromiidae de Haan, 1833, with diagnoses and descriptions of new genera and species (Crustacea, Decapoda, Brachyura). Zoosystema 25: 43–129. Hale, H.M. 1925. The development of two Australian sponge-crabs. Proceedings of the Linnean Society of New South Wales 1: 405–413, pls 39–40. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Hale, H.M. 1941. Decapod Crustacea. British, Australian and New Zealand Antarctic Research Expedition, 1929–1931. Reports–Series B (Zoology and Botany) 4: 257–86,pl.3. Lewinsohn, C. 1984. Dromiidae (Crustacea, Decapoda, Brachyura) from Madagascar and the Seychelles. Bulletin du Muséum National d’Histoire Naturelle, Paris A, 2: 89–129. McLay, C.L. 1988. Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22: 1–403. McLay, C.L. 1993. Crustacea Decapoda: the sponge crabs (Dromiidae) of New Caledonia and the Philippines with a review of the genera. In: Crosnier, A. (ed.), Resultats de Campagnes MUSORSTOM, vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156: 111–251. McLay, C.L. 1998. A new genus and species of dromiid crab (Brachyura, Dromiidae) from the Timor Sea, north-west Australia with records of other species from the China Sea. Zoosystema 20: 339–350.

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McLay, C.L. 2001. A new genus and two new species of unusual dromiid crabs (Brachyura: Dromiidae) from northern Australia. Records of the Australian Museum 53: 1–8. McLay, C.L. 2002. Foredromia rostrata, a new genus and species of sponge crab (Decapoda, Brachyura, Dromiidae) from Southeast Asia. Crustaceana 75: 505–516.

Dynomenidae Ortmann, 1892 Dynomenids are principally tropical reef-dwellers associated with corals but two species occur well down the Australian east coast. They can be recognised by the broad triangular or convex frontal margin and the reduced last walking leg. This last leg, less than one-third the length of the leg in front, is carried alongside the carapace above the other legs and, in males only, ends in a minute claw. Of the five genera and 13 species reviewed in a family revision by McLay (1999), only three species occur in Australia (Davie, 2002). None is common. Dynomenids are thought of as primitive crabs and their relationships to dromiids, homolodromiids and other families was discussed at length by McLay (1999). Diagnosis. Carapace wider than long, convex; surface densely covered in setae; lateral borders distinct, armed with teeth. Front broadly triangular, unarmed; frontal groove well marked; cervical, postcervical, and branchial grooves present. Orbital, antennular and antennal fossae defined. Maxillipeds 3 operculiform; covering buccal cavern; separated at bases by plate at same level as sternum; basis and ischium fused. Sternal sutures 7–8 of female end well apart on low tubercles behind bases of walking legs 2. Chelipeds equal, stout; fingers hollowed at tip. Last pair of legs reduced, dactyli rudimentary, obsoletely subchelate. Abdomen 7–segmented (including telson), without pleurae; both sexes with 5 pairs of pleopods, first pair vestigial in female, last 3 pairs rudimentary in male. Telson much wider than long; retaining mechanism for abdomen present, but not effective. Uropods well-developed dorsal plates. Spermathecal apertures small and rounded, at level of genital openings. Coxa of last walking leg modified by the penis. Male first gonopod with apical plate. (abbreviated and adapted from McLay, 1999)

Key to southern Australian species of Dynomenidae 1. Carapace and chelipeds with few tubercles ...... Dynomene pilumnoidses — Carapace and chelipeds with spines ...... Hirsutodynamene spinosa

Dynomene Desmarest, 1823 McLay (1999) confined the genus to only five species, removing others to new smaller genera. Diagnosis. Carapace wider than long, smooth or minutely granulate, tomentum of short setae. Cheliped dactylus strongly curved. Walking legs with 5 or 6 spines on lower margins of dactyli. Dynomene pilumnoides Alcock, 1900 (Fig. 88a, c). Carapace with 5 conical anterolateral teeth, tomentum obscuring surface, with additional clumps of long setae. Cheliped with spines on upper surface. Red to yellowish. 28 mm. Indo-West Pacific, NSW; subtidal to 400 m depth.

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Fig. 88. Dynomenidae. Carapace: a, Dynomene pilumnoides. b, Hirsutodynomene spinosa. Outer face of right cheliped: c, Dynomene pilumnoides. d, Hirsutodynomene spinosa.

Hirsutodynomene McLay, 1999 McLay (1999) erected the new genus for two species with a ‘shaggy’ appearance. Diagnosis. Carapace wider than long, areolate, granulate and spinose near margins, with dense tomentum of short setae and tufts of longer stiff setae. Cheliped dactylus strongly curved. Walking legs with 5 or 6 spines on lower margins of dactyli. Hirsutodynomene spinosa (Rathbun, 1911) (Fig. 88b, d). Carapace with 5 anterolateral teeth, dense tomentum of short setae, each bent at tip. Cheliped with spine on inner margin. Dark brown. 33 mm. Indo-West Pacific, WA, NT, Qld, NSW (S to Port Jackson); subtidal to 15 m depth.

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. McLay, C.L. 1999. Crustacea: Decapoda: Revision of the family Dynomenidae. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM. Vol. 20. Mémoires du Muséum National d’Histoire Naturelle, Paris 180: 427–569.

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Homolodromiidae Alcock, 1900 Davie (2002) recorded two Australian species in this small family of deep-water carrier crabs. Like dromiids, dorippids, cyclodorippids and cymonomids, they have both the third and fourth walking legs elevated. The surface of the carapace is often covered with long soft hairs, especially anteriorly. The anatomy, taxonomy and relationships of the 20 species in two genera were reviewed in detail by Guinot (1995). Davie (2002) listed two Australian species to which a third is added here.

Diagnosis. Carapace longer than wide; dorsal surface sometimes hairy, branchiostegite decalcified, soft, not defined by lateral line. Eyestalks without orbit. Maxilliped 3 pediform. Cheliped with dactylus closing into cleft tip of fixed finger. Walking legs 4 and 5 reduced, lying obliquely on carapace and directed upwards, subchelate or chelate. Abdomen of both sexes of 7 somites, each with distinct pleura, males with vestigial pleopods 3–5, uropods present and ventral (not visible dorsally). (after Davie, 2002)

cde

Fig. 89. Homolidae. a, Paromolopsis boasi. b, Latreillopsis tetraspinosa (ventral). c, Homolochunia kullar. d, Homola orientalis. Homolodromiidae. e, Homolodromia kai (setae on right side only shown).

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Homolodromia A. Milne Edwards, 1880 Guinot (1995) described all four species known. Diagnosis. Walking legs 1 and 2 elongated, dactyli as long as propodi, falcate; walking legs 4 and 5 chelate. Homolodromia kai Guinot, 1993 (Fig. 89e). Cheliped dactylus without carina. Carapace smooth, with long soft hairs especially in front. 27 mm. South-western Pacific, Indonesia, NSW; 680–850 m depth. New Australian record.

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Guinot, D. 1995. Crustacea Decapoda Brachyura: révision de la famille des Homolodromiidae Alcock, 1900. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 13. Mémoires du Muséum National d’Histoire Naturelle, Paris 163: 155–282.

Subsection Homoloidea De Haan, 1839

Homolidae De Haan, 1839 Davie (2002) called these ‘deep-water carrier crabs’, because of their unusual back walking legs, held above the carapace to grasp a piece of sponge. They are rarely found at shallow shelf depths, most records being from the lower shelf or continental slope at 100 m or more. Members of this family might be confused with others with the fourth walking legs modified in this way: dromiid sponge crabs are much more spherical and latreilliids have three pairs of long spindly legs. Dorippids, cyclodorippids, cymonomids and homolodromiids have both the third and fourth walking legs elevated. The anatomy, taxonomy and relationships of the family were reviewed in detail by Guinot & Richer de Forges (1995), 14 genera and 74 species in all. The Australian fauna comprises eight species in as many genera. Five extend into southern regions (Davie, 2002). The species can be distinguished on the shape of the pseudorostral spines, called supraorbital spines by some – the pair of long spines next to the shorter rostrum.

Diagnosis. Carapace longer than wide, rectangular; dorsal surface granular to spiny; longitudinal suture line running from posterior border to antennal spine (linea anomurica or homoliine line) usually present. Front narrow; prominent slender triangular rostrum, sometimes bifid, often appearing tridentate. Eyestalks without orbit, of 2 moveable articles: first slightly longer than second, slender, conspicuous; and second stout, with terminal cornea. Antennule without socket, basal article subglobular. Antennal flagellum much longer than carapace. Interantennular septum a distinct vertical process. Maxilliped 3 pediform or suboperculiform. Chelipeds and legs long; epipods on chelipeds and walking legs 1–2. Walking leg 4 reduced, lying obliquely on carapace and directed upwards, subchelate-chelate. Abdomen of male, and usually of female, of 7 separate somites. Gills phyllobranchiate; 13–14 pairs. (after Davie, 2002)

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Key to southern Australian species of Homolidae 1. Walking leg 4 reaching to front of carapace ...... 2 — Walking leg 4 reaching well beyond front of carapace ...... 3 2. Rostrum bifid; posterolateral border of carapace lined with spines ...... Homola orientalis — Rostrum simple; posterolateral border of carapace without line of spines ...... Paromolopsis boasi 3. Pseudorostral spines multifid (antler-like) ...... 4 — Pseudorostral spines simple ...... Latreillopsis tetraspinosa 4. Walking leg 4 subchelate; pseudorostral spine less than half carapace length ...... Dagnaudus petterdi — Walking leg 4 chelate; pseudorostral spine longer than carapace length ...... Homolochunia kullar

Dagnaudus Guinot & Richer de Forges, 1995 The genus contains only one large species and was diagnosed in detail by its authors. Diagnosis. Carapace widest at midpoint. Rostrum simple, directed downwards; pseudorostral spine shorter than carapace length, with at least 2 lateral accessory spines. Eyes with cornea folded back. Walking legs cylindrical; walking leg 4 with merus reaching beyond front of carapace; subchelate. Dagnaudus petterdi (Grant, 1905) antlered crab (Fig. 90a, Pl. 18c). Dirty white. 90 mm. New Caledonia, New Zealand, Qld, NSW, Vic., Tas., SA, WA; 81–920 m depth. The species is common in deep-water dredge samples where it can be recognised by the antler-like pseudorostral spines. Accessory spination on the ‘antlers’ is variable. The species has been formerly placed in Latreillopsis and Paramola.

Homola Leach, 1815 Ten species of Homola are known, two in the Atlantic and the rest in the Indo-Pacific. The only one in Australia is widespread in the western Pacific and Indian Ocean. The bifid rostrum identifies members of the genus. Diagnosis. Carapace widest anteriorly, flat, weakly spinulose. Rostrum bifid; pseudorostral spine triangular, directed obliquely. Basal segment of eyestalk not much longer than terminal segment. Walking legs cylindrical; walking leg 4 with merus reaching about middle of carapace; subchelate. Homola orientalis Henderson, 1888 (Fig. 89d). Rostrum clearly bifid. Carapace with posterolateral margins straight, spinose; without a strong subhepatic spine; with 1 anterolateral spine. 38 mm. Indo-West Pacific, Qld, NSW, Tas.,WA (S to Mandurah); 38–650 m depth. Davie & Short (1989) summarised the literature and Guinot & Richer de Forges (1995) also reviewed this widespread species.

Homolochunia Dolfein, 1904 Guinot & Richer de Forges (1995) diagnosed the genus in detail and provided a key to the three species known, all from the Indo-West Pacific. Diagnosis. Carapace more or less parallel-sided. Rostrum simple, directed downwards; pseudorostral spine longer than carapace length, with at least 3 accessory spines. Eyes with cornea folded back. Walking legs extremely thin; walking leg 4 with merus reaching beyond front of carapace; chelate, with long fingers.

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Fig. 90. Homolidae. a, Dagnaudus petterdi. b, Latreillopsis tetraspinosa.

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Homolochunia kullar Griffin & Brown, 1976 (Fig. 89c). Pseudorostral spines diverging, longer than carapace, with 3 or 4 lateral spines and bifid tip. Walking leg 4 with pointed fingers on chela, similar to cheliped. 61 mm. SW Pacific, NSW; 500–825 m depth. The long pseudorostral spines and cheliped-like last legs differentiate this species from the more common Dagnaudus petterdi.

Latreillopsis Henderson, 1888 Seven species from the Indo-West Pacific make up this genus of small (maximum 25 mm) long- legged crabs (Guinot & Richer de Forges, 1995). Diagnosis. Carapace subquadrate. Rostrum simple; pseudorostral spine shorter than carapace length, parallel or incurving, without or with lateral spinules. Eyes with cornea folded back. Walking legs cylindrical; walking leg 4 with merus reaching to front of carapace; subchelate. Latreillopsis tetraspinosa Dai & Chen, 1980. (Fig. 90b). Pseudorostral spines diverging, as long as rostrum, without lateral spinules. Subhepatic region (front of side of carapace) especially inflated, with 4 spines (1 anterolateral spine); dorsal carapace weakly ornamented, without posterolateral spine. Maxilliped 3 distolateral corner pointed. 18 mm. WA (off Cape Naturaliste); 210–250 m depth. The Western Australian record is the first from this region.

Paromolopsis Wood-Mason, 1891 The genus is monotypic and is most similar to Homola from which it differs in having a simple rather than bifid rostrum. Diagnosis. Carapace widest posteriorly, flat, without spines. Rostrum simple; pseudorostral spine triangular, directed forward. Basal segment of eyestalk not much longer than terminal segment. Walking legs cylindrical; walking leg 4 with merus reaching about middle of carapace; subchelate. Paromolopsis boasi Wood-Mason, 1891 (Fig. 89a). 43 mm. Indo-West Pacific, WA, Qld, NSW; 284–1124 m depth. Guinot & Richer de Forges (1995) redescribed this widespread species.

References Davie, P.J.F.2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Davie, P.J.F., & Short, J.W. 1989. Deepwater Brachyura (Crustacea: Decapoda) from southern Queensland, Australia with descriptions of four new species. Memoirs of the Queensland Museum 27: 157–187. Guinot, D., & Richer de Forges, B. 1995. Crustacea Decapoda Brachyura: révision de la famille des Homolidae De Haan, 1839. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 13. Mémoires du Muséum National d’Histoire Naturelle, Paris 163: 283–517. Yaldwyn, J.C., & Dawson, E.W. 1976. First records of the crab genera Homola, Randallia,and Rochinia from New Zealand (Crustacea: Decapoda: Brachyura). National Museum of New Zealand Records 1: 91–103.

Latreilliidae Stimpson, 1858 Like some other families of deep-water crabs, latreilliids have long thin legs. The only Australian species has a small (19 mm long) body and legs about four times as long. The last legs are much shorter, held aloft and are chelate or subchelate, presumably to carry hydroids, gorgonians or other

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disguise. This behaviour is also seen several other families of podotremate crabs such as homolids; and like dromiid sponge crabs, dorippids, cyclodorippids and homolodromiids which all have the third and fourth walking legs elevated. Following the revision of Castro et al. (2003) the family consists of two genera, Latreillia with five species and Eplumula with two. Besides the species listed below, Latreillia valida De Haan, 1839 occurs in north-western Australia. Diagnosis. Carapace longer than broad, pyriform or quadrangular; dorsal surface granular to spiny; linea anomurica (suture line on side) usually present, from posterior border to antennal spine. Front narrow; prominent slender triangular rostrum, sometimes bifid, often appearing tridentate. Epistome and palate distinctly divided, palate shallowly vaulted. Eyestalks without orbit, of 2 movable articles: first much longer than second, slender, conspicuous; and second stout, with terminal cornea. Antennule without socket, basal article subglobular. Antennal flagellum not much longer than carapace. Interantennular septum a distinct vertical process. Maxillipeds 3 suboperculiform. Chelipeds and legs long and slender; without epipods. Leg 4 reduced, lying obliquely on carapace and directed upwards. Sternum of female broad. (after Davie, 2002)

Eplumula Williams, 1982 The only member of the family in southern Australia is a small deep-water spider crab with a long neck. The only other species in the genus, from Japan to the Philippines, carries seaweeds and hydroids with the last walking legs and waves these legs as a display. That species often lives on gorgonians and moves to shallow water during winter, where it becomes associated with sea anemones (Castro et al., 2003). Diagnosis. Walking leg 4 shorter than merus of walking leg 3; propodus half or less that half length of carpus; without conspicuous, feather-like setae on sides, distally flattened, broadened, often distally spinose; dactyli subchelate.

Fig. 91. Latreillidae. Eplumula australiensis.

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Eplumula australiensis (Henderson, 1888) (Fig. 91, Pl. 18d). Branchial region of carapace of adults without spine; dorsal surface of gastric region ‘neck’ without spine except in large females; ventral surface of meri of maxillipeds 3 without spine or tubercle (except in some large females. 19 mm. New Caledonia, northern New Zealand, Qld (N to Moreton Bay), NSW, Vic. (including Port Phillip Bay), Tas., SA, WA (N to Rottnest I.); shelf and slope, 48–300 m depth. The species has most often been referred to as Latreillia australiensis.

References Castro, P., Williams, A.B., & Cooper, L.L. 2003. Revision of the family Latreilliidae Stimpson, 1858 (Crustacea, Decapoda, Brachyura). Zoosystema 25: 601–634. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp.

Subsection Archaeobrachyura Guinot, 1978

Cyclodorippidae Ortmann, 1892 Cyclodorippids are small long-legged crabs from deep water which share with cymonomids, homolodromiids and dromiids the last two pairs of legs subchelate and held over the back. Homolids and latreilliids have only one pair elevated. They differ from Cymonomidae in possessing eyes (absent in cymonomids) and in having the carapace covering the segments bearing the first and second walking legs. The carapace of cymonomids tends to be more square than that of cyclodorippids. Tavares (1991, 1993) has reviewed the taxonomy of this and related families, recognising two subfamilies and ten genera. Only one of the three Australian species occurs in south- eastern Australia. Diagnosis. Carapace subcircular or subpentagonal. Orbits always present. Maxilliped 3 elongate, exopod without flagellum; covering buccal cavity. Walking legs with movable articulation between basis and ischium; first two pairs long and thin; third and fourth reduced and positioned dorsally, usually with special terminal apparatus. Wide sternal plate with short sterno-abdominal cavity more or less defined and deep; female with sternal grooves. Abdominal segments 1 visible dorsally; uropods absent. Male and female genital openings coxal; spermathecal apertures usually at level of genital openings, sometimes located more anteriorly. (adapted from Davie, 2002)

Tymolus Stimpson, 1858 In addition to the species listed below is another from north-western Australia and five more from the west Pacific and Indian oceans (Tavares, 1991, 1997; Tan & Huang, 2000). Diagnosis. Carapace subcircular, dorsal surface indistinctly separated from sides, weakly granular; front with 4 teeth or truncated, less than half as wide as carapace. Antennae longer than half carapace length. Eyestalks retractile, oriented longitudinally. Maxilliped 3 exopod without flagellum. Walking legs 1 and 2 without rows of setae. Abdomen of female broader than carapace; pleonites articulating ventrally.

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Tymolus similis (Grant, 1905) (Fig. 92). Carapace granular and setose. Front with 4 teeth, extraorbital tooth separated from front by deep sinus, without a tooth lateral to this, first tooth (hepatic) weak, second tooth (anterolateral) pointed; merus of walking leg 1 as long as carpus- dactylus. 10 mm. Qld (Cape York), NSW, Vic., Tas.; shelf and slope benthos, 90–540 m depth.

a b c

Fig. 92. Cyclodorippidae. Tymolus similis. a, habitus. b, anterior region. c, abdomen.

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Grant, F.E. 1905. Crustacea dredged off Port Jackson in deep water. Proceedings of the Linnean Society of New South Wales 1905: 312–24, pls10–11. Tan, S.H., & Huang, J.F. 2000. Description of a new species of Tymolus (Crustacea: Decapoda: Brachyura: Cyclodorippidae) from Taiwan. Pp. 135–140 in: Hwang, J.-S., Wang, C.-H., & Chan, T.- Y. (eds), Proceedings of the International Symposium on Marine Biology in Taiwan – Crustacea and zooplankton taxonomy, ecology and living resources, 27–27 May, 1998, Taiwan. National Taiwan Museum: Taipei. Tavares, M. 1991. Révision préliminaire du genre Tymolus Stimpson, avec la description de Tymolus brucei sp. nov. d’Australie occidentale (Crustacea, Brachyura, Cyclodorippoidea). Bulletin du Muséum National d’Histoire Naturelle, Paris 13: 439–456. Tavares, M. 1993. Crustacea Decapoda: Les Cyclodorippidae et Cymonomidae de l’Indo-Ouest Pacifique à l’exclusion du genre Cymonomus. In: Crosnier, A. (ed.), Résultats des campagnes MUSORSTOM, Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156: 253–313. Tavares, M. 1997. Crustacea Decapoda: Cyclodorippidae récoltés dans l’archipel de Vanuatu (Brachyura). In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, vol. 18. Mémoires du Muséum National d’Histoire Naturelle, Paris 176: 261–271.

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Cymonomidae Bouvier, 1897 Cymonomids are long-legged deep-water crabs who hold the last two pairs of legs over the back of the carapace. Other families that do the same are Dorippidae, Cyclodorippidae and Homolodromiidae. Homilids and latreilliids have only the last pair of legs elevated in this way. Members of this family have been treated as part of Cyclodorippidae (refer to this family for differences) and to Tavares (1993a, b) for a discussion of the taxonomy of the genera. The family is represented in Australia by three species in deep water of the eastern coast (Ahyong & Brown, 2003). These last authors have provided a key to 15 Indo-West Pacific species of the family.

Diagnosis. Carapace subquadrate, longer than wide; orbits absent. Maxillipeds 3 elongate, completely covering buccal cavity; exopod of maxilliped 3 with flagellum. Walking legs without movable articulation between basis and ischium; first two pairs long and thin subequal; third and fourth legs reduced and positioned dorsally. Female with sternal grooves. First abdominal segments visible dorsally; uropods absent. Genital openings coxal, female on third legs, male on fifth legs. (adapted from Davie, 2002)

Key to Australian species of Cymonomidae 1. Abdomen with 6 segments; rostrum and lateral frontal projections triangular ...... Cymonomoides delli — Abdomen with 7 segments; rostrum and lateral frontal projections cylindrical ...... 2 2. Carapace, abdomen and legs covered with large stalked globose club-like tubercles; with large hemispherical boss on each posterolateral border ...... Cymonomus kapala — Carapace, abdomen and legs minutely granulate; without boss on posterolateral border ...... Cymonomus soela

Cymonomoides Tavares, 1993 There are four species, most from depths 600 m or more (Lemaitre & Bermúdez, 2000). Diagnosis. Rostrum absent or very short. Eyestalk immoveable, well calcified, fused at bases, without cornea. Chelipeds subequal. Walking legs 1 and 2 long, with flattened dactyli; walking legs 3 and 4 with recurved dactyli. Abdomen of 7 free segments (including telson). Cymonomoides delli (Griffin & Brown, 1976) (Fig. 93b). 9 mm. NSW (off Sydney), Vic., Tas.; 110–675 m depth.

Cymonomus Milne Edwards, 1880 Cymonomus is restricted to species with six-segmented abdomen. Two species occur in Australia. Diagnosis. Rostrum cylindrical. Eyestalk immoveable, well calcified, fused at bases, without cornea. Chelipeds subequal. Walking legs 1 and 2 long, with flattened dactyli; walking legs 3 and 4 with recurved dactyli. Abdomen of 6 free segments (including telson). Cymonomus kapala Ahyong & Brown, 2003 (Fig. 93c). Carapace, abdomen and legs covered with large stalked globose club-like tubercles. Eyestalks fused with carapace, demarcation completely obscured. Carapace with large hemispherical boss on each posterolateral border. 3 mm. NSW; 176 m depth. Cymonomus soela Ahyong & Brown, 2003 (Fig. 93a). Carapace, abdomen and legs granular and tuberculate. Eyestalks fused with carapace, demarcation distinct. Carapace without hemispherical boss on posterolateral border. 7 mm. Tas.; 940–990 m depth.

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Fig. 93. Cymonomidae. a, Cymonomus soela. b, Cymonomoides delli (carapace, with eyestalks). c, Cymonomus kapala (carapace with eyestalks, abdominal somite 1).

References Ahyong, S., & Brown, D.E. 2003. New species of Cymonomus from southeastern Australia (Brachyura, Cymonomidae) with a key to the Indo-West Pacific species. Crustaceana 75: 1363–1374. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Griffin, D.J.G., & Brown, D.E. 1976. Deepwater decapod Crustacea from eastern Australia: brachyuran crabs. Records of the Australian Museum 30: 248–271.

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Lemaitre, R., & Bermúdez, A. 2000. A new cyclodorippoid crab of the genus Cymonomoides Tavares, 1993 (Crustacea: Decapoda: Brachyura: Cymonomidae) from the Caribbean coast of Colombia. Proceedings of the Biological Society of Washington 113: 974–979. Tavares, M. 1993a. Description préliminaire de quatre nouveaux genres et trois nouvelles espèces de Cyclodorippoidea Américains (Crustacea, Decapoda, Brachyura). Vie et Milieu 43: 137–144. Tavares, M. 1993b. Crustacea Decapoda: Les Cyclodorippidae et Cymonomidae de l’Indo-Ouest Pacifique à l’exclusion du genre Cymonomus. In: Crosnier, A. (ed.), Résultats des campagnes MUSORSTOM, Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156: 253–313.

Raninidae De Haan, 1839 The Raninidae are a predominantly tropical family, the members of which are unusual in having a carapace longer than wide, and the abdomen, which is not completely reflexed beneath the carapace as in most other brachyurans. This most uncrab-like shape gives them the common name, frog crabs, and in New Caledonia I have heard children refer to them as ‘tortue’,turtle. This name describes them well. One representative, the spanner crab, Ranina ranina, is a large edible species more frequently seen in fish markets in Queensland than in the wild. Frog crabs live buried in sandy sediments on beaches or on the shelf. The Raninidae are generally assumed to retain a number of primitive characters (assuming that the Brachyura were derived from a more shrimp- or crayfish-like, long tailed, decapod ancestor). Among the primitive characters listed by Bourne (1922) are the presence of a rostrum, the oviduct openings on the coxae of pereopods 3, and the nervous system. Further discussions of the systematics of the Raninidae can be found in S˘tevcˇic´ (1973) but the classification of Brachyura has moved on since their work. Twelve genera of raninids are known worldwide to which can be added a further 21 fossil genera (Dawson & Yaldwyn, 1994, 2000; Tucker, 1998). Guinot (1993) recognised six subfamilies. Four species occur in southern Australia. Other genera and five other species of raninids occur in tropical waters (Davie, 2002). Tyndale-Biscoe & George (1962) and Goeke (1986) are the best available references to identify specimens outside the range of this book. A key to all Recent genera can be found in Dawson & Yaldwyn (2000). The key here is more practical.

Diagnosis. Carapace longer than wide, widest in anterior third. Chelipeds equal, flattened, with fingers nearly at right angles to propodus. Walking legs with flattened propodi and dactyli, last pair of raised above plane of other legs. Abdomen of 7 somites, incompletely reflexed, first 4–5 abdominal somites visible dorsally.

Key to southern Australian species of Raninidae 1. Carapace widest across front, frontal margin with distinct rostrum and three toothed lobes on each side; surface of carapace with numerous anteriorly-directed flattened scales; large species (cl. up to 150 mm) ...... Ranina ranina — Carapace widest back from posterior margin, frontal margin narrow, tridentate (with pointed rostrum and two lateral teeth); surface of carapace smooth or weakly tuberculate; small species (cl. up to 30 mm) ...... 2 2. Carapace widest one-third back, tapering posteriorly, and anteriorly to narrow 3-toothed front ...... Lyreidus tridentatus — Carapace widest about quarter way back, ovate posteriorly, and with broad front ...... 3 3. Carapace about twice as long as greatest width, margin posterior to spines evenly convex; eyestalks folding transversely ...... Notosceles serratifrons

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— Carapace about 1.3 times as long as greatest width, margin posterior to spines parallel and then tapering; eyestalks folding obliquely and ventrally ...... Umalia trirufomaculata

Lyreidus De Haan, 1841 The three species of this genus, from the tropical Atlantic and warmer Indo-West Pacific, are those best known as frog crabs (Griffin, 1970; Goeke, 1986; Dawson & Yaldwyn, 2000). Diagnosis. Carapace elongate-oval, widest one-third way back, anterolateral margins gradually convergent, laterally and dorsally convex, surface smooth and polished. Frontal margin less than half maximum carapace width; rostrum simple, orbit spine with 1 fissure; anterolateral spine well separated. Eyestalks folding transversely, short, broad at base, cornea small. Maxillipeds 3 without oblique ridge. Walking legs 1 and 2 with narrow triangular dactyli, walking leg 3 with dactylus expanded close to base; walking leg 5 very short and arising above and anterior to third pair.

Fig. 94. Raninidae. a, Lyreidus tridentatus. b, Notosceles serratifrons. c, Ranina ranina. d, Umalia trirufomaculata.

321 Marine Decapod Crustacea of Southern Australia

Lyreidus tridentatus De Haan, 1841 (Fig. 94a, Pl. 18e). Carapace with anterolateral spines, evenly tapering anteriorly to equally-triangular rostrum and ocular angles. Carpus of cheliped with 2 spines on upper margin. Colour in life orange, pale cream when preserved. 50 mm. New Zealand, western Pacific, southern Qld, NSW, Vic., northern Tas., Bass Strait, SA, central WA (S to Rottnest I.) ; sandy sediments, 27–380 m depth. Little is known of the habits of L. tridentatus, but since there are no posterior branchial orifices, L. tridentatus probably lies buried in the sediment with only the anterior tip of the carapace exposed.

Notosceles Bourne, 1922 Six species distributed through the Indo-West Pacific comprise this genus (Dawson & Yaldwyn, 2000). The cheliped has a double carina on the upper margin of the hand. Diagnosis. Carapace elongate-oval, widest one-quarter way back, anterolateral margins short, concave, laterally and dorsally convex, surface finely granulate. Frontal margin half maximum carapace width; rostrum trifid, orbital spine with 2 fissures; anterolateral spine separated by distance equal to rostrum-orbital spine. Eyestalks folding transversely, short, broad at base, cornea small. Maxillipeds 3 without oblique ridge. Walking legs 1 and 3 with oval dactyli, walking leg 2 with dactylus pointed; walking leg 5 short and arising above and anterior to fourth pair. Notosceles serratifrons (Henderson, 1893) (Fig. 94b). Rostrum triangular, inner orbital angle a serrated spine, supraorbital spine acute and almost as long as external orbital spine, margin between external orbital and anterolateral spines serrate or spinose. Lower margin of cheliped with 3 spines, carpus serrate dorsally. Indian Ocean, Japan, WA (S to Rottnest I.); shelf depths. The species was formerly referred to the genus Raninoides.

Ranina Lamarck, 1801 The genus is monotypic although other species names have been used. Diagnosis. Carapace shield-shaped, wider than long, widest across front, covered with elongate anteriorly directed flattened scales. Frontal margin comprising rostrum and 2 3-dentate lobes on each anterolateral corner. Eyestalks folding transversely, 3–segmented. Maxillipeds 3 without oblique ridge. Walking legs similar, with flattened blade-like dactyli. Ranina ranina (Linnaeus, 1758) spanner crab (Fig. 94c). Orange to red. 150 mm. Indo-West Pacific, WA (S to Abrolhos Is.), NT, Qld, NSW (S to Port Hacking); sandy bottoms, intertidal to 121 m depth. Ranina ranina is a distinctive crab; examination of the figure should enable quick identification and specimens are unlikely to be confused with any other crab. Males have large chelipeds and more developed anterolateral plates on the carapace. The flattened legs, especially the last pair, are conspicuous adaptations for digging in the soft sediments which this crab usually inhabits. Respiratory openings are present both at the front and back of the carapace and R. ranina can therefore apparently lie buried in the sediment either ‘head up’ or ‘tail up’ (Bourne, 1922). The spanner crab is good eating, and available in fish markets, especially in Queensland (Brown, 1986). It is marketed in southern states and is popular throughout South-East Asia.

Umalia Guinot, 1993 Guinot (1993) placed six Indo-West Pacific species of Ranilia into her new genus, restricting the latter to American species; Davie & Short (1989) provided a key to these six. Diagnosis. Carapace elongate, widest near front, anterolateral margins parallel-sided over anterior half then convergent, dorsally convex, surface finely tuberculate. Frontal margin about half

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maximum carapace width; rostrum simple, orbit spine with 2 fissures; anterolateral spine well separated. Eyestalks folding obliquely, short, broad at base, cornea small. Maxillipeds 3 without oblique ridge. Walking legs 1 and 2 with narrow triangular dactyli, walking leg 3 with dactylus truncate, posteriorly convex; walking leg 5 short and arising dorsally. Umalia trirufomaculata (Davie & Short, 1989) (Fig. 94d). Carapace with 3 red dorsal spots, lateral pair bigger than median one, width three-quarters length, fronto-orbital distance less than half width of carapace. Chela with sharp distal spine on upper border of palm. Qld, WA (possibly not further south than Abrolhos Is); 70–270 m depth.

References Bourne, G.C. 1922. The Raninidae: a study in carcinology. Journal of the Linnean Society of London (Zoology) 35: 25–78, pls 4–7. Brown, I.W. 1986. South Queensland’s spanner crabs –a growing fishery. Australian Fisheries 45: 3–7. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Davie, P.J.F., & Short, J.W. 1989. Deepwater Brachyura (Crustacea: Decapoda) from southern Queensland, Australia with descriptions of four new species. Memoirs of the Queensland Museum 27: 157–187. Dawson, E.W., & Yaldwyn, J.C. 1994. The frog crabs (Crustacea: Brachyura: Raninidae) – a species list and bibliographic index. Occasional Papers of the Hutton Foundation of New Zealand 6: 1–60. Dawson, E.W., & Yaldwyn, J.C. 2000. Description and ecological distribution of a new frog crab (Crustacea, Brachyura, Raninidae) from northern New Zealand waters, with keys to Recent raninid genera and Notosceles species. Tuhinga 11: 47–71. Goeke, G.D. 1986. Decapod Crustacea: Raninidae. In: Crosnier, A. Résultats des campagnes MUSORSTOM I et II. Philippines (1976, 1980) Tome 2. Mémoires du Muséum National d’Histoire Naturelle, Paris 133: 205–228. Griffin, D.J.G. 1970. A revision of the recent Indo-West Pacific species of the genus Lyreidus De Haan (Crustacea, Decapoda, Raninidae). Transactions of the Royal Society of New Zealand, Biological Sciences 12: 89–112. Guinot, D. 1993. Données nouvelles sur les Raninoidea De Haan, 1841 (Crustacea Decapoda Brachyura Podotremata). Comptes Rendus de l’Académie des Sciences, Paris, Sciences de la vie 316: 1324–1331. S˘tevcˇic´, Z. 1973. The systematic position of the family Raninidae. Systematic Zoology 22: 625–632. Tucker, A.B. 1998. Systematics of the Raninidae (Crustacea: Decapoda: Brachyura), with accounts of three new genera and two new species. Proceedings of the Biological Society of Washington 111: 320–371. Tyndale-Biscoe, M., & George, R.W. 1962. The Oxystomata and Gymnopleura (Crustacea, Brachyura) of Western Australia with descriptions of two new species from Western Australia and one from India. Journal of the Royal Society of Western Australia 45: 65–96.

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Section Eubrachyura de Saint Laurent, 1980 The Eubrachyura, or true ‘true crabs’,include two groups, Heterotremata and Thoracotremata.

Subsection Heterotremata Guinot, 1977 The heterotreme crabs are the largest group of crabs. The male genital openings are coxal or coxosternal while the those of the female are sternal. Not all families are represented in southern Australia and there are other views of family versus subfamily ranks for some groups. The families appear in an order that best reflects relationships and is the same as that of Martin & Davis (2001).

Dorippidae MacLeay, 1838 Dorippids are thought to be among the more primitive brachyurans. Distinguishing features of members of this family are the thinner last two pairs of legs, held dorsally and with hooked terminal articles. These enable the crabs to carry a shell or some other object over the carapace, a behaviour which gives the family the name, carrier crabs. Most species are tropical or subtropical, and most occur on soft sediments at continental shelf depths but some occur in deeper water down to about 3000 m. Of the two subfamilies the Ethusinae are represented only in deep water off tropical Australia (see Chen, 1993 for several species off New Caledonia; Davie, 2002). Four species in two genera of Dorippinae have been recorded in Australia. Nine genera and possibly 50 species occur worldwide. Manning and Holthuis (1981) provided keys to all genera and this was updated in their more recent taxonomic account (Holthuis & Manning, 1990) which includes a key to Indo-West Pacific genera and species. Diagnosis. Body longer than wide; subquadrate to rounded, widest posteriorly. Antennules and antennae large; antennules folded obliquely. Endostome produced to form gutter; covered distally with widened distal portion of endopod of maxilliped 1. Maxillipeds 3 not covering anterior part of buccal cavern. Chelipeds with chelae robust; subequal. Walking legs 1 and 2 usually long and stout; last 2 pairs differing from preceding legs, reduced, positioned dorsally, and subchelate. Flat thoracic sternum intercalated between legs; wide, with all sutures between somites 4 and 8 interrupted; sternite 8 visible dorsally, or partly under carapace. Spermathecae absent, paired seminal receptacles being part of genital ducts. First 2 abdominal somites not folded ventrally, visible in dorsal view; uropods absent. (adapted from Davie, 2002)

Paradorippe Serène & Romimohtarto, 1969 Holthuis & Manning (1990) provided a more extensive diagnosis and a key to the four species known, including good illustrations and discussion of the Australian species. Diagnosis. Carapace lateral margins without epibranchial spine, inner suborbital tooth much smaller than exorbital tooth. Walking legs 2 and 3 dactyli without fringes of hairs. Paradorippe australiensis (Miers, 1884) (Fig. 95). Carapace flattish, sparsely, if at all, covered with minute pubescence; granules small, situated near posterior-lateral margins. Walking legs without pubescence. Male pleopod 1 two-thirds length of abdomen. 22 mm. Irian Jaya, Qld (S to Moreton Bay), WA (S to Cockburn Sound); soft sediments, to 22 m depth. Another Australian crab with two pairs of short and clawed posterior legs is Cymonomoides delli, whose carapace is more rectangular, and whose environmental is more shelf edge and slope.

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Fig. 95. Dorippidae. Paradorippe australiensis.

References Chen, H.L. 1993. Crustacea Decapoda: Dorippidae of New Caledonia, Indonesia and the Philippines. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156: 315–345. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Holthuis, L.B., & Manning, R.B. 1990. Crabs of the subfamily Dorippinae MacLeay, 1838, from the Indo-West Pacific region (Crustacea: Decapoda: Dorippidae). Researches on Crustacea, Carcinological Society of Japan Special Number 3: 1–151. Manning, R.B., & Holthuis, L.B. 1981. West African brachyuran crabs. Smithsonian Contributions to Zoology 306: 1–379. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124.

325 Marine Decapod Crustacea of Southern Australia

Calappidae De Haan, 1833 The Calappidae are a distinctive family with highly modified claws which are used to open mollusc shells. Calappids, or box crabs, occur predominantly in tropical shelf and reef habitats and include nine genera and upwards of 20 species. Calappids are shallow-water animals and are usually found on sandy bottoms. Most records are from shallower than 200 metres. Many authors (e.g., Sakai, 1976; S˘tevcˇic´, 1983) use the family name Calappidae in a wider sense than used here and had difficulty in defining it unambiguously. They divided a larger family into four subfamilies: Calappinae, Hepatinae, Matutinae and Orithyinae. Bellwood (1996) reviewed usage and concluded on the basis of a phylogenetic analysis that the family, sensu lato, was polyphyletic as earlier authors had suspected. She showed that the Calappinae and Hepatinae are related to each other but that the Matutinae are closer to Leucosiidae and Orithyinae to Dorippidae. The subfamilies are treated as families here as she suggested and as followed by Davie (2002) Two genera and seven species of Calappidae s.s. occur occasionally in southern Australia. A third genus, Paracyclois, is known from a single specimen (P. grandispinis Etheridge & McCulloch, 1916), a subfossil recovered from dredging operations in clay sediments 6–12 m below the sea floor from Coode Canal, Yarra River Delta, Melbourne. No indications were given as to the likely age of the deposits in which the specimen was found. The only other species in this genus is P. milnedwardsi Miers, 1886, known from north of the Admiralty Islands and Japan, at 80–285 m. Four additional species occur in northern Australian waters and the following key should only be used with caution on material from waters north of Perth and Sydney. Tyndale-Biscoe & George (1962) and Sakai (1976) should be consulted to identify specimens collected from tropical waters of Australia but in some cases names have changed. The most recent taxonomic treatments of calappids on a worldwide basis are by Balss (1957, for a traditional treatment), Galil & Clark (1996, for two genera not known in Australia) and Bellwood (1996, for a phylogenetic treatment).

Diagnosis. Carapace subcircular or transversely ovate; with strongly developed posterolateral expansion entirely or partially covering walking legs. Antennules folding obliquely or nearly vertically. Antennae small. Maxilliped 3 not elongate or acute, not completely closing buccal cavern, not concealing palp when resting; exopod broad. Afferent branchial channels opening in front of bases of maxilliped 3; efferent branchial channels together forming deep channel in endostome, covered by pair of lamellar processes from maxilliped 1. Chelipeds subequal; chelae high, robust, curved, shutting closely against pterygostome; one chela with large hooked tooth near base of gape fitting into cup-like depression on fixed finger. Walking legs with dactyli tapered and pointed, not flattened. Male abdomen with some somites fused. Female openings sternal; male openings coxal. Male gonopod 1 stout, tapered; male gonopod 2 thin, longer than first gonopod, short to long, flagellum may be straight, curved, or crotchet-shaped. (adapted from Davie, 2002)

Key to southern Australian genera and species of Calappidae 1. Lateral margin of carapace with single prominent spine; posterior margin of carapace without thin wing-like projections ...... Mursia curtispina — Lateral margin of carapace without prominent spines; posterior margin of carapace produced into thin extensions under which the walking legs can be hidden . . Calappa … 2 2. Carapace surface smooth posteriorly ...... 3 — Carapace surface, especially posterolateral extensions, dorsally ridged ...... 5 3 Anterior half of carapace with rounded tubercules surrounded by red or pink pigmented rings ...... Calappa japonica — Anterior half of carapace smooth ...... 4 4. Posterolateral margin of carapace continuously beaded with 4 asymmetrical, unevenly-

326 Brachyura – crabs

spaced teeth on each side, separated by shallow indentations; posterior margin unarmed ...... Calappa lophos — Posterolateral margin of carapace with 3 acute teeth on each side, separated by deep indentations; posterior margin of carapace with 9 acute evenly-spaced symmetrical teeth separated by deep indentations ...... Calappa philargius 5. Length of carapace about two-thirds maximum width ...... Calappa hepatica — Length of carapace more than three-quarters maximum width ...... Calappa depressa

Calappa Weber, 1795 Calappa is the most highly modified genus with remarkable pincers whose mechanical advantage enables the crab to crack open mollusc shells (Shoup, 1968). Those species of Calappa which have been studied have been found to eat both molluscs and hermit crabs. Species of Calappa bury themselves swiftly in sandy substrates using the chelipeds to dig (S˘tevcˇic´, 1983). Other species of Calappa not described below occur in northern Australia and can be identified using Tyndale- Biscoe & George (1962) and Sakai (1976). Diagnosis. Posterior half of carapace with lateral wing-like extensions, partly or completely covering walking legs. Maxillipeds 3 do not meet in midline, leaving mandibles exposed. Merus of maxilliped 3 not elongate or acute, never concealing flagellum. Chelipeds dissimilar, right cheliped with dactylus reduced and modified as a small hooked finger. Upper margin of propodus of chelipeds with high denticulate crest. Male pleopod 1 half to two-thirds length of abdomen. Male pleopod 2 about three-quarters length of pleopod 1. Calappa depressa Miers, 1886 little crested crab (Fig. 96b). Length of carapace more than three- quarters maximum width. Anterior two-thirds of carapace strongly tuberculate, posterior third less sculptured but with low transverse beaded ridges across short wing-like extensions. Posterolateral margin of carapace almost smooth, with only 3–4 indistinct shallow teeth on each side. East Africa to New Caledonia, NSW (off Port Jackson), Tas., SA, north-west WA; 2–350 m depth, mostly from soft sediments. Calappa hepatica (Linnaeus, 1758) reef box-crab (Figs 96a, 97c). Length of carapace about two- thirds maximum width. Anterior two-thirds of carapace strongly tuberculate, posterior third less sculptured but with low transverse beaded ridges across definite wing-like extensions. Posterolateral margin of carapace almost smooth, with only 3–4 indistinct shallow teeth on each side. Indo-West Pacific, north-west WA, NT, NSW, Tas.; sand and rubble substrates, intertidal to 100 m depth. Calappa hepatica is similar to C. depressa but more widespread and more commonly collected. Calappa japonica Ortmann, 1892 Japanese box-crab (Figs 96d, 97d). Length of carapace about three-fifths maximum width. Anterior half of carapace covered with rounded tubercules. Tubercules covered with fine, flat granules and surrounded by red-pink pigmented rings. Posterior surface of carapace smooth, with vestigial round or oval tubercules. Walking legs not completely hidden beneath posterior expansions of carapace. Posterolateral margin of carapace with 8 large teeth on each side. Southern Africa, India, Japan, Qld, WA (off Rottnest I.); sand and shell bottoms, 30–380 m depth. Calappa japonica and C. depressa are distinguished from all other Australian species by the poorly developed posterolateral expansions of the carapace (Jones, 1989). The two species are quite dissimilar in other respects and the tubercules on the anterior region of the carapace and the large teeth around the posterior margin of the carapace readily distinguish C. japonica.

327 Marine Decapod Crustacea of Southern Australia

c b

Fig. 96. Calappidae. Carapace: a, Calappa hepatica. b, Calappa depressa. c, Calappa hepatica. d, Calappa japonica. e, Calappa lophos.

328 Brachyura – crabs

b a

d c

Fig. 97. Calappidae. Carapace: a, Calappa philargius. b, Mursia curtispina. Cheliped, lateral: c, Calappa hepatica. d, Calappa japonica. e, Calappa lophos. f, Calappa philargius. g, Mursia curtispina.

329 Marine Decapod Crustacea of Southern Australia

Calappa lophos (Herbst, 1785) red-streaked box-crab (Figs 96e, 97e). Length of carapace about two-thirds maximum width. Carapace almost completely smooth, except for several small bumps near eyes. Posterolateral margin of carapace slightly convex, continuously beaded and with 4 asymmetrical, unevenly-spaced teeth on each side, separated by shallow indentations. Posterior margin unarmed. Crest of propodus of chelipeds with prominent teeth separated by deep indentations. Posterior carapace and chelae with pattern of red-orange lines and spots. Indo-West Pacific from Natal to Japan, Qld, NSW (S to Sydney), WA (S to Fremantle), SA (isolated record); sandy sediments, 5–150 m depth. Calappa philargius (Linnaeus, 1758) red-spotted box-crab, spectacled box-crab (Fig. 97a, f, Pl. 18f). Length of carapace about two-thirds maximum width (three-quarters in small specimens). Carapace smooth except for beaded region near posterior margin, single large red spot on propodus, carpus of chelae and surrounding orbits. Posterolateral margin of carapace with 3 distinct teeth on each side, separated by deep indentations. Posterior margin of carapace almost straight with 9 distinct evenly-spaced symmetrical teeth separated by deep indentations. (Small specimens, 17–23.5 mm, with numerous distinct bumps and points on anterior two-thirds of carapace and lacking red spots.) Indo-Pacific from Persian Gulf to Japan, WA (S to Swan R.), NT, Qld, NSW (S to Port Jackson); sandy sediments, 8–120 m depth. Whitelegge (1900) referred to this species as Calappa cristata.

Mursia Desmarest, 1823 Galil (1993) presented a more detailed diagnosis of the genus and provided a key to all 15 species then known; Crosnier (1997) added another species from the south-west Pacific. Mursia microspina Davie & Short, 1989 is known from tropical Australia to Japan. Diagnosis. Carapace with median spine or prominent tubercle on each side. Propodus of cheliped swollen, granulose, upper margin crested and dentate. Maxilliped 3 merus not elongate or acute, never concealing flagellum, ischium with a granulate row forming a stridulating organ with milled ridge on dactylus of cheliped. Walking legs not flattened, with styliform dactyli. Carina on abdominal somite trilobate. Mursia curtispina Miers, 1886 (Fig. 97b, Pl. 18g). Lateral spines each about quarter of maximum carapace width (excluding spines). Posterior margin with triangular lateral teeth and small median tooth. Cheliped merus with 3 spines, distal one as long as lateral; carpus with weak granules; propodus with 9 tubercles in 3 diagonal rows and 3 more at base of serrate upper palmar crest, lowest row with proximal tubercle sharp, lower margin serrate. Live colour unknown. Western Pacific from Sumatra to Japan, Fiji, WA (S to Rottnest I.), NT, Qld, NSW (S to Cronulla); 80–500 m depth.

References Balss, H. 1957. Decapoda. Dr. H.G. Bronn’s Klassen und Ordnungen des Tierreichs 1: 1505–1672. Bellwood, O. 1996. A phylogenetic study of the Calappidae H. Milne Edwards 1837 (Crustacea: Brachyura) with a reappraisal of the status of the family. Zoological Journal of the Linnean Society 118: 165–193. Crosnier, A. 1997. Une espèce nouvelle de Mursia de Nouvelle-Calédonie (Crustacea, Decapoda, Brachyura, Calappidae). Zoosystema 19: 151–158. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp.

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Galil, B.S. 1993. Crustacea Decapoda: a revision of the genus Mursia Desmarest, 1823 (Calappidae). In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 10. Mémoires du Muséum National d’Histoire Naturelle, Paris 156: 347–379. Galil, B.S., & Clark, P.F. 1996. A revision of Cryptosoma Brullé, 1837 and Cycloes De Haan, 1837 (Crustacea: Brachyura: Calappidae). Zoological Journal of the Linnean Society 117: 175–204. Jones, D.S. 1989. Calappa japonica Ortmann, 1892, a new record for Western Australia (Decapoda, Brachyura, Oxystomata). Crustaceana 56: 133–140. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates. Shoup, J.B. 1968. Shell opening by crabs of the genus Calappa. Science 160: 887–888. S˘tevcˇic´ , Z. 1983. Revision of the Calappidae. Memoirs of the Australian Museum 18: 165–171. Tyndale-Biscoe, M., & George, R.W. 1962. The Oxystomata and Gymnopleura (Crustacea, Brachyura) of Western Australia with descriptions of two new species from Western Australia and one from India. Journal of the Royal Society of Western Australia 45: 65–96. Whitelegge, T. 1900. Scientific results of the trawling expedition of H.M.C.S. ‘Thetis’ off the coast of New South Wales in February and March, 1898. Crustacea. Part I. Memoirs of the Australian Museum 4: 135–199, pls 32–35.

Leucosiidae Samouelle, 1819 The Leucosiidae, aptly known as nut crabs or pebble crabs, are a readily recognisable family of small crabs that are common on soft sediments from the intertidal shore to shelf and slope depths. A study of the decapod crustaceans of the Australian north-west shelf found leucosiids to be the most diverse of all families (Ward & Rainer, 1988). Many leucosiids are apparently nocturnal; during the day they can be found partly buried in the sediment. At night they forage for invertebrates, though they also scavenge dead material (Schembri, 1982). There are 40 extant genera and at least 300 species, plus numerous extinct taxa with a fossil record extending back to the Eocene (Balss, 1957). Leucosiids are most speciose in the tropical Indo-Pacific. Davie (2002) allocated the 75 Australian species to four subfamilies: Cryptocneminae, Ebaliinae, Leucosiinae and Philyrinae but their differentiation is difficult and historically has been extremely fluid (Manning & Holthuis, 1981). In members of Cryptocneminae the carapace is broad and flattened, with lamelliform lateral expansions concealing all or part of walking legs. Leucosiinae have maxilliped 3 with merus shorter than half the length of the ischium, the chelipeds with a swollen palm, and a discoid female abdomen. The remaining two families, which both have the merus of maxilliped 3 half as long or longer than the ischium, are more difficult to distinguish. Ebaliinae have a pronounced ‘head’, or frontal and orbital region differentiated from the body by a lateral depression. The buccal cavern (covered by the endopod of maxilliped 3) reaches beyond the end of the inhalant channel (covered by the exopod of maxilliped 3). The infraorbital lobe is prominent and merges with the lateral border of the inhalant channel (Fig. 98a). Philyrinae, on the other hand have a more circular body, with only a shallow lateral depression at the base of the orbit. The buccal cavern does not end much in advance of the anterior margin of the inhalant channel which itself extends under the lower orbital margin (Fig. 98b). Ten genera and 30 species occur in southern Australian waters, reflecting the essentially tropical diversity of this family. The thoracic sinus in species of Leucosia is a groove on the lateral- underside of the carapace beginning in front of the base of the cheliped and serves as a channel for water to the gills; its margins are often granular. Tyndale-Biscoe & George (1962) provided a key to Australian genera, including tropical forms and discussed some species in detail. Several species have moved to other genera since their publication. Barnard (1950) and Sakai (1976) also provided useful introductions to the generic classifi-

331 Marine Decapod Crustacea of Southern Australia

cation of leucosiids. There are more modern references useful in identifying eastern Pacific taxa, some of which occur in tropical Australia (Chen, 1989; Tan, 1996; Tan & Ng, 1996; Galil, 2001a, b, 2003).

Diagnosis. Carapace subcircular, ovoid, pyriform or pentagonal; regions often not defined; smooth or armed with spines or lobes. Eyes and orbits small. Frontal region narrowed and variously produced anteriorly. Antennules folding obliquely. Antennae small, sometimes obsolete. Maxillipeds 3 elongate, completely closing buccal cavern; palp arising from groove near mesial edge of concealed surface, hidden when maxillipeds closed; exopod broad. Afferent branchial channel opening at bases of maxilliped 3, running along side of buccal cavity, covered by exopod of maxilliped 3. Efferent branchial channel covered by lamellar process of maxilliped 1. Chelipeds symmetrical, robust and elongate. Walking legs with dactyli simple. Abdomen with some somites fused in both sexes. Female genital openings sternal; male openings coxal or sternal. Male gonopod 2 short. (after Davie, 2002)

Key to southern Australian genera of Leucosiidae 1. Carapace with flat lateral projections partly covering walking legs; upper and lower surfaces of carapace separated by a sharp edge ...... Cryptocneminae ...2 — Carapace more or less spherical or sculptured, lacking flat posterolateral projections (may have spines or stout thickened tubercles) ...... 4 2. Carapace about as wide as long, with prominent midlongitudinal and paired oblique ridges ...... Paranursia abbreviata ...p.342 — Carapace wider than long, flat, or with elevated gastric region ...... 3 3. Frontal region of carapace with convex anterior projection between orbits, with single prominent angular projection on each hepatic margin; margins smooth ...... Cryptocnemus vincentianus ...p.333 — Frontal region of carapace with narrow truncate anterior projection between orbits; lateral margins expanded, square anteriorly and almost semicircular posteriorly; margins beaded ...... Leucisca levigena ...p.333 4. Dorsal surface of carapace highly polished; frontal region produced anteriorly; thoracic sinus present; maxilliped 3 with merus shorter than half length of ischium measured along inner border; chelipeds with palm swollen; female abdomen enlarged, discoid ...... Leucosiinae Leucosia ...p.336 — Carapace variously sculptured or smooth, but rarely highly polished; maxilliped 3 with merus half as long or longer than ischium measured along inner border; chelipeds with palm cylindrical, elongate or moderately swollen; female abdomen not discoid ...... 5 5. Buccal cavern (covered by the endopod of maxilliped 3) reaches beyond end of inhalant channel (covered by the exopod of maxilliped 3); anterior part of carapace differentiated from rest by a lateral depression; infraorbital lobe prominent, merging with lateral border of inhalant channel (Fig. 98a); epistome deep in Merocryptus ...... Ebaliinae … 6 — Buccal cavern not ending much in advance of anterior margin of inhalant channel; anterior part of carapace differentiated from rest by only a shallow lateral depression; inhalant channel extending under lower orbital margin (Fig. 98b) ...... Philyrinae … 7 6. Carapace convoluted, gastric regions each with dominating stout conical tubercle; posterior margin with hemispherical tubercle; with 3 distinct projections on each hepatic margin; chelipeds and walking legs strongly tuberculate ...... Merocryptus lambriformis ...p.336

332 Brachyura – crabs

— Carapace more or less spherical, dorsum and margins weakly tuberculate, spinose or smooth ...... Ebalia ...p.335 7. Front bilobed or with single medial V-shaped indentation; carapace with spines or tubercles ...... Arcania ...p.339 — Front bilobed or with 4 small teeth, medial gap narrow; carapace smooth or with granules 8 8. With 3 posterior conical projections of carapace on one horizontal plane ...... Randallia eburnea ...p.343 — Posterior projections absent, or not in one plane if present ...... 9 9. With conical posterior projection above 2 posterolateral projections; carapace granular ...... Myra mammillaris ...p.340 — Without conical projections, or with pairs of triangular lateral projections, weakest an teriorly; carapace smooth or weakly tuberculate ...... 10 10. Roof of orbit with 2 sutures ...... Bellidilia ...p.340 — Roof of orbit with 1 suture ...... Philyra ...p.342

Subfamily Cryptocneminae Stimpson, 1907

Cryptocnemus Stimpson, 1858 Cryptocnemus is one of only two cryptocnemine genera found on the southern Australian coast. Worldwide there are at least 14 species. Only one of the five Australian species is known in this region, and that from only a single specimen. Diagnosis. Carapace pentagonal or rounded, much wider than long and moderately flattened, highest in gastric region, frontal region with convex anterior projection between orbits, posterolateral margins expanded, lamelliform and projecting over part of the walking legs. Orbits small. Outer antennae obsolete or nearly so. Apex of maxilliped 3 sharply triangular. Cryptocnemus vincentianus Hale, 1927 tortoise-crab (Fig. 98c). Carapace 1.5 times as wide as long, minutely pitted, with rounded triangular anterior projection and dorsal median carina; hepatic margin with obtuse angular projection; branchial margin rounded; posterior margin rounded. 7.3 mm. SA (Gulf St Vincent); sublittoral (not recorded since first described).

Leucisca MacLeay, 1838 Two species exist, one on each side of the Indian Ocean (George & Clark, 1976). Diagnosis. Semicircular flattened carapace, with truncate front, with raised beaded margin. Leucisca levigena George & Clark, 1976 (Fig. 98d). Gastric region of carapace elevated; pterygos- tomium and sternum smooth. 5 mm. WA (Yanchep Reef); 1 m depth.

333 Marine Decapod Crustacea of Southern Australia

abe

Fig. 98. Leucosiidae. Left anterior region, ventral: a, Ebalia intermedia. b, Bellidilia laevis. Cryptocneminae. c, Cryptocnemus vincentianus. d, Leucisca levigena. Ebaliinae. e, Ebalia tuberculosa. f, Merocryptus lambriformis. g, Ebalia crassipes. h, Ebalia dentifrons. i, Ebalia intermedia. j, Ebalia spinifera.

334 Brachyura – crabs

Subfamily Ebaliinae Stimpson, 1871

Ebalia Leach, 1817 Australian species of Ebalia have sometimes been referred to the genus or subgenus Phlyxia, finally placed in full synonymy by Manning & Holthuis (1981). Numerous species inhabit the Indo-West Pacific and Atlantic oceans, especially in tropical regions, eleven in Australia. Species are separated on the pattern of major spines and tubercles but juveniles are often more finely spinose than adults. Species of Ebalia use the complex mouthparts to break up their prey, small invertebrates that are captured by the chelae probed into the sediment (Schembri, 1982). Diagnosis. Carapace rhomboidal or nearly so, convex, regions well delineated, surface uneven, tuberculate or carinate; front narrow, produced, margin with 2 or 4 teeth between orbits. Orbits open to antennal fossa. Exopod of maxilliped 3 distinctly shorter than endopod. Abdominal somites 3–6 fused (sometimes 3–5 in male). Male often with tooth at base of telson projecting over abdominal somite 6.

Key to southern Australian species of Ebalia 1. Frontal region with deep median indentation and 2 broad slightly convex tuberculate projections ...... Ebalia tuberculosa — Frontal region with 4 distinct teeth between eyes, frontal margin not strongly tuberculate . . 2 2. Posterior margin of carapace rounded, without any conical or angular projections, anterolateral margin strongly swollen and separated from lateral projection by notch; carapace densely covered with low circular granules ...... Ebalia dentifrons — Posterior margin of carapace with 1 or more projections, anterolateral margin weakly swollen; carapace smooth or covered with slender spines ...... 3 3. Carapace with broad median posterior projection; posterolateral angles rounded ...... Ebalia crassipes — Carapace with angular or conical posterior and posterolateral projections ...... 4 4. Carapace with slender spines, clearly protruding beyond margins ...... Ebalia spinifera — Carapace smooth or granular, with weak triangular angles along margins . . Ebalia intermedia Ebalia crassipes (Bell, 1855) (Fig. 98g). Carapace with 3 or 4 sublateral tubercles, apparently smooth but minutely punctate; hepatic region with slight projection; lateral margins with 3 weak projections, most-posterior projection more widely separated from middle one than anterior one; midposterior projection triangular, posterolateral angles rounded. Male without teeth on abdominal somite 6 or telson; abdominal somite 1 much smaller than 2. 13 mm. NSW (S of Port Stephens) Vic., Tas., SA, WA; sandy and muddy sediments, 4–99 m depth. Ebalia dentifrons Miers, 1886 (Fig. 98h, Pl. 19c). Carapace, excluding frontal region immediately behind eyes, densely covered with low circular granules; hepatic region with prominent bulbous projection; lateral margins with 3 prominent evenly-spaced projections; posterior margin evenly rounded, without angular posterolateral projections. Male with indistinct tooth on abdominal somite 6, directed away from telson. 13 mm. NSW, Vic., SA, WA (N to Dongarra); rocky rubble and sand, intertidal to 102 m depth. The species is especially common in WA. Ebalia intermedia Miers, 1886 smooth nut-crab (Fig. 98i, Pl. 19d). Carapace smooth; hepatic region with obtuse projection; lateral margins with 3 low projections, most-posterior projection more widely separated from middle one than anterior one; posterior margin with median and 2 posterolateral conical projections. Male without teeth on abdominal somite 6 or telson. 13 mm. NSW, Vic., Tas., SA, WA (N to Cockburn Sound); sediments, intertidal to 549 m depth. Ebalia ramsayi (Haswell, 1879) may be a junior synonym on the evidence of its short description.

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Ebalia spinifera Miers, 1886 (Fig. 98j). Carapace covered with slender spines; hepatic region with spiniferous projection; lateral margin with unevenly spaced projections; posterior margin with median cone and 2 posterolateral angles. 10 mm. Qld (S of Cape Moreton), NSW (S to Port Jackson); 6–92 m depth. Ebalia tuberculosa A. Milne Edwards, 1873 nut-crab (Fig. 98e). Carapace strongly sculptured and densely covered with flat-topped and conical granules, with 5 tuberculate projections, with medially notched frontal region; hepatic region with obtuse projection; lateral margins without projections; posterior margin with median cone and strong posterolateral angles. Chelipeds, walking legs and abdomen covered with mosaic of flattened tubercles. Abdomen of male with tooth on telson projecting over abdominal somite 6.10 mm. Indo-West Pacific; NSW, Vic., SA, WA (W to Eucla); 28–800 m depth. The sculptured and strongly convex carapace and indented (not 4-toothed) frontal margin should distinguish this common species from all other southern Australian leucosiids. On the basis of its short description, Balss’ (1935) photograph of the type, and the same type locality, Ebalia quadrata (Milne Edwards, 1873) is a possible junior synonym.

Merocryptus Milne Edwards, 1873 The genus contains a single species, M. lambriformis. Diagnosis. Carapace rhomboidal, strongly but unevenly convex dorsally and heavily tuberculated, concave in hepatic and posterior branchial regions; midlateral margin with wing-like posterolateral projections, convex dorsally, flattened on ventral surfaces; frontal region pronounced, bidentate between orbits. Merocryptus lambriformis Milne Edwards, 1873 (Fig. 98f, Pl. 19e). 14 mm. West Pacific, New Zealand, New Caledonia, Qld, NSW, SA, WA; sandy sediments, 28–274 m depth. The strongly sculptured carapace and wing-like posterolateral projections should be sufficient to distinguish this crab from all other southern Australian leucosiids. Cryptocnemus vincentianus has more rounded lateral lobes on the carapace. Both might be confused with parthenopid crabs.

Subfamily Leucosiinae Samouelle, 1819

Leucosia Weber, 1795 Leucosia would appear to be the only genus in the subfamily. Its species are readily recognised by the highly polished carapace and the thoracic sinus, which is unique to the genus. The thoracic sinus is a lateral groove on the carapace beginning over the base of the cheliped and extending along the side and variously beaded (Figs 99f–h). Over 40 species are known, almost all from the tropical Indo-Pacific. In Australia 17 named species are listed (Davie, 2002). Tyndale-Biscoe & George (1962) and Sakai (1976) remain useful works to identify specimens from waters north of Sydney and Perth and for any tropical species that may stray into southern waters. Diagnosis. Carapace with projecting frontal region, without lateral expansions, with polished dorsal surface. Thoracic sinus present. Maxilliped 3 with merus shorter than half length of ischium measured along inner border. Chelipeds with palm swollen. Male abdomen with somites free. Female abdomen enlarged, discoid.

Key to southern Australian species of Leucosia 1. Dense pubescent patch on lateral margin of carapace and on adjoining posterior margin of merus of cheliped; carapace with c. 40 evenly-spaced small red spots Leucosia haematosticta

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— No pubescence on surface of carapace or cheliped ...... 2 2. Granules on anterolateral margin of carapace continue posteriorly to level of last pair of walking legs ...... Leucosia unidentata — Granules on anterolateral margin of carapace continue to level of first or second pair of walking legs ...... 3 3. Hepatic region of carapace granulated; front tridentate, with median lobe produced into prominent lobe in front of eyes ...... Leucosia anatum — Hepatic region of carapace smooth; front tridentate, without prominent lobe ...... 4 4. Thoracic sinus deep, continuing to last pair of walking legs ...... Leucosia pubescens — Thoracic sinus deep only to level of first pair of walking legs ...... Leucosia perlata Leucosia anatum (Herbst, 1783) (Fig. 99a). Front of carapace rounded, convex and produced beyond eyes, without pubescence; hepatic region with prominent bulge; epibranchial angle (between anterolateral and posterolateral margins) moderately pronounced. Carapace hepatic region granulated; granules on anterolateral margin continue posteriorly to level of walking legs 1. Thoracic sinus with 3–5 white granules. Male with abdominal somites 3–5 fused, somite 2 extremely small. Female with abdominal somites 3–6 fused, somites 1 and 2 similar in size. Carapace with white spots and variable red rings and patterns. 30 mm. Indo- West Pacific, WA (S to Exmouth Gulf) NT, Qld, NSW (S to Port Jackson), SA; soft sediments, 2–80 m depth. Leucosia anatum has many junior synonyms and has been recorded in southern Australia as L. splendida and L. australiensis. Part of the confusion about the identity of this widely distributed species arises from the range of colour patterns observed. Tyndale- Biscoe & George (1962) recognised four colour morphs, with Australian material recognised as ‘morph C’. Leucosia haematosticta Adams & White, 1849 (Fig. 99b). Front of carapace tridentate, median frontal tooth twice as long as lateral teeth; dense patch of pubescence on posterolateral margin of carapace at base of cheliped and on adjoining proximal end of merus of cheliped; hepatic region smooth, without bulge; epibranchial lobe (between anterolateral and posterolateral margins) of carapace strong. Male with abdominal somites 2–5 fused. Female with abdominal somites 2–6 fused. Carapace with 40 or more evenly spaced small red spots; chelae, legs and ventral surface also with small red spots. 15 mm. Indo-West Pacific, WA (S to Shark Bay) NT, Qld, NSW; sediments, 18–100 m depth. Leucosia perlata De Haan, 1841 (Fig. 99b, g). Front of carapace tridentate, without pubescence; hepatic region smooth, without granules, with prominent bulge; epibranchial angle (between anterolateral and posterolateral margins) moderately pronounced; granules on anterolateral margin of carapace continue posteriorly to level of walking legs 1 or 2. Thoracic sinus deep only to level of walking legs 1. Male with abdominal somites 3–5 fused, somite 2 small. Female with abdominal somites 3–6 fused. A pair of dark spots on cardiac region of carapace. 24 mm. Indo-West Pacific, WA (S to Cockburn Sound), Qld; subtidal. Leucosia pubescens Miers, 1877 (Fig. 99c, h). Front of carapace tridentate; without pubescence; hepatic region not granulated, with prominent bulge; epibranchial angle (between anterolateral and posterolateral margins) of carapace not pronounced; granules on anterolateral margin continue posteriorly to level of walking legs 1. Thoracic sinus deep, reaching level of walking leg 4. Male and female with abdominal somites 3–6, fused somite 2 extremely small in male. Light blue with uniform spots and markings of greenish brown, chelipeds and walking legs banded greenish brown. 25 mm. Indo-West Pacific, WA (Shark Bay to Swan R.); soft sediments, 4–11 m depth. Leucosia pubescens (which is not pubescent!) could be confused with L. anatum but can be distinguished by the tridentate front without a prominent lobe and the blue and green colouration.

337 Marine Decapod Crustacea of Southern Australia

g ch

ch a

c b

i j

Fig. 99. Leusosiidae Leucosiinae. a, Leucosia anatum. b, Leucosia haematosticta. c, Leucosia perlata. d, Leucosia pubescens. e, Leucosia unidentata. Thoracic sinus (ch=cheliped base): f, Leucosia anatum (ventral right side). g, Leucosia perlata (lateral left side). h, Leucosia pubescens (lateral left side). Philyrinae. i, Arcania fungilifera. j, Arcania undecimspinosa.

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Leucosia unidentata De Haan, 1841 (Fig. 99e). Front of carapace weakly tridentate; without pubescence; hepatic region granulated, with prominent bulge; epibranchial angle (between anterolateral and posterolateral margins) of carapace not pronounced; granules on anterolateral margin continue posteriorly to level o walking legs 1. Thoracic sinus with 3–5 white granules. Male with abdominal somites 4–6 fused, somites 1 and 2 full width. Female with abdominal somites 3–6 fused, somites 1 and 2 similar in size. 2 pairs of large red spots in centre of carapace. 20 mm. Indo-West Pacific, Qld, WA (S to Rottnest I.); soft sediments, 30–150 m depth. Specimens from WA and identified as Leucosia unidentata differ from this species principally in the structure of the male pleopod, which has 3 full twists and a short tip and may represent an undescribed species.

Subfamily Philyrinae Rathbun, 1913

Arcania Leach, 1817 This genus of usually ornamented crabs with a notched front is distributed in the Indo-West Pacific. Galil (2001a) revised the genus and provided a key to 14 species of which seven occur in Australia. Diagnosis. Carapace about as wide as long (widest near midpoint), variously sculptured, not highly polished, with marginal pointed spines, sometimes 1 pair extremely long, always with median posterior spine. Frontal region of carapace with single indentation or concave excavation, 2 distinct teeth between orbits.

Key to southern Australian species of Arcania 1. Carapace with 1 pair of strong laterally directed spines ...... 2 — Carapace with several pairs of short spines or without spines ...... 4 2. Posterior spine upcurved, with 2 pairs of posterolateral spines ...... Arcania septemspinosa — Posterior spine straight, with 1 pair of posterolateral spines ...... 3 3. Infraorbital lobe incurved, reaching frontal eave; cheliped fingers twice as long as palm; posterior spine sharp, posterolateral spines flattened ...... Arcania gracilis — Infraorbital lobe not reaching frontal eave; cheliped fingers half as long as palm; posterior spine sharp, posterolateral spines flattened ...... Arcania cornuta 4. Lateral and posterior margins of carapace with 10 spines covered with and partly obscured by tubercles ...... Arcania fungilifera — Lateral and posterior margins of carapace with 11 spines covered only basally with smaller teeth ...... Arcania undecimspinosa Arcania cornuta (MacGilchrist, 1905) (Fig. 101f). Carapace with elongate lateral spines, longer than other spines (sometimes blunt), posterior median spine short and not upturned, posterolateral spines conical. Infraorbital lobe not reaching frontal eave. Cheliped fingers half as long as palm. 33 mm. Indo-West Pacific, Qld, WA (off Cape Naturaliste, 140 m); 28–204 m depth. The species was known as Ixoides cornutus until Ixoides was synonymised with Arcania by Galil (2001a). A specimen in WAM is a new record for WA; it has acute lateral spines while some individuals have blunt cylindrical projections (Chen, 1989: fig. 20). Arcania fungilifera Galil, 2001 (Fig. 99i). Lateral margins of carapace with 5 pairs of triangular spines; carapace with numerous stalked tubercles. 12 mm. WA; to 113 m depth.

339 Marine Decapod Crustacea of Southern Australia

Arcania gracilis (Henderson, 1893). Carapace with elongate lateral spines, longer than other spines, posterior median spine long and not upturned, posterolateral spines flattened. Infraorbital lobe incurved, reaching frontal eave. Cheliped fingers twice as long as palm. 18 mm. Indo-West Pacific, WA; 21–366 m depth. Arcania septemspinosa (Fabricius, 1787) (Fig. 101a). Carapace with elongate lateral spines, much longer than other spines, posterior median spine long, sharp and upturned, posterolateral spines acute; with medial granulate ridge running to posterior spine. 27 mm. Indo-West Pacific, NT, Qld, NSW; 7–182 m depth. Arcania undecimspinosa De Haan, 1841 (Fig. 99j). Lateral and posterior margins of carapace with 11 spines covered only basally with smaller teeth. 29 mm. Indo-West Pacific, Qld, NSW, Vic., SA, WA; 7–420 m depth.

Bellidilia Kinahan, 1856 The species assigned to this genus have a complex taxonomic history, both southern Australian species known for many years as members first of Ebalia and then of Philyra (Griffin, 1972), then transferred to a new genus Dittosa (Tan, 1995). But this is an objective synonym of Bellidilia to which they must belong (Davie, 2002). The differences between Bellidilia and Philyra involve male gonopods, male abdomen and orbital sutures (Tan, 1995). Diagnosis. Ebaliinae with front of carapace quadridentate between orbits; roof of orbit with 2 sutures. Carapace dorsally smooth, with 8 or more white dots, lateral margins with 3 low angular projections, with distinct medial posterior spine or ridge. Male with paired lateral swellings basally on each side of fused abdominal somites 3–6 and bump on somite 6 at juncture with telson. Male gonopods 1 and 2 almost equal in length.

Key to southern Australian species of Bellidilia 1. Frontal teeth of carapace unevenly spaced (gap between 2 inner teeth less than outer gap); posterior margin of carapace with 3 distinct teeth ...... Bellidilia undecimspinosa — Frontal teeth of carapace evenly spaced; posterior margin of carapace evenly rounded, smooth and without teeth ...... Bellidilia laevis Bellidilia laevis (Bell, 1855) smooth pebble-crab (Figs 100a, 101c, Pl. 19a). Indentations between apices of all frontal teeth similar in width; posterior margin smoothly rounded. Abdomen of female with somites 2–6 fused. 25 mm. Vic. (E to Corner Inlet), SA, WA (W to Albany); soft sediments, intertidal to 3 m depth. In South Australia, the smooth pebble crab breeds twice a year and reaches maturity after only 4 months. The crabs are active while the tide is ebbing and flowing but remain buried in the sediment at high tide and when exposed (Hale, 1926; McKillup & McKillup, 1994). Bellidilia undecimspinosa Kinahan, 1856 large pebble-crab (Fig. 101b, d, Pl. 19b). Indentation between apices of inner frontal teeth narrower than indentations between these and lateral teeth. Posterior margin of carapace almost straight, with 3 distinct teeth. Abdomen of female with somites 3–6 fused. 38 mm. NSW (S of Lake Macquarie), Vic., Tas., SA (W to Murray R.): soft sediments, 4–95 m depth. Philyra murrayensis (Rathbun, 1923), recorded from South Australia by Hale (1927) is a junior synonym (Griffin, 1972).

Myra Leach, 1817 The Indo-Pacific genus Myra contains about a dozen species revised by Galil (2001b). Tyndale- Biscoe & George (1962) and Campbell & Stephenson (1970) recorded five species from Australia, mostly from tropical waters but Galil (2001b) added others. A single species, M. mammillaris,is collected occasionally in southern Australia but appears to have a spotty distribution.

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a b

Fig. 100. Leucosiidae Philyrinae. a, Bellidilia laevis. b, Myra mammillaris. c, Paranursia abbreviata (habitus and ventral views from Bell, 1885: pl. 34 fig 5). d, Philyra platycheir. e, Philyra scabriuscula. f, Arcania cornuta.

341 Marine Decapod Crustacea of Southern Australia

Diagnosis. Carapace subglobular, with median spine higher than posterolateral spines; front produced anteriorly, truncate, and bidentate between orbits. Opposing edges of cheliped armed with fine teeth only. Myra mammillaris Bell, 1855 (Fig. 100b). Carapace margin beaded; hepatic region with raised, beaded ridge; lateral margin rounded, without angular projections; posterior margin with 3 spines. Carapace and chelipeds evenly covered with numerous globules. WA (S to Swan R.), Qld (S to Keppel Bay), Tas., SA; intertidal to 36 m depth; sandy sediments.

Paranursia Sèrene & Soh, 1976 Diagnosis. Philyrinae with flattened carapace; front broadly pointed; carapace with oblique ridges running from midlongitudinal ridge across epibranchial regions; without hepatic ridges; posterior margins not bilobed. Cheliped with granular ridge on upper and lower edges. Paranursia abbreviata (Bell, 1855) (Fig. 100c). 10 mm. India, Thailand, Qld, NSW; subtidal. The dorsal carapace pattern immediately distinguish this species, rare in southern Australia.

f e

Fig. 101. Leucosiidae Philyrinae. a, Arcania septemspinosa. b, c, Bellidilia undecimspinosa (with anterior region). d, Bellidilia laevis (anterior region). e, Philyra orbicularis. f, Randallia eburnea.

Philyra Leach, 1817 The carapace in species of Philyra frequently has a beaded margin. Balss (1957) listed 33 species of Philyra but some of these have been removed to other genera (Tan, 1995). Two of the Australian species reviewed by Tyndale-Biscoe & George (1962) now belong in Bellidilia. Diagnosis. Carapace without distinct medial posterior spine or ridge; roof of orbit with 1 suture. Carapace dorsally smooth, lateral margins with 3 low angular projections. Male abdominal somites 3–6 fused,. Male gonopod 1, 2 or 3 times as long as 2. Male abdominal somites 5–6 with limited mobility, suture notched laterally, 6 wider than 5.

342 Brachyura – crabs

Key to southern Australian species of Philyra 1. Fixed finger of cheliped with fringe of hair ...... Philyra platycheir — Fixed finger of cheliped without fringe of hair ...... 2 2. Surface of carapace smooth, polished ...... Philyra orbicularis — Surface of carapace (excluding anterior-most region) covered with low rounded granules ...... Philyra scabriuscula Philyra orbicularis (Bell, 1855) (Fig. 101e). Carapace front bidentate between orbits; margins without angular projections or teeth, beaded laterally and posteriorly. Cheliped fingers without fringe of hairs. Male abdomen with somites 2–6 fused, sides parallel until extreme distal region. Female abdominal somites 3–6 fused. 21 mm. NSW, WA (S to Rottnest I.); 12–35 m depth. Philyra platycheir De Haan, 1841 (Fig. 100d). Carapace front shallowly concave between orbits; margins beaded, most prominently on each side which each have about 20 larger marginal beads; carapace dorsally shiny; hepatic region with beaded ridge; branchial and posterior regions with indistinct granulations. Cheliped fixed finger with fringe of hairs. 14 mm. Indo- West Pacific, Qld, NSW, WA (S to Cockburn Sound); 35–150 m depth. The fringe of hairs on the cheliped are conspicuous and distinguish this species from all other Australian leucosiids. Philyra scabriuscula (Fabricius, 1798) (Fig. 100e). Carapace front divided by deep groove; margins crenellate; surface of carapace (excluding anterior-most region) covered with low rounded granules; hepatic region with obscure ridge. Cheliped fingers denticulate. 15 mm. Indian Ocean, WA; intertidal and subtidal.

Randallia Stimpson, 1857 Chen (1989) provided a key to the six species of Randallia and figured the only Australian species The alignment of the three posterior projections of carapace on one horizontal plane is characteristic. Diagnosis. Carapace about as wide as long, lacking posterolateral projections (carapace widest near midpoint). Carapace variously sculptured or smooth but not highly polished. Frontal region of carapace with a single indentation or concave excavation; lateral margins rounded, smooth or with a row of evenly-spaced small granules. All 3 posterior projections of carapace on 1 horizontal plane. Randallia eburnea Alcock, 1896. (Fig. 101f). Body not tomentose, regions indistinct; posterior border with 3 tubercles; lateral border with 1 tubercle. Chelipeds elongate. 33 mm. Indo-West Pacific, Qld, WA; muddy sediments, 30–210 m depth.

References Balss, H. 1935. Brachyura of the Hamburg Museum expedition to south-western Australia, 1905. Journal of the Royal Society of Western Australia 21: 113–151. Balss, H. 1957. Decapoda. Dr. H.G. Bronn’s Klassen und Ordnungen des Tierreichs 1: 1505–1672. Barnard, K.H. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38: 1–837. Bell, T. 1855. Horae carcinologicae, or notices of crustacea. I. A monograph of the Leucosiidae, with observations on the relations, structure, habits and distribution of the family; a revision of the generic characters; and descriptions of new genera and species. Transactions of the Linnean Society of London 21: 277–314,pls30–34. Campbell, B.M., & Stephenson, W. 1970. The sublittoral Brachyura (Crustacea: Decapoda) of Moreton Bay. Memoirs of the Queensland Museum 15: 235–301, pl.22.

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Chen, H. 1989. Leucosiidae (Crustacea, Brachyura). In: Forest, J. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144: 181–264. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Galil, B. 2001a. A revision of the genus Arcania Leach, 1817 (Crustacea: Decapoda: Leucosioidea). Zoologische Mededelingen, Leiden 75: 169–206. Galil, B. 2001b. A revision of Myra Leach, 1817 (Crustacea: Decapoda: Leucosioidea). Zoologische Mededelingen, Leiden 75: 409–446. Galil, B. 2003. Four new genera of leucosiid crabs (Crustacea: Brachyura: Leucosiidae) for three new species and nine species previously in the genus Randallia Stimpson, 1857, with a description of the type species, R. ornata (Randall, 1939). Proceedings of the Biological Society of Washington 116: 395–422. George, R.W., & Clark, M. 1976. Two new species of pebble crab (Oxystomata: Leucosiidae) from Western Australia. Records of the Western Australian Museum 4: 303–309. Griffin, D.J.G. 1972. Brachyura collected by Danish expeditions on south-eastern Australia (Crustacea, Decapoda). Steenstrupia 2: 49–90. Hale, H.M. 1926. Habits of the smooth pebble-crab (Philyra laevis). South Australian Naturalist 8: 67–69. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Manning, R.B., & Holthuis, L.B. 1981. West African brachyuran crabs. Smithsonian Contributions to Zoology 306: 1–379. McKillup, S.C., & McKillup, R.V. 1994. Reproduction and growth of the smooth pebble crab Philyra laevis (Bell 1885) at two sites in South Australia during 1990–91. Transactions of the Royal Society of South Australia 118: 245–251. Miers, E.J. 1886. Report on the Brachuyra collected by H.M.S. Challenger during the years 1873–76. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76. Zoology 17: 1–362 pls 1–29. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates pp. Schembri, P.J. 1982. The functional morphology of the feeding and grooming appendages of Ebalia tuberosa (Pennant) (Crustacea: Decapoda: Leucosiidae). Journal of Natural History 16: 467–480. Tan, C.G.S. 1995. Dittosa, a new genus of leucosiid (Crustacea: Decapoda: Brachyura) from southern Australia and New Zealand. Proceedings of the Biological Society of Washington 108: 465–476. Tan, C.G.S. 1996. Leucosiidae of the Albatross expedition to the Philippines, 1907–1910 (Crustacea: Brachyura: Decapoda). Journal of Natural History 30: 1021–1058. Tan, C.G.S., & Ng, P.K.L. 1996. A revision of the Indo-Pacific genus Oreophorus Rüppell, 1830 (Crustacea: Decapoda: Brachyura: Leucosiidae). Pp. 101–189 in: Richer de Forges, B. (ed.) Les fonds meubles des lagons de Nouvelle-Calédonie (sédimentologie, benthos). ORSTOM Editions: Paris. Tyndale-Biscoe, M., & George, R.W. 1962. The Oxystomata and Gymnopleura (Crustacea, Brachyura) of Western Australia with descriptions of two new species from Western Australia and one from India. Journal of the Royal Society of Western Australia 45: 65–96. Ward, T.J., & Rainer, S.F. 1988. Decapod crustaceans of the North West Shelf, a tropical continental shelf of north-western Australia. Australian Journal of Marine and Freshwater Research 39: 751–765.

