STUDIES ON THE SYSTEMATICS OF AUSTRALIAN AMPITHOID AMPHIPODS (Crustacea) WITH A CHECKLIST AND AN INTERACTIVE KEY TO WORLD SPECIES Rachael A. Evans B.Sc. (Hons) Crustacean Section Division of Invertebrate Zoology Australian Museum 6 College St Sydney South Study carried out in fulfilment of award of Masters of Science (by research). University of New South Wales December, 1997 'I hereby declare that this submission is my own work and that, to the best of , my knowledge and belief, it contains no material previously published or written by another person nor material which to a sustantial extent has been accepted for the award of any other degree or diploma of the university or other institute of higher learning, except where due acknowledgement is made in the text.' CERTIFICATE OF ORIGINALITY I hereby declare that this submission is my own work and to the best of my knowledge it contains no materials previously published or written by another person, nor material which to a substantial extent has been accepted for the award of any other degree or diploma at UNSW or any other educational institution. except where due acknowledgement is made in the thesis. Any contribution made to the research by colleagues, with whom I have worked at UNSW or elsewhere. during my candidature. is fully acknowledged. I also declare that the intellectual content of this thesis is the product of my own work, except to the extent that assistance from others in the project's design and conception or in style, presentation and linguistic expression is acknowledged. ii TABLE OF CONTENTS Title Page Disclaimer ii Table of Contents iii Acknowledgments vi Abstract vii CHAPTER 1: SYSTEMATICS OF SOME AUSTRALIAN AMPITHOID AMPHIPODS INTRODUCTION 1 Status of Australian species 2 A~s 4 MATERIAL AND METHODS 5 SYSTEMATICS 6 Key to Ampithoid Taxa known from Australian waters 6 Species Taxonomy 10 Ampithoe eremitis sp. nov. 10 Ampithoe gallaharae sp. nov. 15 Cymadusa aestos sp. nov. 19 Cymadusa chuawe sp. nov. 25 Cymadusa elegantis sp. nov. 30 Cymadusa imbroglio Rti.'o,no-r""o:t"' 34 Cymadusa setosa (Ha.6w{.,\I) 39 Exampithoe ecklonicola sp. nov. 58 iii Paragrubia edgari sp. nov. 62 Pseudopleonexes nexis sp. nov. 67 Sunamphitoe angrox sp. nov. 72 Sunamphitoe fantome sp. nov. 75 DISCUSSION Generic distributions 80 Species diversity 83 Conservation 84 REFERENCES 87 ILLUSTRATIONS 95 CHAPTER 2: CHECKLIST TO THE AMPITHOID SPECIES OF THE WORLD INTRODUCTION Aims 138 Taxonomic History 138 World Systematics 141 CHECKLIST Amphithoides 146 Amphitholina 147 Ampithoe 148 Cymadusa 173 Exampithoe (Exampithoe) 181 Exampithoe (Melanesius) 182 Macropisthopous 184 Paragrubia 184 iv Peramphithoe 186 Plumithoe 193 Pseudoamphithoides 195 Pseudopleonexes 196 Sunamphitoe 197 REFERENCES 199 CHAPTER 3: HANDBOOK TO INTERACTIVE KEY TO THE WORLD SPECIES OF AMPITHOID AMPHIPODS Title page 217 Introduction 218 How to Install this Program 218 The Data 219 Start: How to get to lntkey 220 How to ID animals 221 Glossary 223 V ACKNOWLEDGMENTS I would like to thank Dr Jim Lowry and Dr lain Suthers for all their help and encouragement throughout the duration of this project. Also special thanks to Alistair Poore for his advice and help; Helen Stoddart for all her advice on dissecting and examining amphipods; Wendy and Craig Taylor for inking the plates, and to Roger Springthorpe for advice on the making of the plates. vi ABSTRACT Twelve species of common ampithoid amphipods, including ten new species are reported from Australian waters. This brings the total number of valid species known from this area to twenty-one. The genus Pseudopleonexes is recorded for the first time from Australian waters. Ten species, Ampithoe eremitis sp. nov., A. gallaharae sp. nov., Cymadusa aestos sp. nov., C. chuawe sp. nov., C. elegantis sp. nov.; Exampithoe ecklonicola sp. nov., Paragrubia edgari sp. nov., Pseudopleonexes nexis sp. nov., Sunamphitoe angrox sp. nov., and Sunamphitoe fantome sp. nov., are described. Cymadusa imbroglio Rabindranath, 1972 is redescribed and recorded for the first time from Australian waters. Cymadusa setosa (Haswell, 1879a) is a poorly described species from Australian waters. The taxonomic status of this species is discussed, it is redescribed and illustrated and a neotype is established. A traditional dichotomous key is provided for the twenty one known species from Australian waters. In this thesis the taxonomic problems for a number of the more common species from Australian waters are discussed. An annotated checklist and an interactive key to the world species are presented. vii CHAPTER 1 SYSTEMATICS