Matutidae De Haan, 1841 Matutids are flat circular-bodied crabs with all the walking legs flattened for swimming and burrowing in sand. Many are highly coloured and patterned. The family is predominantly tropical but three can be found in southern Australia. The behaviour of species of Matuta is not well

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known but one species has been observed to consume polychaete worms, shrimps and amphipods as well as molluscs and hermit crabs (Stevcic, 1983). Species of Matuta can bury themselves in sandy substrates using the flattened walking legs which are also used for swimming. Many species are brightly coloured. The Matutidae have traditionally been regarded as a subfamily of Calappidae, albeit a rather ill-fitting one (S˘tevcˇic´, 1983), but the historical account and phylogenetic analysis of Bellwood (1996) provided evidence that they were more closely related to Leucosiidae. The diagnosis is based on the autapomorphies defined by her. Four genera are included in the Matutidae, three in Australia (Galil & Clark, 1994). The family tends to be tropical but two members of one genus occur in southern Australia.

Diagnosis. Carapace more or less circular, with spine at most lateral point. Antennal 1 basal article less than 5% of orbital width and not reaching the inner orbital gap. Chelipeds equal; merus with row of 3–4 spines on lower margin. Walking legs flattened, especially distally. Male gonopod 1 with distal finger-like dorsal process. Male genital openings coxal.

Key to southern Australian species of Matutidae 1. Cheliped with midpalmar ridge oblique, outer dactylar ridge in male strongly milled throughout; carpus of walking leg 3 with 1 keel on upper edge ...... Matuta planipes — Cheliped with midpalmar ridge parallel to lower margin, outer dactylar ridge in male slightly milled distally only; carpus of walking leg with 2 keels on upper edge ...... Ashtoret … 2 2. Carapace with red spots, each with pale centre, except in curved band around a red patch at base of lateral spine; lateral spine 0.15 as long as width of carapace . . . . . Ashtoret granulosa — Carapace with red spots, more dense anteriorly, propodi and dactyli of walking legs with red patches, no red patch at base of lateral spine; lateral spine 0.2 as long as width of carapace ...... Ashtoret lunaris

Ashtoret Galil & Clark, 1994 Galil & Clark (1994) separated eight species from Matuta to their new genus and provided a key. Diagnosis. Carapace with strong lateral spine. Cheliped with midpalmar ridge parallel to lower margin, dactylar ridge in male tuberculate distally or smooth. Carpus of walking leg with 2 keels on upper edge. Abdominal somite 3 carinate. Ashtoret granulosa (Miers, 1877) armed crab (Fig. 102a, b). Lateral spine 0.15 as long as width of carapace. Carapace with midposterolateral tubercle. Outer surface of propodus of cheliped coarsely granulate. Carapace with red spots, each with pale centre, except in curved band around a red patch at base of lateral spine. 58 mm. Indonesia, Tahiti, WA (S to Fremantle), Qld (S to Port Curtis), isolated record from Port Adelaide, SA; intertidal to 55 m depth. Ashtoret lunaris (Forskål, 1775) (Fig. 102c, e, Pl. 19f). Lateral spine 0.2 as long as width of carapace. Carapace with midposterolateral tubercle. Outer surface of propodus of cheliped a 5-lobed ridge, second and fourth prominent. Carapace with red spots, more dense anteriorly, propodi and dactyli of walking legs with red patches, no red patch at base of lateral spine. 55 mm. Indian Ocean, WA, Qld, NSW (S to Green Cape); intertidal to 15 m depth. In tropical Queensland where this species is common it feeds on small crustaceans and gastropods, preferring larger prey like hermit crabs when adult (Perez & Bellwood, 1988).

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d c b

f e

Fig. 102. Matutidae. a, b, Ashtoret granulosa. c, Ashtoret lunaris. d, Matuta planipes. Face of left chela: e, Ashtoret lunaris. f, Matuta planipes.

Matuta Weber, 1795 Galil & Clark (1994) restricted what was a larger genus to just three species. Diagnosis. Carapace with strong lateral spine. Cheliped with midpalmar ridge oblique, dactylar ridge in male strongly tuberculate throughout. Carpus of walking leg 3 with 1 keel on upper edge. Abdominal somite 3 carinate. Matuta planipes Fabricius, 1798 flower moon-crab, reticulated surf-crab (Fig. 102d, f) Front of carapace with lateral lobes nearly straight. Cheliped with lower proximal angle of palm tuberculate, without a laterally-directed spine; midpalmar ridge with a prominent proximal spine. Carapace with reticulating brown lines forming small rings anteriorly and larger loops posteriorly. 61 mm. Indo-West Pacific, WA,NT,Qld, NSW (S to Sydney region); subtidal to 40 m depth.

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References Bellwood, O. 1996. A phylogenetic study of the Calappidae H. Milne Edwards 1837 (Crustacea: Brachyura) with a reappraisal of the status of the family. Zoological Journal of the Linnean Society 118: 165–193. Galil, B.S., & Clark, P.F. 1994. A revision of the genus Matuta Weber, 1795 (Crustacea: Brachyura: Calappidae). Zoologische Verhandelingen, Leiden 294: 1–55. Perez, O.S., & Bellwood, O. 1988. Ontogenetic changes in the natural diet of the sandy shore crab Matuta lunaris (Forskal) (Brachyura: Calappidae). Australian Journal of Marine and Freshwater Research 39: 193–199. S˘tevcˇic´ , Z. 1983. Revision of the Calappidae. Memoirs of the Australian Museum 18: 165–171.

Majidae Samouelle, 1819 Commonly, majids are known as spider crabs or decorator crabs because of their long thin legs and their ability to camouflage themselves. Naturally, in a family so large as this one not all species have long legs and not all use camouflage. Nevertheless, these are useful common names for many seen in southern Australia. Leptomithrax gaimardii is a typical long-legged species, a large spider crab with a body up to 165 mm long, a leg-span of about 700 mm, and often reported by divers in large numbers in a few metres depth along the Victorian and Tasmanian coasts. Species of Notomithrax are smaller, typical decorator crabs with relatively short legs, but furnished with strong hooked setae on the carapace and legs used by the crab to attach disguise in the form of pieces of algae, bryozoans and sponges. While they too are moderately abundant even in the lower intertidal they are difficult to find in the algal habitats they favour. Members of this family are usually best recognised by the pear-shaped body, tapering anteriorly to a more or less prominent rostrum, often bifurcated. Many species have a spiny carapace and the eyestalks are usually protected by spines whose arrangement is critical to their classification. Of course there are exceptions, some species are flattened, others are more rectangular, and a few are poorly spinose. The chelipeds are never massive, usually only slightly stronger and shorter than the walking legs. Some families that might be confused with Majidae are Hymenosomatidae in shallow water and some primitive families in the deep sea. Hymenosomatids generally have a flattened carapace and never a complex orbit. Primitive crabs such as Homolidae, Homolodromiidae, Cymonomidae, Dorippidae and Cyclodorippidae have the last one or two pairs of walking legs held over the top of the carapace and are largely confined to outer shelf or slope waters. Majid crabs are especially diverse in tropical Indo-Pacific waters and 63 genera and over 150 species are known from Australia (Davie, 2002). About half of the species occur in southern Australia. While a few species can be found in the lower intertidal, usually camouflaged species in beds of algae, most are from shelf depths on sedimentary and reef environments. Majids are one of the few families of crabs well represented on the continental slope, especially by some large spider crabs. The taxonomy of the family is fluid with at least 50 family or subfamily names proposed (Davie, 2002). Griffin & Tranter (1986) and the earlier works by Desmond Griffin are widely accepted as the authoritative studies on the taxonomy of this complex family, especially in Australia and New Zealand. They recognised seven subfamilies and some more recent authors have elevated some of these to family rank (four families in Clark & Webber, 1991 on the basis of larval studies). Martin & Davis (2001) discussed the history of various views and concluded with a superfamily Majoidea and six families which don’t overlap completely with those of Griffin & Tranter. Early studies of numerous majid species reared in the laboratory shed little light on the phylogeny of the family (e.g. Webber & Wear, 1981) but more recent work casts doubt on the

347 Marine Decapod Crustacea of Southern Australia

monophyly of some of the subfamilies/families. Building on studies by Marques & Pohle (1988), these same authors used larval features to investigate cladistic relationships of the subfamilies (Pohle & Marques, 2000; Marques et al., 2003). On the basis of 21 majid genera, and in the second study of seven mithracine genera, they concluded that the Oregoninae, Majinae and Inachinae are monophyletic, the last only if genera assigned to Inachoidinae are included. They did not believe that Epialtinae, Pisinae or Mithracinae are monophyletic but remained uncertain of precise relationships. The degree of protection offered to the eyestalk by ridges, spines and lobes is the critical feature distinguishing them. This key to subfamilies follows earlier ones (Griffin, 1966a; Griffin & Tranter, 1986) which non-specialists find exceptionally hard to master. This is in part because of the difficulty in understanding the terminology applied to the orbit and the armature surrounding it. It is essential to clean hairs or camouflaging algae and encrusting fauna away from the rostrum and orbit of decorating species before attempting an identification. Read the couplets carefully and refer to the illustrations (Figs 103–118). The eyestalk sits on the side of the carapace and is either clearly visible from above and below, without an orbit and unprotected by spines, or sits in an orbit. The orbit comprises four components which together offer the eye various degrees of concealment. First, dorsally is a supraorbital eave (se) extending laterally to more or less obscure the eye from above. The supraorbital eave may be simple but more often is ornamented anteriorly by a preorbital spine (pre) at the front and an antorbital spine (ant) behind. Second is a postorbital spine (post) or lobe sitting behind the eyestalk and sometimes cupped to receive it. The third component is the so-called intercalated spine (int) which lies between the supraorbital eave and the postorbital spine. The orbit is fully completed underneath only in Mithracinae where the basal antennal article (baa) is expanded laterally and is the fourth component of the orbit. The basal antennal article is usually not free (as might be expected of a limb) but is part of the lower carapace surface, an area between the two antennal slots (as) housing the pair of antennules and the eyestalks on the side. A pore (antennal gland) (ag) sits near the proximal, most posterior, end of the basal antennal article. The rest of the antenna attaches to the distal, most anterior, margin of the basal article. The shape and armature of the basal antennal article feature in keys. Five of the subfamilies occur in Australia naturally and another only because of introduction of an exotic species. Species are listed here alphabetically within subfamilies and genera. The boxes list the most common shallow-water decorator crabs and spider crabs, and species found only mainly at >80 metres depth, that is on the shelf edge, slope or seamounts.

Diagnosis. Carapace widest posteriorly, with well-developed branchial regions; typically pear- shaped or triangular, rarely more circular; dorsal surface convex, smooth to highly ornamented, hooked setae often present on carapace and legs. Front narrow, produced as rostrum: typically 2 long, horn-like projections, fused at base, but sometimes a short, strongly deflexed flat plate. Orbits complete, poorly developed or absent. Anterolateral margins of carapace often armed with well- developed spines. Basal article of antenna well developed, usually fused with epistome, and often also with front. Epistome large, buccal cavity quadrate. Legs often spiny and/or granular, often with stiff setae. Chelipeds highly mobile, usually shorter than walking legs. Male and female abdomen of 7 free somites, rarely some fused, sometimes somite 6 fused to telson, to form pleotelson. Female genital openings sternal; male openings coxal. Gonopod 1 greatly exceeding gonopod 2. (after Davie, 2002)

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SPIDER CRABS AND DECORATOR CRABS COMMON IN SHALLOW-WATER Huenia australis Microhalimus deflexifrons Paramithrax barbicornis Achaeus curvirostris Naxia aries Prismatopus spatulifer Dumea latipes Naxia aurita Pseudomicippe maccullochi Pyromaia tuberculata Naxia spinosa Schizophrys rufescens Anacinetops stimpsoni Naxia tumida Tumulosternum wardi Leptomithrax gaimardii Notomithrax minor Micippa platipes Leptomithrax sternocostulatus Notomithrax ursus Micippa spinosa

SPIDER CRABS AND DECORATOR CRABS FOUND MAINLY AT DEPTHS > 80 M OR ON SEAMOUNTS Griffinia lappacea Platymaia remifera Leptomithrax waitei Pugettia tasmanensis Platymaia wyvillethomsoni Prismatopus spatulifer Achaeus pugnax Vitjazmaia latidactyla Teratomaia richardsoni Cyrtomaia maccullochi Kimbla franklini Naxioides robillardi Cyrtomaia murrayi Leptomithrax depressus Phalangipus australiensis Cyrtomaia suhmi Leptomithrax gaimardii Phalangipus hystrix Dorhynchus ramusculus Leptomithrax globifer Rochinia fultoni Macropodia trigonus Leptomithrax sternocostulatus Rochinia mosaica Platymaia fimbriata Leptomithrax tuberculatus

Key to subfamilies of Majidae in southern Australia 1. Eyes without orbits or with a supraorbital eave and postorbital spine, eyestalks clearly visible in dorsal view; basal antennal article slender (usually more than twice as long as wide) (Fig. 103a–f) ...... 2 — Eyes with orbits variously well protected by an eave and spines anterior and posterior to the eyestalk, eyestalk hidden under these spines; basal antennal article broad (at most twice as long as wide) (Fig. 103g–j) ...... 5 2. Eyes without orbits; eyestalks generally long, either non-retractile or retractile against an acute postorbital spine affording no concealment; basal antennal article usually extremely slender and sometimes free distally (Fig. 103c, d); merus of maxilliped 3 often not as wide as ischium ...... 3 — Eyes with at most a supraorbital eave and postorbital lobe; eyestalks short and stout, not completely concealed in dorsal view; basal antennal article not extremely slender, always fused distally (Fig. 103a, b, e, f); merus of maxilliped 3 as wide as ischium ...... 4 3. Male abdomen with somite 7 subtriangular and not inserted into somite 6 (rostrum bifid) ...... Inachinae ...p.355 — Male abdomen with somites 6 and 7 fused; (rostrum triangular in only Australian species) ...... Inachoidinae Pyromaia tuberculata ...p.365 4. Eyes without true orbits, supraorbital eave weakly developed, sometimes with small preorbital spine and/or postorbital spine (or eyes sunk in sides of huge beak-like rostrum); intercalated spine absent (Fig. 103a, b) ...... Epialtinae ...p.251

349 Marine Decapod Crustacea of Southern Australia

rostrum rostrum

pre pre baa se se as

ag ab

rostral spine rostral spine

baa as

ag cd

rostral spine rostral spine baa as se se post post ag ef

rostral spine rostral spine baa baa se ant ant post post int int as g h ag

rostral spine rostral spine pre pre baa se se ant ant post post as i j ag Fig. 103. Majidae. Subfamily differentiation. Dorsal and ventral views of anterior regions: a, b, Epialtinae (Huenia halei). c, d, Inachinae (Achaeus podocheloides). e, f, Pisinae (Hyastenus elatus). g, h, Majinae (Notomithrax minor). i, j, Mithracinae (Tiarinia angusta). See p. 348 for abbreviations.

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— Eyes with partial orbits comprising a supraorbital eave sometimes produced into a preorbital spine, plus a prominent postorbital lobe, often cupped to house retracted eyestalk; intercalated spine sometimes present (Fig. 103e, f) ...... Pisinae ...p.384 5. Basal antennal article not expanded laterally to form a floor to the orbit; orbit dorsally comprises a supraorbital eave and postorbital lobe usually separated by a hiatus such that upper eave extends more than lower margin; intercalated spine usually present (Fig. 103g, h) ...... Majinae ...p.366 — Basal antennal article expanded laterally to form a floor to the orbit; orbit dorsally comprises supraorbital eave (often with antorbital spine) and adjacent hollowed postorbital lobe such that eye is in deep circular socket; intercalated spine sometimes present (Fig. 103i, j) ...... Mithracinae ...p.381

Subfamily Epialtinae Macleay, 1838 The orbits of members of this subfamily are undeveloped, the often immoveable eyestalks sunken into the sides of the carapace. Of the 18 species recorded from Australia, only seven occur along southern coasts. Griffin & Tranter (1986) provided a key to most genera now known. Most shallow-water species do not decorate but use leaf-like body form as camouflage in algal habitats.

Diagnosis. Eyes without true orbits; intercalated spine not present. Eyestalks short, occasionally obsolescent, scarcely movable; either concealed beneath a forwardly produced supra-ocular spine, or sunk in the sides of a beak-like rostrum; postocular lobe sometimes present but not excavated to house retracted eye. Basal antennal article truncate, triangular, fused distally. Merus of maxilliped 3 as wide as ischium. Dactyli of walking legs prehensile or subchelate; last 3 pairs often disproportionately short compared with first pair. Male abdomen not terminally broadened, somite 7 subtriangular and not inserted deeply into somite 6. Male gonopod 1 usually slender, weakly curved; aperture usually terminal, apex simple, curved or expanded into lobes, rarely with a slender process. (after Griffin & Tranter, 1986)

Key to southern Australian genera and species of Epialtinae 1. Rostrum single or secondarily bifurcate ...... 2 — Rostrum of 2 spines or lobes; if spines then generally divergent ...... 5 2. Branchial margin with 2 small lobes or tubercles; a small lateral lobe just behind eye; propodus of’ first walking leg smooth on lower margin, without lobe or tuft of setae; sexes similar ...... Menaethius monoceros — Branchial margin with 1 lobe, often very large; no lateral lobe behind eye; propodus of first walking leg with ventral tuft of setae and sometimes lobe in distal half; sexes dissimilar ...... Huenia ...3 3. Rostrum distally flattened dorsoventrally, bifurcate ...... Huenia bifurcata — Rostrum cylindrical, at most weakly notched ...... 4 4. Walking legs cylindrical; rostrum narrowed distinctly in distal quarter . . . . Huenia australis — Walking legs with at least carpus prominently carinate on upper margin; rostrum evenly tapering distally ...... Huenia halei 5. Orbit a continuous rim enclosing eye, without hiatus; eye immoveable; meri of first 2 pairs of walking legs with 2 spines on superior margin and tubercles on outer face . . . Perinia tumida

351 Marine Decapod Crustacea of Southern Australia

— Orbit not comprising a continuous rim, distinct hiatus above and below, or eye not retractile against postorbital lobe; meri of walking legs smooth or with only a terminal tubercle or lobe ...... 6 6. Rostral spines one-fifth postrostral carapace length; hepatic tooth weak ...... Pugettia tasmanensis — Rostral spines two-thirds postrostral carapace length; hepatic spine laterally flattened and curved anteriorly and upwards ...... Griffinia lappacea

Griffinia Richer de Forges, 1994 The genus of rare small deep-water crabs, not more than 20 mm long, comprises only three Indo- West Pacific species (Richer de Forges, 1994). Diagnosis. Carapace smooth, with few setae; rostrum of 2 diverging spines. Orbit open, defined by strong, acute preocular spine joining narrow supraocular eave ending in blunt postocular spine. Basal antennal article longer than broad. Lateral border with granules; 1 hepatic spine. Cheliped carinate in male. Abdomen with 7 somites in male, 6 in female. Griffinia lappacea (Rathbun, 1918) burr crab (Fig. 105a). Rostral spines two-thirds length of carapace; hepatic spine prominent, flattened, curved anteriorly and upwards; cheliped of male with high leaf-like crest on carpus. 20 mm. Philippines, Indonesia, WA, SA (Great Australian Bight); 366–548 m depth. This is a rare species, reported as Antilibinia lappacea but reclassi- fied and redescribed by Richer de Forges (1994).

Huenia De Haan, 1837 Nine species belong to this Indo-West Pacific genus (Griffin & Tranter, 1986) of which three occur in southern Australia. Members of this genus are not immediately recognisable as majids, with their characteristically irregular leaf-like flattened carapace mimicking the leaves of the brown algae in which they live. Body shape is variable within species and highly sexually dimorphic. Diagnosis. Rostrum single or secondarily bifurcate. Carapace dorsally flat; preorbital angle dorsoventrally flattened, produced as a triangular lobe; hepatic lobe (more anterior of 2) rounded or triangular; branchial margin with 1 lobe, often very large (triangular in southern Australian species); no lobe behind eye. Sexes dissimilar. Huenia australis Griffin & Tranter, 1986 (Fig. 104a, Pl. 20a). Carapace with hepatic lobe small, triangular, and branchial lobe larger, triangular; rostrum narrowed distinctly in distal half. Walking legs cylindrical. 26 mm. Vic. (Western Port), Tas., SA, WA (Perth); 6–30 m depth. The carapace shape is variable; specimens can be bright-leaf-green or red with a white patch, depending presumably on the algae on which they are found. Huenia bifurcata Streets, 1870 (Fig. 105b, c). Carapace with branchial lobe broad, weakly bilobed, and hepatic lobe truncate or blunt in female, absent in male; rostrum distally flattened dorsoventrally, bifurcate. Walking legs compressed and carinate. 29 mm. Qld, NSW (Bega); 4–15 m depth. Huenia halei Griffin & Tranter, 1986 (Fig. 103a, b). Carapace variable, with hepatic lobe broad and rounded, and branchial lobe larger and weakly bilobed; 2 low tubercles dorsally. Rostrum evenly tapering distally. Walking legs flattened, carinate. 21 mm. Vic. (Western Port), SA, WA (Perth); 5–33 m depth. The record of H. proteus from South Australia (Hale, 1927) has now been referred to H. halei.

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Fig. 104. Majidae Epialtinae. a, Huenia australis. b, Perinia tumida.

Menaethius Milne Edwards, 1834 The genus comprises only two species, both widespread in the Indo-West Pacific (Griffin & Tranter, 1986). Diagnosis. Carapace smooth; rostrum slender; preorbital angle dorsoventrally flattened, produced as a triangular lobe; hepatic margin with 1 and branchial margin with 2 small lateral lobes; small lateral lobe just behind eye. Walking leg 1smooth on lower margin. Sexes similar. Menaethius monoceros (Latreille, 1825) (Fig. 105d). Rostrum very slender throughout its length. Dorsal branchial region with rounded elevations and sometimes a tubercle. Walking legs not carinate. 27 mm. Japan to Red Sea, Qld, NSW, Vic., WA (S to Rottnest I.), Lord Howe I.; coral and rocky reef, low intertidal and subtidal. The species which has numerous synonyms is exceptionally rare in this region.

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d e

Fig. 105. Majidae Epialtinae. a, Griffinia lappacea. b, c, Huenia bifurcata (female, male). d, Menaethius monoceros. e, Pugettia tasmanensis.

Perinia Dana, 1852 This is a monotypic genus at one time placed in Pisinae but is similar to Menaethius. Diagnosis. Carapace with low tubercles; rostrum short, bifid; margin of orbit continuous, without sutures separating postocular lobe, enclosing immoveable eyestalk. Walking legs with spines on upper margins. Sexes similar. Perinia tumida Dana, 1852 (Fig. 104b). Rostrum one-tenth carapace length, comprising 2 broad flattened lobes separated by U-shaped gap; each lobe subacute and slightly incurved. 10 mm. Indo-West Pacific from Japan to Red Sea, Qld, NSW, SA, WA; low intertidal and subtidal reef.

Pugettia Dana, 1851 Pugettia is a Pacific genus of a dozen species for which a key was provided by Griffin & Tranter (1986) and at least this one has been added since. In Australia it is known now only from one offshore species. Diagnosis. Carapace with few tubercles or spines; rostral spines slender, at least one-quarter (usually not more than half) carapace length; orbit with distinct hiatus above and below eyestalk; postorbital lobe distinct from eave; hepatic lobe directed laterally.

354 Brachyura – crabs

Pugettia tasmanensis Richer de Forges, 1993 (Fig. 105e). Rostral spines one-fifth postrostral length. Carapace hepatic region with a broad flattened blunt tooth, branchial region with 2 tubercles and longer tooth, with middorsal hump posteriorly. 5 mm. Off NSW, Taupo Seamount; 132 m depth.

Subfamily Inachinae Macleay, 1838 The absence of an orbit and the elongate exposed eyestalks are the defining features of members of this subfamily. While some shallow water representatives are found in some genera (Achaeus, Dumea, Litosus) others like Cyrtomaia, Macropodia, Platymaia and Vitjazmaia are confined to the deep sea. Diagnosis. Eyes without orbits; eyestalks generally long, either non-retractile, or retractile against sides of carapace, or against an acute postocular spine affording no concealment. Basal antennal article usually long and extremely slender; sometimes free distally. Merus of maxilliped 3 often not as wide as ischium. Male abdomen not terminally broadened, somite 7 subtriangular and not inserted deeply into somite 6. Pleopod 1 not usually very stout, usually straight or weakly curved, rarely twisted, aperture terminal or subterminal, sometimes a slit, often protected by a flap, apex simple or with a lobe. (after Davie, 2002)

Key to southern Australian genera of Inachinae 1. Distal margin of basal antennal article extending forward no further than half diameter of eyestalk, free; rostrum of 2 blunt spines; carapace without spines ...... Oncinopus — Distal margin of basal antennal article at least level with and usually extending beyond anterior margin of eyestalk, free or fused to front; rostrum variable; carapace with or without spines ...... 2 2. Antennal gland at proximal end of basal antennal article, in contact with anterior margin of epistome ...... 3 — Antennal gland removed from proximal end of basal antennal article, distant from posterior margin of epistome (Fig. 108j) ...... 7 3. Rostrum of 2 short blunt lobes, sometimes with spinules; interantennal partition lacking spine; carpus of cheliped with 1 large spine on inner margin; shallow subtidal ...... 4 — Rostrum of 2 slender spines, sometimes with accessory spines; interantennal partition with ventral spine, sometimes bifid; carpus of cheliped with many spines or tubercles; deep-water ...... 5 4. Anterolateral margin with 3 large spines and 2 smaller spines between; subhepatic region spinose; cheliped with acute granules on outer surface ...... Dumea — Anterolateral margin with 1 broadly triangular tubercle and anteriorly directed spines; subhepatic region smooth; cheliped smooth on outer surface ...... Litosus 5. Basal antennal article free distally (Fig. 109i); interantennular spine horizontal, long; propodi of walking legs 2–4 flattened ...... Platymaia — Distal margin of basal antennal article fused to front; interantennular spine deflexed, short; propodi of walking legs cylindrical ...... 6 6. Rostrum bifid, with antennal spine visible below ...... Cyrtomaia — Rostrum long, with pair of shorter pseudorostral spines ...... Vitjazmaia

355 Marine Decapod Crustacea of Southern Australia

Fig. 106. Majidae Inachinae. a, Achaeus curvirostris. b, Dumea latipes.

356 Brachyura – crabs

7. Maxilliped 3 with merus as broad as ischium; basal antennal article flattened distally; rostral spines separate from base ...... Ephippias — Maxilliped 3 with merus narrower than ischium; basal antennal article cylindrical distally; rostral spines separate, contiguous or obsolete ...... 8 8 Upper orbit with postorbital spine or lobe against which eyestalk retracts . . . . . Dorhynchus — Upper orbit without postorbital spine or lobe; eyestalk immoveable or retracting against hepatic margin ...... 9 9. Rostrum of 2 contiguous spines produced well in front of antennal fossae; basal antennal article fused with epistome, free anteriorly; deep-sea ...... Macropodia — Rostrum of 2 short lobes, apices rounded or blunt, or rostrum obsolete; basal antennal article fused with epistome anteriorly; mostly shelf species ...... Achaeus

Achaeus Leach, 1817 Several species of these small crabs with long, thin legs are recognisable by having a narrowed carapace posterior to the eyes, giving the appearance of a neck. They differ from species of Dorhynchus in lacking a prominent postorbital spine. The genus Achaeus includes almost 30 species worldwide, most of which inhabit continental shelf waters; five species are known to occur in southern Australia. More species occur in northern waters, and these can be identified using Griffin & Tranter’s (1986) key to Indo-West Pacific species. More detailed descriptions of the Australian species can be found in Griffin (1970). Diagnosis. Carapace pear-shaped to elongate subtriangular, narrowed behind orbit almost as a neck, sometimes with few tubercles but never long spines. Rostrum of 2 short, acute or rounded spines. Orbit a narrow supraorbital eave, laterally spinulated or smooth; postorbital spine if present not prominent. Eyestalks long, non-retractile. Basal antennal article extremely slender, smooth, weakly tuberculate, spinulous or armed with spines laterally. Merus of maxilliped 3 subovate, not notched distally, palp articulating terminally. Chelipeds greatly enlarged in adult male, usually spinous. Walking legs long and slender, dactyli of legs 3 and 4 usually falcate and spinulated on lower margin. Abdomen with 6 somites in both sexes, male abdomen widest at middle of third somite. Male pleopod 1 bluntly pointed, aperture located subterminally in a groove. (after Griffin & Yaldwyn, 1965)

Key to southern Australian species of Achaeus 1. Carapace smooth; rostrum deeply divided by V-shaped notch, apically rounded, with spinulose apex ...... Achaeus lacertosus — Carapace with 2 or more prominently tubercles or spines in midline and several laterally; rostral lobes acute or blunt, spinulose laterally and/or medially but not apically ...... 2 2. Supraorbital eave with 1–3 spines; walking leg 4 dactyl weakly curved . . . Achaeus curvirostris — Supraorbital eave smooth or spinulose; walking leg 4 dactyl falcate to semicircular ...... 3 3. Carapace with long spines on mesogastric and cardiac regions; rostral lobes narrow, contiguous, separated by narrow slit ...... Achaeus pugnax — Carapace with tubercles on mesogastric and cardiac regions; rostral lobes broad at base, separated by broad notch ...... 4 4. Rostral lobes apically blunt, smooth, separated by shallow notch; eyestalk with prominent tubercle on anterior margin ...... Achaeus brevirostris — Rostral lobes subacute, separated by deep notch; eyestalk with 2 spinules on anterior margin ...... Achaeus podocheloides

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Achaeus brevirostris (Haswell, 1879) (Fig. 108a). Carapace with prominent neck in male, constricted behind orbit, shorter and unconstricted in female, with curved setae along margins. Rostral lobes apically blunt, smooth, separated by shallow notch; supraorbital eave smooth; eyestalk with prominent tubercle on anterior margin. 12 mm. NT, Qld, NSW (S to Port Jackson), WA (S to Perth), Indonesia–Thailand, Indian Ocean; 6–60 m depth. Achaeus curvirostris (Milne Edwards, 1873) (Fig. 106a). Carapace with prominent neck in male, constricted behind orbit, shorter and unconstricted in female, with curved setae along margins. Rostral lobes apically acute, smooth, separated by shallow notch; supraorbital eave with a prominent spine, shorter spinules and postorbital spine; eyestalk with prominent tubercle on anterior margin. 11 mm. Qld, NSW, Tas. (Bass Strait), WA (S to Cape Naturaliste), New Zealand, Indo-West Pacific; 10–150 m depth. The species was described as Achaeus fissifrons (Haswell, 1879) by Griffin & Yaldwyn (1965). Achaeus lacertosus Stimpson, 1858 (Fig. 108b). Carapace smooth, with short unconstricted region behind orbit, with curved setae dorsally. Rostral lobes short, rounded, spinulose with curled marginal setae, separated by broad V-shaped notch; supraorbital eave smooth; eyestalk without tubercle on anterior margin. 12 mm. NT, Qld, NSW (S to Port Jackson), WA, Indo-West Pacific, Japan-South Africa; 6–90 m depth. Achaeus podocheloides Griffin, 1970 (Fig. 108c). Carapace with triangular neck, constricted behind orbit, with curved setae on lateral regions. Rostral lobes triangular, separated by broad V-shaped notch; supraorbital eave smooth; eyestalk with 2 spinules on anterior margin. 8 mm. WA (S to Cape Naturaliste); to 140 m depth. Achaeus pugnax (de Man, 1928) (Fig. 108d). Carapace broadening evenly behind orbit, with curved setae along margins. Rostral lobes short, close, separated by narrow slit; supraorbital eave spinulose; eyestalk with small tubercle on anterior margin. 12 mm. WA (S to Perth), Japan, Indonesia; 80–540 m depth.

Cyrtomaia Miers, 1886 Species of Cyrtomaia are long-legged, subspherical-bodied deep-water crabs that can best be recognised by the pale colour and spinose or tubercular carapace. They do not decorate but often have epizoic stalked barnacles on the carapace. The cylindrical (not flattened) propodi and dactyli of the last three walking legs and prominent slender submarginal spine on the anterior part of the protogastric region distinguish them from similar genera like Platymaia. Griffin & Tranter (1986) recognised 17 species of which four occur in Australia, three in our region but with scarcely overlapping distributions. The species that occur in the tropical Indo-West Pacific can be identified using the key in Griffin & Tranter (1986). Guinot & Richer de Forges (1982; 1985) dealt with more species not recognised by Griffin & Tranter. Diagnosis. Carapace generally subcircular, broader than long, convex, usually bearing several long spines. Rostrum of 2 short, close spines. Interantennular spine sometimes projecting horizontally forward below rostrum so that rostrum appears trispinose. Orbit consisting of narrow supraorbital eave and well developed, simple postorbital spine; eyestalks long, slender, cornea terminal, ovoid. Basal antennal article cylindrical, slender, not firmly fixed to side of rostrum. Chelipeds spinous, elongate in male. Walking legs long, slender, cylindrical, spinous. Abdomen of 7 free somites.

Key to southern Australian species of Cyrtomaia 1. Supraorbital eave with anterior spine ...... Cyrtomaia murrayi — Supraorbital eave unarmed ...... 2

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Fig. 107. Majidae Inachinae. a, Ephippias endeavouri. b, Oncinopus aranea (from De Haan, 1839: pl. 29 fig. 2). c, Platymaia aff. wyvillethomsoni.

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2. Rostral spines subparallel, straight; eyestalk occupying more than half orbit; tubercle midway between protogastric and anterior branchial spine ...... Cyrtomaia maccullochi — Rostral spines divergent, curved; eyestalk occupying less than half orbit; spine midway between protogastric and anterior branchial spine ...... Cyrtomaia suhmi Cyrtomaia maccullochi Rathbun, 1918 slender-handed spider crab (Fig. 108e). Carapace densely and finely granular; posterior mesogastric spine without flanking spine. Rostral spines subparallel and horizontal. Supraorbital eave unarmed. 54 mm. South China Sea, WA, SA; 347–820 m depth. This is the only species on the southern Australian coast. Cyrtomaia murrayi Miers, 1886 (Fig. 108f). Carapace spinose, especially laterally and with 2 on posterior margin, otherwise almost smooth. Rostral spines short and triangular. Supraorbital eave with anterior spine. 31 mm. Indo-West Pacific, WA (S to Rottnest I.); 120–204 m depth. Cyrtomaia suhmi Miers, 1886 (Fig. 108g). Carapace smooth; posterior mesogastric spine without flanking spine. Rostral spines divergent and arched. Supraorbital eave unarmed. 90 mm. Indo- West Pacific; Qld, WA, NSW (S to Nowra); 460–1125 m depth.

Dorhynchus Thomson, 1873 The species assigned to this genus have been commonly assigned to Achaeopsis Stimpson, 1857, a junior synonym (Davie, 2002). Its four species are small crabs with long, thin and frail legs and slender postorbital spines, inhabiting continental shelf and slope waters to a depth of 1000 m. Four species are known worldwide and can be identified using the keys to Achaeopsis (Griffin, 1966b; Griffin & Tranter, 1986). Only one species occurs in southern Australia, recognisable from the thin rostral spines separated by a long narrow slit and two strong dorsal spines in the midline. Diagnosis. Carapace ovate-triangular, convex, spinous; interorbital carapace narrow; orbit with supraorbital and postorbital spines. Rostrum bifid. Eyes retractile to sides of carapace. Basal antennal article long and narrow, in an almost vertical plane; movable portion much longer than rostrum and inserted beside it. Maxilliped 3 spinous; merus narrower than ischium, constricted at base, rounded at extremity, palp articulating distally. Abdomen of 6 somites. Chelipeds strong, spinous; merus trigonal; palm dilated; fingers broad. Walking legs long and slender. Dorhynchus ramusculus (Baker, 1906) slender spider crab (Fig. 108j) Carapace with protogastric region and anteromedial angle of branchial region smooth. Rostral spines extending beyond end of peduncle of antenna. Walking legs with spine on upper margin of meri. 15 mm. New Zealand, NSW (off Wollongong), Vic., Tas., SA, WA (south coast); to 738 m depth. A detailed description of this species (as Achaeopsis ramusculus) can be found in Griffin (1966b).

Dumea Loh & Ng, 1999 Dumea was differentiated from the larger genus Paratymolus Miers, 1879 by Loh & Ng (1999). Both genera include small and compact majids, not immediately recognisable as members of this family. It is the broad rostrum and shallow lateral (rather than anterior) orbits that place the species in Majidae. Diagnosis. Carapace and posterolateral margin without tubercles; anterolateral margin with 3 large spines and 2 smaller spines between; subhepatic region spinose; rostral lobes weak. Eyestalks short. Basal antennal article not fused anteriorly to carapace. Cheliped with acute granules on outer surface; carpal spine short, stout. Abdomen triangular in both males and females, abdominal somites 3–5 fused and laterally concave in male. Dumea latipes (Haswell, 1880) velvet crab (Fig. 106b, Pl. 20b). No spinule at base rostral notch, rostral spinules equal. Orbital region not toothed. 10 mm. Qld, NSW, Vic., SA, WA; reef flats to 4 m depth.

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b c a

h g

Fig. 108. Majidae Inachinae. a, Achaeus brevirostris. b, Achaeus lacertosus. c, Achaeus podocheloides. d, Achaeus pugnax. e, Cyrtomaia maccullochi. f, Cyrtomaia murrayi. g, Cyrtomaia suhmi. h, Dorhynchus ramusculus. i, Dorhynchus ramusculus (left profile, carapace). j, Dorhynchus ramusculus (anterior region, ventral).

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Ephippias Rathbun, 1918 This monotypic genus is restricted to Australian waters. Diagnosis. Supraorbital eave not expanded; preorbital and antorbital spines absent; postorbital spine small and remote from eye. Basal antennal article slender with 1 anterior spine, antenna excluded from orbit. Pleopod 1 of male curves outward distally and aperture a subterminal split. Abdomen of 7 somites. Ephippias endeavouri Rathbun, 1918 Endeavour crab (Fig. 107a). 115 mm. S from northern NSW, SA, to northern WA (not yet known from Vic. or Tas.); to 50 m depth. This large crab from shelf depth is distinguished by the first walking legs being much longer than the chelipeds and other legs, and twice as long as the carapace. It is usually covered with sponges.

Litosus Loh & Ng, 1999 The genus of two species was differentiated from Paratymolus and Dumea by Loh & Ng (1999). Diagnosis. Carapace and posterolateral margin without tubercles; anterolateral margin with 1 broadly triangular tubercle and anteriorly directed spine; subhepatic region smooth; rostral lobes weak. Eyestalks short. Basal antennal article not fused anteriorly to carapace. Cheliped smooth on outer surface; carpal spine short, stout. Abdomen triangular in both males and females, abdominal somites 3–5 fused and laterally concave in male. Litosus sexspinosus (Miers, 1884) (Fig. 109a). Chela of male not bulbous; cutting edges of fingers evenly toothed. 8 mm. Indo-West Pacific, Qld, NSW, WA; reef, subtidal. The species may be confused with Dumea latipes but has much smoother anterolateral margins.

Macropodia Leach, 1814 The genus is well represented in the eastern Atlantic and Mediterranean (Forest, 1978) and less so in the Indian Ocean (Griffin & Tranter, 1986). It has not been recently diagnosed but the long contiguous rostral spines, appearing as if a single structure, distinguish the genus from Achaeus. The only Australian species is from deep water like the others. The small size and extremely spindly legs separate it from other majids. Diagnosis. Carapace pear-shaped to elongate subtriangular, narrowed behind orbit almost as a neck, sometimes with few tubercles but never long spines. Rostrum of 2 contiguous spines produced well in front of antennal fossae. Orbit a narrow supraorbital eave. Eyestalks long, non-retractile. Macropodia trigonus Richer de Forges, 1993 (Fig. 109b). Carapace triangular. Legs extremely thin, length 5 times carapace width. 7.5 mm. NSW (Taupo Seamount); 244 m depth.

Oncinopus De Haan, 1839 Members of Oncinopus are distinguished by the short rounded rostral lobes, together forming a broad projection divided by a parallel-sided slit and extending backwards over the eyestalks. Distal articles of the first two walking legs are dilated and the last two are subchelate. The four species can be identified using the key in Griffin & Tranter (1986). Their key relies on the reliable structure of the male pleopods and female gonopore (see diagnoses) and carapace shapes are subtly different. Diagnosis. Basal antennal article slender, distal margin extending forward no further than half diameter of eyestalk, free. Interantennal spine absent. Rostrum of 2 blunt spines; carapace without spines. Male gonopod 1 twisted 3 times.

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b a c

e g

i j

Fig. 109. Majidae Inachinae. a, Litosus sexspinosus. b, Macropodia trigonus. c, d, Platymaia fimbriata (dorsal and lateral rostral and interantennal spines). e, f, Platymaia maoria (dorsal and lateral rostral and interantennal spines). g, h, i, Platymaia aff. wyvillethomsoni (dorsal and lateral rostral and interantennal spines, ventral orbit and antennae). j, Vitjazmaia latidactyla.

Key to southern Australian species of Oncinopus 1. Female gonopore distinctly elevated and opening mesially; anterior half of carapace clearly tapering (hepatic width half branchial width) ...... Oncinopus aranea

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— Female gonopore on surface; anterior half of carapace moderately tapering (hepatic width two-thirds branchial width) ...... Oncinopus neptunus Oncinopus aranea De Haan, 1839 thin-shelled spider crab (Fig. 107b). Male pleopod 1 with beak- like lateral subapical lobe. Female gonopore distinctly elevated and opening mesially. 11 mm. West Pacific, Qld, SA, WA (S to Bunbury); coral and shallow rocky reef and slope depths. Oncinopus neptunus Adams & White, 1848. Male pleopod 1 slightly widened subapically. Female gonopore on surface. 13 mm. Indo-West Pacific, Qld, NSW (S to Port Jackson); subtidal coral and rocky reef to 90 m depth.

Platymaia Miers, 1886 Like Cyrtomaia, this is a long-legged deep-water genus comprising seven or more Indo-West Pacific species. It differs from Cyrtomaia principally in having flattened rather than cylindrical walking legs. Identity of southern Australian species remains in doubt. Diagnosis. Carapace subcircular, convex, sloping steeply posteriorly, poorly calcified, spinose and setose; rostrum with pair of pseudorostral spines; supraocular spine present or absent; 1 postorbital spine. Eyes small. Basal antennal article spinose, moveable. Abdomen with 7 somites. Walking leg 1 long, especially spinose; walking legs 2–4 longer, with carpus-dactylus flat, with setose margin.

Key to southern Australian species of Platymaia 1. Carapace surface with larger spines plus numerous smaller spines . . . . Platymaia fimbriata — Carapace surface smooth except for granules and few short spines ...... 2 2. Interantennal and rostral spines dagger-like (in lateral view); with row of 3 short supraorbital spines ...... Platymaia maoria — Interantennal and rostral spines short, triangular (in lateral view); without supraorbital spines ...... Platymaia aff. wyvillethomsoni Platymaia fimbriata Rathbun, 1916 (Fig. 109c, d). Carapace surface with larger spines plus numerous smaller spines; postorbital spine equal to rostral spines; interantennal spine about 3 times length of rostral spines. 51 mm. West Pacific, WA, Qld, NSW; continental slope, 400–620 m depth. Davie & Short (1989) recognised three sympatric forms off the coast of Queensland differing in the degree of spination. Platymaia maoria Dell, 1963 (Fig. 109e, f). Carapace surface granulate; interantennal and rostral spines dagger-like; with row of 3 short supraorbital spines. 60 mm. New Zealand, Qld, NSW; continental slope, 405–600 m depth. Platymaia aff. wyvillethomsoni Miers, 1886 three-spined spider crab (Figs 107c, 109g–i). Carapace surface granulate; interantennal and rostral spines short, triangular; without supraorbital spines. 52 mm. West Pacific, Qld, NSW, SA; continental slope, 200–820 m depth. Guinot & Richer de Forges (1985) believed that the species on the east coast of Australia is Platymaia remifera Rathbun, 1916 and those from SA an undescribed species similar to P. wyvillethomsoni. These were assigned to P. wyvillethomsoni by Griffin & Tranter (1986); they included P. remifera and P. alcocki as junior synonyms. The illustrated specimen is from Queensland.

Vitjazmaia Zarenkov, 1994 The only species of Vitjazmaia is one of a group of long-legged deep-water spider crabs (Webber & Richer de Forges, 1995). Diagnosis. Carapace subcircular, convex, sloping steeply posteriorly, poorly calcified, spinose and setose; rostrum long with pair of pseudorostral spines; 1 strong postorbital spine; 1 supraorbital

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spine at base of rostrum. Eyes small. Basal antennal article short, broad, spinose, moveable; 2 distal spines. Abdomen with 7 somites. Cheliped propodus long, thickest distally. Walking legs long, depressed, oval in cross-section, with small recurved spinules. Vitjazmaia latidactyla Zarenkov, 1994 (Fig. 109j). Western Indian Ocean, New Zealand, Lord Howe Rise, NSW, Tas.; continental slope, 497–1290 m depth.

Subfamily Inachoidinae Dana, 1851 A group of ten genera of majid-like crabs comprise the Inachoidinae. Drach & Guinot (1983) advocated family status, a view shared by Lemaitre et al. (2001), because of the unique relationship between the carapace and the rest of the skeleton. Guinot & Richer de Forges (1997) went so far as to suggest that its affinities lay not within Majidae but with Hymenosomatidae. This was not the view of Pohle & Marques (2000) whose work with larval stages lead them to doubt the monophyly of the subfamily and place its genera back in Inachinae. Such is the uncertainty about phylogenetic relationships within the majoid families. Diagnosis. Eyes without orbits, unconcealed; eyestalks generally long, non-retractile, or retractile against carapace sides or against acute postocular spine. Male abdomen with telson fused to somite 6 to form pleotelson.

Fig. 110. Majidae Inachoidinae. Pyromaia tuberculata.

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Pyromaia Stimpson, 1871 The genus is essentially an American and northern Pacific genus (Lemaitre et al., 2001) with five species. Diagnosis. Carapace pyriform, with tubercles and spines; rostrum simple. Latero-external parts of pleurites 5–8 extend beyond the side of the carapace. Postorbital spine curving around eye. Walking legs elongate, with curved dactyli. Cheliped of male short and inflated. Pyromaia tuberculata (Lockington, 1877) (Fig. 110, Pl. 20c). Basal antennal article without strong spine on ventrolateral margin. 30 mm. Eastern Pacific (California to Columbia) but introduced to Japan before 1976 (Sakai, 1976; Furota, 1988, 1990, 1996; Furota & Furose, 1988), New Zealand by 1978 (Webber & Wear, 1981), Cockburn Sound, WA by 1978 (Morgan, 1990), Port Phillip Bay and Gippsland coast, Vic., by 1990 (Poore & Storey, 1999), Brazil by 1989 and Argentina (Schejter et al., 2002); subtidal muddy sediments to 412 m depth. Pyromaia tuberculata is immediately recognisable in the shallow environments in which it occurs by being the only southern Australian majid with single rostrum – all other common shallow-water species have a pair of rostral spines. This is one of four species of introduced crabs in southern Australia, the others being Metacarcinus novaezelandiae (Cancridae), Halicarcinus innominatus (Hymenosomatidae), and Carcinus maenas (Portunidae), but the only one to have invaded soft sediments. It has become locally abundant in Port Phillip Bay at depths below 12 m since its first appearance there in about 1990. It is certain now to be an important food source for demersal fishes (Parry et al., 1995). The larval stages are known (Fransozo & Negreiro-Fransozo, 1970).

Subfamily Majinae Samouelle, 1819 The Majinae are the most diverse and the largest subfamily of decorator crabs in southern Australia and in fact in all the Indo-West Pacific where they dominate the majid fauna. They are represented in this region by more than 20 genera and more than 70 species. Griffin & Tranter (1986) provided keys to the species of the Indo-West Pacific and their study is the basis of this treatment and the catalogue of Australian species in Davie (2002). Majine decorator crabs have well developed orbits surrounding the deeply set eyestalks differing from the mithracines in possessing a spine between the eave and postorbital spine, and the eye not protected below by an expanded basal antennal article. The most commonly encountered shallow-water decorator crabs belong to the majine genera Notomithrax and Naxia and the southern ‘great spider crab’ to the genus Leptomithrax. Identification to genus and species often involves cleaning algae and epifauna away from the dorsal surface of the carapace, rostral spines and orbit to see details of ornamentation. Diagnosis. Eyes with nearly complete, or complete, orbits formed by a supraorbital eave (often with an antorbital spine on posterior corner), a postorbital lobe, and usually an intercalated spine between. Basal antennal article broad but not expanded to form a floor to the orbit; often with a strong spine at 2 anterior angles. Male gonopod 1 slender, weakly curved, its aperture usually subterminal and usually protected by lobes, its apex slender and simple. (after Griffin & Tranter, 1986)

Key to southern Australian genera of Majinae 1. Eyestalks very long, flattened, with weak lobe on anterior and posterior margins; supraorbital eave weakly expanded, barely covering but curved over bulbous base of eyestalk; carapace with numerous compound tubercles ...... Pippacirama ...p.378

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— Eyestalks not flattened or lobed; supraorbital eave not curved over base of eyestalk; carapace variously ornamented (if eyestalks long, then not flattened; carapace with few tubercles) . . 2 2. Eyestalks with more than two-thirds exposed in dorsal view ...... 3 — Eyestalks with not more than half exposed in dorsal view ...... 4 3. Postorbital lobe apex blunt; without intercalated spine; carapace with few dorsal tubercles; rostral spines with subparallel lateral margins, separate by broad V . . . Pseudomicippe ...p.379 — Postorbital lobe and intercalated spine subequal, broad and blunt; carapace without dorsal tubercles; rostral spines symmetrical ...... Anacinetops ...p.368 4. Antennal peduncle arising within orbit; carapace with numerous distinct flat blunt tubercles ...... Choniognathus ...p.368 — Antennal peduncle excluded from orbit by basal antennal article process; carapace variously ornamented ...... 5 5. Postorbital spine not cupped, eye not protected from above or below ...... 6 — Postorbital lobe cupped anteriorly to receive and protect eye, at least from above ...... 9 6. Gastric, cardiac and intestinal regions of carapace smooth; basal antennal article narrowing slightly distally and notched before anterolateral angle produced as short crenulate tooth; rostral spines not much longer than wide, with convex margins . . . . Paramithrax ...p.376 — Gastric, cardiac and intestinal regions of carapace with tubercles or spines; basal antennal article without distal notch, with anterolateral spine; rostral spines usually longer than wide, evenly tapering ...... 7 7. Orbital eave broadly rounded anteriorly, with triangular antorbital spine; basal antennal article broad, with 2 distal spines; first moveable article about as wide as long ...... Notomithrax ...p.376 — Orbital eave acute anteriorly, with acute or no antorbital spine; basal antennal article narrow, with distolateral spine only; first moveable article longer than wide ...... 8 8. Walking legs with propodi having expanded lower distal margin; carapace with at least 1 mesogastric tubercle; surface bare between groups of strong curled setae; intercalated spine present ...... Naxia ...p.374 — Walking legs with propodi cylindrical; carapace without mesogastric tubercle; surface setose between groups of strong curled setae; intercalated spine absent . . . Microhalimus ...p.372 9. Preorbital angle of eave smooth and rounded ...... 10 — Preorbital angle of eave produced as spine or tubercle ...... 13 10. Rostral spines with 1 or 2 lateral accessory spines ...... Schizophrys ...p.380 — Rostral spines simple ...... 11 11. Carapace densely spinose or tuberculate dorsally; intercalated spine engaging with or opposing a tubercle on postorbital spine in adults; adults greater than 40 mm ...... Leptomithrax ...p.368 — Carapace with few blunt tubercles but no spines dorsally (lateral spines only); intercalated spine clearly separate from postorbital spine; adults less than 35 mm ...... 12 12. Carapace scarcely broader posteriorly; basal antennal article with 2 slender anterior spines; intercalated spine larger than antorbital spine, not excluded from margin of orbit ...... Schizophroida ...p.379 — Carapace much wider posteriorly; basal antennal article with 2 short blunt spines; intercalated spine smaller than antorbital spine, excluded from margin of orbit; ...... Tumulosternum ...p.381

367 Marine Decapod Crustacea of Southern Australia

13. Rostral spines less than one-third postrostral carapace length ...... Teratomaia ...p.380 — Rostral spines more than one-third postrostral carapace length ...... 14 14. Hepatic and branchial regions with long smooth or knobbed spines, and sometimes lobes extending on to dorsal surface ...... Prismatopus ...p.378 — Hepatic and branchial region with short spines laterally only ...... Kimbla ...p.368

Anacinetops Miers, 1879 This monotypic genus has long unprotected eyestalks, like some inachines but differs from them in the orbit armature. Diagnosis. Eyestalks with more than two-thirds exposed in dorsal view. Rostral spines broad, deflexed; postorbital lobe and intercalated spine subequal, broad and blunt; carapace without dorsal tubercles. Basal antennal article with strong simple distolateral spine. Anacinetops stimpsoni (Miers, 1879) shaggy seaweed crab (Fig. 114f). 29 mm. Indonesia, WA, NT, Qld, SA; subtidal to 22 m depth. The species was figured with and without a dense coat of filamentous algae as Eruma hispidum by Hale (1927). The eyestalks are elongate and exposed. The rostrum is strongly deflexed and more elongate in smaller individuals.

Choniognathus Rathbun, 1932 Choniognathus is known from four Indo-West Pacific species, recognisable by the pear-shaped carapace covered with low tubercles, and the distinct notch behind the hepatic region which is produced as a lobe or spine (Griffin & Tranter, 1986). Diagnosis. Carapace pear-shaped, with constriction behind hepatic region, with low tubercles. Rostral spines short, strongly diverging. Supraorbital margin comprising unarmed eave, intercalated spine and postorbital lobe. Choniognathus granulosa (Baker, 1906) granulate spider crab (Fig. 113a). 25 mm. Carapace with rounded tubercles; hepatic region expanded laterally as a notched lobe, slot-like constriction behind. Maxilliped 3 with free ischium and merus. SA, WA (Rottnest I.); subtidal to 180 m depth. The species was figured as Eurynome granulosa by Hale (1927), said by him to be covered with hairs and mud.

Kimbla Griffin & Tranter, 1986 The two species of this deep-water genus differ from Leptomithrax and Teratomaia in their small size and absence of carapace spines. Diagnosis. Carapace oval, scarcely narrowing anteriorly, with weak tubercles and spines. Rostral spines strongly diverging. Supraorbital margin comprising eave without preorbital spine, intercalated spine and postorbital lobe. Kimbla franklini Richer de Forges, 1993 (Fig. 113b). Carapace with 2 small hepatic, 1 mesobranchial, 3 intestinal, 2 cardiac, 1 mesogastric and 1 protogastric spines; rostral spines longer the one-third of carapace; postocular spine tapered, with accessory spine. 8 mm. NSW (Taupo Seamount); 244 m depth.

Leptomithrax Miers, 1876 The dozen or so species of large spider crabs that comprise the genus Leptomithrax are found in the Western Pacific from Japan to New Zealand at shelf and slope depths (Griffin, 1966b; Griffin & Tranter, 1986). Species of Tumulosternum are similar (both lack a preorbital spine) but the very

368 Brachyura – crabs

small completely free intercalated spine distinguishes them from Leptomithrax. Except in juveniles, the intercalated spine in Leptomithrax engages a lobe or at least abuts a tubercle on the postorbital spine (Fig. 114g, h). Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, with dense tubercles and spines. Rostral spines not or slightly diverging. Supraorbital margin comprising eave without preorbital and with antorbital spine, with intercalated spine and postorbital lobe.

Key to southern Australian species of Leptomithrax 1. Intercalated spine not reaching end of antorbital spine on supraorbital eave, sitting dorsally so that eave and postorbital lobe almost in contact; with pits on sternum of male; walking legs not much longer than carapace length ...... 2 — Intercalated spine reaching end of antorbital spine on supraorbital eave and in same plane (shorter in juveniles); with only shallow depressions on sternum of male; walking legs 1.5 times or more carapace length ...... 4 2. With 2 branchial spines on margin of carapace and third spine more posterior and dorsal; pits on sternum of male deep, with sharp edges lined with beads ...... Leptomithrax sternocostulatus — With 3 branchial spines on margin of carapace and fourth spine more posterior and dorsal; pits on sternum of male without sharp rims ...... 3 3. Postorbital lobe spine-like or slightly truncate ...... Leptomithrax tuberculatus — Postorbital lobe excavate apically and with broad lobe on posterior margin ...... Leptomithrax depressus 4. Rostral spines slender (length about 3 times basal width); postorbital lobe with sharp tip and smaller subapical spine and tubercles near base; maxilliped 3 with proximal margin of merus with swollen smooth boss ...... Leptomithrax globifer — Rostral spines stout (length about twice basal width); postorbital lobe triangular, with subapical spine and second tubercle on posterior margin; maxilliped 3 with smooth ridge on both sides of merus–ischium suture ...... 5 5. Postorbital lobe with subapical spine prominent in adult, smaller in juvenile; middorsal regions of carapace with many spines and tubercles ...... Leptomithrax gaimardii — Postorbital lobe bifid, subapical spine equal to main apex; middorsal regions of carapace smooth except for few spines ...... Leptomithrax waitei Leptomithrax depressus Richer de Forges, 1993 (Fig. 114g). Carapace flattened overall, with 3 branchial spines on margin and fourth more posterior and dorsal; rostral spines short and slightly divergent. Intercalated spine not reaching end of antorbital spine on supraorbital eave, sitting dorsally so that eave and postorbital lobe almost in contact; postorbital lobe excavate apically, with lobe on posterior margin. 20 mm. Off NSW (Gascoyne Seamount); 143 m depth (Richer de Forges, 1993). Leptomithrax gaimardii (Milne Edwards, 1834) great spider crab (Fig. 111a, Pl. 20d, e). Carapace, middorsal regions with many spines and tubercles; rostral spines stout but more slender in juveniles. Basal antennal article prominent reaching end of eye. Intercalated spine reaching end of antorbital spine on supraorbital eave and in same plane (shorter in juveniles); postorbital lobe with subapical spine prominent in adult, smaller in juvenile, plus second spine or tubercle on posterior margin. Generally orange with fouling of sponges, ascidians, some algae. 165 mm. NSW, Vic., Tas., SA, WA; subtidal to 820 m depth. Juveniles of Leptomithrax gaimardii, which have more slender rostral spines, can be difficult to distinguish from L. globifer which is a narrower species, does not occur at shallow shelf

369 Marine Decapod Crustacea of Southern Australia

Fig. 111. Majidae Majinae. a, Leptomithrax gaimardii. b, c, Leptomithrax sternocostulatus (with detail of orbital spines).

370 Brachyura – crabs

Fig. 112. Majidae Majinae. a, Naxia aurita. b, Pseudomicippe varians.

371 Marine Decapod Crustacea of Southern Australia

depths, and has small tubercles along the posterior margin of the postorbital spines proximal to the accessory spine, instead of a single tubercle. Leptomithrax gaimardii is the most commonly encountered species of large spider crab on the southern coast. Divers have reported masses of individuals entangled in shallow water (Pl. 20e). Similar aggregations have been reported for other spider crabs such as Maja squinado (S˘tevcˇic´, 1971; Jones & Hartnoll, 1997; González-Gurriarán et al., 1998; Sampredo & González-Gurriarán, 2004). Facilitation of mating and protection during moulting at other times have been implicated in causes for aggregation. Leptomithrax globifer Rathbun, 1918) swollen spider crab (Figs 113c, 116a). Carapace, regions without spinules between major spines; rostral spines slender. Intercalated spine reaching end of antorbital spine on supraorbital eave and in same plane; postorbital lobe with sharp tip and smaller subapical spine and tubercles near base. 87 mm. Qld, Vic., Tas., SA, ?WA; shelf edge, 150–220 m depth. The species was figured by Hale (1927). The smooth swollen boss on the merus of maxilliped 3 is diagnostic. Leptomithrax sternocostulatus (Milne Edwards, 1851) ribbed spider crab (Fig. 111b, c). Carapace with 2 branchial spines on margin and third more posterior and dorsal, rostrum slender and diverging. Intercalated spine not reaching end of antorbital spine on supraorbital eave, sitting dorsally so that eave and postorbital lobe almost in contact; postorbital lobe truncate apically, with subapical lobe. Sternum of male with deep pits, each with sharp edges lined with beads. 50 mm. Qld, NSW,Vic., Tas.,SA, WA; shelf, intertidal to 140 m depth. The species distribution overlaps with that of L. gaimardii and could be almost as common. The pitted sternum of the male is distinctive. Leptomithrax tuberculatus (Whitelegge, 1900) (Fig. 114h). Carapace convex overall, with 3 branchial spines on margin of carapace and fourth more posterior and dorsal; rostral spines short and divergent. Intercalated spine not reaching end of antorbital spine on supraorbital eave, sitting dorsally so that eave and postorbital lobe almost in contact; postorbital lobe spine-like or slightly truncate. 40 mm. Qld, NSW; shelf to 144 m depth (Richer de Forges, 1993). Leptomithrax waitei (Whitelegge, 1900) (Fig. 114i). Carapace almost circular, generally spinose, middorsal regions of carapace smooth except for few spines; rostral spines stout. Intercalated spine reaching end of antorbital spine on supraorbital eave; postorbital lobe bifid, subapical spine equal to main apex. 140 mm. Qld, NSW, Tas.; slope, 200–540 m depth.

Microhalimus Haswell, 1880 The only species has been referred in the past to the genus Naxia from which it differs in the absence of intercalated spine and having cylindrical propodi on the walking legs. Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, without mesogastric tubercle; surface setose between groups of strong curled setae. Rostral spines diverging. Supraorbital margin comprising eave without preorbital, with weak or no antorbital spine, without intercalated spine, and with a postorbital spine remote from the eye and offering no protection. Walking legs with propodi cylindrical. Microhalimus deflexifrons (Haswell, 1880) (Fig. 113d). 15 mm. NSW, Vic., Tas., SA; subtidal to 71 m depth. The generic characters are sufficient to identify the species, notable for a strongly domed gastric region. The clusters of strong hooked setae often carry foliose bryozoan colonies.

372 Brachyura – crabs

ab c

def

ghi

Fig. 113. Majidae Majinae. a, Choniognathus granulosa. b, Kimbla franklini. c, Leptomithrax globifer. d, Microhalimus deflexifrons. e, Notomithrax minor. f, Notomithrax ursus. g, Paramithrax barbicornis. h, Pippacirama tuberculosa. i, Prismatopus goldsboroughi.

373 Marine Decapod Crustacea of Southern Australia

ab c

de i

g hj

Fig. 114. Majidae Majinae. a, Prismatopus occidentalis. b, Prismatopus spatulifer. c, Schizophroida hilensis. d, Tumulosternum parvispinosus. e, Tumulosternum wardi. Anterior region: f, Anacinetops stimpsoni. g, Leptomithrax depressus. h, Leptomithrax tuberculatus. i, Leptomithrax waitei. j, Pseudomicippe maccullochi.

Naxia Latreille, 1825 Decorator crabs of the genus Naxia are second in abundance to species of Notomithrax along the southern coast. All are highly cryptic, using stiff curved setae as hooks to attach seaweeds, sponges, bryozoans and hydroids to the carapace and legs. The thin chelipeds remain clear of camouflage. Species of Naxia all have expanded propodi on the walking legs, more obvious in some species

374 Brachyura – crabs

than others. Microhalimus deflexifrons is similar but has cylindrical legs and lacks an intercalated spine. The species can be distinguished by careful reference to the orbital region and the basal antennal article. The taxonomy and subfamily placement of the genus are now better clarified than before (Griffin, 1966b; Griffin & Tranter, 1986). Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, with mesogastric tubercle; surface smooth between groups of strong curled setae. Rostral spines diverging. Supraorbital margin comprising eave with preorbital spine, acute antorbital and intercalated spines, and with a postorbital spine remote from the eye and offering no protection. Basal antennal article broad, with distolateral spine only. Walking legs with propodi having expanded lower distal margin.

Key to Australian species of Naxia 1. Intercalated spine separated from supraorbital eave by notch or fused to it; hepatic spine short, blunt; anterolateral spine of basal antennal article with an accessory spinule ...... Naxia tumida — Intercalated spine well separated from supraorbital eave; hepatic spine sharp; anterolateral spine of basal antennal article unarmed ...... 2 2. Rostral spines slender, strongly curved outwards; preorbital and antorbital angles not produced; postorbital and hepatic spines close at base ...... Naxia aries — Rostral spines stout, straight, divergent; preorbital and antorbital angles spine-like; postorbital and hepatic spines well separated at base ...... 3 3. Preorbital and antorbital spines broad and triangular; intercalated spine directed laterally; walking legs with propodi only slightly expanded ...... Naxia aurita — Preorbital spine broad, upturned, antorbital spine sharp; intercalated spine directed obliquely forward; walking legs with propodi strongly expanded ...... Naxia spinosa Naxia aries (Guérin-Méneville, 1834) ramshorn crab (Figs 112a, 115a, b, 116b, Pl. 21b). Rostral spines slender, strongly curved outwards, hepatic spine sharp. Anterolateral angle of basal antennal article unarmed. Preorbital and antorbital angles not produced; intercalated spine well separated from supraorbital eave; postorbital and hepatic spines close at base. Walking legs with propodi expanded. 37 mm. Vic., Tas., SA; low intertidal to 85 m depth in Bass Strait. The rostral spines are more divergent in this species than in others. It is commonly seen. Naxia aurita (Latreille,1825) smooth seaweed crab (Figs 112a, 115c, d, Pl. 21a). Rostral spines stout, straight, divergent; hepatic spine sharp. Anterolateral angle of basal antennal article with short spine. Preorbital and antorbital spines broad and triangular; intercalated spine well separated from supraorbital eave, directed laterally; postorbital and hepatic spines well separated at base. Walking legs with propodi only slightly expanded. Yellow-brown. 70 mm. NSW, Vic., Tas., SA, WA; low intertidal to 108 m in Bass Strait. This is the most smooth of the four species and the most common in Victoria and South Australia. Naxia spinosa (Hess, 1865) spiny seaweed crab (Figs 115e, f, 116c). Rostral spines stout, straight, divergent; hepatic spine sharp. Anterolateral angle of basal antennal article with spine. Preorbital spine broad, upturned and antorbital spine sharp; intercalated spine well separated from supraorbital eave, directed obliquely forward. Walking legs with propodi strongly expanded. 70 mm. NSW, Vic., Tas., SA, WA; low intertidal to 10 m depth. Naxia tumida (Dana, 1852) little seaweed crab (Figd 115g, h, 116d). Rostral spines stout, straight, divergent; hepatic spine short, blunt. Anterolateral angle of basal antennal article a spine with an accessory spinule. Preorbital and antorbital angles not produced; intercalated spine separated from supraorbital eave by notch or fused to it; postorbital and hepatic spines well separated at base. Walking legs with propodi strongly expanded. 25 mm. Qld, NSW, SA; low intertidal to 25 m depth.

375 Marine Decapod Crustacea of Southern Australia

Notomithrax Griffin, 1963 Species of Notomithrax are heavily camouflaged with algae, sponges and hydroids. They differ from species of Naxia with which they co-occur in Victoria and Tasmania by the rounded front to the supraorbital eave. Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, dorsally tuberculate, with hooked setae. Rostral spines diverging. Supraorbital margin comprising eave without (rarely with small) preorbital, with prominent antorbital and intercalated spines, and with a postorbital spine remote from the eye and offering no protection. Basal antennal article broad, with 2 distal spines. Walking legs with propodi cylindrical.

Key to southern Australian species of Notomithrax 1. Carapace with tubercles and spines in dorsal midline; 7 marginal branchial spines ...... Notomithrax minor — Carapace with tubercles only in dorsal midline; 3 marginal branchial spines ...... Notomithrax ursus Notomithrax minor (Filhol, 1885) (Fig. 113e, Pl. 21c). Carapace with tubercles and spines in dorsal midline; protogastric region smooth; 7 marginal branchial spines. Cheliped with carpus having poorly developed ridge on upper surface and entire well developed lateral ridge. 40 mm. New Zealand, NSW, Vic., Tas.; low intertidal to 90 m depth. This is the most common decorator crab in Museum Victoria collections, usually covered with sponges rather than algae to the extent that the camouflage is bigger than the crab itself. Notomithrax ursus (Herbst, 1788) (Fig. 113f, Pl. 21d). Carapace with tubercles only in dorsal midline; 3 marginal hepatic spines and 3 marginal branchial spines. Cheliped with carpus having serrate crests. 52 mm. New Zealand, NSW,Vic., Tas.; under rocks and in rock pools on the low intertidal of rocky exposed coasts. The diet of N. ursus in New Zealand is made up of mostly of alga but guts also contain a wide variety of small gastropods and crustaceans (Woods, 1993). Algae are the usual camouflage.

Paramithrax Milne Edwards, 1834 There is only one species, with a characteristic crenulate tooth on the basal antennal article (Griffin, 1963) which, together with the shorter rostral spines, distinguishes it from species of Notomithrax. Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, dorsally weakly tuberculate, with hooked setae. Rostral spines subparallel, partially fused. Supraorbital margin comprising eave with rounded preorbital angle, obsolete antorbital spine, intercalated spine, and short conical postorbital spine separated from eave by narrow fissures. Basal antennal article notched before anterolateral angle produced as short crenulate tooth, with short distomesial spine. Walking legs with propodi cylindrical. Paramithrax barbicornis (Latreille, 1825) sea toad (Fig. 115i). 30 mm. NSW, Vic., SA, WA; rocky subtidal reef. The notched and crenulate basal antennal article and the good protection provided by the rounded anterolateral region of the orbital eave distinguish this species from the similar Notomithrax ursus.

376 Brachyura – crabs

ab cd

ef gh

i jkl

p n o

Fig. 115. Majidae Majinae. Left anterior region, dorsal and ventral: a, b, Naxia aries. c, d, Naxia aurita. e, f, Naxia spinosa. g, h, Naxia tumida. Left anterior region, ventral: i, Paramithrax barbicornis. j, Pseudomicippe varians. k, Schizophrys aspera. l, Schizophrys dama. m, Schizophrys rufescens. n, o, Teratomaia richardsoni. p, Leptomithrax sternocostulatus (anterior male sternum).

377 Marine Decapod Crustacea of Southern Australia

abcd

Fig. 116. Majidae Majinae. Maxilliped 3: a, Leptomithrax globifer. Walking leg 4: b, Naxia aries. c, Naxia spinosa. d, Naxia tumida.

Pippacirama Griffin & Tranter, 1986 The only species is endemic to southern Australia. Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, dorsally spinose and tuberculate, with hooked setae. Rostral spines deflexed, basally fused. Supraorbital margin comprising eave with short antorbital lobe only, without intercalated spine, and with lamellate postorbital spine with lobe on anterior margin, well separated from eave. Basal antennal article broad, with short anterolateral spine. Walking legs with propodi cylindrical, tuberculate. Pippacirama tuberculosa (Milne Edwards, 1834) (Fig. 113h). 21 mm. NSW, Vic., SA, WA; rocky reef from intertidal to 7 m depth. Depressed short rostral lobes immediately identify this species but size of the rostral lobes and degree of tuberculation on the carapace vary. The long flattened eyestalks are also extraordinary. The species was figured as Paramicippe tuberculosa by Hale (1927).

Prismatopus Ward, 1933 Griffin & Tranter (1986) treated species included in Prismatopus as belonging to their new genus, Thacanophrys. Earlier, several of its species had been included in Chlorinoides Haswell, 1880. ICZN rules dictate that Ward’s name has priority at least for the three treated here (Davie, 2002). Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, branchial region with 1–3 smooth or knobbed spines, sometimes lobes. Rostral spines slender, diverging, more than one-third postrostral carapace length. Supraorbital margin comprising eave with preorbital and antorbital spines, with intercalated spine, and postorbital lobe excavated anteriorly to receive eyestalk; all closely approximated. Basal antennal article with 2 distal teeth. Male chela palm about 1.5 times as long as high.

Key to southern Australian species of Prismatopus 1. Mesial margin of rostral spine armed with short lobe or spines; preorbital lobe lamellate, wide, truncate or concave; antorbital spine short or absent ...... Prismatopus spatulifer — Mesial margin of rostral spine with tubercles; preorbital and antorbital lobes spine-like . . 2 2. Rostral spines no more than third postrostral carapace length, tips separated by less than twice width at base ...... Prismatopus goldsboroughi

378 Brachyura – crabs

— Rostral spines half postrostral carapace length, tips separated by more than twice width at base ...... Prismatopus occidentalis Prismatopus goldsboroughi (Rathbun, 1906) (Fig. 113i). Rostral spines no more than third postrostral carapace length, without mesial lobes, with tubercles, tips separated by less than twice width at base. Preorbital and antorbital lobes spine-like. 19 mm. Hawaii, Kermadec Is, NSW, Vic.; 126–401 m depth. Prismatopus occidentalis (Griffin, 1970) (Fig. 114a). Rostral spines half postrostral carapace length, without mesial lobes, with tubercles, tips separated by more than twice width at base. Preorbital and antorbital lobes spine-like. 17 mm. WA (Geraldton to Rottnest I.); 111–187 m depth. Prismatopus spatulifer (Haswell, 1882) (Fig. 114b). Rostral spines half postrostral carapace length, each with mesial lobe or spines, tips separated by 3 times width at base. Preorbital lobe lamellate, wide, truncate or concave; antorbital spine short. 25 mm. NSW, Vic., Tas., SA, WA; shelf and slope, to 460 m depth. The species immediately stands out with its bifid plate standing vertically near the posterior margin of the carapace and the leaf-shaped plate directed posteriorly.

Pseudomicippe Heller, 1861 Griffin & Tranter (1986) reviewed the taxonomy of this Indo-West Pacific genus of a dozen species. All have long eyestalks protruding from an egg-shaped body. Two of the four Australian species occurs along the southern coast. Diagnosis. Carapace egg-shaped, barely narrowing anteriorly, dorsally tuberculate, with few spines, with hooked setae. Rostral spines sometimes deflexed, sometimes basally fused. Supraorbital margin comprising eave expanded anteriorly, with short antorbital lobe, without intercalated spine, and with postorbital spine, separated from eave by narrow fissure. Basal antennal article broad, with prominent anterolateral spine. Walking legs with propodi cylindrical, tuberculate.

Key to southern Australian species of Pseudomicippe 1. With pair of tubercles near midline between eyes; eyestalk well exceeding postorbital spine ...... Pseudomicippe varians — With single tubercles in midline between eyes; eyestalk barely exceeding postorbital spine ...... Pseudomicippe maccullochi Pseudomicippe maccullochi Griffin & Tranter, 1986 (Fig. 114j). Carapace with prominent branchial tubercles and middorsal row of 4–5 mesogastric tubercles, second the largest. Rostral spines strongly deflexed. Supraorbital eave with preorbital lobe expanded. Basal antennal article with spine on anterolateral angle, laterally concave. Eyestalk barely exceeding postorbital spine. 24 mm. Lord Howe I., Norfolk I., Qld, Vic.; low rocky intertidal. Pseudomicippe varians Miers, 1879 (Fig. 112b). Carapace with 4–5 branchial tubercles; pair of mesogastric tubercles between eyes followed by 3 in midline. Rostral spines strongly deflexed. Supraorbital eave with preorbital lobe expanded. Basal antennal article with lobe on anterolateral angle, laterally concave. Eyestalk well exceeding postorbital spine. 24 mm. Qld, WA (S to Rottnest I.); low rocky intertidal to shelf.

Schizophroida Sakai, 1933 Griffin & Tranter (1986) reviewed the taxonomy of this Indo-West Pacific genus with one widespread species and one or two from Japan.

379 Marine Decapod Crustacea of Southern Australia

Diagnosis. Carapace pear-shaped, narrowing anteriorly, dorsally weakly tuberculate, with hooked setae. Rostral spines subparallel. Supraorbital margin comprising eave with preorbital lobe rounded, with short antorbital lobe, with intercalated spine, and with postorbital spine cupped to receive eyestalk. Basal antennal article with 2 slender anterior spines. Walking legs with propodi cylindrical. Schizophroida hilensis (Rathbun, 1906) (Fig. 114c). 33 mm. West Pacific, Tasman Sea, Qld, NSW; low intertidal reef. The combination of parallel thin rostral spines and two strong distolateral spines on the basal antennal article are diagnostic.

Schizophrys White, 1848 Three of the four species known occur in Australia (Griffin & Tranter, 1986). The lateral accessory spines on the rostrum identify the genus and differentiate the species. Diagnosis. Carapace pear-shaped, narrowing anteriorly, dorsally weakly tuberculate, with hooked setae. Rostral spines subparallel, each with 1 or 2 lateral accessory spines. Supraorbital margin comprising eave rounded anterolaterally, with short antorbital lobe, with intercalated spine, and with postorbital spine bearing anterior accessory spine, cupped to receive eyestalk. Basal antennal article with 2 slender anterior spines. Walking legs with propodi cylindrical.

Key to Australian species of Schizophrys 1. Rostral spines each with 2 lateral accessory spines ...... Schizophrys dama — Rostral spines each with 1 lateral accessory spine ...... 2 2. Lateral accessory spine far removed from rostrum apex; gastric region with transverse row of 4 tubercles or spines ...... Schizophrys aspera — Lateral accessory spine and rostrum apex subequal; gastric region granular, without tubercles or spines ...... Schizophrys rufescens Schizophrys aspera (Milne Edwards, 1834) red sea toad – so-called because it is red, not because it is found in the Red Sea! (Fig. 115k). Rostral spine with 1 lateral accessory spine, almost equal to rostral apex. Gastric region with transverse row of 4 tubercles or spines. Reddish. 45 mm. Indo-West Pacific, Qld, NSW, SA, WA; reefs, shallow subtidal, with coral and sponge. Schizophrys dama (Herbst, 1804) (Fig. 115l). Rostral spine with 2 lateral accessory spines. Gastric region granular, without tubercles or spines. 59 mm. WA, NT, Qld; coral reef, subtidal, with coral and sponge. Essentially a tropical species, the species was collected once in the Great Australian Bight in 1888. Schizophrys rufescens Griffin & Tranter, 1986 (Fig. 115m, Pl. 21e). Rostral spine with 1 lateral accessory spine far removed from rostrum apex. Gastric region granular, without tubercles or spines. Reddish. 58 mm. NT, Qld, SA, WA; under rocks, reef to 40 m depth.

Teratomaia Griffin & Tranter, 1986 The single species of Teratomaia is distinguished from species of Leptomithrax by the possession of an preorbital spine. It is common in deep-sea trawl by-catch. Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, with tubercles and long spines. Rostral spines slender, diverging, one-third or less postrostral length. Supraorbital margin comprising eave with preorbital and antorbital spines, with intercalated spine and postorbital lobe excavated anteriorly to receive eyestalk; all closely approximated. Basal antennal article with 2 distal teeth. Teratomaia richardsoni (Dell, 1960) (Fig. 115n, o). 102 mm. New Zealand, NSW, Vic., Tas.; 224–648m depth. Griffin (1966b) described the rostrum as one-fifth the postrostral length; in at least some Australian material it is longer, almost one-third postrostral length.

380 Brachyura – crabs

Tumulosternum McCulloch, 1913 Tumulosternum is a genus of three species endemic to eastern Australia. The species may easily be confused with species of Leptomithrax but the broader body strongly tapering anteriorly, and the very small intercalated spine completely free of the postorbital spine are diagnostic. Diagnosis. Carapace pear-shaped, strongly narrowing anteriorly, with tubercles and short spines, hepatic margin with strong spine or tubercle. Rostral spines short, broad. Supraorbital margin comprising eave without preorbital spine, with antorbital spine directed outwards, with small intercalated spine (sometimes absent) and angular postorbital lobe excavated anteriorly to receive eyestalk. Basal antennal article bilobed anteriorly.

Key to species of Tumulosternum 1. Carapace without lateral branchial spines; subhepatic region with subcircular flattened plate; similar plates elsewhere on lower surface and margins of carapace ...... Tumulosternum longimanus — Carapace with 2 or more lateral branchial spines, without flattened plates ...... 2 2. Branchial margin with 2 spines; anterolateral spine of basal antennal article as wide as long; rostral spines fused for basal third, then abruptly tapering . . . Tumulosternum parvispinosus — Branchial margin with 4 spines (3 lateral and fourth more dorsal); anterolateral spine of basal antennal article longer than wide; rostral spines triangular, tapering evenly from base ...... Tumulosternum wardi Tumulosternum longimanus (Haswell, 1880). Carapace without lateral branchial spines; subhepatic region with subcircular flattened plate; similar plates elsewhere on lower surface and margins of carapace. Rostral spines triangular, tapering evenly from base. 19 mm. NSW, WA (Rottnest I.); low intertidal and subtidal reef, under rocks. Tumulosternum parvispinosus Ward, 1933 (Fig. 114d). Carapace with 2 lateral branchial spines, without flattened plates. Rostral spines fused for basal third, then abruptly tapering. 23 mm. Qld, NSW; low intertidal and subtidal reef, under coral and rocks. Tumulosternum wardi Griffin & Tranter, 1986 (Fig. 114e). Carapace with 4 lateral branchial spines, without flattened plates. Rostral spines triangular, tapering evenly from base. 26 mm. NSW (Sydney), Vic. (Western Port); subtidal reef.

Subfamily Mithracinae MacLeay, 1838 Mithracine crabs are more rectangular than most other majids, essentially because of the tubular orbits sheltering the eyestalks below (with an expanded basal antennal article) as well as from above and front and back. Griffin & Tranter (1986) did not believe that these were necessarily one evolutionary lineage, a view supported by cladistic analyses of larval characters (Pohle & Marques, 2000; Marques et al., 2003). Unlike other subfamilies, the Indo-West Pacific fauna is relatively impoverished compared with that in the Americas. Three genera and six species inhabit southern Australia; four genera and 14 species in the continent as a whole. Diagnosis. Carapace broadened anteriorly by the outstanding, often tubular, nearly complete, or complete, orbits. Basal antennal article moderately broad, expanded to form floor to orbit which is formed above by an arched supraorbital eave or semi-tubular horn, and an adjacent hollowed postorbital lobe; intercalated spine sometimes present. Retracted eye can be completely concealed. Rostrum often more or less deflexed. Male gonopod 1 usually slender, straight or weakly curved; aperture terminal; apex simple or curved, or expanded into 2 or 3 processes. (after Griffin & Tranter, 1986)

381 Marine Decapod Crustacea of Southern Australia

Key to southern Australian genera of Mithracinae 1. Rostral spines broad, flattened, depressed, fused for at least proximal two-thirds; intercalated spine present (sometimes fused to postorbital spine) ...... Micippa — Rostral spines slender, cylindrical, separate from base; intercalated spine absent ...... 2 2. Rostral spines separate, with single mesial distal accessory spine ...... Paranaxia — Rostral spines fused over most of length, with 2–4 lateral accessory spines (unarmed in some species outside region) ...... Tiarinia

Micippa Leach, 1817 Half of the ten species of this Indo-West Pacific genus are represented in southern Australia (Griffin & Tranter, 1986). All are bizarre crabs with a broad depressed rostral plate. Diagnosis. Carapace scarcely narrowed anteriorly, tuberculate. Rostral spines broad, flattened, depressed, fused for at least proximal two-thirds; intercalated spine present (sometimes fused to postorbital spine).

Key to southern Australian species of Micippa 1. Rostrum with distal notch ...... 2 — Rostrum divided for distal quarter or more ...... 3 2. Carapace with 3 spines or tubercles along dorsal midline; intercalated spine separated from postorbital lobe by narrow fissure; rostrum directed anteroventrally ...... Micippa spinosa — Carapace without spines along dorsal midline; intercalated spine adjacent to postorbital lobe; rostrum directed posteroventrally ...... Micippa curtispina 3. Rostrum bifid, apices diverging ...... Micippa thalia — Rostrum with 2 short apical teeth or lobes, and lateral tooth on each side ...... 4 4. Rostral lateral tooth acute, recurved; basal antennal article smooth; orbit usually open below, with a hiatus between basal antennal article and postorbital lobe; anterolateral borders of carapace with 8–10 spines ...... Micippa platipes — Rostral lateral tooth directed laterally; basal antennal article tuberculate; orbit closed below, with basal antennal article and postorbital lobe in contact; anterolateral borders of carapace with 3–6 spines ...... Micippa philyra Micippa curtispina Haswell, 1880 (Fig. 117a, b). Rostrum with distal notch. Carapace without spines along dorsal midline; intercalated spine adjacent to postorbital lobe. 19 mm. Singapore, Qld, NSW, WA (S to Rottnest I.); subtidal reef. Micippa philyra (Herbst, 1803) (Fig. 117a). Rostrum ending in 2 medial triangular teeth and short lateral pair directed laterally. Basal antennal article tuberculate; orbit closed below, with basal antennal article and postorbital lobe in contact; anterolateral borders of carapace with 3–6 spines. 23 mm. Indo-West Pacific, NT, Qld, NSW, Vic., SA, WA; intertidal reef to about 90 m depth. Micippa platipes Rüppell, 1830 (Fig. 117d). Rostrum divided for distal quarter, with short broad recurved spine on each side. Basal antennal article smooth; orbit open below, with a hiatus between basal antennal article and postorbital lobe; anterolateral borders of carapace with 8–10 spines. 36 mm. Indo-West Pacific, Qld, NSW; subtidal reef. Micippa spinosa Stimpson, 1858 spiny flat-beaked crab (Fig. 117e). Rostrum with distal notch. Carapace with 3 spines along dorsal midline; intercalated spine separated from postorbital lobe by narrow fissure. Basal antennal article smooth, orbit closed below. 21 mm. NSW, Vic.,

382 Brachyura – crabs

b c

f d e

h i

Fig. 117. Majidae Mithracinae. a, b, Micippa curtispina (carapace, anterior view). c, Micippa philyra (anterior view of rostrum). d, Micippa platipes (anterior view of rostrum). e, Micippa spinosa (anterior region, dorsal). f, Micippa thalia (anterior view of rostrum). g, h, Paranaxia serpulifera (carapace and lateral view of right orbit and base of rostrum). i, Tiarinia angusta (rostrum and orbits).

SA; subtidal to 64 m depth. Hale (1927) figured this species and referred to a variety without strong dorsal tubercles described originally as Paramicippe affinis Miers, 1879. Specimens from Bass Strait are of this form. Micippa thalia (Herbst, 1803) (Fig. 117f). Rostrum divided for almost half length, distal teeth diverging. Basal antennal article smooth; orbit open below, with gaps between basal antennal article, postorbital lobe, intercalated spine and antorbital spine. Red. 26 mm. Indo-West Pacific, NT, Qld, NSW, Vic., SA, WA; 20–100 m depth.

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Paranaxia Rathbun, 1924 Paranaxia is a monotypic tropical Australian genus. Diagnosis. Carapace width two-thirds postrostral carapace length, branchial region 1.5 times as wide as hepatic region. Rostral spines separate, slender, cylindrical, separate from base, with single mesial accessory spine. Preorbital spine large; intercalated spine absent. Paranaxia serpulifera (Guérin-Méneville, 1829) (Fig. 117g, h). 116 mm. Indonesia, NT, Qld, WA (S to Cockburn Sound); 7–30 m depth. The crab is covered with a sponge of fine hairs and has characteristic rostral spines which converge distally; the mesial accessory spines meet in the middle. A pelagic larval phase is lacking and young crabs are brooded directly (Morgan, 1987).

Tiarinia Dana, 1851 Of eleven species of this Indo-West Pacific genus only one reaches southern Australia. But that too is essentially a tropical species. Diagnosis. Carapace greatest width twice interorbital width. Rostral spines fused or contiguous, with or without lateral spines. Preorbital lobe blunt; intercalated spine absent. Tiarinia angusta Dana, 1852 (Fig. 117i). Rostral spines fused along most of length, with lateral accessory spines. 50 mm. Japan to Indonesia, NT, Qld, WA (S to Rottnest I.); intertidal and shallow subtidal. Related species from north of Australia are distinguished only by examining reproductive structures (Griffin & Tranter, 1986).

Subfamily Pisinae Alcock, 1895 The characteristic feature of pisine spider crabs is the prominent postorbital tooth or lobe which is cupped to receive the retracted eyestalk but which never completely conceals the cornea in dorsal view. Although the subfamily is the most diverse throughout the Indo-West Pacific and tropical Australia it is represented on the southern coast by only five genera and ten species. Herbstia crassipes (Milne Edwards, 1873) was described from Bass Strait but its identity is doubtful (Griffin & Tranter, 1986). Diagnosis. Orbit comprising supraorbital eave, with or without preorbital spine; with or without intercalated spine; and large prominent postorbital tooth or lobe, usually cupped to receive retracted eyestalk, but which never completely conceals cornea in dorsal view. Eyestalks short. Basal antennal article broad, at least at base; always fused distally; anterior angle usually produced as tooth or spine. Merus of maxilliped 3 expanded at distolateral angle; wider than ischium; palp at distomesial angle. Rostrum usually 2-spined. Male abdomen not terminally broadened, somite 7 subtriangular and not inserted deeply into somite 6. (after Griffin & Tranter, 1986)

Key to southern Australian species of Pisinae 1. Carapace subcircular; rostrum a short triangular fused plate for most of length, with short apical spines; legs long and slender ...... Phalangipus … 2 — Carapace pear-shaped; rostral spines of various lengths, rarely fused; legs not long and slender ...... 3 2. Posterior margin with short median tubercle, not overlapping carapace edge; male chelipeds smooth ...... Phalangipus australiensis — Posterior margin with long median spine, well overlapping carapace edge; male chelipeds spiny ...... Phalangipus hystrix

384 Brachyura – crabs

3. Rostrum lyre-shaped (curving outwards distally), spines separated by parallel-sided slit ...... Lyramaia elegans — Rostrum rostral spines diverging from base (sometimes curving outwards distally) ...... 4 4. Rostral spines with dorsal accessory spine in distal half; with intercalated spine ...... Naxioides robillardi — Rostral spines without dorsal accessory spine; without intercalated spine ...... 5 5. Supraorbital eave with weakly developed antorbital angle; postorbital lobe convex laterally ...... Hyastenus ...6 — Supraorbital eave with preorbital spine or plate, without antorbital angle; postorbital lobe sometimes laterally with a plate adjacent to plate on hepatic region ...... Rochinia . . . 7 6. Rostrum one-quarter postrostral length; carapace strongly tuberculate; branchial region with 3 or 4 marginal and 3–5 submarginal tubercles plus epibranchial spine . . . . . Hyastenus sebae — Rostrum three-quarters postrostral length; carapace mostly smooth; branchial region with epibranchial spine and no other tubercle ...... Hyastenus elatus 7. Rostral spines about one-sixth of total carapace length; with prominent medial cardiac spine; walking legs with meri dorsally carinate ...... Rochinia mosaica — Rostral spines about one-quarter of total carapace length; with medial cardiac tubercle conical; walking legs with meri not carinate ...... Rochinia fultoni

Hyastenus White, 1847 The genus was discussed at length by Griffin & Tranter (1986) who recognised 34 species throughout the Indo-West Pacific. One of the southern Australian species is tuberculate while the other is smooth. Both have long diverging rostral spines. Diagnosis. Carapace pear-shaped, variously tuberculate or spinous. Orbit comprising supraorbital eave with preorbital spine small or slender, weak antorbital spine, without intercalated spine. Postorbital lobe separated by narrow hiatus, anterior margins sometimes with proximal lobe, posterior margin convex, not keeled nor lobed. Basal antennal article narrow, distolateral angle blunt or a small spine, lateral margin convex or a triangular lobe; article separated from postorbital lobe by U-shaped hiatus. Hyastenus elatus Griffin & Tranter, 1986 (Fig. 118a). Carapace mostly smooth; rostrum three- quarters postrostral length; branchial region with epibranchial spine and no other tubercle. 88 mm. Indonesia, Singapore, NT, Qld, NSW (S to Botany Bay), WA (S to Perth); intertidal to 72 m depth. Hyastenus sebae White, 1847. Carapace strongly tuberculate on most regions; rostrum one-quarter postrostral length; branchial region with 3 or 4 marginal and 3–5 submarginal tubercles plus epibranchial spine. 33 mm. Eastern Indo-West Pacific, NT, Qld, WA (S to Rottnest I.); intertidal to 120 m depth. H. oryx Milne Edwards, 1872 is a junior synonym.

Lyramaia Griffin & Tranter, 1986 The only species resembles species of Hyastenus but differs in the lyre-shape of the rostral spines. Diagnosis. Carapace pear-shaped, slightly tuberculate and spinous. Orbit comprising supraorbital eave with weak preorbital and antorbital spines, without intercalated spine; postorbital lobe weakly excavate, posterior margin convex, not keeled nor lobed. Basal antennal article distolateral angle a small spine, lateral margin produced and directed dorsally; article separated from postorbital lobe by hiatus.

385 Marine Decapod Crustacea of Southern Australia

Lyramaia elegans (Haswell, 1881) (Fig. 118b). Rostrum with median slit and spines diverging apically. 14 mm. Qld, NSW; subtidal to 80 m depth.

Naxioides Milne Edwards, 1865 The combination of a small dorsal accessory spine on the rostral spines and an intercalated spine identify the six species of this Indo-West Pacific genus (Griffin & Tranter, 1986). Diagnosis. Carapace pear-shaped, slightly tuberculate and spinous. Rostral spine with small dorsal accessory spine. Orbit comprising supraorbital eave with preorbital and antorbital spines, with intercalated spine; postorbital lobe weakly excavate, posterior margin convex, not keeled nor lobed. Basal antennal article with distolateral spine, lateral margin with lobe; article separated from postorbital lobe by hiatus. Naxioides robillardi (Miers, 1882) (Fig. 118c). Carapace with 14–20 long dorsal spines; cardiac region with 1 medial spine; hepatic margin with 1 spine. Supraorbital eave with upright preorbital spine, antorbital spine directed laterally. Basal antennal article with 1 spine each on distolateral angle and on lateral margin. 43 mm. Indo-West Pacific, NSW; 30–360 m depth.

e g d

Fig. 118. Majidae Pisinae. a, Hyastenus elatus. b, Lyramaia elegans. c, Naxioides robillardi (carapace, lateral). d, Phalangipus australiensis. e, Phalangipus hystrix. f, Rochinia fultoni. g, Rochinia mosaica.

386 Brachyura – crabs

Guinot & Richer de Forges (1985) included photographs of this very spiky species. The small accessory spine on the upper side of the rostral spine identifies the genus, not to be confused with the spines on the mesial faces of the rostral spines of Paranaxia serpulifera.

Phalangipus Latreille, 1828 Species of Phalangipus have a broad carapace with a rostrum of two short partially fused spines, and extremely long slender walking legs. There are nine Indo-West Pacific species (Griffin, 1973; Griffin & Tranter, 1986). Diagnosis. Carapace subcircular, with tubercles and spines. Rostral spines short, fused for half length spine. Orbit comprising supraorbital eave with supraorbital spine midway along, sometimes antorbital spine, with intercalated spine; postorbital lobe weakly excavate, posterior margin convex, not keeled nor lobed. Basal antennal article with distolateral lobe, lateral margin with suborbital lobe; article. Phalangipus australiensis Rathbun, 1918 (Fig. 118d). Posterior margin with short median tubercle, not overlapping carapace edge; branchial region with 2 lateral and 1 submarginal tubercles; lateral clusters of tubercles dorsally; male chelipeds smooth. 25 mm. Indonesia, WA, NT, Qld, NSW (S to Port Stephens); 5–120 m depth. Phalangipus hystrix (Miers, 1886) (Fig. 118e). Posterior margin with long median backward- directed spine, well overlapping carapace edge; branchial region with 3 marginal and 1 submarginal tubercles; protogastric regions with 4 pairs of tubercles in diverging rows, median mesogastric spine anteriorly; male chelipeds spiny. 35 mm. Indo-West Pacific, WA (S to Rottnest I.); 50–353 m depth.

Rochinia Milne Edwards, 1875 The absence of an intercalated spine and the collusion, more or less, of the postorbital spine with a spine or lobe on the hepatic region identify members of this large genus. It is represented by about 30 species throughout all the world’s tropical and temperate oceans (Griffin & Tranter, 1986) including two recently described from seamounts off the eastern Australian coast (Davie & Short, 1989; Richer de Forges, 1992). Diagnosis. Carapace pear-shaped to elongate, variously tuberculate or spinous. Orbit comprising supraorbital eave with small preorbital spine or plate, without antorbital spine, without intercalated spine. Postorbital lobe separated by U-shaped hiatus, hardly excavate, posterior margin sometimes with plate fused to plate on hepatic region. Basal antennal article narrow, distolateral angle a small tooth, lateral margin entire lobe; article separated from postorbital lobe by narrow U-shaped hiatus. Rochinia fultoni (Grant, 1905) (Fig. 118f). Rostral spines about one-quarter of total carapace length; with medial cardiac tubercle conical. Walking legs with meri not carinate. 30 mm. NSW, Tas.; subtidal. Rochinia mosaica (Whitelegge, 1900) little thorn-back crab (Fig. 118g). Rostral spines about one- sixth of total carapace length; with prominent medial cardiac spine. Walking legs with meri dorsally carinate. 15 mm. Qld, NSW, Vic., SA; 20–200 (890) m depth.

References Clark, P.F., & Webber, W.R. 1991. A redescription of Macrocheira kaempferi (Temminck, 1836) zoeas with a discussion of the classification of the Majoidea Samouelle, 1819 (Crustacea: Brachyura). Journal of Natural History 25: 1259–1279.

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Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Davie, P.J.F., & Short, J.W. 1989. Deepwater Brachyura (Crustacea: Decapoda) from southern Queensland, Australia with descriptions of four new species. Memoirs of the Queensland Museum 27: 157–187. Drach, P., & Guinot, D. 1983. Les Inachoididae Dana, famille de Majoidea caractérisée par des connex- ions morphologiques d’un type nouveau entre carapace, pleurites, sternites, pléon. Comptes Rendus Hebdomadaires de Séances de l’Académie des Sciences, Paris 297: 37–42. Forest, J. 1978. Le genre Macropodia Leach dans les eaux atlantiques européennes (Crustacea Brachyura Majidae). Cahiers de Biologie Marine 19: 323–342. Fransozo, A., & Negreiro-Fransozo, M.L. 1970. Larval stages of Pyromaia tuberculata (Lockington, 1877) (Decapoda, Majidae, Inachinae) reared in the laboratory. Crustaceana 70: 304–323. Furota, T. 1988. The ecology of the introduced spider crab Pyromaia tuberculata in an organically polluted bay (preliminary report). Benthos Research 33/34: 79–89. Furota, T. 1990. Population structure of the introduced spider crab Pyromaia tuberculata in the upper- most region of Tokyo Bay. Benthos Research 39: 1–7. Furota, T. 1996. Life cycle studies on the introduced spider crab Pyromaia tuberculata (Lockington) (Brachyura: Majidae). I. Egg and larval stages. Journal of Crustacean Biology 16: 71–76. Furota, T., & Furose, K. 1988. Distribution of the introduced spider crab, Pyromaia tuberculata, along the coast of Japan. Benthos Research 33/34: 75–78. González-Gurriarán, E., Fernández, L., Freire, J., & Muiño, R. 1998. Mating and role of seminal receptacles in the reproductive biology of the spider crab Maja squinado (Decapoda, Majidae). Journal of Experimental Marine Biology and Ecology 220: 269–285. Griffin, D.J.G. 1963. Notomithrax gen. nov. and the status of the genus Paramithrax H. Milne Edwards (Crustacea, Brachyura, Majidae). Transactions of the Royal Society of New Zealand 3: 229–237. Griffin, D.J.G. 1966a. A review of the Australian majid spider crabs (Crustacea, Brachyura). Australian Zoologist 13: 259–98, pls 15–17. Griffin, D.J.G. 1966b. The marine fauna of New Zealand: spider crabs, family Majidae (Crustacea, Brachyura). Bulletin, New Zealand Department of Scientific and Industrial Research 172: 1–111. Griffin, D.J.G. 1970. The Australian majid spider crabs of the genus Achaeus (Crustacea, Brachyura). Journal of the Royal Society of Western Australia 53: 97–119. Griffin, D.J.G. 1973. A revision of the spider crabs of the genus Phalangipus (Crustacea, Brachyura, Majidae). Journal of Natural History 7: 165–207. Griffin, D.J.G., & Tranter, H.A. 1986. The Decapoda Brachyura of the Siboga Expedition. Part VIII Majidae. Siboga Expéditie Monographie 39: 1–335. Griffin, D.J.G., & Yaldwyn, J.C. 1965. A record of the majid brachyuran genus Achaeus from New Zealand with notes on the Australian species. Transactions of the Royal Society of New Zealand 6: 33–51. Guinot, D., & Richer de Forges, B. 1982. Révision du genre Indo-Pacifique Cyrtomaia Miers, 1886: Campagnes océanographiques du Challenger, de l’Albatross,du Siboga et du Vauban (Crustacea Decapoda Brachyura). Annales de l’Institut Océanographique, Monaco et Paris 58: 5–88. Guinot, D., & Richer de Forges, B. 1985. Crustacés Décapodes: Majidae (genres Platymaia, Cyrtomaia, Pleistacantha, Sphenocarcinus et Naxioides). In: Résultats des Campagnes MUSORSTOM I et II –Philippines (1976, 1980). Tome 2. Mémoires du Muséum National d’Histoire Naturelle, Paris A (Zoologie) 133: 83–179. Guinot, D., & Richer de Forges, B. 1997. Affinités entre les Hymenosomatidae MacLeay, 1838 et les Inachoididae Dana, 1851 (Crustacea, Decapoda, Brachyura). Zoosystema 19: 453–502. de Haan, W. 1839. Crustacea. Pp. 1–243, pls 1–55, A–Q in: de Siebold, P.F. (ed.) Fauna Japonica sive descriptio animalium ... Leiden. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp.

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Jones, D.R., & Hartnoll, R.G. 1997. Mate selection and mating behaviour in spider crabs. Estuarine, Coastal and Shelf Science 44: 185–193. Lemaitre, R., Campos, N.H., & Bermúdez, A. 2001. A new species of Pyromaia from the Caribbean Sea, with a rediscription of P. propinqua Chace, 1940 (Decapoda: Brachyura: Majoidea: Inachoididae). Journal of Crustacean Biology 21: 760–773. Loh, L.W., & Ng, P.K.L. 1999. A revision of the spider crabs of the genus Paratymolus Miers, 1879, with descriptions of two new genera and six new species (Crustacea: Decapoda: Brachyura: Majidae). Raffles Bulletin of Zoology 47: 365–407. Marques, F.P.L., & Pohle, G. 1988. The use of structural reduction in phylogenetic reconstruction of decapods and a phyloenetic hypothesis for fifteen genera of Majidae: testing previous hypotheses and assumptions. Invertebrate Reproduction and Development 33: 241–262. Marques, F.P.L., Pohle, G.W., & Vrbova, L. 2003. On the larval stages of Macrocoeloma diplacanthum (Decapoda: Brachyura: Majidae), with a review of mithracine phylogenetic aspects. Journal of Crustacean Biology 23: 187–200. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. Miers, E.J. 1879. Crustacea. In: An account of the petrological, botanical, and zoological collections made in Kerguelen’s Land and Rodriguez during the Transit of Venus Expeditions ... in the years 1874–75. Philosophical Transactions of the Royal Society of London 168: 200–214, pl. 11. Morgan, G.J. 1987. Brooding of juveniles and observations on dispersal of young in the spider crab Paranaxia serpulifera (Guérin)(Decapoda, Brachyura, Majidae) from Western Australia. Records of the Western Australian Museum 13: 337–343. Morgan, G.J. 1990. An introduced eastern Pacific majid crab from Cockburn Sound, southwestern Australia. Crustaceana 58: 316–317. Parry, G.D., Hobday, D.K., Currie, D.R., Officer, R.A., & Gason, A.S. 1995. The distribution, abundance and diets of demersal fish in Port Phillip Bay. CSIRO, Port Phillip Bay Environmental Study Technical Report 21: 1–107. Pohle, G., & Marques, F.P.L. 2000. Larval stages of Paradasygyius depressus (Bell, 1835) (Crustacea: Decapoda: Brachyura: Majidae) and a phylogenetic analysis for 21 genera of Majidae. Proceedings of the Biological Society of Washington 113: 739–760. Poore, G.C.B., & Storey, M. 1999. Soft sediment Crustacea of Port Phillip Bay. Centre for Research on Introduced Marine Pests, CSIRO Marine Research, Technical Report 20: 150–170. Richer de Forges, B. 1992. A new species of Sphenocarcinus A. Milne Edwards, 1875 trom Tasmantid guyots, S. lowryi n. sp. (Crustacea, Decapoda, Brachyura). Records of the Australian Museum 44: 1–5. Richer de Forges, B. 1993. Deep sea crabs of the Tasman seamounts (Crustacea: Decapoda: Brachyura). Records of the Australian Museum 45: 11–24. Richer de Forges, B. 1994. A new genus of deep-sea majid crab: Griffinia gen. nov. (Crustacea, Decapoda, Brachyura). The Beagle, Occasional Papers of the Northern Territory Museum of Arts and Sciences 11: 65–72. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates. Sampredo, M.-P., & González-Gurriarán, E. 2004. Aggregating behaviour of the spider crab Maja squinado in shallow waters. Journal of Crustacean Biology 24: 168–177. Schejter, L., Spivak, E.D., & Luppi, T.A. 2002. Presence of Pyromaia tuberculata (Lockington, 1877) adults and larvae in the Argentine continental shelf (Crustacea: Decapoda: Majoidea). Proceedings of the Biological Society of Washington 115: 605–610. S˘tevcˇic´, Z. 1971. Laboratory observations on the aggregations of the spiny spider crab Maja squinado Herbst. Animal Behaviour 19: 18–25. Webber, W.R., & Richer de Forges, B. 1995. Deep sea Majidae (Decapoda: Brachyura) new to New Zealand with a description of Oxypleurodon wanganella sp. nov. Journal of the Royal Society of New Zealand 25: 501–516.

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Webber, W.R., & Wear, R.G. 1981. Life history studies on New Zealand Brachyura 5. Larvae of the family Majidae. New Zealand Journal of Marine and Freshwater Research 15: 331–383. Woods, C.M.S. 1993. Natural diet of the crab Notomithrax ursus (Brachyura: Majidae) at Oaro, South Island. New Zealand Journal of Marine and Freshwater Research 27: 309–315.

Hymenosomatidae MacLeay, 1838 Hymenosomatids look more like ‘spider crabs’ than majids, the family to which the name ‘spider crab’ is also applied. Davie (2002) referred to two species as ‘sea-spiders’, not to be confused with pycnogonids to which this popular name also applies. The name derives from their small round or triangular flat bodies and long spindly legs. Most hymenosomatids are small and inconspicuous, many rarely more than a centimetre across, but of course there are exceptions. The largest reach 26 mm but the family may include one of the smallest crabs known, Elamenopsis minima Lucas & Davie, at 1.7 mm across. Most species in southern Australia live in muddy environments, either muddy or poorly sorted sandy sediments or with the fine particles trapped among tufting reef algae, seagrasses, or sessile invertebrates. While hymenosomatids may be inconspicuous to the casual observer, some species are exceptionally abundant and species of Halicarcinus are the most common crabs seen in soft sediments and algal samples along the southern Australian coast. Worldwide almost half of the species known tolerate low salinity and at least eight species live in permanent fresh water (Lucas, 1980). One genus, Amarinus, contains the only freshwater crabs in southern Australia. There is evidence for symbiotic associations with echinoderms, and with abalone (Haliotis species, own observations of Elamena producta Kirk, 1879 in New Zealand). As with any small benthic invertebrate of soft sediments, hymenosomatids feature in the diet of demersal fishes. Definitive studies of taxonomy and biology of the family in New Zealand and Australia were by Melrose (1975) and Lucas (1980). The similarity between hymenosomatids and majids is more than accidental as there is compelling evidence from morphology that some majids, inachoidines in particular, may be more related to hymenosomatids than they are to other majids (Guinot & Richer de Forges, 1997). The structure of spermatozoa of two hymenosomatids studied is unusual and does not help resolve relationships (Richer de Forges et al., 1997). The family is confined to the Indo-West Pacific and subantarctic; no species are known from the Americas (southern tip excepted) or Atlantic Ocean. Its 90 species in 16 genera range from the deep sea (one species in this region), through the shelf and reefs of the subtidal, into the intertidal and estuaries. While hymenosomatids have zoeal larvae like other brachyurans the megalopa stage is skipped. Detritivory and predation are the favoured feeding strategies. Davie (2002) listed nine genera and 30 species described from Australia, of which about half occur in temperate waters. At least one undescribed species has been discovered recently (O’Hara, 2002). The key is based on that of Lucas (1980) to Australian species with additions and correc- tions to nomenclature.

Diagnosis. Cephalothorax thin, cuticle not heavily calcified, without hooked setae; dorsal carapace surface flat or nearly so, often with strong gastro-cardiac groove and contiguous grooves; without orbits or almost so, eyes exposed; antennular fossae poorly defined. Antennae fused with epistome (when present), article 2 slender. Maxillipeds 3 ischium well developed, palp articulating near anterolateral angle of merus. Chelipeds not longer than twice carapace width in females and most males, fingers not bent at an angle to palm. Male and female abdomens with not more than 5 somites, excluding telson. Male genital apertures sternal; female genital apertures on sternites of first walking legs. (after Davie, 2002)

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Key to southern Australian species of marine and freshwater Hymenosomatidae 1. Rostrum absent (rare species) ...... Halicarcinides nuytsi — Rostrum present ...... 2 2. Rostrum continuous with dorsal carapace surface, not separated by a ridge or groove; rostrum triangular or truncated; dorsal carapace surface without grooves or with only faint grooves ...... 3 — Rostrum separated from dorsal carapace surface by a ridge or groove; rostrum of various forms, often with several lobes; dorsal carapace surface usually with distinct transverse groove ...... 4 3. Carapace, with rostrum, approximately triangular ...... Trigonoplax longirostris — Carapace, with rostrum, approximately polygonal ...... Elamena abrolhensis 4. Epistome absent (maxillipeds 3 and antennae in contact, Fig. 121c) . . . Hymenosoma hodgkini — Epistome present (maxillipeds 3 and antennae separated by plate, Fig. 121b) ...... 5 5. Maxillipeds 3 narrow, not covering three-quarters of mouth-field . . Neorhynchoplax octagonalis — Maxillipeds 3 broad, covering most of mouth-field ...... 6 6. Rostrum trilobate (lateral lobes may be reduced or fused to narrow medial lobe); dactyli of walking legs with 1 or 2 rows of teeth ...... Halicarcinus ...7 — Rostrum a single broad lobe; dactyli of walking legs with a single inconspicuous subterminal tooth or lacking teeth entirely ...... Amarinus ...12 7. Medial rostral lobe much longer than lateral lobes (lateral lobes may be fused with medial lobe) ...... 8 — Medial rostral lobe about as long as lateral lobes ...... 10 8. Without an acute tooth or spine on lateral carapace walls at posterior lateral angle of carapace; lateral rostral lobes completely fused to medial lobe ...... 9 — With an acute tooth or spine on lateral carapace walls at posterior lateral angles of carapace (approximate to base of walking leg 1); lateral rostral lobes not fused to medial lobe ...... Halicarcinus rostratus 9. Carapace with circular lamelliform plate marked on anterior half; deep-sea ...... Halicarcinus lucasi — Carapace without circular lamelliform plate marked on anterior half; subtidal ...... Halicarcinus bedfordi 10. Without acute spine on lateral carapace walls or carapace rim at posterior lateral angle; rostral lobes arising well below carapace rim (New Zealand and Tasmania) ...... Halicarcinus innominatus — With acute spine on lateral carapace walls or carapace rim at posterior lateral angle (approximately dorsal to base of walking leg 1); rostral lobes arising at level of carapace rim or slightly below ...... 11 11. Median rostral slightly longer than lateral lobes, all lobes project straight forward; dactyli of walking legs with 2 rows of teeth so close together as to appear as a single row (widespread in southern Australia) ...... Halicarcinus ovatus — Median rostral slightly shorter than lateral lobes, lateral lobes slope down and out; dactyli of ...... walking legs with 2 distinctly separate rows of teeth (New Zealand and subantarctic islands) ...... Halicarcinus planatus 12. Mature females generally > 10 mm; antennal spines acute and prominent (ventral view of carapace) ; chelae of large males with a pulvinus (sac arising between bases of fingers) ...... Amarinus laevis

391 Marine Decapod Crustacea of Southern Australia

— Mature females generally < 10 mm; antennal spines small and obtuse (ventral view of carapace); chelae of large males without pulvinus ...... 13 13. Female genital apertures 0.4–0.5 mm wide; telson approximately 0.35 of total abdomen length in mature females ...... Amarinus paralacustris — Female genital apertures 0.65–0.8 mm wide; telson approximately 0.25 of total abdomen length in mature females ...... Amarinus lacustris

bc d e

Fig. 119. Hymenosomatidae. a, Halicarcinus ovatus. b, Amarinus lacustris. c, Amarinus laevis. d, Amarinus paralacustris. e, Elamena abrolhensis.

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Amarinus Lucas, 1980 Species of Amarinus are confined to estuarine and freshwater environments of islands in the south-western Pacific and Australia. The broad rostrum is distinctive. Remnants of uropods inserted laterally between the last abdominal somite and the telson of males (also present in Hymenosoma hodgkini) is an unusual brachyuran feature seen elsewhere only in dromiacean families. Diagnosis. Carapace circular to oval, wider than long; surface flat or slightly concave with well defined grooves; rostrum spade-like, deflexed, concave laterally, extending over eyestalks, separated from dorsal surface by ridge or groove or both. Maxillipeds 3 slightly separated medially. Chelipeds stouter than walking legs, especially in male. Amarinus lacustris (Chilton, 1882) fresh-water spider crab (Fig. 119b). Antennal spines small and obtuse (ventral view of carapace); chelae of large males without a pulvinus. Female genital apertures 0.65–0.8 mm wide, without curved medial process. Telson approximately 0.25 of abdomen length in mature females. 10 mm. North I., New Zealand; Norfolk I., Lord Howe I., Vic., Tas., SA; freshwater and salt lakes, lagoons. Amarinus lacustris is a widespread inland brackish water species whose distribution raises questions about how this came to be (Lucas, 1970; McLay, 1988). It occurs as far inland as Lake Colac in Victoria and at salinities as high as 9.6 parts per thousand. It does occur but less frequently in fresh water. Its preferred habitats are lakes and fast-flowing streams where it inhabits vegetation in muddy sediments (Walker, 1969). There is no free larval stage. This species and the similar Australian species, A. paralacustris, can only be distinguished as mature females. Amarinus laevis (Targioni Tozzetti, 1877) (Figs 119c, 121b, Pl. 21f). Antennal spines acute and prominent (ventral view of carapace); chelae of large males with a pulvinus (a sac arising between bases of fingers). 22 mm. Qld, NSW,Vic., Tas., SA, southern WA; estuaries and brackish water (Lucas & Hodgkin, 1970a, b). Amarinus paralacustris (Lucas, 1970) (Fig. 119d). Antennal spines small and obtuse (ventral view of carapace); chelae of large males without a pulvinus. Female genital apertures 0.4–0.5 mm wide, with curved medial process. Telson approximately 0.35 of abdomen length in mature females. 10 mm. Qld, NSW, to central Vic.; estuaries and brackish water.

Elamena Milne-Edwards, 1837 The angular rostrum characterises the dozen or so species of Elamena, all found in the Indo-West Pacific. Diagnosis. Carapace plus rostrum polygonal to pear-shaped; dorsal surface without well-defined grooves; rostrum truncate, continuous with carapace surface, concealing antennae and most of eyes; epistome as long as broad; pterygostomial regions with prominent lobules. Maxillipeds 3 broad, meeting medially. Chelipeds stouter than walking legs, especially in male. Elamena abrolhensis Gordon, 1940 (Fig. 119e). Carapace wider than long; rostrum truncate, with deep rostral keel continuous with interantennal septum; postocular lobe with small tooth; posterolateral angles pointed and obtuse; male pleopod 1 S-shaped, with 12 subterminal setae. 8.5 mm. SA (W of Port Noarlunga), WA (N to Abrolhos Is); subtidal, rocky reef. Earlier texts have erroneously referred this species to E. truncata (Stimpson, 1858) (Lucas, 1980).

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Halicarcinides Lucas, 1980 This monotypic genus is the only one in Australia without a rostrum. Diagnosis. Carapace subcircular, dorsal carapace surface with sharply defined grooves; rostrum absent; epistome present; eyes completely concealed in dorsal view. Maxillipeds 3 broad, meeting in midline. Maxilliped 3 broad, almost meeting medially. Chelipeds stouter than walking legs, especially in male. Halicarcinides nuytsi (Hale, 1927) beakless sea spider (Fig. 120a). NSW (Port Jackson), SA (St Francis I.); subtidal reef, rare (known from only two localities but not collected since 1930).

Halicarcinus White, 1846 Halicarcinus is the largest and most frequently encountered genus in Australia. Two species, H. ovatus and H. rostratus are abundant in surveys of soft sediments of bays and the shelf in south- eastern Australia and H. ovatus is also common in algal samples from rocky reefs. The genus is represented throughout the western Pacific as far south as the subantarctic islands. Diagnosis. Carapace subcircular; surface usually with distinct H-shaped pattern of grooves; postero- and anterolateral angles often with acute spine; rostrum trilobate (lateral lobes may be reduced), separated from dorsal carapace surface by ridge or groove; epistome broad. Maxilliped 3 broad. Walking legs with dactyli having 1 or 2 rows of teeth. Halicarcinus bedfordi Montgomery, 1931 (Fig. 120b). Rostrum approximately triangular, narrowing abruptly halfway or further along its length, rounded apex; lateral lobes fused with medial lobe. Posterolateral angle of carapace without spine. 7 mm. WA (S to Swan River but rare), NT, Qld (S to Moreton Bay); estuarine. Halicarcinus innominatus Richardson, 1949 (Fig. 120c). Rostrum trilobed, lobes short, subequal, close together, apically rounded, arising well below carapace rim. Postero- and anterolateral angles obtuse; lateral carapace wall without acute spine. Walking legs with dactyli having 1 row of teeth. 19 mm. New Zealand, eastern Tas.; intertidal, often associated with mussels. This is almost certainly a species introduced to Tasmania from New Zealand, probably with oyster shipments as early as the end of the nineteenth century. The introduced species is very similar to H. ovatus with which it co-occurs but can be distinguished by the absence of a spine on the carapace and subtle differences in the rostral lobes. Halicarcinus lucasi Richer de Forges, 1993. Rostrum simple, long, curved back towards top; carapace without anterolateral tooth, with round lamelliform plate between gastric and cardiac zones. 2 mm. NSW (Taupo Seamount, Tasman Sea); 244 m depth. This exceptionally small species is known only from slope depths (Richer de Forges, 1993); all other Australian species are intertidal or shallow shelf inhabitants. Halicarcinus ovatus Stimpson, 1858 three-pronged sea spider (Fig. 119a, Pl. 21g). Rostrum trilobate, medial lobe slightly longer than lateral lobes, arising level with or below carapace rim. Postero- and anterolateral angles obtuse; lateral carapace wall with acute spine. Walking legs with dactyli having 2 rows of teeth. Variable colouration in one population: white, orange, black, mottled. 10 mm. NSW, Vic., Tas., SA, WA (N to Geraldton); intertidal to 85 m depth. This is one of the most common crabs in algal samples, in Victoria at least, but its small size and cryptic behaviour makes it inconspicuous except after careful sample sorting. It co-occurs with the similar H. innominatus in algal habitats in Tasmania and with H. rostratus in soft sediments in Victoria. Lucas (1971) reported three different kinds of larvae from different populations in south-western Australia suggesting that more than one species are involved.

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Halicarcinus planatus (Fabricius, 1775) (Fig. 121d). Rostrum trilobate, lateral lobes slightly longer than medial lobe, arising at level of carapace rim or slightly below. Lateral carapace wall with acute spine. Walking legs with dactyli having 2 rows of teeth. 19 mm. New Zealand, southern South America, Macquarie I. and most other subantarctic islands; intertidal–subtidal, rocky shores. This is the only species of intertidal crab found in the Australian subantarctic. It does not occur on the Australian mainland or Tasmania. Halicarcinus rostratus (Haswell, 1881) beaked sea spider (Fig. 120d). Rostrum trilobate, lateral lobes reduced, fused with medial lobe, medial lobe longer than eyes, tapering, arising below level of carapace. Postocular teeth conspicuous. Lateral carapace wall with an acute spine Walking legs with dactyli having 2 irregular rows of teeth. 5.5 mm. Central NSW, Vic., Tas., SA, WA (N to Swan R.); uncommon intertidally but frequently encountered in soft benthic samples, 2–130 m depth.

Hymenosoma Desmarest, 1825 The absence of an epistome (Fig. 121c or a minute epistome) and broad rostrum distinguish this genus in Australia. Diagnosis. Carapace oval, surface with well defined or very weak grooves; rostrum triangular with rounded apex, concave laterally, deflexed, shorter than eyes, separated from dorsal carapace surface by a groove; epistome absent or reduced. Maxillipeds 3 not meeting medially. Chelipeds stouter than walking legs, especially in male. Hymenosoma hodgkini Lucas, 1980 (Fig. 121c, f). Carapace with well-defined grooves; rostrum much shorter than eyes. Walking legs dactyli with row of fine teeth. Abdominal somite 3 of mature females with 2 large bulges projecting posteriorly. 7.5 mm. Qld, NSW, Vic. (W to Western Port); intertidal to 9 m, estuaries and marine bays, muddy sediments.

Neorhynchoplax Sakai, 1938 Ng & Chuang (1996) reviewed this genus, recognisable usually by the combination of trilobate rostrum, narrow maxilliped 3, toothed dactyli and tooth on the posterolateral angle of the carapace. Females carry the eggs freely in a brood chamber formed by the abdomen, not attached to pleopods. Diagnosis. Carapace circular or octagonal, surface with well defined grooves; rostrum trilobate, separated from dorsal carapace surface by a groove; epistome absent. Maxillipeds 3 narrow, not meeting medially. Chelipeds stouter than walking legs, especially in male; walking legs slender, not distinctly flattened. Abdomen without lobes on posterior margin of somite 1; male somites 3–4, 3–5 or 4–5 fused. Neorhynchoplax octagonalis (Kemp, 1917) (Fig. 120f). Posterolateral angle of carapace with spine; anterolateral border without teeth. Walking legs having dactyli with subterminal tooth only. Male abdomen with somites 3–5 fused. 3.4 mm. India, Qld, NSW (Port Hacking); intertidal, subtidal, muddy habitats. Lucas & Davie (1982) referred to this species as a member of the genus Elamenopsis. This is the first record of the species in NSW. There is some doubt that the Australian population belongs to this species, first described from India.

Trigonoplax Milne Edwards, 1853 This western Pacific genus is represented in Australia by two species, one tropical and one temperate. Its triangular carapace is highly diagnostic.

395 Marine Decapod Crustacea of Southern Australia

ab cd

Fig. 120. Hymenosomatidae. a, Halicarcinides nuytsi. b, Halicarcinus bedfordi. c, Halicarcinus innominatus. d, Halicarcinus rostratus. e, Hymenosoma hodgkini. f, Neorhynchoplax octagonalis.

b c

Fig. 121. Hymenosomatidae. a, Trigonoplax longirostris. b, Amarinus laevis (anterior region, anteroventral view). c, Hymenosoma hodgkini (anterior region, anteroventral view). d, Halicarcinus planatus (anterior region, dorsal). e, Amarinus laevis (cheliped). f, Hymenosoma hodgkini (dactylus of walking leg 4).

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Diagnosis. Carapace triangular, anterolateral angles almost absent, surface without defined grooves; rostrum triangular, not separated from dorsal carapace surface by a groove, concealing antennae and most of eyes, with keel; epistome long and bulging. Maxillipeds 3 meeting medially. Chelipeds similar to walking legs in both sexes. Trigonoplax longirostris McCulloch, 1908 (Fig. 121a, Pl. 21h). Carapace as long as wide, carapace walls strongly converging ventrally; rostrum deflexed, with shallow rostral keel. Chelipeds with weakly spatulate fingers. Walking legs with dactyli widest at midpoint. Vic. (Western Port), SA, WA (N to Mandurah); shallow reef.

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Guinot, D., & Richer de Forges, B. 1997. Affinités entre les Hymenosomatidae MacLeay, 1838 et les Inachoididae Dana, 1851 (Crustacea, Decapoda, Brachyura). Zoosystema 19: 453–502. Lucas, J.S. 1970. Breeding experiments to distinguish two sibling species of Halicarcinus (Crustacea, Brachyura). Journal of Zoology, London 160: 267–278. Lucas, J.S. 1971. The larval stages of some Australian species of Halicarcinus (Crustacea, Brachyura, Hymenosomatidae) I. Morphology. Bulletin of Marine Science 21: 471–490. Lucas, J.S. 1980. Spider crabs of the family Hymenosomatidae (Crustacea; Brachyura) with particular reference to Australian species: systematics and biology. Records of the Australian Museum 33: 148–247. Lucas, J.S., & Davie, P.J.F. 1982. Hymenosomatid crabs of Queensland estuaries and tidal mud flats, including descriptions of four new species of Elamenopsis A. Milne Edwards and a new species of Amarinus Lucas. Memoirs of the Queensland Museum 20: 401–419. Lucas, J.S., & Hodgkin, E.P. 1970a. Growth and reproduction of Halicarcinus australis (Haswell) (Crustacea, Brachyura) in the Swan Estuary, Western Australia. 1. Crab instars. Australian Journal of Marine and Freshwater Research 21: 149–162. Lucas, J.S., & Hodgkin, E.P. 1970b. Growth and reproduction of Halicarcinus australis (Haswell) (Crustacea, Brachyura) in the Swan Estuary, Western Australia. II. Larval stages. Australian Journal of Marine and Freshwater Research 21: 163–173. McLay, C.L. 1988. Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22: 1–403. Melrose, M.J. 1975. The marine fauna of New Zealand: family Hymenosomatidae (Crustacea, Decapoda, Brachyura). New Zealand Oceanographic Institute Memoir 34: 1–123. Ng, P.K.L.,& Chuang, C.T.N. 1996. The Hymenosomatidae (Crustacea: Decapoda: Brachyura) of southeast Asia, with notes on other species. Raffles Bulletin of Zoology, Supplement 3: 1–82. O’Hara, T. 2002. Endemism, rarity and vulnerability of marine species along a temperate coastline. Invertebrate Systematics 16: 671–684. Richer de Forges, B. 1993. Deep sea crabs of the Tasman seamounts (Crustacea: Decapoda: Brachyura). Records of the Australian Museum 45: 11–24. Richer de Forges, B., Jamieson, B.G.M., Guinot, D., & Tudge, C.C. 1997. Ultrastructure of the spermatozoa of Hymenosomatidae (Crustacea: Brachyura) and the relationships of the family. Marine Biology 130: 233–242. Walker, K. 1969. The ecology and distribution of Halicarcinus lacustris (Brachyura: Hymenosomatidae) in Australian inland waters. Australian Journal of Marine and Freshwater Research 20: 163–173.

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Parthenopidae Macleay, 1838 Parthenopids are among the most ornamented crabs, often with highly sculptured triangular carapace with sharp edges. The family is best represented in the tropics but even there are rarely seen because of their cryptic habits. They live on coral reefs and shelly benthic environments. Only one definitely, possibly three, of almost 30 Australian species occurs along southern coasts (Davie, 2002). Some leucosiids are ornamented and might superficially be confused with parthenopids. The family has a complex taxonomic history, with several subfamilies, some elevated to family rank from time to time, and sometimes thought not to represent a monophyletic clade (see discussion by Davie, 2002). This key to species is highly artificial. Diagnosis. Carapace variable; regions usually demarcated with prominent grooves, dorsal surface usually tuberculate, granulate to eroded; branchial regions usually swollen and distinctly separated from swollen gastric and cardiac regions; front narrow, simple or trilobed. Eyes small, retracting into small, circular orbits (rare exceptions). Chelipeds robust, often longer than walking legs; merus and chela often prismatic in cross-section. Male abdominal somites 3–5 fused, although sutures may be visible. Male gonopod 1 stout, simple, sometimes with strong spines; gonopod 2 usually short but sometimes slightly longer than first. Male genital openings coxal; female genital openings sternal. (adapted from Davie, 2002)

Key to southern Australian species of Parthenopidae 1. Carapace strongly laterally produced, concealing walking legs . . . . Cryptopodia spatulifrons — Carapace not laterally produced; walking legs visible from above ...... 2 2. Chela palm with row of conical spikes on outer margin; rostrum triangular ...... Rhinolambrus contrarius — Chela palm unarmed on outer margin; rostrum apically emarginate . Thyrolambrus excavatus

Cryptopodia Milne Edwards, 1834 Davie & Turner (1995) provided a key to Australian species, six out of 15 known in the Indo-West Pacific. The only southern species may be a doubtful record for this essentially tropical genus revised by Chiong & Ng (1998). Diagnosis. Carapace triangular, with extensive lateral wings completely covering short walking legs in ventral cavity, with deep triangular depression in middle; rostrum triangular and horizontal; pterygostomial region smooth. Cheliped merus with wing-like expansion distally. (after Chiong & Ng, 1998) Cryptopodia spatulifrons Miers, 1879 (Fig. 122c). Carapace granulated and pitted, rostrum triangular, with convex crenulate margins. Maxilliped 3 not swollen, external surface heavily pitted. 50 mm. Northern WA, northern Qld, possibly from NSW; rare, to 13 m depth (Chiong & Ng, 1994; Davie & Turner, 1995).

Rhinolambrus Milne Edwards, 1879 This is essentially an Indo-West Pacific genus whose several species of grotesque crabs are recognised by their elongate ornamented chelipeds and carapace constriction behind the eyes (Flipse, 1930). Diagnosis. Carapace triangular globular, rough or spiky, without extensive lateral wings; rostrum triangular. Cheliped elongate, spinose or tuberculate. Walking legs slender, visible in dorsal view.

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Fig. 122. Parthenopidae. a, Thyrolambrus excavatus. b, Rhinolambrus contrarius. c, Cryptopodia spatulifrons.

Rhinolambrus contrarius (Herbst, 1796) (Fig. 122b). Carapace including rostrum longer than broad; rostrum longer than wide, laterally spinose, flat. Carapace and chelipeds covered with rough tubercles and spikes. 55 mm. Indo-West Pacific (India to Japan and Hawaii), Qld, NSW; 10–30 m depth, rocky benthos.

Thyrolambrus Rathbun, 1894 Flipse (1930) listed four species from the Indo-West Pacific. Diagnosis. Carapace globular, as wide as long, granulated and spiny, with postocular constriction, without extensive lateral wings; rostrum broad, emarginate. Cheliped rough, merus stout. Walking legs compact, visible in dorsal view. Thyrolambrus excavatus Baker, 1905 eroded crab (Fig. 122a). 19 mm. SA; to 50 m depth. This, an endemic species, may in truth be the only parthenopid in the region.

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References Chiong, W.L., & Ng, P.K.L. 1994. The identity of Cryptopodia spatulifrons Miers, 1879, and description of a new species, Cryptopodia fistulosa (Crustacea: Decapoda: Brachyura: Parthenopidae). Raffles Bulletin of Zoology 42: 949–959. Chiong, W.L., & Ng, P.K.L. 1998. A revision of the buckler crabs of the genus Cryptopodia H. Milne Edwards, 1834 (Crustacea: Decapoda: Brachyura: Parthenopidae). Raffles Bulletin of Zoology 46: 157–216. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Davie, P.J.F., & Turner, P.A. 1995. New records of Cryptopodia (Crustacea: Decapoda: Parthenopidae) from Australia. Memoirs of the Queensland Museum 38: 447–462. Flipse, H.J. 1930. Decapoda F. (Decapoda Brachyura continued) Oxyrhyncha: Parthenopidae. Siboga Expéditie Monographie 39c: 1–96.

Atelecyclidae Ortmann, 1893 Atelecyclids are rare deep-water crabs, usually with a longitudinally oval or circular carapace with an uneven row of triangular teeth along the anterior margin. Ng (1993) and Salva & Feldmann (2001) treated the one-time atelecyclid subfamily, Thiinae, as a distinct family confined to the Atlantic Ocean, and excluded other genera. Salva & Feldmann (2001) used morphological characters in a cladistic analysis to illuminate relationships between the five atelecyclid and two thiid genera and four genera of Belliidae. Two genera occur in Australia, both with one described species restricted to shelf or deep water, but only one has been found in south-eastern Australia, none in the south-west. Diagnosis. Carapace usually rounded to longitudinally ovate or pentagonal; regions moderately well defined; anterolateral margins usually armed with teeth or spines; anterolateral and posterolateral margins with dense setae; front with 2–4 teeth or lobes; orbits complete. Antennular fossae quadrate to longer than broad, antennules fold longitudinally or almost so. Antennae well developed, short, basal article either fixed or only slightly moveable; flagellum distinctly but not strongly setose. Walking leg 4 shorter but similar to other legs. (adapted from Davie, 2002)

Trichopeltarion Milne Edwards, 1880 The oval shape, trifid rostrum and five more or less complex spines along the anterolateral margin immediately differentiates these species from any other at shelf or slope depths (Dell, 1968). Guinot (1989) listed the four species then known and discussed the close relationship of the genus to other atelecyclids. Diagnosis. Carapace oval, dorsal surface tuberculate; lateral margins armed with spines. Basal antennal article without a projection filling the orbital hiatus. Front 3–spined. Trichopeltarion wardi Dell, 1968 (Fig. 123a). Rostral spines similar, narrow; 5 more or less lateral spines triangular, complex; carapace spines complex, all over surface. 26 mm. Eastern Vic., Tas.; 73–510 m depth. Trichopeltarion sp. (Fig. 123b, Pl. 22a). Medial rostral spines broader than lateral spines, rounded; anterolateral margin with sharp spines more or less clustered on 5 lobes; carapace with few short spines on anterior half. 30 mm. Tasmanian Seamounts (44°S, 147°E), 900–1700 m depth. The undescribed species from seamounts of south-eastern Tasmania has been trawled in large numbers (Poore et al., 1998).

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Fig. 123. Atelecyclidae. a, Trichopeltarion wardi. b, Trichopeltarion sp.

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Dell, R.K. 1968. A new crab of the genus Trichopeltarion from Australia. Australian Zoologist 14: 275–276, pl. 16. Guinot, D. 1989. Les genres Trachycarcinus Faxon et Trichopeltarion A. Milne Edwards (Crustacea, Brachyura: Atelecyclidae). In: Forest, J. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144: 347–385. Ng, P.K.L. 1993. Krausiinae, a new subfamily for the genera Kraussia Dana, 1852, Palpedia, new genus, and Garthasia new genus (Crustacea: Decapoda: Brachyura: Xanthidae), with descriptions of two new species from Singapore and the Philippines. Raffles Bulletin of Zoology 41: 133–157. Poore, G.C.B., Hart, S., Taylor, J., & Tudge, C. 1998. Decapod crustaceans from Tasmanian seamounts. Pp. 65–78 in: Koslow, J.A., & Gowlett-Holmes, K. (eds), The seamount fauna of southern Tasmania: benthic communities, their conservation and impacts of trawling. Final report to Environment Australia and The Fisheries Research Development Corporation. CSIRO Marine Research: Hobart. Salva, E.W., & Feldmann, R.M. 2001. Reevaluation of the family Atelecyclidae (Decapoda: Brachyura). Kirtlandia 52: 9–62.

Cancridae Latreille, 1803 Until recently all species were placed in one genus, Cancer Linnaeus. The European edible crab, Cancer pagurus, is a member of this family and is still frequently seen in the European and British markets. The genus name Cancer was used for most species of crabs by Linnaeus and others early in the history of crustacean taxonomy. Nations (1975, 1979) organised the species of a more restricted Cancer into subgenera elevated to genera by Schweitzer & Feldmann (2000). Only two species occur in Australia (Davie, 2002) but only one naturally. On the basis of the chronology of abundant fossil records, Schweitzer & Feldmann (2000) believed that the Cancrinae, the subfamily to which the Australian species belong, evolved in the Southern Hemisphere and subsequently dispersed northward along the coast into the North Pacific Ocean. From this area of origin almost all cancrine genera dispersed.

Diagnosis. Carapace broadly oval; anterolateral margins strongly convex, dentate; posterolateral margins short; front narrow, usually with teeth each side of median tooth which may be displaced ventrally; orbits small. Antennules folding longitudinally or a little obliquely; basal antennal arti-

401 Marine Decapod Crustacea of Southern Australia

cle longer than broad, flagellum short, setose. Epistome narrow. Maxillipeds 3 may overlap endostome. Chelipeds massive, subequal, similar in both sexes. Walking legs laterally compressed to more or less cylindrical; dactyli styliform, Male abdomen with somites 3–5 fused. Sternum narrow; female genital openings sternal; male openings coxal. Male gonopod 1 stout, straight, tapered; gonopod 2 slender, and of similar length. (adapted from Davie, 2002)

Key to Australian species of Cancridae 1. Carapace regions separated by weak grooves or indistinct; carapace region ornamentation not densely granular; low intertidal to subtidal ...... Metacarcinus novaezelandiae — Carapace regions separated by very deep, narrow, smooth grooves; surface of regions densely granular; deep-water ...... Platepistoma nanum

Metacarcinus Milne Edwards, 1862 The genus inhabits most of the Pacific and the North Atlantic Oceans. Only one species occurs in Australia, and that is introduced from New Zealand. Diagnosis. Carapace ovate, about two-thirds as wide as long. Front with 5 spines including inner- orbital spine, inner 3 spines closely spaced; front usually not produced beyond orbits. Fronto- orbital width about 0.26–0.34 maximum carapace width; orbits shallow, directed forward. Anterolateral margin with 9 or 10 variable spines. Posterolateral margins rimmed, sometimes with 1 spine; carapace regions poorly developed, smooth or ornamented with fine granules. Palm of chelipeds usually with 4 granular or smooth ridges on outer surface; fixed finger with upper and lower ridges; cutting edges of fingers with teeth. Metacarcinus novaezelandiae (Jacquinot, 1846) pie-crust crab (Fig. 124a, Pl. 22b). 150 mm. New Zealand, introduced to eastern Tas., Vic. (Flinders Reef, Gippsland Lakes, unconfirmed old record from Port Phillip Bay), NSW (Eden); low intertidal to subtidal. All recent literature refers to this species as Cancer novaezelandiae. The species was first recorded in Hobart harbour in about 1930 and is now common in eastern Tasmania. The species was recorded in Port Phillip Bay about the same time (McNeill & Ward, 1930) but has never been seen there since. An individual was collected by a fisher in the Gippsland Lakes in 1991, reporting the species because it had ‘never been seen before.’ A local population has resided on Flinders Reef, Vic., since at least 1995 but there are few records elsewhere in Victoria. Adult moulting and mating probably occur late in the year and eggs are probably carried for five or six months (McLay, 1988). The hatchling passes through five zoeal stages and a megalopa before settling (Wear & Fielder, 1985). Maturity is probably reached after one year but maximum size not until the age of 4 or 5 years. The adult crab lives often buried in sand and feeds at night, mostly on bivalve molluscs but will take other prey (McLay, 1988).

Platepistoma Rathbun, 1906 Davie (1991) discussed the genus, which is relatively well defined by its strong carapace regions, and differentiated it from Cancer. Platepistoma is known only from the Indo-Pacific region where its species are distinguished by subtle sculptural differences. Diagnosis. Carapace oval, three-quarters as long as wide. Front with 5 poorly developed, short, spines including inner-orbital spines. Fronto-orbital width 0.38–0.44 maximum carapace width; orbits shallow, very small, directed forward, with 2 fissures. Anterolateral margin with 9 or 10 small spines. Posterolateral margin rimmed, nearly straight, entire or with 1 spine; carapace regions well defined by deeply excavated, smooth grooves, regions with dense tubercles. Palm of

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Fig. 124. Cancridae. a, Metacarcinus novaezelandiae. b, Platepistoma nanum.

chelipeds, outer surface with 4 or 5 rows of granules; fixed finger with 2 rows of granules, cutting edges with blunt denticles or nearly smooth. Platepistoma nanum Davie, 1991 (Fig. 124b). 29 mm. Western Tasman Sea (Norfolk Ridge and Britannia Seamount), Qld, Vic., Tas.; 425–560 m depth on seamounts. The species is smaller

403 Marine Decapod Crustacea of Southern Australia

than most others in the genus and confined to seamounts throughout the Tasman Sea (Davie, 1991). Jamieson et al. (1997) used a description of the spermatozoa to illuminate relationships of the genus.

References Davie, P.J.F. 1991. Crustacea Decapoda: the genus Platepistoma Rathbun, 1906 (Cancridae) with the description of three new species. In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 9. Mémoires du Muséum National d’Histoire Naturelle, Paris 152: 493–514. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Jamieson, B.G.M., Guinot, D., Tudge, C.C., & Richer de Forges, B. 1997. Ultrastructure of the spermatozoa of Corystes cassivelaunus (Corystidae), Platepistoma nanum (Cancridae) and Cancer pagurus (Cancridae) supports recognition of the Corystoidea (Crustacea, Brachyura, Heterotremata). Helgoländer Meeresuntersuchungen 51: 83–93. McLay, C.L. 1988. Brachyura and crab-like Anomura of New Zealand [Crabs of New Zealand]. Leigh Laboratory Bulletin 22: 1–403. McNeill, F.A., & Ward, M. 1930. Carcinological notes. No. 1. Records of the Australian Museum 17: 357–383. Nations, D. 1975. The genus Cancer (Crustacea: Brachyura): systematics, biogeography and the fossil record. Natural History museum of Los Angeles County, Science Bulletin 23: 1–104. Nations, D. 1979. The genus Cancer and its distribution in time and space. Bulletin of the Biological Society of Washington 3: 153–187. Schweitzer, C.E., & Feldmann, R.M. 2000. Re-evaluation of the Cancridae Latreille, 1802 (Decapoda: Brachyura) including three new genera and three new species. Contributions to Zoology 69: 223–250. Wear, R.G., & Fielder, D.R. 1985. The marine fauna of New Zealand: larvae of the Brachyura (Crustacea, Decapoda). New Zealand Oceanographic Institute Memoir 92: 1–90.

Corystidae Samouelle, 1819 Of three genera, two, Gomeza Gray and Jonas Jacquinot & Lucas occur in Australia (Sakai, 1976). Both inhabit soft sediments but are rarely caught. The produced front and longer-than-wide body are characteristic. Diagnosis. Carapace longitudinally oval; dorsal surface convex from side to side, regions ill- defined; lateral margins of carapace convex, often armed with teeth or spines along entire length, or teeth confined to anterior half; front with 2 or 3 teeth or lobes; orbits incomplete. Antennules fold longitudinally. Antennal flagellum often very long and setose; flagella interlock to form respiratory tube. Epistome absent. Buccal cavern elongate. Thoracic sternum narrow, elongate, with sutures complete and parallel. Maxillipeds 3 elongate, subpediform, extending almost to antennules. Chelipeds strongly sexually dimorphic. Walking legs 5 with dactylus sometimes broadened for swimming. Male abdomen of 5 somites; male and female abdomens with basal somites visible dorsally; female abdomen not covering genital openings. (adapted from Davie, 2002)

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Fig. 125. Corystidae. Gomeza bicornis.

Gomeza Gray, 1831 Of two species (Sakai, 1976) one occurs in Australia. Diagnosis. Carapace longitudinally oval, convex on anterior third, rostrum bifid, orbits small, lateral borders armed with 10 tubercles. Antenna flagellum much more than half length of carapace. Last walking leg with styliform dactylus. Gomeza bicornis Gray, 1831 masked burrowing crab (Fig. 125). 21 mm. Indo-West Pacific, Japan to Sri Lanka; Qld, WA, SA Gulfs; coral reefs, burrowing in sandy bottoms; rare. The oval shape and long antennae distinguish this species from most other crabs.

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Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates.

Geryonidae Colosi, 1923 Geryonids are large deep-water crabs, some of which are the basis of small fisheries in many parts of the world and so are the attention of some published biological studies (e.g., Attrill et al., 1991) and numerous reports. The only Australian species is the basis of a small fishery in Western Australia. According to Manning & Holthuis (1989) there are three genera and 24 species, but only one in Australia. Geryonids belong to a group of families of crabs that carcinologists have found difficult to distinguish. For example, Sakai’s (1976) key to families relies on a triplet in which Geryonidae are said to be an ‘intermediate family between Xanthidae and Goneplacidae.’ Luckily, crab taxonomy has moved on from here but these families still remain problematic.

Diagnosis. Carapace hexagonal, smooth to granular, with 3–5 weak anterolateral teeth on each side; suborbital margin with inner tooth only. Chelipeds unequal, merus with subdistal spine on upper margin. Walking legs with naked dactyli, T-shaped in cross-section. Abdomen with 7 segments apparent in both sexes, but 3–5 fused in male.

Chaceon Manning & Holthuis, 1989 With 21 species worldwide, the genus dominates the family. For a key to the Pacific species see Ng & Manning (1998). Diagnosis. Carapace with 5 anterolateral teeth on each side, second and fourth weak; almost as long as wide; branchial regions not markedly inflated; frontal teeth well developed; orbits shallow, rounded. Chaceon bicolor Manning & Holthuis, 1989 Pacific golden crab (Fig. 126). Anterior carapace purple, branchial regions tan, legs yellowish. 165 mm. Japan, West Pacific, New Caledonia, WA, Qld, NSW, Tas.; 275–1620 m depth. The species was widely reported as Geryon affinis. Davie (2002) believed the WA population belongs to another species, as yet undescribed. The only other large deep-water crab in the region is the eriphiid, Pseudocarcinus gigas, but this is wider than long rather than longer than wide.

Fig. 126. Geryonidae. Chaceon bicolor.

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References Attrill, M.J., Hartnoll, R.G., & Rice, A.L. 1991. Aspects of the biology of the deep-sea crab Geryon trispinosus from the Porcupine Seabight. Journal of the Marine Biological Association of the United Kingdom 71: 311–328. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Manning, R.B., & Holthuis, L.B. 1989. Two new genera and nine new species of geryonid crabs (Crustacea, Decapoda, Geryonidae). Proceedings of the Biological Society of Washington 102: 50–77. Ng, P.K.L., & Manning, R.B. 1998. A new deep-water crab from Belau, Micronesia, with a key to the Pacific species of Chaceon (Crustacea: Decapoda: Brachyura: Geryonidae). Proceedings of the Biological Society of Washington 111: 389–397. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates.

Portunidae Rafinesque-Schmaltz, 1815 Portunids are better known as swimming crabs. Most spend much of their time burrowed into sandy or muddy sediments of many diverse habitats. Many species are efficient swimmers, using the flattened last pair of legs as paddles. These swimming legs and the pointed toothed and spined claws enable them to be highly active and aggressive predators. Portunids may be confused with spanner crabs (Raninidae) and moon crabs (Calappidae, Matutidae) which have most of the legs flattened for swimming – only the last legs are paddle-like in portunids. Larger species of portunids (e.g. Ovalipes australiensis, Scylla serrata, Portunus pelagicus, P. sanguinolentus and Charybdis feriata) contribute to a tropical and subtropical Australian crab fishing industry as they do in other countries (Ng, 1998). Mud crabs (S. serrata) are sold live throughout Australia. Other species are sold fresh or cooked. The Portunidae are predominantly a tropical Indo-West Pacific family but representatives occur in all seas. In Australia, there are some 20 genera and 100 species; of the 43 species which have been included here, fewer than a dozen are common in southern Australian waters. The remainder are tropical species which are collected occasionally as far south as Sydney or Perth. Seven subfamilies are recognised, of which six occur in southern Australia. Stephenson (1972) provided keys to most Australian portunids that can be used to identify material from northern Australia or material which does not fit the diagnoses given here. This paper summarised numerous earlier papers by the same author whose taxonomic studies established our knowledge of the Australian fauna. Also relevant today, but more for northern Australian taxa, is the review of Apel & Spiridonov (1998). The terminology used in the past to describe portunids is inconsistent. Here, the number of frontal lobes or teeth of the carapace excludes the inner supraorbital lobes or teeth (unless otherwise stated). Its lobes or teeth are referred to as median, submedian and lateral frontal (in the case of six lobes). The orbit is defined mesially by the supraorbital tooth, laterally by the first anterolateral tooth but there may be a tooth in between. The number of anterolateral teeth includes the first adjacent to the orbit, pilose describes a dense covering of short hairs, easily rubbed off with one’s fingers, and pubescent or hairy describes relatively longer hairs which are less easily scraped off. The last leg has been referred to as the natatory (swimming) leg or leg 5 (if the cheliped is counted) – here it is referred to as leg 4 as in other crabs. The family includes the introduced species, Carcinus maenas. It has one of the least widened paddles of all portunids and may be confused with grapsid shore crabs whose habitat it shares. Another introduced species, Charybdis japonica, has been recorded only once.

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Diagnosis. Carapace typically subhexagonal or transversely oval, flat, regions not well defined, usually wider than long, and widest between last pair of anterolateral teeth; 1–9 pairs of anterolateral teeth; front horizontal, wide, lobulate or dentate. Eyes typically short, well protected by sunken rounded orbits; rarely greatly lengthened and largely unprotected. Lobe on endite of maxilliped 1 (‘portunid lobe’) usually present. Chelipeds typically robust; fingers usually sharply pointed, occasionally spoon-shaped (no southern Australian portunids have this character). Last pair of legs generally modified for swimming, with propodus and dactylus paddle-like; dactylus rarely styliform. Abdominal somites 3–5 typically fused in males; male genital openings coxal; female openings sternal. (after Davie, 2002)

Key to subfamilies of Portunidae in southern Australia 1. Chelipeds longer than all walking legs ...... 2 — One or more pairs of walking legs same length or longer than chelipeds ...... 4 2. Orbits and eyestalks extremely long ...... Podophthalminae ...p.412 — Orbits and eyestalks short ...... 3 3. Male first gonopod without subterminal spines ...... Portuninae ...p.418 — Male first gonopod with subterminal spines ...... Thalamitinae ...p.422 4. Last legs paddle-shaped, dactylus distally rounded or acute ...... 5 — Last legs typically more leg-like than paddle-like; if swimming paddles, dactylus distally acute ...... Carcininae . ..p.409 5. Flagellum of antenna included in orbit; carapace moderately wider than long ...... Polybiinae ...p.413 — Flagellum of antenna excluded from orbit; carapace slightly wider than long, if at all ...... Caphyrinae ...p.408

Subfamily Caphyrinae Paul’son, 1875 Two genera belong to Caphyrinae in Australia, only one represented in southern Australia. Diagnosis. Carapace subcircular or subhexagonal, usually not much wider than long; surface smooth or traversed on each side by ridges; eyestalks of normal length; 5–6 anterolateral teeth or nearly entire. Basal antennal article lying obliquely, with anterolateral lobe entering orbital hiatus, peduncle and flagellum excluded from orbit. Chelipeds as long or little longer than walking legs, bearing a reduced set of spines on merus, carpus and palm; fingers not or weakly grooved. Walking legs short. Male first gonopod with subterminal spines. (after Davie, 2002)

Lissocarcinus Adams & White, 1849 This genus comprises four Australian species of which only two are found in southern Australia (Davie, 2002). Diagnosis. Carapace as wide as long or slightly wider than long; anterolateral border with 5 lobes or teeth or nearly entire; front subentire or with 2 lobes. Eyes not reaching end of carapace. Chelipeds short, subequal; fingers strongly dentate on inner margin. Walking leg 4 with paddle- shaped dactylus.

Key to southern Australian species of Lissocarcinus 1. Front with 2 lobes divided by median notch; anterolateral teeth blunt; cheliped palm upper surface smooth ...... Lissocarcinus laevis

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Fig. 127. Portunidae Caphyrinae. a, b, Lissocarcinus laevis (with anterior region, ventral). c, Lissocarcinus polybioides.

— Front subentire; anterolateral teeth acuminate; cheliped palm upper surface with 2 carinae ...... Lissocarcinus polybioides Lissocarcinus laevis Miers, 1886 (Fig. 127a, b, Pl. 22c). Carapace wider than long, minutely granulate; carapace ridges indistinct; 2 frontal lobes separated by small notch; 5 anterolateral teeth, first small, second, third and fourth subequal and not acute, fifth small and dentiform. Cheliped merus with small tubercle on inner margin distally; chela palm smooth without carinae. cw. 11.5 mm. Indo-West Pacific (South Africa to Japan and Hawaii), WA, NSW; sublittoral to 50 m depth, often associated with cerianthid anemones. Lissocarcinus polybioides Adams & White, 1849 (Fig. 127c). Carapace as wide as long; epibranchial ridge distinct; front subentire; 5 anterolateral teeth. Cheliped palm upper surface bearing 2 carinae. cw. 20 mm. Widespread tropical Indo-West Pacific, Qld, NSW (Port Jackson), SA, WA (Rottnest I.); intertidal to 200 m depth, free living or associated with madreporarian corals (Stephenson, 1972).

Subfamily Carcininae MacLeay, 1838 The last leg of members of this subfamily are typically more leg-like than paddle-like, and if modified at all, the dactylus is acute rather than truly paddle-like. Carcinines are poor swimmers. Diagnosis. Carapace subhexagonal, as wide as long or moderately wider than long; not obviously convex in both directions; eyestalks short; 5 or fewer anterolateral teeth, supraorbital fissures reduced, infraorbital fissure reduced or absent. Basal antennal article narrow, cylindrical, longer than wide, lying longitudinally; lobe on maxilliped 1 (‘portunid lobe’) inconspicuous. Chelipeds long and stout; usually without spines except for inner carpal spine. At least 1 pair of walking legs

409 Marine Decapod Crustacea of Southern Australia

longer than chelipeds; dactyli of last pair of legs mostly lanceolate or styliform. Male gonopod 1 without subterminal spines. (after Davie, 2002)

Key to southern Australian genera of Carcininae 1. 5 anterolateral teeth; cheliped fingers greenish (dactyl) and white (fixed finger) . . . Carcinus — 4 anterolateral teeth; both cheliped fingers black ...... Nectocarcinus

Carcinus Leach, 1814 Carcinus is only represented by one species in southern Australia. It was introduced to Australia from the north-western Atlantic and has invaded every state of Australia except for the Northern Territory. Diagnosis. Carapace wider than long, moderately convex; 5 sharp anterolateral teeth. Basal antennal article immoveable; no gap between antennal article and inner lower orbital margin. Walking leg 4 not widened. (Ingle, 1980) Carcinus maenas (Linnaeus, 1758) European shore crab, green crab (Fig. 128a, Pl. 22d) Dorsal surface granular; frontal region produced, median lobe most advanced and not setose, submedians wide and obtuse; posterior margins of carapace almost straight. Chelipeds slightly unequal. Walking legs 1–4 smooth, moderately stout; dactylus of last styliform, compressed and with margins setose. cw. 65 mm. Introduced to Australia: NSW, Vic. (Mallacoota to Port Phillip Bay), Tas. (east and north-eastern coasts), SA (S Gulf coast), WA (Swan R., single record from 1965); originally from European and north-west African coast, North and Baltic seas, now established in southern Australia, South Africa, Japan, and northern Atlantic and Pacific coasts of North America (Carlton & Cohen, 2003); estuarine, mud flats, rocks, sand flats, seagrass, subtidal, burrowing predators of molluscs and fin-fish species, intertidal to 60 m depth. The species has been well researched in Australia because of the perceived impacts on native species (e.g. Proctor & Thresher, 1997; Sinclair, 1997; Zeidler, 1997; Thresher et al., 2002). The crab was first reported in Port Phillip Bay in 1900 (Fulton & Grant, 1900) but was not reported in South Australia until the 1970s (Zeidler, 1978) and Tasmania in the 1990s (Gardner et al., 1994). Distribution appears to have remained static for long periods but has moved long distances, naturally or human-assisted, on rare occasions. The species was recorded in the Red Sea, Pakistan, Sri Lanka, Burma, Madagascar, Panama, Brazil and Hawaii in the nineteenth century but established populations do not exist now presumably because water temperatures are too high (Carlton & Cohen, 2003). The same may be true of the species in the Swan River.

Nectocarcinus Milne Edwards, 1860 Dell et al. (1970) provided a key to this temperate genus with species from the New Zealand, Australian and eastern Pacific. Like others in the subfamily, the swimming legs are weakly developed. Diagnosis. Carapace wider than long; carapace ridges well defined; front entire or divided into lobes; 4 anterolateral teeth. Basal antennal article fixed or free, not wider than long. Maxilliped 3 elongate, ischium hollowed on outer surface. Chelipeds short, robust. Walking leg 4 weakly modified for swimming.

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b d

Fig. 128. Portunidae Carcininae. a, Carcinus maenas. b, Nectocarcinus integrifrons. c, d, Nectocarcinus spinifrons (front with orbits, outer left cheliped). e, Nectocarcinus tuberculosus (anterior region).

Key to southern Australian species of Nectocarcinus 1. Front divided into 4 lobes ...... Nectocarcinus aff. bullatus — Front entire or divided into 2 lobes ...... 2 2. Front without row of tubercles; cheliped merus with 4–5 spines on lower outer border; chela palm inner margin of upper surface with large distal spine ...... Nectocarcinus spinifrons — Front with at least 1 row of tubercles; cheliped merus lacking prominent spines on lower outer border; chela palm inner margin of upper surface with small distal spine ...... 3 3. Front entire or with very shallow medial notch; front (in larger specimens) with single row of numerous small rounded tubercles (usually on soft bottoms on sheltered coasts) ...... Nectocarcinus integrifrons — Front with V-shaped median notch; front (in larger specimens) with double row of prominent blunt tubercles (usually on more exposed rocky coasts) ...... Nectocarcinus tuberculosus

411 Marine Decapod Crustacea of Southern Australia

Nectocarcinus aff. bullatus Balss, 1924. Front divided into 4 lobes. 20 mm. This undescribed species found at 140–570 m depth in the Great Australian Bight is close to, but four times as big as, a species known only from Juan Fernandez I. off Chile (Museum Victoria collections, P. Davie, pers. comm.). Nectocarcinus integrifrons (Latreille, 1825) rough rock crab (Fig. 128b). Carapace convex, uninterrupted or sometimes with a very shallow and minute central notch; dorsal surface covered with tubercles; front (in larger specimens) with single row of numerous small rounded tubercles; epibranchial region with groups of tubercles merging with those on dorsal surface of anterolateral teeth; 4 anterolateral teeth; chela palm outer surface with tubercles in transverse rows, none more prominent or more darkly coloured than any other. cw. 80 mm. NSW (Port Jackson), Vic., Tas., SA, WA (N to Fremantle); intertidal to 15 m depth. Nectocarcinus spinifrons Stephenson, 1961 (Fig. 128c, d). Carapace uniformly covered with setae; protogastric region with granules; 4 sharp anterolateral teeth; cheliped merus bearing 4–5 spines (excluding spinules) on lower outer border, distal spine inflated, inner border with single prominent spine near tip; cheliped carpus inner border bearing large spine with secondary spinules on distal surface; chela palm outer surface with irregular rows of spiniform granules, inner margin of upper surface large spine subdistally. cw. 21 mm. SA (Greenly I.), WA (Shark Bay); 75–80 m depth. Nectocarcinus tuberculosus Milne Edwards, 1860 rough rock crab, velvet crab (Fig. 128e, Pl. 22e) Carapace densely pilose, particularly anteriorly; frontal margin almost straight, with a distinct narrow V-shaped notch centrally, bordered by double row of prominent blunt tubercles; epibranchial regions with groups of tubercles merging with groups surrounding anterolateral teeth; 4 anterolateral teeth; chela palm outer surface with tubercles in transverse rows, some more prominent and darker than others. cw. 90 mm. NSW (N to Eden), Vic., Tas., SA (W to off Murray R.); to 50 m depth. The species sometimes enter rock lobster pots and are sold in markets as velvet crabs, so-called because of the furry coating of the carapace. For details on the structure of the mouthparts and foregut see Salindeho & Johnston (2003).

Subfamily Podophthalminae Dana, 1851 The subfamily is easily recognised by the very long orbits and eyestalks, resembling ocypodid sentinel crabs. It is represented by one genus in Australia (Davie, 2002). Diagnosis. Carapace wide, widest anteriorly; front T-shaped, lying below eyestalks; orbits and eyestalks very long, occupying whole of anterior margin of carapace; supraorbital fissures reduced; 2–4 anterolateral teeth. Antennule not completely retractile in fossae beneath front. Basal antennal article short; flagellum long and slender, included in orbital hiatus. Epistome short. Chelipeds much longer than walking legs, bearing spines on merus, carpus and palm. Walking leg 4 with propodus and dactylus flattened, paddle-shaped. (after Davie, 2002)

Podophthalmus Lamarck, 1801 Of the three species of Podophthalmus, only one is found in southern Australia (Ng, 1998). Diagnosis. As for subfamily. Podophthalmus vigil (Weber, 1795) (Fig. 129). Carapace green; chelipeds and parts of legs violet to maroon in adults (Ng, 1998). cw. 142 mm. Indo-West Pacific (South Africa to Hawaii); Qld, NSW, WA; sublittoral; reported to swim to lights at night (Davie, 2002), to 70 m depth. Podophthalmus vigil is a large species with a distinctive colour pattern and is the only species of the genus of commercial value (Ng, 1998).

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Fig. 129. Portunidae Podophthalminae. Podophthalmus vigil.

Subfamily Polybiinae Ortmann, 1893 Polybiinae tend to be not much wider than long and are distinguished from the similar Caphyrinae by having the antennal flagellum included in the orbit. Stephenson & Campbell (1960) used the subfamily name, Macropipinae. Diagnosis. Carapace only moderately wide, not obviously convex in both directions; eyestalks of normal length; 4–5 anterolateral teeth. Antenna arising from orbital hiatus and lying almost along longitudinal axis of carapace; antennal flagellum in orbital hiatus. Chelipeds and walking legs stout and long, first pair of legs as long as chelipeds; last legs with rounded or moderately lanceolate dactylus. (after Davie, 2002)

Key to southern Australian genera of Polybiinae 1. Basal antennal article anterolateral angle without a lobule ...... Parathranites — Basal antennal article anterolateral angle with a lobule ...... 2 2. Dactyli of last pair of legs suboval, with apex rounded ...... Ovalipes — Dactyli of last pair of legs broadly lanceolate, with apex acute ...... Liocarcinus

Liocarcinus Stimpson, 1871 Only one species from the genus, otherwise widespread in the Atlantic and Indo-Pacific oceans, occurs in Australia. Diagnosis. Carapace wider than long; front with 3 lobes; dorsal margin of orbit with 2 closed incisions; 5 anterolateral teeth. Basal antennal article anterolateral angle produced into a lobule. Cheliped merus without distal tooth on inner border; carpus with strong tooth on inner angle, generally lacking outer tooth; chela dactylus with 3 dorsal ridges. Walking legs 1–3 each with 1 ventral ridge; walking leg 5 dactylus lanceolate. Abdominal somites 3–5 fused in males. (after Manning & Holthuis, 1981) Liocarcinus corrugatus (Pennant, 1777) ridged swimming crab (Fig. 130a, Pl. 22f, g). cw. 26 mm. Atlantic, Indo-West Pacific, southern Qld, NSW, Vic., Tas., SA, south-western WA, subtidal to about 100 m depth. This species has been broadly reported in Australia as Macropipus corru-

413 Marine Decapod Crustacea of Southern Australia

gatus (e.g. Hale, 1927). In the opinion of Davie (2002), the Indo-West Pacific and Australian species may prove to be as many as four separate species, none L. corrugatus.

Ovalipes Rathbun, 1898 A cladistic analysis of morphology of species of Ovalipes, corroborated by the fossil record, has shown that they have evolved from ancestors with stridulatory structures. The more primitive living species of Ovalipes, which inhabit shallow waters, retain the efficient stridulatory structures (Parker et al., 1998). However, some species of Ovalipes have evolved iridescence rather than sound as a means of signalling. The highly iridescent species live beyond 200 m depth, where only the blue portion of sunlight exists. Ovalipes australiensis is the most commonly encountered swimming crab in southern Australia. Shallow trawls in marine bays often catch many individuals, the cast carapaces with the characteristic pair of red spots frequently wash up on sandy beaches and markets sell sand crabs alongside other crustaceans. The key does not include O. elongatus Stephenson & Rees, 1968 from Lord Howe Island and the Kermadec Islands.

Diagnosis. Carapace slightly wider than long; front armed with 1–2 teeth or lobes, obsolete submedian frontal lobes often present; 5 anterolateral teeth. Basal antennule article short, not filling orbital hiatus, not fused distally with front; basal antennal article often visible and flat dorsally, sometimes iridescent. Stridulatory apparatus present on either mesobranchial or pterygostomial area of carapace (anterior underside) or on under surface of chela palm. (after Stephenson & Rees, 1968b)

Key to southern Australian species of Ovalipes 1. Front with 1 tooth (longer than pair of acute inner supraorbital teeth alongside) ...... Ovalipes iridescens — Front divided into 2 or 4 teeth (paired supraorbital teeth alongside) ...... 2 2. Front with 2 median teeth close together, fused, projecting in front of obsolete submedian pair and acute supraorbital teeth; upper orbital border unarmed; anterolateral teeth bordered by fine granules ...... Ovalipes molleri — Front with 2 teeth close together, separate, not longer than supraorbital teeth, without submedians; upper orbital border with tooth inside first anterolateral; anterolateral teeth bordered by coarse granules ...... 3 3. Basal antennal article without flattened dorsal area, upper surface with dense long hairs ...... Ovalipes australiensis — Basal antennal article with flattened dorsal area, upper surface sparsely hirsute ...... 4 4. Inner supraorbital notch deep; very finely granular between carinae on upper surface of chela palm; lower surface of chela palm with stridulating ridge; mesobranchial area of carapace without stridulating apparatus ...... Ovalipes catharus — Inner supraorbital notch feebly developed; very coarsely granular between carinae on upper surface of chela palm; lower surface of chela palm without stridulating ridge; mesobranchial area of carapace with stridulating apparatus ...... Ovalipes georgei Ovalipes australiensis Stephenson & Rees, 1968 sand crab (Fig. 130b, Pl. 23a, b). Front divided into 2 short teeth, not longer than triangular inner supraorbital teeth; upper orbital border toothed, inner supraorbital notch deep; 5 anterolateral teeth bordered by coarse granules. Basal antennal article without flattened dorsal area, largely obscured dorsally by hirsute frontal border, upper surface with long dense hairs. Chela palm, upper surface with 3 distinct

414 Brachyura – crabs

Fig. 130. Portunidae Polybiinae. a, Liocarcinus corrugatus. b, Ovalipes australiensis.

415 Marine Decapod Crustacea of Southern Australia

carinae, innermost ending in a spine, between carinae coarsely granular; chela palm, lower surface with stridulating ridge. Red carapace with distinctive pair of large red spots posteriorly. cw. 98 mm. Lord Howe I., Qld (N to Wide Bay), NSW, Vic., Tas., SA, WA (N to Rottnest I.); sandy beaches to 60 m depth. Sand crabs are often available as a local delicacy in spite of being much smaller than mud crabs or blue swimmers. They are caught commercially in most southern states and usually sold cooked. The two red spots on the carapace are distinctive in living animals. They have a reputation among fishers for stealing bait. Alternative common names are surf crab or two-spot crab. Ovalipes catharus White, 1843 (Fig. 131b). Front divided into 2 short teeth, not longer than triangular inner supraorbital teeth; upper orbital border toothed, inner supraorbital notch deep; 5 anterolateral teeth, bordered by coarse granules. Basal antennal article with small flattened dorsal area, clearly visibly dorsally, upper surface sparsely hirsute. Chela palm, upper surface with 3 distinct carinae, innermost ending in a spine, between carinae very finely granular; chela palm, lower surface with stridulating ridge. cw. 89 mm. New Zealand (Wear & Haddon, 1987), Vic., SA; shallow water. Ovalipes georgei Stephenson & Rees, 1968 (Fig. 131c). Front divided into 2 short teeth, scarcely longer than acute inner supraorbital teeth; upper orbital border toothed, inner supraorbital notch feebly developed; 5 anterolateral teeth, bordered by coarse granules; mesobranchial area of carapace with stridulating apparatus. Basal antennal article with flattened dorsal area (sparsely hirsute). Chela palm, upper surface with 3 distinct carinae, innermost ending in a spine, between carinae very coarsely granular. cw. 46 mm. WA (Rottnest I., Cottesloe) (Stephenson, 1972). This species possesses a unique stridulating apparatus. It involves the modified leg 4 rubbing against the granular ridge on the mesobranchial area of the carapace (Stephenson & Rees, 1968b). Ovalipes iridescens (Miers, 1886) (Fig. 131d). Front with 1 median tooth, longer than acute inner supraorbital teeth; upper orbital border unarmed, inner supraorbital notch fairly deep; 5 anterolateral teeth, fine granules between teeth; pterygostomial area of carapace with stridulating apparatus. Basal antennal article protruding well forwards, with flattened dorsal area and small hairs. Chela palm, upper surface with 3 distinct carinae, innermost distally bearing 4 spines, between outer and central carina very sparsely granular, between central and inner, granular. cw. 63 mm. South Africa, Indonesia, Philippines, Japan, Vic.; 80–440 m depth (rare). Ovalipes molleri (Ward, 1933) (Fig. 131e, Pl. 23c). Front with 2 median teeth close together, fused, projecting in front of obsolete submedian pair and acute supraorbital teeth; upper orbital border unarmed, inner supraorbital notch fairly deep; 5 anterolateral teeth, between teeth bordered by very fine granules; pterygostomial area of carapace with stridulating apparatus. Basal antennal article protruding well forwards, with flattened, hirsute, dorsal area. Chela palm, upper surface with 3 granular carinae, inner most distally bearing 4 spines, between outer and central carinae smooth, between central and inner sparsely granular. cw. 62 mm. New Zealand, Qld, NSW, Vic.; 150–270 m depth.

416 Brachyura – crabs

Fig. 131. Portunidae Polybiinae. a, Parathranites orientalis. Front and orbits: b, Ovalipes catharus. c, Ovalipes georgei. d, Ovalipes iridescens. e, Ovalipes molleri.

Parathranites Miers, 1886 Eight species are known, all from the Indo-West Pacific (see revision and key by Crosnier, 2002). Diagnosis. Carapace wider than long; front with 4 subequal teeth (including inner supraorbital teeth); 5 anterolateral teeth, first four subequal, the last spiniform and directed laterally; carapace with a curved spine at lateral margin of posterior border. Basal antennal article straight; flagellum communicating with orbit. Parathranites orientalis (Miers, 1886) (Fig. 131a). External orbital tooth entire. Carapace 1.30–1.65 times as wide as long, smooth; cardiac region without median posterior tubercle. 35 mm. Indo-West Pacific, Qld, NSW (S to Port Stephens); 80–455 m depth (Stephenson, 1972; confirmed by Crosnier, 2002). This is the only species of the genus found in Australian waters.

417 Marine Decapod Crustacea of Southern Australia

Subfamily Portuninae Rafinesque-Schmaltz, 1815 Members of Portuninae distinctly have the cheliped longer than all walking legs. Three genera are found in southern Australia and most are shallow water species. The subfamily includes the two largest edible species in Australia, the mud crab and the blue swimmer. Diagnosis. Carapace subhexagonal, usually wider than long, rarely subcircular; 4–9 pairs of anterolateral teeth. Basal antennal article usually wide, elongate, lying obliquely, with anterolateral lobe not filling orbital hiatus; antennal peduncle and flagellum included in orbital hiatus. Chelipeds longer than walking legs, bearing a set of spines on merus, carpus and palm; walking leg 4 with paddle-shaped propodus and dactylus. Male gonopod 1 without subterminal spines. (after Davie, 2002)

Key to southern Australian genera of Portuninae 1. Anterolateral teeth alternately large and small; basal antennal article with no distal lobule at anterolateral angle ...... Lupocyclus — Anterolateral teeth subequal, or if smaller teeth present, not alternating with larger ones; basal antennal article produced into a small lobule at anterolateral angle ...... 2 2. Chela palm inflated and smooth ...... Scylla — Chela palm prismatic and ribbed ...... Portunus

Lupocyclus Adams & White, 1849 Diagnosis. Carapace slightly wider than long, front prominent, divided into 4 lobes; inner supraorbital teeth smaller; 5 anterolateral teeth, smaller denticles may be in intervening notches. Male abdomen of 5 somites, somites 2–3 keeled. (after Barnard, 1950) Lupocyclus sp. (Fig. 132a). One individual of an undescribed species, cw. 19 mm, has been discovered in western Bass Strait, Vic., 84 m depth.

Portunus Weber, 1795 Portunus, known in years past as Neptunus, is a large genus with many species of commercial importance in the tropical Indo-West Pacific (Ng, 1998). Most inhabit sandy shelf environments where they burrow in the sediments emerging only to feed. Davie (2002) allocated the 22 Australian species to six subgenera not referred to here. The species occurring in southern Australia are generally at the southern extent of the range and, except for P. pelagicus locally, none is common. Diagnosis. Carapace always wider than long; surface of carapace divided into regions; front divided into 3–6 teeth (plus inner supraorbital lobes); 9 anterolateral teeth but sometimes as few as 4, subequal, or if smaller teeth present, not alternating with larger ones. Chela palm prismatic and ribbed.

Key to southern Australian species of Portunus 1. Posterior area of carapace bearing 3 red spots ...... Portunus sanguinolentus — Posterior area of carapace without spots ...... 2 2. Maxilliped 3 anterolateral angle produced more forwards than laterally (Fig. 133g, i) . . . . 3 — Maxilliped 3 anterolateral angle produced laterally (Fig. 133f, h) ...... 4

418 Brachyura – crabs

3. Carapace distinctly less than twice as wide as long (measured from largest anterolateral teeth) ...... Portunus pubescens — Carapace twice as wide as long (measured from largest anterolateral teeth ) ...... 7 4. Cheliped merus with 1 spine on outer border ...... Portunus rubromarginatus — Cheliped merus with 2 spines on outer border ...... 5 5. Last anterolateral spine slightly larger than preceding spines; 2 median frontal lobes fused . . 6 — Last anterolateral spine distinctly larger than preceding spines; 2 median frontal lobes separate ...... Portunus haanii 6. Granules on carapace covering almost entire surface; penultimate somite of male abdomen slightly convex, maximum width twice that of ultimate somite ...... Portunus granulatus — Granules on carapace in patches; penultimate somite of male abdomen with bulging curved sides, 3 times wider than ultimate somite ...... Portunus orbitosinus 7. Posterolateral junction of carapace rounded; front divided into 4 lobes . . Portunus pelagicus — Posterolateral junction of carapace elevated to form a short ridge bearing an almost right-angled border; front divided into 3 lobes ...... Portunus rugosus Portunus granulatus (Milne Edwards, 1834) (Fig. 132b, 133j). Carapace covered with close-packed granulated areas with a fine pubescence among the granules; front 4-lobed (plus supraorbital lobes), 2 medial lobes fused, shorter than lateral lobes; last anterolateral spine slightly the most protruding; posterolateral junction rounded. Cheliped merus with 2 spines on outer border. Maxilliped 3 anteroexternal angle acutely produced. cw. 31 mm. Tropical Indo-West Pacific, Qld, NSW (Port Jackson), WA; 4–120 m depth. Portunus haanii (Stimpson, 1858) (Fig. 132c). Front 4-lobed (plus supraorbital lobes); last anterolateral spine distinctly larger than preceding spines and directed laterally; posterolateral junction rounded; weak tubercle between last anterolateral tooth and posterolateral junction. Cheliped merus with 2 large spines on outer border distally. Maxilliped 3 anteroexternal angle produced. cw. 63 mm. Indo-West Pacific, NT, WA; 30–350 m depth. Portunus orbitosinus Rathbun, 1911 (Figs 132d, 133f, k). Carapace covered with widely spaced, conspicuous granulated areas, separated by regions with dense pubescence; front 4-lobed (plus supraorbital lobes), 2 median lobes fused, shorter than lateral lobes; last anterolateral spine slightly larger than preceding spine; posterolateral junction rounded. Cheliped merus with 2 spines on outer border. Maxilliped 3 anteroexternal angle produced acutely. cw. 33 mm. Indo-West Pacific, Qld, NSW, WA; sublittoral to 75 m depth. Portunus pelagicus (Linnaeus, 1758) blue swimming crab, blue swimmer, blue manna crab, sand crab (Fig. 132e, Pl. 23d, e) Carapace twice as wide as long; front 4-lobed (plus supraorbital lobes); last anterolateral spine distinctly larger than preceding spines and directed laterally; posterolateral junction rounded. Cheliped merus with 1 spine on outer border distally. Maxilliped 3 anteroexternal angle produced forwards. Males with blue patterns, especially posteriorly; females dull green and patterned all over; chelipeds and legs especially bright blue. cw. 190 mm. Widespread in Indo-West Pacific, New Zealand, throughout Australia, but rare in southern NSW,Vic., Tas., SA (except Gulfs) and southern WA; to 65 m depth. The blue swimming crab is by far the most broadly commercially caught swimming crab in Australia, and throughout south-eastern Asia. Major fisheries exist in the estuaries of southern Qld (where they are targeted or are a significant bycatch of prawn trawlers), NSW and in western SA. Pots or drop-nets are the main fishing methods for professional and amateur crabbers. Female crabs are known as jennies in NSW. The species prefers muddy or sandy bottoms but can be found with seagrass. It tends to move offshore as it grows, reaches maturity in about one year, and is more abundant, for fishers, during summer (Sumpton & Smith, 1990; Sumpton et al., 1994).

419 Marine Decapod Crustacea of Southern Australia

Portunus pubescens (Dana, 1852) (Figs 132f, 133g). Carapace 1.6 times as wide as long; front 4-lobed (plus supraorbital lobes); last anterolateral spine distinctly the longest and stoutest. Cheliped merus outer border unarmed. Maxilliped 3 anteroexternal angle not produced laterally. cw. 33 mm. Indo-West Pacific, Qld, NSW (S to Port Jackson); 20–30 m depth (Stephenson, 1972). Portunus rubromarginatus (Lanchester, 1900) (Figs 132g, 133h, Pl. 23f). Front 4-lobed (plus supraorbital lobes); last anterolateral spine distinctly the widest and most protruding and directed slightly forwards; posterolateral junction rounded; sharp posterolateral spine between last anterolateral tooth and posterolateral junction. Cheliped merus with 1 spine on outer border. Maxilliped 3 merus anteroexternal angle produced laterally. cw. 54 mm. Indo- West Pacific, Qld, NSW (S to Port Jackson), WA; to 38 m depth.

a b

d e

g h

Fig. 132. Portunidae Portuninae. a, Lupocyclus sp. b, Portunus granulatus. c, Portunus haanii. d, Portunus orbitosinus. e, Portunus pelagicus. f, Portunus pubescens. g, Portunus rubromarginatus. h, Portunus rugosus.

420 Brachyura – crabs

Portunus rugosus (Milne Edwards, 1861) (Figs 132h, 133c, e, i, Pl. 23g). Carapace twice as wide as long; front 3-lobed (plus supraorbital lobes); last anterolateral spine distinctly larger than preceding spines and directed laterally; posterolateral junction elevated to form a short ridge bearing an almost right-angled border. Cheliped merus with 1 spine on outer border distally. Maxilliped 3 anteroexternal angle with a flattened expansion directed more forwards than laterally. cw. 30 mm. Indo-West Pacific, WA (S to Cockburn Sound); 18–20 m depth.

a b

c d

Fig. 133. Portunidae Portuninae. a, Portunus sanguinolentus. b, Scylla serrata. c, Portunus rugosus (antenna). Cheliped : d, Scylla serrata. e, Portunus rugosus. Maxilliped 3: f, Portunus orbitosinus. g, Portunus pubescens. h, Portunus rubromarginatus. i, Portunus rugosus. Male abdomen: j, Portunus granulatus. k, Portunus orbitosinus.

421 Marine Decapod Crustacea of Southern Australia

Portunus sanguinolentus (Herbst, 1783) red-spotted swimming crab (Fig. 133a, Pl. 23h). Front 4-lobed (plus supraorbital lobes); last anterolateral spine distinctly larger than preceding spines and directed laterally; posterolateral junction rounded. Cheliped merus outer border unarmed. Maxilliped 3 anteroexternal angle not produced laterally. Posterior area of carapace with 3 blood-red spots. cw. 120 mm. Indo-West Pacific, New Zealand, WA (S to Geraldton), Qld, NSW (S to Botany Bay), SA; to 30 m depth. The species is of some commercial interest in South-East Asia and Australia (Sumpton et al., 1989). The three red spots on the carapace are diagnostic.

Scylla De Haan, 1833 The taxonomy of the genus Scylla has been uncertain for quite some time. Keenan et al. (1998) revised the genus to resolve four species of Scylla common throughout the Indo-Pacific. Two of these are found in Australian waters (Davie, 2002). The genus is readily distinguished from other southern Australian portunines by the inflated chela palm. Diagnosis. Carapace wider than long, surface smooth; gastrocardiac grooves poorly to moderately well-defined; front divided into 4 lobes; 9 anterolateral teeth. Cheliped longer than walking legs; chela palm inflated and without carinae; cheliped, carpus and palm with spines. Male abdomen somites 3–5 fused; female abdomen broadly oval. (after Keenan et al., 1998) Scylla serrata (Forskål, 1755) mud crab, giant mud crab, muddie (Fig. 133b, d). Cheliped carpus with 2 spines on distal half of outer margin; chela palm with pair of distinct spines on dorsal margin distally and spine at carpus articulation. Colour variable, from purple through green to browny-black depending on habitat (Keenan et al., 1998) Last legs marbled in both females and males (Ng, 1998). cw. 280 mm. Indo-West Pacific, northern New Zealand, northern WA, NT, Qld, NSW (S to Port Hacking), NSW; isolated records from Mallacoota estuary, Vic., Swan R., WA, and south-western WA estuaries; introduced to Hawaii; intertidal to 7 m depth, usually in mangroves (Stephenson, 1972). The species colonised estuaries in south-western WA by means of larval recruitment from populations in north-west of Australia in 1999–2000 (Gopurenko et al., 2003). There is a major fishery for this, the largest of the commercial swimming crabs at up to 3.5 kg in weight. Crab pots are the main fishing method in Qld and NT. The crabs are marketed live as far south as the major cities in Australia. In Qld, biology and fisheries potential have been of considerable interest.

Subfamily Thalamitinae Paul’son, 1875 Thalmatines are like portunines but differ in that the male first gonopod has subterminal spines. Most representatives of the subfamily are found in shallow water but because of their small size are not commercially exploited. The extent of the frontal border distinguishes two genera. Both have several species in tropical Australia but are found only occasionally in southern states. Diagnosis. Carapace subhexagonal or subtrapezoidal, usually markedly wider than long; 4–7 anterolateral teeth. Basal antennal article widened, usually filling orbital hiatus, with ridges, granules or spines; antennal peduncle and flagellum usually excluded from orbit. Chelipeds longer than walking legs, bearing a set of spines on merus, carpus and palm. Walking leg 4 with paddle- shaped propodus and dactylus. Male gonopod 1 with subterminal spines (Fig. 138). (after Davie, 2002)

422 Brachyura – crabs

Fig. 134. Portunidae Thalamitinae. a, Charybdis miles. b, Thalamita crenata.

Key to southern Australian genera of Thalamitinae 1. Frontal-orbital border distinctly less than greatest width of carapace (Fig. 134a); anterolateral border oblique and arched, with 6 (rarely 7 or 8) anterolateral teeth ...... Charybdis — Frontal-orbital border not much less than greatest width of carapace (Fig. 134b); anterolateral border not markedly arched, with 4 or 5 anterolateral teeth (including subsidiary tooth present in some species) ...... Thalamita

423 Marine Decapod Crustacea of Southern Australia

Charybdis De Haan, 1833 Species of Charybdis tend to be oval crabs with weak lateral spines and the front about half the total width. Fewer than half the Australian species reach southern latitudes. Diagnosis. Carapace hexagonal, wider than long; ridges distinct; front divided into 6 lobes (not including inner supraorbital lobes; posterolateral margins form an even curve or meet in a distinct projecting angle with posterior border; 6 anterolateral teeth (rarely with 7–8 teeth). Maxilliped 3 with outer distal angle produced laterally. Chelipeds unequal; cheliped merus with spines, cheliped carpus with large spine on inner angle and 3 spines on outer angle; chela propodus with spines on upper surface, and 1 proximal spine near carpus articulation, outer surface costate; fingers grooved. Walking legs compressed; leg 4 merus with strong spine at outer border; dactylus and propodus leaf-shaped, propodus generally with small spinules. Male abdomen somites 3–5 fused. Male gonopod 1 slender, elongate, with terminal bristles on inner and outer borders. (after Wee & Ng, 1995)

Key to southern Australian species of Charybdis 1. Antennal flagellum sitting in orbital hiatus between produced anterolateral lobe of basal antennal article and supraorbital lobe ...... Charybdis bimaculata — Antenna excluded from orbit; anterolateral extension of basal antennal article and supraorbital lobe in contact ...... 2 2 . Carapace with distinct ridges behind epibranchial ridges ...... 3 — Carapace without distinct ridges behind epibranchial ridges ...... 4 3. Orbit with strong dorsal inclination; chela palm without longitudinal sulcus on lower surface ...... Charybdis granulata — Orbit without strong dorsal inclination; chela palm with longitudinal sulcus on lower surface ...... Charybdis natator 4. First anterolateral tooth truncate or notched; second anterolateral tooth subequal to first . . 5 — First anterolateral tooth not truncate or notched; second anterolateral tooth rudimentary, attached to posterior border of first ...... Charybdis orientalis 5. Cheliped merus with 4 spines on inner border; leg 4 propodus with 2–4 spinules on outer border ...... Charybdis miles — Cheliped merus with 3 spines on inner border; leg 4 propodus with smooth outer border . . 6 6. Chela propodus with 4 spines on upper surface (including spine at carpus articulation); male abdomen somite 4 with keel ...... Charybdis feriata — Chela propodus with 5 spines on upper surface (including spine at carpus articulation); male abdomen somite 4 not keeled ...... Charybdis japonica Charybdis bimaculata (Miers, 1886) (Figs 135a, 137a). Carapace wider than long; first anterolateral tooth pointed, second not as sharp as others. cw. 41 mm. Tropical Indo-West Pacific, Qld (Moreton I.), NSW (S to Botany Bay); 47–275 m depth; common in deep trawls in southern Queensland (Stephenson, 1972). This is the only species of Charybdis among those recorded here in which the antennal flagellum sits in a gap between the basal antennal article and the supraorbital margin. Charybdis feriata (Linnaeus, 1758) (Figs 135b, 137f). Carapace convex and smooth; no ridges posterior to epibranchials; first anterolateral tooth truncate and strongly notched. Chelipeds smooth; cheliped merus with 3 strong spines on inner border; chela propodus with 4 spines on upper surface. Leg 4 propodus with smooth outer border. Male abdomen somite 4 with keel. cw. 116 mm. Indo-West Pacific, WA (S to Fremantle), NT, Qld, NSW, rare in Vic.; to 30 m depth.

424 Brachyura – crabs

Charybdis granulata (De Haan, 1833) (Figs 135c, 137k). Carapace surface convex, covered unevenly with dense short setae; 3 pairs of short mesobranchial ridges behind epibranchials, cardiac ridge sinuous (sometimes interrupted in middle); anterolateral teeth 1–5 stout; increasing posteriorly. Orbit with strong dorsal inclination. Chelipeds strongly granular and pubescent; cheliped merus with 3 sharp spines and several spiniform tubercles on inner border; chela palm without longitudinal sulcus on lower surface. Leg 4 propodus with serrations on outer border. cw. 67 mm. East Africa to Japan (Wee & Ng, 1995), possibly WA; 15–35 m depth. Charybdis japonica (A. Milne-Edwards, 1861) (Figs 135d, 137a, g). Carapace pilose; no ridges behind epibranchials; first anterolateral tooth with a slightly sinuous border. Cheliped merus with 3 spines on inner border; chela propodus with 5 spines on upper surface. Leg 4 propodus with smooth outer border. cw. 80 mm. Red Sea, Malaysia, China, Thailand, Taiwan and Japan; introduced to South Australia and New Zealand; 10–15 m depth. In December 2000, a single mature male of C. japonica was found in the Port River, Adelaide. The crab was caught by a recreational fisher in a hoop net. An extensive 3-day survey of the area by fisheries officers using crab pots and hoop nets failed to collect any additional specimens (R. Webber, pers. comm.). Charybdis miles (De Haan, 1835) (Fig. 134a). Carapace surface convex, with dense short setae; no ridges behind epibranchials; first anterolateral tooth notched. Chelipeds elongate, surface finely pubescent; cheliped merus with 4 spines and a spinule on inner border. Leg 4 propodus with 2–4 spinules on outer border. Pale pink, stronger pink distally on limbs (Stephenson & Rees, 1968a). cw. 67 mm. Indo-West Pacific (Arabian Gulf to Japan), NT, Qld, NSW (S to Botany Bay); 20–100 m depth. Charybdis natator (Herbst, 1794) (Figs 135e, 137l, p, Pl. 23i). Carapace uniformly pilose, sparse granules on anterolateral surface; epibranchials interrupted by unbroken metagastric ridge; 1 pair of cardiac ridges and 3 short pairs of mesobranchial ridges; first anterolateral tooth truncate. Orbit without strong dorsal inclination. Chelipeds granular and pilose; cheliped merus with 3–4 spines on inner border; chela palm with longitudinal sulcate on lower surface. Leg 4 propodus with serrations on outer border. cw. 145 mm. Tropical Indo-Pacific, Lord Howe I., Qld, NSW, WA; intertidal to 60 m depth (Sumpton, 1990a, b). Charybdis orientalis Dana, 1852 (Fig. 135f). Carapace densely pilose except elevated ridges; no ridges behind epibranchials; first anterolateral tooth sharp; second tooth rudimentary, attached to posterior border of first anterolateral tooth. Chelipeds outer surface pubescent; cheliped merus with 3 spines on inner border. Leg 4 propodus with serrations on outer border. cw. 46 mm. Tropical Indo-West Pacific, Qld (N to Cairns), NSW; intertidal to 50 m depth.

Thalamita Latreille, 1829 The genus is recognised by the extremely wide frontal-orbital border, almost the same width as the carapace. Eleven species of Thalamita are found in southern Australian waters, less than half those known from Australia. Diagnosis. Carapace wider than long; ridges distinct; front divided into 2, 4 or 6 lobes (not including inner supraorbital lobes); posterolateral borders form an even curve with posterior border; 4 or 5 anterolateral teeth. Chelipeds unequal; merus with 3 spines; carpus with large spine on inner angle and 3 spines on outer angle; chela palm with spines on upper surface, and 1 proximal spine near carpus articulation, outer surface ribbed; fingers grooved. Walking legs compressed; leg 4 merus with strong spine at posterior border, dactylus and propodus wide, propodus generally with small spinules. Male abdomen somites 3–5 fused. (after Wee & Ng, 1995)

425 Marine Decapod Crustacea of Southern Australia

a b

Fig. 135. Portunidae Thalamitinae. a, Charybdis bimaculata. b, Charybdis feriata. c, Charybdis granulata. d, Charybdis japonica. e, Charybdis natator. f, Charybdis orientalis. g, Thalamita admete. h, Thalamita bouvieri.

426 Brachyura – crabs

Fig. 136. Portunidae Thalamitinae. a, Thalamita danae. b, Thalamita intermedia. c, Thalamita macropus. d, Thalamita parvidens. e, Thalamita picta. f, Thalamita prymna. g, Thalamita sima

427 Marine Decapod Crustacea of Southern Australia

Key to southern Australian species of Thalamita 1. Carapace with 4 anterolateral teeth (including rudimentary teeth) . . . . . Thalamita bouvieri — Carapace with 5 or more anterolateral teeth (including rudimentary teeth) ...... 2 2. Front divided into 2 lobes (not including inner supraorbital lobes) ...... 3 — Front divided into more than 2 lobes (not including inner supraorbital lobes) ...... 5 3. Chela palm with 6 spines or tubercles on upper surface; male gonopod 1 bilobed ...... Thalamita admete — Chela palm with fewer than 6 spines or tubercles on upper surface; male gonopod 1 not bilobed ...... 4 4. Chela palm with squamiform markings on lower surface; male gonopod 1 distal tip broadly flared ...... Thalamita sima — Chela palm with smooth lower surface; male gonopod 1 distal tip strongly recurved ...... Thalamita parvidens 5. Basal antennal article with 1or 2 broadly fused spines and several spiniform tubercles on each side of ridge ...... Thalamita prymna — Basal antennal article with granulated or smooth ridge ...... 6 6. Frontal submedian lobes not distinctly wider than median lobes ...... 7 — Frontal submedian lobes wider than median lobes ...... 8 7. Chela palm smooth with single ridge running to fixed finger on outer surface; anterolateral tooth 4 large ...... Thalamita crenata — Chela palm granular with 3 ridges on outer surface; anterolateral tooth 4 rudimentary ...... Thalamita danae 8. Male gonopod 1 distal tip tapering; anterolateral tooth 2 rudimentary . . Thalamita macropus — Male gonopod 1 distal tip flared; anterolateral tooth 2 subequal to other anterolateral teeth ...... 9 9. Frontal median lobes with rounded borders; front divided into 6 lobes . . . . Thalamita picta — Frontal median lobes with truncate borders; front divided into 4 lobes Thalamita intermedia Thalamita admete (Herbst, 1803) (Figs 135g, 137h, 138a). Carapace finely pilose, wider (c. 1.8) than long; all anterior carapace ridges present, cardiac and mesobranchial ridges less granular; front divided into 2 lobes, separated by a distinct notch; 5 anterolateral teeth, fourth tooth rudimentary. Cheliped merus with 3–4 spines on inner border, granulated outer border; cheliped palm upper surface granulated and pilose, bearing 4 spines and 2 tubercles, lower surface extremely smooth. cw. 37 mm. Indo-West Pacific, NT, Qld, NSW, WA; intertidal to 15 m depth. Thalamita bouvieri Nobili, 1906 (Fig. 135h). Carapace finely pilose, wider (c. 1.5) than long; epibranchial ridges present, curving anteriorly from last anterolateral tooth; no ridges posterior to epibranchials; front divided into 4 lobes, separated by small notch; 4 anterolateral teeth, third rudimentary. cw. 21 mm. Indo-West Pacific, Qld, NSW (S to Port Jackson); intertidal to 40 m depth. Thalamita crenata Rüppell, 1830. (Figs 134, 137m, 138b). Carapace surface smooth, sparsely pilose; all anterior carapace ridges present and finely distinct, none behind epibranchial ridges; front divided into 6 lobes, broadly rounded; 5 anterolateral teeth subequal, decreasing slightly in size posteriorly. Cheliped merus bearing 3–4 spines on inner border; cheliped palm upper surface bearing 5 spines, outer surface smooth with single ridge running to fixed finger. cw. 75 mm. Indo-West Pacific, NT, Qld, NSW, WA; intertidal, under stones, mud flats, mangroves, occasionally amongst corals (Davie, 2002).

428 Brachyura – crabs

Thalamita danae Stimpson, 1858 (Fig. 136a, 137n). Carapace densely pilose (easily removed) except for raised transverse ridges; smooth ridge across cardiacs and each mesobranchial regions; front divided into 6 lobes, medians with truncate anterior borders, separated by narrow notch; 5 anterolateral teeth, fourth tooth rudimentary, all stout. Cheliped merus inner border bearing 3 spines on distal half, several large granules on proximal half; chela palm upper surface with 5 spines, 2 on inner border, 1 on middle of outer border, reduced tubercle at distal end and spine at carpus articulation; chela palm outer surface with 3 costae. cw. 73 mm. Indo-West Pacific, NT, Qld, NSW, WA; littoral to 40 m depth. Thalamita intermedia Miers, 1886 six-lobed crab (Figs 136b, 137o). Carapace densely pilose; cardiac region with 2 indistinct ridges, mesobranchial ridge indistinct; front divided into 4 lobes, submedians wider than medians; 5 anterolateral teeth. Chela palm with 4 spines on upper surface. cw. 38 mm. Qld, SA, WA; to 15 m depth. The ‘six lobes’ of the common name refer to the four frontal lobes plus the pair of inner supraorbital teeth. Thalamita macropus Montgomery, 1931 (Fig. 136c). Carapace pilose; all anterior ridges present; front divided into 6 lobes, medians rounded, submedians flattened and almost twice as wide as medians; 6 anterolateral teeth, decreasing posteriorly, anterolateral tooth 2 rudimentary. Cheliped merus inner border bearing 3 spines; chela palm with 4 spines and 1 tubercle on upper surface. cw. 27 mm. WA (S to Abrolhos Is), NT, Qld, NSW (S to Manly); 7–100 m depth. Thalamita parvidens (Rathbun, 1907) (Figs 136d, 138c). Mesobranchial and cardiac ridges present posterior to epibranchials; front with 2 lobes, separated by distinct notch; 5 anterolateral teeth. Chela palm upper surface with 2 inner spines and 2 outer spines or tubercles and a spine at carpus articulation. cw. 36 mm. Indo-West Pacific, Qld, NSW, WA; littoral to 50 m depth. Thalamita picta Stimpson, 1858 (Figs 136e, 137d, 138d). Carapace surface pilose; carapace ridges distinct including cardiac and pair of mesobranchial ridges; front divided into 6 lobes, medians mostly projecting, rounded and distinctly separate, laterals narrowest and separated from wide submedians by deep notch; 5 anterolateral teeth, fourth rudimentary. Cheliped merus inner border bearing 3–4 spines; chela palm with 4 spines on upper surface and outer distal tubercle, inner surface smooth or with squamiform markings. cw. 30 mm. Indo-West Pacific, Qld, NSW (S to Port Jackson); coral reef and rocky shores. Thalamita prymna (Herbst, 1803) (Figs 136f, 137e). Carapace wider than long; dorsal surface smooth and shiny, pubescence at bases of anterolateral spines and frontolateral carapace ridges; no ridges behind epibranchials; front straight, divided into 6 short and truncated lobes, submedians widest, often fused to medians; 5 anterolateral teeth, fourth rudimentary. Cheliped merus inner border bearing 3 sharp spines, lower surface smooth; chela palm with 5 stout spines on upper surface, 2 on inner border of upper surface, spiniform tubercles at proximal end, 2 on outer border and spine at carpus articulation. cw. 72 mm. Indo-West Pacific, WA (S to Abrolhos Is), NT, Qld, NSW (S to Shell Harbour); intertidal to 30 m depth. Thalamita sima Milne Edwards, 1834 four-lobed swimming crab (Figs 136g, 137g, 138e). Carapace surface pilose; all anterior carapace ridges present; cardiac ridge interrupted in midline; front divided into 2 lobes, separated by shallow notch; 5 anterolateral teeth. Cheliped merus with 3 spines on inner border; chela palm with 5 spines on upper surface, lower surface covered by squamiform markings. cw. 85 mm. Indo-West Pacific, WA (S to Swan River), NT, Qld, NSW (S to Botany Bay), SA; intertidal to 50 m depth. An entoniscid isopod parasite has been reported from this crab (Shields & Earley, 1993).

429 Marine Decapod Crustacea of Southern Australia

a b

e f

Fig. 137. Portunidae Thalamitinae. Antenna: a, Charybdis bimaculata. b, Charybdis japonica. c, Thalamita crenata. d, Thalamita picta. e, Thalamita prymna. Cheliped: f, Charybdis feriata. g, Charybdis japonica. h, Thalamita admete. i, Thalamita picta. j, Thalamita sima. Chela palm, under surface: k, Charybdis granulata. l, Charybdis natator. Chela palm, outer surface: m, Thalamita crenata. n, Thalamita danae. o, Thalamita intermedia (chela). p, Charybdis natator (swimming leg).

430 Brachyura – crabs

c d e ab

Fig. 138. Portunidae Thalamitinae. Male gonopod apex: a, Thalamita admete. b, Thalamita crenata. c, Thalamita parvidens. d, Thalamita picta. e, Thalamita sima.

References Apel, M., & Spiridinov, V.A. 1998. Taxonomy and zoogeography of the portunid crabs (Crustacea: Decapoda: Brachyura: Portunidae) of the Arabian Gulf. Fauna Arabica 17: 159–331. Barnard, K.H. 1950. Descriptive catalogue of South African decapod Crustacea (crabs and shrimps). Annals of the South African Museum 38: 1–837. Carlton, J.T., & Cohen, A.N. 2003. Episodic global dispersal in shallow water marine organisms: the case history of the European shore crabs Carcinus maenas and C. aestuarii. Journal of Biogeography 30: 1809–1820. Crosnier, A. 2002. Révision du genre Parathranites Miers, 1886 (Crustacea, Brachyura, Portunidae). Zoosystema 24: 799–825. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Dell, R.K., Griffin, D.J.G., & Yaldwyn, J.C. 1970. A new swimming crab from the New Zealand subantarctic and a review of the genus Nectocarcinus A. Milne Edwards. Transactions of the Royal Society of New Zealand 12: 49–68. Fulton, S.W., & Grant, F.E. 1900. Note on the occurrence of the European crab, Carcinus maenas,Leach, in Port Phillip. Victorian Naturalist 17: 145–146. Gardner, N.C., Kwa, S., & Paturusi, A. 1994. First recording of the European shore crab Carcinus maenas in Tasmania. Tasmanian Naturalist 116: 26–28. Gopurenko, D., Hughes, J.M., & Bellchambers, L. 2003. Colonisation of the south-west Australian coastline by mud crabs: evidence for a recent range expansion or human-induced translocation? Marine and Freshwater Research 54: 833 – 840. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Ingle, R.W. 1980. British Crabs. Oxford University Press, Inc.: New York. 222 pp. Keenan, C.P., Davie, P.J.F., & Ng, P.K.L. 1998. A revision of the genus Scylla De Haan, 1833 (Crustacea: Decapoda: Brachyura: Portunidae). Raffles Bulletin of Zoology 46: 217–245. Lanchester, W.F. 1900. On some malacostracous crustaceans from Malaysia in the collection of the Sarawak Museum. Annals and Magazine of Natural History 6: 249–265. Manning, R.B., & Holthuis, L.B. 1981. West African brachyuran crabs. Smithsonian Contributions to Zoology 306: 1–379.

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Miers, E.J. 1886. Report on the Brachuyra collected by H.M.S. Challenger during the Years1873-76. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873-76. Zoology 17: 1–362, pls 1–29. Montgomery, S.K. 1931. Report on the Crustacea Brachyura of the Percy Sladen Trust expedition to the Abrolhos Islands under the leadership of Prof. W.J. Dakin in 1913, along with other crabs from Western Australia. Journal of the Linnean Society of London (Zoology) 37: 405–464, pls 24–30. Ng, P.K. 1998. Crabs. Pp. 1045–1155 in: Carpenter, K.E., & Niem, V.H. (eds), The living resources of the western central Pacific. FAO: Rome. Parker, A.R., McKenzie, D.R., & Ahyong, S. 1998. A unique form of light reflector and the evolution of signalling in Ovalipes (Crustacea: Decapoda: Portunidae). Proceedings of the Royal Society of London 265: 861–867. Proctor, C., & Thresher, R.E. 1997. The invasive history, distribution and abundance of Carcinus maenas in Australia. In Thresher, R.E. (ed.) Proceedings of the First International Workshop on the demography, impacts and management of introduced populations of the European crab, Carcinus maenas. 20–21 March 1997, CSIRO Marine Laboratories, Hobart. Centre for Research on Introduced Marine Pests, CSIRO Marine Research, Technical Report 11: 34–36. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates. Salindeho, I.R., & Johnston, D.J. 2003. Functional morphology of the mouthparts and proventriculus of the rock crab Nectocarcinus tuberculosus (Decapoda : Portunidae). Journal of the Marine Biological Association of the United Kingdom 83: 821–834. Shields, J.D., & Earley, C.G. 1993. Cancrion australiensis new species (Isopoda: Entoniscidae) found in Thalamita sima (Brachyura: Portunidae) from Australia. International Journal for Parasitology 23: 601–608. Sinclair, M.A. 1997. Interactions between native grapsid crabs and Carcinus maenas in Victoria. In: Thresher, R.E. (ed.) Proceedings of the First International Workshop on the demography, impacts and management of introduced populations of the European crab, Carcinus maenas. 20–21 March 1997, CSIRO Marine Laboratories, Hobart. Centre for Research on Introduced Marine Pests, CSIRO Marine Research, Technical Report 11: 38–45. Stephenson, W. 1961. The Australian portunids (Crustacea: Portunidae). Australian Journal of Marine and Freshwater Research 12: 92–128. Stephenson, W. 1972. An annotated check list and key to the Indo-west–Pacific swimming crabs (Crustacea: Decapoda: Portunidae). Bulletin of the Royal Society of New Zealand 10: 1–64. Stephenson, W., & Campbell, B.M. 1959. The Australian portunids (Crustacea: Portunidae). Australian Journal of Marine and Freshwater Research 10: 84–124. Stephenson, W., & Campbell, B. 1960. The Australian portunids (Crustacea: Portunidae) IV: Remaining Genera. Australian Journal of Marine and Freshwater Research 11: 73–122. Stephenson, W., & Cook, S. 1973. Studies of Portunus gladiator complex and related species of Portunus (Crustacea: Decapoda). Memoirs of the Queensland Museum 16: 415–434. Stephenson, W., Hudson, J.J., & Campbell, B.M. 1957. The Australian portunids (Crustacea; Portunidae). Australian Journal of Marine and Freshwater Research 8: 491–507. Stephenson, W., & Rees, M. 1968a. A revision of the Charybdis miles ‘group’ of species (Crustacea: Portunidae), with description of a new species from Queensland waters. Memoirs of the Queensland Museum 15: 91–109. Stephenson, W., & Rees, M. 1968b. A revision of the genus Ovalipes Rathbun, 1898 (Crustacea, Decapoda, Portunidae). Records of the Australian Museum 27: 213–261, pls 35–42, figs 1–4. Sumpton, W.D. 1990a. Biology of the rock crab Charybdis natator (Herbst) (Brachyura: Portunidae). Bulletin of Marine Science 46: 425–431. Sumpton, W.D. 1990b. Morphometric growth and fisheries biology of the crab, Charybdis natator (Herbst) in Moreton Bay, Australia (Decapoda, Brachyura). Crustaceana 59: 113–120.

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Sumpton, W.D., Potter, M.A., & Smith, G.S. 1994. Reproduction and growth of the commercial sand crab, Portunus pelagicus (L.) in Moreton Bay, Queensland. Asian Fisheries Science 7: 103–113. Sumpton, W.D., & Smith, G.S. 1990. Effect of temperature on the emergence, activity and feeding of male and female sand crabs (Portunus pelagicus). Australian Journal of Marine and Freshwater Research 41: 545–550. Sumpton, W.D., Smith, G.S., & Potter, M.A. 1989. Notes on the biology of the portunid crab, Portunus sanguinolentus (Herbst), in subtropical Queensland waters. Australian Journal of Marine and Freshwater Research 40: 711–717. Thresher, R., Proctor, C., Ruiz, G., Gurney, R., MacKinnon, C., Walton, W., Rodriguez, L., & Bax , N. 2002. Invasion dynamics of the European shore crab, Carcinus maenas, in Australia. Marine Biology 142: 867–876. Wear, R.G., & Haddon, M. 1987. Natural diet of the crab Ovalipes catharus (Crustacea, Portunidae) around central and northern New Zealand. Marine Ecology Progress Series 35: 39–49. Wee, D.P.C.,& Ng, P.K.L.1995. Swimming crabs of the genera Charybdis De Haan, 1833, and Thalamita Latreille, 1829 (Crustacea: Decapoda: Brachyura: Portunidae) from Peninsular Malaysia and Singapore. Raffles Bulletin of Zoology, Supplement 1: 1–128. Zeidler, W. 1978. Note on the occurrence of the European shore crab Carcinus maenas (Linn., 1758) in Australia. South Australian Naturalist 53: 11–12. Zeidler, W. 1997. The European shore crab (Carcinus maenas) in southern Australian waters. In: Thresher, R.E. (ed.) Proceedings of the First International Workshop on the demography, impacts and management of introduced populations of the European crab, Carcinus maenas. 20–21 March 1997, CSIRO Marine Laboratories, Hobart. Centre for Research on Introduced Marine Pests, CSIRO Marine Research, Technical Report 11: 37.

Goneplacidae MacLeay, 1838 Goneplacids are typical undistinguished crabs, some easily confused with grapsid shore crabs or xanthoid crabs. The most reliable character to distinguish species from members of Grapsidae is the attachment of the palp of maxilliped 3: at or near the distal-internal angle of the merus in goneplacids and at the disto-external angle or middle of the anterior border in grapsids. Only one species is at all common in shallow water on the southern Australian coast – Litocheira bispinosa. The remainder are either deep-sea species or rare in this region. Of the 16 species listed by Davie (2002), nine or ten species occur in southern Australia but, as Davie has documented, genera and family-level groups have drifted in and out of this family and xanthoid families over the last decades. Even today, differentiation of the six subfamilies now recognised is difficult and some of the genera are hard to diagnose. The genera are not listed in the subfamilies to which they were assigned by Davie and only diagnostic characters are given for the genera. The key given here is an artificial one relying more on visible carapace characters than on any that might be phylogenetically significant.

Diagnosis. Carapace usually wider than long; dorsal surface convex, smooth, regions poorly defined; frontal margin usually entire, sometimes bi- to multilobate; anterolateral margins usually armed with 1–4 teeth or lobes. Orbits complete, sometimes moderately or greatly elongate. Interantennular septum typically narrow. Maxillipeds 3 with carpus articulating at or near disto- internal angle of merus. Walking legs ambulatory, never flattened. Male abdomen with 7 free somites or with somites 3–5 immoveable. Male gonopod 1 moderately stout, weakly curved or sinuous; gonopod 2 usually shorter than first. Male genital openings sternal or sternocoxal, with vas deferens passing along open or closed groove in sternum. (adapted from Davie, 2002)

433 Marine Decapod Crustacea of Southern Australia

Key to southern Australian species of Goneplacidae 1. Carapace twice as wide as long; anterior margin ending laterally in acute spine, comprising narrow front and long orbits grooved to receive extremely long eyestalks ...... Ommatocarcinus macgillivrayi — Carapace rectangular or slightly wider than long; anterior margin various but not with extremely long eyestalks ...... 2 2. Carapace hexagonal (posterolateral margin tapering), with 1 sharp laterally- or obliquely- directed spine between anterolateral and posterolateral margin ...... 3 — Carapace hexagonal, with >1 anterolateral teeth; or ovoid, with obscure lobes; or square, with 1 small lateral spine or knob ...... 5 3. Front with notch; lateral spine laterally-directed; mid-shelf species . . . . . Psopheticus insignis — Front entire; lateral spine obliquely directed forward; deep-shelf species ...... 4 4. First anterolateral spine after orbital tooth, acute and set back about one-third way to second anterolateral spine ...... Carcinoplax meridionalis — First anterolateral spine after orbital tooth, blunt and scarcely set back from orbital tooth ...... Carcinoplax victoriensis 5. Front with distinct notch in middle; anterolateral margin with 2 triangular teeth, second defining widest point of carapace ...... 6 — Front without distinct notch in middle or obscurely bilobed; anterolateral margin with 1 small spine or with blunt lobes ...... 7 6. Carapace regions defined by ridges and grooves; anterolateral border with lobe followed by 2 triangular teeth ...... Planopilumnus orientalis — Carapace regions not defined by ridges and grooves, with bilobed ridge near front; anterolateral border with 2 triangular teeth at widest point ...... Neopilumnoplax heterochir 7. Carapace as wide or slightly wider than long ...... 8 — Carapace at least 1.5 times as wide as long ...... 9 8. Anterolateral margin straight, defined by sharp spine; front double-edged ...... Litocheira bispinosa — Anterolateral margin curved, defined by slight knob; front single-edged . .Georgeoplax glabra 9. Carapace about 1.5 times as wide as long, distinctly rugose along anterior and anterolateral quarter; 4 anterolateral notches clearly delineate lobes; thoracic sternum tapering at about 45° anteriorly ...... Flindersoplax vincentiana — Carapace about 1.7 times as wide as long, not or barely rugose along anterior and anterolateral margin; anterolateral lobes poorly delineated by notches; thoracic sternum tapering at about 20° anteriorly ...... Pseudozius caystrus

Carcinoplax Milne Edwards, 1852 These pale typical crab-shaped crabs with one spine on the anterolateral corner of the rectangular carapace and another between it and the orbit can immediately be differentiated from other crabs in deep water, which are often of unusual forms. Specimens are sometimes abundant in deep- water dredge samples. Guinot (1989) described 20 species in detail without diagnosing the genus or describing the two species from southern Australia. Rathbun (1923) described the southern Australian species of which the first is more common in Museum Victoria samples. Diagnosis. Carapace widest at junction of anterolateral and posterolateral borders; anterolateral border with 2 spines; eyestalk occupying about one-quarter of frontal border; front deflexed, straight, without median notch.

434 Brachyura – crabs

Fig. 139. Goneplacidae. a, Litocheira bispinosa. b, Neopilumnoplax heterochir. c, Planopilumnus orientalis.

435 Marine Decapod Crustacea of Southern Australia

Carcinoplax meridionalis Rathbun, 1923 (Fig. 140c, Pl. 24a). First anterolateral spine after orbital tooth, acute and set back about one-third way to second anterolateral spine; usually with brown fingers. 30 mm. NSW, Vic., Tas. (including seamounts), SA, WA; shelf and slope, 126–640 m depth. Carcinoplax victoriensis Rathbun, 1923 (Fig. 140d, Pl. 24b). First anterolateral spine after orbital tooth, blunt and scarcely set back from orbital tooth; fingers only brown at tips. 37 mm. NSW, Vic., Tas.; shelf and slope, 125–765 m depth.

Flindersoplax Davie, 1989 This monotypic genus is very close to Pseudozius, both looking more like typical xanthoids (Davie, 1989; Ng & Wang, 1994). It differs from xanthoids in possessing short male gonopods 2 with a curved tip, half as long as the stout gonopod 1. In xanthoids gonopod 2 is elongate or whip-like. Diagnosis. Carapace wider than long, oval, flat or slightly convex, regions poorly defined; anterolateral margin with 4 blunt teeth; front bilobed. Thoracic sternum tapering at about 45° anteriorly. Flindersoplax vincentiana (Rathbun, 1929) (Figs 140e, 141c). 30 mm. SA (Gulf St Vincent); subtidal.

e c d b

Fig. 140. Goneplacidae. a, Psopheticus insignis. b, Georgeoplax glabra. c, Carcinoplax meridionalis. d, Carcinoplax victoriensis. e, Flindersoplax vincentiana.

436 Brachyura – crabs

a b

e c d f

Fig. 141. Goneplacidae. a, Ommatocarcinus macgillivrayi. b, Pseudozius caystrus. Thoracic sternum: c, Flindersoplax vincentiana. d, Pseudozius caystrus. e, f, Litocheira bispinosa (abdomen, front).

Georgeoplax Türkay, 1983 There is only one species, distinguished from a similar one in another genus, Litocheira bispinosa, by Türkay (1983). Diagnosis. Carapace about as long as wide; anterolateral border defined by slight knob; eyestalk occupying about one-quarter of frontal border; front deflexed, straight, single edged, without median notch. Walking legs without setae. Georgeoplax glabra (Baker, 1906) (Fig. 140b). 12 mm. SA, WA (Rottnest I.); 10–20 m depth.

Litocheira Kinahan, 1856 Diagnosis. Carapace wider than long; sharp spine at posterior end of oblique anterolateral margin; eyestalk occupying about one-quarter of frontal border; front deflexed, straight, double-edged, without median notch; walking legs setose. Litocheira bispinosa Kinahan, 1856 two-spined slender-clawed crab (Figs 139a, 141e, f, Pl. 24c). Mottled purple-brown. 11 mm. Vic., Tas. SA, WA; intertidal muddy environments to about 15 m depth. This small crab is the only common goneplacid from intertidal and shallow environments in southern Australia and is distinguishable from grapsids and others by the simple sharp lateral spines and the medial position of the palp on maxilliped 3. It is common in muddy bays, like Western Port.

Neopilumnoplax Serène, 1969 Diagnosis. Carapace more or less hexagonal; anterolateral border with teeth; eyestalk occupying about one-quarter of frontal border; front deflexed, straight, with median notch. Neopilumnoplax heterochir (Studer, 1883) (Fig. 139b). 13 mm. South-eastern Atlantic, NSW,Vic., Tas.; 108–520 m depth. A rare deep-water species identifiable on the basis of the carapace.

437 Marine Decapod Crustacea of Southern Australia

Ommatocarcinus White, 1851 The extraordinarily long eyes of members of this genus resemble and even exceed those of ocypodids. Diagnosis. Carapace much wider than long; orbital angle a laterally-directed sharp spine; eyestalks in groove occupying most of frontal border and sometimes exceeding width; front straight, without median notch. Ommatocarcinus macgillivrayi White, 1851 (Fig. 141a). 60 mm. New Zealand, Japan, Qld, NSW; subtidal to 100 m. This is a tropical species rare off NSW.

Planopilumnus Balss, 1933 Diagnosis. Carapace wider than long, regions well defined by grooves, with setae in rows; anterolateral border with broad triangular teeth; eyestalks very short, sunken; front with median notch and distinct preorbital lobe or tooth next to eye. Planopilumnus orientalis Balss, 1933 (Fig. 139c). 13 mm. Indo-West Pacific, Qld. Vic., SA; shallow subtidal, but rare in southern Australia.

Pseudozius Dana, 1851 This genus of three species is very close to Flindersoplax, both looking more like typical xanthoids (Davie, 1989; Ng & Wang, 1994). Diagnosis. Carapace wider than long, oval, frontal margin strongly deflexed, regions poorly defined; anterolateral margin with 1 prominent tooth and 3 obsolete blunt teeth; front bilobed. Thoracic sternum tapering at about 20° anteriorly. Pseudozius caystrus Adams & White, 1849 (Fig. 141b, d). 21 mm. Indo-West Pacific, Qld, Tas.; coral and rocky reef, rarely recorded.

Psopheticus Wood-Mason, 1890 The genus would appear to differ from Carcinoplax only in being without the more anterior anterolateral tooth. Diagnosis. Carapace widest at junction of anterolateral and posterolateral borders; anterolateral border ending in 1 spine; eyestalk occupying about one-third of frontal border; front deflexed, with median notch. Psopheticus insignis Alcock, 1900 (Fig. 140a). Carapace about 1.4 times as wider as long; purplish- red with large deep red spot with white edge and centre in middle of carapace. 27 mm. West Pacific, NSW; 30–50 m depth. There is only one unconfirmed record from NSW in Australian Museum collections.

References Davie, P.J.F. 1989. A re-appraisal of Heteropanope Stimpson, and Pilumnopeus A. Milne Edwards (Crustacea: Decapoda: Pilumnidae) with descriptions of new species and genera. Memoirs of the Queensland Museum 27: 129–156. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Griffin, D.J.G., & Campbell, B.M. 1969. The sublittoral Goneplacidae and Pinnotheridae (Crustacea: Brachyura) of Moreton Bay). Memoirs of the Queensland Museum 15: 141–164.

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Guinot, D. 1989. Les genre Carcinoplax H. Milne Edwards, 1852 (Crustacea, Brachyura: Goneplacidae). In: Forest, J. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 5. Mémoires du Muséum National d’Histoire Naturelle, Paris 144: 265–345. Ng, P.K.L., & Wang, C.H. 1994. Notes on the enigmatic genus Pseudozius Dana, 1851 (Crustacea, Decapoda, Brachyura). Journal of the Taiwan Museum 41: 83–99. Rathbun, M.J. 1923. Report on the crabs obtained by the F.I.S.‘Endeavour’ on the coasts of Queensland, New South Wales, Victoria, South Australia and Tasmania. Report on the Brachyrhyncha, Oxystomata and Dromiacea. Biological Results of the Fishing Experiments carried on by the F.I.S. ‘Endeavour’, 1909–14 5: 95–156, pls 116–142. Türkay, M. 1983. Georgeplax, new genus for Litocheira glabra Baker, 1906 (Crustacea: Decapoda: Brachyura). Australian Museum Memoir 18: 101–105.

Hexapodidae Miers, 1886 Three pairs of walking legs instead of the usual four are characteristic of the Hexapodidae, as the name suggests. These are not the only crab-like crustaceans with six legs in southern Australia and the key in the box (Six-legged Crabs) will distinguish them. Another striking feature is the very broad carapace. Hexapodid genera have often in the past been treated as members of Goneplacidae but deserve family status (Manning & Holthuis, 1981). These authors provided a key to 11 genera; about 20 species were listed by Huang et al. (2002). Two species in different genera occur in Australia. Diagnosis. Carapace much broader than long; regions poorly defined; lateral margins convex anteriorly, more or less divergent posteriorly; front narrow. Eyes small, reduced, with cornea scarcely wider than stalk. Chelipeds robust; 3 well-developed pairs of walking legs; last pair of legs and sternite absent. Male abdomen not filling space between last pair of walking legs; with several somites fused. Female openings sternal. (abbreviated from Davie, 2002)

Key to Australian genera and species of Hexapodidae 1. Sternum in male without grooves at end of abdomen (anterolateral triangular excavations at most) ...... Hexapinus granuliferus — Sternum in male with broad deep oblique grooves running between bases of maxillipeds 3 and end of abdomen ...... Hexapus sexpes

Hexapinus Manning & Holthuis, 1981 Manning & Holthuis (1981) listed only three species but countenanced the possibility of others. Diagnosis. Eyes reduced, moveable. Walking legs short, dactyli not enlarged. Maxilliped 3 ischium width equal to merus length; propodus not strongly dilated. Sternum in male without grooves at end of abdomen (anterolateral triangular excavations in largest individuals). Male abdominal somites 3–5 fused, telson rounded. Hexapinus granuliferus (Campbell & Stephenson, 1970) (Fig. 142a, Pl. 24d). Carapace 1.7 times as wide as long. 15 mm. Southern Qld, NSW, Vic., Tas.; intertidal to 50 m depth, bays and shelf, often found in the papery tubes of species of the polychaete genus Chaetopterus. This is the more common of the two hexapodid species. The fine dorsal granulation on the carapace is obscured by a mat of hairs in larger individuals. The legs are also extremely hairy, with a fringe of longer setae on the margins.

439 Marine Decapod Crustacea of Southern Australia

Fig. 142. Hexapodidae. a, Hexapinus granuliferus. b, Hexapus sexpes (sternum and abdomen).

Hexapus De Haan, 1833 Campbell & Stephenson (1970) and Manning & Holthuis (1981) discussed the number of species and differences between them. One species has been recorded rarely from eastern Australia; it differs from the more common Hexapinus granuliferus by the grooves on the sternum of the male, and the carapace being twice as wide long. Diagnosis. Eyes reduced, moveable. Walking legs short, dactyli not enlarged. Maxilliped 3 ischium width equal to merus length; propodus not strongly dilated. Sternum in male with broad deep oblique grooves running between bases of maxillipeds 3 and end of abdomen. Male abdominal somites 3–5 fused, telson rounded. Hexapus sexpes (Fabricius, 1798) (Fig. 142b). 12 mm. Indo-West Pacific, southern Qld, NSW; muddy habitats, intertidal to 50 m depth; rare in southern Australia.

References Campbell, B.M., & Stephenson, W. 1970. The sublittoral Brachyura (Crustacea: Decapoda) of Moreton Bay. Memoirs of the Queensland Museum 15: 235–301, pl.22.

440 Brachyura – crabs

Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B CSIRO Publishing: Melbourne. xiv, 641 pp. Huang, J.-F., Hsueh, P.-W., & Ng, P.K.L. 2002. Crabs of the family Hexapodidae (Decapoda: Brachyura) from Taiwan, with description of a new genus and new species. Journal of Crustacean Biology 22: 651–660. Manning, R.B., & Holthuis, L.B. 1981. West African brachyuran crabs. Smithsonian Contributions to Zoology 306: 1–379.

Eriphiidae MacLeay, 1838 The family placement of genera now included in Eriphiidae has been confusing, partly because of poor understanding of relationships and partly because of the recent revival of long forgotten names under laws of priority. Even now some doubt remains about the applicability of some family and subfamily names (Davie, 2002). Some genera have been variously placed in Xanthidae, Menippidae or Oziidae. The first of these is a valid and different family but the other two are here treated as subfamilies of Eriphiidae. Three of the four subfamilies of Eriphiidae occur in southern Australia and together are represented by nine species. Nineteen species in 11 genera occur in Australia as a whole. The eriphiids are difficult to characterise: their broad, oval carapace of so-called ‘xanthoid’ shape places them with xanthids and pilumnids from which they can only be distinguished by examining the male gonopods. One of the subfamilies, Menippinae, includes in southern Australia three species of large commercially significant edible deep-water crabs. Oziines and eriphiines are more the size of typical shore crabs and live in shallow cryptic rocky environments.

Diagnosis. Carapace hexagonal to transversely oval or subrectangular; frontal margin bi- or quadrilobed. Anterolateral margins with 4 lobes or broad teeth, but sometimes spines, rarely entire; usually clearly differentiated from posterolateral margins. Endostome with crests defining efferent branchial channels well developed, reaching anterior border of buccal cavity. Chelipeds unequal; larger chela often with prominent molariform tooth. Thoracic sternum narrow, never prominently widened. Male abdomen with 7 somites distinct, moveable. Male gonopod 1 stout, straight to weakly curved; gonopod 2 longer than, or as long as first, with distinct long flagellum. (adapted from Davie, 2002)

Key to southern Australian species of Eriphiidae 1. Carapace subquadrate, dorsally convex; antennae not adjacent to orbits and antennules, flagellum in prominent cleft in frontal margin ...... Eriphiinae . . . Eriphia ...2 — Carapace subovate to quadrate; antennae adjacent to orbits and antennules, flagellum free . . 3 2. Carapace, chelipeds and walking legs covered with setae; frontal lobes smooth ...... Eriphia scabricula — Carapace, chelipeds and walking legs smooth; frontal lobes tuberculate . . . . Eriphia sebana 3. Large deepwater crabs (in southern Australia); male gonopod 2 with distal part distinctly longer than subdistal part, abruptly becoming slender in terminal section ...... Menippinae ...4 — Small intertidal or subtidal crabs; male gonopod 2 with distal part subequal or little longer than subdistal part, tapering gradually ...... Oziinae ...6 4. Giant crab with oval carapace, wider than long, anterolateral margins roughly dentate ...... Pseudocarcinus gigas — Large crab with hexagonal carapace, about as wide as long, front and anterolateral margins spinose ...... Hypothalassia ...5

441 Marine Decapod Crustacea of Southern Australia

5. Scattered long setae present on frontal region behind 2 frontal spines; dactylus of last walking leg 4.7–6.1 times as long as wide; reddish brown; eastern Australia . . . Hypothalassia armata — Numerous short setae present on frontal region; dactylus of last walking leg 3.7–4.3 times as long as wide; beige to cream; southern and western Australia ...... Hypothalassia acerba 6. Orbit closed by meeting of supra- and infraorbital angles, excluding antenna (Fig. 144c) ...... Bountiana norfolcensis — Orbit open, supra- and infraorbital angles not meeting (Fig. 144d) ...... 7 7. Front emarginate in midline, each half with small tubercles; basal article of antenna broad and short, not in contact with front ...... Eupilumnus laciniatus — Front sinuous, with 3 similar shallow excavations; basal article of antenna broadly in contact with front ...... Ozius . . . 8 8. Epibranchial ridge not broken by lateral sulcus; frontal lobes swollen, appearing deflexed, first anterolateral tooth rounded, finely granulate and almost striated behind ...... Ozius deplanatus — Epibranchial ridge interrupted by a lateral sulcus; frontal region forming a concave shelf, not deflexed, first anterolateral tooth with large granules behind ...... Ozius truncatus

Subfamily Eriphiinae MacLeay, 1838

Eriphia Latreille, 1817 Davie & Ng (2000) tabulated characteristics of this genus and listed seven species. Two occur in Australia and are essentially tropical species sometimes found as far south as Sydney. Diagnosis. Carapace subhexagonal, convex to almost flat, orbit open. Eyes large, bulbous. Efferent branchial openings circular, distinct. Antennule with second article almost as long as basal article. Walking legs with slender dactyli. Eriphia scrabricula Dana, 1852 (Fig. 144b). Carapace, chelipeds and walking legs covered with setae; frontal lobes smooth. 26 mm. Widespread in Indo-West Pacific, WA, NT, Qld, to central NSW; intertidal reef in crevices. Eriphia sebana (Shaw & Nodder, 1803) (Fig. 144a). Carapace, chelipeds and walking legs smooth; frontal lobes tuberculate. 50 mm. Widespread in Indo-West Pacific, NT, Qld, to central NSW; intertidal reef. This species is one of several in the Indo-West Pacific which are known to be toxic when eaten by humans (Garth & Alcala, 1977). Luckily it is rarely encountered in southern Australia. Toxins in reef crabs have been shown to be often transient and unpredictable, so it is wise advice not to indulge in culinary experiment with any small reef crabs in tropical waters.

442 Brachyura – crabs

Fig. 143. Eriphiidae. a, Hypothalassia acerba. b, Ozius deplanatus. c, Ozius truncatus (lateral carapace).

443 Marine Decapod Crustacea of Southern Australia

c d b a

f e

Fig. 144. Eriphiidae. a, Eriphia sebana. b, Eriphia scrabricula. Orbit: c, Bountiana norfolcensis (closed). d, Ozius deplanatus (open). e, Eupilumnus laciniatus. f, Bountiana norfolcensis.

Subfamily Menippinae Ortmann, 1893

Hypothalassia Gistel, 1848 There is a minor fishery for these species which are sold as ‘champagne crab.’ The specific distinction of the western and eastern Australian species has only just been reported (Koh & Ng, 2000). These authors also gave more extended diagnoses of the genus and species. Diagnosis. Carapace hexagonal, dorsal surface smooth, with spines anteriorly, regions well defined by grooves. Front notched, with 2 spines adjacent to each side of notch, orbital margin spinose, internal and external orbital spines well developed, anterolateral margin spinose. Efferent branchial openings partially covered by maxillipeds 3. Chelipeds asymmetrical, spinose. Walking legs with stiff setae. Hypothalassia acerba Koh & Ng, 2000 champagne crab (Fig. 143a). Numerous short setae present on frontal region; dactylus of last walking leg 3.7–4.3 times as long as wide; beige to cream. 150 mm. SA, southern WA; muddy habitats, 120–220 m depth. The fisheries potential of this population was first recognised by George (1966) who reported it as H. armata. Hypothalassia armata (De Haan, 1833) champagne crab Scattered long setae present on frontal region behind 2 frontal spines; dactylus of last walking leg 4.7–6.1 times as long as wide; reddish brown. 150 mm. West Pacific Ocean, Qld, NSW; muddy and rocky habitats, 30–600 m depth.

444 Brachyura – crabs

Pseudocarcinus Milne Edwards, 1834 The single species of this genus is so familiar to most Australian carcinologists that neither it nor its genus have ever been diagnosed since Henri Milne Edwards described it on the basis of specimens collected at the end of the eighteenth century. Pseudocarcinus gigas (Lamarck, 1818) giant crab (Pls 4, 5). Carapace oval, 1.4 times as wide as long, areas well marked, smooth in adults but more tuberculate in juveniles; front with 4 short spines, anterolateral margin with 4 obscure lobes, each with 2 or 3 short teeth. Small individuals are yellow with red spots (or vice versa) with well marked regions on the carapace while larger ones are more uniformly red. 400 mm. Southern NSW, Vic., Tas., SA, southern WA; rocky reef, 30–820 m. The Tasmanian giant crab may be the heaviest brachyuran crab, reaching at least 17.6 kg but possibly more. This weight is exceeded only by estimates of fishers who have reported even larger animals falling back into the sea from crab pots! It is only males with huge claws that reach this weight; the heaviest females are about 6 kg. Males can reach 40 cm across the carapace and the larger claw is longer than this. For comparison, the anomuran Alaskan king crab, Macrocheira kaempferi, reaches more than 18 kg and its legs have a 3 metres span. The species occurs along the southern coast of Australia from 18 m to a depth of 400 m but is common only on the continental shelf break, 140–270 m on bryozoan substrates. Migration into deeper water takes place in autumn, especially among more mature individuals. Its distribution appears limited to the temperature range, 10–18°C. Gastropod molluscs and decapods dominate the diet; males use the large claw for crushing food. Juveniles moult every three or four years but as adults this interval increases to nine years in adult females. Although crabs have been tagged as part of biological and fisheries research, recapture rates from the field have been generally low and growth rate estimates unreliable. Growth rate is low, as expected for a crab living in cold water. Eggs appear in late autumn and hatch six months later in spring. The largest females produce a batch of up to 2.5 million eggs, but not every year. The hatched larva spends about 60 days in the plankton before settlement. The commercial giant crab pot fishery commenced in 1992 after a 20-year period when crabs were taken as a bycatch with rock lobsters. Giant crabs are predominantly taken from depths between 150 m and 300 m. Export markets followed the introduction of holding tanks for live crabs. Annual catches peaked at 260 tonnes in 1994 and declined to less than half that within seven years. Strict management regimes are now in place. Much has been written on fisheries biology (Levings et al., 1996, 2001), and on growth, reproduction and larval development (Gardner, 1996, 1997, 1998, 2001; Gardner & Maguire, 1998; Gardner & Quintana, 1998; McGarvey et al., 2002).

445 Marine Decapod Crustacea of Southern Australia

Subfamily Oziinae Dana, 1851

Bountiana Davie & Ng, 2000 The genus is monotypic. Diagnosis. Body transversely oval, strongly vaulted anteriorly; regions poorly defined, smooth. Front obliquely deflexed, with transverse ridge each side of median notch. Anterolateral margin with 3 small blunt spines. Orbit closed by meeting of supra- and infraorbital angles, excluding antenna. Efferent branchial openings almost completely covered by maxilliped 3, endostomial ridges weak. Maxillipeds 3 gaping. Chelipeds asymmetrical. Bountiana norfolcensis (Grant & McCulloch, 1907) (Fig. 144c, f). Norfolk and Lord Howe islands, southern NSW; intertidal, under stones. This crab has a typical xanthoid oval shape with one enlarged black-fingered cheliped, perhaps best recognised by its relatively smooth carapace (Davie & Ng, 2000).

Eupilumnus Kossmann, 1877 Yang et al. (1998) provided a key to eight species of Globopilumnus, now believed to be a junior synonym. Members of the genus may best be recognised by the tendency for the palm of the cheliped to be globular and with short spines or tubercles. Diagnosis. Carapace slightly wider than long, oval, convex, with few small dorsal spines. Front obliquely deflexed, bilobed. Efferent branchial openings not covered by maxilliped 3, not well defined. Maxillipeds 3 gaping. Chelipeds asymmetrical. Orbit open, supra- and infraorbital angles not meeting; basal article of antenna broad and short, not in contact with front. Eupilumnus laciniatus (Sakai, 1980) (Fig. 144e). Carapace covered with dense coarse pubescence, obscuring outline; anterolateral teeth lined with accessory granules. Chelipeds and walking legs without stridulating granules, propodi unarmed. 9 mm. Western Pacific Ocean, Vic.; continental slope, 258–624 m depth.

Ozius Milne Edwards, 1834 Two of five Australian species occur in the south with some overlap, O. deplanatus on the eastern coast and reaching western Victoria and O. truncatus on the southern and south-western coast. Diagnosis. Carapace transversely oval, strongly vaulted anteriorly; regions well defined, smooth. Front obliquely deflexed, lobular or sinuous. Efferent branchial openings not covered by maxilliped 3, endostomial ridges sharp. Maxillipeds 3 gaping. Chelipeds asymmetrical. Orbit open, supra- and infraorbital angles not meeting; basal antennal article broadly in contact with front. Dactylus of small cheliped as long as upper margin of propodus. Ozius deplanatus (White, 1847) reef crab (Fig. 143b). Epibranchial ridge not broken by lateral sulcus; frontal lobes relatively swollen, appearing deflexed; first anterolateral tooth rounded, hardly defined, finely granulate and almost striated behind; carapace 1.5–1.65 times as broad as long. 50 mm. New Zealand, southern NSW, Norfolk and Lord Howe islands, eastern Vic., Tas. (Flinders and Erith islands); intertidal reef. Ozius truncatus Milne Edwards, 1834 reef crab (Fig. 143c). Epibranchial ridge interrupted by a lateral sulcus; frontal region forming a concave shelf, not deflexed; first anterolateral tooth moderately well defined and with large granules behind; carapace 1.4–1.5 times as broad as long. 58 mm. Southern NSW, Vic., SA, southern WA; intertidal reef.

446 Brachyura – crabs

References Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Davie, P.J.F., & Ng, P.K.L. 2000. Bountiana, a new genus for Eriphia norfolcensis Grant & McCulloch, 1907 (Crustacea: Brachyura: Eriphiidae). Memoirs of the Queensland Museum 45: 267–272. Gardner, C. 1996. Behavioural basis of depth regulation in the first zoeal stage of the giant crab Pseudocarcinus gigas (Brachyura: Oziidae). In: High latitude crabs -biology, management, and economics. Proceedings of the international symposium on biology, management, and economics of crabs from high latitude habitats, Anchorage, Alaska, October 11–13, 1995. University of Alaska Sea Grant College Program Report 96–02: 229–253. Gardner, C. 1997. Effect of size on reproductive output of giant crabs Pseudocarcinus gigas (Lamarck) (Oziidae). Marine and Freshwater Research 48: 581–587. Gardner, C. 1998. First record of larvae of the giant crab Pseudocarcinus gigas in the plankton. Papers and Proceedings of the Royal Society of Tasmania 132: 47–48. Gardner, C. 2001. Composition of eggs in relation to embryonic development and female size in giant crabs Pseudocarcinus gigas (Lamarck). Marine and Freshwater Research 52: 333–338. Gardner, C., & Maguire, G.M. 1998. Effect of photoperiod and light intensity on survival, development and cannibalism of larvae of the Australian giant crab Pseudocarcinus gigas (Lamarck). Aquaculture 165: 51–63. Gardner, C., & Quintana, R. 1998. Larval development of the Australian giant crab Pseudocarcinus gigas (Lamarck, 1818) (Decapoda: Oziidae) reared in the laboratory. Journal of Plankton Research 20: 1169–1188. Garth, J.S., & Alcala, A. 1977. Poisonous crabs of the Indo-West Pacific coral reefs, with special reference to the genus Demania Laurie. Pp. 645–651, figs 1–7 in: Proceedings of the the Third International Coral Reef Symposium, Rosential School of Marine and Atmospheric Science. University of Miami: Miami. Garth, J.S., & Kim, H.S. 1983. Crabs of the family Xanthidae (Crustacea: Brachyura) from the Philippine Islands and adjacent waters based largely on collections of the U.S. Fish Commission steamer Albatross in 1908-1909. Journal of Natural History 17: 663–729. George, R.W. 1966. Hypothalassia armata (De Haan) in Western Australia. Crustaceana 10: 223–224. Grant, F.E.,& McCulloch, A.R. 1907. Decapod Crustacea from Norfolk Island. Proceedings of the Linnean Society of New South Wales 32: 151–6,pl.1. Koh, S.K., & Ng, P.K.L. 2000. A revision of the spiny crabs of the genus Hypothalassia Gistel, 1848 (Crustacea: Decapoda: Brachyura: Eriphiidae). Raffles Bulletin of Zoology 48: 123–141. Levings, A., Mitchell, B.D., Heeren, T., Austin, C., & Matheson, J. 1996. Fisheries biology of the giant crab (Pseudocarcinus gigas, Brachyura, Oziidae) in southern Australia. In: High latitude crabs -biology, management, and economics. Proceedings of the international symposium on biology, management, and economics of crabs from high latitude habitats, Anchorage, Alaska, October 11–13, 1995. University of Alaska Sea Grant College Program Report 96–02: 125–151. Levings, A., Mitchell, B.D., McGarvey, R., Mathews, J., Laurenson, L., Austin, C., Heeren, T., Murphy, N., Miller, A., Rowsell, M., & Jones, P. 2001. Fisheries biology of the Giant Crab, Pseudocarcinus gigas. Fisheries Research and Development Corporation Final Report 93/220 & 97/132. McCoy, F. 1885–1890. Prodromus of the Zoology of Victoria: or figures and descriptions of the living species of all classes of the Victorian indigenous animals. Vol 2. Government Printer: Melbourne. McGarvey, R., Levings, A., & Matthews, J.M. 2002. Moulting growth of the Australian giant crab, Pseudocarcinus gigas. Marine and Freshwater Research 53: 869–881. Yang, S.-L., Dai, A.-Y., & Ng, P.K.L. 1998. Globopilumnus fragaria, a new species of eriphiid crab (Crustacea: Decapoda: Brachyura: Xanthoidea) from the South China Sea, with a key to the genus Globopilumnus Balss, 1933. Raffles Bulletin of Zoology 46: 93–99.

447 Marine Decapod Crustacea of Southern Australia

Pilumnidae Samouelle, 1819 Pilumnids are small cryptic crabs found under rocks, in crevices or in coral. Many are wider than long. Some species are readily recognised as belonging to Pilumnidae because of the dense mat and/or fringe of hair on the carapace and legs. Others are not densely hairy and may seem similar to Xanthidae (both families usually have distinctive black-tipped chelae). When in doubt it is necessary to examine the male abdomen and gonopods (pleopods 1 and 2). In pilumnids the male abdomen has seven somites free (rarely of six) while in xanthids there are only five. Another family that might be confused is Eriphiidae, also with seven abdominal somites in the male but in these, the male pleopod 1 is about as long as pleopod 2. In pilumnids, the first gonopod is slender and sinuous while the second is very short and sigmoid. Pilumnids are among the most difficult of crabs to identify to species in southern Australia, and close attention to the structure of the first pleopod (gonopod 1) of male specimens will be required to separate some closely related species. This section relies heavily on information provided by Griffin & Yaldwyn (1971). The family diagnosis is from Davie (2002) who recognised and diagnosed six subfamilies for about 87 species recorded in Australia. Half the subfamilies occur in southern Australia and some are treated as full families by other authors. The eumedonines are atypical curiously-shaped smooth crabs whose systematic position has varied from full family rank to subfamily of Pilumnidae or Parthenopidae (Ng & Clark, 2000). The key to subfamilies and genera will work for species in southern Australia but may not be applicable for other faunas. Genera are arranged alphabetically within subfamilies.

Diagnosis. Carapace hexagonal, transversely rectangular or transversely ovate; dorsal surface convex, smooth or granular; frontal margin entire, bilobate or multilobate; anterolateral margins typically with 1–4 teeth, lobes, or spines. Endostome with crests defining efferent branchial channels more or less well developed and continued to anterior border of buccal cavern. Claws subequal or markedly unequal. Legs stout, short to moderately long. Abdomen of male with 7 moveable somites; elongate and narrow, covering most of sternite 4; abdominal-sternal locking mechanism present; genital openings coxal or coxal-sternal. Male gonopod 1 slender, sinuous, tip typically simple, recurved; gonopod 2 very short and sigmoid. (from Davie, 2002)

Key to southern Australian subfamilies and genera 1. Carapace flat, angular, with spine between anterolateral and posterolateral angles; front projecting ...... Eumedoninae ...2 — Carapace convex, oval or subquadrate, anterolateral and posterolateral angles weakly differentiated; front scarcely extended anteriorly ...... 5 2. Rostrum of 2 prominent well separated spines (and narrow obscured medial lobes); first walking legs more slender and longer than others; floor of orbit in contact with front, excluding antennal peduncle ...... Ceratocarcinus ...p.449 — Rostrum of 2 close lobes, either slightly notched or deeply separated; walking legs similar; floor of orbit not in contact with front, separated by gap filled by antennal peduncle . . . . . 3 3. Rostrum of 2 deeply separated lobes; lateral angle of carapace directed forward; walking legs compressed and lobed ...... Zebrida ...p.451 — Rostrum triangular, medially notched; lateral angle of carapace directed laterally; walking legs not compressed ...... 4 4. Merus of walking legs cylindrical, granulate ...... Gonatonotus ...p.451 — Merus of walking legs distinctly crested, almost entire ...... Eumedonus ...p.449

448 Brachyura – crabs

5. Carapace with dense mat of hairs obscuring regions and margins (except Pilumnopeus with scattered hairs); eyestalks fully formed in deep orbits ...... Pilumninae ...6 — Carapace smooth; eyestalks small, immobile in shallow orbits with weak lower margins, eyes reduced or obsolete ...... Rhizopinae ...8 6. Carapace naked or with only scattered hairs in anterolateral region (sides and ventral surfaces densely covered with long hairs) ...... Pilumnopeus ...p.452 — Carapace usually well covered with long hairs or felt (at least on anterior and lateral regions) 7 7. Carapace totally covered with an even velvet of short thick hairs; posterolateral region of carapace subtly concave ...... Actumnus ...p.452 — Carapace covered totally, or on anterior and anterolateral regions only, with stiff or shaggy long hairs, sometimes mixed with short clubbed hairs; posterolateral region of carapace subtly convex ...... Pilumnus ...p.452 8. Eyes obsolete; carapace semicircular, widest posteriorly ...... 9 — Eyes embedded in orbits, moveable, cornea well formed; carapace subrectangular ...... 10 9. Merus of maxilliped 3 anterolaterally rounded ...... Xenophthalmodes ...p.458 — Merus of maxilliped 3 produced at anterolateral angle ...... Mertonia . ..p.458 10. Carapace dorsally highly polished; with more or less single marginal row of long setae ...... Ceratoplax ...p.457 — Carapace dorsally with some setae and with dense band of long setae marginally and extending on to front and sides of carapace ...... Heteropilumnus ...p.458

Subfamily Eumedoninae Dana, 1853 The subfamily has been treated at full family rank and as a subfamily of Parthenopidae. Its members are superficially not at all similar to the typical ovate furry pilumnid.

Ceratocarcinus White, 1847 Chia & Ng (1998) redefined the genus in detail and included three species. Diagnosis. Carapace broader than long; rostrum short, deflexed; inner supraorbital tooth long, produced beyond rostrum; anterolateral margin lamelliform, with 3 blunt teeth separated by narrow fissures and strong oblique fourth tooth defining anterior end of posterolateral margin. Chelipeds elongate, granular. Ceratocarcinus longimanus White, 1847 gouty crab (Fig. 145a). Red-brown with 4 thin yellow transverse stripes; borders of carapace yellow. 18 mm. Indo-West Pacific, NT, Qld, SA, WA; sublittoral, associated with comatulid crinoids. The distinctive hexagonal shape with the four anterior corners defined by stout blunt spines, immediately identifies this species. Hale’s (1927) record of C. dilatatus (a synonym) from the Great Australian Bight is the only record of this essentially tropical species so far south (Castro et al., 1995).

Eumedonus Milne Edwards, 1834 The five species of Eumedonus are commensal with sea-urchins (Chia & Ng, 2000), recognised by the pentagonal shape, bilobed rostrum and laterally directed carapace spines. The crests on the walking legs distinguish the genus from Gonatonotus. The shape of the carapace varies within species with age and may not be reliable for the separation of some species.

449 Marine Decapod Crustacea of Southern Australia

d h

e f

Fig. 145. Pilumnidae Eumedoninae. a, Ceratocarcinus longimanus. b, Eumedonus niger. c, Eumedonus vicinus. d, Gonatonotus nasutus. Walking leg 4: e, Eumedonus niger. f, Gonatonotus nasutus. g, Zebrida brevicarinata. Pilumninae. h, Actumnus setifer. i, Pilumnopeus serratifrons.

450 Brachyura – crabs

Diagnosis. Carapace pentagonal, with strong lateral spine; inner supraorbital teeth absent. Chelipeds with dorsal margin of palm usually with lamelliform crests in females. Upper margins of merus and carpus of walking legs distinctly crested, propodus slightly so; lower margin of merus with 2 parallel crests, stronger only on merus 4.

Key to southern Australian species of Eumedonus 1. Carapace and walking legs with few granules; lateral spines on carapace symmetrically triangular, directed laterally ...... Eumedonus vincinus — Carapace and walking legs granular; lateral spines on carapace asymmetrical, directed slightly posteriorly ...... Eumedonus niger Eumedonus niger Milne Edwards, 1834 (Fig. 145b, e). Carapace and walking legs granular; lateral spines on carapace asymmetrical, directed slightly posteriorly. Male abdominal somites 3 and 4 separated by shallow depression. 15 mm. Japan, China, Philippines, WA, Qld, NSW; shelf to 138 m depth. Eumedonus vincinus Rathbun, 1918 (Fig. 145c). Carapace and walking legs with few granules; lateral spines on carapace symmetrically triangular, directed laterally. Male abdominal somites 3 and 4 separated by deep depression. 14 mm. Qld, NSW; shelf to 200 m depth.

Gonatonotus Adams & White, 1847 Species of Gonatonotus, like Eumedonus, are associated with sea-urchins (Chia & Ng, 2000). The shape of the rostrum distinguishes its three species. Diagnosis. Carapace pentagonal, with strong lateral spine; inner supraorbital teeth absent; surfaces of carapace, chelipeds and ambulatory legs covered with numerous granules. Chelipeds with dorsal margin of palm granular, without crests. Upper margins of merus and carpus of walking legs granular, not crested; lower margin of merus with 2 faint parallel crests. Gonatonotus nasutus Chia & Ng, 2000 (Fig. 145d, f). Rostrum 1.2 times as long as wide, with ventral ridge; anterior part of interantennular septum depressed. 16 mm. Philippines, Vanuatu, New Caledonia, WA (S to Lancelin I.), NSW (Byron Bay); 20–210 m depth. The species has been seen in NSW with the echinoid Phyllacanthus parvispinus.

Zebrida White, 1847 The zebra-like stripes running front to back are one of the features of species of Zebrida.The crabs, often in pairs, attach to the spines of sea urchins using the subchelate walking legs. The genus, long thought to be monotypic, is now considered to comprise three species (Ng & Chia, 1999). The ecology of Z. adamsi White, 1847 has been well studied. Ten species of echinoid hosts have been identified and the crabs do eat the urchin’s tissue (Suzuki & Takeda, 1974; Ng & Chia, 1999). Diagnosis. Carapace flat, smooth, with longitudinal brown and white stripes; rostrum of 2 deeply separated lobes; lateral angle of carapace flat, directed forward. Walking legs compressed and lobed, propodus and dactylus subchelate. Zebrida brevicarinata Ng & Chia, 1999 (Fig. 145g, Pl. 24e). Eyestalk without lamelliform expansion. Rostrum rounded. Lateral carapace tooth rounded. Walking legs 1–3 with meri unarmed on upper margins. 8 mm. WA (S of Perth; a specimen from Albany identified as Z. adamsi is this species).

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Subfamily Pilumninae Samouelle, 1819 Most typical ovate hairy or velvety pilumnids belong to this subfamily, distinguished in southern Australia from rhizopines by having well developed eyes. Differences between the genera are subtle, except to crab specialists.

Actumnus Dana, 1851 Actumnus and Pilumnus are similar genera and traditional keys rely on the former having a more convex carapace than the latter, the setation being more even, and the posterolateral border being concave to house the last walking leg. These are all relative characters difficult to understand unless specimens of Pilumnus are alongside. Davie (2002) listed five species for Australia but only one occurs in the south. It might be confused with P. rufopunctatus but the carapace regions are indicated by a clear pattern of grooves not seen in Pilumnus. Diagnosis. Carapace strongly convex, areas well marked, totally covered with an even velvet of short thick hairs; posterolateral region of carapace subtly concave. Actumnus setifer (De Haan, 1835) short-haired crab (Fig. 145h, Pl. 24f). Carapace subcircular, 1.3 times as wide as long; front with a shallow concavity and tuberculate lobe close to orbit. Cheliped with evenly-spaced smooth round granules protruding through felt on outer surface of propodus and dorsal surface of carpus. Walking legs with dense felt and conspicuous fringe of long hairs. Creamy-white. 19 mm. Indo-West Pacific, SA, WA, NT, Qld; coral reefs, subtidal reef to 200 m depth.

Pilumnopeus Milne Edwards, 1863 Nine or so species occur through the Indo-West Pacific and eastern Atlantic, only one in Australia (Davie, 1989). Diagnosis. Carapace anteroposteriorly convex, dorsal surface with hairy granular crests. Front bilobed, lateral lobule separate from supraorbital angle; anterolateral margin with 4 teeth or lobes, first a broad lobe confluent with outer orbital angle; suborbital margin with a large tooth at inner end, visible in dorsal view. Sternum with fused somites 3–4 long, such that telson reaches less than half distance to suture 2–3; sternite 8 visible laterally in male. Pleopod 1 of male slender, sinuous, tip recurved. Pilumnopeus serratifrons (Kinahan, 1856) smooth-handed crab (Fig. 145i, Pl. 24g). Carapace length:width ratio 3:4. First 2 anterolateral lobes truncate, third sharp and fourth right- angled, margins of all marked with minute beads. Few scattered hairs on anterolateral regions of carapace. New Zealand, southern Qld, NSW, Vic., SA; estuarine, mud flats under stones, lower intertidal. This is the only Australian species in the genus, described in detail by Davie (1989). The four anterolateral teeth and rough ridges on the anterior carapace are distinctive. Larval stages are known (Greenwood & Fielder, 1984).

Pilumnus Leach, 1816 This is perhaps the most difficult crab genus of all despite the optimistic comment of Hale (1927: 162) that ‘identification is easy’ and his straightforward key. Rathbun (1923), who described several of the species included here, also provided a key that takes quite some effort to master. I agree with Griffin & Yaldwyn (1971) who pointed out that identification of species of Pilumnus ‘can be difficult’, in particular with the earlier keys. Species characterisation is complicated by the fact that small specimens tend to be less tuberculate and spinous. Ornamentation characters should therefore be used cautiously when specimens of different sizes are being compared. Any

452 Brachyura – crabs

carapace spination is usually obscured by the mat of short hairs but should be looked for in the anterolateral region inside the carapace margin. The difference between stiff simple bristles and soft fat clubbed setae is real and important. Clubbed setae may be short and form a velvety mat, especially anteriorly, or long and scattered in groups or short rows. Griffin & Yaldwyn provided a table of characters in which most species can be compared. Their illustrations of male gonopods, some reproduced here, are an aid to the identification of difficult species. Takeda & Miyake (1968) provided a key to species from the western Pacific and redescribed some Australian species. This new key builds on all of these works. It does not include P. humilis Miers, 1884 described from Qld but recorded from Tasmania by Davie (2002).

Diagnosis. Carapace covered totally, or on anterior and anterolateral regions only, with stiff or shaggy long hairs, sometimes mixed with short clubbed hairs; usually with 3 anterolateral spines or lobes; posterolateral region of carapace subtly convex.

Key to southern Australian species of Pilumnus 1. Carapace and legs covered with a dense pile of long shaggy thick hairs over all surfaces, especially long on margins of carapace and walking legs, obscuring all tubercles and spines; carapace regions divided by deep grooves between clusters of hairs . . . . Pilumnus vespertilio — Carapace and legs with at most a velvety pile of short hairs or scattered stiff hairs, sometimes smooth posteriorly ...... 2 2. Carapace with ridges, tubercles or spines near anterolateral margin in addition to 3 marginal spines; with soft pile or sparse shaggy hairs over anterior region and sometimes posterior region also ...... 3 — Carapace without ridges or numerous spines (or with few small spines and tubercles) parallel to and near anterolateral margin in addition to 3 marginal spines; with stiff hairs on carapace and legs and few soft hairs ...... 6 3. Submarginal ornamentation comprises 1–3 tall erect spines near anterolateral margin and sometimes 1 behind orbit; frontal lobes separated by deep narrow notch, each lobe with tubercles; stiff hairs over most of carapace and chelipeds ...... Pilumnus tomentosus — Submarginal ornamentation comprises tubercles or ridges near anterolateral margin and sometimes behind orbit; frontal lobes separated by broad V, each lobe smooth or with obscure tubercles; setation often absent or reduced posteriorly, usually mostly soft velvet or shaggy hairs ...... 4 4. Carapace with dense pile of short clubbed hairs, without tufts of long hairs; male gonopod 1 tip bent at less than right angle ...... Pilumnus rufopunctatus — Carapace with dense pile of low clubbed hairs mixed with tufts of long hairs; male gonopod 1 bent at right angle or hooked ...... 5 5. Carapace with clubbed setae, at least close to anterior and anterolateral margins, plus few scattered clusters of 2–3 longer soft hairs elsewhere; with group of 3–12 tubercles on a curved ridge near anterolateral margin (obscured ridge or small tubercles only on small individuals); male gonopod 1 with hooked tip, less bent in juveniles, with long setae at right angle near tip ...... Pilumnus monilifer — Carapace with dense pile of clubbed setae at least over anterior two-thirds, with long hairs; tubercles near anterolateral margin scattered; male gonopod 1 with tip bent at right angles, without long marginal setae ...... Pilumnus fissifrons 6. Anterolateral margin with 3 long sharp conical spines, with bases not markedly swollen . . 7 — Anterolateral margin with 3 triangular or conical spines, with bases broad and swollen . . . 8

453 Marine Decapod Crustacea of Southern Australia

7. Carapace with about 12 hairs forming a fringe behind frontal margin; longest of anterolateral spines about as long as orbit ...... Pilumnus acer — Carapace without fringe behind frontal margin orbit; longest of anterolateral spines shorter than orbit ...... Pilumnus australis 8. Carapace with numerous stiff hairs, smooth; upper margin of meri, propodi and carpi of walking legs without spines ...... Pilumnus kingstoni — Carapace with scattered long stiff hairs; larger specimens with small spinules close to anterolateral margin of carapace; upper margin of meri of walking legs with few tubercles or spines, of carpi with spines ...... 9 9. Outer face of palm of cheliped with short, low, acute tubercles; meri of walking legs with dorsal spines; propodi with spines ...... Pilumnus vestitus — Outer face of palm of cheliped with elongate spines, higher than basal width, meri of walking legs without dorsal spines; propodi with dorsal spines ...... Pilumnus etheridgei Pilumnus acer Rathbun, 1923 long-spined hairy crab (Fig. 146a, k, Pl. 25a). Carapace anterolateral margin with 3 long sharp conical spines; carapace with numerous scattered stiff hairs, about 12 hairs forming a fringe behind frontal margin. Cheliped with spines on outer face; walking legs with sharp spines on meri, carpi, but not propodi. 22 mm. Vic., Tas., SA, WA (S coast); subtidal to 220 m depth. Pilumnus australis Whitelegge, 1900 (Fig. 146b, l). Similar to P. acer but without frontal fringe of hairs, without spine on external orbital angle, orbit with mesial spine on ventral margin. 12 mm. NSW (Coogee to Batemans Bay), 66–146 m depth; rocky substrates. Pilumnus etheridgei Rathbun, 1923 (Fig. 146c, m, Pl. 25b). Carapace anterolateral margin with 3 sharp spines, usually secondarily tuberculate; carapace with scattered long stiff hairs, especially anteriorly; larger specimens with small spinules close to anterolateral margin of carapace. Cheliped with small stout spines on outer face; walking legs with few tubercles or smooth on upper margin of meri, with spines on carpi and propodi. 17 mm. NSW (S of Port Kembla), Vic., Tas., SA, WA (N to Fremantle); intertidal (rarely), subtidal to 100 m (common in Bass Strait). Pilumnus fissifrons Stimpson, 1858 tasselled crab (Figs 146d, 147k, l). Like P. monilifer but with few granules near anterolateral border, male gonopod 1 with straight tip bent almost at a right angles, and strongly depressed bilobed front, each lobe deeply excavate laterally into a rounded mesial part and granule-tipped lateral part, and walking legs with carpi and propodi with sharp granules. 12 mm. Qld, NSW, SA, WA; subtidal (Takeda & Miyake, 1968). Pilumnus kingstoni (Rathbun, 1923) downy crab (Fig. 146e, n). Like P. etheridgei but with prominent (hepatic) spine (in place of tubercles) in front of first anterolateral spine; walking legs without spines or tubercles on upper margins. 19 mm. Vic., SA; 24–560 m depth (rare). Pilumnus monilifer Haswell, 1881 beaded hairy crab (Figs 146f, g, o, 147m, n, Pl. 25c). Carapace with clubbed setae, at least close to anterior and anterolateral margins, plus few scattered clusters of 2–3 longer soft hairs elsewhere; carapace with group of 3–12 tubercles on a curved ridge near anterolateral margin (see below). Male gonopod 1 with hooked tip, less bent in juveniles, with long setae at right angle near tip. 19 mm. Vic., Tas., SA; intertidal to 69 m depth. This is the most common species of Pilumnus in Bass Strait, recognisable from the mat of short clubbed hairs and scattered longer hairs, plus the curved row of short tubercles protruding through this mat. As presently understood it is highly variable (Griffin & Yaldwyn, 1971). The curved row of tubercles of larger specimens is represented in juveniles simply as a low ridge buried in the soft velvety fur. The hooked male gonopod 1 with prominent setae is characteristic but this is less bent in what are supposed to be smaller males, although the setae are present. In some of these smaller specimens the meri of the walking legs are spinose.

454 Brachyura – crabs

d a

b e

f g

i k

j l

m p o

Fig. 146. Pilumnidae Pilumninae. Right anterolateral carapace and front: a, Pilumnus acer. b, Pilumnus australis. c, Pilumnus etheridgei. d, Pilumnus fissifrons. e, Pilumnus kingstoni. f, g, Pilumnus monilifer (with front view). h, Pilumnus rufopunctatus. i, Pilumnus tomentosus. j, Pilumnus vestitus. Walking leg 3: k, Pilumnus acer. l, Pilumnus australis. m, Pilumnus etheridgei. n, Pilumnus kingstoni. o, Pilumnus monilifer. p, Pilumnus vestitus.

455 Marine Decapod Crustacea of Southern Australia

c b

e j

g i

o k l m n p

Fig. 147. Pilumnidae Rhizopinae. a, Heteropilumnus fimbriatus. b, Ceratoplax luteus. c, Mertonia integra. d, Xenophthalmodes dolichophallus. Cheliped: e, Ceratoplax glaberrima (male). f, g, Ceratoplax luteus (male and female). h, Mertonia integra. i, Xenophthalmodes dolichophallus. Maxilliped 3: j, Mertonia integra. Pilumninae. Male gonopods 1 and 2 (from Griffin & Yaldwyn, 1971): k, l, Pilumnus fissifrons. m, n, Pilumnus monilifer. o, p, Pilumnus rufopunctatus.

Pilumnus rufopunctatus Stimpson, 1858 red-spotted hairy crab (Figs 146h, 147o, p). Similar to P. monilifer but carapace without tufts of long hairs (short hairs only) and with scattered sharp tubercles over anterolateral regions (8 or 9 pairs not in curved row) projecting through hairs. 17 mm. NSW (S of Clarence R.), SA, WA; subtidal (Takeda & Miyake, 1968) (not present in large Museum Victoria collections).

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Pilumnus tomentosus Latreille, 1825 common hairy crab (Fig. 146i, Pl. 25a). Carapace with submarginal 1–3 tall erect spines near anterolateral margin and sometimes 1 behind orbit; frontal lobes separated by deep narrow notch, each lobe with tubercles; carapace, chelipeds and walking legs with even covering of stiff hairs, no clubbed setae. 38 mm. NSW (Newcastle), Vic., Tas., SA, WA (W to Albany); intertidal and (more commonly) subtidal to 310 m depth. The coat of stiff bristles identifies this common species. Records of P. tomentosus from northern Australia and elsewhere in the Indo-West Pacific have been referred to another species, P. terraereginae Haswell, 1882 (Griffin, 1970). Pilumnus vespertilio (Fabricius, 1793). Carapace and legs covered with a dense pile of long shaggy thick hairs over all surfaces, especially long on margins of carapace and walking legs, obscuring all tubercles and spines; carapace regions divided by deep grooves between clusters of hairs. 34 mm. Widespread in Indo-West Pacific, tropical WA, NT, Qld; on coral and rocky reefs. The species is instantly recognisable but a record of its occurrence near Sydney seems improbable (Takeda & Miyake, 1968). Pilumnus vestitus Haswell, 1882 (Fig. 146j, p). Carapace anterolateral margin with 3 sharp spines, with bases broad and swollen; carapace with scattered long stiff hairs, especially anteriorly; larger specimens with small spinules and tubercles close to anterolateral margin of carapace. Cheliped with short low, acute tubercles on outer face; walking legs with spines on upper margin of meri and carpi but not propodi. 18 mm. NSW (near Angourie Point, S to Port Jackson); subtidal. This species hatches at an advanced megalopa stage (Hale, 1931).

Subfamily Rhizopinae Stimpson, 1858 While some rhizopines are similar in overall form to typical oval pilumnids, others assume more unusual shapes. Those with rectangular bodies with rounded corners resemble hexapodids, but have four pairs of walking legs. All have reduced eyes. Several are characterised by a fringe of long setae along the front and anterolateral margins. Of 20 genera reviewed by Ng (1987), ten genera and about 20 species inhabit Australia. Differences between them are subtle and their taxonomic history complex (Ng, 1987). In spite of examining type material, not all material at hand could be confidently identified.

Ceratoplax Stimpson, 1858 These pale crabs are best recognised by the single fringe of long setae bordering the carapace, chelipeds and walking legs. Davie (2002) recorded three of the five Australian species from the southern coast but they are rarely taken. Ceratoplax luteus (McNiell, 1929) and Ceratoplax glaberrima (Haswell, 1881) are illustrated here from type material. The granulation of the cheliped distinguishes these two species but specimens in Museum Victoria and Western Australian Museum can not be easily assigned to one species or the other. Diagnosis. Carapace clearly wider than long, most of dorsal surface polished smooth, with more or less single marginal row of long setae, few shorter setae submarginally. Eyes embedded in orbits, moveable, cornea well formed. Maxilliped 3 merus sometimes produced distolaterally. Ceratoplax glaberrima (Haswell, 1881) (Fig. 147e). Carapace anterolateral margins with 4 obscure lobes. Cheliped palm in male, deep and with granules along upper surface, in rows on outer surface and ridge along lower edge; in female with 2 regular longitudinal rows of granules and close to lower margin an acute granulated ridge. 16 mm. NSW, SA, WA; subtidal. Baker (1907) described the female of C. punctata with a granulate ridge on the lower margin and spinules on the outer face of the cheliped palm; it is said to be a junior synonym of C. glaberrima (Davie, 2002).

457 Marine Decapod Crustacea of Southern Australia

Ceratoplax inermis (Haswell, 1881). Cheliped palm in male ‘finely and closely granulous above, nearly smooth externally, but with a few obscure granules and a low longitudinal ridge near the lower margin,’ in the female ‘finely granular above, on the proximal portion of the outer surface and below, and with a longitudinal row of granules on the outer surface close to the lower border, becoming a well marked entire ridge on the digital portion.’ 23 mm. NSW; subtidal. The diagnoses quotes from Haswell (1881); the chela of the female holotype seems similar to that of C. luteus and the name may be a senior synonym. Ceratoplax luteus (McNiell, 1929) (Fig. 147b, f, g, Pl. 25f). Cheliped palm of female, outer face with few granules over lower half, with granular lower ridge; palm of large males smooth. 21 mm. Qld, NSW; subtidal. The species was reillustrated as Rhizopa gracilipes by Griffin & Campbell (1969).

Heteropilumnus de Man, 1895 Most authors agree that the genus is poorly defined although more than a dozen species have been assigned to it (Davie & Humpherys, 1997). The only southern Australian species is easy to recognise from the ragged dark grey fringe of long setae. Diagnosis. Carapace clearly wider than long, dorsal surface with few or many setae, becoming a dense band of long setae marginally and extending on to front and sides of carapace. Eyes embedded in orbits, moveable, cornea well formed. Maxilliped 3 merus not produced distolaterally. Heteropilumnus fimbriatus (Milne Edwards, 1834) bearded crab (Fig. 147a, Pl. 25e). Anterolateral margins of carapace divided by notches into lobes. Dark dirty grey. 25 mm. Qld, NSW, Vic., Tas., SA; intertidal to 69 m depth (common).

Mertonia Laurie, 1906 The Australian species is one of only two, the other being recorded from Sri Lanka to Japan. Diagnosis. Carapace semicircular, widest at posterolateral angles. Eyes obsolete. Antenna with flagellum thickly setose; maxilliped 3 with merus produced distolaterally. Mertonia integra (Haswell, 1881) (Fig. 147c, h, j). 8.5 mm. NSW (Port Jackson). The only specimen ever collected was redescribed by McNiell (1929).

Xenophthalmodes Richters, 1880 The semicircular carapace characterises this Indo-West Pacific genus of about five species. The Australian species might be easily confused with Mertonia integra. Diagnosis. Carapace semicircular, widest at produced posterolateral angles. Eyes obsolete. Antenna with flagellum slender and naked; maxilliped 3 with merus not produced distolaterally. Xenophthalmodes dolichophallus Tesch, 1918 (Fig. 147d, i). 25 mm. Indonesia, Qld (Moreton Bay), NSW (Botany Bay). Griffin & Campbell (1969) described specimens from Moreton Bay.

References Baker, W.H. 1907. Notes on South Australian decapod Crustacea. Part V. Transactions of the Royal Society of South Australia 31: 173–190, pls 23–25. Castro, P.,Chia, D.G.B., & Ng, P.K.L.1995. On the systematic status of Ceratocarcinus longimanus White, 1847 (Crustacea: Decapoda: Brachyura: Eumedonidae), a crab symbiotic with comatulid crinoids. Raffles Bulletin of Zoology 43: 239–250.

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Chia, D.G.B., & Ng, P.K.L.1998. A revision of Ceratocarcinus White, 1847, and Harrovia Adams & White, 1849 (Crustacea: Decapoda: Eumedonidae), two genera of crabs symbiotic with crinoids. Raffles Bulletin of Zoology 46: 493–563. Chia, D.G.B., & Ng, P.K.L. 2000. A revision of Eumedonus H. Milne Edwards, 1834 and Gonatonotus White, 1847 (Crustacea: Decapoda: Brachyura: Eumedonidae), two genera of crabs symbiotic with sea urchins. Journal of Natural History 34: 15–56. Davie, P.J.F. 1989. A re-appraisal of Heteropanope Stimpson, and Pilumnopeus A. Milne Edwards (Crustacea: Decapoda: Pilumnidae) with descriptions of new species and genera. Memoirs of the Queensland Museum 27: 129–156. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Davie, P.J.F.,& Humpherys, A. 1997. New species of rhizopine crabs (Crustacea: Brachyura) from northern Australia. Memoirs of the Queensland Museum 42: 97–103. Greenwood, J.G., & Fielder, D.F. 1984. The zoeal stages of Pilumneopeus serratifrons (Kinahan, 1856) (Brachyura: Xanthidae) reared under laboratory conditions. Journal of Natural History 18: 31–40. Griffin, D.J.G. 1970. The status of Pilumnus terraereginae Haswell (Crustacea, Decapoda, Xanthidae) and a giant new Pilumnus from Queensland waters. Memoirs of the Queensland Museum 15: 323–330, pls 26–27. Griffin, D.J.G., & Campbell, B.M. 1969. The sublittoral Goneplacidae and Pinnotheridae (Crustacea: Brachyura) of Moreton Bay). Memoirs of the Queensland Museum 15: 141–164. Griffin, D.J.G., & Yaldwyn, J.C. 1971. Port Phillip Bay Survey 2. Brachyura (Crustacea, Decapoda). Memoirs of the National Museum of Victoria 32: 43–63, table 1. Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Hale, H.M. 1931. The post-embryonic development of an Australian xanthid crab (Pilumnus vestitus Haswell). Records of the South Australian Museum 4: 321–331. Haswell, W.A. 1881. On some new Australian Brachyura. Proceedings of the Linnean Society of New South Wales 6: 540–551. McNeill, F.A. 1929. Studies in Australian carcinology. No. 3. Records of the Australian Museum 17: 144–156, pls 35–37. Ng, P.K.L. 1987. The Indo-Pacific Pilumnidae II. A revision of the genus Rhizopa Stimpson, 1858, and the status of the Rhizopinae Stimpson, 1858 (Crustacea, Decapoda, Brachyura). Indo-Malayan Zoology 4: 69–111. Ng, P.K.L., & Chia, D.G.B. 1999. Revision of the genus Zebrida White, 1847 (Crustacea: Decapoda: Brachyura: Eumedonidae). Bulletin of Marine Science 65: 481–495. Ng, P.K.L., & Clark, P.F. 2000. The eumedonid file: a case study of systematic compatibility using larval and adult characters (Crustacea: Decapoda: Brachyura). Invertebrate Reproduction and Development 38: 225–252. Rathbun, M.J. 1923. Report on the crabs obtained by the F.I.S.‘Endeavour’ on the coasts of Queensland, New South Wales, Victoria, South Australia and Tasmania. Report on the Brachyrhyncha, Oxystomata and Dromiacea. Biological Results of the Fishing Experiments carried on by the F.I.S. ‘Endeavour’, 1909–14 5: 95–156, pls 16–42. Suzuki, K., & Takeda, M. 1974. On a parthenopid crab, Zebrida adamsii on the sea urchins from Suruga Bay, with a special reference to their parasitic relations. Bulletin of the National Science Museum, Tokyo, Ser. A (Zoology) 17: 287–296, pl. 1. Takeda, M., & Miyake, S. 1968. Pilumnid crabs of the family Xanthidae from the West Pacific. I. Twenty- three species of the genus Pilumnus, with description of four new species. Zoological Laboratory, Faculty of Agriculture, Kyushu University 1: 1–60.

459 Marine Decapod Crustacea of Southern Australia

Xanthidae MacLeay, 1838 Many compact, black-fingered, cryptic crabs are xanthids, members of a family that possibly contains more species in the Indo-West Pacific than any other. While essentially tropical, Xanthidae are well represented in southern Australia. Some are endemic but many of those recorded are at the southern limit of their ranges. While almost 40 species have been recorded within the region covered by this book, only one species could be considered common. That one is Actaea peronii, one of the most abundant of all crabs on the shallow southern shelf. Others are rarely found or have been reported at the southern limit of coral reef, particularly at Rottnest I. Typically, xanthid crabs are found intertidally or just subtidally, hiding under rocks or in sponges and other sessile invertebrates. Many are passive grazers on algae or feed on detritus. Generally, they continue their cryptic behaviour when disturbed by playing dead. But like all generalities, there are exceptions – some species are active predators (Milke & Kennedy, 2001). In 1968 Holthuis (1968) asked the question ‘are there poisonous crabs?’ The answer now seems to be certainly ‘yes’ but so far only in the tropics and xanthids are the main culprits. Poisoning and death of humans after eating xanthid crabs, especially of the genus Zosimus are well documented (Llewellyn & Davie, 1987; Llewellyn et al., 2002). The Xanthidae have been treated in the past as a large family incorporating many genera now considered part of a wider xanthoid grouping. As in most recent treatments (e.g. Guinot, 1978; Martin, 1988), Xanthidae are here defined more narrowly. Other xanthoid families, Pilumnidae, Eriphiidae, Goneplacidae, Trapeziidae and Domeciidae in southern Australia, are covered sepa- rately. This reappraisal followed the critical revision of Serène (1984). Pilumnids are the most likely to be confused but males of that family have an abdomen with 7 moveable elongate somites. Eriphiids are similar also but eriphiid males have a gonopods of similar lengths. Keys below are based on those in Serène’s monograph and from which most of Davie’s (2002) subfamily diagnoses were derived. Many of the couplets used by Serène (1984) do not have match- ing characters and are difficult to interpret. He used many relative terms, like elongate or short – descriptions that are not always helpful to a novice. Often, this kind of characterisation is difficult to avoid. Serène (1984) did, however, publish valuable photographs of many tropical species. Alain Crosnier, who edited Serène’s volume after his death, noted that his subfamily key was provisional and he had intended other characters be included. Many species have a pattern of regions (areolae), areas of the dorsal carapace, more or less separated by grooves (and labelled with letter–number codes by Serène. These regions are sometimes subdivided or the grooves between them obscure. Generic diagnosis presented here include only what are believed to be the states of diagnostic characters; characters (many) that are ambiguous or unknown are omitted. Many species have been included here on the basis of only a single record over a century or more of collecting by dedicated crab taxonomists employed in museums. It would not be surpris- ing if they were never seen again but, if global warming continues to facilitate the southerly spread of tropical species, they may become more frequent. Equally likely are future discoveries of other species from more northern localities. Davie (2002) catalogued 168 species in 50 genera in eleven subfamilies. All the subfamilies are represented in southern Australia and at least half of the genera.

Diagnosis. Carapace hexagonal, transversely oval or sometimes subcircular; dorsal regions generally well-defined; surface usually ridged or granular. Frontal margin bilobed. Anterolateral margins generally with 2–6 teeth or lobes, sometimes weakly differentiated. Epistome with ridges that define efferent branchial channels absent or confined to the posterior part of the buccal cavern, not continuing on to meet the anterior buccal frame. Thoracic sternum narrow. Male abdomen consisting of 5 moveable somites with somites 3–5 fused (sutures sometimes evident). Male gonopod 1 slender, curved or sinuous, simple apically, typically with long setae distally or subdistally; gonopod 2 less than quarter length of first. (adapted from Davie, 2002)

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Key to subfamilies of Xanthidae Couplets are simplified from those in Serène (1984) with modification to include a new subfamily and revised nomenclature. Atypical taxa are eliminated first. Additional unpaired characters that might be useful are given in parentheses in lieu of diagnoses. 1. Ambulatory legs short, laterally compressed, especially distal articles; dactyli blade-like (carapace dorsal surfaces evenly and moderately convex, regions poorly defined or absent; anterolateral margin strongly convex, serrated, lined with sharp granules or spines; posterolateral margin converging; carapace and leg fringed with long setae) . . Kraussiinae ...p.469 — Ambulatory legs not broadly flattened, cylindrical or carinate at most ...... 2 2. Chelipeds equal, similar, slender; chelae with fingers long, thin, cutting edges armed with pointed teeth for gripping anemones (carapace rounded to quadrate, smooth, regions weakly indicated, sometimes covered with thick coat of setae; sternum elongate for entire length; abdominal somites 1 and 2 visible in dorsal view; telson of male covering only small part ofsternite 4) ...... Polydectinae ...p.470 — Chelae with fingers various, not especially long or thin or armed with pointed teeth . . . . . 3 3. Carapace subcircular or oval, not much broader than long, flat, without distinct regions, naked, more or less granular; lateral margins convex, weakly dentate (chelipeds unequal and dissimilar in both sexes; larger cheliped with short fixed finger, dactylus convex, shorter than width of palm; smaller cheliped with fingers similar, conical, thin apically; in contact throughout length, dactylus as long as width of palm) ...... Cymoinae ...p.466 — Carapace usually broader than long, dorsally convex, margins entire or dentate ...... 4 4. Chelipeds subequal and similar, or sometimes moderately unequal, fingers thin and acute, usually crossing at their extremities; carapace surface with regions well defined; usually thickly setose; buccal cavern narrow and oval or quadrangular (anterolateral and posterolateral margins well delineated; anterolateral margin with 4 lobes or teeth, but may be entire and granular (Calvactaea), last lobe often continued onto posterobranchial region by transverse crest; maxilliped 3 sometimes elongate and pediform) . . Zalasiinae ...p.476 — Cheliped with fingers various, not crossing at their extremities; carapace surface various, rarely with thick tomentum; buccal cavern broad or quadrangular ...... 5 5. Chelipeds with ends of fingers rounded and spoon-shaped; walking legs with dactylo- propodal articulation formed by rounded prolongation of propodal lateral margin sliding against and beneath projecting button situated proximally on lateral margin of dactylus (if not clearly differentiated then dactylus apically bifid) ...... 6 — Chelipeds with ends of fingers acute; walking legs without dactylo-propodal articulation so-modified ...... 7 6. Front wider than one-third maximum width; front typically with 2 large submedian lobes and 2 small lateral lobules, or nearly straight; anterolateral margin typically with 4 teeth behind exorbital angle (sometimes 2 or 3) ...... Chlorodiinae ...p.465 — Front narrower than one-third maximum width; front 2- or 4-lobed, with median fissure, and lateral sinus separating supraorbital angles; 4–8 anterolateral teeth behind exorbital angles ...... Etisinae ...p.468 7. Carapace transversely oval, generally about twice as broad as long; anterolateral margins entire or weakly dentate; chelipeds with palm elongate; fingers elongate and dentate, fixed finger without proximal or median tooth on cutting edge ...... Liomerinae ...p.469 — Carapace usually less than twice as broad as long; anterolateral margins usually dentate; chelipeds with palm and fingers not especially elongate ...... 8

461 Marine Decapod Crustacea of Southern Australia

8. Chelipeds subequal, with carpus, palm and fingers fitting against carapace; walking legs similarly folded into strongly concave posterolateral margins of carapace; sternum often grooved longitudinally, sometimes with cavities (carapace regions strongly marked; anterolateral margin convex, distinctly separated from concave posterolateral margin, not meeting exorbital angle but continuing below orbit) ...... Euxanthinae ...p.468 — Chelipeds and walking legs not fitting against lower carapace ...... 9 9. Carapace granular, tuberculate or spinose, regions well defined and lobulate; anterolateral margins with 3 teeth or lobes behind exorbital angles; front bilobed, hardly projecting; chelipeds and walking legs usually granular or spinose (lateral margin of front with ventral prolongation meeting basal antennal article such that antennal flagellum included within orbit; thoracic sternite 4 with longitudinal furrow; male abdomen locking on to sternite 6 with buttons) ...... Actaeinae ...p.462 — Carapace rarely granular or spinose, regions not or well-defined; 4 or 5 anterolateral teeth (sometimes indistinct or subdivided); front nearly straight or bilobed; chelipeds rarely granular ...... 10 10. Chelipeds with carina on upper margin of palm; walking legs with anterior margins crested, particularly on carpus and propodus; anterolateral margins rounded or crested, typically with 4 lobes or broad teeth behind exorbital angle, but may be obscure . . . . Zosiminae ...p.477 — Chelipeds strong, smooth, granular, tuberculate, or spinous but not carinate; walking legs rounded in section, smooth or granular, rarely carinate; anterolateral margins, typically 4 or 5 teeth or broad lobes, but some may be subdivided into as many as 10 teeth (sternum with fourth sternite large, such that telson covers only proximal half) . . . . . Xanthinae ...p.470

Subfamily Actaeinae Alcock, 1898 Member of this subfamily usually have a granular, tuberculate or spinose carapace and legs with the regions well defined and lobulate.

Key to southern Australian genera and species of Actaeinae 1. Cheliped palms swollen, fingers curved and hook-like, with setae in gape ...... Pseudoliomera helleri — Cheliped palms not swollen, fingers scarcely curved ...... 2 2. Carapace with posterolateral excavation to accommodate walking legs 3–5 ...... Actaeodes tomentosus — Carapace with straight or barely concave posterolateral margins ...... 3 3. Carapace, chelipeds and legs with well-spaced granules and fine short setae; carapace regions weakly defined ...... Forestia abrolhensis — Carapace, chelipeds and legs smooth, tuberculate or spinose, without setae; carapace regions defined by deep grooves ...... Actaea ...4 4. Carapace with plate-like tubercles anteriorly; legs covered with blunt spines . Actaea peronii — Carapace covered with mosaic of coalesced tubercles; legs barely knobbed ...... 5 5. Carapace with few large tubercles barely projecting ...... Actaea calculosa — Carapace with sharpened elevated tubercles; male cheliped with black on fixed finger extending on to palm ...... Actaea carcharias

462 Brachyura – crabs

Actaea De Haan, 1833 About 20 species of Actaea are known (Serène, 1984), all from the Indo-West Pacific and many from tropical Australia. The genus is divided into two groups depending on the nature of the ornamentation, both groups represented in southern Australia (Guinot, 1976). Diagnosis. Carapace with areas well separated, separate by deep grooves, with mosaic-like tubercles or squamiform or nodiform tubercles, without setae; posterolateral margins nearly straight. Chelipeds not swollen, fingers without rigid setae on cutting edges. Walking legs granular or spinose. Male sternites 3 and 4 separated by suture only laterally (Fig. 148a). Male abdominal somite 6 with pair of depressions on posterolateral corners (part of locking mechanism). Actaea calculosa (Milne Edwards, 1834) facetted crab (Fig. 148c, Pl. 26b). Carapace covered with mosaic of coalesced tubercles; anterolateral margin scarcely lobed. Legs barely knobbed. 20 mm. NT, Qld, NSW, Vic., SA, WA; subtidal to at least 55 m depth. Actaea carcharias White, 1848. Carapace with sharpened tubercles all over; anterolateral margin with sharp tubercles, forming 4 obscure lobes. Chelipeds and walking legs with similar sharpened tubercles. 22 mm. Qld, WA; coral reef, subtidal (Guinot, 1976). Actaea peronii Milne Edwards, 1834 thorn-legged crab (Fig. 148d, Pl. 26c, d). Carapace with plate- like tubercles anteriorly; anterolateral margin with 5 rounded lobes. Chelipeds with shiny tubercles. Walking legs covered with numerous long blunt spines. Carapace red or orange, often with patches of white on legs. 20 mm. Qld, NSW, Vic., Tas., SA, WA (south-western coast); subtidal to at least 110 m depth. Actaea peronii is one of the commonest and most colourful small crabs in the sediments of the southern Australian shelf but is rarely seen intertidally or in bays. It is certainly one of the easiest to recognise. A subspecies, Actaea peronii occidentalis Odhner, 1925 was described from south-western WA. Actaeodes Dana, 1851 Guinot (1976) recognised five species in her revision of the genus. Actaeodes is similar to euxan- thine genera in having posterolateral excavations of the carapace accommodating the last three pairs of walking legs. Diagnosis. Carapace with areas well separated, separate by deep grooves, with tubercles; anterolateral margin with 4 short lobes. Carapace with posterolateral excavation to accommodate walking legs 2–4. Chelipeds not swollen, fingers dentate, with teeth interacting. Walking legs granular. Male sternites 3 and 4 separated by continuous groove (fig. 148b). Male abdominal somite 6 with pair of depressions on posterolateral corners (part of locking mechanism). Actaeodes tomentosus (Milne Edwards, 1834). Carapace 1.55 times as wide as long, with felt of fine maroon setae over areas and in grooves. Cheliped fingers strongly curved, with thick, hoof- like, excavated tips. 33 mm. Common in Indo-West Pacific, WA, Qld, NSW (S to Sydney); intertidal reef. Forestia Guinot, 1976 Only three species are known, all from the Indo-West Pacific (Guinot, 1976; Serène, 1984). Diagnosis. Carapace with areas well separated only anteriorly, separated by deep grooves, with small granules, covered with fine short yellow setae. Posterolateral margins of carapace nearly straight. Chelipeds not swollen, fingers without rigid setae on cutting edge. Walking legs granular or spinose. Male sternites 3 and 4 separated by suture only laterally. Male abdominal somite 6 with pair of depressions on posterolateral corners (part of locking mechanism). Forestia abrolhensis (Montgomery, 1931). Carapace behind front and eyes divided by grooves longitudinally into 2 lateral paired areas and median ridge; all areas finely tuberculate. 28 mm. Zanzibar, WA (S to Rottnest I.); shallow subtidal.

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Pseudoliomera Odhner, 1925 Serène (1984) provided a key to eight species of which three occur in Australia. The brushes of setae on the fingers of the chelipeds are diagnostic. Diagnosis. Chelipeds swollen; fingers short, hook-shaped, with brushes of rigid setae near teeth on cutting edge. Walking legs granular. Pseudoliomera helleri (Milne Edwards, 1865) (Fig. 148e). Carapace 1.5 times as wide as long; regions well defined by shallow grooves, granular; anterolateral margins convex, granular, without carinae. Walking legs without carinae; walking leg 1 dactylus without stiff comb of setae. 29 mm. Indo-West Pacific, NT, WA; subtidal reef to 35 m depth.

a b

e c

m n d

k l j

Fig. 148. Xanthidae Actaeinae. Sternites: a, Actaea. b, Actaeodes. c, Actaea calculosa. d, Actaea peronii. e, Pseudoliomera helleri (chela). Chlorodiinae. f, Chlorodiella laevissima. g, Chlorodiella nigra. x, Fronts: h, Cyclodius obscurus. i, Cyclodius ungulatus. j, k, Pilodius areolatus (with left antenna, basal article stippled, and orbit, anterior view). Cymoinae. l, Cymo cerasma. Male pleopod 1: m, Cymo cerasma. n, Cymo andreossyi.

464 Brachyura – crabs

Subfamily Chlorodiinae Dana, 1851 The chlorodiines are a group associated usually with coral reefs but some may be found on rocky environments. The subfamily shares with Etisinae rounded and spoon-shaped cheliped fingers but differs from them the narrower front and fewer frontal and anterolateral lobes. Serène recognised five genera, all occurring in tropical Australia, one albeit under another generic name (Davie, 2002). Records in temperate Australia are few. The key to species is based on Serène’s keys.

Key to southern Australian species of Chlorodiinae 1. Antennal basal article with anterolateral external angle prolonged and in orbital hiatus, excluding antennal flagellum ...... Pilodius areolatus — Antennal basal article without anterolateral external angle prolonged; antennal flagellum free in hiatus ...... 2 2. Carapace dorsally with well defined regions; front half or less greatest carapace width ...... Cyclodius ...3 — Carapace smooth, regions poorly defined; front half or more greatest carapace width ...... Chlorodiella ...4 3. Front almost straight, submedian lobes obscure, external angles obtuse . . Cyclodius obscurus — Front with strong submedian lobes and external angles ...... Cyclodius ungulatus 4. Anterolateral margins of carapace with first tooth often very reduced, second–fourth teeth well developed, blunt lobes in large specimens; walking legs with numerous plumose setae; male gonopod 1 with short recurved distal setae ...... Chlorodiella nigra — Anterolateral margins of carapace with first and fourth teeth small, second and third teeth well developed; walking legs with sparse plumose setae; male gonopod 1 with recurved setae much longer than diameter of gonopod ...... Chlorodiella laevissima

Chlorodiella Rathbun, 1897 Serène (1984) listed and provided a key to nine species, all of which live on coral reefs associated with living corals. One or both of the species listed below may occur on Rottnest I., WA; the differences in spination between them are subtle and may be size-dependent. Diagnosis. Carapace dorsally smooth, regions not defined; front straight, half carapace width or more, not projecting, with weak median notch; weak lateral lobes separated by shallow lateral sinus from orbital angle, with 4 lateral teeth (sometimes obscure or fused). Antennal basal article without anterolateral projection, flagellum free in orbit. Male gonopod 1 with subapical setae directed backwards. Chlorodiella laevissima (Dana, 1852) (Fig. 148f). Anterolateral margins of carapace with first and fourth teeth small, second and third teeth well developed. Walking legs with sparse plumose setae. Male gonopod 1 with recurved setae much longer than diameter of gonopod. 7 mm. Indo-West Pacific, Qld, WA (S to Rottnest I.); coral reef. This and the larger following species are widely reported throughout the Indo-West Pacific. Two subspecies have been described. Chlorodiella nigra (Forskål, 1775) (Fig. 148g). Anterolateral margins of carapace with first tooth often very reduced, second–fourth teeth well developed, blunt lobes in large specimens. Walking legs with numerous plumose setae. Male gonopod 1 with short recurved distal setae. 21 mm. Indo-West Pacific, WA (S to Rottnest I.), NT, Qld, NSW (S to Sydney region); shallow rocky beach or coral reef.

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Cyclodius Dana, 1851 Serène (1984) treated this genus as Phymodius Milne Edwards, 1863, a junior synonym. Two of its six species reach southern Australia. Diagnosis. Carapace with well defined regions; front half or less greatest carapace width, with clear median notch, with 4 lateral teeth. Antennal basal article without anterolateral external angle prolonged; antennal flagellum free in hiatus. Male gonopod 1 with long subapical setae directed distally. Cyclodius obscurus (Jacquinot, 1846) (Fig. 148h). Front with obsolete submedian lobes; external angle obtuse. 27 mm. Indo-West Pacific, WA (S to Rottnest I.); coral and rocky reef, intertidal to shallow subtidal. The species has previously been reported as Phymodius monticulosus (Dana, 1852). Cyclodius ungulatus (Milne Edwards, 1834) (Fig. 148i). Front with open median notch, blunt submedian lobes; external angle strong, separated by concave notch from submedial lobe. 22 mm. Indo-West Pacific, Qld (Great Barrier Reef), Lord Howe I., WA (S to Rottnest I.); coral and rocky reef, intertidal to shallow subtidal.

Pilodius Dana, 1851 Clark & Galil (1993) revised the genus and provided a key to the 15 species they recognised. Many other species previously assigned were removed to other genera. Several species inhabit northern Australia of which Pilodius moranti Clark & Galil, 1993 lives on Lord Howe I. The prolongation of the basal antennal article is the critical character defining the genus (Fig. 148k). Diagnosis. Carapace regions defined; front more than one-third carapace width, hardly projecting, with 2 large submedian lobes and 2 small lateral lobes, with 3 or 4 anterolateral lobes or teeth. Antennal basal article with anterolateral external angle prolonged into orbital hiatus, excluding antennal flagellum. Chelipeds unequal, fingers massive, spoon-shaped. Pilodius areolatus (Milne Edwards, 1834) (Fig. 148j, k). Carapace regions elevated as domes covered with shiny granules separated by dense tomentum, central region tripartite. Anterolateral margin with 4 granular lobes behind exorbital angle. Carapace, chelipeds and limbs with pearly granules; walking legs with well-defined fringes of long setae along anterior and posterior margins. 25 mm. Indo-West Pacific, WA, NT, Qld (including Great Barrier Reef), NSW (S to Sydney region); coral and rocky reef, intertidal to shallow subtidal.

Subfamily Cymoinae Alcock, 1898 Crabs of the only genus, Cymo, are flat, without distinct regions and are symbionts of corals.

Cymo De Haan, 1833 The genus has six species in the Indo-West Pacific, the last described from Western Australia (Morgan, 1990). Both species from southern Australia have a bilobed front, unlike others, and are weakly spinose or tuberculate.

Key to southern Australian species of Cymo 1. Chelipeds with white fingers; male pleopod 1 with simply recurved apex . . Cymo andreossyi — Chelipeds with black fingers; male pleopod 1 with sinuous flattened apex . . . Cymo cerasma

466 Brachyura – crabs

e c b

Fig. 149. Xanthidae Etisinae. a, Etisus sp. b, Etisus electra. Euxanthinae. c, d, e, Euxanthus exsculptus (with upper and lateral views of propodus and dactylus of walking leg). Kraussiinae. f, g, Palapedia pelsartensis (with detail of front). Liomerinae. h, Liomera cinctimana. i, Liomera pallida.

467 Marine Decapod Crustacea of Southern Australia

Cymo andreossyi (Audouin, 1826) white-fingered coral clinger (Fig. 148n). Carapace with bilobed, spinose or tuberculate front. Chelipeds with white fingers. Male pleopod 1 with simply recurved apex. 26 mm. Indo-West Pacific, WA (S to Rottnest I. (Black & Prince, 1983)), Qld, NSW; intertidal and shallow subtidal, between branches of coral. Cymo cerasma Morgan, 1990 (Fig. 148l, m). Carapace with bilobed, spinose or tuberculate front. Chelipeds with black fingers. Male pleopod 1 with sinuous flattened apex. 15 mm. WA (S to Rottnest I.); intertidal to 15 m depth, coral and rock.

Subfamily Etisinae Ortmann, 1893 The combination of spoon-shaped cheliped fingers and specialised leg articulation are similar to Chlorodiinae but the two genera of this subfamily differ in having a narrower front and more anterolateral teeth or lobes. Only one genus reached southern Australia.

Etisus Milne Edwards, 1834 Several of the 15 or so described species have well developed anterolateral teeth or lobes. At least ten species occur in tropical Australia; few individuals have been taken in southern Western Australia. Diagnosis. Carapace feebly convex; frontal margin weakly promoted in front of supra-orbital angles. Buccal cavity not elongated.

Key to southern Australia species of Etisus 1. Front 4-lobed ...... Etisus electra — Front 2-lobed ...... Etisus sp. Etisus electra (Herbst, 1801) (Fig. 149b). Carapace 1.4 times as wide as long; anterolateral margins with 4 simple teeth. Front 4-lobed, submedian lobes separated from lateral lobes by deep notch, all extending well in front of supra-orbital angles. 25 mm. Indo-West Pacific, Qld, WA (S to Rottnest I.); coral reef, intertidal to shallow subtidal. Etisus sp. (Fig. 149a). Front bilobed, straight, separated from supra-orbital lobes angles by narrow groove; anterolateral margin with 4 complex large teeth, each basically tridentate. 28 mm. WA (Fremantle); under rocks, 2 m depth (single specimen, 1971, in Western Australian Museum). The specimen cannot be identified from the key in Serène (1984) or later publications.

Subfamily Euxanthinae Alcock, 1898 The upper surface of the chela and the lower surface of the carapace, each defined by a sharp margin, fit closely. The anterolateral margin is well below the orbits. The posterolateral margin of the carapace is straight or obtuse-triangularly excavated for the walking legs. The carapace regions are particularly obvious. Only one species of about ten known from tropical Australia is represented in the south and then only rarely.

Euxanthus Dana, 1851 Species are highly sculptured and generally shiny, almost like a shelled walnut. Diagnosis. Carapace smooth, strongly convex, without subhepatic cavities; walking legs folded into concave posterolateral margin.

468 Brachyura – crabs

Euxanthus exsculptus (Herbst, 1790) (Fig. 149c–e). Exorbital angle without tooth; anterolateral margin with 5 teeth, first 3 weaker than last 2. 61 mm. Indo-West Pacific, WA (S to Rottnest I.), Qld; coral reef under rubble, intertidal to shallow subtidal.

Subfamily Kraussiinae Ng, 1993 The genus Kraussia was once placed in Atelecyclidae, also roughly circular, but, even since settling in Xanthidae, its affinities have been debated. In a cladistic analysis of Atelecyclidae, Salva & Feldmann (2001) treated Kraussia as one of two genera of Thiidae. While establishing his new family, Ng (1993) moved all eight Australian species except one from Kraussia to his new genus, Palapedia. The flattened walking legs, presumably for digging in sediment, distinguish it and similar more recent genera from other xanthids and converge on those in matutids and calappids.

Palapedia Ng, 1993 Of eight Australian species, four from tropical Western Australia were described by Serène (1972) who provided a key (to Kraussia). Diagnosis. Carapace with anterolateral margin longer than concave posterolateral. Cheliped fingers sharp or flattened. Walking leg 4 less than twice as long as wide. Male abdominal somite 2 2.3 times as wide as long. Palapedia pelsartensis (Serène, 1972) (Fig. 149f). Front 4-lobed, with open median slit, half width of carapace. Major cheliped with swollen palm ornamented with transverse rows of granules, and short fingers. Minor cheliped with tapering fingers, fixed finger bent down. Walking legs with dactyli with straight, carinate, granular anterior borders. 20 mm. WA (Shark Bay to Rottnest I.); subtidal sediments.

Subfamily Liomerinae Sakai, 1976 A broad flattened carapace with broad anterolateral lobes characterise the two genera of liomer- ines, themselves separated by only subtle differences (Serène, 1984).

Liomera Dana, 1851 Liomera is an abundant Indo-West Pacific genus with more than 20 species, of which at least 12 occur in Australia on coral reefs. Most are wider than other xanthids. The only records in southern Australia are from Rottnest I., WA; other species might be expected on its coral reefs and keys in Serène (1984) and Sakai (1976) should be referred to. Diagnosis. Carapace regions convex both laterally and longitudinally.

Key to southern Australian species of Liomera 1. Carapace with weakly marked regions, smooth; to 70 mm ...... Liomera cinctimana — Carapace with well marked regions, finely granulate; to 12 mm ...... Liomera pallida Liomera cinctimana (White, 1847) (Fig. 149h). Carapace with weakly-marked regions, smooth; anterolateral margins with 4 feebly rounded lobes. Chelipeds of male with pigment often encircling propodus. Adults reddish-violet, juveniles with white areas on carapace. 70 mm. Indo-West Pacific, East Pacific, WA (S to Rottnest I.), Qld; coral and rocky reef, intertidal to 35 m depth.

469 Marine Decapod Crustacea of Southern Australia

Liomera pallida Borradaile, 1900 (Fig. 149i). Carapace with well-marked regions, finely granulate; anterolateral margins with 4 obsolete granulate lobes. Chelipeds of male with pigment confined to fingers. Purple with white granulate legs. 12 mm. Indo-West Pacific, WA (S to Rottnest I.); coral and rocky reef, intertidal to shallow subtidal.

Subfamily Polydectinae Dana, 1851 Poydectine xanthids have a carapace about as wide as long and with regions weakly defined. They carry a pair of anemones in their fine chelae and are commonly called boxer crabs. The crab presumably deters predators by threatening with its poisonous mittens. With its chelae so occupied, the crab is unable to catch its own food and may steal that caught by the anemones. Few species are recognised, in two genera, all in the Indo-West Pacific.

Lybia Milne Edwards, 1834 One species, Lybia tessellata, is widespread in the Indo-West Pacific and of interest to divers and keepers of marine aquaria. It not only has an interesting association with anemones and unusual behaviour but is brilliantly coloured. The carapace is divided into reddish and pale polygons by black lines and the legs are transversely striped (Pl. 26a). This species occurs in tropical Australia but another was described from Port Jackson. Diagnosis. Carapace and legs without tomentum (present in Polydectus); without orbital tubercles. Chelipeds elongate. Lybia australiensis (Ward, 1933) (Fig. 150a). Carapace anterolateral margin with prominent lobe; lateral angle rounded. 7 mm. NSW (Port Jackson); subtidal among bryozoans (known only from type specimen collected in 1928).

Subfamily Xanthinae MacLeay, 1838 Members of the Xanthinae are the typical xanthids, best defined by the absence of any of the defining characteristics of other families. The sternum has a large sternite 4, such that the telson covers only the proximal half. The chelipeds are always strong; sometimes subequal and similar, while others are unequal and dissimilar. Like other subfamilies, xanthines are most diverse in the tropics with almost 30 species reported from Australia. Most southern records are rare and limited but the three species of Megametope are southern endemics in a genus reported elsewhere only in New Zealand and Japan. A few specimens in museum collections are identified with uncertainty – the genera to which they have been attributed are included in the key.

Key to southern Australian species of Xanthinae 1. Front strongly and evenly produced in front of eyes ...... Megametope ...2 — Front not strongly produced, straight or irregularly toothed ...... 4 2. Front with straight margin ...... Megametope punctata — Front with convex margin ...... 3 3. Front evenly convex, thin, with small notch near orbit ...... Megametope carinata — Front with median obtuse angle, thickened, without notch near orbit ...... Megametope rotundifrons 4. Larger cheliped with fingers with rounded, enlarged hollow tips not crossing when closed; anterolateral margin of carapace with more than 4 teeth ...... Leptodius exaratus

470 Brachyura – crabs

— Larger cheliped with pointed fingers, barely crossing; anterolateral margin of carapace with 4 or fewer teeth or lobes ...... 5 5. Male chelipeds subequal; male pleopod 1 elongate, with curved row of subdistal long setae on one margin and short setae on other ...... 6 — Male chelipeds subequal or unequal; male pleopod 1 not as above ...... 9 6. Carapace regions of strongly projecting regions separated by deep grooves; 4 anterolateral lobes blunt ...... Neoxanthias impressus — Carapace regions obscure or weakly separated by grooves; 4 anterolateral lobes blunt or some sharp ...... Xanthias ...7 7. Carapace smooth; upper surface of cheliped smooth with longitudinal groove on external surface ...... Xanthias pachydactylus — Carapace and upper–outer surface of chelipeds granular ...... 8 8. Outer surface of cheliped with 1 longitudinal furrow; upper surface with rows of large contiguous tubercles forming a crest ...... Xanthias elegans — Outer surface of cheliped with 3 longitudinal furrows ...... Xanthias lamarcki 9. Carapace regions indistinct, smooth; anterolateral margin with first and second lobes rounded, third and fourth teeth well defined ...... Lachnopodus bidentatus — Carapace regions distinct, smooth or granular; anterolateral margin with 3 or 4 well defined teeth ...... 10 10. Anterolateral margin with 3 teeth ...... Miersiella haswelli — Anterolateral margin with 4 teeth ...... 11 11. Carapace regions smooth; anterolateral margin with last two teeth more acute than first two; chelipeds unequal and dissimilar, smooth globular tubercles on outer face of larger palm, acute spines on smaller palm ...... Paraxanthias notatus — Carapace regions regularly granular; anterolateral margin with second and third teeth bigger than first and fourth; chelipeds dissimilar ...... 12 12. Male chelipeds subequal, with upper margin of carpus and palm granular, granules arranged in irregular rows; first anterolateral tooth smaller than other, submarginal . . . . . Paraxanthodes sp. — Male chelipeds unequal, with upper margin of carpus and palm with small regular granules; first anterolateral tooth not submarginal ...... Nanocassiope sp.

Lachnopodus Stimpson, 1858 One of about six species has been recorded from southern WA. Diagnosis. Carapace regions indistinct, granular or smooth; anterolateral margin with lobes 1 and 2 rounded, teeth 3 and 4 well defined. Male chelipeds subequal, similar. Lachnopodus bidentatus (Milne Edwards, 1867) Carapace smooth; anterolateral margin with lobes 1 and 2 obtuse, lobes 3 and 4 rounded. Walking legs without spines, rugose on lower margins. 21 mm. Indo-West Pacific, WA (south-western coast); subtidal.

Leptodius Milne Edwards, 1863 Most species (Serène, 1984) have been recorded from coral reefs in Australia but only one reaches as far south along the eastern and southern coasts to be included here. Diagnosis. Carapace regions well defined; anterolateral margin of carapace with 4 teeth or irregularly toothed. Chelipeds with fingers having rounded, enlarged hollow tips not crossing when

471 Marine Decapod Crustacea of Southern Australia

closed. Basal article of antenna elongated. Antennular fossa narrow. Maxilliped 3 merus broad, projecting distolaterally. Leptodius exaratus (Milne Edwards, 1834) (Fig. 150b). Carapace 1.5 times as wide as long, regions rugose; anterolateral margin with 4 simple teeth. 24 mm. Indo-West Pacific, WA (south-western coast), NT, Qld, NSW (S to Sydney region); rocky shore, intertidal.

Megametope Filhol, 1886 Megametope is only one of two xanthid genera with endemic species on the southern coast; the other is Actaea. Another species occurs in New Zealand and another in Japan. The strongly produced front and smooth carapace immediately characterise it. Diagnosis. Carapace smooth; front strongly and evenly produced in front of eyes; anterolateral margins with obscure lobes if any. Megametope carinata (Baker, 1907) crested forehead crab (Fig. 150c, Pl. 26e). Front evenly convex, thin, with small notch near orbit; 4 anterolateral lobes divided by small notches. Walking legs crested. 30 mm. SA, WA (south-western coast); shallow subtidal. Megametope punctata Haswell, 1882) (Fig. 150d). Front with straight margin, without notch; anterolateral lobes obscure. 18 mm. NSW (southern coast); shallow subtidal. Megametope rotundifrons (Milne Edwards, 1834) smooth forehead crab (Fig. 150e, Pl. 26f). Front with median obtuse angle, thickened, without notch near orbit; 4 anterolateral lobes divided by small notches. Walking legs rounded in section. 28 mm. Vic. (W of Western Port), SA; shallow subtidal. Lioxantho haswelli Fulton & Grant, 1906 is a junior synonym of this species (cf. Davie, 2002).

Miersiella Guinot, 1967 Davie (2002) included the only species with the Euxanthinae but Xanthinae seems more appropriate. It differs from Nanocassiope only in having fewer anterolateral teeth (Guinot, 1967). There is only one species. Diagnosis. Carapace flat, regions distinct, granular; front broad, with slight median notch; anterolateral margin with 3 teeth. Chelipeds unequal, elongated; with fingers crossing, with small triangular teeth. Miersiella haswelli (Miers, 1886) (Fig. 150f). 9 mm. Indian Ocean, Philippines, Japan, New Caledonia, Loyalty Is, NSW (S to Eden); continental shelf, 80–527 m depth.

Nanocassiope Guinot, 1967 A single specimen identified to this genus has been recorded in Western Australia. The genus is similar to Miersiella but has four anterolateral teeth, the second and third bigger than the first and fourth. N. alcocki (Rathbun, 1902) occurs in the Timor Sea. Nanocassiope sp. WA (off Rottnest I.); 182 m depth.

Neoxanthias Ward, 1933 Three species are known in this essentially tropical genus; one reaches southern WA, but rarely. Diagnosis. Carapace regions of strongly projecting, regions separated by deep grooves, especially obvious heart-shaped region in centre; frontal lobes oblique, excavate; anterolateral margin of 4 blunt lobes. Chelipeds subequal, with pointed fingers, barely crossing. Male pleopod 1 elongate, with curved row of subdistal long setae on one margin and short setae on other.

472 Brachyura – crabs

d c

h g

Fig. 150. Xanthidae Polydectinae. a, Lybia australiensis. Xanthinae. b, Leptodius exaratus. c, Megametope carinata. d, Megametope punctata. e, Megametope rotundifrons. f, Miersiella haswelli. g, Neoxanthias impressus. h, Paraxanthodes cumatodes.

473 Marine Decapod Crustacea of Southern Australia

Neoxanthias impressus (Lamarck, 1818) (Fig. 150g). Carapace regions, chelipeds and walking legs smooth; heart-shaped central region with anteriorly directed submedial lobes. 66 mm. Indo- West Pacific, Qld, WA; coral reef, intertidal to shallow subtidal.

Paraxanthias Odhner, 1925 The genus is monotypic. Diagnosis. Carapace regions smooth; anterolateral margin with 2 posterior teeth more acute than 2 anterior teeth; chelipeds unequal and dissimilar, smooth globular tubercles on outer face of larger palm, acute spines on smaller palm. Paraxanthias notatus (Dana, 1852). Indo-West Pacific, Qld, NSW; coral reef, intertidal to 35 m depth. Specimens in the Australian Museum from southern NSW have been identified as this species.

Paraxanthodes Guinot, 1967 The submarginal first anterolateral tooth is diagnostic. One species lives in Japan and another in the Persian Gulf. Museum collections contain material from southern Australia attributed to both species. Diagnosis. Carapace regions regularly granular; anterolateral margin with first tooth smaller than others, submarginal, second and third teeth bigger than first and fourth. Chelipeds subequal, with upper margin of carpus and palm granular, granules arranged in irregular row. Paraxanthodes cumatodes MacGilchrist, 1905 (Fig. 150h). Front straight, hidden in dorsal view by parallel granular row; 2 small blunt spines under first anterolateral tooth. 8 mm. Persian Gulf, WA (off Lancelin); 183 m depth. Paraxanthodes obtusidens Sakai, 1965. Front straight, not hidden in dorsal view; granular crest under first anterolateral tooth. 43 mm. Japan, NSW (unconfirmed); 15–80 m depth.

Xanthias Rathbun, 1897 Xanthias is not clearly defined, especially from Neoxanthias, but nevertheless is home to at least 15 species (Serène, 1984). The colour photo is of an undetermined species from NSW (Pl. 26g). Diagnosis. Carapace regions obscure or weakly separated by grooves; front bilobed, each lobe indistinctly notched near inner orbital tooth, not strongly produced; 4 anterolateral lobes blunt, or some sharp. Chelipeds of male unequal, with pointed fingers, barely crossing. Male pleopod 1 elongate, with curved row of subdistal long setae on one margin and short setae on other. Xanthias elegans (Stimpson, 1858) (Fig. 151a). Carapace granular. Outer surface of cheliped with 1 longitudinal furrow; upper surface with rows of large contiguous tubercles forming a crest. 13 mm. Indo-West Pacific, WA (S to Rottnest I.), Qld, NSW (S to Port Jackson); coral and rocky reefs, intertidal to 35 m depth. Xanthias lamarcki (Milne Edwards, 1834). Carapace granular. Outer surface of cheliped with 3 longitudinal furrows. 23 mm. Indo-West Pacific, WA, ?NSW; coral reef, intertidal to shallow subtidal. Xanthias pachydactylus (Milne Edwards, 1867). (Fig. 151b, c). Carapace smooth. Upper surface of cheliped smooth with longitudinal groove on external surface. 19 mm. Indo-West Pacific, Qld, WA.

474 Brachyura – crabs

h j

Fig. 151. Xanthidae Xanthinae. a, Xanthias elegans. b, c, Xanthias pachydactylus (with chela). Zalasiinae. d, Banareia villosa. e, Calvactea tumida. f, g, Zalasius australis (with lateral view of front). Zosiminae. h, Atergatis floridus. i, Lophozozymus pictor. j, Lophozozymus pulchellus.

475 Marine Decapod Crustacea of Southern Australia

Subfamily Zalasiinae Serène, 1968 Members of this subfamily are best recognised by the thick coating of hairs over the carapace which is strongly domed. In some literature it is called Trichiinae (Serène, 1984).

Key to southern Australian species of Zalasiinae 1. Carapace without obvious regions, minutely granular or smooth; anterolateral margins and front continuously convex ...... Calvactaea tumida — Carapace with regions more or less obvious, granulate; anterolateral margins rectangular or toothed ...... 2 2. Carapace with regions separated by deep grooves, covered with many shiny tubercles, with dense long setae especially in grooves ...... Zalasius australis — Carapace with regions weakly separated, covered with few granules, especially laterally, with felt of short setae ...... Banareia villosa

Banareia Milne Edwards, 1869 Serène (1984) provided a key for 15 species of Banareia, a genus of crabs characteristically covered with dense long setae. All species are rare, apparently rather immobile, and found in cavities or at the base of colonies of alcyonarians or corals. Diagnosis. Carapace convex, granular, covered with dense long setae. Anterolateral margin with 4 lobes, sometimes weakly marked, always granular. Banareia villosa Rathbun, 1906 (Fig. 151d). Carapace regions weakly separated; anterolateral margins granular, largely parallel, with strong first tooth, smaller second tooth, and tubercle between anterolateral and posterolateral margins. 9 mm. Hawaii, NSW; shelf. Australian material may belong to a similar undescribed species (Davie, 2002).

Calvactaea Ward, 1933 There is a single species, highly sculptured. Diagnosis. Carapace with indistinct regions, strongly globose, finely and regularly granular, covered with felt of short setae. Anterolateral margin evenly curved, granular. Calvactaea tumida Ward, 1933 (Fig. 151e). 23 mm. Japan, Qld, NSW (S to Port Jackson), WA; inside cavities in the walls of, and climbing between, alcyonarians (soft corals), 10–90 m depth.

Zalasius Rathbun, 1897 The genus was previously known as Trichia, a preoccupied name (Holthuis, 1996). Six species occur throughout the Indo-West Pacific, three in Australia (Guinot, 1976; Sakai, 1976). Diagnosis. Carapace with regions deeply divided, strongly globose, granular, covered with felt of short setae. Anterolateral margin produced. Buccal cavity narrow, oval. Zalasius australis (Baker, 1936) long-haired crab (Fig. 151f, g). Carapace 1.1 times as wide as long; anterolateral lobes obtusely triangular, granular; triangular lobe between anterolateral and posterolateral margins. 18 mm. SA; shelf. This rarely taken species was figured as Trichia dromiaeformis australis in Parthenopidae (by Hale, 1927) and in Banareia (by Serène, 1984).

476 Brachyura – crabs

Subfamily Zosiminae Alcock, 1898 Crabs of this subfamily are distinguished from other xanthids in having carinae or crests on the upper margin of the cheliped palms and flattened crested walking legs, and sometimes with crests along the margins of the carapace. Serène (1984) recognised seven genera, at least one, Zosimus, containing species of tropical poisonous crabs.

Key to south Australian species of Zosiminae 1. Carapace with anterolateral margins smoothly rounded, crested, separated from posterior margin by blunt corner ...... Atergatis floridus — Carapace with 2 lobes and 2 triangular teeth on anterolateral margins ...... 2 2. Anterolateral margin with first rounded lobe separated from orbital angle by deep gap; cheliped palm with strong carina ...... Lophozozymus pictor — Anterolateral margin with first lobe confluent with orbital angle; cheliped palm without carina ...... Lophozozymus pulchellus

Atergatis De Haan, 1835 Eleven species were listed by Serène (1984). All live below low water on coral reefs or rocky substrates in the Indo-West Pacific. Some are associated with corals. Only one species extends its range far enough south in Australia to appear at Rottnest Island. Diagnosis. Carapace dorsally smooth or punctate, convex, transversely semicircular or oval; anterolateral margin with crest, not or weakly emarginate. Walking legs short, flattened, with crests on upper and lower margins of carpus and propodus at least. Atergatis floridus (Linnaeus, 1767) (Fig. 151h). Carapace smooth, 1.8 times as wide as long, with weak tooth and furrow at the lateral angle. Front sinuous, with submedian lobes separated by deep sinus. Crimson red or red mottled with yellow. 51 mm. Indo-West Pacific, WA (S to Rottnest I.), NT, Qld, possibly to NSW; coral reef, intertidal to shallow subtidal.

Lophozozymus Milne Edwards, 1863 Serène (1984) listed 13 species from various parts of the Indo-West Pacific and figured some. There is some variation in the expression of the four lobes and teeth on the anterolateral margin of the carapace. Species inhabit rocky bottoms or coral debris down to 250 m depth. At least five species occur in northern Australia. Diagnosis. Carapace dorsally smooth or areolated, convex, transversely oval; anterolateral margin with 2 broad anterior lobes and 2 posterior carinate teeth. Walking legs with crests on upper margin of carpus and propodus at least. Lophozozymus pictor (Fabricius, 1798) (Fig. 151i). Carapace dorsally weakly lobate, smooth. Anterolateral margin with first rounded lobe separated from orbital angle by deep gap, second lobe subtriangular, third lobe triangular, last triangular and smallest. Cheliped palm with strong carina. Carapace with reticulated pattern of dull red lines. 80 mm. Indo-West Pacific, WA (S to Rottnest I.); 0–30 m depth. Lophozozymus pulchellus Milne Edwards, 1867 (Fig. 151j). Carapace dorsally weakly lobate, smooth. Anterolateral margin with first lobe confluent with orbital angle, second to fourth lobes triangular. Cheliped palm without carina. Carapace patterned in brick red and paler shades. 25 mm. Indo-West Pacific, WA (S to Rottnest I.); coral reef, shallow subtidal.

477 Marine Decapod Crustacea of Southern Australia

References Black, R., & Prince, J. 1983. Fauna associated with the coral Pocillopora damicornis at the southern limit of its distribution in Western Australia. Journal of Biogeography 10: 135–152. Clark, P.F., & Galil, B. 1993. A revision of the xanthid genus Pilodius Dana, 1851 (Crustacea: Brachyura: Xanthoidea). Journal of Natural History 97: 1119–1206. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Guinot, D. 1958. Sur une collection de Décapodes Brachyoures (Portunidae et Xanthidae) d l’ile Mayotte. II. Xanthidae (suite). Bulletin du Muséum National d’Histoire Naturelle, Paris (Ser. 2) 30: 175–183. Guinot, D. 1967. Recherches préliminaires sur les groupements naturels chez les Crustacés décapodes brachyoures. II. Les anciens genres Micropanope Stimpson et Medaeus Dana. Bulletin du Muséum National d’Histoire Naturelle, Paris 39: 345–374. Guinot, D. 1976. Constitution de quelques groupes naturels chez les Crustacés Décapodes Brachyoures. I. La superfamille des Bellioidea et trois sous-familles de Xanthoidae (Polydectinae Dana, Trichiinae de Haan, Actaeinae Alcock). Mémoires du Muséum National d’Histoire Naturelle, Paris, ser. A (Zoologie) 97: 1–308, pls 1–19. Guinot, D. 1978. Principes d’une classification évolutive des Crustacés Décapodes Brachyoures. Bulletin Biologique de la France et de la Belgique 112: 211–292. Hale, H.M. 1927. The crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Holthuis, L.B. 1968. Are there poisonous crabs? Crustaceana 15: 216–222. Holthuis, L.B. 1996. The nomenclatural status of the generic names Trichia De Haan, 1839, and Zalasius Rathbun, 1897 (Decapoda, Brachyura). Crustaceana 69: 122–124. Llewellyn, L.E., & Davie, P.J.F. 1987. Crabs and other crustaceans. Pp. 126–135 in: Covacivech, J., Davie, P.,& Pearn, J. (eds), Toxic plants and animals: a guide for Australia. Queensland Museum: Brisbane. Llewellyn, L.E., Dodd.M.J., Robertson, A., Ericson, G., de Koning, C., & Negri, A.P. 2002. Post-mortem analysis of samples from a human victim of a fatal poisoning caused by the xanthid crab, Zosimus aeneus. Toxicon 40: 1463–1469. Martin, J.W. 1988. Phylogenetic significance of the brachyuran megalopa: evidence from the Xanthidae. Symposia of the Zoological Society of London 59: 69–102. Milke, L.M., & Kennedy, V.S. 2001. Mud crabs (Xanthidae) in Chesapeake Bay: claw characteristics and predation on epifaunal bivalves. Invertebrate Biology 120: 67–77. Morgan, G.J. 1990. A collection of Thalassinidea, Anomura and Brachyura (Crustacea: Decapoda) from the Kimberley region of northwestern Australia. Zoologische Verhandelingen, Leiden 265: 1–90. Ng, P.K.L. 1993. Krausiinae, a new subfamily for the genera Kraussia Dana, 1852, Palpedia, new genus, and Garthasia new genus (Crustacea: Decapoda: Brachyura: Xanthidae), with descriptions of two new species from Singapore and the Philippines. Raffles Bulletin of Zoology 41: 133–157. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates. Salva, E.W., & Feldmann, R.M. 2001. Reevaluation of the family Atelecyclidae (Decapoda: Brachyura). Kirtlandia 52: 9–62. Serène, R. 1972. Observations on the Indo-Pacific species of Kraussia Dana 1852 (Decapoda: Brachyura). Journal of the Royal Society of Western Australia 55: 41–62. Serène, R. 1984. Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par Crosnier, A. : Carpiliidae et Menippidae. Faune Tropicale 24: 1–400. Ward, M. 1933. New genera and species of marine Decapoda Brachyura from the coasts of New South Wales and Queensland. Australian Zoologist 7: 377–394, pls 21–23.

478 Brachyura – crabs

Trapeziidae Miers, 1886 Trapeziids are small colourful associates of corals. Most species take shelter between the branches of branching reef-building corals, typically of the family Pocilloporidae, while others live with soft corals. The crabs live in unstable male-female pairs, feeding on the coral mucus or zooplankton (Castro, 1988; Patton, 1994). Their smooth colourful carapace, long chelipeds and association with corals distinguish the family. Serène (1984) provided the definitive study and keys to the family, recognising two subfamilies and seven genera. Most inhabit the tropical Indo-West Pacific and a few in the eastern Pacific. The second of his subfamilies, Domeciinae, was elevated to family rank by Davie (2002) and its only southern Australian species treated under Domeciidae. The close association between coral crabs and coral reefs means that the family is not as well represented in southern Australia as it is in the tropics. Of many species found on the Great Barrier Reef, only two were recorded as far south as temperate Australia by Davie (2002) but collections from Rottnest I., WA, in the Western Australian Museum include others associated with the coral Pocillopora damicornis at the southern end of its range (Black & Prince, 1983).

Diagnosis. Carapace flattened, subquadrilateral, pentagonal or transversely oval; dorsal surface smooth or finely granular, regions not or feebly marked. Front almost straight; 4- or 2-toothed. Basal antennal article slender. Antennules folding transversely. Eyes exposed, not well protected by orbits. Crests of endostome defining efferent channels well developed. Maxilliped 3 with merus shorter than ischium. Maxilliped 2 with endopod of 4 articles. Chelipeds long, meri projecting beyond carapace when folded; slightly unequal, palm smooth or finely granular. Walking legs with dactylo-propodal articulation formed by a rounded prolongation of propodal lateral margin sliding against and beneath a projecting button situated proximally on lateral margin of dactylus. (adapted from Davie, 2002)

Key to southern Australian species of Trapeziidae 1. Chelipeds unequal, larger propodus with seta-filled depression on upper proximal surface; front weakly convex, separated from intraorbital angle by indistinct furrow . . Tetralia cinctipes — Chelipeds equal, with upper margin of merus dentate; front, divided into 4 lobes, separated from intraorbital angle by conspicuous furrow...... Trapezia ...2 2. Upper and outer border of chelipeds with tomentum of long setae; chela with keel-like upper edge ...... Trapezia cymodoce — Chelipeds without tomentum of long setae along entire length; chela upper edge rounded . . 3 3. Carapace without conspicuous dorsal pattern; dark brown ...... Trapezia digitalis — Carapace and chelipeds with honeycomb-like dorsal pattern of red-brown lines ...... Trapezia septata

Tetralia Dana, 1851 Two dissimilar chelipeds, the larger with a pit filled with setae, characterise the seven known species of Tetralia (Galil, 1986a). Diagnosis. Carapace trapezoidal, lateral spines at about or posterior to midpoint; front weakly convex, finely denticulate, and separated from intraorbital angle by indistinct antennal furrow. Basal antennal article with lateral prolongation closing orbit. Chelipeds similar but unequal, larger propodus with seta-filled depression on upper proximal surface. Male abdomen with 7 free somites.

479 Marine Decapod Crustacea of Southern Australia

c b

e f d

Fig. 152. Trapeziidae. Carapace and chelipeds: a, Tetralia cinctipes (larger cheliped only). b, Trapezia cymodoce. c, Trapezia digitalis. d, Trapezia septata. Domeciidae. e, f, Domecia hispida (with maxilliped 3).

Tetralia cinctipes Paul’son, 1875. (Fig. 152a). Frontal margin denticulate. Chelipeds with many granules. Carapace light brown, frontal border red-brown with thin brown band; chelipeds darker on upper surface, with spots; pereopods banded (Jones & Morgan, 2002). 9 mm. Indo- West Pacific, WA (S to Rottnest I.), Qld; coral reef, subtidal, commensal with species of the coral genus Acropora. The species has been commonly misidentified as T. glaberrima (Galil, 1986b).

Trapezia Latreille, 1828 Trapezia is the largest genus with more than 20 species, several illustrated in colour by Castro (1997) and in picture books on coral reefs. Diagnosis. Carapace trapezoidal, lateral spines at about or posterior to midpoint; front sinuous, divided into 4 lobes or teeth, and separated from intraorbital angle by conspicuous antennal furrow. Basal antennal article straight, excluded from closed orbit. Chelipeds similar in form, equal or barely unequal; with upper margin of merus dentate. Male abdomen somites 3–5 fused. Trapezia cymodoce (Herbst, 1801) (Fig. 152b). Upper and outer border of chelipeds with tomentum of long setae along entire length; chela with keel-like upper edge. Purplish-blue with 2 transverse rows of orange dots radiating across anterior half from larger dots in centre; chelipeds orange. 20 mm. WA (S to Rottnest I.); subtidal, coral reef, symbiotic with pocilloporid corals. Trapezia cymodoce is close to T. bidentata (Forskål, 1775), often referred to as T. ferruginea Latreille, 1828, a species that might also occur on Rottnest I. Both are large species, the latter differing in being uniformly orange, having a tomentum only on the carpus, chela with rounded upper margin, etc. (Castro, 1997).

480 Brachyura – crabs

Trapezia digitalis Latreille, 1828 (Fig. 152c). Chelipeds without tomentum of long setae along entire length; chela upper edge rounded. Carapace without conspicuous dorsal pattern; dark brown with a light brown line along anterior border of carapace and cheliped merus. 12 mm. Indo-Pacific, WA (S to Rottnest I.), NT, Qld (Great Barrier Reef); subtidal, coral reef, symbiotic with pocilloporid corals. Trapezia septata Dana, 1852 (Fig. 152d). Chelipeds orange. without tomentum of long setae along entire length; chela upper edge rounded. Carapace and chelipeds with honeycomb-like dorsal pattern of red-brown lines on cream to pink background. 17 mm. Indo-West Pacific, WA (S to Rottnest I.), NT, Qld (Great Barrier Reef); subtidal, coral reef, symbiotic with pocilloporid corals. The colour pattern is distinctive (for colour photos see Healy & Yaldwyn, 1970; Jones & Morgan, 2002). The species has been reported widely as T. areolata.

References Black, R., & Prince, J. 1983. Fauna associated with the coral Pocillopora damicornis at the southern limit of its distribution in Western Australia. Journal of Biogeography 10: 135–152. Castro, P. 1988. Animal symbioses in coral reef communities: a review. Symbiosis 5: 161–184. Castro, P. 1997. Trapeziid crabs (Brachyura: Xanthoidea: Trapeziidae) of New Caledonia, eastern Australia, and the Coral Sea. Pp. 59–107 in: Richer de Forges, B. (ed.) Les fonds meubles des lagons de Nouvelle-Calédonie (sédimentologie, benthos). ORSTOM Editions: Paris. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Galil, B. 1986a. Tetraloides – a new genus of coral-inhabiting crabs. Crustaceana 50: 68–77. Galil, B. 1986b. On the identity of Tetralia cinctipes Paulson, 1975 (Decapoda, Brachyura). Crustaceana 51: 97–102. Healy, A., & Yaldwyn, J.C. 1970. Australian Crustaceans in Colour. Reed: Sydney. 112 pp. Jones, D.S., & Morgan, G.J. 2002. A Field Guide to Crustaceans of Australian Waters. Reed New Holland: Sydney. 224 pp. 2nd edn. Patton, W.K. 1994. Distribution and ecology of animals associated with branching corals (Acropora spp.) from the Great Barrier Reef, Australia. Bulletin of Marine Science 55: 193–211. Serène, R. 1984. Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par Crosnier, A. : Carpiliidae et Menippidae. Faune Tropicale 24: 1–400.

Domeciidae Ortmann, 1893 Domeciids are small crabs associated with corals, typically residents of Acroporidae and Pocilloporidae. Many publications treat them as a members of a subfamily of Trapeziidae, others as xanthids, but Davie (2002) chose to give them family rank. They differ from trapeziids in not being so flattened, having a bilobed rather than 4-lobed front, having spinose or granular chelipeds and specialised maxillipeds 2 and 3. Most are confined to the Indo-Pacific but the species dealt with below is found in the western Atlantic also. Only one of the four genera, Domecia, inhabits Australia. Diagnosis. Carapace with rounded front, tapering posteriorly, not flattened; regions not marked. Front bilobed. Antennules folding transversely. Maxilliped 3 with merus very short, much broader than long. Maxilliped 2 with endopod having propodus and dactylus fused as large endite. Cheliped with merus short; palm granular or spinose. Walking legs with dactylo-propodal articulation formed by a rounded prolongation of propodal lateral margin sliding against and beneath a projecting button situated proximally on lateral margin of dactylus; dactylus claw-like. (adapted from Davie, 2002)

481 Marine Decapod Crustacea of Southern Australia

Domecia Eydoux & Souleyet, 1842 The genus differs from other domeciids principally in having a spinose front, whereas in others it is smooth or denticulate (Galil & Takeda, 1986). There are three species (Guinot, 1964). The crabs favour species of Acropora with flat plates but do live in other genera and even in sponges. The crab induces the coral to grow so that a crevice or pit is formed in which it lives (Patton, 1967). Although the crab prefers to remain hidden, it can move between coral hosts. Diagnosis. Carapace with front and anterolateral margins spinose. Chelipeds and walking legs spinose. Domecia hispida Eydoux & Souleyet, 1842 (Fig. 152e, f). Front with 1 or 2 spines behind margin. Walking leg 4 merus spinose along length. Sternite sharply pointed anteriorly. Patches of purple, especially on anterior carapace and chelipeds; stripes across legs. 9 mm. Indo-Pacific, WA (S to Rottnest I.), NSW; intertidal and subtidal coral (Black & Prince, 1983).

References Black, R., & Prince, J. 1983. Fauna associated with the coral Pocillopora damicornis at the southern limit of its distribution in Western Australia. Journal of Biogeography 10: 135–152. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Galil, B., & Takeda, M. 1986. Resurrection of the genus Jonesius and establishment of a new genus: commensal crabs associated with corals from the Indo-Pacific Ocean. Bulletin of the National Science Museum, Tokyo, Ser. A (Zoology) 12: 163–171. Guinot, D. 1964. Les trois espèces du genre Domecia (Decapoda: Brachyura): D. hispida Eydoux & Souleyet, D. glabra Alcock et D. acanthophora (Desbonne & Schram). Crustaceana 7: 267–283. Patton, W.K. 1967. Studies on Domecia acanthophora, a commensal crab from Puerto Rico, with particular reference to modifications of the coral host and feeding habits. Biological Bulletin 132: 56–67.

Cryptochiridae Paul’son, 1875 The coral gall crabs are a family of unusual crabs, obligate associates of scleractinian corals. They live in cavities, tunnels or pits, in the coral where they feed on mucus and the living coral tissues. Evidence of the presence of crabs is usually a swollen area of the coral skeleton in which there is a small opening providing some ventilation. Females of the largest species grows to only 6 mm. But males are smaller and look more like large megalopas or juveniles. In some genera, the male and female live trapped together but in others the male can wander about the surface of the coral. Cryptochirids are naturally more diverse in the tropics along with the corals in which they live, especially in the western Pacific. Only one of the four species reported from Australia has been recorded as far south as southern Australia, and that rarely. In some older literature the family was erroneously called Hapalocarcinidae. Essential contributions to the systematics of the family have been by Roy Kropp and others (Kropp & Manning, 1987, 1996; Kropp, 1990, 1994). Diagnosis. Carapace narrow, oblong, front more or less truncate, without central tooth. Mandibular palp absent, mandibular incisor lacking processes. Maxilla 1 consisting of undivided body and exopod. Maxilla 2 reduced to finger like structure. Maxilliped 3 with reduced exopod or absent, merus narrower than ischium. Chelipeds small. Gills absent from walking legs 4 and 5. Adult female larger than male, with paired pleopods 2–4, with abdomen expanded posteriorly to form egg chamber, visible in dorsal view. (adapted from Kropp & Manning, 1987)

482 Brachyura – crabs

Cryptochirus Heller, 1861 The genus was diagnosed in detail by Kropp (1990). The only species to occur in southern Australia is widespread in the Indo-West Pacific. Diagnosis. Carapace deflected anteriorly, with clusters of rounded tubercles in middle region. Anterior extension of sternite of cheliped granulate. Walking leg 1 without a distomesial expansion. Cryptochirus coralliodytes Heller, 1861 (Fig. 153). Female 6.5 mm. Indo-West Pacific, Qld, (Great Barrier Reef), NSW (S to Sydney); coral reefs to 30 m depth.

Fig. 153. Cryptochiridae. Cryptochirus coralliodytes.

References Kropp, R.K. 1988. The status of Cryptochirus coralliodytes Heller and Lithoscapthusu paradoxus Milne Edwards (Brachyura: Cryptochiridae). Proceedings of the Biological Society of Washington 101: 872–882. Kropp, R.K. 1990. Revision of the genera of gall crabs (Crustacea: Cryptochiridae) occurring in the Pacific Ocean. Pacific Science 44: 417–448. Kropp, R.K. 1994. The gall crabs (Crustacea: Decapoda: Brachyura: Cryptochiridae) of the Rumphius Expeditions revisited, with descriptions of three new species. Raffles Bulletin of Zoology 42: 521–538. Kropp, R.K., & Manning, R.B. 1987. The Atlantic gall crabs, family Cryptochiridae (Crustacea: Decapoda: Brachyura). Smithsonian Contributions to Zoology 462: 1–21. Kropp, R.K., & Manning, R.B. 1996. Crustacea Decapoda: two new genera and species of deep water gall crabs from the Indo-West Pacific (Cryptochiridae). In: Crosnier, A. (ed.), Résultats des Campagnes MUSORSTOM, Vol. 15. Mémoires du Muséum National d’Histoire Naturelle, Paris 168: 531–539.

483 Marine Decapod Crustacea of Southern Australia

Subsection Thoracotremata Guinot, 1977 The thoracotreme crabs have sternal male and female genital openings. The families appear in an order that best reflects relationships (Martin & Davis, 2001).

Pinnotheridae De Haan, 1833 Pea crabs are specialised crabs, often found living insides the cavity of bivalve molluscs. Bivalves are not the only associates of pinnotherids – other molluscs (Geiger & Martin, 1999), holothurians (Takeda et al., 1997; Hamel et al., 1999; Ng & Manning, 2003), echinoids (Campos & Solís-Marín, 1998) and tube-dwelling chaetopterid polychaetes (Grove et al., 2000) all play host to pinnotherid crabs. Not all pinnotherids are obligate associates; free-living species exist. The exact nature of the association between pea crabs and their hosts was reviewed by Cheng (1967) and research has continued. While the relationship has been viewed as symbiotic or commensal, the benefits for the partners are not clear. It is certainly true that the crab derives considerable protection from predation by living in a shell or a holothurian gut or worm tube and it is probable that it gets some nutrient from mucus and the food trapped by the host as well. The crabs are also know to feed on the gill tissue of host bivalves. A consequence for the host mussel or oyster is reproductive stress resulting in reduced numbers of gametes (Bologna & Heck, 2000). In at least one bivalve species with a life-span of less than one year, crabs must reach maturity quickly and find an alternative host or become free-living for part of the year (Hseuh, 2003). On the other hand, one species of pea crab induces the atrophy of one of the respiratory trees of its holothurian host and is thought to be an obligate permanent parasite (Hamel et al., 1999). One of the consequences of these associations is coordinated life cycles, at least in molluscs (Soong, 1997). The Pinnotheridae comprise about 250 species in numerous genera currently being revised (e.g. Campos, 1996; Campos et al., 1998; Ng & Manning, 2003). The Australian fauna, especially in the tropics, is more diverse than the currently listed 13 species would indicate (Ahyong & Brown, 2003).

Diagnosis. Carapace typically more or less round or transversely oval, often poorly calcified especially in commensal females; anterolateral margins smooth or minutely toothed; front narrow; orbits and eyes very small; cornea sometimes obsolescent. Antennules and antennae usually very small. Interantennular septum reduced to a thin plate, or absent. Buccal cavern broad, short, usually semicircular. Maxilliped 3 with merus often very large; ischium usually small, sometimes completely lost or fused with merus; exopod small, more or less concealed. (from Davie, 2002)

Key to genera and species of southern Australian Pinnotheridae 1. Maxilliped 3 palp of 2 articles ...... Ostracotheres . . . 2 — Maxilliped 3 palp of 3 articles ...... 3 2. Carapace almost circular, longer than wide ...... Ostracotheres holothuriensis — Carapace subrectangular, posterolateral angles pronounced . . . . . Ostracotheres subglobosus 3. Last article of maxilliped 3 palp styliform and attached medially . . . . . Pinnotheres hickmani — Last article of maxilliped 3 palp small and terminal ...... Orthotheres haliotidis

Orthotheres Sakai, 1969 The six known species of Orthotheres are all associated with gastropod mollusc hosts, usually in pairs (Geiger & Martin, 1999).

484 Brachyura – crabs

c b d

efg

Fig. 154. Pinnotheridae. a, Pinnotheres hickmani (female). Carapace of female: b, Orthotheres haliotidis. c, Ostracotheres holothuriensis. d, Ostracotheres subglobosus. Maxilliped 3: e, Ostracotheres holothuriensis. f, Orthotheres haliotidis. g, Pinnotheres hickmani.

Diagnosis. Carapace of female considerably wider than long. Maxilliped 3 palp of 3 articles, last article small and terminal. Walking leg 4 more slender that 1–3, with dactyli short and sharply hooked. Orthotheres haliotidis Geiger & Martin, 1999 (Fig. 154b, f). WA (Fremantle), Qld (Great Barrier Reef, Port Denison); in mantle cavity of abalones Haliotis asinina, H. coccoradiata, H. squamata (Geiger & Martin, 1999). The species has been incorrectly referred to Pinnixia faba (Ahyong & Brown, 2003).

485 Marine Decapod Crustacea of Southern Australia

Ostracotheres Milne Edwards, 1853 Ostracotheres is one of six genera of pinnotherids with a 2–articled palp on maxilliped 3 (Campos, 1996). Diagnosis. Carapace broadly rounded, without sharp lateral margins or sulci. Antennae minute. Maxilliped 3 with inner distal angle of merus curved, carpus rounded; palp 2–articled. Walking leg 4 articulated to body not dorsally to leg 3; dactyli simple. Ostracotheres holothuriensis (Baker, 1907) (Fig. 154c, e). Carapace of both sexes almost circular, about 1.1 times as long as wide, with longitudinal grooves on anterior third. Chela deeper distally, fixed finger and dactylus each with irregular teeth in proximal half. 12 mm. NSW (Sydney region), Vic., Tas., SA; in the gut cavities of holothurians and in the ascidian Herdmania momus. Other species of pea crabs residing in holothurians have been shown to adversely affect their hosts (Takeda et al., 1997; Hamel et al., 1999). The species was described in detail (Pregenzer, 1988) but has been recorded in Vic. as Pinnotheres obesa (Fulton & Grant, 1906). Ng & Manning (2003) believed that the species’ generic affinities should be reexam- ined. Ostracotheres subglobosus (Baker, 1907) pea crab (Fig. 154d, Pl. 26h). Carapace of female subrectangular, posterolateral angles pronounced, smooth. Chela cylindrical, fixed finger with small proximal teeth, dactylus with obscure teeth. Male not known. 9 mm. ?Japan, SA, WA (Perth, new record); in bivalve molluscs, mytilids and pectinids (Pregenzer, 1988).

Pinnotheres Bosc, 1802 Pinnotheres is certainly the largest genus but recent revisions have put some of its species in other genera. Diagnosis. Carapace of female wider than long. Maxilliped 3 palp of 3 articles, last article styliform and attached along inner side of second last. Pinnotheres hickmani (Guiler, 1950) pea crab (Fig. 154a, g). Qld (Moreton Bay), NSW, Vic., Tas., SA, WA (N to Shark Bay); in bivalve molluscs, mytilids, trigoniids and donacids. The species has been recorded from Australia as P. pisum, a European species, and as P. novaezelandiae,a situation corrected by Pregenzer (1979) who redescribed the species and distinguished the three species. Pregenzer (1978) found that individuals in cultured mussels grew faster than in wild mussels but that growth in both paralleled that in the host. Infestation rates were up to 70% in wild mussels but 100% in cultured individuals.

References Ahyong, S., & Brown, D.E. 2003. Description of Durckheimia lochi n. sp., with an annotated check-list of Australian Pinnotheridae (Crustacea: Decapoda: Brachyura). Zootaxa 254: 1–20. Bologna, P.A.X.,& Heck, K.L. 2000. Impact of pea crab parasitism on bay scallop reproductive potential. Gulf and Caribbean Research Reports 12: 43–46. Campos, E. 1996. Partial revision of pinnotherid crab genera with a two-segmented palp on the third maxilliped (Decapoda: Brachyura). Journal of Crustacean Biology 16: 556–563. Campos, E., Díaz, V.,& Gamboa-Contrera, J.A. 1998. Notes on distribution and taxonomy of five poorly known species of pinnotherid crabs from the eastern Pacific (Crustacea: Brachyura: Pinnotheridae). Proceedings of the Biological Society of Washington 111: 372–381. Campos, E., & Solís-Marín, F. 1998. New records of crabs (Pinnotheridae) symbiotic with irregular echinoids in Cuba. Caribbean Journal of Science 34: 329–330. Cheng, T.C. 1967. Marine mollusks as hosts for symbiosis with a review of known parasites of commercially important species. Advances in Marine Biology 5: 1–424.

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Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Fulton, S.W., & Grant, F.E. 1906. Some little known Victorian decapod Crustacea with descriptions of new species. No. III. Proceedings of the Royal Society of Victoria 19: 5–15, pls 3–5. Geiger, D.L., & Martin, J.W. 1999. The pea crab Orthotheres haliotidis new species (Decapoda: Brachyura: Pinnotheridae) in the Australian abalone Haliotis asinina Linnaeus, 1758 and Haliotis squamata Reeve, 1846 (Gastropoda: Vetigastropoda: Haliotidae). Bulletin of Marine Science 64: 269–280. Grove, M.W., Finelli, C.M., Wethey, D.S., & Woodin, S.A. 2000. The effects of symbiotic crabs on the pumping activity and growth rates of Chaetopterus variopedatus. Journal of Experimental Marine Biology and Ecology 246: 31–52. Hamel, J.F., Ng, P.K.L., & Mercier, A. 1999. Life cycle of the pea crab Pinnotheres halingi sp. nov., an obigate symbiont of the sea cucumber Holothuria scabra Jaeger. Ophelia 50: 149–175. Hseuh, P.-W. 2003. Responses of the pea crab Pinnotheres taichungae to the life history patterns of its primary bivalve host Laternula marilina. Journal of Natural History 37: 1453–1462. Martin, J.W., & Davis, G.E. 2001. An updated classification of the Recent Crustacea. Natural History Museum of Los Angeles County, Science Series 39: 1–124. Ng, P.K.L.,& Manning, R.B. 2003. On two new genera of pea crabs parasitic in holothurians (Crustacea: Decapoda: Brachyura: Pinnotheridae) from the Indo-West Pacific, with notes on allied genera. Proceedings of the Biological Society of Washington 116: 901–919. Pregenzer, C. 1978. Pinnotheres hickmani (Guiler) in wild and cultured Mytilus edulis in Port Phillip Bay, Victoria. Australian Journal of Marine and Freshwater Research 29. Pregenzer, C. 1979. A redescription of Pinnotheres hickmani (Guiler) and comparison with Pinnotheres novaezelandiae Filhol and Pinnotheres pisum (L.) (Decapoda Brachyura). Crustaceana Supplement 5: 22–30. Pregenzer, C. 1988. A redescription of Pinnotheres holothuriensis Baker, 1907 and Pinnotheres subglobosus Baker, 1907 with a reassignment to the genus Ostracotheres (Decapoda, Brachyura). Crustaceana 55: 17–28. Soong, K. 1997. Some life-history observations on the pea crab, Pinnotheres tsingtaoensis, symbiotic with the bivalve mollusk, Sanguinolaria acuta. Crustaceana 70: 855–866. Takeda, S., Tamura, S., & Washio, M. 1997. Relationship between the pea crab Pinnixa tumida and its endobenthic holothurian host Paracaudina chilensis. Marine Ecology Progress Series 149: 143–154.

Mictyridae Dana, 1851 The mictyrids are one of the most loved crabs in Australia; known as soldier crabs, they are distinguished by their semi-spherical bodies, bright blue colour, and their habit of swarming in vast numbers on flat sandy beaches exposed at low tide. Unlike other brachyurans they always walk forwards. They possess broad external maxillipeds which completely close the buccal cavity. Three species have been recorded from Australia but two more from the tropics were foreshadowed by Davie (2002). Diagnosis. Carapace deep, longer than wide; dorsal surface with cervical and cardiobranchial grooves well defined; eyes exposed, orbits obsolete, with small postocular spine. Interantennular septum narrow. Antennular flagellum rudimentary, folding vertically. Antennae small. Buccal cavern large, oval, closed by convex expanded, foliaceous maxillipeds 3. Maxilliped 3 with palp articulating at distolateral angle of merus; exopod slender, concealed, without flagellum. Chelipeds slender, subequal, similar in males and females. Walking legs slender. Male abdomen with somites free, broad, similar to female; base entirely covering sternum. (after Davie, 2002)

487 Marine Decapod Crustacea of Southern Australia

Key to Australian species of Mictyridae 1. Tooth on the dactylus of the male chela obsolete; anterolateral spines not connected by ridges to anterior branchial regions ...... Mictyris livingstonei — Tooth on the dactylus of the male chela conspicuous; anterolateral spines connected by ridges to anterior branchial regions ...... 2 2. Carapace and maxillipeds smooth; posterior border conspicuously produced beyond curve of abdomen; rostrum subpentagonal ...... Mictyris longicarpus — Carapace and maxillipeds covered by large granules; posterior border conspicuously produced but not overlapping curve of abdomen; rostrum triangular . . . Mictyris platycheles

Mictyris Latreille, 1806 – soldier crabs Mictyris is the only genus in the family, represented by one widespread Indo-West Pacific species (M. longicarpus), two endemic Australian species, and another in Japan and nearby (Takeda, 1978). The Australian species were described in detail by McNeill (1926), from whose work the key and diagnoses were adapted. Mictyris livingstonei McNiell, 1926. Carapace and maxillipeds smooth, except for granular ridges on branchial regions. Rostrum with broad base and triangular or truncate tapering apex. Posterior border produced but not overlapping curve of abdomen. Anterolateral spines not connected by ridges to anterior branchial regions. Tooth on the dactylus of the male chela obsolete. Slaty-blue. 14 mm. Qld, NSW (S to Trial Bay); intertidal estuarine and mangrove environments. Unlike the other species this does not swarm on the sediment surface at low tide. This species does not occur within ‘southern Australia’ as used elsewhere in this guide. Mictyris longicarpus Latreille, 1806 (Fig. 155a, b, d, Pl. 27a, b). Carapace and maxillipeds smooth. Rostrum subpentagonal. Posterior border greatly produced and overlapping curve of abdomen. Anterolateral spines connected by ridges to anterior branchial regions. Male with small proximal tooth on dactylus of cheliped. Sky-blue, branchial regions cream-white, leg joints red-purple in live animals. 25 mm. Indo-West Pacific, WA (N of Perth), NT, Qld, NSW, Vic. (E of Wilsons Promontory); intertidal sand or mudflats. The species occurs in large numbers swarming on surface and emerging from their shallow temporary burrows when exposed by the falling tide (Cameron, 1966). Mictyris platycheles Milne Edwards, 1852 (Fig. 155c, Pl. 28a, b). Carapace and maxillipeds covered by large granules. Rostrum triangular. Posterior border produced but not overlapping curve of abdomen. Anterolateral spines connected by ridges to anterior branchial regions. Male with broad triangular proximal tooth on dactylus of cheliped passing behind fixed finger when closed. Blue with underside and branchial regions paler, legs brown-yellow. 16 mm. Qld (Moreton Bay), NSW, Vic. (W to Port Phillip), Tas. (north coast); intertidal sand and mudflats. This is the dominant species in central Victoria, swarming in large numbers on the surface of the sand when the tide recedes.

References Cameron, A.M. 1966. Some aspects of the behaviour of the soldier crab, Mictyris longicarpus. Pacific Science 20: 224–234. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp.

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McNeill, F.A. 1926. Studies in Australian carcinology. No. 2. A revision of the family Mictyridae, the status and synonymy of the genus Megametope and its contained species. Records of the Australian Museum 15: 100–131, pls 9, 10. Takeda, M. 1978. Soldier crabs from Australia and Japan. Bulletin of the National Science Museum, Tokyo, Ser. A (Zoology) 4: 31–38.

b cd

Fig. 155. Mictyridae. a, Mictyris longicarpus (lateral). Rostrum: b, Mictyris longicarpus. c, Mictyris platycheles. d, Mictyris longicarpus (right chela, male).

489 Marine Decapod Crustacea of Southern Australia

Ocypodidae Rafinesque-Schmaltz, 1815 Ocypodids are principally intertidal crabs so are well known, especially in more tropical regions. There, fiddler crabs, sentinel crabs or ghost crabs are commonly encountered at low tide on sandy or muddy intertidal flats. Species diversity is lower on southern shores but the ocypodids remain important inhabitants of mud flats. Most species excavate permanent burrows. A characteristic of many species is the long eyestalks enabling the eyes to be held erect while exploring away from the burrow. The front of the carapace (the area between the eyes) is short relative to the entire width, and the carapace is widest at this point or shortly behind; this combination of characters could be the most reliable way to distinguish the family from the other important intertidal family, Grapsidae. Grapsids have the front occupying at least one-third of the width and the carapace is usually wider more posteriorly. Having said that, the two families are poorly differentiated and recently molecular and morphological analyses have found that neither is monophyletic. In particular, the ocypodid Macrophthalmus appears on the basis of molecular evidence to be closer to some varunine and grapsine genera of Grapsidae than to other ocypodids (Kitaura et al., 2002). Males of several species of ocypodids display arm-waving behaviour, generally thought of as courtship and aggressive behaviour (e.g. Zucker, 1988; Wada & Wang, 1998; Moriito & Wada, 2000). In the genus Uca, more diverse in the tropics than in southern Australia, species can be differentiated on the basis of the bright colours of the male large cheliped (George & Jones, 1982). Species of Ocypode are rapid runners on sandy shores while Macrophthalmus are slower inhabitants of the muddiest environments. Diagnosis. Carapace quadrangular to pentagonal; deep and strongly vaulted, or flattened. Front narrow, often in form of deflexed lobe between bases of eyestalks. Orbits broad, occupying whole anterior margin of carapace each side of front. Eyestalks long. Maxilliped 3 usually completely closing buccal cavern, sometimes leaving a gap but never rhomboidal (cf. Grapsidae); exopod slender and usually concealed; palp articulating at or near distolateral angle of merus. Male abdomen narrow, with all somites distinct. Often with hairy-edged pouch leading into branchial cavity between bases of walking legs 2 and 3 or 1 and 2. (after Davie, 2002)

Key to southern Australian species of Ocypodidae 1. Carapace almost twice as wide as long, deep, regions weakly marked, lateral margins strongly convex; chelipeds equal, with fingers twisted; thoracic sternite narrowed posteriorly ...... Heloeciinae . . . Heloecius cordiformis — Carapace quadrilateral or wider than long, deep or flat, regions more or less marked, lateral margins curved or straight, often with tooth or teeth; chelipeds equal in both sexes or larger in male; fingers not twisted; thoracic sternite wider posteriorly ...... 2 2. Carapace deeply convex, with anterolateral tooth behind orbital tooth; merus of walking legs with tympanum (flat membrane) ...... Dotillinae . . . Scopimera inflata — Carapace flat, with or without anterolateral tooth; merus of walking leg without tympanum . . 3 3. Chelipeds subequal in both sexes; carapace often with lateral teeth; without brushes of long setae at bases of walking legs 2 and 3 ...... Macrophthalminae ...4 — Chelipeds unequal in male, equal and smaller in female; carapace with straight lateral margin; with brush of long setae edging pouch leading to branchial cavity at bases of walking legs 2 and 3 ...... Ocypodinae ...10 4. Male chela globose, with cutting edges of fingers obscured by dense hair, dactylus without large differentiated tooth on margin ...... Australoplax tridentata — Male chela not globose, with cutting edges of fingers without hair, dactylus with large differentiated tooth on margin ...... 5

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Fig. 156. Ocypodidae. a, Heloecius cordiformis. b, Macrophthalmus punctulatus. c, Macrophthalmus latifrons. Anterolateral carapace and front: d, Macrophthalmus latreillei. e, Macrophthalmus setosus. Maxilliped 3: f, Macrophthalmus latifrons. g, Macrophthalmus crassipes.

491 Marine Decapod Crustacea of Southern Australia

5. Front 0.4 anterior width; maxilliped 3 merus smaller than ischium; epistome with convex middle region; fingers of chela pointed ...... Enigmaplax littoralis — Front at most 0.3 anterior width; maxilliped 3 merus smaller or subequal to ischium; epistome with straight or concave middle region or with protuberance; fingers of chela spoon-tipped (at least in female) ...... Macrophthalmus ...6 6. Epistome with central protuberance ...... 7 — Epistome straight or excavated ...... 8 7. Merus of maxilliped 3 markedly smaller than ischium (Fig. 156g); male chela palm with spines on inner surface near articulation with carpus ...... Macrophthalmus crassipes — Merus and ischium of maxilliped 3 subequal (Fig. 156f); male chela palm without spines on inner surface ...... Macrophthalmus latifrons 8. Merus and ischium of maxilliped 3 subequal; front 0.3 total width ...... Macrophthalmus punctulatus — Merus of maxilliped 3 markedly smaller than ischium; front narrow ...... 9 9. Carapace widest across external orbital angles; male chela and fixed finger with longitudinal ridge near lower margin on outer surface ...... Macrophthalmus setosus — Carapace widest posterior to external orbital angle; male chela and fixed finger without longitudinal ridge near lower margin on outer surface ...... Macrophthalmus latreillei 10. Carapace wider than long, broader anteriorly; chelipeds unequal in male only . . Uca vomeris — Carapace as broad as long, slightly tapering posteriorly; chelipeds unequal in both sexes ...... Ocypode ...11 11. Palm of large cheliped without stridulating organ ...... Ocypode cordimanus — Palm of large cheliped with stridulating organ on inner face ...... 12 12. Stridulating organ composed of elongated tubercles in upper part and more closely space transverse ridges in the lower; lower orbital ridge without notch (specimens > 24 mm wide with horned eyestalk) ...... Ocypode ceratophthalma — Stridulating organ solely composed of 10–25 tubercles; lower orbital ridge with deep lateral and median notches ...... Ocypode convexa

Subfamily Dotillinae Dana, 1851

Scopimera De Haan, 1883 The tympanum, a thin area of the exoskeleton on the face of the merus of the walking legs, is a diagnostic feature of the genus and its only species in southern Australia. The crabs sits on its long fingers with the body oriented upwards such that the large maxillipeds are clearly visible from above. The Australian species is one of few in the Indo-West Pacific. Diagnosis. Carapace globose. Frontal width about one-eighth total width. Orbits with prominent lower border, with infraorbital crest. Maxillipeds 3 dominant and closing buccal cavern. Chelipeds with fingers slender and bent. Walking legs slender, dactyli curved, unarmed; merus with tympanum (flat membrane); with hairy ridged pouch between bases of walking legs 1 and 2. Scopimera inflata A. Milne Edwards, 1873 sand-bubbler crab (Fig. 157h) 20 mm. Qld, NSW; midintertidal sheltered sand beaches. The sand-bubbler crab is well known on beaches north of Sydney where it lives in vertical burrows. It feeds by leaving the burrow and collecting sand grains from which it scrapes microbes. The sand is rolled into balls by the mouthparts and

492 Brachyura – crabs

then discarded on the surface of the beach in an idiosyncratic pattern of radiating rows leading away from the mouth of the burrow (Fielder, 1970). Similar crabs in the Northern Territory and Western Australia belong to another species in the same genus (Davie, 2002).

Subfamily Heloeciinae Milne Edwards, 1852

Heloecius Dana, 1851 The only species in the subfamily Heloeciinae is an Australian endemic (Türkay, 1983). Diagnosis. Carapace narrowing posteriorly, lateral margins convex; front narrow, deflexed, a lobe between eyestalks. Interantennal septum broad. Chelipeds equal, smaller in female. With brush of setae edging pouch leading to branchial cavity between bases of walking legs 2 and 3. Thoracic sternum narrowed posteriorly. Heloecius cordiformis (Milne Edwards, 1837) semaphore crab (Fig. 156a, Pl. 28g) Purple, chelipeds lighter purple, orange-red in juveniles. 25 mm. Qld (N to Brisbane), NSW, Vic. (mainly central bays), Tas.; midintertidal estuaries, mangroves, seagrass mudflats. The twisted fingers of the chelipeds are characteristic. The species can be the most abundant crab on the muddy environments in south-eastern Australia and is sometimes found together with Macrophthalmus latifrons. The crab’s tunnel is vertical into the mud and ends in an enlarged chamber. The common name, semaphore crab, derives from the behaviour of the male which display at the mouth of its burrow by raising and lowering its chelipeds in unison (Griffin, 1965).

Subfamily Macrophthalminae Dana, 1851

Australoplax Barnes, 1966 Barnes (1966) based his new genus for one species on the globose male chela with dense hairs on the fingers and correlated these features with others in the male abdomen, carapace and maxillipeds. Diagnosis. Carapace flat, wider than long, lateral margins subparallel, with anterolateral tooth plus 2 lateral teeth. Frontal width about 0.3 times total width, not basally constricted. Orbits reaching greatest width of carapace, sloping backwards, upper border smooth, lower border with central concavity, smooth in male and tuberculate in female; without infraorbital crest. Antennae in orbital hiatus. Maxillipeds 3 not closing the buccal cavern, merus about as large as ischium; ischium with oblique row of hairs. Chelipeds subequal, well developed in male; merus triangular in cross-section; palm globose, cutting margins evenly serrated, obscured externally by dense hairs. Walking legs slender, dactyli curved, unarmed; merus with subdistal tooth on upper margin; no hairy ridged pouch between bases of walking legs 1 and 2. Australoplax tridentata (Milne Edwards, 1873) furry-clawed crab, tuxedo crab (Fig. 157a). 15 mm. NT, Qld, NSW (S to Sydney); intertidal mangroves, estuaries (Barnes, 1967).

Enigmaplax Davie, 1993 The genus contains only one species, common in eastern Australia (Davie, 1993). Diagnosis. Carapace flat, quadrangular, lateral margins subparallel, dentate. Frontal width 0.4 times total width. Postfrontal lobes (elevations immediately behind front) sharply defined,

493 Marine Decapod Crustacea of Southern Australia

e g

Fig. 157. Ocypodidae. a, Australoplax tridentata. b, Ocypode convexa. c, Enigmaplax littoralis. d, Uca vomeris. Cheliped: e, Australoplax tridentata. f, Macrophthalmus punctulatus. g, Uca vomeris. h, Scopimera inflata. i, Ocypode ceratophthalma (stridulatory organ, inside of cheliped).

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straight. Orbits with lower border complete, without infraorbital crest. Antennae included in orbital hiatus. Maxillipeds 3 not closing buccal cavern. Chelipeds weak, cylindrical, cutting margins evenly serrated, with basal tooth in adult males. Walking legs slender, dactyli curved, unarmed; merus with subdistal tooth on upper margin; no hairy ridged pouch between bases of walking legs 1 and 2. Enigmaplax littoralis Davie, 1993 (Fig. 157c). Parallel lateral margins with 3 teeth, including most dominant anterolateral tooth. 9 mm. Qld (N to Cairns), NSW (S to Port Hacking); intertidal and shallow subtidal Zostera beds. The species often co-occurs with, and is easily confused with the grapsid, Ilyograpsus paludicola (Rathbun, 1909); both occur in Port Hacking, NSW. Enigmaplax littoralis is more rectangular. The similarity between the two species would seem to confirm the poor distinction between these two families.

Macrophthalmus Desmarest, 1823 Macrophthalmus is a widespread and diverse genus of tropical intertidal crabs, all burrowers on intertidal mudflats especially in mangrove forests. Most of the species recorded in southern Australia are rare and at the southern limits of their range near Sydney or Perth. The only exception is M. latifrons, whose distribution does not overlap with the others. The Australian fauna was revised by Barnes (1967) who also reviewed the biogeography of the subfamily Macrophthalminae (Barnes, 1968). He concluded that the Australian fauna was more related to the species in South- East Asia rather than those further east or west. The quarter of the species that are endemic are confined to the eastern and south-eastern coasts of the continent and include the temperate species. Later, Barnes (1971) provided a key to all the species of Macrophthalmus then known. The genus is divided into subgenera and the five southern Australian species represent five subgenera as follows: M. (Macrophthalmus) crassipes, M. (Tasmanoplax) latifrons, M. (Venetus) latreillei, M. (Chaenostoma) punctulatus, M. (Mareotis) setosus.

Diagnosis. Carapace flat, wider than long, lateral margins subparallel or tapering posteriorly, with lateral teeth. Frontal width up to 0.3 times total width, often narrower, sometimes basally constricted. Orbits reaching greatest width of carapace, transverse or sloping backwards, upper border usually granulate, lower border complete, projecting, tuberculate; without infraorbital crest. Antennae in orbital hiatus. Maxillipeds 3 not closing the buccal cavern, merus about as large as or smaller than ischium; ischium without row of hairs. Chelipeds subequal, well developed in male; merus triangular in cross-section; palm never globose, cutting margins serrated and with larger tooth, not obscured externally by dense hairs. Walking legs slender, dactyli curved, unarmed; merus with subdistal tooth on upper margin; no hairy ridged pouch between bases of walking legs 1 and 2. Macrophthalmus crassipes Milne Edwards, 1852 smooth sentinel crab (Fig. 156g). Front narrow, deflexed, constricted. Epistome with central protuberance. Merus of maxilliped 3 markedly smaller than ischium. Male chela palm with spines on inner surface near articulation with carpus, fixed finger deflexed. 37 mm. Eastern Indo-West Pacific, WA, NT, Qld, NSW; intertidal mangrove mudflats. Macrophthalmus latifrons (Haswell, 1881) southern sentinel crab (Fig. 156c, f). Front narrow, deflexed, slightly constricted. Carapace with granules in branchial region. Epistome with central protuberance. Male chela palm without spines but with longitudinal ridge on inner surface, without larger tooth on deflexed fixed finger. Yellow-brown. 26 mm. Vic., Tas., SA; intertidal mudflats, seagrass beds. In Victoria the southern sentinel crab is abundant on the intertidal mudflats of Western Port, Andersons Inlet, Corner Inlet, Vic., but virtually absent elsewhere in the state (Phillips et al., 1984).

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Macrophthalmus latreillei (Desmarest, 1822) smooth sentinel crab (Fig. 156d). Front narrow, deflexed, constricted. Carapace granular. Epistome straight. Merus of maxilliped 3 markedly smaller than ischium. Male chela and fixed finger without longitudinal ridge near lower margin on outer surface, inner surface of male dactylus hairy, fixed finger straight. 60 mm. Indo-West Pacific, WA, NT, Qld; intertidal mangrove mudflats. Macrophthalmus punctulatus Miers, 1884 smooth sentinel crab (Fig. 156b) Front 0.3 total width. Carapace not granular. Epistome straight. Merus and ischium of maxilliped 3 subequal. Male chela palm without spines on inner surface, fixed finger moderately deflexed. 11 mm. Qld, NSW, WA; intertidal mangrove mudflats. Macrophthalmus setosus Milne Edwards, 1852 smooth sentinel crab (Fig. 156e). Front narrow, deflexed, constricted. Carapace smooth. Epistome excavated. Merus of maxilliped 3 markedly smaller than ischium. Male chela and fixed finger with longitudinal ridge near lower margin on outer surface, fixed finger deflexed. 40 mm. Qld, NSW, WA; intertidal mangrove mudflats.

Subfamily Ocypodinae Rafinesque-Schmaltz, 1815

Ocypode Weber, 1795 Ghost crabs of the genus Ocypode are mainly inhabitants of tropical beaches but two species reach as far south as central NSW and one to Perth, WA. All make deep burrows above the high tide mark and rarely leave the burrow during the day. At night however they run quickly over the beach where they scavenge and are known to forage almost a kilometre away from the sea. All have long eyes, held erect when the crab is active. Most species are a pale sandy colour, darker on the carapace. Ocypode ceratophthalma is easily recognised as adults by the long horn extending beyond the eye but this feature is not present in smaller individuals. The stridulating organ, a rough area on the inside of the large cheliped is a more reliable character to differentiate the species (George & Knott, 1965). Diagnosis. Carapace as broad as long, slightly tapering posteriorly. Antenna 1 flagellum rudimentary. Eyestalks not elongated, cornea swollen, occupying whole ventral part of stalk. Chelipeds unequal in both sexes. Ocypode ceratophthalma (Pallas, 1772) horn-eyed ghost crab (Fig. 157i, Pl. 28c). Stridulating organ composed of well spaced elongated tubercles in upper part and closely spaced transverse ridges in the lower. Lower orbital ridge without notch. Specimens >24 mm wide with horned eyestalk. 40 mm. Indo-West Pacific; WA (north-west), NT, Qld, NSW (S to Sydney); supratidal and intertidal, very active on ocean beaches. Ocypode convexa Quoy & Gaimard, 1824 (Fig. 157b). Stridulating organ solely composed of 10–25 tubercles. Lower orbital ridge with deep lateral and median notches. Eyestalk without horn. 42 mm. WA (S to Yallingup); supratidal and intertidal, ocean beaches. Ocypode cordimanus Desmarest, 1825. Palm of large cheliped without stridulating organ. Lower orbital ridge with broad key-hole shaped lateral notch and shallow median notch. Eyestalk without horn. 28 mm. Indo-West Pacific; in Australia from Cocos (Keeling) Islands, WA (north-west), NT, Qld, NSW; supratidal and intertidal, ocean beaches.

Uca Leach, 1814 Species of Uca are the true fiddler or semaphore crabs. They are notable, the males at least, for one grossly enlarged colourful cheliped used in signalling other males and potential mates. The genus is diverse in the tropics and taxonomically confused. Crane’s (1975) major revision scarcely dealt

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with the Australian fauna. Although the literature abounds with other genera and subgenera, George & Jones (1982) who revised the Australian fauna recognised only Uca. Their detailed observations and colour photographs are essential to identify the almost 20 species known from tropical Australia. About half the species are Australian endemics but only one species is resident in Sydney. Two more occur in northern NSW. Uca crassipes (Adams & White, 1848) recorded only once from Port Jackson as ‘washed up in a gale’ (George & Jones, 1982) is not included here. Diagnosis. Carapace wider than long, much broader anteriorly. Antenna 1 flagellum small, not hidden beneath front. Eyestalks slender and long, cornea small and terminal. Chelipeds unequal in male only, one grossly enlarged. Uca vomeris McNiell, 1920 (Fig. 157d, g). Front narrow. Major cheliped with groove on outer surface of fixed finger extending from a triangular depression, without groove on dactylus, and without groove parallel to lower margin. Female chela without serration or teeth in gape. Major chela above and dactylus grey to pink, chela below and fixed finger orange; walking legs orange-brown, in female with cream patch on first. 30 mm. NT, Qld, NSW (S to Port Jackson); intertidal, unshaded sandy mud along lower tide levels of bays and creeks (Hagen, 1993).

References Barnes, R.S.K. 1966. The status of the crab genus Euplax H. Milne Edwards, 1852; and a new genus Austroplax of the subfamily Macrophthalminae Dana, 1851 (Brachyura: Ocypodidae). Australian Zoologist 13: 370–376, pl. 1. Barnes, R.S.K. 1967. The Macrophthalminae of Australasia: with a review of the evolution and morphological diversity of the type genus Macrophthalmus (Crustacea: Brachyura). Transactions of the Zoological Society of London 31: 197–261. Barnes, R.S.K. 1968. Aspects of the Australasian zoogeography of the Macrophthalminae (Brachyura, Ocypodidae). Proceedings of the Linnean Society of London 179: 67–75. Barnes, R.S.K. 1971. Biological results of the Snellius expedition. XXIII. The genus Macrophthalmus (Crustacea, Brachyura). Zoologische Verhandelingen, Leiden 115: 1–40. Crane, J. 1975. Fiddler Crabs of the World. Ocypodidae: Genus Uca. Princeton University Press: Jersey. 736 pp. Davie, P.J.F. 1993. A new genus of macrophthalmine crab (Crustacea: Decapoda: Ocypodidae) from eastern Australia. Records of the Australian Museum 45: 5–9. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Fielder, D.R. 1970. The feeding behaviour of the sand crab Scopimera inflata (Decapod, Ocypodidae). Journal of the Zoological Society of London 160: 35–49. George, R.W., & Jones, D.S. 1982. A revision of the fiddler crabs of Australia (Ocypodinae: Uca). Records of the Western Australian Museum, Supplement 14: 1–99. George, R.W., & Knott, M.E. 1965. The ocypode ghost crabs of Western Australia (Crustacea, Brachyura). Journal of the Royal Society of Western Australia 48: 15–21. Griffin, D.J.G. 1965. The behaviour of shore crabs. Australian Natural History 13: 87–91. Hagen, H.O.v. 1993. Waving display in females of Uca polita and of other Australian fiddler crabs. Ethology 93: 3–20. Kitaura, J., Wada, K., & Nishida, M. 2002. Molecular phylogeny of grapsoid and ocypodoid crabs with special reference to the genera Metaplax and Macrophthalmus. Journal of Crustacean Biology 22: 682–693. Moriito, M., & Wada, K. 2000. The presence of neighbors affects waving waving display frequency by Scopimera globosa (Decapoda, Ocypodidae). Journal of Ethology 18: 43–45.

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Phillips, D.A.B., Handreck, C.P., Bock, P.E., Burn, R., Smith, B.J., & Staples, D.A. 1984. Coastal invertebrates of Victoria. An atlas of selected species. Marine Research Group of Victoria in association with the Museum of Victoria: Melbourne. 168 pp. 1st edn. Türkay, M. 1983. The systematic position of an Australian mangrove crab Heloecius cordiformis (Crustacea: Decapoda: Brachyura). Memoirs of the Australian Museum 18: 107–111. Wada, K., & Wang, C.-H. 1998. Territorial and sexual behaviors in Ilyoplax formosensis and I. tansuiensis (Crustacea, Brachyura, Ocypodidae). Journal of the Taiwan Museum 51: 119–125. Zucker, N. 1988. Preliminary observation of cheliped use during social activities in sentinel crabs (Brachyura, family Ocypodidae, genus Macrophthalmus) from northern Queensland, Australia. Bulletin of Marine Science 43: 98–102.

Palicidae Bouvier, 1898 Although diverse in tropical benthic collections, palicids are uncommon and were poorly understood until revised by Castro (2000). Following his revision of the Indo-West Pacific fauna 58 species are now known, 13 in Australia (Castro & Davie, 2003). Palicids, at least in southern Australia, are small flat crabs, about as wide as long, with weak chelipeds, and reduced last legs. The only southern records are from Western Australia. The southern species have a highly sculptured carapace. Castro (2000) provided an extended diagnosis (see too Davie, 2002) and keys to genera and species from which the following was abbreviated. Southern Australian species belong to one of the two subfamilies, Palicinae. Most palicids live on muddy or sandy benthic environments on the shelf or slope and some are associated with coral reefs. Diagnosis. Carapace depressed but with various high bosses; frontal carapace border with 2–4 lobes; anterolateral border with 1–4 teeth; supraorbital and suborbital borders long, usually with lobes; orbits wide and deep. Chelipeds small. Buccal cavity square, mostly but not completely covered by maxillipeds 3; maxilliped 3 merus small, with palp on inner border. Walking leg 4 smaller than other legs, smaller than carapace length, similar in form to other legs (Crossonotoninae) or more slender (Palicinae). Abdomen of 7 somites.

Pseudopalicus Moosa & Serène, 1981 The combination of walking leg characters serves to define the genus. Eleven or 12 species are known, most in the Indo-West Pacific but only one occurs on the southern coast and another as far south as Perth, distinguished by carapace armature. Diagnosis. Carapace hexagonal, with dorsal bosses, wider than long; frontal carapace border with 4 lobes; anterolateral borders curved and armed with 3 or 4 teeth. Walking leg 4 reduced, shorter, more slender, and articulating more dorsally than other walking legs. Walking legs 1–3 not especially filiform, with flattened carpi and propodi, meri with tuberculate margins; walking leg 3 much longer than carapace width. Abdominal segments 1–2 dorsoventrally compressed and narrower than others.

Key to southern Australian species of Pseudopalicus 1. Anterolateral border with 4 pointed teeth; posterior border with 8–10 tubercles ...... Pseudopalicus investigatoris — Anterolateral border with 3 granular teeth; posterior border with 4–6 tubercles ...... Pseudopalicus macromeles

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Fig. 158. Palicidae. a, Pseudopalicus investigatoris. b, Pseudopalicus macromeles.

Pseudopalicus investigatoris (Alcock, 1900) (Fig. 158a) Anterolateral border of carapace with 4 pointed teeth (excluding exorbital tooth); posterior border with 8–10 pointed or elongate tubercles; supraorbital border with 2 triangular lobes. Reddish-brown. 26 mm. Indo-West Pacific, WA (S to Garden I.); shelf and slope, 43–426 m depth. Pseudopalicus macromeles Castro, 2000 (Fig. 158b). Anterolateral border of carapace with 3 granular teeth (excluding exorbital tooth); posterior border with 4–6 low elongate tubercles; supraorbital border with 2 triangular lobes. 13 mm. Vic. (eastern slope), SA, WA (N to Dongara); shelf and slope, 128–570 m depth.

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References Castro, P. 2000. Crustacea Decapoda: a revision of the Indo-West Pacific species of palicid crabs (Brachyura Palicidae). In: Crosnier, A. (ed), Résultats des Campagnes MUSORSTOM, Vol. 21. Mémoires du Muséum National d’Histoire Naturelle, Paris 184: 437–610. Castro, P., & Davie, P.J.F. 2003. New records of palicid crabs (Crustacea: Brachyura: Palicidae) from Australia. Memoirs of the Queensland Museum 49: 153–157. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp.

Grapsidae MacLeay, 1838 The Grapsidae are a large family of principally intertidal crabs – the crabs most commonly met with by fossickers on the seashore in southern Australia. Most of the crabs that flee when disturbed by turning stones on rocky coasts are grapsids. The most distinctive feature characterising members of the family is the wide front between short eyestalks, and the rhomboidal gap between the third maxillipeds (Fig. 163c). With at least 40 genera and 300 species described worldwide (Rice, 1980) the family is one of the most diverse known, especially in shallow waters. Of about 60 Australian species 15 occur in southern Australia. Most species in southern Australia are active species of rocky shores or burrow in muddy environments intertidally (see detailed review of several species by Griffin, 1971). Several live in muddy environments in tropical regions. In mangrove forests, burrowing grapsids are important nutrient recyclers (Smith et al., 1991). Species of the genus Geograpsus are inhabitants of tropical forests. The family includes the bromeliad crab who makes the water held by these plants its home. Crabs of the genus Planes are pelagic, especially in the tropics where they ride on rafts of algae and other flotsam. The Grapsidae have traditionally included the plagusiids as a distinctive subfamily but these have been removed to their own family by Sternberg & Cumberlidge (1998), a decision followed here. Davie (2002) treated the remainder as four subfamilies but acknowledged that phylogenetic analyses do not provide support for the usual allocation of genera. Sternberg & Cumberlidge (1998) concluded on the basis of a cladistic analysis of morphological characters that the Grapsinae as usually understood are monophyletic, that Sesarminae are paraphyletic with Gecarcinidae and others, although part could be treated as monophyletic, and that Varuninae were found to be polyphyletic and mixed with ocypodids and other grapsid genera. American genera of Grapsidae were investigated by a molecular analysis of the 16s rRNA gene (Schubart et al., 2000) who concluded that several genera did not fit well in the subfamilies in which they are traditionally included. Kitaura et al. (2002) also caste doubt on the monophyly of the Grapsidae. Each of these subfamilies is represented in southern Australia but the differences between them, Grapsinae and Varuninae for example, can be difficult to understand. These two subfamilies for example are separated on the absence (Grapsinae) or presence (Varuninae) of a suborbital crest, a ridge subparallel to and below the actual infraorbital margin, a lesser ridge running on to the orbital tooth. The suborbital crest ends on the pterygostomial face. Some Australian genera are hard to place – and some look more like ocypodids than grapsids! The subfamilies are identified in the key and the descriptions of Australian species are arranged below by subfamily. More extended family and subfamily diagnoses can be found in Davie’s (2002) catalogue. Ambiguous characters have been edited from his family diagnosis in the short version presented here.

Diagnosis. Carapace quadrilateral to quadrate; front wide, much broader than orbits. Interantennular septum broad, dividing orbit into 2 fossae. Buccal cavern quadrate. Maxillipeds 3 usually separated by a distinct rhomboidal gap; with palp articulating at distolateral angle or at

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middle of distal border of merus. Chelipeds usually stout and robust, subequal, sexes similar. Male abdomen usually with 7 free somites, somites 4–6 sometimes fused. Male and female genital openings sternal. Male gonopod 1 often with corneous tip or process well developed, usually obscured by dense distal brush of long setae.

Key to southern Australian species of Grapsidae 1. Outer surface of maxilliped 3 with oblique hairy ridge (Fig. 160d) ...... 2 — No hairy ridge on maxilliped 3 (Fig. 162d) ...... 10 2. Carapace quadrangular or parallel or divergent margins, with anterolateral angles being widest point of carapace; pterygostomial region (underside of carapace above legs) with hairs and granules arranged in reticulate pattern (Fig. 163c) . . Sesarminae ...(p.508) ...3 — Carapace smoothly convex or subparallel margins, with anterolateral angles not being widest point of carapace; pterygostomial region (underside of carapace above legs) sometimes granular, without reticulate lines of setae ...... Cyclograpsinae ...(p.503) ...4 3. Lateral margin of carapace with 1 postorbital tooth ...... Perisesarma longicristatum — Lateral margin of carapace without postorbital tooth ...... Parasesarma erythodactyla 4. Lateral margins convex, without teeth ...... Cyclograpsus ...5 — Lateral margins convex or straight, with at least 1 notch and blunt tooth ...... 6 5. Sternum and maxilliped 3 conspicuously hairy, prominent tufts of hairs between bases of walking legs (Fig. 160e); frontal region of carapace smooth to minutely granulate ...... Cyclograpsus audouinii — Sternum, maxilliped 3 and bases of walking legs with short, sparse hairs (Fig. 160f); frontal region of carapace distinctly granulate ...... Cyclograpsus granulosus 6. Frontal width (frontal width of carapace, excluding orbital region) much less than half fronto- orbital width (frontal width of carapace, including orbital region) (orbits relatively wide); abdominal somite 3 of male much wider than other abdominal somites ...... Helice leachii — Frontal width (frontal width of carapace, excluding orbital region) half or more fronto- orbital width (frontal width of carapace, including orbital region) (orbits relatively small); abdominal somite 3 of male only slightly wider than other abdominal somites ...... 7 7. Carapace deep, vaulted (c. 0.65 times length); front of carapace curving evenly downwards; legs slender; telson of female 1.5 times as wide as long ...... Helograpsus haswellianus — Carapace flat (< 0.6 times length); front of carapace projecting forwards shelf-like, not curving down; legs stout; telson of female 2 times as wide as long ...... Paragrapsus ...8 8. Anterolateral margins with 1 tooth posterior to outer orbital angle ...... Paragrapsus quadridentatus — Anterolateral margins with 2 teeth posterior to outer orbital angle ...... 9 9. First walking legs with felt on anterior surface of carpus, propodus and dactylus (Fig. 160b); suture between sternites 1 and 2 not marked by prominent ridge ...... Paragrapsus laevis — First walking legs with only inner edge felted (felting visible only by extending legs in ventral view) (Fig. 160a); sternites 1 and 2 of male separated by a distinct ridge ...... Paragrapsus gaimardii 10. Carapace with 3 weak anterolateral teeth, first well separated from orbital tooth, third defining widest point of carapace; fingers of cheliped with mat of hairs on inner and outer surfaces (Fig. 163e) [subfamily character: suborbital crest subparallel with and below infraorbital margin over most of orbit length] ...... Varuninae … (p. 509) … Brachynotus spinosus

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c d e

Fig. 159. Grapsidae Cyclograpsinae. a, Cyclograpsus granulosus. b, Helice leachii. c, Helograpsus haswellianus. d, Paragrapsus gaimardii. e, Paragrapsus quadridentatus.

— Carapace with 1 or 2 weak anterolateral teeth close to orbital tooth or 2 strong teeth behind orbital tooth; fingers of cheliped without mat of hairs on inner and outer surfaces [subfamily character: suborbital crest dominates inner half of orbital margin, true infraorbital margin ending far from inner end of margin, sometimes interrupted by a deep notch] ...... Grapsinae ...(p.505) ...11 11. Anterolateral margins of carapace with 2 strong teeth behind orbital tooth followed by minor notches ...... Ilyograpsus paludicola — Anterolateral margins of carapace with 1 or 2 weak teeth close to orbital tooth ...... 12 12. All walking legs, or at least second pair of walking legs, with dense mat of hairs on outer margin ...... 13 — Walking legs naked or with sparse line of hairs, never densely hairy ...... 14 13. Carapace smoothly convex; lateral margin with a single rounded indentation near orbit but without acute teeth; anterior edge of merus, propodus and dactylus of all walking legs with dense fringe of hairs; pelagic ...... Planes major — Carapace depressed, almost flat; lateral margin with 1 acute tooth behind outer orbital angle; posterior surface of propodus of walking legs with dense mat of hairs (most conspicuous on walking legs 2); benthic ...... Pachygrapsus laevimanus

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d a c b

e f g

Fig. 160. Grapsidae Cyclograpsinae. Right front, eyestalk, cheliped and walking leg 1: a, Paragrapsus gaimardii. b, Paragrapsus laevis. c, Paragrapsus quadridentatus. Maxilliped 3: d, Cyclograpsus audouinii. Thoracic sternum: e, Cyclograpsus audouinii. f, Cyclograpsus granulosus. g, Paragrapsus gaimardii.

14. Carapace with 10 or more prominent transverse ridges connecting to lateral margins; propodus and dactylus of walking legs conspicuously setose; fixed finger not markedly bent; abdomen and telson of male with evenly tapering sides, telson slightly narrower than abdominal somite 6 ...... Leptograpsus variegatus — Carapace smooth or with several faint ridges near lateral margins; propodus and dactylus of walking legs with sparse minute setae; fixed finger markedly bent downwards; telson markedly narrower than abdominal somite 6 of male ...... Leptograpsodes octodentatus

Subfamily Cyclograpsinae H. Milne Edwards, 1853

Cyclograpsus H. Milne Edwards, 1837 The genus Cyclograpsus includes at least 17 species, which are distributed throughout the world in temperate and tropical latitudes. Two species occur in southern Australia and another in northern Australian waters. Campbell & Griffin (1966) provided a key to all Indo-Pacific species and more detailed descriptions of the two species covered here. Both species are among the most common intertidal grapsid crabs in south-eastern Australia. Both prefer the higher intertidal of rocky boulder-covered shores and in areas of overlap C. audouinii is often found higher on the shore than C. granulosus and was able to survive a greater degree of desiccation than the other (Jessop, 1995). Also, in these areas of overlap the two species hybridise, hybrids showing intermediate morpholo- gies in granulation, hairs and felting on walking legs, inter-leg hairiness (Wescott, 1979; Jessop, 1995). In New Zealand, Cyclograpsus lavauxi is well studied (Jones, 1976; Taylor et al., 1984). Diagnosis. Carapace smooth, with at most 2–3 faint ridges obliquely connecting to posterolateral margin; infraorbital margin open, antenna 2 free to enter orbit; anterolateral corners rounded; lateral margin behind orbital angle without teeth. Pterygostomial region with granules but lacking distinct reticulate pattern.

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Cyclograpsus audouinii (H. Milne Edwards, 1837) (Fig. 160d, e, Pl. 28e). Frontal region of carapace smooth to minutely granulate. Sternum and maxilliped 3 conspicuously hairy, prominent tufts of hairs between bases of walking legs. Carapace mottled variably with red-brown or purple on yellow. 40 mm. New Guinea, Qld, NSW, Vic., SA, WA (N to Shark Bay); upper intertidal to supratidal, rarely subtidal, but limited to supratidal fringe when coexisting with C. granulosus (Wescott, 1979); sheltered or moderately-exposed rocky shores, common. Cyclograpsus granulosus Milne Edwards, 1853 (Figs 159a, 160f, Pl. 28f). Frontal region of carapace distinctly granulate. Sternum, maxilliped 3 and bases of walking legs with short, sparse hairs. Carapace mottled variably with red-brown or purple on yellow. 38 mm. Vic., Tas., SA (W to Kangaroo I.); intertidal, rocky and boulder shores, common.

Helice De Haan, 1833 Currently Helice comprises seven species distributed throughout the Indo-Pacific (Campbell & Griffin, 1966) but according to Davie (2002) the only Australian representative deserves to be called by an older species name and warrants a generic status separate from the rest. Not much is known about the Australian species but the ecology, physiology and reproduction of Helice crassa in New Zealand is well documented (Jones, 1980, 1981; Hawkins et al., 1982; Jones & Simons, 1983). Diagnosis. Carapace with frontal width (frontal width of carapace, excluding orbital region) markedly less than half fronto-orbital width (frontal width of carapace, including orbital region) (orbits relatively wide); frontal border merging smoothly with orbits, with no angle between them; anterolateral margin with 2 teeth behind orbital angle. Epistome visible in dorsal view. Pterygostomial region granulate but not reticulate. Abdominal somite 3 of male much wider than other somites. Telson of mature females about as broad as long. Helice leachii Hess, 1865 (Fig. 159b). Strongly hirsute between coxae and sternum; gape of chelae also with mat of hairs at articulation of propodus and dactyl. Purple-red with scattered light cream mottling. 24 mm. Indo-West Pacific, Qld, NSW (S to Port Jackson); burrowing near and above high tide level in muddy and stony environments, extending into estuaries (Campbell & Griffin, 1966).

Helograpsus Campbell & Griffin, 1966 The genus is monotypic, its species being endemic to Australia. Diagnosis. Carapace deep and vaulted (height c. 0.65 length); frontal width (frontal width of carapace, excluding orbital region) half fronto-orbital width (frontal width of carapace, including orbital region) (orbits relatively small); with angle between frontal border and orbits; anterolateral margin with 1 tooth behind orbital angle. Epistome not visible in dorsal view. Pterygostomial region granulate but not reticulate. Abdominal somite 3 of male not wider than other somites. Telson of mature females about as broad as long. Helograpsus haswellianus (Whitelegge, 1889) Haswell’s shore crab (Fig. 159c, Pl. 28g). Olive- brown or reddish. 25 mm. Qld, NSW, Vic., Tas., SA (W to Adelaide); high intertidal and supratidal, bays, estuaries almost to fresh water, burrowing under rocks and in seagrass on muddy environments. The deep body profile, microscopically granular carapace and chelae, and single anterolateral notch are distinguishing features. Crabs construct their burrows using the chelae as shovels and sometimes cover the surface of the sediment with pellets of excavated mud. The crab itself inhabits an enlarged space just a short distance from the surface (Hale, 1927).

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Paragrapsus H. Milne Edwards, 1853 The genus Paragrapsus contains three species; all are restricted to south-eastern Australia. Only P. laevis reaches as far north as southern Queensland (Campbell & Griffin, 1966). The three species are the most common shore crabs on exposed rocky coasts of the region. Diagnosis. Carapace flat (height < 0.6 length); frontal width (frontal width of carapace, excluding orbital region) more than half fronto-orbital width (frontal width of carapace, including orbital region) (orbits relatively small); with curve or weak angle between frontal border and orbits; anterolateral margin with 1 or 2 teeth behind orbital angle. Epistome not visible in dorsal view. Pterygostomial region smooth. Abdominal somite 3 of male not wider than other somites. Telson of mature females twice as broad as long. Paragrapsus gaimardii (H. Milne Edwards, 1837) (Figs 159d, 160a, Pl. 29a). Lateral margin of carapace with 2 teeth behind orbital angle. First walking leg with felting restricted to inner edge of propodus and dactylus. Suture between sternites 1 and 2 with prominent ridge. Yellow-brown, with dark red spots. 45 mm. NSW (S of Narooma), Vic., Tas., SA: intertidal to 1.5 m depth, estuaries and sheltered coasts, under stones or in burrows, common. Paragrapsus laevis (Dana, 1851) (Fig. 160b, Pl. 29b). Lateral margin of carapace with 2 teeth behind orbital angle. First walking legs with extensive felting on anterior surface of carpus, propodus and dactylus. Suture between sternites 1 and 2 without ridge.Yellow-brown, with dark red spots. 41 mm. Qld (S of Moreton Bay), Vic. (W to Warrnambool), Tas.; mid-intertidal on sheltered coasts. Paragrapsus quadridentatus (H. Milne Edwards, 1837) (Figs 159e, 160c, Pl. 29c). Lateral margin of carapace with 1 tooth behind orbital angle. First walking leg with sparse felting restricted to inner edge of propodus and some grooves on dactylus. Suture between sternites 1 and 2 without ridge. Greenish or brownish grey. 32 mm. Vic. (W of Mallacoota), Tas., SA (E to Cape Marino); lower intertidal, under rocks on sheltered and exposed coasts, very common.

Subfamily Grapsinae MacLeay, 1838

Ilyograpsus Barnard, 1955 The genus was included in Varuninae by Davie (2002) but I agree with Crosnier (1965) that the absence of subparallel infraorbital margin and suborbital crest excludes it from that subfamily. The presence in the female of spoon-shaped fingers on the cheliped suggest a strong affinity to ocypodids. Diagnosis. Carapace subquadrate, widest in more posterior half; anterolateral margins slightly oblique, dentate; frontal width (frontal width of carapace, excluding orbital region) much less than half fronto-orbital width (frontal width of carapace, including orbital region) (orbits relatively wide); infraorbital margin running vertically from orbital angle to meet end of suborbital crest directly, no overlap. Maxilliped 3 merus wider than long. Male cheliped with acute fingers, female with spoon-shaped fingers. Ilyograpsus paludicola (Rathbun, 1909) (Fig. 161a). Carapace with broad orbital tooth, square first anterolateral tooth, longer acute second anterolateral tooth, and minor teeth and notches on posterolateral margin. 9 mm. South Africa, Indian Ocean, Malaysia, Qld, NSW; estuarine mangrove sediments. The species often co-occurs with, and is easily confused with, the ocypodid Enigmaplax littoralis (see Davie, 1993b); both occur in Port Hacking, NSW. Crosnier (1965) discussed and illustrated this species, in particular its infraorbital structure, and placed it in Grapsinae rather than Varuninae.

505 Marine Decapod Crustacea of Southern Australia

a b c

d e h

Fig. 161. Grapsidae Grapsinae. a, Ilyograpsus paludicola. b, Leptograpsodes octodentatus. c, Leptograpsus variegatus. d, Pachygrapsus laevimanus. e, Planes major. Left orbit and antennae: f, Leptograpsodes octodentatus. g, Leptograpsus variegatus. h, Pachygrapsus laevimanus.

Leptograpsodes Montgomery, 1931 The genus is monotypic, in spite of several available names, and restricted to southern Australia. The orbital structure, maxillipeds and male gonopods are very similar to that of the varunine genus Brachynotus but two pairs of postfrontal lobes and tuberculation on the anterior top of the carapace and striations on the meri of the walking legs are more grapsine (Griffin, 1969). Diagnosis: Carapace with strongly convex lateral margins, with 1 prominent tooth immediately behind orbital angle and 2 or 3 small teeth more posterior; dorsally a pair of oblique lateral grooves meeting in transverse groove. Infraorbital margin continuous from near base of orbital spine to near inner limit of orbit, paralleled by more dominant suborbital crest ending on pterygostomial face. Chelae smooth on outer surface, tuberculate on inner surface. Telson of male narrower than abdominal somites. Leptograpsodes octodentatus (H. Milne Edwards, 1837) burrowing shore crab (Figs 161b, f, 162d, e, Pl. 29d) Fixed finger of cheliped bent strongly downwards. Carapace dark brown to purple, with extensive yellow mottling. 70 mm. NSW (S of Cape Jervis), Vic., Tas., SA, WA (N to Abrolhos Is); supratidal, boulder beaches and rocky platforms. This is a large fast moving crab of the upper shore, usually found well above high tide mark and often in burrows near fresh or brackish water. It is highly resistant to desiccation. It usually leaves the burrow to forage at night on the beach (Hale, 1927; George, 1962). While the species has been figured and described in detail its subfamily position remains a puzzle (Montgomery, 1931; Griffin, 1969).

506 Brachyura – crabs

c d b

Fig. 162. Grapsidae Grapsinae. a, Leptograpsodes octodentatus (chela, lateral). b, Pachygrapsus laevimanus (walking leg). c, Planes major (walking leg). d, Leptograpsodes octodentatus (maxilliped 3). e, Leptograpsodes octodentatus (male abdomen). f, Leptograpsus variegatus (male abdomen).

Leptograpsus H. Milne Edwards, 1853 The genus is said to contain a single species widespread in southern Australia, New Zealand, Peru and Chile, and in numerous islands in the South Pacific (Griffin, 1973). Campbell & Mahon (1974) found on the basis of morphological analysis that the orange and blue colour morphs in Australia warrant separate specific status but none of the eight available specific names have been applied. Diagnosis. Carapace depressed and strongly granulate in frontal region; 2 parallel longitudinal grooves in gastric region and 1 transverse groove in cardiac region; 8–10 oblique ridges cover all but central region; lateral margin with 2 strong teeth immediately behind outer orbital angle. Infraorbital margin with deep notch near orbital tooth; suborbital crest denticulate, merges with infraorbital margin. All articles of walking legs flattened, spinose. Abdomen of male regularly triangular. Leptograpsus variegatus (Fabricius, 1793) (Figs 161c, g, 162f). Carapace usually purple with lighter flecks; juveniles more uniform dark grey. 80 mm. Indo-West Pacific, Qld, NSW, Vic., Tas., SA, WA (N to Shark Bay); high intertidal exposed rocky coasts. This fast-moving crab occurs in the upper intertidal region on more exposed rocky coasts than Leptograpsodes octodentatus, is more rectangular in shape, and does not have the deflexed cheliped fingers. The species feeds on algae, limpets and barnacles (Skilleter & Anderson, 1986).

Pachygrapsus Randall, 1840 Of about a dozen species of Pachygrapsus, three occur in Australia. Useful keys do exist (Tesch, 1918; Crosnier, 1965; Sakai, 1976) but some names have changed since these were published. Diagnosis. Front of carapace depressed; frontal width (frontal width of carapace, excluding orbital region) half or more fronto-orbital width (frontal width of carapace, including orbital region) (orbits relatively small); frontal and anterolateral regions with faint lateral lines; lateral margin

507 Marine Decapod Crustacea of Southern Australia

almost straight. Chelipeds with dark, spooned tips. Posterior surface of propodus of walking legs with dense mat of hairs (most conspicuous on walking legs 2). Abdomen of male regularly triangular. Pachygrapsus laevimanus Stimpson, 1858 (Fig. 161d, h, 162 b). Lateral margin with 1 postorbital tooth. Red spots arranged in a pattern of transverse striations. 16 mm. Qld, NSW, Vic. (W to Western Port); intertidal rocky shores, under rocks. The species was known for many years as P. transversus Gibbes, 1850, a name now restricted to an American species.

Planes Bowdich, 1825 Chace (1951) recognised two similar species, both of which have a long list of junior synonyms, but for the one found in southern Australia he chose the wrong one. What has long been known in the Pacific as Planes cyaneus Dana 1853 should be called P. major. Both species are pelagic, living on drifting wrack on the ocean and rare close to shore and in southern latitudes. Occasionally individuals will wash ashore, especially on the eastern seaboard of Australia. Diagnosis. Carapace dorsally smooth and strongly convex; frontal region slightly depressed; lateral margins convex. Chelipeds with dark, spooned tips. Carpus, propodus and dactylus of walking legs with fringe of long hairs on anterior margin. Abdomen of male regularly triangular. Planes major (Macleay, 1838) (Figs 161e, 162c). Convex carapace; flattened, hairy walking legs. Carapace mottled with light green-yellow on darker green background. 25 mm. Atlantic and Pacific Oceans; Qld, NSW, Tas. (Flinders I.); pelagic in oceanic waters on flotsam. There few records of this species in southern Australia; the most southern is at 40°S from a glass net float found on Flinders Island. There have been no records from Western Australian waters.

Subfamily Sesarminae Dana, 1851 Members of the subfamily Sesarminae have a conspicuous transverse hairy ridge on the outer surface of maxilliped 3. Frequently the pterygostomial region is covered with a reticulated pattern of granulae. This is the most diverse of the four subfamilies, with eight species recorded from southern Australia.

Parasesarma de Man, 1895 Parasesarma was treated as a subgenus by Sakai (1976). Five species occur in Australia. Diagnosis. Carapace about as wide as long; front deflexed and bordered with a sharp postfrontal ridge; anterolateral corner slightly acute; lateral border smooth behind orbital angle, with about 8 diagonal ridges connecting with lateral margin. Infraorbital margin open, so that flagellum of antenna 2 not excluded from orbit. Cheliped with 2 or 3 pectinate crests on upper surface; opposing margins of fixed and moveable fingers tuberculate. Pterygostomial region with granules and setae arranged in regular reticulate pattern. Parasesarma erythodactyla (Hess, 1865) (Fig. 163a, c, Pl. 29e). Male cheliped palm with 2 oblique pectinate crests. Walking legs with a felt of short setae on the anterior margins of the carpi and propodi. Green-black–bright green, with characteristic bright orange chelipeds. 25 mm. Qld, NSW, Vic. (W to Western Port); mangroves and mudflats. The species has been referred to in the literature as Sesarma erythrodactyla, that is, in the wrong genus and the species name spelled not as Hess wrote it, whatever he intended! The reticulate pattern on the pterygostomial region (the front of the carapace) is shared with Perisesarma, both genera having been subgenera of Sesarma in the past. Records of this species from Japan and Taiwan are now

508 Brachyura – crabs

b e a

c d

Fig. 163. Grapsidae Sesarminae. a, Parasesarma erythodactyla. b, Perisesarma longicristatum. c, Parasesarma erythrodactyla (anterior view). Varuninae. d, e, Brachynotus spinosus (carapace, chela).

referred to another species (Davie, 1993a). The crab feeds on decaying mangrove leaves (Camilleri, 1989).

Perisesarma de Man, 1895 In most earlier literature species of Perisesarma were treated as members of the subgenus Sesarma (Chiromanthes) (Davie, 2002; Rahayu & Davie, 2002). Campbell (1967) provided a key and described new Australian species under that name. Diagnosis. Carapace wider than long; front deflexed and bordered with a sharp postfrontal ridge; anterolateral corner slightly acute; lateral border with 1 tooth behind orbital angle, with diagonal ridges connecting with lateral margin. Infraorbital margin open, so that flagellum of antenna 2 not excluded from orbit. Cheliped with propodus with 2 or 3 transverse pectinate crests; dactylus with stridulatory tuberculation. Pterygostomial region with granules and setae arranged in regular reticulate pattern. Perisesarma longicristatum (Campbell, 1967) (Fig. 163b). Cheliped dactylus with 8 or 9 asymmetrical tubercles, most distal not elongate; distal propodal crest with c. 25 teeth. Carapace with scattered tufts of hair. 20 mm. NT, Qld, NSW; high intertidal, mangrove and muddy sediments, burrowing.

Subfamily Varuninae Alcock, 1900 Only one species of varunine grapsid is native to southern Australia. Three east Asian species have been introduced to the eastern USA and Europe and might be anticipated in Australia (see Box: Grapsid crabs with potential for introduction to Australia).

509 Marine Decapod Crustacea of Southern Australia

Brachynotus De Haan, 1833 The genus Brachynotus includes over a dozen species (according to Balss, 1957) or four restricted to the western Atlantic and Mediterranean (Schubart et al., 2001) but the generic concept seems rather vague. The latter view would exclude the Australian species. Tesch (1918) provided a key to Indo-West Pacific species from which the genus and species diagnoses are derived. Diagnosis. Carapace wider than long; frontal width (frontal width of carapace, excluding orbital region) half fronto-orbital width (frontal width of carapace, including orbital region); widest in more posterior half; anterolateral margins slightly oblique, with 3 blunt teeth. Merus of maxilliped 3 as long as wide, not expanded distolaterally; exopod narrow. Brachynotus spinosus (H. Milne Edwards, 1853) little shore crab (Fig. 163d, e, Pl. 29f). Carapace finely granulate anteriorly, strongly depressed, with 2 postfrontal lobes; with 3 teeth on anterolateral margin behind orbital angle. Walking legs without setae, merus with strong distal spine on upper margin. Brown-green with lighter mottling, sometimes with banded legs. 17 mm. South-western Pacific, Vic., Tas., SA; sheltered coasts especially in bays, intertidal to 10 m depth. The thick mat of hairs on both inner and outer surfaces of the fingers of the chelipeds are unique among southern Australian grapsids (Griffin, 1969).

References Balss, H. 1957. Decapoda. Dr. H.G. Bronn’s Klassen und Ordnungen des Tierreichs 1: 1505–1672. Camilleri, J. 1989. Leaf choice by crustaceans in a mangrove forest in Queensland. Marine Biology 102: 453–459. Campbell, B.M. 1967. The Australian Sesarminae (Crustacea: Brachyura): five species of Sesarma (Chiromantes). Memoirs of the Queensland Museum 15: 1–19. Campbell, B.M., & Griffin, D.J.G. 1966. The Australian Sesarminae (Crustacea: Brachyura): genera Helice, Helograpsus nov., Cyclograpsus, and Paragrapsus. Memoirs of the Queensland Museum 14: 127–174, pls 20–23. Campbell, N.A., & Mahon, R.J. 1974. A multivariate study of variation in two species of rock crab of the genus Leptograpsus. Australian Journal of Zoology 22: 417–425. Chace, F.A. 1951. The oceanic crabs of the genera Planes and Pachygrapsus. Proceedings of the United States National Museum 101: 65–103. Crosnier, A. 1965. Crustacés Décapodes Grapsidae et Ocypodidae. Faune de Madagascar 18: 1–143, pls 1–11. Davie, P.J.F. 1993a. A new species of sesarmine crab (Brachyura: Grapsidae) from Japan and Taiwan, previously known as Sesarma erythodactyla Hess, 1865. Crustacean Research 22: 65–74. Davie, P.J.F. 1993b. A new genus of macrophthalmine crab (Crustacea: Decapoda: Ocypodidae) from eastern Australia. Records of the Australian Museum 45: 5–9. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K. (eds), Zoological Catalogue of Australia. Vol.19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. George, R.W. 1962. The burrowing shore crab of southern Australia. Australian Natural History 14: 71–74. Griffin, D.J.G. 1969. Notes on the taxonomy and zoogeography of the Tasmanian grapsid and ocypodid crabs (Crustacea, Brachyura). Records of the Australian Museum 27: 323–347. Griffin, D.J.G. 1971. The ecological distribution of grapsid and ocypodid shore crabs (Crustacea: Brachyura) in Tasmania. Journal of Animal Ecology 40: 597–621. Griffin, D.J.G. 1973. A revision of the two southern temperate shore crabs Leptograpsus variegatus (Fabricius) and Plagusia chabrus (Linnaeus) (Crustacea, Decapoda, Grapsidae). Journal of the Royal Society of New Zealand 3: 415–440.

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Hale, H.M. 1927. The Crustaceans of South Australia. Part 1. South Australian Government Printer: Adelaide. 201 pp. Hawkins, A.J.S., Jones, M.B., & Marsden, I.D. 1982. Aerial and aquatic respiration in two mud crabs, Helice crassa Dana (Grapsidae) and Macrophthalmus hirtipes (Jacquinot) (Ocypodidae), in relation to habitat. Comparative Biochemistry and Physiology 73A: 341–347. Jessop, M. 1995. Hybridization between two species of grapsid crab in the Westernport Bay area, Victoria. B.Sc.(Hons) thesis. University of Melbourne. 32 pp. Jones, M.B. 1976. Limiting factors in the distribution of intertidal crabs (Crustacea: Decapoda) in the Avon-Heathcote estuary, Christchurch. New Zealand Journal of Marine and Freshwater Research 10: 577–587. Jones, M.B. 1980. Reproductive ecology of the estuarine burrowing mud crab Helice crassa (Grapsidae). Estuarine and Coastal Marine Science 11: 433–443. Jones, M.B. 1981. Effect of temperature, season, and stage of life cycle on salinity tolerance of the estuarine crab Helice crassa Dana (Grapsidae). Journal of Experimental Marine Biology and Ecology 52: 271–282. Jones, M.B., & Simons, M.J. 1983. Latitudinal variation in reproductive characteristics of a mud crab, Helice crassa (Grapsidae). Bulletin of Marine Science 33: 656–670. Kitaura, J., Wada, K., & Nishida, M. 2002. Molecular phylogeny of grapsoid and ocypodoid crabs with special reference to the genera Metaplax and Macrophthalmus. Journal of Crustacean Biology 22: 682–693. Montgomery, S.K. 1931. Report on the Crustacea Brachyura of the Percy Sladen Trust expedition to the Abrolhos Islands under the leadership of Prof. W.J. Dakin in 1913, along with other crabs from Western Australia. Journal of the Linnean Society of London (Zoology) 37: 405–464, pls 24–30. Rahayu, D.L., & Davie, P.J.F. 2002. Two new species and a record of Perisesarma (Decapoda, Brachyura, Grapsidae, Sesarminae) from Indonesia. Crustaceana 75: 597–608. Rice, A.L. 1980. Crab zoeal morphology and its bearing on the classification of the Brachyura. Transactions of the Zoological Society of London 35: 271–424. Sakai, K. 1976. Crabs of Japan and Adjacent Seas. Kodansha Ltd: Tokyo. 773 pp. plus volume of 251 plates pp. Schubart, C.D., Cuesta, J.A., Diesel, R., & Felder, D. 2000. Molecular phylogeny, taxonomy, and evolution of nonmarine lineages within the American grapsoid crabs (Crustacea: Brachyura). Molecular Phylogenetics and Evolution 15: 179–190. Schubart, C.D., Cuesta, J.A., & Rodríguez, A. 2001. Molecular phylogeny of the crab genus Brachynotus (Brachyura: Varuninae) based on the 16S rRNA gene. Hydrobiologia 449: 41–46. Skilleter, G.A., & Anderson, D.T. 1986. Functional morphology of the chelipeds, mouthparts and gastric mill of Ozius truncatus (Milne Edwards) (Xanthidae) and Leptograpsus variegatus (Fabricius) (Grapsidae) (Brachyura). Australian Journal of Marine and Freshwater Research 37: 67–79. Smith, T.J., Boto, K.G., Frusher, S., & Giddins, R.I. 1991. Keystone species and mangrove forest dynamics: the influence of burrowing by crabs on soil nutrient status adn forest productivity. Estuarine and Coastal Marine Science 33: 419–432. Sternberg, R.v., & Cumberlidge, N. 1998. Taxic relationships within the Grapsidae Macleay, 1838 (Crustacea: Decapoda: Eubrachyura). Journal of Comparative Biology 3: 115–136. Taylor, H.H., Forster, M.E., Jones, M.B., & Marsden, I.D. 1984. Aquatic and aerial respiration in a high shore crab, Cyclograpsus lavauxi. Proceedings of the Physiological Society of New Zealand 4: 40. Tesch, J.J. 1918. Decapoda F. (Decapoda Brachyura continued) Hymenosomatidae, Retropilumnidae, Ocypodidae, Grapsidae and Gecarcinidae. Siboga Expéditie Monographie 39c: 1–148, pls 1–6. Türkay, M. 1978. Zwei neue Grapsiden-Arten aus dem Indopazifik (Euchirograpsus madagariensis n. sp.) mit Einführung von Miersiograpsus n. gen. (Crustacea: Decapoda: Grapsidae). Senckenbergiana Biologica 59: 133–141. Wescott, G.C. 1979. Analysis of marine hybridizaton between two species of the genus Cyclograpsus (Crustacea: Brachyura) in south-eastern Australia. Australian Journal of Marine and Freshwater Research 30: 469–484.

511 Marine Decapod Crustacea of Southern Australia

GRAPSID CRABS WITH POTENTIAL FOR INTRODUCTION TO AUSTRALIA Three species of grapsid crabs have been introduced to Europe and North America from eastern Asia and might be expected to arrive in southern Australia in the future. All live within the temperature ranges experienced here and are suspected to have been transported by shipping over the last few decades. All are members of the subfamily Varuninae distinguishable from the only native species, Brachynotus spinosus, by their more oval shape. All have larger lateral teeth than B. spinosus. Eriocheir sinensis Milne Edwards, 1853, the Chinese mitten crab, has four sharp spines on the front and four strong lateral spines. The mitten crab is so-called because of the thick coating of long hairs over the chelipeds. The crab grows to 70 mm wide. The species is native to China and Korea between 26° and 40°N where their normal habitat is the bottoms and banks of freshwater rivers and estuaries. The crab was found in 1912 in a tributary of the River Aller, Germany and spread rapidly during the 1930s to low-lying areas of Germany, Scandinavia, Holland, France, Belgium and the UK. In North America, specimens were first seen in the Detroit River, Ontario, Canada in 1965 and in Lake Erie in 1973. It reached California in 1992. The mitten crab is considered edible in its home countries. Unlike the following species, it would be expected in southern Australia in estuarine and freshwater environments rather than marine. Hemigrapsus sanguineus (De Haan, 1835) has a 30-mm wide square carapace with three spines on each side. Colour is variable, green to purple to orange-brown to red, with bands along its legs and red spots on the cheliped claws. Males crabs have a distinctive fleshy, bulb-like structure at the base of the dactylus of the cheliped. There is a finely striated ridge under the eye and lateral margin. The species is a native of Japan, where it is one of the commonest intertidal species, Korea and China. The species lives on exposed rocky coasts. It has been introduced to the Atlantic coast of the USA where it was first discovered in 1988, France and The Netherlands on the Atlantic European coast and to Croatia in the Mediterranean. The species is now well established from Maine to North Carolina and breeds in the USA. Hemigrapsus penicillatus De Haan 1835 is similar to H. sanguineus, but the male has a tuft of hair at the base of the dactylus cheliped instead of a fleshy bulb. The suborbital ridge is smooth but in three unequal parts. The species is a native of Japan, Korea, China and Hawaii where it favours more protected shores of lagoons and estuaries. The crab was recorded for the first time in north-western France in 1994 and in The Netherlands in 2000.

Plagusiidae Dana, 1851 The deep antennular notches dividing the front of the carapace immediately define the most well known plagusiids in southern Australia. Plagusia chabrus, the active rock crab of exposed shores is its most frequently seen member. Until recently, species of Plagusia were treated as members of Grapsidae, the family that includes most shore crabs. Now however, the genus and Percnon (not seen in southern Australia) are thought sufficiently different to warrant being placed in another family (Sternberg & Cumberlidge, 1998). Inclusion of Miersiograpsus and one other genus (which lack the antennal notches) in the same family is based in part on molecular evidence (Cuesta & Schubart, 1997), and on the unique fused male abdominal somites 3–6 which lock on to a button on the sternite (Davie, 2002).

512 Brachyura – crabs

Diagnosis. Carapace subcircular to quadrate; surface smooth or tuberculate and tomentose. Front either narrow, not overhanging epistome, with 3 lobes separated by antennular notches, such that antennae 1 are visible in dorsal view; or broader, overhanging epistome, with only shallow antennular grooves. Anterolateral margins with 1 to several teeth or spines behind exorbital angle. Maxilliped 3 not completely closing buccal cavern; merus and ischium without an oblique setose crest; exopod slender; palp articulating near anteroexternal angle of merus. Male abdomen entirely covering sternum between last pair of legs; somites 3–6 or 3–5 fused. Abdominal locking mechanism of press-button type.

Key to southern Australian species of Plagusiidae 1. Antennules transverse, in groove under front ...... Miersiograpsus australiensis — Antennules extending vertically into notch reaching surface of carapace . . . . . Plagusia … 2 2. Carapace tuberculate ...... Plagusia speciosa — Carapace not tuberculate, smooth or tomentose ...... 3 3. Carapace and abdomen with dense felt of fine hairs; walking legs with numerous short spines on upper edge of merus ...... Plagusia chabrus — Carapace and abdomen smooth, without hairs; walking legs with single spine on upper edge of merus ...... Plagusia glabra

Miersiograpsus Türkay, 1978 Griffin & Brown (1976) reported the only Australian species from deep water off NSW as a species of the grapsid genus, Litocheira, which it superficially resembles. There is only one other species (Türkay, 1978). Diagnosis. Carapace without velvety covering, but with scattered long stiff setae. Anterolateral margin with 2 teeth, including exorbital tooth. Miersiograpsus australiensis Türkay, 1978 (Fig. 164e). Chela long and narrow, palm almost twice as long as high. Frontal carapace lobes separated by a broad U-shaped groove. 9.5 mm. New Caledonia, NSW; 459–825 m depth (Crosnier, 2001).

Plagusia Latreille, 1804 Plagusia is a genus of eight species and Dawson’s (1987) key is useful in spite of some incorrect names and synonymies (Davie, 2002). Three species occur in southern Australia, readily distinguished by the antennal clefts in the anterior margin of the carapace. Oceanic dispersal of some species of Plagusia, often found clinging to hulls of ships and driftwood, accounts for their wide distribution (Dawson, 1987). Diagnosis. Carapace length approximately equal to width, lateral margins dentate behind orbital angle and slightly divergent to base of third pair of walking legs. Antennules visible dorsally in clefts in anterior margin of carapace between orbits. Merus of maxilliped 3 as broad as ischium. Plagusia chabrus (Linnaeus, 1758) red rock crab, cleft-fronted shore crab (Fig. 164a, b, Pl. 29g). Carapace and abdomen with dense felt of fine hairs. Chela palm strongly granulate, granulae arranged in longitudinal lines. Walking legs with numerous small spines on upper margin of merus, with smooth ridges separating grooves felted with fine hairs. Abdomen of male tapering regularly towards telson. Orange-red, darker crimson on felted areas of carapace and walking legs. 73 mm. Indo-West Pacific, New Zealand, Qld (N to Rockhampton), NSW, Vic., Tas., SA, WA (N to Shark Bay); exposed lower intertidal rocky shores to 50 m depth. The red

513 Marine Decapod Crustacea of Southern Australia

rock crab is active and can seen out of the water at extreme low water or by SCUBA divers on wharf piles and in rock crevices which it often shares with rock lobsters (Jasus edwardsii). Plagusia glabra Dana, 1852 (Fig. 164c, d). Carapace and abdomen smooth, without hairs. Chela palm with sparse, small granulae dorsally, otherwise smooth. Walking legs with 1 spine on upper margin of merus, carpus, propodus and dactyl with narrow lines of long hairs, without ridges and felted grooves. Abdominal somites 3–5 of male with almost parallel sides. Deep red, carapace patterned with intricate red maze on a lighter fawn to orange background. 50 mm. Southern Qld, NSW, Vic. (W to Wilsons Promontory); intertidal.

e d

Fig. 164. Plagusiidae. a, b, Plagusia chabrus (with chela). c, d, Plagusia glabra (carapace, chela). e, Miersiograpsus australiensis.

514 Brachyura – crabs

Plagusia speciosa Dana, 1851. Carapace covered with squamiform tubercules not fringed with short hairs, under side of body smooth. Walking legs with merus with single row of long hairs, with lobe on coxa not dentate. Abdominal somite 7 of male rounded, of female pointed. 45 mm. Indo-West Pacific, NT, Qld, NSW; intertidal and subtidal, also on flotsam.

References Crosnier, A. 2001. Grapsidae (Crustacea, Decapoda, Brachyura) d’eau profonde du Pacifique sud-ouest. Zoosystema 23: 783–796. Cuesta, J.A., & Schubart, C.D. 1997. The first zoeal stage of Glyptograpsus impressus, with comments on the subfamilial arrangement of Grapsidae (Crustacea: Brachyura). Cahiers de Biologie Marine 38: 291–299. Davie, P.J.F. 2002. Crustacea: Malacostraca: Eucarida (Part 2): Decapoda – Anomura, Brachyura. In: Wells, A., & Houston, W.W.K., Zoological Catalogue of Australia. Vol. 19.3B. CSIRO Publishing: Melbourne. xiv, 641 pp. Dawson, E.W. 1987. A key to the world species of Plagusia (Crustacea: Brachyura), with a new record of P. depressa tuberculata Lamarck from New Zealand. National Museum of New Zealand Records 3: 37–45. Griffin, D.J.G., & Brown, D.E. 1976. Deepwater decapod Crustacea from eastern Australia: brachyuran crabs. Records of the Australian Museum 30: 248–271. Sternberg, R.v., & Cumberlidge, N. 1998. Taxic relationships within the Grapsidae Macleay, 1838 (Crustacea: Decapoda: Eubrachyura). Journal of Comparative Biology 3: 115–136. Türkay, M. 1978. Zwei neue Grapsiden-Arten aus dem Indopazifik (Euchirograpsus madagariensis n. sp.) mit Einführung von Miersiograpsus n. gen. (Crustacea: Decapoda: Grapsidae). Senckenbergiana Biologica 59: 133–141.

515 This page intentionally left blank 12. STOMATOPODA – MANTIS SHRIMPS Shane T. Ahyong

The order Stomatopoda includes the crustaceans commonly known as mantis shrimps or mantis prawns. The carapace is broad and shield-like, and the body is elongate and usually dorsoventrally flattened. Unlike decapods, the pleopods of stomatopods are broad and carry foliaceous gills. One of the most distinctive features of the modern stomatopods, however, is the second maxilliped which is greatly enlarged as a raptorial claw. The raptorial claws may be divided into two functional forms: ‘spearers’ and ‘smashers’ (Caldwell & Dingle, 1976). ‘Spearing’ claws have a series of long, curved teeth on the dactylus on which soft-bodied prey is impaled. ‘Smashing’ claws have an inflated, calcified heel at the base of the dactylus which is used as a hammer to break open hard bodied prey such as snails and crabs. As a rule, stomatopods are cryptic, benthic predators and are seldom encountered despite their abundance. Little is known of the biology of Australian species. Vision in stomatopods is remarkably well-developed (Caldwell, 1991). In most species, the cornea is divided in half by a central band of ommatidia such that each eye is capable of binocular vision. Further, this central band of ommatidia is specialised for colour vision and detecting polarised light (Marshall, 1988; Marshall et al., 1991). Stomatopods live in burrows or crevices intertidally and subtidally on all types of substrate. Most gonodactyloids occur on hard or rough substrates and usually live in preformed rock or coral crevices. Squilloids, eurysquilloids, parasquilloids and lysiosquilloids burrow in soft, level substrates. Most occur in shallow water, but species of the superfamily Bathysquilloidea occur in deep water, down to 1500 m (Manning, 1991). Whereas most stomatopods are solitary, lysiosquil- lids and several nannosquillids live in monogamous pairs. In stomatopods, sexes are readily distinguished by external features. Males have a pair of long, slender penes articulating at the bases of the last pereopods (third walking legs). The female gonopores are located on the sternum between the first pereopods (first walking legs). Sexual dimorphism is present in many species and is usually marked in mature males by inflation of the raptorial claws and dorsal armature of the telson, giving the males a slightly more robust, bulkier appearance than females. Most decapods carry eggs on the pleopods, but female stomatopods carry the egg mass with the maxillipeds until hatching. Stomatopods do not have a nauplius larval stage. Initial larval stages are either benthic or pelagic but all subsequent stages are pelagic. Lysiosquilloid larvae hatch at the earliest stage of development, known as antizoeae, bearing five pairs of biramous thoracic appendages, but without abdominal appendages. The antizoea develops into an erichthus, bearing two or fewer intermediate denticles on the telson and pleopods appearing progressively from front to rear. Squilloid and gonodactyloid larvae hatch as a pseudozoea, bearing two pairs of uniramous thoracic appendages and four or five pairs of pleopods respectively. Squilloid pseudozoeae develop into an alima, a stage with four or more intermediate denticles on the telson, whereas gonodactyloid pseudozoeae develop into an erichthus larva (Tirmizi & Kazmi, 1984). Stomatopod larvae may be abundant in the plankton, particularly during summer, and may be important prey for pelagic fish. Settling postlarvae resemble and assume the lifestyle of adults. Larval development of

517 Marine Decapod Crustacea of Southern Australia

members of the other superfamilies, Erythrosquilloidea, Eurysquilloidea, Bathysquilloidea and Parasquilloidea, is poorly known. Stephenson & McNeill (1955) and Manning (1966) provided the first major studies of Australian stomatopods. Since then, the systematics of the Stomatopoda has been revised considerably (e.g. Manning, 1980, 1995; Ahyong & Harling, 2000; Ahyong, 2001). The most recent phylogenetic study (Ahyong & Harling, 2000) and taxonomic study of the Australian Stomatopoda (Ahyong, 2001) recognised seven superfamilies and 17 families and 146 species from Australia. Ahyong (2001) provided keys to the world genera and all species in genera represented in Australian waters. More than 480 species are known worldwide and nearly one-third of these occur in Australian waters. Thirty-seven species are known from the southern half of the continent between Perth in the west and Newcastle in the east. Approximately half of the southern species are temperate water endemics, whereas the other half comprises more widespread tropical species. Owing to the effect of the southern flowing East Australian and Leeuwin Currents, additional future records of tropical species from temperate waters are likely. Lengths given are maximum adult total length measured dorsally along the midline from the tip of the rostral plate to the apices of the submedian teeth of the telson. General morphological features are illustrated in Fig. 165. Diagnosis. Malacostraca with cephalothorax partly covered by carapace. Ophthalmic and antennular somites articulated. Eyes stalked. Antennules triflagellate. With 5 pairs of maxillipeds, 6- segmented, uniramous, subchelate. Second maxilliped in extant forms greatly enlarged to form raptorial claw. Pereopods biramous, endopod reduced, protopod 3-segmented. Pleopods biramous; rami flap-like; exopod bearing gills.

Key to superfamilies of Stomatopoda 1. Propodi of maxillipeds 3–4 subquadrate, broader than long, usually with ventral ribbing (Fig. 165f) ...... 2 — Propodi of maxillipeds 3–4 ovate, usually longer than broad, without ventral ribbing (Fig. 165d, e) ...... 3 2. Telson with distinct, slender median carina. Dorsal processes of antennular somite short, anteriorly compressed, rounded laterally ...... Erythrosquilloidea ...p.520 — Telson without median carina, at most with low, median boss or swelling. Dorsal processes of antennular somite spiniform or dorsoventrally flattened, not anteriorly compressed or rounded laterally ...... Lysiosquilloidea ...p.526 3. All primary teeth of telson with movable apices ...... Bathysquilloidea ...p.520 — At most, submedian teeth of telson with movable apices ...... 4 4. Telson with 4 or more closely spaced intermediate denticles arranged in regular row ...... Squilloidea ...p.535 — Telson with no more than 3 (usually 2) ‘intermediate’ denticles ...... 5 5. Body depressed. [Dactylus of raptorial claw with 4 or more well formed teeth] ...... Eurysquilloidea (not known from southern Australia) — Body subcylindrical, strongly convex ...... 6 6. Uropodal protopod terminating in 3 primary spines. Cornea asymmetrically bilobed, with outer margin of eye longer than inner margin; with 2–3 rows of hexagonal ommatidia in the midband ...... Parasquilloidea ...p.535 — Uropodal protopod terminating in 1 or 2 primary spines. Cornea subglobular or symmetrically bilobed, with 6 rows of rectangular ommatidia in midband ...... Gonodactyloidea . . . p. 522

518 Stomatopoda – mantis shrimps

c b

e d f

Fig. 165. Stomatopoda. a, general morphology. b, raptorial claw. c, telson. d, third maxilliped (Squillidae). e, third maxilliped (Gonodactylidae). f, third maxilliped (Lysiosquillidae).

519 Marine Decapod Crustacea of Southern Australia

Superfamily Bathysquilloidea Manning, 1967 The superfamily comprises two families, of which only Bathysquillidae is known from Australian waters. Bathysquilloids occupy deep, outer-shelf habitats and for this reason, are seldom seen. Diagnosis. Propodi of maxillipeds 3–5 longer than broad, without ventral ribbing. Telson with distinct median carina; with 4 or 5 pairs or primary teeth, each with movable apices; intermediate denticles absent.

Key to southern Australian species of Bathysquilloidea 1. Rostral plate longer than broad. Eyes large, subglobular. Carpus of raptorial claw with 2 dorsal spines ...... Bathysquilla crassispinosa — Rostral plate broader than long. Eyes small, stubby. Carpus of raptorial claw with 1 dorsal spine ...... Bathysquilla microps

Bathysquillidae Manning, 1967 Bathysquillidae includes two genera, Bathysquilla and Altosquilla. Both genera are known from Australia, but only Bathysquilla is represented in southern waters. Diagnosis. Abdominal somite 5 without long dorsal median spine. Telson broader than long. Uropodal exopod with distal and proximal segments fully articulating.

Bathysquilla Manning, 1963 Both of the two species of Bathysquilla are known from southern Australia. Diagnosis. Telson with 4 pairs of primary teeth on posterior margin. Bathysquilla crassispinosa (Fukuda, 1909) (Fig. 166a–c). Rostral plate longer than broad. Eyes large, subglobular. Raptorial claw with 2 dorsal spines on carpus. Lower posterior margins of abdominal somites not lined with short spines. 297 mm. Indo-West Pacific from Madagascar to Japan, Qld, NSW (S to Montague I.); 170–420 m depth. Bathysquilla microps (Manning, 1961) (Fig. 166d–f, Pl. 30a). Rostral plate broader than long. Eyes small, stubby. Raptorial claw with 1 dorsal spine on carpus. Lower posterior margins of abdominal somites lined with short spines. 255 mm. Western Atlantic, central to western Pacific including Philippines, New Zealand, Qld, NSW (S to Kiama); 604–1519 m depth.

Superfamily Erythrosquilloidea Manning & Bruce, 1984 The superfamily comprises one family, one genus and two species.

Diagnosis. Propodi of maxillipeds 3–5 subquadrate, with ventral ribbing. Telson with distinct median carina; intermediate denticles present; submedian teeth with movable apices.

Erythrosquillidae Manning & Bruce, 1984 Diagnosis. As for superfamily.

520 Stomatopoda – mantis shrimps

c a

Fig. 166. Bathysquillidae. a–c, Bathysquilla crassispinosa. d–f, Bathysquilla microps. a, d, anterior region. b, e, raptorial claw. c, f, posterior region.

521 Marine Decapod Crustacea of Southern Australia

Erythrosquilla Manning & Bruce, 1984 Of the two known species of Erythrosquilla, one is known from southern Australia. Diagnosis. As for superfamily. Erythrosquilla hamano Ahyong, 2001 (Fig. 172d–f). Dactylus of raptorial claw with 8 or 9 teeth. Telson with accessory median carina flanking median carina; with four triangular denticles between submedian and intermediate teeth. 27 mm. Japan, southern WA.

Superfamily Gonodactyloidea Giesbrecht, 1910 The gonodactyloids are principally tropical and are poorly represented in southern waters. Gonodactyloidea include both species that burrow in soft substrates and species that occupy preformed cavities in rock or coral. Gonodactylids and pseudosquillids may be found in preformed cavities or under rock and coral. Hemisquillids construct U-shaped burrows in sandy- mud. Ahyong (2001) provided keys to all gonodactyloid families and genera. Three gonodactyloid families are represented in southern Australia. Diagnosis. Propodi of maxillipeds 3–5 ovate, longer than broad, without ventral ribbing. Body subcylindrical, strongly convex in cross-section. Telson with distinct median boss or carina and 2 (rarely 3) intermediate denticles.

Key to southern Australian species of Gonodactyloidea 1. Rostral plate without long, slender median spine. Dactylus of raptorial claw without strongly inflated, calcified heel at base. Uropodal exopod with distal and proximal segments articulating terminally (end-to-end) ...... 2 — Rostral plate with long, slender median spine. Dactylus of raptorial claw with strongly inflated, calcified heel at base. Uropodal exopod with distal segment articulating on inner distal margin of proximal segment ...... Gonodactylidae … 3 2. Rostral plate triangular. Dactylus of raptorial claw without teeth on inner margin ...... Hemisquillidae … Hemisquilla australiensis — Rostral plate broadly ovate. Dactylus of raptorial claw with 3 slender teeth on inner margin ...... Pseudosquillidae … Pseudosquilla ciliata 3. Telson with 3 middorsal carinae. Ocular scales broader than high. Uropodal protopod without lobes between terminal spines ...... 4 — Telson with 5 middorsal carinae. Ocular scales small, narrow. Uropodal protopod with 1 or 2 lobes between terminal spines ...... Gonodactylaceus falcatus 4. Rostral plate with acute anterolateral angles; anterior margins straight; lateral margins strongly divergent ...... Gonodactylus smithii — Rostral plate with rounded anterolateral angles; anterior margins concave; lateral margins moderately divergent ...... Gonodactylus chiragra

522 Stomatopoda – mantis shrimps

Gonodactylidae Giesbrecht, 1910 Gonodactylids use their ‘smashing’ claws to break open hard-bodied prey such as gastropods and crabs. Gonodactylids are abundant in tropical waters, particularly on coral reefs where they live in or under rock and coral, and compete aggressively for living space. Gonodactylids include some of the most colourful of stomatopods, frequently with red, blue, or green highlighting. The three species reported below are rare in southern waters and are best considered extralimital. Diagnosis. Rostral plate with long median spine. Body subcylindrical, compact. Uropodal exopod with distal segment articulating on inner distal margin of proximal segment. Movable spines on outer margin of uropod exopod not recurved anteriorly. Raptorial claw with subterminal ischiomeral articulation; dactylus strongly inflated basally, inner margin unarmed.

Gonodactylaceus Manning, 1995 Ahyong (2001) recognised five species, all confined to the Indo-West Pacific. Diagnosis. Eye with subcylindrical cornea. Ocular scales small, narrower than basal width of rostral spine. Anterior margin of lateral plates of carapace convex, rounded, extending beyond base of rostral plate. Mandibular palp present. Telson with 5 middorsal carinae; intermediate carinae without accessory mesial, longitudinal carina. Gonodactylaceus falcatus (Forskål, 1775) (Fig. 167a–d). Rostral plate with divergent lateral margins; anterolateral angles blunt, rounded. Dorsal depression of merus of raptorial claw yellow in life with a black anterior spot. Telson with 5 middorsal carinae. Abdominal somites smooth, without transverse groove near posterior margin. Margin of uropodal endopod with single row of setae. 70 mm. Western Indian Ocean to Japan, Hawaii, New Caledonia, WA, NT, Qld, NSW (S to Sydney); shallow rocky and coral reefs.

Gonodactylus Berthold, 1827 The genus includes five Indo-West Pacific species of which two are known from southern Australia. Diagnosis. Eye with subcylindrical cornea. Ocular scales large, broader than basal width of rostral spine. Anterior margin of lateral plates of carapace convex, rounded, extending beyond base of rostral plate. Mandibular palp present. Telson with 3 middorsal carinae or bosses; intermediate carinae without accessory mesial, longitudinal carina. Gonodactylus chiragra (Fabricius, 1781) (Fig. 167e–g). Rostral plate with subparallel or slightly divergent lateral margins; anterolateral angles rounded; anterior margins lateral to apical spine concave. Dorsal depression of merus of raptorial claw white with a black spot anteriorly. Median carina of telson unarmed. 90 mm Tropical Indo-West Pacific, WA (S to Perth) NT, Qld, NSW (S to Sydney); shallow rocky and coral reefs. Gonodactylus smithii Pocock, 1893 (Fig. 167h–j). Rostral plate with strongly divergent lateral margins; anterolateral angles angular or blunt; anterior margins lateral to apical spine straight. Dorsal depression of merus of raptorial claw dark red to deep purple with white margin. Median carina of telson usually with posterior spine. 75 mm Tropical Indo-West Pacific, NT, Qld, NSW, Vic. (S to Bass Strait); shallow rocky and coral reefs.

523 Marine Decapod Crustacea of Southern Australia

b c

a d

Fig. 167. Gonodactylidae. a–d, Gonodactylaceus falcatus. e–g, Gonodactylus chiragra. h–j, Gonodactylus smithii. a, e, h, anterior region. b, f, i, ocular scales. d, raptorial claw. c, g, j, posterior region.

524 Stomatopoda – mantis shrimps

i h

Fig. 168. Pseudosquillidae. a–c, Pseudosquilla ciliata. d–f, Hemisquilla australiensis. Parasquillidae. g–i, Pseudosquillopsis dofleini. a, d, g anterior region. b, e, h, posterior region. c, f, i, raptorial claw.

Hemisquillidae Manning, 1980 The family includes a single genus and four species. Ahyong (2001) provided a key to all species. Unlike other families, species of Hemisquillidae occur only in temperate waters. Hemisquillids are active during the day and burrow in sandy substrates. One species is known from Australia. Diagnosis. Eye with subcuboidal cornea. Rostral plate triangular. Body strongly convex in cross- section, compact, non-carinate. Raptorial claw with terminal ischiomeral articulation; dactylus inflated basally, inner margin unarmed. Uropodal protopod terminating in 1 primary spine; exopod segments articulating terminally (end-to-end).

525 Marine Decapod Crustacea of Southern Australia

Hemisquilla Hansen, 1895 Hemisquilla australiensis Stephenson, 1967 (Fig. 168d–f, Pl. 30b). Rostral plate as long as or longer than broad. Raptorial claw propodus blue in life. Telson with 2 or 3 rounded lobes between submedian and intermediate teeth, at most with minute apical point. Lateral denticle absent, or represented at most by minute round tubercle. Uropodal exopod with 4 (rarely 3) movable spines on outer margin of proximal segment. 174 mm. New Zealand, central NSW, Vic., SA; 18–c. 480 m depth.

Pseudosquillidae Manning, 1977 Pseudosquillidae include four genera, principally restricted to the tropical Indo-West Pacific. Only the monotypic genus Pseudosquilla is represented in southern waters and is rarely encountered. All genera and species of Pseudosquillidae can be identified from Ahyong (2001). Diagnosis. Rostral plate broadly ovate. Body compact, subcylindrical, non-carinate. Raptorial claw ischiomeral articulation terminal; dactylus slender, uninflated basally, armed with 3 teeth. Uropodal protopod terminating in 2 primary spines; exopod segments articulating terminally (end-to-end).

Pseudosquilla Dana, 1852 Diagnosis. Eye with subcylindrical cornea. Rostral plate broadly ovate, without anterior spine. Carapace without large circular spots. Telson with 3 carinae lateral to median carina. Uropodal protopod with inner spine longer than outer. Pseudosquilla ciliata (Fabricius, 1787) (Fig. 168a–c). 100 mm. Tropical Atlantic, Indo-West Pacific, WA, NT, Qld, NSW (S to Sydney area); all types of substrate; shore to around 100 m depth.

Superfamily Lysiosquilloidea Giesbrecht, 1910 Members of this superfamily are easily recognised by their smooth, flattened dorsum. Many species bear alternating light and dark transverse banding. Lysiosquilloids seldom leave their burrows and are consequently less frequently encountered than squilloids which actively forage. They are ambush predators and feed on passing soft bodied prey such as small fish, cephalopods and shrimps. Three families are represented in southern waters. Diagnosis. Propodi of maxillipeds 3–5 subquadrate, as broad as long, usually ribbed ventrally. Telson without distinct median carina.

Key to southern Australian species of Lysiosquilloidea 1. Uropodal endopod with strong dorsal proximal fold ...... Nannosquillidae … 2 — Uropodal endopod without strong dorsal proximal fold ...... 9 2. Dactylus of raptorial claw with 4–6 teeth; ischium without distal spine. Posterior margin of telson with row of short teeth above ‘true’ marginal armature; 2 intermediate denticles present ...... 3 — Dactylus of raptorial claw with 8 or more teeth; ischium with distal spine. Posterior margin of telson without row of spinules above marginal armature; 4 intermediate denticles present ...... 4

526 Stomatopoda – mantis shrimps

3. Rostral plate with rounded anterolateral corners. Posteromedian projection of telson blunt, rounded ...... Hadrosquilla edgari — Rostral plate with angular anterolateral corners. Posteromedian projection of telson truncate, emarginate ...... Hadrosquilla perpasta 4. Rostral plate quadrate; with short median spine. Posteromedian projection of telson a short, slender spine ...... Austrosquilla osculans — Rostral plate cordiform or triangular. Posteromedian projection of telson blunt, rounded 5 5. Abdominal somite 6 with ventrolateral spine and triangular lobe anterior to uropod articulation ...... 6 — Abdominal somite 6 with either a ventrolateral spine or triangular lobe anterior to uropod articulation, never with both ...... 8 6. Ocular scale with blunt, rounded apex. Basal segment of pereopods 1–3 without slender, ventrally directed spine on inner or outer margins ...... Austrosquilla middletoni — Ocular scale with pointed apex. Basal segment of pereopods 1–3 with slender, ventrally directed spine on inner and outer margins ...... 7 7. Ocular scale a small slender spine. Telson intermediate denticles on same plane ...... Austrosquilla rachelae — Ocular scale triangular. Telson intermediate denticles on different planes ...... Austrosquilla melanocauda 8. Dactylus of raptorial claw with 11–13 teeth. Telson without distinct rounded lobe on lateral proximal margin ...... Austrosquilla vercoi — Dactylus of raptorial claw with 19–21 teeth. Telson with distinct rounded lobe on lateral proximal margin ...... Austrosquilla tsangi 9. Posterior margin of telson without rows of slender, sharp teeth ...... (Lysiosquillidae) Lysiosquilla colemani — Posterior margin of telson with rows of slender, sharp teeth ...... Tetrasquillidae ... 10 10. Rostral plate with median spine. Dactylus of raptorial claw with 6 or more teeth. Ventral surface of telson with strong postanal spine; 4 or more intermediate marginal denticles present ...... 11 — Rostral plate without median spine. Dactylus of raptorial claw with 5 teeth. Ventral surface of telson without postanal spine; 2 intermediate marginal denticles present ...... Heterosquilla pentadactyla 11. Rostral plate broader than long; median spine short, not extending beyond anterior margin of eyes; with ventral carina or spine ...... Acaenosquilla brazieri — Rostral plate longer than broad; median spine long, extending well beyond anterior margin of eyes; ventral carina absent ...... Kasim insuetus

Tetrasquillidae Manning & Camp, 1993 Ahyong (2001) recognised eight genera of tetrasquillids worldwide. Three genera are known from southern Australia. Diagnosis. Body depressed, compact. Pereopodal endopods elongate, ovate or subcircular. Uropodal endopod without strong, dorsal proximal fold. Raptorial claw dactylus not inflated basally, with 4 or more teeth on opposable margin; propodus fully pectinate. Telson with raised median prominence dorsally; submedian teeth with movable apices; posterior margin with rows of slender spines or denticles.

527 Marine Decapod Crustacea of Southern Australia

Acaenosquilla Manning, 1990 The genus includes two Indo-West Pacific species; one is known from Australia. Diagnosis. Eye with bilobed cornea, broader than stalk. Rostral plate broader than long; apical spine short; with ventral carina or spine. Raptorial claw dactylus with 6 teeth; ischium without outer distal spine. Pereopods 1–3 basal segment with ventrally directed posterior spine and angular lobe on inner margin. Abdominal somite 6 with posterolateral spines. Telson dorsal surface with posteriorly armed carinae and broad low median elevation terminating in 3 slender spines; posterior margin with 2 pairs of fixed primary teeth and 4 intermediate denticles; ventral surface with postanal spine. Acaenosquilla brazieri (Miers, 1880) (Fig. 169h–k). Cornea of eye with prominent mesial tubercle. Telson dorsal surface with at least 3 pairs of posteriorly armed carinae; posterior margin with spinules between dorsal and marginal spines. 100 mm. Central Qld, NSW (S to Port Jackson); to 60 m depth.

Heterosquilla Manning, 1962 The genus includes five species of which one is known from Australia. Diagnosis. Telson posterior margin with 2 pairs of fixed primary teeth; 2 or more intermediate denticles present; ventral surface without postanal spine. Heterosquilla pentadactyla Ahyong, 2001 (Fig. 169a–c). Eye partially concealed by rostral plate; cornea subglobular. Rostral plate trianguloid; apex faintly sulcate; with sinuous margins. Raptorial claw dactylus with 5 teeth; ischium without outer distal spine. Pereopods 1–2 basal segment with ventrally directed inner and outer spine; pereopod 3 with outer spine only. Abdominal somite 6 with short, triangular posterolateral spines. Telson dorsal surface with blunt submedian carinae and broad low median elevation terminating in a blunt tooth. 43 mm. Vic. (Bass Strait); 95 m depth.

Kasim Manning, 1995 Diagnosis. Rostral plate longer than broad; apical spine long, slender. Posterior margin of telson with 2 pairs of fixed primary teeth and ventral postanal spine; 4 or more intermediate denticles present; ventral surface with postanal spine. Kasim insuetus (Manning, 1970) (Fig. 169d–g). Eye with cornea bilobed, broader than stalk, mesial lobe conical. Rostral plate broad basally, apical spine extending beyond eyes. Antennal protopod with 1 ventral papilla. Mandibular palp present. Dactylus of raptorial claw with 6 teeth. Pereopods 1–3 basal segment with ventrally directed posterior spine; pereopod 1 also with inner spine. Abdominal somite 6 with 3 pairs of posteriorly directed spines. Telson dorsal surface with numerous longitudinal carinae; low median prominence terminating in acute spine; intermediate denticles slender; ventral surface with median postanal spine and 2 pairs of carinae, outer armed. 50 mm. WA (Great Australian Bight to Rottnest I.); 70–100 m depth.

Lysiosquillidae Giesbrecht, 1910 The family includes three genera, all of which occur in tropical Australian waters. One genus is represented in southern waters. Diagnosis. Eye with cornea bilobed, broader than stalk. Body depressed, loosely articulated. Pereopodal endopods slender. Uropodal endopod without strong fold on outer dorsal margin.

528 Stomatopoda – mantis shrimps

b c

e g

h j

Fig. 169. Tetrasquillidae. a–c, Heterosquilla pentadactyla. d–g, Kasim insuetus. h–k, Acaenosquilla brazieri. a, d, h, anterior region. b, e, i, raptorial claw. c, f, j, posterior region. g, k, ventral surface of telson.

529 Marine Decapod Crustacea of Southern Australia

Lysiosquilla Dana, 1852 Diagnosis. Antennular somite dorsal processes produced into anteriorly directed spines. Antennal scale slender, length about 3 or more times width. Telson of adults with fixed submedian teeth. Lysiosquilla colemani Ahyong, 2001 (Fig. 172a–c, Pl. 30c). Rostral plate cordiform, smooth dorsally, without median carina or sulcus; apex acute. Mandibular palp absent. Dactylus of raptorial claw with 11–14 teeth. Base of endopodal articulation with small ventral spine. 170 mm. Southern Qld, NSW (S to Wollongong area).

Nannosquillidae Manning, 1980 This family includes 14 genera, of which two are known from southern Australia. Diagnosis. Body depressed, loosely articulated. Pereopodal endopods ovate or subcircular. Uropodal endopod with strong, dorsal proximal fold. Telson submedian teeth with movable apices.

Austrosquilla Manning, 1966 The genus includes six species endemic to southern Australia. Diagnosis. Rostral plate with apical spine. Antennal protopod without dorsal tooth. Mandibular palp absent. Raptorial claw with distal spine on ischium. Thoracic somite 5–7 lateral processes single. Abdominal somite with posterolateral spines. Telson dorsal surface, without longitudinal carinae or spines. Telson posterior margin with 1 pair of movable submedian teeth and 2 pairs of fixed marginal teeth; 1 marginal and 4 intermediate denticles present. Uropodal protopod with inner branch longer than outer. Austrosquilla melanocauda (Kunze, 1981) (Fig. 170g–i). Eye with subglobular cornea. Rostral plate broad basally; lateral margins strongly convergent, sinuous; apex slender, acute. Antennal protopod with 1 ventral papilla. Dactylus of raptorial claw unknown. Pereopods 1–3 basal segment with slender, inner and outer spines. Abdominal somite 6 with 2 ventrolateral spines anterior to uropod articulation. Telson with low median prominence terminating in blunt projection; lateral proximal margin without distinct semicircular lobe; intermediate denticles short, conical, not on the same plane. 47 mm. NSW (known only from type locality, Botany Bay). Austrosquilla middletoni Ahyong, 2001 (Fig. 171a–c). Eye with subglobular cornea. Rostral plate triangular, broad basally; lateral margins strongly convergent, sinuous; apex slender, acute. Antennal protopod with 1 ventral papilla. Dactylus of raptorial claw with 10–14 teeth. Pereopods 1–3 basal segment unarmed on outer margin; pereopod 1 inner margin with broad triangular tooth. Abdominal somite 6 with 2 ventrolateral spines anterior to uropod articulation. Telson with low median prominence terminating in blunt projection; lateral proximal margin without distinct semicircular lobe; intermediate denticles short, slender. 42 mm. Known with certainty only from SA; intertidal and shallow subtidal; constructs burrows in fine sand, usually found in pairs. Austrosquilla osculans (Hale, 1924) (Fig. 170d–f). Eye with subglobular cornea. Rostral plate broadly quadrate with apical spine. Antennal protopod with 2 mesial and 1 ventral papilla. Dactylus of raptorial claw with 8–11 teeth. Pereopods 1–2 basal segment with inner and outer spine; pereopod 3 with outer spine only. Abdominal somite 6 with 2 ventrolateral spines anterior to uropod articulation. Telson with low median prominence terminating in acute spine; lateral proximal margin without distinct semicircular lobe; intermediate denticles elongate, triangular. 42 mm. Vic., SA; shallow, soft substrates.

530 Stomatopoda – mantis shrimps

h i

k m

Fig. 170. Nannosquillidae. a–c, Austrosquilla vercoi. d–f, Austrosquilla osculans. g–i, Austrosquilla melanocauda. j–m, Austrosquilla tsangi. a, d, g, j, anterior region. b, e, l, raptorial claw. c, f, i, m, posterior region. h, k, ventral surface of uropod.

531 Marine Decapod Crustacea of Southern Australia

a c

l j

Fig. 171. Nannosquillidae. a–c, Austrosquilla middletoni. d–f, Austrosquilla rachelae. g–i, Hadrosquilla edgari. j–l, Hadrosquilla perpasta. a, d, g, j, anterior region. b, e, h, k, raptorial claw. c, f, i, l, posterior region.

532 Stomatopoda – mantis shrimps

Fig. 172. Lysiosquillidae. a–c, Lysiosquilla colemani. Erythrosquillidae. d–f, Erythrosquilla hamano. a, d, anterior region. b, e, raptorial claw. c, f, posterior region.

533 Marine Decapod Crustacea of Southern Australia

Austrosquilla rachelae Ahyong, 2001 (Fig. 171d–f). Eye with faintly bilobed cornea. Rostral plate broad basally; lateral margins strongly convergent, sinuous; apex long, slender, acute. Antennal protopod with 1 ventral papilla. Dactylus of raptorial claw with 16 teeth. Pereopods 1–3 basal segment with slender, inner and outer spines. Abdominal somite 6 with 2 ventrolateral spines anterior to uropod articulation. Telson with low median prominence terminating in blunt projection; lateral proximal margin without distinct semicircular lobe; intermediate denticles short, slender. 57 mm. Vic., SA, WA (N to Cottesloe); intertidal and shallow subtidal; constructs burrows in fine sand. Austrosquilla tsangi Ahyong, 2001 (Fig. 170j–m). Eye with subglobular cornea. Rostral plate triangular, broad basally; lateral margins convergent, sinuous; apex acute. Antennal protopod with 1 ventral papilla. Dactylus of raptorial claw with 19–21 teeth. Pereopods 1–2 basal segment with inner and outer spine; pereopod 3 with outer spine only. Abdominal somite 6 with 1 ventrolateral spine anterior to uropod articulation. Telson with low median prominence terminating in blunt projection; lateral proximal margin with distinct semicircular lobe; intermediate denticles short, slender. 50 mm. NSW, intertidal, burrows in clean, fine sand, usually found in pairs. Austrosquilla vercoi (Hale, 1924) (Fig. 10a–c). Eye with subglobular cornea. Rostral plate subcordi- form; lateral margins subparallel for more than half median length, converging on short apical spine. Antennal protopod with 1 ventral papilla. Dactylus of raptorial claw with 8–12 teeth. Pereopods 1–2 basal segment with inner and outer spine; pereopod 3 with outer spine only; pereopod 1 inner margin with broad triangular tooth. Abdominal somite 6 with 1 ventrolateral spine anterior to uropod articulation. Telson with low median prominence terminating in blunt projection; lateral proximal margin without distinct semicircular lobe; intermediate denticles short, slender. 55 mm. Known with certainty only from Port Phillip Bay, Vic., Tas., and the type locality, Robe, SA.

Hadrosquilla Manning, 1966 The genus includes two endemic species. Diagnosis. Eye with subglobular cornea. Rostral plate quadrate, with triangular apical point. Antennal protopod unarmed dorsally, with 1 ventral and 2 mesial and ventral papillae. Thoracic somites 5–7 lateral margin flattened. Mandibular palp absent. Abdominal somite 6 posterolater- ally armed. Hadrosquilla edgari Ahyong, 2001 (Fig. 171g–i). Rostral plate quadrate, lateral margins straight, anterolateral corners rounded, apex obtuse, sharp. Dactylus of raptorial claw with 6 teeth. Pereopods 1–3 basal segment with triangular tooth on inner margin. Telson posterior margin with upper armature consisting of 1–2 slender, acute teeth above intermediate denticles; median projection blunt, rounded. Proximal segment of uropod with 4 movable spines. 36 mm. Tas.; intertidal to shallow subtidal; burrows in sand and under rocks. Hadrosquilla perpasta (Hale, 1924) (Fig. 171j–l, Pl. 30d). Rostral plate quadrate, lateral margins convex, anterolateral corners acute, apex obtuse, sharp. Dactylus of raptorial claw with 4–6 teeth. Pereopods 1–3 basal segment unarmed. Telson posterior margin with upper armature consisting of 4–5 stubby teeth above intermediate denticles; median projection flattened, emarginate. Proximal segment of uropod with 5–6 movable spines (rarely 4). 55 mm. Moreton Bay, Qld, NSW, Vic., SA; burrows in sandy, mud, among Zostera or under rocks, intertidal to 10 m depth.

534 Stomatopoda – mantis shrimps

Superfamily Parasquilloidea Manning, 1995 One family is included. Diagnosis. Eye with bilobed cornea. Maxillipeds 3–5 propodi longer than or as long as broad, without ventral ribbing. Raptorial claw with ischiomeral articulation terminal; dactylus with 3 teeth, not inflated basally. Telson with distinct median carina and 2 intermediate denticles. Uropodal protopod with three primary spines; articulation of exopod segments terminal.

Parasquillidae Manning, 1995 The family includes three genera, Faughnia, Parasquilla and Pseudosquillopsis. Faughnia is known from the Indo-West Pacific, Parasquilla from the Atlanto-Eastern Pacific and Pseudosquillopsis is known from both major regions. Only Pseudosquillopsis is represented in southern Australia. Keys to the Indo-West Pacific species of the Parasquillidae were given by Ahyong (2001). Diagnosis. As for superfamily.

Pseudosquillopsis Serène, 1962 The genus includes four species and one, P. dofleini, is known from Australia. Presently, the only known specimens P. dofleini from Australian are larvae or postlarvae. Therefore, a specimen from Japan is illustrated. Diagnosis. Rostral plate with long median spine. Carpus of raptorial claw with blunt dorsal tooth. Abdominal somites 1–5 without dorsal carinae. Telson without submedian denticles in adults. Pseudosquillopsis dofleini (Balss, 1910) (Fig. 168g–i). Rostral plate rounded laterally. Thoracic somites 6–7 lateral processes without posterolateral spine. Inner proximal margin of uropodal protopod with spinules on inner margin, increasing in size distally, with inner primary spine of uropodal protopod not markedly larger than distalmost spine. 85 mm. Japan, southern WA; shore to about 40 m depth.

Superfamily Squilloidea Latreille, 1802 One family is recognised. Diagnosis. Maxillipeds 3–5 propodi longer than or as long as broad, without ventral ribbing. Telson with distinct median carina and 4 or more intermediate denticles. Dorsum of thorax and abdomen longitudinally carinate. Uropodal protopod terminating in one or two primary spines.

Squillidae Latreille, 1802 Squillidae presently include 45 genera worldwide; 12 genera and 17 species are known from southern Australia. Most temperate water stomatopods are squilloids and are easily recognised by the dorsally carinate abdomen and distinct median carina of the telson. They actively forage at night and for this reason are the most frequently encountered, particularly in night-time demersal trawls. Most squilloids construct U-shaped burrows in soft level substrates where they remain during the day. Diagnosis. As for superfamily.

535 Marine Decapod Crustacea of Southern Australia

Key to southern Australian species of Squillidae 1. Propodus of raptorial claw pectinate, without row of erect spines. Posterolateral margin of carapace entire, not deeply excavate ...... 2 — Propodus of raptorial claw with row of erect spines. Posterolateral margin of carapace deeply excavate ...... 15 2. Lateral process of thoracic somite 6 bilobed ...... 3 — Lateral process of thoracic somite 6 single ...... 10 3. Lateral process of thoracic somite 5 a slender spine, directed laterally, with a ventral spine directed anterolaterally; spines not connected at base ...... Kempina mikado — Lateral process of thoracic somite 5 comprising two spines, connected basally ...... 4 4. Anterior margin of ophthalmic somite with median spinule. Carapace dorsal surface smooth; anterior width exceeding half median length ...... 5 — Anterior margin of ophthalmic somite without median spinule; truncate or emarginate. Carapace dorsal surface pitted; anterior width less than half median length ...... 6 5. Outer inferodistal angle of raptorial claw with stout tooth. Anterior margin of ophthalmic somite broadly rounded, usually with median spinule. Antennular peduncle uniformly maroon in life ...... Erugosquilla woodmasoni — Outer inferodistal angle of raptorial claw without tooth, at most obtusely angled. Anterior margin of ophthalmic somite trapezoid, with median spinule. Antennular peduncle banded blue and yellow/orange in life ...... Erugosquilla grahami 6. Merus of raptorial claw without spine on outer distal margin ...... Quollastria kapala — Merus of raptorial claw with spine on outer distal margin ...... 7 7. Median carina of carapace with distinct anterior bifurcation, uninterrupted basally ...... Oratosquilla oratoria — Median carina of carapace with or without anterior bifurcation; if bifurcation present, interrupted basally ...... 8 8. Telson with numerous longitudinal carinae on dorsal and ventral surface. Rostral plate with short median carina ...... Oratosquillina berentsae — Telson without numerous longitudinal carinae on dorsal and ventral surface. Rostral plate without short median carina ...... 9 9. Lobe between spines of uropodal protopod small and rounded, with convex or straight margin. Thoracic somite 6 lateral process with triangular anterior lobe ...... Oratosquillina interrupta — Lobe between spines of uropodal protopod small and rounded, with concave margin. Thoracic somite 6 lateral process with quadrate anterior lobe . . . . . Oratosquillina inornata 10. Thoracic somite 5 lateral process bilobed. Propodus of raptorial claw with distal tooth ...... Belosquilla laevis — Thoracic somite 5 lateral processes single. Propodus of raptorial claw without distal tooth ...... 11 11. Dorsal surface of telson with numerous longitudinal carinae either side of the median carina ...... 12 — Dorsal surface of telson without numerous longitudinal carinae either side of the median carina ...... 14 12. Carapace without anterolateral spines. Dactylus of raptorial claw with 4 teeth ...... Distosquilla miles

536 Stomatopoda – mantis shrimps

— Carapace with anterolateral spines. Dactylus of raptorial claw with more than 4 teeth . . . 13 13. Telson submedian teeth with fixed apices; ventral surface with numerous longitudinal carinae ...... Anchisquilla fasciata — Telson submedian teeth with movable apices; ventral surface without numerous longitudinal carinae ...... Levisquilla jurichi 14. Abdominal somites 1–5 with median and submedian carinae. Inner margin of uropodal protopod crenulate. Submedian teeth of telson with movable apices . . Anchisquilloides mcneilli — Abdominal somites 1–5 without median carina; abdominal somites 1–3 without submedian carinae. Inner margin of uropodal protopod lined with spines. Submedian teeth of telson with fixed apices ...... Lenisquilla lata 15. Rostral plate with evenly rounded apex. Intermediate carinae of thoracic somites with posterior spines ...... Harpiosquilla sinensis — Rostral plate with bluntly pointed or angular apex. Intermediate carinae of thoracic somites without posterior spines ...... 16 16. Carapace without median carina. Distal segment of uropodal exopod black ...... Harpiosquilla melanoura — Carapace with median carina. Distal segment of uropodal exopod black or brown on inner half only ...... Harpiosquilla harpax

Anchisquilla Manning, 1968 The genus includes three species of which one is known from southern Australia Diagnosis. Eye with bilobed cornea, broader than stalk. Rostral plate triangular; apex rounded. Carapace with anterolateral spines, median carina absent. Mandibular palp present. Dactylus of raptorial claw with 6 teeth; dorsal carina of carpus entire; merus without outer, inferodistal spine. Thoracic somite 5 lateral process a single, triangular spine, directed anterolaterally. Thoracic somites 6–7 lateral processes single, rounded. Abdominal somites 1–5 without submedian carinae. Telson dorsal surface with numerous carinae either side of median carina; submedian teeth with fixed apices. Uropodal protopod with spinous inner margin. Anchisquilla fasciata (De Haan, 1844) (Fig. 174a–e). Telson with accessory median carina entire or subdivided proximally; dorsolateral carina over entire surface; prelateral lobe with blunt apex; ventral surface with longitudinal carinae lateral to postanal carina. 100 mm. Western Indian Ocean to Japan, New Caledonia, NT, Qld, NSW, Vic., SA; rare off southern Australia, sand and mud on level substrates; subtidal to approximately 30 m depth.

Anchisquilloides Manning, 1977 The genus includes two Indo-West Pacific species, of which one is known from Australia. Diagnosis. Eye with bilobed cornea, broader than stalk. Rostral plate triangular; apex acute. Carapace with anterolateral spines and median carina, anterior bifurcation absent. Mandibular palp present. Raptorial claw carpus dorsal carina entire; merus without outer inferodistal spine; dactylus with 5–6 teeth. Abdomen with median, submedian, intermediate, lateral and marginal carinae. Thoracic somite 5–7 lateral processes single, that of thoracic somite 5 slender, recurved anteriorly, that of thoracic somite 6–7 broad, triangular, acute posteriorly. Telson dorsal surface without carinae on either side of median carina; submedian teeth with movable apices. Uropodal protopod with crenulate inner margin.

537 Marine Decapod Crustacea of Southern Australia

Anchisquilloides mcneilli (Stephenson, 1953) (Fig. 173e–h, Pl. 30e). Dorsal surface smooth. Carapace with anterolateral spines. Raptorial claw dactylus with 5–6 teeth. Abdominal somites 2 and 5 without dark median and submedian rectangular patches dorsally. Uropodal protopod with large rounded lobe on outer margin of inner spine, broader than adjacent spine, margin concave. 120 mm. Central Qld, NSW, Vic., Tas., SA, WA (N to Perth); level bottoms, 14–300 m depth.

Belosquilla Ahyong, 2001 The genus is monotypic and endemic to Australia. Diagnosis. Eye with bilobed cornea, broader than stalk. Carapace with anterolateral spines and distinct median carina; median carina with distinct anterior bifurcation; posterolateral margins entire, not excavate. Dactylus of raptorial claw with 6 teeth; dorsal carina of carpus entire; propodus with stout distal tooth, pectinate on opposable margin; merus without outer, inferodistal spine. Thoracic somite 5 lateral process bilobed. Thoracic somites 6–7 single. Abdomen with submedian, intermediate, lateral and marginal carinae. Abdominal somites 2 and 5 with dark median and submedian rectangular patches dorsally. Telson submedian teeth with fixed apices. Uropodal protopod inner spine outer margin with rounded lobe, margin straight or concave; inner margin of inner spine crenulate. Belosquilla laevis (Hess, 1865) (Fig. 177a–d, Pl. 30f). 120 mm. Southern Qld, NSW, Vic., SA, WA; estuarine and shallow coastal waters on sandy mud; intertidal to 40 m depth. This is the most common shallow water stomatopod in southern Australia.

Distosquilla Manning, 1977 This monotypic genus is endemic to southern Australia. Diagnosis. Dorsal surface smooth. Eye with bilobed cornea, broader than stalk, outer lobe longer than inner. Rostral plate ovoid, apex rounded; median carina absent. Carapace without anterolateral spines; median carina absent. Mandibular palp absent. Dactylus of raptorial claw with 4 teeth; carpus dorsal carina entire; merus without outer inferodistal spine. Thoracic somite 5–7 single, lateral processes rounded. Abdomen with numerous additional longitudinal carinae. Telson dorsal surface with numerous carinae either side of the median carina; submedian teeth with movable apices. Uropodal protopod with inner margin lined with spines; lobe on outer margin of inner spine rounded, margin concave or straight. Abdominal somites 2 and 5 without dark median and submedian rectangular patches dorsally. Distosquilla miles (Hess, 1865) (Fig. 173a–d). 151 mm. Vic., Tas.,WA (N to Perth); sand and mud; usually shallow subtidal but to 920–1040 m depth.

Erugosquilla Manning, 1995 The genus includes at least six Indo-West Pacific species. Diagnosis. Dorsal surface smooth. Eye with bilobed cornea, broader than stalk. Anterior margin of ophthalmic somite usually with apical spinule. Carapace anterior width exceeding half midlength. Carapace with anterolateral spines and median carina, anterior bifurcation obsolete. Mandibular palp present. Dactylus of raptorial claw with 6–7 teeth; carpus dorsal carina tuberculate; merus with or without outer inferodistal spine. Abdomen with submedian, intermediate, lateral and marginal carinae. Thoracic somite 5–7 lateral processes strongly bilobed. Telson submedian teeth with fixed apices. Uropodal protopod with crenulate inner margin crenulate.

538 Stomatopoda – mantis shrimps

Erugosquilla grahami Ahyong & Manning, 1998 (Fig. 176a–d, m, Pl. 31a). Rostral plate short, trapezoid; apex flattened, without median carina. Anterior margin of ophthalmic somite trapezoid with small median spinule. Raptorial claw dactylus with 6 teeth; merus without outer inferodistal spine. Telson dorsal surface with rows of shallow pits, without rows of tubercles flanking median carina. Uropodal protopod with small rounded lobe on outer margin of inner spine. Antennular peduncle banded blue and yellow in life. Uropods blue. Abdominal somites 2 and 5 with dark median and submedian rectangular patches dorsally. 150 mm. Taiwan, WA (Exmouth Gulf), NT, Qld, NSW (S to Port Jackson); sand and mud substrates, subtidal to 20 m depth. Erugosquilla woodmasoni (Kemp, 1911) (Fig. 176e–h, n). Rostral plate short, trapezoid; apex flattened, without median carina. Anterior margin of ophthalmic somite broadly rounded, usually with small median spinule. Raptorial claw dactylus with 6 teeth; merus with outer inferodistal spine. Telson dorsal surface with rows of shallow pits, without rows of tubercles flanking median carina. Uropodal protopod with small rounded lobe or spine on outer margin of inner spine. Antennular peduncle uniformly maroon in life. Uropods blue. Abdominal somites 2 and 5 without dark median and submedian rectangular patches dorsally. 140 mm. Western Indian Ocean to Japan, WA, NT, Qld, NSW (S to Port Jackson); sand and mud substrates; subtidal to 30 m depth.

f g

Fig. 173. Squillidae. a–d, Distosquilla miles. e–h, Anchisquilloides mcneilli. a, e, anterior region. b, f, thoracic somites 5–7, lateral region. d, h, raptorial claw. c, g, posterior region.

539 Marine Decapod Crustacea of Southern Australia

d a c

b f

k h j

l n

Fig. 174. Squillidae. a–e, Anchisquilla fasciata. f–j, Levisquilla jurichi. k–n, Lenisquilla lata. a, f, k, anterior region. b, g, l, thoracic somites 5–7, lateral region. d, i, n, raptorial claw. c, h, m, posterior region. e, j, ventral surface of telson.

540 Stomatopoda – mantis shrimps

d a c b

e g

k h

i j

Fig. 175. Squillidae. a–d, Harpiosquilla harpax. e–g, Harpiosquilla melanoura. h–k, Harpiosquilla sinensis. a, e, h, anterior region. b, i, thoracic somites 5–7, lateral region. d, g, k, raptorial claw. c, f, j, posterior region.

541 Marine Decapod Crustacea of Southern Australia

a m d c

e h n

j k

Fig. 176. Squillidae. a–d, m, Erugosquilla grahami. e–h, n, Erugosquilla woodmasoni. i–l, Oratosquilla oratoria. a, e, i, anterior region. b, f, j, thoracic somites 5–7, lateral region. d, h, l, raptorial claw. c, g, k, posterior region. m, n, anterior margin of ophthalmic somite.

542 Stomatopoda – mantis shrimps

Harpiosquilla Holthuis, 1964 Diagnosis. Raptorial claw propodus with erect spines on opposable margin. Eye with strongly bilobed cornea, much broader than stalk. Mandibular palp present. Carapace with anterolateral spines; median carina without anterior bifurcation. Carapace posterolateral margins deeply excavate. Thoracic somite 5 lateral processes single. Thoracic somites 6–7 single or faintly bilobed. Telson submedian teeth with fixed apices. Uropodal protopod with crenulate inner margin. Harpiosquilla harpax (De Haan, 1844) (Fig. 175a–d, Pl. 31b). Rostral plate trianguloid; apex elongate. Carapace with median carina. Dactylus of raptorial claw with 8 teeth. Thoracic somite 6–7 lateral processes single, triangular. Thoracic somite 5 lateral process rounded. Thoracic somites 6–8 intermediate carinae unarmed posteriorly. Telson dorsal surface with pair of maroon spots proximally. Uropod exopod distal segment dark on inner half. 250 mm. Western Indian Ocean to Japan, New Caledonia, WA, NT, Qld, NSW (S to Botany Bay); sand and mud; subtidal to 90 m depth. Harpiosquilla melanoura Manning, 1968 (Fig. 175e–g, Pl. 31c). Rostral plate trianguloid; apex elongate. Carapace without median carina. Dactylus of raptorial claw with 8 teeth. Thoracic somite 5 lateral process single, rounded. Thoracic somites 6–7 lateral processes faintly bilobed. Thoracic somites 6–8 intermediate carinae unarmed posteriorly. Abdominal somites 1–5 without submedian carinae. Telson dorsal surface with pair of red-maroon spots proximally; uropod exopod distal segment black. 160mm. Western Indian Ocean to Japan, NT, Qld, NSW (S to Botany Bay); sand and mud; subtidal to around 75 m depth. Harpiosquilla sinensis Liu & Wang, 1998 (Fig. 175h–k, Pl. 31d). Rostral plate semicircular; apex rounded, without anterior projection. Carapace with median carina. Dactylus of raptorial claw with 8 teeth. Thoracic somite 5 lateral process single, apex acute. Thoracic somites 6–7 lateral processes faintly triangular; thoracic somite 7 posterior lobe usually secondarily bifid. Thoracic somites 6–8 intermediate carinae armed posteriorly. Telson dorsal surface with pair of dark spots proximally; uropod exopod distal segment dark on inner half. 200 mm. Taiwan, southern South China Sea, Qld, NSW (S to Newcastle Bight); 63–212 m depth.

Kempina Manning, 1978 The genus includes two Indo-West Pacific species one from Australia. Diagnosis. Dorsal surface rugose, eroded and pitted. Eye with bilobed cornea, broader than stalk. Carapace with anterolateral spines and distinct median carina. Mandibular palp present. Dactylus of raptorial claw with 6 teeth; carpus dorsal carina entire; merus without outer inferodistal spine. Abdomen with submedian, intermediate, lateral and marginal carinae. Thoracic somite 5 lateral process a single spine, directed laterally, with ventral spine directed anterolaterally. Thoracic somites 6–7 strongly bilobed. Telson submedian teeth with fixed apices. Uropodal protopod with crenulate inner margin. Kempina mikado (Kemp & Chopra, 1921) (Fig. 177e–h, Pl. 32a). Rostral plate trapezoid; apex flattened to rounded; median carina present. Abdominal somites 2 and 5 with dark median and submedian rectangular patches dorsally. 182 mm. Western Indian Ocean to Japan, WA, NT, central Qld, NSW (S to Sydney). This species occurs in deep offshore waters and off NSW is frequently taken with Anchisquilloides mcneilli in depths between 70 and 150 m.

Lenisquilla Manning, 1977 The genus includes two species, of which one is known from Australia. Diagnosis. Eye with bilobed cornea, broader than stalk. Carapace with anterolateral spines; median carina absent. Mandibular palp present. Dactylus of raptorial claw with 5–6 teeth; carpus dorsal

543 Marine Decapod Crustacea of Southern Australia

carina entire; merus without outer inferodistal spine. Abdomen with intermediate, lateral and marginal carinae, with or without submedian carinae. Thoracic somite 5 lateral process a single spine, directed anterolaterally. Thoracic somites 6–7 lateral processes broadly rounded. Telson submedian teeth with fixed apices; dorsal surface without numerous dorsal carinae either side of the median carina. Uropodal protopod with spinous inner margin. Lenisquilla lata (Brooks, 1886) (Fig. 174k–n). Dorsal surface very minutely punctate. Rostral plate triangular; apex rounded. Raptorial claw dactylus with 5 or 6 teeth. Abdominal somites 1–3 without submedian carinae. Uropodal protopod inner spine outer margin with large rounded lobe, margin concave. Abdominal somites 2 and 5 without dark median and submedian rectangular patches dorsally. 80 mm. Red Sea to Japan, WA, NT, Qld, NSW (S to Newcastle).

Levisquilla Manning, 1977 The genus includes three Indo-West Pacific species of which one is known from southern Australia. Diagnosis. Eye with bilobed cornea, broader than stalk. Carapace with anterolateral spines; without median carina. Mandibular palp absent. Dactylus of raptorial claw with 6 teeth; carpus dorsal carina entire; merus without outer inferodistal spine. Abdomen with intermediate, lateral and marginal carinae. Abdominal somites 1–5 without submedian carinae. Thoracic somite 5 lateral process a single spine, directed anterolaterally. Thoracic somites 6–7 lateral processes broadly rounded. Telson submedian teeth of telson with movable apices. Uropodal protopod with spinous inner margin. Levisquilla jurichi Makarov, 1979 (Fig. 174f–j). Rostral plate triangular; apex rounded; median carina absent. Telson dorsal surface with accessory median and numerous dorsal carinae either side of the median carina. 50 mm. Eastern Indian Ocean, South China Sea, New Caledonia, WA, Qld, NSW (S to Port Jackson); soft level substrates to around 20 m depth.

Oratosquilla Manning, 1968 The genus includes four Indo-West Pacific species. Diagnosis. Eye with bilobed cornea, broader than stalk. Carapace with anterolateral spines and distinct median carina, anterior bifurcation uninterrupted basally. Dactylus of raptorial claw with 6 teeth; carpus dorsal carina tuberculate. Thoracic somites 5–7 lateral processes bilobed. Abdomen with submedian, intermediate, lateral and marginal carinae. Telson submedian teeth with fixed apices. Uropodal protopod with crenulate inner margin. Oratosquilla oratoria (De Haan, 1844) (Fig. 176i–l, Pl. 32b). Dorsal surface minutely pitted. Rostral plate trapezoid; apex flattened to rounded; median carina absent. Raptorial claw merus with outer inferodistal spine. Thoracic somites 5–7 with strongly bilobed lateral processes. Abdominal somite 4 submedian carinae unarmed. Uropodal protopod with small rounded lobe on outer margin of inner spine; exopod distal segment dull yellow with dark outer margin. Abdominal somites 2 and 5 without dark median and submedian rectangular patches dorsally. 185 mm. Japan, China, Vietnam, NSW (Sydney area), Vic. (Lakes Entrance); sand and mud substrates, subtidal to 20 m depth. Oratosquilla oratoria appears to have been introduced to eastern Australia (Ahyong, 2001).

544 Stomatopoda – mantis shrimps

a c

b e

f g

i l j

Fig. 177. Squillidae. a–d, Belosquilla laevis. e–h, Kempina mikado. i–l, Quollastria kapala. a, e, i, anterior region. b, f, j, thoracic somites 5–7, lateral region. d, h, l, raptorial claw. c, g, k, posterior region.

545 Marine Decapod Crustacea of Southern Australia

a c

f g

j k

Fig. 178. Squillidae. a–d, Oratosquillina berentsae. e–h, Oratosquillina inornata. i–l, Oratosquillina interrupta. a, e, i, anterior region. b, f, j, thoracic somites 5–7, lateral region. d, h, l, raptorial claw. c, g, k, posterior region.

546 Stomatopoda – mantis shrimps

Oratosquillina Manning, 1995 Oratosquillina includes 13 species, all from the Indo-West Pacific; two are known from southern Australia. Diagnosis. Eye with bilobed cornea, broader than stalk. Carapace with anterolateral spines and distinct median carina, anterior bifurcation interrupted basally. Merus of raptorial claw with outer inferodistal spine. Abdomen with submedian, intermediate, lateral and marginal carinae. Thoracic somite 5–7 lateral processes strongly bilobed. Telson submedian teeth with fixed apices. Uropodal protopod with crenulate inner margin. Oratosquillina berentsae Ahyong, 2001 (Fig. 178a–d, Pl. 32c). Dorsal surface smooth, without fine pitting. Rostral plate trapezoid; apex flattened; with median carina. Carapace median carina without anterior bifurcation. Mandibular palp absent. Dactylus of raptorial claw with 6 teeth; carpus dorsal carina entire. Thoracic somite 6 lateral process anterior lobe slender. Thoracic somite 7 lateral process anterior lobe low, angular. Telson dorsal and ventral surface with numerous carinae. Uropodal protopod inner spine outer margin with rounded lobe, margin concave. Abdominal somites 2 and 5 without dark, median and submedian rectangular patches dorsally. 100 mm. NSW (Evans Head to Port Jackson); level muddy sand substrates to 51 m depth. Oratosquillina inornata (Tate, 1883) (Fig. 178e–h). Dorsal surface evenly and minutely pitted. Rostral plate quadrate, elongate; apex flattened or rounded; median carina absent. Carapace median carina with distinct anterior bifurcation. Dactylus of raptorial claw with 6 teeth; carpus dorsal carina entire. Thoracic somite 6 lateral process anterior lobe quadrate. Thoracic somite 7 lateral process anterior lobe triangular, slender. Telson dorsal surface without numerous carinae. Uropodal protopod inner spine outer margin with rounded lobe, margin concave. Abdominal somites 2 and 5 without dark, median and submedian rectangular patches dorsally. 112 mm. Western Pacific to Eastern Indian Ocean, WA (north), NT, Qld (Gulf of Carpentaria), SA; burrows in mud; intertidal and shallow sublittoral. Oratosquillina interrupta (Kemp, 1911) (Fig. 178i–l). Dorsal surface minutely pitted. Rostral plate quadrate, short; apex flattened; median carina absent. Anterior bifurcation of median carina distinct. Dactylus of raptorial claw with 6 teeth; carpus dorsal carina tuberculate. Thoracic somite 6 lateral process anterior lobe triangular. Thoracic somite 7 lateral process anterior lobe triangular. Telson dorsal surface without numerous carinae. Uropodal protopod inner spine outer margin with small rounded lobe, margin convex or straight. Abdominal somites 2 and 5 without dark, median and submedian rectangular patches dorsally. Telson median carina with dark, maroon patch proximally. 160 mm. Western Indian Ocean to Taiwan, Vietnam, WA, NT, Qld, NSW (S to Sydney area); sand and mud; subtidal to 20 m depth.

Quollastria Ahyong, 2001 The genus includes nine Indo-West Pacific species. Diagnosis. Eye with bilobed cornea, broader than stalk. Carapace with anterolateral spines and distinct median carina, anterior bifurcation interrupted basally. Dactylus of raptorial claw with 5 or 6 teeth; carpus dorsal carina undivided; merus without outer inferodistal spine. Thoracic somites 5–7 lateral processes bilobed. Abdomen with submedian, intermediate, lateral and marginal carinae. Telson submedian teeth with fixed apices. Uropodal protopod inner spine inner margin crenulate. Quollastria kapala Ahyong, 2001 (Fig. 177i–l, Pl. 32d). Dorsal surface pitted. Rostral plate quadrate, elongate; apex flattened; median carina absent. Carapace median carina with distinct anterior bifurcation. Dactylus of raptorial claw with 6 teeth. Maxillipeds 1–4 with

547 Marine Decapod Crustacea of Southern Australia

epipod. Telson dorsal surface without numerous carinae; submedian denticles widely spaced, with gap between denticles exceeding half denticle width. Uropodal protopod with rounded lobe between terminal spines; distal segment of exopod dark on outer proximal margin and inner three-quarters of inner half. 110 mm. Southern Qld, NSW (S to Sydney); 131–411 m depth.

References Ahyong, S. 2001. Revision of the Australian stomatopod Crustacea. Records of the Australian Museum, Supplement 26: 1–326. Ahyong, S., & Harling, C. 2000. The phylogeny of the stomatopod Crustacea. Australian Journal of Zoology 48: 607–642. Caldwell, R.L. 1991. Stomatopods: the better to see you with my dear. Australian Natural History 23: 696–705. Caldwell, R.L., & Dingle, H. 1976. Stomatopods. Scientific American 234 (1): 80–89. Manning, R.B. 1966. Notes on some Australian and New Zealand stomatopod Crustacea, with an account of the species collected by the Fisheries Investigation Ship Endeavour. Records of the Australian Museum 27: 79–137. Manning, R.B. 1980. The superfamilies, families and genera of Recent stomatopod Crustacea, with diagnoses of six new families. Proceedings of the Biological Society of Washington 93: 362–372. Manning, R.B. 1991. Stomatopod Crustacea collected by the Galathea Expedition, 1950–1952, with a list of Stomatopoda known from depths below 400 meters. Smithsonian Contributions to Zoology 521: 1–18. Manning, R.B. 1995. Stomatopod Crustacea of Vietnam: the legacy of Raoul Serène. Crustacean Research, Special Number 4: 1–339. Marshall, N.J. 1988. A unique colour and polarization vision system in mantis shrimps. Nature 333: 557–560. Marshall, N.J., Land, M.F., King, C.A., & Cronin, T.W. 1991. Compound eye structure: the detection of polarized light. The compound eyes of mantis shrimps (Crustacea, Hoplocarida, Stomatopoda). Philosophical Transactions of the Royal Society of London B 334: 33–56. Stephenson, W., & McNeill, F. 1955. The Australian Stomatopoda (Crustacea) in the collections of the Australian Museum, with a check list and key to the known Australian species. Records of the Australian Museum 33: 239–265. Tirmizi, N.M., & Kazmi, Q.B. 1984. A handbook on the Pakistani mantis shrimp Oratosquilla. University of Karachi, Centre of Excellence in Marine Biology Publication 4: 1–101.

548 GLOSSARY

Crustaceans, especially eucarideans, are built on essentially the same body-plan, a head and thorax covered by a carapace plus an abdomen (Fig. 179). Homologies exist between the major taxa for all the limbs and for the regions into which the carapace is sculptured. Terminology is much the same for all taxa but there are minor differences according to body type. The glossary includes most of the specialised terms but hybrid words, like gastrofrontal, are not defined. Plurals and adjectival forms are given for some, as are alternatives used in other literature.

Orientation Anterior and posterior refer to the head and tail ends of a typical decapod, dorsal and ventral to the top and bottom surfaces. In some literature dorsal and ventral are also used to refer to one or other edge of a limb when the terms upper and lower or extensor and flexor are anatomically more correct. Extensor and flexor refer to the action of the internal muscles along each margin. Similarly, strictly speaking legs do not have a anterior or posterior suface or edge but the terms are applied to describe the position of spines or ridges. Medial refers to a structure in the midline of the animal and mesial to the face of a paired limb facing the midline. Lateral is the opposite of mesial or refers to a position away from the midline. Sublateral is used to indicate, for example, a spine between the medial and lateral spines. Proximal and distal describe the attached and remote ends of a limb. Adjectives describing orientation are frequently combined to indicate intermediate conditions, e.g. posterolateral, anteroventral etc. The body is divided into segments, here called somites, arranged in three sections (tagmata), cephalon (head), thorax and abdomen (Fig. 179). Each appendage or limb is also segmented, the segments called articles here. The word segment is used for both somite and article in some literature.

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Fig. 179. The names of the regions, 20 somites and paired appendages of Malacostraca. Alternative names for thoracic limbs are given for lobsters and shrimps (five pereopods), crabs (cheliped and four walking legs), and stomatopods (five maxillipeds and 3 pereopods). Pleopods 1 and 2 are called gonopods in males when modified for sperm transfer. The carapace covers the cephalon and thorax in decapods.

549 Marine Decapod Crustacea of Southern Australia

Prawns, lobsters and shrimps Identification of families, genera and species of prawns, lobsters and shrimps relies on differences in the sculpture and spination of the carapace and the rostrum. Basic nomenclature is the same for all groups (Fig. 180). The rostrum may carry dorsal and ventral teeth and may have an adrostral ridge running parallel along each side. The orbit (concavity on each side of the rostrum) is defined at its lower end by an antennal spine. in some carideans an orbital spine sits between the rostrum and antennal spine. The free side of the carapace, called the pterygostomian, or branchiostegite when it covers the gills, may have a pterygostomial spine at its anteroventral corner and a branchiostegal spine nearby but away from the margin. Running usually obliquely across the carapace is the cervical ridge and groove. The hepatic spine is situated at or near the lower edge of the cervical groove. The cervical groove may run completely across the dorsal part of the carapace or may be visible only laterally. Ridges may run posteriorly from the antennal, orbital and branchiostegal spines. Other spines and grooves are important in some groups but not others. Spines on the dorsal regions of the carapace are labelled according to the area where they attach. Males and females differ in the shape and number of the first two pairs of pleopods. This is most obvious in Dendrobranchiata where males possess a petasma, a complex formed from the paired endopods of the first pleopods. In some groups, males may lack or have reduced first pleopods and have an additional lobe, the appendix masculina, on pleopod 2. Secondary sexual characters of the chelipeds or other limbs differentiate the sexes of some groups.

r st e as bs pt bg

B AS

al 1 2

sc 3

a2 mp3 4

i p1 5 m

p2 p3 T u p p4

d p5

Fig. 180. Typical caridean shrimp (Palaemon serenus). a1, antennule; a2, antenna; AS, abdominal somites (1–6); as, antennal spine; b, basis; B, branchiostegite; bg, branchiostegal groove; bs, branchiostegal spine; c, carpus; d, dactylus; e, eyestalk; i, ischium; m, merus; mp3, maxilliped 3; p, propodus; p1–p5, pereopod 1–pereopod 5; pl1, pl2, pleopod 1, pleopod 2; pt, pterygostomial angle; R, rostrum; sc, scaphocerite; st, stylocerite; T, telson; u; uropod. Hepatic spine not shown in this species (see Crangonidae).

550 Glossary

Fig. 181. Diagramatic dorsal (a) and ventral (b) views of typical brachyuran crab (from Rathbun, M.J. 1918. The grapsoid crabs of America. Bulletin of the United States National Museum 97: 1–461, pls 1–161) show carapace regions and names of articles of chelipeds and walking legs.

551 Marine Decapod Crustacea of Southern Australia

Crabs Identification of crabs to family, genus and species relies on understanding crab anatomy and the special terms applied. The dorsal surface of the carapace is consistently divided into regions more or less obviously divided by grooves. The names of the regions, or areolae, reflects the internal organs – gastric for stomach, cardiac for heart, hepatic for ‘liver’ and intestinal for intestine. Prefixes like epi- and pro- refer to divisions of these (Fig. 181). Ornamentation of the carapace margin is also critical. The front is between the eyes and may be lobed; the eyestalks have a terminal cornea and are in orbits, or sockets which may be protected by spines; the anterolateral and posterolateral margins run from the orbit to the back of the carapace and are more or less distinct. The anterolateral margin is frequently dentate. The five pairs of pereopods include the chelipeds plus four pairs of walking legs, which are the terms used here. The buccal field or cavern is the ventral region between the eyes, often defined by ridges, and covered by the pair of third maxillipeds. In most crabs the antennules (first antennae) and antennae (second antennae) are small. The mobile articles of the antennules usually sit in a pair of slits (fossae) in the first article which is part of the underside. The antennae are larger and lie between the antennules and next to the orbit. The basal article of the antenna is usually fused to the carapace and can be overlooked. Its relationship to the orbit can be significant. Males and females are easily told. The narrow abdomen of the male may have fused segments; the shape of its two pairs of pleopods (rarely more), called gonopods 1 and 2, can be critical. Females have a broader and freer abdomen and five pairs of pleopods. The sexes may be further differentiated by secondary sexually characters of size, spination or development of chelipeds.

Glossary abdomen posterior part of body, of six segments or somites, sometimes including tailfan abyssopelagic pelagic at abyssal depths, greater than 2000 m depth acicle small scaphocerite or antennal scale adrostral ridge ridge parallel to and separated by narrow groove from rostrum (Penaeoidea) afferent branchial channels pair of channels between ridges, supplying water to branchial cavity and opening in front of chelipeds or alongside epistome (some Brachyura) antenna (pl. antennae) second cephalic appendage, usually with peduncle of five articles and multiarticulate flagellum (sometimes antenna 2); plural sometimes used to refer to both antennules and antennae antennal scale see scaphocerite antennal spine spine on anterior margin of carapace ventral or lateral to orbit (antennal angle if spine absent) antennule first cephalic appendage, usually with peduncle of three articles and two multiarticulate flagella (sometimes antenna 1) anterolateral margin margin of carapace between orbit and lateral angle or spine appendix interna (pl. appendices internae) small branch on inner margin of pleopod 2, triangular or rod-like, usually with clusters of hooks to attach to its pair appendix masculina (pl. appendices masculinae) small branch on inner margin of pleopod 2 of males, usually rod-like, proximal to appendix interna arthrobranch gill attaching to membrance between coxa and side of body, 1 or 2 pairs per somite and usually pointing upwards article any of the segments of a limb or appendage (see coxa, basis, ischium, merus, carpus, propodus, dactylus) basis (pl. bases; adj. basial or basal) second article of limb bathypelagic pelagic at bathyal depths, 200 to 2000 m depth

552 Glossary

benthic living on the sea floor; infaunal species burrow, epibenthic species walk over the surface (cf. pelagic) branchiocardiac (ridge or groove) line along dorsal limit of branchiostegite branchiostegite (adj. branchiostegal) expanded ventrolateral part of carapace covering gills branchia (pl. of branchos, rarely used; adj. branchial) gills, usually thoracic branchial cavity space between thorax and lateral flap of carapace (branchiostegite) enclosing branchia or gills branchial region paired regions of dorsal carapace between hepatic regions in front posterior margin behind, sometimes bearing branchial spine(s), defined laterally in crabs by posterolateral margin buccal cavern space, especially in crabs, enclosing the mouthparts and more or less covered by third maxillipeds, defined anteriorly by epistome and laterally by pterygostomial region, sometimes by ridges (sometimes buccal frame) carapace shield arising from posterior margin of head and comprising lateral extensions of thoracic somites covering all or most of cephalothorax carcinisation tendency for decapods to evolve towards a crab-like form cardiac a median region of the carapace, between the gastric and intestinal regions carina (adj. carinate) keel, acute ridge, crest carpus fifth article of limb (first article of palp in maxilliped 3) cephalon head, body segments bearing eyes, antennules, antennae, mandibles, maxillules and maxillae cephalothorax all fused somites of the cephalon (head) and thorax, covered in Decapoda by carapace cervical groove groove across the middle of the carapace between gastric and cardiac regions, curving forwards laterally, sometimes incomplete medially, sometimes accompanied by cervical ridge and ending laterally in cervical incision in some lobsters, sometimes with paired postcervical spines chela (adj. chelate) claw or pincer, derived from interacting fixed finger (extension from palm of propodus) and moveable finger (dactylus); palm refers chromatophore pigmented cell in exoskeleton cincinnuli minute hooked processes interlocking halves of petasma (Dendrobranchiata) cladistics (adj. cladistic) study of evolutionary branching patterns continental shelf sea floor between low water level and shelf break, usually about 200 m depth continental slope sea floor between edge of continental shelf, usually about 200 m depth, and top of continental rise, about 1000 m depth (often refers to slope and rise) continental rise sea floor between lower limit of continental slope and abyssal plain (about 1000 to 4000 m) cornea facetted and pigmented part of eyestalk coxa first article of limb, usually short dactylus seventh article of limb (last article interacting with fixed finger to form chela in chelate limb) (sometimes dactyl) dendrobranchiate gill gill with paired primary branches subdivided dorsal oval dorsal region of carapace defined by linea thalassinica laterally, cervical groove posteriorly and oblique semicircle anteriorly (Thalassinidea) efferent branchial channels pair of channels between ridges, venting branchial cavity and opening lateral to endostome (some Brachyura)

553 Marine Decapod Crustacea of Southern Australia

endite lobe, usually mesial, of proximal articles of limbs, especially mouthparts endopod inner of two branches of biramous limb, comprising ischium through to dactylus; in many the dominant or only branch (sometimes endopodite) endostome plate inside buccal cavern surrounding mouth epipod lateral branch of limb attached sometimes with gill (podobranch) to coxa, coxal endite (sometimes epipodite) epistome region of carapace between front of carapace, antennules and buccal region exopod outer of two branches of biramous limb, comprising one or several short articles arising from basis; absent from many limbs (sometimes exopodite) eyestalk articulating eye comprising peduncle and (usually) pigmented cornea fixed finger prolongation of propodus of chelate limb (sometimes pollex) flagellum (pl. flagella) multiarticulate appendage of antennule (2 present) and antenna (1 only); also appendage of expods of maxillipeds front anterior median region of dorsal carapace between orbits and gastric region; or in crabs the margin between the orbits gastric region median region of dorsal carapace between frontal region and cardiac region; divided from anterior in midline into mesogastric, metagastric and urogastric regions, and with protogastric and epigastric regions laterally hepatic region paired regions of dorsal carapace between orbits and branchial regions, defined laterally in crabs by anterolateral margin, sometimes bearing hepatic spine(s), and sometimes (prawns and lobsters) with a curved hepatic groove or ridge ventrally (see subhepatic) incisor terminal process of mandible, cutting blade, often toothed (more properly, incisor process) Indo-West Pacific biogeographic region extending from eastern coast of Africa, throughout the Indian Ocean, to the western coast of the Pacific Ocean and the Pacific islands, including the northern half of Australia intestinal region most posterior median region of the dorsal carapace, sometimes with intestinal spine(s) ischiomerus fused ischium and merus (maxilliped 3 of Gnathophyllidae) ischium (adj. ischial) third article of limb (first article of endopod) linea anomurica, linea thalassinica longitudinal lateral groove defining upper margin of branchiostegite in Anomura and Thalassinidea mandible (adj. mandibular) third limb of cephalon, comprising a thick body with mesial molar process, terminal incisor process and mandibular palp, usually of 3 articles maxilla third limb of cephalon, small, complex and membranous (sometimes maxilla 2) maxilliped one of three pairs of complex thoracic mouthparts; maxilliped 1 is membranous like the maxilla, maxilliped 2 a little more robust, and maxilliped 3 is much larger and may be more or less opercular over the other mouthparts in crabs or pereopod-like in shrimps and prawns maxillule second limb of cephalon, small and membranous (sometimes maxilla 1) megalopa terminal larval stage in Brachyura and Anomura merus (adj. meral) fourth article of limb mesopelagic midwater over continental slope, beyond 200 m depth midwater pelagic

554 Glossary

monophyletic describing a higher taxon comprising all the species descendant from a common ancestor mouthparts all cephalothoracic limbs involved with feeding: mandibles through to third maxillipeds ocular scale articulating plate attached mesially on eyestalks (some Anomura, Stomatopoda) ocular spineor projection spine or lobe on carapace above eyes (some Palinura or Caridea) orbitorostral groove paired grooves between median ridge extending to rostrum and ridges above eyes (Alpheidae) palm body of propodus of chelate limb (excluding fixed finger) paraphyletic describing a higher taxon comprising some species descendant from a common ancestor but excluding other descendants of the same ancestor peduncle proximal segments of biramous limb, especially of antennule, antenna, pleopods and uropod (sometimes protopod or sympod) pelagic free-swimming in water mass, not associated with sea floor (cf. benthic) pereopod non-maxilliped thoracic limbs; five pairs in Decapoda (thoracopods 4–8) and three pairs in Stomatopoda (thoracopods 6–8); includes one pair of chelipeds and four pairs of walking legs in Brachyura. Each pereopod comprises the seven articles coxa, basis and endopodal articles ischium to dactylus, plus sometimes an exopod (attached to the basis) petasma male genital structure comprising the enlarged and coupled endopods of pleopods 1 (Dendrobranchiata); usually composed of complex lobes and lobules photophore luminescent integumental organ, sometimes with lens phyllobranchiate gill gill with plate-like or leaf-like branches arranged in paird series along main axis phyllosoma pelagic larva of Palinura (Fig. 56), develops eventually into peurulus stage before settlement pleopod one of paired limbs on abdominal somites 1–5, usually biramous pleurobranch gill attached to body wall inside branchial cavity dorsal to coxae of pereopods, usually 1 pair per somite and pointing down between arthrobranchs pleuron (pl. pleura) lateral part of body wall, especially of abdomen, produced ventrally podobranch gill attached to coxa of pereopod or its epipod polyphyletic describing a higher taxon comprising species descendant from a common ancestor that belongs to another taxon post- prefix meaning posterior to, as in postrostral, postcervical etc. propodus sixth article of limb (palm of chela with fixed finger attached) prosartema process arising from mesial margin of first article of antennule (Dendrobranchiata) pterygostomial region anteroventral region of carapace (anterior part of branchiostegite of Caridea) or ventral region of carapace (of Brachyura); bearing pterygostomial spine at anteroventral angle (in Caridea and Dendrobranchiata) ramus (pl. rami) branch of limb; endopod is inner branch and exopod is outer branch rostrum (adj. rostral) prolongation of median part of anterior carapace between eyes scaphocerite lateral plate attached to distal margin of second article of antenna, equivalent to exopod (sometimes antennal scale or acicle when small) seamount submarine mountain, usually volcanic and steep-sided, reaching much shallower water than surrounding continental slope or abyssal depths seminal receptacles sacs for storage of spermatophores or sperm, usually on sternite of thoracic

555 Marine Decapod Crustacea of Southern Australia

somite 7 (sometimes spermathecae), associated with thelycum in Dendrobranchiata setal row row of regularly spaced plumose setae, attached in pits on carapace and some limbs (Thalassinidea) somite body segment, as in abdominal somites 1–6 spermatophore mass of spermatozoa deposited by gonopods of male on to thoracic sternum or into seminal receptacle of female statocyst cavity filled with fluid and particles acting as organ of orientation sternite ventral plate of integument (cf. tergite) sternum ventral surface of body, to which limbs are attached, or between limbs stridulating organ sound-producing organ formed by interacting areas of regularly ridged or beaded integument (e.g. paired chelipeds in Strigopagurus or cheliped and carapace in ocypodids) stylamblys specialised second endopodal article of pleopod 2 of female Palinura stylocerite scale arising from lateral base of first article of antennular peduncle sub- prefix meaning almost or close to subchelate claw-like, formed of transverse end of propodus and opposing dactylus (cf. chelate) tailfan telson plus pair of uropods telson plate attached medially to terminal segment of abdomen tergite dorsal plate of integument (cf. sternite) tergum arched dorsal part of abdominal somites thelycum female genitalia comprising modified posterior two or three sternal plates (Dendrobranchiata) thorax middle body region of eight somites bearing maxillipeds and pereopods tomentum mat of fine hairs trichobranchiate gill gill with fine branches on a central axis uropod paired limb of sixth abdominal somite, combining with telson to form tailfan walking leg one of last four pairs of thoracic limbs, or thoracic limbs exclusive of maxillipeds and cheliped(s) zoea (adj. zoeal) one of several larval stages, first stage hatching from the egg; transforming to final megalopa stage

556 CREDITS

Line drawings have been obtained from several sources as indicated below for each figure. Original illustrations by Kate Nolan (KN), Graham Milledge (GM), Gary Poore (GP), Shane Ahyong (SA) and Sherri Lehmann (SL) were prepared from specimens in Museum Victoria, Australian Museum, South Australian Museum and Western Australian Museum. Published photographs or line drawings have been adapted and copied by one or other of the above and are indicated by reference to the source paper cited in the relevant section. Direct scans of published drawings are annotated with *. Figures Fig. 3. a, KN. b–e, Dall, 2001. Fig. 4. a, Dall, 2001. Fig. 5. a, d, e, j, KN. b, f, Crosnier, 1991. c, Hanamura, 1989. g, h, Crosnier, 1985. i, Perèz-Farfante, 1980. k, Perèz-Farfante & Kensley, 1997. Fig. 6. a, b, e, Manning, 1988. c, Grey et al., 1983. d, f, h, KN. g, Perèz-Farfante, 1980. Fig. 7. a, b, Crosnier, 2003. c, e, h, Dall, 1999. d, KN. f, g, KN. i, j, Dakin & Colefax, 1940. Fig. 8. KN. Fig. 9. a, Colefax, 1940. b, Hale, 1941. c–i, Vereshchaka, 2000. Fig. 10. a, b, KN. c, Bruce, 1991. d, Bruce, 1973. d, Noël, 1985. e, Chace, 1985. f, g, j, k, Chace, 1986. h, i, Okuno, 1997. l, k, Chace, 1984. Fig. 11. Kensley et al., 1987. Fig. 12. a, b, d, e, KN. c, Crosnier & Forest, 1973. f–h, SL. i–m, Kensley et al., 1987. Fig. 13. KN. Fig. 14. a, Bate, 1888. b, f, g, Crosnier & Forest, 1973. c, e, Hanamura, 1989. d, Holthuis, 1993. h–n, KN. Fig. 15. KN. Fig. 16. a, f, *Bruce, 1990a. b, c, *Kensley, 1983. d, e, *Bruce, 1990b. Fig. 17. a, c–e, KN. b, Chace, 1986. Fig. 18. a, KN. b, Okuno, 1997. c, d, Hale, 1941. Fig. 19. Chace, 1983/KN. Fig. 20. KN. Fig. 21. a, h, e, Mannning, 1963/KN. b, SA. c, Bruce, 1988a. d, g, Bruce, 1973. f, Schmitt, 1933. Fig. 22. KN. Fig. 23. a, b, Holthuis, 1950. c, f–i, KN. d, de Man, 1908. e, Chace, 1972. j, k, Bray, 1976. Fig. 24. a, *Bruce, 1991a. b, *Bruce, 1977. c, *Bruce, 1980. Fig. 25. a, *Bruce & Cropp, 1984, b, *Bruce, 2003. c, Bruce, 2002. d, KN. e, Bruce, 1992a. Fig. 26. KN except e, j, k, Banner & Banner, 1982a. Fig. 27. KN except a, t, u, *Banner & Banner, 1982a; k–o, *Banner & Banner, 1973. Fig. 28. KN. Fig. 29. KN except c, d, g, h, b', c', *Banner & Banner, 1982a; z, a', *Banner & Banner, 1973. Fig. 30. KN. Fig. 31. a, Bruce, 1990. b, c, Taylor & Poore, 1998. Fig. 32. a–d, KN. e–i, Kensley et al., 1987. Fig. 33. a, b, KN. c, Crosnier & Forest, 1973. Fig. 34. KN. Fig. 35. a, g, i–k, KN. b, Hanamura & Evans, 1996. c, Crosnier & Forest, 1973. d, Kensley et al., 1987. e, f, h, Chan & Crosnier, 1991. Fig. 36. a, *Dardeau & Heard, 1983. b, *Hanamura et al., 1999. c, e–i, KN. d, *Duris 1992.

557 Marine Decapod Crustacea of Southern Australia

Fig. 37. a, KN. b, c, Kensley et al., 1987. Fig. 38. a, Borradaile, 1910. b, c, Baba, 1979. d, e, Holthuis, 1946. Fig. 39. GM. Fig. 40. a–c, e–g, i, Galil, 2000. d, h, Ahyong & Brown, 2002. Fig. 41. a, b, GM. c–i, Galil, 2000. Fig. 42. GM. Fig. 43. a, Chan & Yu, 1991. b, c, Watabe, & Lizuka, 1999. d, e, Macpherson, 1990. f, GM. g, Holthuis, 1946. Fig. 44. a, Sakai, 1988. b, Kensley, 1996. c, f, Poore, 1997. d, h, i, Poore, 1994. e, Poore & Griffin, 1979. g, Yaldwyn, 1972. j–l, GM. Fig. 45. a, b, e–h, Poore & Griffin, 1979. c, d, GM. Fig. 46. a, b, GM. c, d, Poore & Griffin, 1979. Fig. 47. GP. Fig. 48. GM. Fig. 49. GM. Fig. 50. a, b, e, f, i–l, KN. c, d, Poore & Griffin, 1979. g, h, Sakai, 2000. Fig. 51. GM. Fig. 52. GM. Fig. 53. a–d, g, GM. e, g, Poore & Griffin, 1979. Fig. 54. a, d–g, GM. b, c, Poore & Griffin, 1979. Fig. 55. a, b, e–h, GM. c, d, *Poore & Griffin, 1979. Fig. 56. Atkinson & Bousted, 1982. Fig. 57. Holthuis, 1991. Fig. 58. GM. Fig. 59. GM. Fig. 60. a, de Saint Laurent & Macpherson, 1990a. b, Baba, 2000. c, GM. d–f, Ahyong & Poore, 2004. Fig. 61. a, KN, b–h, Ahyong & Poore, 2004. Fig. 62. Ahyong & Poore, 2004. Fig. 63. a–c, Ahyong & Poore, 2004. d–h, GM. Fig. 64. Ahyong & Poore, 2004 except d, GM. Fig. 65. a, b, d, f, GM. c, e, g–l, Ahyong & Poore, 2004. Fig. 66. a, c, d, Ahyong & Poore, 2004. b, GM. Fig. 67. GM. Fig. 68. a, Sankolli, 1963. b, Grant & McCulloch, 1906. c, d, GM. Fig. 69. GM. Fig. 70. a, Efford & Haig, 1968. b, Davie, 2002. c, GM. Fig. 71. a–d, g–j, GM. e, f, k, Morgan & Forest, 1991a. Fig. 72. a, *Alcock, 1905. b, e, GM. c, McLaughlin & Holthuis, 2001. d, Morgan, 1989. d, McLaughlin & Holthuis, 2001. Fig. 73. GM except i, Morgan, 1987b. Fig. 74. GM. Fig. 75. GM. Fig. 76. a, Sakai, 1976. b–e, SL. Fig. 77. a, McLaughlin, 1994, b, McLaughlin, 2003b. c, Lemaitre & McLaughlin, 2003. d–q, GM. Fig. 78. a–c, GM. d–f, McLaughlin, 1997. Fig. 79. GM. Fig. 80. a, c, Lemaitre, 1996. b, d–f, McLaughlin, 2003. Fig. 81. a–d, Lemaitre, 1996. e, f, KN. g-i, l, Lemaitre, 1999. j, k, Lemaitre & McLaughlin, 1992. Fig. 82. a–c, d, f, g, Lemaitre, 1996. e, Lemaitre, 1999. Fig. 83. GM. Fig. 84. KN. Fig. 85. KN. Fig. 86. a–c, KN. d, GP. e, f, Lewinsohn, 1984.

558 Glossary

Fig. 87. a, Hale, 1927. b, d, GP. c, Griffin, 1972. e, Guinot & Tavares, 2003. Fig. 88. McLay, 1999. Fig. 89. a, KN. b, SL. c, Guinot & Richer de Forges, 1995. d, Yaldwyn & Dawson, 1976. e, Guinot, 1995. Fig. 90. a, GM. b, KN. Fig. 91. KN. Fig. 92. a, Tavares, 1991/Grant, 1905. b, c, *Tavares, 1991. Fig. 93. a, KN. b, Griffin & Brown, 1976. c, Ahyong & Brown, 2003. Fig. 94. a–b, GM. c, KN. d, Davie & Short, 1989. Fig. 95. GM. Fig. 96. GM. Fig. 97. GM except b, g, Galil, 1993. Fig. 98. a, b, GP. c, SL. d, George & Clark, 1976. e–i, GM. j, Miers, 1886. Fig. 99. a, GM. b–f, h–j, KN. g, GP. Fig. 100. a, b, d, GM. e, KN. c, *Bell, 1885. f, GP. Fig. 101. a, f, Chen, 1989. b, GM. c, d, GP. e, KN. Fig. 102. a, GM. b–f, Galil & Clark, 1984. Fig. 103. GP. Fig. 104. a, KN. b, Miers, 1879. Fig. 105. a, Richer de Forges, 1994. b, c, Griffin, 1966b. d, KN. e, Richer de Forges, 1993. Fig. 106. KN. Fig. 107. a, c, KN. b, *De Haan, 1839. Fig. 108. a–d, Griffin, 1970. e, f, Guinot & Richer de Forges, 1982. g, Griffin & Tranter, 1986. h–j, KN. Fig. 109. a, Loh & Ng, 1999. b, Richer de Forges, 1993. c–i, Guinot & Richer de Forges, 1985. j, Webber & Richer de Forges, 1995. Fig. 110. KN. Fig. 111. a, b, KN. c, GP. Fig. 112. KN. Fig. 113. a, d–f, h, KN, b, Richer de Forges, 1993. c, GP. g, Griffin, 1963. i, Griffin, 1970. Fig. 114. a, Griffin, 1970. b, c, f, KN. g, h, Richer de Forges, 1993. d, i, j, GP. e, Griffin & Tranter, 1986. Fig. 115. a–h, k–m, Griffin & Tranter, 1986. i, n–p, GP. j, KN. Fig. 116. a, GP. b–d, KN. Fig. 117. a, b, KN. c, d, Sakai, 1976. e–i, GP. Fig. 118. a, b, Griffin & Tranter, 1986. c, Guinot & Richer de Forges, 1985. d, e, Griffin, 1973. f, g, GP. Fig. 119. a, KN. b–e, Lucas, 1980. Fig. 120. a, Lucas, 1980. b, d–f, KN. c, Melrose, 1975. Fig. 121. a, d, KN. b, c, e, GP. d, Melrose, 1975. Fig. 122. b, e, GP. c, KN. Fig. 123. GP. Fig. 124. a, KN. b, Davie, 1991. Fig. 125. KN. Fig. 126. GP. Fig. 127. a, b, Miers, 1886. c, Stephenson, 1972. Fig. 128. KN. Fig. 129. Ng, 1998. Fig. 130. KN. Fig. 131. a, KN. b–e, Stephenson & Rees, 1968b. Fig. 132. a, SL. b, e, Sakai, 1976. c, Stephenson & Cook, 1973. d, f, Stephenson & Campbell, 1959. g, Lanchester, 1900. h, Stephenson, 1961. Fig. 133. a, Sakai, 1976. b, KN. d, Keenan et al., 1998. c, e, SL. f–h, j, k, Stephenson & Campbell, 1959. i, Stephenson, 1961. Fig. 134. KN. Fig. 135. a, Stephenson et al., 1957. b–d, g, Wee & Ng, 1995. e, f, KN. h, Barnard, 1950. Fig. 136. a, e–g,Wee & Ng, 1995. b, Miers, 1886. c, Montgomery, 1931. d, Stephenson, 1961.

559 Marine Decapod Crustacea of Southern Australia

Fig. 137. a, Stephenson et al., 1957. b–j, m, n, p, Wee & Ng, 1995. k, l, KN. o, Miers, 1886. Fig. 138. Wee & Ng, 1995. Fig. 139. a, KN. b, Miers, 1886. c, Davie, 2002. Fig. 140. a, Sakai, 1976. b, Türkay, 1983. c, d, GP. e, Davie, 1989. Fig. 141. a, Griffin & Campbell, 1969. b, d, Ng & Wang, 1994. c, Davie, 1989. e, f, GP. Fig. 142. a, KN. b, Manning & Holthuis, 1981. Fig. 143. a, b, KN. c, GP. Fig. 144. a, b, Serène, 1984. c, d, GP. e, Garth & Kim, 1983. f, Davie & Ng, 2000/Grant & McCulloch, 1907. Fig. 145. a, g, i, KN. b–f, Chia & Ng, 1998. h, GP. Fig. 146. a, e, j, k, p, n, SA. b–d, f–i, l, m, o, GP. Fig. 147. a, KN. b, d, e, g, GP. c, h, j, McNiell, 1929, f, i, Griffin & Campbell, 1969. k–p, *Griffin & Yaldwyn, 1971. Fig. 148. a, b, Guinot, 1976. c, d, GM. e–g, GP.h–k, Serène, 1984. l, Morgan, 1990. m, n, Guinot, 1958. Fig. 149. a, f, g, i, KN. b, h, GP. c–e, Serène, 1984. Fig. 150. a, Ward, 1933. b, g, Serène, 1984. c, e, GM. d, GP. f, Guinot, 1967. h, KN. Fig. 151. a–d, f, g, i, j, GP. e, Sakai, 1976. h, Serène, 1984. Fig. 152. a–d, GP. e, f, Guinot, 1964. Fig. 153. Kropp, 1988. Fig. 154. a, KN. b, f, Geiger & Martin, 1999. c, e, Pregenzer, 1988. g, Pregenzer, 1979. Fig. 155. a–c, KN. d, SL. Fig. 156. a–c, KN. d, e, GP. f, g, Barnes, 1967. Fig. 157. GP. Fig. 158. a, Castro, 2000. b, KN. Fig. 159. GM. Fig. 160. GM except d, SL. Fig. 161. a, GP. b–h, GM. Fig. 162. GM except d, SL. Fig. 163. GM except b, Campbell, 1967. Fig. 164. a–d, GM. e, Türkay, 1978. Figs 165–178. SA. Fig. 180. KN. Fig. 181. *Rathbun, 1918.

Plates Photographers who allowed use of their 35 mm slides are credited for plates and images as follows: Shane Ahyong (photographer and copyright): 31a, 32b. Bill Boyle (photographer and copyright): 8c, 18b, 20d, e, 22g. Neville Coleman (photographer and copyright): 30c, f, 31b, 32d. David Evans (photographer, copyright CSIRO Marine Laboratories): e–g. Karen Gowlett-Holmes (photographer and copyright): 7a, d, e, 8b, 9a, d, g, 10d, e, g–i, 11a, h, 12g, 13e–g, 14a, c, 15a–d, f, g, 16a–d, g, h, 17a–e, h, 18a, c, e, g, 19e, f, 20a, 21a, e, 22a, b, f, 23c–e, 24a–c, e–g, 25d, f, 26b, c, f–h, 28a, c, e, 29a. Rudie H. Kuiter (photographer and copyright): 7f, 8a, 10c, f, 11b–d, 12c, 13b, c, 15e, 17g, 18d, f, 22c, 26a, e, 27a, b, 29g. Michael Marmach (photographer, copyright Museum Victoria): 1c, 7c, 8d, e, 9b, c, e, f, h, 10a, b, 12a, b, d–f, 13d, 14b, d, 16e, f, 17f, 19a–d, 20b, c, 21b–d, f–g, 22d, e, 23a, f–i, 24d, 25a–c, e, 26d, 28b, d, f, g, 29b–f, 30d. Mark Norman (photographer and copyright): 1e, 7b, 23b. Gary Poore (photographer, copyright Museum Victoria): 1b, d. Roger Springthorpe (photographer): 30a (copyright Australian Museum); 30b, e, 31 c, d, 32a, c (copyright Shane Ahyong).

560 INDEX

Note that page numbers in bold refer to an illustration abbreviata, Paranursia 332, 341, 342 Amphionidacea 14, 16, 22 abrolhensis, Calcinus 254 Amphipoda 22 abrolhensis, Dromidiopsis 305 amplectipes, Leontocaris 118, 122, 123 abrolhensis, Elamena 391, 392, 393 amsa, Pagurixus 273, 275 abrolhensis, Forestia 462, 463 Anacinetops 367 Acaenosquilla 528 Anaspidacea 14, 22 Acanthephyra 63, 64, 66 anatum, Leucosia 337, 338 acantholepis, Micropagurus 217, 273, 274–75, Pl. 16 Anchisquilla 537 acanthura, Nephropsis 163, 165, 166 Anchisquilloides 537–38 acer, Pilumnus 454, 455, Pl. 25 Anchistioididae 81 acerba, Hypothalassia 442, 443, 444 Anchistus 92 Acetes 46, 47 Ancylocheles 244 Achaeopsis 360 andamanensis, Stereomastis 156 Achaeus 355, 357–58, 362 andamanicus, Metanephrops 164, 165 Achelata 16, 22, 159, 197–213 andreossyi, Cymo 464, 466, 468 aciculus, Paguristes 260, 261 angelikae, Neocallichirus 181, 183, 184 Acrothoracica 21 angulatus, Puerulus 201, 204 Actaea 462, 463, 472 angusta, Tiarinia 350, 383, 384 Actaeinae 462–64 Anomura 9, 16, 17, 22, 159, 169, 215–87, 290 Actaeodes 463 Anostraca 21 Actumnus 449, 452 Antarctus 207, 209 aculeata, Galathea 232 antipodarum, Arctides 208, 209 aculeatus, Polycheles 152, 154, 154 antipodarum, Bresilia 71, 71,72 acutifrons, Acanthephyra 63, 64, 66 antipodes, Paramunida 238, 239 acutifrons, Pasiphaea 61 appendiculis, Dorphinaxius 175 Acutigebia 190 arafurensis, Metanephrops 162, 163 adamsi, Zebrida 451 aranea, Oncinopus 359, 363, 364 admete, Thalamita 426, 428, 430, 431 Arcania 333, 339–41 Aegaeon 137, 140 Archaeobrachyura 290, 298, 316–23 Aeglidae 215, 219 arcticus, Sergestes 48 aequimanus, Eucalliax 170, 180, 183, 184 Arctides 207, 209 aesopius, Periclimenes 86, 93, 95, Pl. 8 arenosus, Biffarius 181, 182, Pl. 12 affinis, Geryon 406 areolata, Trapezia 481 affinis, Lucifer 45 areolatus, Pilodius 464, 465, 466 affinis, Palaemon 89 Arete 99, 111 affinis, Paramicippe 383 aries, Naxia 296, 349, 375, 377, 378, Pl. 21 Agononida 9, 229–31 Aristaeomorpha 25 Alainopasiphaea 61 Aristaeopsis 25 Albunea 247 Aristeidae 24–27, 26 Albuneidae 218, 247–48, 248 Aristeus 25, 26 alcocki, Nanocassiope 472 armata, Hypothalassia 442, 444 alcocki, Platymaia 364 armatus, Sergestes 48, 49 alcocki, Plesionika 130, 132, 133 arrosor, Dardanus 255, 256, Pl. 15 Allogalathea 229, 231 Ascidiophilus 298 Alope 119 Ascothoracica 21 Alpheidae 9, 56, 57, 58, 98–117, 103, 107–109, Pl. 8, Ashtoret 345 Pl. 9 aspera, Schizophrys 377, 380 Alpheoidea 56, 98–127 asprosoma, Munida 232, 233, 233 Alpheopsis 99 assimilis, Glyphocrangon 142 Alpheus 99–106, 110, 113 Astacida 159 alticrenatus, Ibacus 198, 208, 210, 211 Astacidae 159 Altosquilla 520 Astacidea 16, 17, 22, 159–67 Alvinocaris 70 Astacoidea 159 Amarinus 390, 391, 393 Astacura 159 Ambidexter 128 astacus, Astacus 159 americanum, Gnathophyllum 81, 82, 83, Pl. 7 astrinx, Alpheus 100, 102, 103, 109, Pl. 8 americanus, Homarus 162 Atelecyclidae 291, 292, 400–01, 401, 469, Pl. 22

561 Marine Decapod Crustacea of Southern Australia

Atergatis 477 Axiidae 9, 161, 170, 171, 172, 173–76, 174, 175, Pl. 11 Athanas 99, 111–12 Axioidea 170, 173–78 Athanopsis 99, 112–13 Axiopsis 173–74 atlanticus, Discias 72 Axius 176 atlanticus, Sergestes 48 babai, Uroptychus 222, 223, 225, 227 atrinubes, Palaemonetes 86, 88,91 baccatus, Polycheles 152, 154, 155 Atyidae 56, 58, 69–70, 69 bakeri, Synalpheus 115 Atyoidea 56 balssi, Pasiphaea 61 audouinii, Cyclograpsus 501, 503, 503, 504, Pl. 28 balssi, Rhynchocinetes 76 auriculatus, Notostomus 63, 65,67 Banareia 476 aurita, Naxia 296, 349, 371, 375, 377, Pl. 21 barbicornis, Paramithrax 349, 373, 376, 377 Austinogebia 190 Barbouriidae 98 australe, Macrobrachium 89 barnardi, Pasiphaea 59, 60,61 australiensis, Alpheus 101, 102, 103, 109 bathamae, Trizocheles spinosus 287 australiensis, Axiopsis 173, 174, 174, 175 Bathyarctus 207, 209 australiensis, Dromidiopsis 300, 302, 303, 305, Pl. 17 Bathynellacea 14, 22 australiensis, Eplumula 316, Pl. 18 Bathypaguropsis 271, 272 australiensis, Galathea 220, 230, 231, 232, Pl. 13 Bathypalaemonellidae 80 australiensis, Haliporoides sibogae 44 Bathysquilla 520 australiensis, Hemisquilla 522, 525, 526, Pl. 30 Bathysquillidae 520, 521, Pl. 30 australiensis, Hippolyte 119, 120, 121, Pl. 10 Bathysquilloidea 517, 518, 520 australiensis, Latreillia 316 bedfordi, Halicarcinus 391, 394, 396 australiensis, Leucosia 337 Bellidilia 333, 340, 342 australiensis, Lybia 470, 473 Belliidae 400 australiensis, Metanephrops 162, 163, 164, Pl. 11 Belosquilla 538 australiensis, Miersiograpsus 513, 514 bennettae, Metapenaeus 33, 36, 36,41 australiensis, Ovalipes 296, 407, 414, 415, 416, Pl. 23 Benthesicymidae 9, 24, 27–31, 29 australiensis, Paradorippe 324, 325 Benthesicymus 27, 30 australiensis, Parapenaeus 33, 34,37 berentsae, Oratosquillina 536, 546, 547, Pl. 32 australiensis, Paratya 58, 69–70, 69 berentsae, Pasiphaea 61 australiensis, Petrocheles 218, 242, 244, 245 Betaeus 99, 113 australiensis, Phalangipus 349, 384, 386, 387 bicavernosa, Haledromia 298, 300, 306, Pl. 17 australiensis, Processa 127, 128 bicolor, Chaceon 406, 406 australiensis, Sicyonia 41, 42 bicornis, Gomeza 405, 405 australiensis, Trypaea 180, 182, 183, 185, Pl. 12 bicuspidatus, Scyllarus 210 australiensis, Upogebia 190, 191, 192, 193 bidens, Alpheus 99, 101, 102, 103, Pl. 9 australis, Acetes 47 bidentata, Trapezia 480 australis, Alainopasiphaea 61 bidentatus, Lachnopodus 471 australis, Alope 119 Biffarius 181 australis, Athanopsis 107, 113, Pl. 9 bifurca, Plesionika 130, 132, 133 australis, Austrodromidia 298, 299, 300, 302, 303, bifurcata, Huenia 351, 352, 354 303, 304 bimaculata, Charybdis 424, 426, 430 australis, Betaeus 107, 113 birsteini, Paralomis 266, 269, 269 australis, Dardanus 255, 257 bispinoculata, Munidopsis 235, 236, 237 australis, Dromidia 303 bispinosa, Litocheira 297, 433, 434, 435, 437, 437, Pl. 24 australis, Eumunida 221, 222 bispinosus, Panulirus longipes 202 australis, Hippa 248, 249 bituberculatus, Synalpheus 114, 115 australis, Huenia 349, 351, 352, 353, Pl. 20 Blepharipodidae 247 australis, Palaemonetes 85, 88,91 boasi, Paromolopsis 310, 312, 314 australis, Paracrangon 136, 138, 139, Pl. 10 bonaspei, Willemoesia 157 australis, Pasiphaea 59, 60,61 boschmai, Metanephrops 163, 164, 165, Pl. 11 australis, Pilumnus 454, 455 Bountiana 446 australis, Rhynchocinetes 76, 77, 77, Pl. 7, Pl. 13 bouvieri, Gennadas 28, 29,30 australis, Trichia dromiaeformis 476 bouvieri, Parapagurus 280, 282, 283, 284, Pl. 17 australis, Uroptychus 222, 223, 224, 227 bouvieri, Thalamita 426, 428 australis, Zalasius 475, 476 bowerbankii, Upogebia 190, 191, 192–93, 193 Australoplax 493 Brachynotus 506, 510 Australoremus 274 Brachypoda 21 australosulcatus, Alpheus 101, 102, 103, 109 Brachyura 9, 16, 17, 22, 159, 218, 289–515 Austrodromidia 9, 300, 303–04 Branchiopoda 13, 21 Austrolepidopa 247–48 Branchiura 21 Austropenaeus 27 brazieri, Acaenosquilla 527, 528, 529 Austrosquilla 530, 534 Bresilia 71, 72 Axianassidae 169, 187 Bresiliidae 56, 70–72, 71

562 Index

Bresilioidea 56, 70–73 chiragra, Gonodactylus 522, 523, 524 brevicarinata, Zebrida 450, 451, Pl. 24 Chiromanthes 509 brevirostris, Achaeus 357, 358, 361 Chirostylidae 9, 215, 219, 220–28, 222, 224, 227 brevirostris, Paguristes 260, 261, 262, Pl. 15 Chirostylus 220 brocki, Paraxiopsis 170, 173, 174, 175, 176 Chlorinoides 378 brodiei, Neolithodes 267, 268 Chlorodiella 465 brownae, Discias 71, 71,72 Chlorodiinae 461, 465–66 brucei, Ibacus 210, 211 Chlorotocella 129, 130–31 bulga, Leontocaris 118, 122, 123 Chlorotocus 132 bullatus, Nectocarcinus 411, 412 Choniognathus 367 bunburius, Alpheus 100, 101, 102, 103, 104, 109 chydaea, Munida 232, 233, 234 burkenroadi, Sympagurus 284–85, 285 cidaris, Oncopagurus 281, 282, 283 Bythiopagurus 271, 272, 277 ciliata, Pseudosquilla 522, 525, 526 Calappa 326, 327, 330 cinctimana, Liomera 467, 469 Calappidae 291, 292, 326–31, 328, 329, 345, 407, Pl. 18 cinctipes, Tetralia 479, 480, 480 Calappinae 326 Cirripedia 21 calcar, Uroptychus 222, 223, 225 Cladocera 21 Calcinus 218, 252, 253–54 clavicarpus, Gennadas 31 calculosa, Actaea 462, 463, 464, Pl. 26 Clibanarius 218, 252, 254, 256 caledonica, Austrolepidopa 248 Coenobitidae 250, 251 Callianassa 181 Coenobitoidea 215, 216, 250 Callianassidae 169, 170, 171, 172, 180–86, 182, 183, colemani, Lysiosquilla 527, 530, 533, Pl. 30 Pl. 12 collaroy, Corallianassa 180, 183, 184 Callianassoidea 170, 180–95 collini, Ctenocheles 187, 187 Calliax 184 collumianus, Alpheus 99, 100, 104 Calocarididae 169, 173 comatularum, Synalpheus 107, 109, 114, 115, 116 Calvactaea 476 compressus, Latreutes 119, 120, 121, 124, Pl. 10 Cambaridae 159 contrarius, Rhinolambrus 398, 399, 399 Campylonotidae 56, 80–81, 80 convexa, Ocypode 492, 494, 496 Campylonotoidea 56, 80–81 Copepoda 21 Campylonotus 80 Corallianassa 184 camtschatica, Paralithodes 266 coralliodytes, Cryptochirus 483, 483 Cancellus 252, 254 cordiformis, Heloecius 297, 490, 491, 493, Pl. 28 Cancer 5, 401 cordimanus, Ocypode 492, 496 Cancridae 291, 292, 401–04, 403, Pl. 22 cornuta, Arcania 339, 341 Cancrinae 401 cornutus, Ixoides 339 capensis, Gennadas 28, 29,30 corrugatus, Liocarcinus 413–14, 415, Pl. 22 Caphyrinae 408–09, 413 corrugatus, Macropipus 413 caradina, Hippolyte 119, 124 Corystidae 291, 292, 404–06, 405 carcharias, Actaea 462, 463 Crangon 99 Carcininae 408, 409–12 Crangonidae 9, 54, 55, 58, 136–41, 138, Pl. 10 Carcinoplax 434, 436, 438 Crangonoidea 55, 136–43 Carcinus 410 crassa, Helice 504 cardus, Uroptychus 223, 224, 225 crassimanus, Dardanus 256, 257, 258 Caridea 9, 17, 22, 53–143, 54, 57, 145 crassipes, Ebalia 334, 335 carinata, Megametope 470, 472, 473, Pl. 26 crassipes, Herbstia 384 carinidactylus, Periclimenes 86, 93,95 crassipes, Macrophthalmus 491, 492, 495 carnifex, Cardisoma 289 crassipes, Uca 497 catharus, Ovalipes 414, 416, 417 crassispinosa, Bathysquilla 520, 521 caystrus, Pseudozius 434, 437, 438 crassissima, Metapenaeopsis 33, 34,35 celebensis, Leandrites 87 cremnus, Alpheus strenuus 110 Cephalocarida 13, 21 Crenarctus 207, 209–10 ceramicus, Biffarius 181, 182 crenata, Thalamita 423, 428, 430, 431 cerasma, Cymo 464, 466, 468 crenatus, Crenarctus 208, 210, 210, Pl. 13 Ceratocarcinus 448, 449 cristata, Calappa 330 ceratophthalma, Ocypode 492, 494, 496, Pl. 28 cristatus, Haliporoides 42,43 Ceratoplax 449, 457–58 crosnieri, Notostomus 67 chabrus, Plagusia 512, 513–14, 514, Pl. 29 Crossonotoninae 498 chacei, Ibacus 210, 211 Cryptochiridae 291, 293, 482–83, 483 chacei, Merhippolyte 118, 123, 124 Cryptochirus 483 Chaceon 406 Cryptocneminae 331, 332, 333 Chaenostoma 495 Cryptocnemus 333 challengeri, Metanephrops 162, 163, 164 Cryptodromia 304 Charybdis 423, 424–25 Cryptopodia 398

563 Marine Decapod Crustacea of Southern Australia

Ctenocheles 186–87 Dittosa 340 Ctenochelidae 170, 171, 186–87, 187 dochmiodon, Thaumastocheles 160 Cumacea 14, 22 dofleini, Pseudosquillopsis 525, 535 cumatodes, Paraxanthodes 473, 474 dolichophallus, Xenophthalmodes 456, 458 curtirostris, Acanthephyra 63, 64, 66 dolospina, Palaemon 85, 87, 88, 89–90, Pl. 8 curtispina, Micippa 382, 383 Domecia 481, 482 curtispina, Mursia 326, 329, 330, Pl. 18 Domeciidae 291, 294, 460, 477, 480, 481–82 curvirostris, Achaeus 349, 356, 357, 358 Domeciinae 477 curvirostris, Trachypenaeus 33, 34,38 Dorhynchus 357, 360 custos, Anchistus 86, 92 Dorippidae 291, 318, 324–325, 325, 326, 347 custos, Diogenes 258, 259 Dorippinae 324 cyaneus, Planes 508 Dorphinaxius 175 Cyclodius 465, 466 dorsalis, Arete 107, 109, 111 Cyclodorippidae 290, 291, 316–17, 317, 318, 347 Dotillinae 490, 492–93 Cyclograpsinae 501, 503–05 dromana, Upogebia 189, 192, 193, 194 Cyclograpsus 501, 503–04 Dromia 304, 305 cygnus, Panulirus 199, 201, 202, 203, 204 Dromiacea 290, 298–311 Cymo 466, 468 dromiaeformis, Trichia 476 cymodoce, Trapezia 479, 480, 480 Dromidia 303 Cymoinae 461, 466, 468 Dromidiopsis 304–05 Cymonomidae 290, 291, 316, 318–20, 319, 347 Dromiidae 9, 290, 292, 298–308, 299, 301, 302, 303, Cymonomoides 318 Pl. 17, Pl. 18 Cymonomus 318 Dromiinae 298 Cyrtomaia 355, 358, 360 dubius, Diogenes 259 Dagnaudus 312 Dumea 355, 360 dalli, Metapenaeus 33, 36–37, 41 Dynomene 308 dama, Schizophrys 377, 380 Dynomenidae 218, 290, 292, 308–09, 309 danae, Macrobrachium 89 Ebalia 9, 333, 335–36, 340 danae, Thalamita 427, 428, 429, 430 Ebaliinae 331, 332, 335–36 dapsiles, Calcinus 253, 255 eburnea, Randallia 333, 342, 343 Dardanus 218, 252, 256–58 echinus, Synalpheus 114, 115 darwinii, Upogebia 190, 191, 192, 193–94, 193 edgari, Hadrosquilla 527, 532, 534 dasypus, Munidopsis 237 eduardoi, Penaeopsis 33, 34, 36,38 debilis, Palaemon 86, 88,89 edwardsiana, Aristaeopsis 25, 26 debilis, Systellaspis 64, 66,68 edwardsianus, Plesiopenaeus 25 Decapoda 2, 14, 16–17, 22 edwardsii, Alpheus 102, 103, 104, 109 deflexifrons, Microhalimus 349, 372, 373, 375 edwardsii, Jasus 200, 204, Pl. 2 deformis, Dardanus 257 edwardsii, Munidopsis 235, 236, 237 delli, Cymonomoides 318, 319, 324 edwardsii, Plesionika 130, 132, 133–34 delli, Ogyrides 58, 127, 127 Elamena 393 demani, Petrarctus 206 Elamenopsis 395 Dendrobranchiata 9, 16, 17, 22, 23–51 elatus, Hyastenus 350, 385, 386 dentifrons, Ebalia 334, 335, Pl. 19 electra, Etisus 467, 468 deplanatus, Ozius 442, 443, 444, 446 elegans, Allogalathea 220, 230, 231, Pl. 13 depressa, Calappa 327, 328 elegans, Lyramaia 385, 386, 386 depressa, Harpiliopsis 92 elegans, Notostomus 63, 66,67 depressa, Petalomera 306 elegans, Xanthias 471, 474, 475 depressus, Leptomithrax 349, 369, 374 elegantissima, Raymunida 239 deprofundis, Propagurus 277–78, 277 elongatum, Tozeuma 119, 124, 125, Pl. 10 deuteropus, Alpheus 100, 104 elongatus, Petrolisthes 4, 218, 242, 244–45, 245, Pl. 14 digitalis, Alpheus 99, 101, 104 elongatus, Strigopagurus 263, 264, Pl. 16 digitalis, Trapezia 479, 480, 481 eminens, Agononida 229–30, 230 dilatatus, Ceratocarcinus 449 empheres, Uroptychus 223, 224, 225 dimorphus, Athanas 107, 111, 112 endeavourae, Munida 233, 233, 234 dimorphus, Sympagurus 284, 285–86, 285, Pl. 17 endeavouri, Ephippias 359, 362 Diogenes 218, 252, 259–60 endeavouri, Metapenaeus 37, 40 diogenes, Paragiopagurus 281, 282, 283, Pl. 17 enigma, Pseudocheles 70 Diogenidae 9, 217, 250, 251–65, 255, 258, 264, 270, enigma, Rhynchocinetes 76, 77 Pl. 15, Pl. 16 Enigmaplax 493, 495 Diplostraca 21 Enoplometopidae 160, 161–62, 165, Pl. 11 Disciadidae 70 Enoplometopoidea 159–60 disjunctus, Sergestes 48 Enoplometopus 159, 161–62 dispar, Pisidia 218, 243, 243, 245–46, Pl. 14 ensis, Metapenaeus 33, 37, 40 Distosquilla 538 enthrix, Polycheles 152, 154

564 Index

Ephippias 357 foresti, Palaemonella 86, 94, 94 Epialtinae 348, 349, 351–55 Forestia 463 Epigodromia 305 formosanus, Raymunida 239, 239 Epipedodromia 305 fossor, Synalpheus 108, 109, 114, 115 Eplumula 315–16 Fractosternalia 159 Eriphia 441, 442 franklini, Kimbla 349, 368, 373 Eriphiidae 291, 294, 441–47, 443, 444, 448, 460, frontalis, Paguristes 218, 260, 261, 262, Pl. 15 Pl. 4, Pl. 5 Fultodromia 298, 300, 305–06 Eriphiinae 441, 442 fultoni, Rochinia 349, 385, 386, 387 Erugosquilla 538–39 Funchalia 32, 34–35 Eryoneicus 151 fungilifera, Arcania 338, 339 erythodactyla, Parasesarma 297, 501, 508–09, 509, fusca, Metapenaeopsis 33, 35, 36 Pl. 29 gaimardii, Leptomithrax 296, 347, 349, 369, 370, erythrodactyla, Sesarma 508 372, Pl. 20 Erythrosquilla 522 gaimardii, Paragrapsus 297, 501, 502, 503, 505, Pl. 29 Erythrosquillidae 520, 522, 533 Galathea 9, 220, 229, 231–32 Erythrosquilloidea 518, 520, 522 Galatheidae 9, 215, 219, 220, 228–40, 230, 233, Pl. 13 esculentus, Penaeus 33, 38, 40 Galatheoidea 215, 216, 219–46 etheridgei, Pilumnus 454, 455, Pl. 25 Galearctus 207, 210–11 Ethusinae 324 Gastroptychus 220, 221 Etisinae 461, 465, 468 Gebiacantha 190 Etisus 467, 468 Gecarcinidae 289, 500 Eualus 118, 119 Gennadas 27, 28, 30–31 Eubrachyura 290, 291, 324–515 Geograpsus 500 Eucalliax 184 georgei, Ovalipes 414, 416, 417 Eucarida 2, 14, 22 Georgeoplax 437 Eugonatonotidae 73 germanus, Synalpheus neptunus 116 Eumalacostraca 2, 14, 22 Geryonidae 291, 293, 406–07, 406 Eumedoninae 293, 448, 449, 451 gibber, Chlorotocella 130, 131 Eumedonus 448, 449, 451 gibber, Parapandalus leptorhynchus 131 Eumunida 220, 221 gibbosa, Vercoia 136, 138, 141 Eupasiphae 59 gibbosus, Notostomus 63, 66,67 Euphausiacea 14, 15, 16, 22 gigas, Pseudocarcinus 406, 441, 445, Pl. 4, Pl. 5 euphrosyne, Alpheus 106 gilchristi, Gennadas 28, 29, 29,30 Eupilumnus 446 gilesii, Eupasiphae 59, 60 Eurysquilloidea 518 glaberrima, Ceratoplax 456, 457 Eutrichocheles 176 glaberrima, Tetralia 480 Euxanthinae 462, 468–69 glabra, Georgeoplax 434, 436, 437 Euxanthus 468–69 glabra, Plagusia 513, 514, 514 exaratus, Leptodius 470, 472, 473 glacialis, Hymenodora 64, 66,67 excavata, Dromidiopsis 306 globifer, Leptomithrax 349, 369, 372, 373, 378 excavatus, Thyrolambrus 398, 399, 399 Globopilumnus 446 exsculptus, Euxanthus 467, 469 globosa, Lamarckdromia 298, 300, 301, 302, 306, Pl. 18 exul, Discias 72 Glypheidae 159 faba, Pinnixia 485 Glypheoidea 159 Facetotecta 21 Glyphocrangon 142–43 facetus, Alpheus 99, 100, 104 Glyphocrangonidae 55, 141–43 falcatus, Gonodactylaceus 522, 523, 524 Glyphus 60 fasciata, Anchisquilla 537, 540 Gnathophyllidae 56, 81–84, 82, Pl. 7 Faughnia 535 Gnathophylloides 81, 83 faxoni, Solenocera 42, 43, 44 Gnathophyllum 81, 83 feriata, Charybdis 407, 424, 426, 430 goldsboroughi, Prismatopus 373, 378, 379 ferruginea, Trapezia 480 Gomeza 404, 405 filholi, Pagurus 276 Gonatonotus 448, 449, 451 fimbriata, Platymaia 349, 363, 364 Goneplacidae 9, 291, 294, 297, 406, 433–39, 435, fimbriatus, Heteropilumnus 456, 458, Pl. 25 436, 437, 439, 460, Pl. 24 fissifrons, Achaeus 358 Gonodactylaceus 523 fissifrons, Pilumnus 453, 454, 455, 456 Gonodactylidae 519, 522, 523, 524 flindersi, Philocheras 137, 138, 140 Gonodactyloidea 518, 522–26 flindersi, Uroptychus 223, 224, 225 Gonodactylus 523 Flindersoplax 436 Goreopagurus 272, 274 floridus, Atergatis 475, 477 gracilimanus, Uroptychus 223, 224, 225 foliacea, Aristaeomorpha 25, 26 gracilipes, Rhizopa 457 foliaceum, Petalidium 47, 50 gracilis, Alpheus 100, 104

565 Marine Decapod Crustacea of Southern Australia

gracilis, Arcania 339, 340 Hepatinae 326 gracilis, Hymenodora 64, 66,67 hesperius, Uroptychus 223, 224, 225 gracilis, Nematocarcinus 73, 74,75 Heterocarpus 133 gracilis, Parapontophilus 137, 139 heterochir, Neopilumnoplax 434, 435, 437 gracilis, Processa 128 Heteropilumnus 449, 458 grahami, Erugosquilla 536, 539, 542, Pl. 31 Heterosquilla 528 grahami, Plesionika 130, 132, 134 Heterotremata 290, 291, 324–483 graminea, Coralliocaris 92 hexaceras, Upogebia 194 grandispinis, Paracyclois 326 Hexapinus 439 granti, Athanas 107, 112 Hexapodidae 218, 291, 439–41, 440, Pl. 24 granti, Pachycheles 242, 243, 244 Hexapus 440 granulata, Charybdis 424, 425, 426, 430 hickmani, Pinnotheres 484, 485, 486 granulatus, Portunus 419, 420, 421 hilensis, Schizophroida 374, 380 granuliferus, Hexapinus 218, 439, 440, Pl. 24 Hippa 218, 249 granulimanus, Pagurixus 273, 275 Hippidae 218, 247, 248, 249–50 granulosa, Ashtoret 345, 346 Hippoidea 215, 216, 247–50 granulosa, Choniognathus 368, 373 Hippolyte 121 granulosa, Eurynome 368 Hippolytidae 9, 56, 57, 58, 118–126, 120, 122, 124, granulosus, Cyclograpsus 297, 501, 502, 503, 503, Pl. 9, Pl. 10 504, Pl. 28 Hirsutodynamene 309 Grapsidae 9, 290, 291, 293, 297, 433, 490, 500–12, hirta, Lomis 216, 218, 241, 241, Pl. 14 502, 503, 506, 507, 509, 512, Pl. 28, Pl. 29 hirtimanus, Pagurus 276 Grapsinae 502, 505–08 hispida, Domecia 480, 482 gravelei, Ancylocheles 243, 243, 244 hispidum, Eruma 368 gravelei, Porcellana 244 hispidus, Stenopus 147, 148, 149, Pl. 11 gregaria, Munida 232, 234, Pl. 13 histrix, Paralomis 266, 269, 269 Griffinia 352 hiyashii, Heterocarpus 133 gunnii, Patiriella 10 hodgkini, Hymenosoma 391, 393, 395, 396 haanii, Portunus 419, 420 holothuriensis, Ostracotheres 484, 485, 486 haanii, Scyllarides 210, 212 holthuisi, Lipkius 73 Hadrosquilla 534 holthuisi, Nephropsis 162, 163, 165, 166 haematosticta, Leucosia 336, 337, 338 holthuisi, Periclimenes 92 haigae, Propagurus 277, 277 homarus, Panulirus 201, 202 hailstonei, Alpheus 100, 103, 104–05, 109 Homola 312 Haledromia 306 Homolidae 218, 290, 291, 310, 311–14, 313, 347, Pl. 18 halei, Huenia 350, 351, 352 Homolochunia 312 Halicarcinides 394 Homolodromia 311 Halicarcinus 9, 390, 391, 394–95 Homolodromiidae 290, 292, 310–11, 310, 3183, 347 haliotidis, Orthotheres 484, 485, 485 Homoloidea 290, 298, 311–16 Haliporoides 43–44 Hoplocarida 2, 14, 21 halli, Hymenopenaeus 42, 43, 44 hortus, Michelea 177, 177 hamano, Erythrosquilla 522, 533 huegelii, Jasus 201 handrecki, Pagurixus 273, 275, 276, Pl. 16 Huenia 351, 352 hanseni, Lucifer 42,45 humilis, Pilumnus 453 Hapalocarcinidae 482 Hyastenus 385 Haplogastrinae 266 Hymenoceridae 81 harpagatrus, Synalpheus 108, 109, 114, 115 Hymenodora 64, 66–67 harpax, Harpiosquilla 537, 541, 543, Pl. 31 Hymenopenaeus 43, 44 Harpiosquilla 543 Hymenosoma 395 hasswelli, Athanas 111, 112 Hymenosomatidae 9, 291, 292, 296, 347, 365, haswelli, Lioxantho 472 390–97, 392, 396, Pl. 21 haswelli, Miersiella 471, 472, 473 Hypochonchinae 298 haswelli, Munida 220, 232, 234 Hypothalassia 441, 444 haswellianus, Helograpsus 297, 501, 502, 504, Pl. 28 hystrix, Phalangipus 349, 384, 386, 387 healyi, Laomedia 170, 172, 188, 188 Ibacus 207, 211–12 Helice 504 ignobilis, Hippolyte 121 helleri, Pseudoliomera 462, 464, 464 Ilyograpsus 505 Heloeciinae 490, 493 imperialis, Calcinus 253, 255 Heloecius 493 impressus, Neoxanthias 471, 473, 474 Helograpsus 504 Inachinae 348, 349, 355–65, 365 Hemisquilla 526 Inachoidinae 348, 349, 365–66 Hemisquillidae 522, 525–26, Pl. 30 incerta, Agononida 229, 230 hendersoni, Gastroptychus 221 incertus, Gennadas 28, 29,30 hepatica, Calappa 327, 328, 329 incisa, Austrodromidia 300, 303

566 Index

incisus, Ibacus 212 lacustris, Amarinus 392, 392, 393 indicus, Arete 109, 111 laevimanus, Pachygrapsus 502, 506, 507, 508 indicus, Oncopagurus 281, 282, 283 laevis, Amarinus 296, 391, 392, 393, 396, Pl. 21 indicus, Penaeus 40 laevis, Bellidilia 297, 334, 340, 341, 342, Pl. 19 indicus, Thenus 207 laevis, Belosquilla 536, 538, 545, Pl. 30 inermis, Ceratoplax 458 laevis, Lissocarcinus 408, 409, 409, Pl. 22 inflata, Scopimera 490, 492–93, 494 laevis, Paragrapsus 297, 501, 503, 505, Pl. 29 innominatus, Halicarcinus 4, 366, 391, 394, 396 laevis, Pentacheles 152, 153, 154, 155 inopinata, Neoglyphea 159 laevissima, Chlorodiella 464, 465 inornata, Oratosquillina 536, 546, 547 Lamarckdromia 306 insignis, Austrodromidia 300, 302, 303 lamarcki, Xanthias 471, 474 insignis, Psopheticus 434, 436, 438 lambriformis, Merocryptus 332, 334, 336, Pl. 19 insuetus, Kasim 527, 528, 529 lamellata, Stimdromia 298, 300, 303, 307 integra, Mertonia 456, 458 laminata, Sergia 50, 50,51 integrifrons, Nectocarcinus 411, 411, 412 Laomedia 188 intermedia, Ebalia 297, 334, 335, Pl. 19 Laomediidae 169, 170, 171, 187–89, 188 intermedia, Thalamita 427, 428, 429, 430 Laperousazi, Uroptychus 223, 224, 225, 227 intermedium, Macrobrachium 89, 90 lappacea, Antilibinia 352 intermedius, Palaemon 85, 88, 89, 90 lappacea, Griffinia 349, 352, 354 intermedius, Philocheras 137, 138, 140, Pl. 10 lata, Lenisquilla 537, 540, 544 interrupta, Oratosquillina 536, 546, 547 latens, Calcinus 253–54, 255 investigatoris, Benthesicymus 28, 29,30 lateralis, Stimdromia 298, 300, 303, 307 investigatoris, Pseudopalicus 498, 499, 499 latidactyla, Vitjazmaia 349, 363, 365 iocosta, Synalpheus 114, 115 latifrons, Macrophthalmus 297, 491, 492, 493, 495 iridescens, Ovalipes 414, 416, 417 latimanus, Parapagurus 282, 283, 284 Isopoda 2, 22 latipes, Dumea 349, 356, 360, 362, Pl. 20 isos, Munida 233, 233, 234 latisulcatus, Penaeus 33, 34, 38, 40, Pl. 7 Ixoides 339 latreillei, Macrophthalmus 491, 492, 496 janitor, Pagurus 276, 277 Latreillia 315 japonica, Calappa 326, 328, 329 Latreilliidae 290, 291, 314–16, 315, Pl. 18 japonica, Charybdis 4, 407, 424, 425, 426, 430 Latreillopsis 312, 314 japonica, Munida 234 Latreutes 121 japonica, Sergia 50, 50,51 latro, Birgus 250 Jasus 199, 200–01 latus, Uroptychus 223, 224, 225–26, 227 jerviensis, Pagurixus 273, 275–76 laurentae, Paguristes 260, 261, 262 Jonas 404 laurentae, Plesionika 130, 132, 133, 134 jurichi, Levisquilla 537, 540, 544 lavauxi, Cyclograpsus 503 kaempferi, Macrocheira 445 leachii, Helice 501, 502, 504 kai, Homolodromia 310, 311 Leander 88, 90 Kakaducarididae 81, 84 Leandrites 87 kapala, Cymonomus 318, 319 Lebbeus 118, 121 kapala, Glyphocrangon 142, 142 Lenisquilla 543–44 kapala, Munida 232, 233, 234 Leontocaris 9, 118, 123 kapala, Pasiphaea 59, 60,61 Leptochela 61 kapala, Quollastria 536, 545, 547–48, Pl. 32 leptodactyla, Willemoesia 157 Kasim 528 Leptodius 471–72 kempi, Gennadas 29, 29,30 Leptograpsodes 506 Kempina 543 Leptograpsus 507 kensleyi, Munidopsis 235, 236, 237 Leptomithrax 9, 366, 367, 368–69, 372, 380, 381 kermadecensis, Dorphinaxius 173, 174, 175, 175 leptorhynchus, Parapandalus 131 kermadecensis, Polycheles 152, 154 Leptostraca 14, 21 kimberi, Tozeuma 119, 125 Leucifer 45 Kimbla 368 Leucisca 333 kingstoni, Pilumnus 454, 455 Leucosia 331, 332, 336–37, 339 Kraussia 469 Leucosiidae 9, 291, 292, 297, 326, 331–44, 334, 338, Kraussiinae 461, 469 341, 342, 345, Pl. 19 kuiteri, Rhynchocinetes 76, 77 Leucosiinae 331, 336–39 kullar, Homolochunia 310, 312, 314 leura, Michelea 176 lacazei, Aegaeon 137, 138 levigena, Leucisca 332, 333, 334 laccadivensis, Periclimenes 86, 94,95 Levisquilla 544 lacertosus, Achaeus 357, 358, 361 limosus, Biffarius 181, 182, Pl. 12 lacertosus, Pagurus 276 lindae, Metapenaeopsis 33, 34, 35, 36 Lachnopodus 471 Linuparus 199, 201–02 laciniatus, Eupilumnus 442, 444, 446 Liocarcinus 413–14

567 Marine Decapod Crustacea of Southern Australia

Liomera 469–70 macropus, Thalamita 427, 428, 429 Liomerinae 461, 469–70 madreporae, Periclimenes 92 Lipkius 73, 74 maenas, Carcinus 4, 296, 366, 407, 410, 411, Pl. 22 Lissocarcinus 408–09 magnifica, Galathea 230, 231, 232, Pl. 13 Lissosabinea 139 Majidae 9, 290, 291, 292, 347–90, 350, 353, 354, Lithodes 266, 268 356, 359, 361, 363, 365, 370, 371, 373, 374, 377, Lithodidae 215, 216, 250, 266–70, 267, Pl. 16 378, 383, 386, Pl. 20, Pl. 21 Lithodinae 266 Majinae 348, 351, 366–81 Litocheira 437, 513 Majoidea 347 litoreus, Palaemon 86, 88,90 major, Planes 502, 506, 507, 508 Litosus 355, 362 Malacostraca 2, 14, 21 litosus, Uroptychus 222, 223, 226, 227 mammillaris, Myra 333, 340, 341, 342 littoralis, Enigmaplax 492, 495, 494, 505 maoria, Platymaia 363, 364 livingstonei, Mictyris 488 Mareotis 495 lobidens, Alpheus 99, 101, 102, 103, 105, 109 marginata, Munidopsis 235, 236, 237 Lomidae 240 marini, Agononida 229, 230, 231 Lomis 241 marionis, Nauticaris 118, 124, 124, Pl. 9 Lomisidae 215, 216, 218, 240–42, 241 marmoratus, Saron 118, 124, 125, Pl. 10 longicarpus, Mictyris 488, 489, Pl. 27 marsupialis, Glyphus 59, 60, 60 longicristatum, Perisesarma 501, 509, 509 martia, Plesionika 130, 131, 134 longimanus, Ceratocarcinus 449, 450 martini, Polycheles 152, 154, 156 longimanus, Tumulosternum 381 Matuta 344–45, 346 longipes, Panulirus 202, 203 Matutidae 291, 292, 344–347, 346, 407, Pl. 19 longirostris, Trigonoplax 391, 396, 397, Pl. 21 Matutinae 326 longisetosus, Paguristes 260, 262 mawsoni, Antarctus 208, 209, 210 longispina, Lithodes 266, 268, 269, Pl. 16 Maxillipoda 13, 21 longistylus, Penaeus 40 mcneilli, Anchisquilloides 537, 538, 539, 543, Pl. 30 longitaenia, Pasiphaea 59, 60,62 Megametope 470, 472 longvae, Uroptychus 223, 226 megistos, Dardanus 257 lophochir, Diogenes 258, 259 melanocauda, Austrosquilla 527, 530, 531 lophodactylus, Synalpheus 114, 116 melanoura, Harpiosquilla 537, 541, 543, Pl. 31 Lophogastrida 14, 16, 22 Melicertus 38 Lophopagurus 272, 274 Menaethius 353 lophos, Calappa 327, 328, 329, 330 Meningodora 67 Lophozozymus 477 Menippidae 441 lottini, Alpheus 100, 103, 105, 109 Menippinae 441, 444–45 lucasi, Halicarcinus 391, 394 merguiensis, Penaeus 40 Lucifer 45–46 Merhippolyte 123 Luciferidae 24, 42, 45–46 meridionalis, Carcinoplax 434, 436, 436, Pl. 24 lunaris, Ashtoret 345, 346, Pl. 19 Merocryptus 332, 336 Lupocyclus 418, 420 Mertonia 449, 458 luteus, Ceratoplax 456, 457, 458, Pl. 25 Metacarcinus 402 Lybia 470 Metacrangon 137, 138, 139 Lyramaia 385–86 Metalpheus 99, 113 Lyreidus 321–22 Metanephrops 162, 163–65 Lysiosquilla 530 Metapenaeopsis 32, 35–36 Lysiosquillidae 519, 527, 528, 530, 533, Pl. 30 Metapenaeus 32, 33, 36–37, 40 Lysiosquilloidea 518, 526–34 metavitulans, Sicyonia 41, 42 mabahissae, Aristeus 25, 26, 26 Michelea 176–77 maccullochi, Cyrtomaia 349, 360, 361 Micheleidae 170, 171, 176–77 maccullochi, Pseudomicippe 349, 374, 379 Michelopagurus 272 maccullochi, Synalpheus 117 Micippa 9, 382–83 macgillivrayi, Ommatocarcinus 434, 437, 438 Microhalimus 367, 372 macleayi, Metapenaeus 33, 36, 37, 41 Micropagurus 272, 274–75 macphersoni, Nephropsis 163, 165, 166 microphthalma, Physetocaris 127, 136 Macrobrachium 85, 89, 90 microphylla, Michelea 170, 176, 177, 177 macrocolus, Bythiopagurus 272, 273 Microprosthema 146 macrodactylus, Alpheus 101, 105 microps, Bathysquilla 520, 521, Pl. 30 macrodactylus, Palaemon 4, 86, 88,90 microps, Munida 234 macromeles, Pseudopalicus 498, 499, 499 microspina, Mursia 330 Macrophthalminae 490, 493, 495–96 Mictacea 14, 22 Macrophthalmus 490, 492, 495–96 Mictyridae 291, 293, 296, 487–89, 489, Pl. 27, Pl. 28 Macropipinae 413 Mictyris 488 Macropodia 355, 357, 362 middletoni, Austrosquilla 527, 530, 532

568 Index

Miersiella 472 neptunus, Oncinopus 364 Miersiograpsus 512, 513 neptunus, Synalpheus 114, 116 mikado, Kempina 536, 543, 545, Pl. 32 neptunus, Synalpheus neptunus 116 miles, Charybdis 423, 424, 425 niger, Eumedonus 450, 451 miles, Diogenes 259 nigra, Chlorodiella 464, 465 miles, Distosquilla 536, 538, 539 Nikoides 128 milnedwardsi, Paracyclois 326 nilandensis, Synalpheus 114, 116 mindoro, Meticonaxius 176 nitidus, Austropenaeus 25, 26,27 mineri, Gnathophylloides 81, 82,83 nodipes, Fultodromia 300, 301, 306 minima, Elamenopsis 390 norfolcensis, Bountiana 442, 444, 446 minor, Notomithrax 296, 349, 350, 373, 376, Pl. 21 norvegicus, Nephrops 162, 163 minuta, Pseudopontonia 86, 94, 95–96 notatus, Paraxanthias 471, 474 minutus, Oncopagurus 281, 282, 283 Notomithrax 347, 366, 367, 374, 376 minutus, Pagurus 276 Notopontonia 92 Mithracinae 348, 351, 381–84 Notosceles 322 Mixtopagurus 286 Notostomus 63, 67–68 molleri, Ovalipes 414, 416, 417, Pl. 23 Notostraca 21 mollis, Meningodora 67 novacastellum, Glyphocrangon 142, 142, 143 monilifer, Pilumnus 297, 453, 454, 455, 456, 456, Pl. 25 novaeguinae, Metapenaeopsis 33, 35, 36 monoceros, Menaethius 351, 353, 354 novaehollandiae, Jasus edwardsii 200, Pl. 13 moranti, Pilodius 466 novaehollandiae, Macrobrachium 85, 88,89 mosaica, Rochinia 349, 385, 386, 387 novaezealandiae, Alpheus 101, 103, 105, 109 multidentatus, Stylodactylus 78–79, 79 novaezealandiae, Chlorotocus 130, 132, 132 Munida 9, 220, 229, 232–35, 238 novaezeelandiae, Oplophorus 64, 65, 66,68 Munidopsis 9, 220, 228, 235–38 novaezelandiae, Cancer 402 murrayensis, Philyra 340 novaezelandiae, Metacarcinus 4, 366, 402, 403, Pl. 22 murrayi, Cyrtomaia 349, 358, 360, 361 novaezelandiae, Pinnotheres 486 murrayi, Lithodes 266, 268 novaezelandiae, Spongicoloides 146–47, 147 Mursia 330 nowra, Uroptychus 223, 224, 226 Myodocopida 14 nuytsi, Halicarcinides 391, 394, 396 Myra 340, 342 obesa, Pinnotheres 486 Mysida 16, 22 obliquus, Philocheras 137, 138, 140 Mysidacea 14 obscurus, Cyclodius 464, 465, 466 Mystacocarida 21 obtusidens, Paraxanthodes 474 Nannosquillidae 526, 530, 531, 532, 534, Pl. 30 occidentalis, Actaea peronii 463 Nanocassiope 471, 472 occidentalis, Enoplometopus 162, 165, Pl. 11 nanum, Platepistoma 403–04, 403 occidentalis, Prismatopus 374, 379 nanus, Lophopagurus 217, 273, 274, Pl. 16 octagonalis, Neorhynchoplax 391, 395, 396 nanus, Polycheles 152, 153, 155, 156 octodentata, Austrodromidia 300, 303, 303, 304, Pl. 17 nasutus, Gonatonotus 450, 451 octodentata, Cryptodromia 304 natalis, Geocarcoides 289 octodentatus, Leptograpsodes 297, 503, 506, 506, natator, Charybdis 424, 425, 426, 430, Pl. 23 507, 507, Pl. 29 Nauticaris 123–24 Ocypode 490, 492, 496 Naxia 9, 296, 366, 367, 374–75, 376 Ocypodidae 291, 293, 297, 490–98, 491, 494, Pl. 28 Naxioides 386–87 Ocypodinae 490, 496–97 Neaxius 178 Ogyrides 127, 127 Nectipoda 21 Ogyrididae 56, 58, 126–128, 127 Nectocarcinus 410–12 Ommatocarcinus 438 neglecta, Upogebia 190, 192, 193, 194 Oncinopus 355, 362–64 Nematocarcinidae 9, 56, 73–75, 74 Oncopagurus 281–82 Nematocarcinoidea 56, 73–78 Oplophoridae 9, 54, 55, 63–69, 65, 66 Nematocarcinus 73, 74,75 Oplophoroidea 55, 63–69 Neocallichirus 181, 184 Oplophorus 64, 68 Neolithodes 268 oratoria, Oratosquilla 4, 536, 542, 544, Pl. 32 neomeris, Synalpheus 114, 116 Oratosquilla 544 Neopilumnoplax 437 Oratosquillina 547 Neorhynchoplax 395 orbicularis, Philyra 342, 343 Neoxanthias 472, 474 orbitosinus, Portunus 419, 420, 421 Nephropidae 9, 160, 161, 162–65, 165, Pl. 11 Oregoninae 348 Nephropoidea 160 orientalis, Alope 119, 120, 124, Pl. 9 Nephrops 163 orientalis, Charybdis 424, 425, 426 Nephropsis 163, 166 orientalis, Homola 310,312 Neptunus 418 orientalis, Parathranites 417 neptunus, Metanephrops 162, 163 orientalis, Planopilumnus 434, 435, 438

569 Marine Decapod Crustacea of Southern Australia

orientalis, Sergestes 48 Parapaguridae 9, 251, 270, 279–86, 280, 281, 283, orientalis, Thenus 207 285, Pl. 17 Orithyinae 326 Parapagurus 281, 282, 284 ornatus, Panulirus 202, 203 Parapasiphae 9, 61 ornithorhynchus, Athanas 107, 112 Parapenaeus 33, 37 Orthotheres 484–85 Parapontophilus 139 oryx, Hyastenus 385 Parasesarma 508–09 osculans, Austrosquilla 527, 530, 531 parasocialis, Alpheus 100, 106, 107, 109, Pl. 8 Ostracoda 14, 21 Parasquilla 535 Ostraconotus 271 Parasquillidae 525, 535 Ostracotheres 486 Parasquilloidea 518, 535 Ovalipes 413, 414, 416 Parastacidae 159 ovatus, Halicarcinus 296, 391, 392, 394, Pl. 21 Parastacoidea 159 Oziidae 441 Parathelphusidae 289 Oziinae 441, 446 Parathranites 413, 417 Ozius 442, 446 Paratya 69–70 Pachycheles 244 Paratymolus 360 pachydactylus, Xanthias 471, 474, 475 Paraxanthias 474 Pachygrapsus 507–08 Paraxanthodes 471, 474 pacifica, Hippa 248, 249 Paraxiopsis 175–76 pacifica, Willemoesia 152, 154, 155, 157 parkeri, Projasus 201, 203 pacificus, Alpheus 99, 101, 103, 105, 109 Paromolopsis 314 Pagurapseudidae 217 Parthenopidae 291, 292, 398–400, 399, 449, 476 Paguridae 9, 217, 250, 251, 270–79, 273, 277 parvidens, Thalamita 427, 428, 429, 431 Paguristes 9, 218, 252, 260, 261, 261–63 parvispinosus, Tumulosternum 374, 381 Pagurixus 217, 272, 275–76 parvus, Athanas 112 Paguroidea 215, 216, 217, 250–65 paschalis, Thor 119, 125 Pagurus 215, 217, 271, 272, 276 Pasiphaea 59, 61–62 pagurus, Cancer 401 Pasiphaeidae 55, 57, 58–59, 60 Palaemon 89–91 Pasiphaeoidea 55, 58–62 Palaemonella 92, 94 patulus, Uroptychus 223, 226, 227 Palaemonetes 91 pavoninum, Tozeuma 119, 125 Palaemonidae 56, 58, 84–96, 87, 88, 93, 94, Pl. 8 pectinatus, Sergestes 48 Palaemoninae 84, 85, 86–91 Pedunculata 21 Palaemonoidea 56, 81–97 pedunculatus, Dardanus 257, 258, Pl. 15 Palapedia 469 pelagica, Acanthephyra 66 Palicidae 291, 292, 498–500, 499 pelagicus, Portunus 407, 418, 420, Pl. 23 Palicinae 498 pelsartensis, Palapedia 469 Palinura 16, 17, 151, 159, 169, 197 Penaeidae 24, 31–41, 34, 36, Pl. 7 Palinuridae 16, 197, 198–206, 201, Pl. 2, Pl. 3, Pl. 13 Penaeoidea 24–44 Palinuroidea 197 Penaeopsis 37–38 pallida, Liomera 467, 469, 470 Penaeus 32, 38, 40 palmensis, Metapenaeopsis 33, 35–36 pencillatus, Hemigrapsus 5, 512 paludicola, Ilyograpsus 495, 502, 505, 506 penicillatus, Panulirus 201 Pandalidae 9, 56, 129–35, 131, 132, Pl. 10 penicillifer, Lucifer 45 Pandaloidea 56, 129–35 Pentacheles 152, 154 Panulirus 199, 202–03 pentadactyla, Heterosquilla 527, 528, 529 papillosus, Alpheus 101, 106 Pentastomida 21 papposus, Sympagurus 285 Peracarida 14, 22, 53 Paracrangon 139 Percnon 512 Paracyclois 326 Periclimenes 86, 95 Paradorippe 324 Perinia 354 Paragiopagurus 281, 282 Perisesarma 509 Paraglypturus 184 perlata, Leucosia 337, 338 paragracilis, Metalpheus 107, 109, 113 peronii, Actaea 460, 462, 463, 464, Pl. 26 Paragrapsus 297, 501, 505 peronii, Ibacus 204, 206, 208, 210, 211, 212, Pl. 6 paralacustris, Amarinus 392, 392, 393 perpasta, Hadrosquilla 527, 532, 534, Pl. 30 paraleura, Michelea 176 Petalidium 46, 47 Paralomis 266, 269 Petalomera 307 Paramithrax 367, 376 Petrocheles 244 Paramola 312 Petrochirus 252 Paramunida 229, 238 Petrolisthes 242, 244 Paranaxia 382, 384 petterdi, Dagnaudus 312, 313, Pl. 18 Paranursia 342 Phalangipus 384, 387

570 Index

philargius, Calappa 327, 329, 330, Pl. 18 Pontoniinae 84, 85, 91–96 Philocheras 9, 58, 137, 140, 141 Pontophilus 140 Philyra 333, 340, 342–43 poorei, Biffarius 181, 183 philyra, Micippa 382, 383 poorei, Goreopagurus 273, 274 Philyrinae 331, 332, 339–44 Porcellana 246 Phlyxia 335 Porcellanidae 215, 218, 219, 242–46, 243, 245, Pl. 14 phrixa, Paralomis 266, 269, 269 Porcellanopagurus 271, 276 Phylladiorhynchus 229, 238 Portunidae 290, 291, 293, 296, 407–33, 411, 413, 415, Phyllocarida 21 417, 420, 421, 423, 426, 427, 430, 431, Pl. 22, Pl. 23 Phyllopoda 21 Portuninae 9, 408, 418–22 Phymodius 466 Portunus 418–22 Physetocarididae 56, 127, 135–36 potens, Sergia 50, 50,51 Physetocaridoidea 135–36 prehensilis, Sergia 49, 50, 51 Physetocaris 136 Prismatopus 368, 378–79 picta, Thalamita 427, 428, 429, 430, 431 proales, Munidopsis 235, 236, 237 pictor, Lophozozymus 475, 477 procera, Agononida 229, 230, 231 Pilodius 466 Processa 128 pilosus, Uroptychus 222, 226, 227 Processidae 56, 127, 128–129 Pilumnidae 9, 291, 293, 294, 296, 448–59, 450, 455, Processoidea 56, 128–129 456, 460, Pl. 24, Pl. 25 producta, Elamena 390 Pilumninae 449, 452–57 profundus, Parapontophilus gracilis 139 pilumnoidses, Dynomene 308, 309 Projasus 199, 203 Pilumnopeus 449, 452 Propagurus 271, 277–78 Pilumnus 9, 449, 452–54, 456–57 propensalata, Pontocaris 137, 140 Pinnotheres 486 propinquus, Gennadas 28, 29, 29, 30–31 Pinnotheridae 291, 293, 484–87, 485, Pl. 26 propinquus, Hymenopenaeus 42, 43, 44 Pippacirama 366, 378 proteus, Huenia 352 Pisidia 245–46 prymna, Thalamita 427, 428, 429, 430 Pisinae 348, 351, 384–87 Pseudaristeus 27 pisum, Pinnotheres 486 Pseudocarcinus 445 Plagusia 512, 513–15 Pseudoliomera 464 Plagusiidae 291, 293, 512–13, 514, Pl. 29 Pseudomicippe 367, 379 planatus, Halicarcinus 391, 395, 396 Pseudomunida 220 Planes 500, 508 Pseudopalicus 498–99 planipes, Matuta 345, 346, 346 Pseudopontonia 95–96 Planopilumnus 438 Pseudosquilla 526 Platepistoma 402–04 Pseudosquillidae 522, 525, 526 platipes, Micippa 349, 382, 383 Pseudosquillopsis 535 platycheir, Philyra 341, 343 Pseudozius 436, 438 platycheles, Mictyris 296, 488, 489, Pl. 28 Psopheticus 438 platycheles, Notopontonia 86, 92, 93 pubescens, Leucosia 337, 338 Platymaia 355, 358, 364 pubescens, Portunus 419, 420, 420, 421 plebejus, Penaeus 33, 34, 36, 38, 40, Pl. 7 Puerulus 199, 203–04 plectrorhynchus, Strahlaxius 178, 179 pugettensis, Upogebia 189 Pleocyemata 16, 17, 22, 23, 53 Pugettia 354 Plesionika 130, 133–34 pugil, Paguristes 260, 261, 262 plumifera, Bresilia 71, 71,72 pugnax, Achaeus 349, 357, 358, 361 podocheloides, Achaeus 350, 357, 358, 361 pulchellus, Lophozozymus 475, 477 Podocopida 14 punctata, Ceratoplax 457 Podophthalminae 408, 412 punctata, Megametope 470, 472, 473 Podophthalmus 412 punctulatus, Macrophthalmus 491, 492, 494, 496 Podotremata 290, 298–323 purpureantennatus, Paguristes 260, 261, 262 Polybiinae 408, 413–17 pusillus, Phylladiorhynchus 220, 238, 239 polybioides, Lissocarcinus 409, 409 Pylochelidae 250, 251, 286–87, 287, Pl. 17 Polycheles 152, 154–57 Pylojacquesidae 250, 251 Polychelida 16, 17, 22, 151–57, 220 Pylopaguropsis 271 Polychelidae 9, 16, 151–57, 153, 154, 220 Pyromaia 366 Polydectinae 461, 470 quadrata, Ebalia 336 Polydectus 470 quadraticoxa, Cancellus 254, 255 Polyonyx 246 quadridentatus, Paragrapsus 297, 501, 502, 503, polyphagus, Panulirus 202 505, Pl. 29 Pomatochelidae 270, 286 quadrispinosa, Acanthephyra 64, 65, 66 Pontocaris 137, 140 Quollastria 547–48 Pontonia 95 rachelae, Austrosquilla 527, 532, 534

571 Marine Decapod Crustacea of Southern Australia

ramsayi, Ebalia 335 Scylla 418, 422 ramusculus, Achaeopsis 360 Scyllaridae 16, 197, 206–13, 208, 210, Pl. 6, Pl. 13 ramusculus, Dorhynchus 349, 360, 361 Scyllarides 207, 212 Randallia 343 Scyllarinae 206 Ranilia 322 Scyllarus 206 Ranina 322 Scytoleptus 175 ranina, Ranina 320, 321, 322 sebae, Hyastenus 385 Raninidae 290, 292, 320–23, 321, 407, Pl. 18 sebana, Eriphia 441, 442, 444 Raninoides 322 semidentatus, Aristeus 25, 26 rathbunae, Aegaeon 137 seminudus, Sergestes 47, 48 rathbunae, Campylonotus 80, 80 semisulcatus, Penaeus 40, 101 rathbunae, Pasiphaea 59, 62 senex, Diogenes 218, 258, 259 raymondi, Uroptychus 223, 226, 227 septata, Trapezia 479, 480, 481 Raymunida 229, 239 septemspinosa, Arcania 339, 340, 342 reflexa, Plesionika 130, 132, 134 serenei, Diogenes 259–60 remifera, Platymaia 349, 364 serenus, Palaemon 86, 87, 88, 90–91, Pl. 8 Remipedia 13, 21 Sergestes 46, 47 reynaudii, Amphionides 16 Sergestidae 9, 24, 46–51, 49, 50 Rhinolambrus 398–399 Sergestoidea 24, 45–51 Rhizocephala 21 Sergia 46, 47, 49–51 Rhizopinae 449, 457–58 serpulifera, Paranaxia 383, 384, 387 Rhynchocinetes 9, 58, 76–77 serrata, Nephropsis 163, 164, 166 Rhynchocinetidae 9, 56, 58, 73, 75–77, 77, Pl. 7 serrata, Scylla 407, 421, 422 richardi, Stylopandalus 132, 134 serratifrons, Notosceles 297, 320, 321, 322 richardsoni, Alpheus 101, 106, 107, 108, 109, Pl. 9 serratifrons, Pilumnopeus 450, 452, Pl. 24 richardsoni, Teratomaia 349, 377, 380 serratus, Axius 173 richeri, Lithodes 268 serratus, Rhynchocinetes 76, 77, 77, Pl. 7 richeri, Parapagurus 282, 283, 284 serricornis, Munidopsis 235, 236, 237, 238 Rimicaris 70 serripes, Scytoleptus 175 robillardi, Naxioides 349, 385, 386–87, 386 Sesarma 508, 509 robustus, Gnathophylloides 81, 82,83 Sesarminae 500, 501, 508–09 Rochinia 385, 387 Sessilia 21 rogeri, Gastroptychus 221, 222 setifer, Actumnus 450, 452, Pl. 24 rogeri, Munida 232, 233, 234 setifer, Dardanus 256, 257, 258, 258 rosenbergii, Macrobrachium 84 setosus, Macrophthalmus 491, 492, 496 rostrata, Munidopsis 235, 236, 237 sexpes, Hexapus 218, 439, 440, 440 rostratus, Halicarcinus 296, 391, 394, 395, 396 sexspinosus, Litosus 362, 363 rotumana, Palaemonella 86, 94, 94 sextuberculatus, Parapenaeus 33, 34,37 rotundifrons, Megametope 470, 472, 473, Pl. 26 sibogae, Acetes 47, 50 rubens, Bathyarctus 209, 210 sibogae, Athanas 112 rubridigitalis, Munida 232, 234 sibogae, Haliporoides 42, 43, 44 rubromarginatus, Portunus 419, 420, 420, 421, Pl. 23 sibogae, Heterocarpus 129, 131, 133 rufescens, Schizophrys 349, 377, 380, Pl. 21 sibogae, Metanephrops 162, 163, Pl. 11 rufopunctatus, Pilumnus 452, 453, 456, 455, 456 sibogae, Pseudaristeus 25, 27 rugosus, Portunus 419, 420, 421, 421, Pl. 23 sibogae, Strobopagurus 284, 285 rugulosus, Rhynchocinetes 77 sica, Acanthephyra 63, 64, 66, 66 Sagmariasus 199, 200–01 Sicyonia 41–42, 42 sanguineus, Hemigrapsus 5, 512 Sicyoniidae 24, 41–42 sanguinolentus, Portunus 407, 418, 421, 422, Pl. 23 sigmoideus, Nematocarcinus 73, 74,75 Sarcostraca 21 Silentes 199 sargassi, Sergestes 48 sima, Thalamita 427, 428, 429, 430, 431 Saron 125 similis, Tymolus 317, 317 scabricula, Eriphia 441, 442, 444 simsoni, Acutigebia 189, 190, 191, 192, 193, Pl. 12 scabriuscula, Philyra 341, 343 sinensis, Eriochier 5, 512 Schizophroida 367, 379–80 sinensis, Harpiosquilla 537, 541, 543, Pl. 31 Schizophrys 367, 380 sinuatus, Pagurus 276, 277, Pl. 16 schmitti, Austrolepidopa 248 smithi, Acanthephyra 64, 66 scintillans, Sergia 50, 50,51 smithii, Gonodactylus 522, 523, 524 Scopimera 492–93 socialis, Alpheus 100, 106, 107, 109 scotiae, Pasiphaea 61 soela, Cymonomus 318, 319 sculpta, Epigodromia 300, 305 soela, Sympagurus 284, 285, 286 sculptus, Polycheles 152, 154, 155, 156 Solenocera 44 sculptus, Scyllarus 209 Solenoceridae 24, 42, 43–44 scutatus, Gennadas 28, 29,31 sordidus, Linuparus 201, 201, 202

572 Index

soror, Periclimenes 92 suhmi, Cyrtomaia 349, 360, 361 spatulifer, Prismatopus 349, 374, 378, 379 suhmi, Nephropsis 163, 165, 166 spatulifrons, Cryptopodia 398, 399 suhmi, Polycheles 152, 154, 155, 156 speciosa, Plagusia 515 sulcata, Nephropsis 162, 163 Spelaeogriphacea 14, 22 sulcatifrons, Parapasiphae 59, 60,61 Sphaerodrominae 298 sulcatus, Alpheus 99, 110 spinicaudus, Chlorotocella 130, 131, 131, Pl. 10 sulcatus, Paguristes 260, 261, 263 spinicruris, Munida 233, 233, 235 superba, Coralliocaris 92 spinifera, Ebalia 334, 335, 336 surdus, Polycheles 152, 154, 155, 156 spinifera, Fultodromia 300, 306 sydniensis, Leptochela 58, 59, 60, 61, 72 spinifrons, Austinogebia 189, 190, 191, 192, 193 Sympagurus 281, 281, 282, 284–86 spinifrons, Nectocarcinus 411, 411, 412 Synalpheus 98, 99, 113–117 spinipes, Plesionika 130, 132, 133, 134 Synaxidae 197, 199 spinosa, Hirsutodynamene 308, 309, 309 Syncarida 14, 22 spinosa, Micippa 349, 382–83, 383 Systellaspis 68 spinosa, Naxia 349, 375, 377, 378 taeniatus, Clibanarius 255, 256, Pl. 15 spinosus, Brachynotus 297, 501, 509, 510, 512, Pl. 29 Tanaidacea 14, 22, 217 spinosus, Oplophorus 64, 66,68 Tantulocarida 21 spinosus, Trizocheles 287, 287, Pl. 17 tarda, Pasiphaea 59, 62 spinosus, Trizocheles spinosus 287 tasmanensis, Pugettia 349, 352, 354, 355 spongiarum, Alpheus 101, 106 tasmaniae, Munidopsis 235, 236, 237 Spongicola 146 tasmanicus, Alpheus 101, 110 Spongicolidae 145, 146–48, 147, Pl. 11 Tasmanoplax 495 Spongicoloides 146–47 taylori, Gnathophyllum 81, 82, 83, Pl. 7 squamifera, Thalassina 169 tenuicornis, Leander 88, 88 squammosus, Scyllarides 212 tenuimana, Acanthacaris 162, 163 squamosa, Agononida 229, 230, 231 tenuirostris, Hippolyte 121 squamosus, Paguristes 218, 260, 261, 262 tenuirostris, Stenopus 147, 148, 149, Pl. 11 Squillidae 519, 535–48, 539-42, 545-46, Pl. 30, Pl. Teratomaia 368, 380 31, Pl. 32 terraereginae, Calcinus 254 Squilloidea 518, 535–48 terraereginae, Pilumnus 457 squinado, Maja 372 tessellata, Lybia 470, Pl. 26 steintizi, Albunea 247 testudinarius, Remipes 249 Stenopodidae 145, 147, 148–49, Pl. 11 Tetralia 479–80 Stenopodidea 17, 22, 53, 145–49 tetraspinosa, Latreillopsis 310, 312, 313, 314 Stenopus 148–49 Tetrasquillidae 527–28, 529 stephensoni, Alpheus 101, 104, 107, 109, 110 Thacanophrys 378 Stereomastis 154 Thalamita 423, 425, 428–29 sternocostulatus, Leptomithrax 349, 369, 370, 372, 377 Thalamitinae 408, 422–31 sternoornatus, Gastroptychus 221 Thalassinidae 169 stewarti, Nephropsis 162, 163 Thalassinidea 16, 17, 22, 159, 169–95, 215 Stimdromia 300, 307 Thalassinoidea 169, 215 stimpsoni, Anacinetops 349, 368, 374 Thalassocarididae 129 stimpsonii, Synalpheus 108, 109, 114, 116 thalia, Micippa 382, 383, 383 stimulator, Sergestes 48 Thaumastochelidae 160 Stomatopoda 2, 14, 21, 517–48 theano, Synalpheus 114, 117 Strahlaxiidae 169, 170, 171, 172, 178, 179, Pl. 12 Thecostaca 21 Strahlaxius 173, 178 Thenus 206, 207 strenuus, Alpheus 102, 110 Thermosbaenacea 14, 22 strenuus, Alpheus strenuus 110 Thiidae 469 streptodactylus, Synalpheus 108, 109, 114, 116 Thiinae 400 Stridentes 199 Thomassinia 169 strigimanus, Strigopagurus 263, 264, Pl. 16 Thomassiniidae 169 strigimanus, Trizopagurus 218, 263 thomsoni, Epipedodromia 298, 300, 302, 305 Strigopagurus 252, 263 thomsoni, Platydromia 305 Strobopagurus 280, 281, 284 Thor 125 Stylodactylidae 55, 78–79, 79 Thoracica 21 Stylodactyloidea 55, 78–79 Thoracotremata 290, 291, 324, 484–515 Stylodactylus 78–79 Thyrolambrus 399 Stylopandalus 134 Tiarinia 382, 384 subglobosus, Ostracotheres 484, 485, 486, Pl. 26 tinayrei, Gennadas 28, 29, 31 subsolanus, Uroptychus 223, 226–27, 227 Tisea 251 subsquamata, Galathea 231, 232 tomentosum, Tozeuma 119, 124, 125 subsquamosa, Munidopsis 235, 236, 237 tomentosus, Actaeodes 462, 463

573 Marine Decapod Crustacea of Southern Australia

tomentosus, Pilumnus 453, 455, 457, Pl. 25 Uroptychus 9, 220, 221–27 tooradin, Eucalliax 184 ursus, Notomithrax 349, 373, 376, Pl. 21 tooradin, Paraglypturus 180, 183, 184 valida, Latreillia 315 Tozeuma 9, 125 validum, Microprosthema 146, 147, Pl. 11 Trachypenaeus 32, 38 validus, Pentacheles 152, 154, 155 Trachysalambria 38 variabilis, Coenobita 250 tractabilis, Upogebia 190, 192, 193, 194, Pl. 12 varians, Pseudomicippe 371, 377, 379 transversa, Austrothelphusa 289 variegatus, Leptograpsus 503, 506, 507, 507 transversus, Pachygrapsus 508 Varuninae 500, 501, 505, 509–10, 512 transversus, Polyonyx 218, 243, 243, 246, Pl. 14 velutina, Metapenaeopsis 33, 34,36 Trapezia 480–81 velutinus, Metanephrops 163, 164, 165, 165 Trapeziidae 291, 294, 460, 479–81, 480 Venetus 495 treis, Munidopsis 235, 236, 238 ventricosa, Hippolyte 121 Trichia 476 vercoi, Austrosquilla 527, 531, 534 Trichiinae 476 Vercoia 141 Trichopeltarion 400, Pl. 22 verreauxi, Jasus 200–01, 204, Pl. 3 tridentata, Australoplax 490, 493, 494 verrilli, Munidopsis 235, 236, 238 tridentata, Lissosabinea 136, 138, 139 versicolor, Panulirus 203 tridentatus, Lyreidus 320, 321, 322, Pl. 18 vesca, Meningodora 63, 66,67 tridentatus, Porcellanopagurus 276, 277, 286, Pl. 16 vespertilio, Pilumnus 453, 457 Trigonoplax 395, 397 vestitus, Pilumnus 454, 455, 457 trigonops, Austrolepidopa 218, 248, 248 victoriae, Munidopsis 235, 236, 238 trigonus, Linuparus 201, 201, 202 victoriensis, Carcinoplax 434, 436, 436, Pl. 24 trigonus, Macropodia 349, 362, 363 victoriensis, Philocheras 137, 138, 140 trirufomaculata, Umalia 321, 321, 323 vigil, Podophthalmus 412, 413 triserratus, Lophopagurus 273, 274 villosa, Banareia 475, 476 trispinosa, Alpheopsis 99, 103, 109 villosa, Funchalia 33, 34,35 Trizocheles 287 villosus, Alpheus 100, 107, 109, 110, Pl. 9 truncata, Elamena 393 villosus, Sympagurus 284, 286 truncatus, Ozius 442, 443, 446 vincentiana, Flindersoplax 434, 436, 436, 437 truncifrons, Latreutes 121 vincentianus, Cryptocnemus 332, 333, 334, 336 Trypaea 184–85 vincinus, Eumedonus 450, 451 tsangi, Austrosquilla 527, 531, 534 virescens, Clibanarius 255, 256 tuberculata, Pyromaia 4, 349, 365, 366, Pl. 20 virilis, Aristeus 25, 26 tuberculatus, Leptomithrax 349, 369, 372, 374 Vitjazmaia 355, 364–65 tuberculatus, Paguristes 260, 261, 263 vomeris, Uca 492, 494, 497 tuberculosa, Ebalia 334, 335, 336 waitei, Leptomithrax 349, 369, 372, 374 tuberculosa, Paramicippe 378 wardi, Thaumastochelopsis 160 tuberculosa, Pippacirama 373, 378 wardi, Trichopeltarion 400, 401 tuberculosus, Nectocarcinus 411, 411, 412, Pl. 22 wardi, Tumulosternum 349, 374, 381 tumida, Calvactaea 475, 476 waroona, Strahlaxius 170, 178, 179, Pl. 12 tumida, Cryptodromia 300, 304 werribee, Axiopsis 170, 173, 174, 174, 175, Pl. 11 tumida, Naxia 349, 375, 377, 378 whiteleggii, Galathea 230, 231, 232 tumida, Perinia 351, 354 Willemoesia 157 tumidomanus, Synalpheus 108, 109, 114, 117, Pl. 9 wilsoni, Dromia 298, 299, 300, 304, Pl. 17 tumidus, Jasus 201 wilsoni, Petalomera 304 Tumulosternum 367, 368, 381 woodmasoni, Erugosquilla 536, 539, 542 Tymolus 316–17 woodwardi, Funchalia 34,35 typhlops, Polycheles 152, 155, 156–57 wyvillethomsoni, Platymaia 349, 359, 363, 364 typhlops, Polycheles typhlops 157 Xanthias 471, 474, Pl. 26 typus, Cancellus 217, 254, 255, Pl. 15 Xanthidae 9, 291, 293, 294, 406, 441, 448, 460–78, typus, Lucifer 42,45 464, 467, 473, 475, Pl. 26 Uca 490, 496–97 Xanthinae 462, 470–74 Umalia 322–23 Xenophthalmodes 449, 458 umbilicatus, Galearctus 210, 211 yaldwyni, Bathypaguropsis 272, 273 undecimspinosa, Arcania 338, 339, 340 yaldwyni, Lebbeus 121, 124 undecimspinosa, Bellidilia 340, 342, Pl. 19 yaldwyni, Periclimenes 86, 94,95 undulatipes, Nematocarcinus 73, 74,75 yarramundi, Leontocaris 118, 122, 123 ungulatus, Cyclodius 464, 465, 466 Zalasiinae 461, 476 unidentata, Leucosia 337, 338, 338 Zalasius 476 Unipeltata 21 Zebrida 448, 451 Upogebia 189, 192–94 zeidleri, Uroptychus 223, 227, 227 Upogebiidae 9, 169, 171, 172, 189–95, 191, 192, Zosiminae 462, 477 193, Pl. 12 Zosimus 460, 477

574