OF SOME AUSTRALIAN AMPITHOID AMPHIPODS INTRODUCTION Recent and current studies (Freewater and Lowry, 1994; Poore and Lowry, 1997) have begun to show that ampithoid amphipods are among the most abundant and diverse amphipod crustaceans inhabiting algae in coastal Australian waters. Poore and Lowry (1997), in a study of subtidal macroalgae in Port Jackson, New South Wales, Australia, showed that about 50% of individuals examined were of the family Ampithoidae and five of the ten most abundant species in their study were ampithoids. However, only 14.6% of the total number of species observed in the study were ampithoids. The abundance and diversity of ampithoids has also been shown in various other studies on macroalgae and seagrasses ( Hay et al., 1987; Duffy, 1990; Edgar, 1983a, b; Nelson, 1979, 1980). In the past ten years the number of species described from Australian waters has risen sharply. Before 1994, only six species of ampithoids were known from Australian waters. Of these six only three were ide-v'l-\ifii,.b\~. In the last five years, sixteen new species have been described (Freewater and Lowry, 1994; Poore and Lowry, 1997; Evans, present study). Also, two species described originally from other locations have been reported for the first time in Australian waters (Poore and Lowry, 1997; Evans, present study). These studies plus observations of collections indicate that a large 1 number of ampithoid species remain to be documented and described from Australian waters. Ampithoid amphipods have played important roles in a number of ecological studies (Edgar, 1983a, b; Nelson, 1979, 1980). However, due to inadequate systematic work, the ecological studies often lack essential taxonomic information. An example is the ecological study of Edgar (1983) in southeastern Tasmania. The taxa used in this work were unidentified to species level and the generic level identifications were to some extent inaccurate. Two of Edgar's undescribed species are documented in this paper. Studies such as Edgar ( 1983) indicate the need for a good systematic basis before ecological studies proceed. This basis can provide a useful and comprehensive guide to Australian ampithoid systematics for studies in areas such as conservation, ecology, biogeography and evolution. Recent research is starting to remedy this situation, but there is still a lot to be done. For example, of the 14 species recorded recently from two studies in the Sydney area, (on Sargassum sp. from Port Jackson - Poore and Lowry, 1997; on Ecklonia sp. from Botany Bay - present study, and N. Gallaher, unpublished) only three of the species overlap. Status of Australian Species Before Barnard and Karaman ( 1991) only six species were recorded from Australian waters (Table 1). However, Ampithoe cinerea Haswell, 1879a, Ampithoe flindersi Stebbing, 1888 and Ampithoe australiensis Bate, 1862 are unidentifiable. This is either because the original description was based on only 2 female specimens or the original description was incomplete and poorly illustrated. Occasionally this means that the species have been placed in the wrong genus. Poore and Lowry (1997) transferred Ampithoe quadrimana to the genus Plumithoe, and Cymadusa variata (Sheard, 1936) to the genus Paragrubia. Table 1: Ampithoid species recorded from Australian Waters. Ampithoe australiensis Bate, 1862 (unidentifiable). Ampithoe cinerea Haswell, 1879a (unidentifiable). Ampithoe caddi Poore and Lowry, 1997. Ampithoe eremitis sp.nov. Ampithoe flindersi Stebbing, 1888 (unidentifiable). Ampithoe gal/aharae sp. nov. Ampithoe kava Myers, 1985 (recorded by Poore and Lowry, 1997). Ampithoe ngana Poore and Lowry, 1997. Cymadusa aestos sp. nov. Cymadusa chuawe sp. nov. Cymadusa elegantis sp. nov. Cymadusa imbroglio Rabindranath, 1972 (recorded here). Cymadusa munnu Poore and Lowry, 1997. Cymadusa setosa (Haswell, 1879a) (recorded here). Exampithoe (Melanesius) kutti Poore and Lowry, 1997. Exampithoe (Melanesius) ecklonicola sp. nov. Paragrubia edgari sp. nov. 3 Paragrubia variata (Sheard, 1936). Peramphithoe parmerong Poore and Lowry, 1997. Plumithoe quadrimana (Haswell, 1879b) (recorded by Poore and Lowry, 1997). Pseudopleonexes nexis sp. nov. Sunamphitoe angrox sp. nov. Sunamphitoe fantome sp. nov. Sunamphitoe graxon Freewater and Lowry, 1994. Aims Ampithoid amphipods are diverse and abundant herbivores in Australian waters, but they are not well known taxonomically. The aim of chapter 1 is to describe the more common and problematical species from Australian waters. As an aid to identification a dichotomous key to the known and new Australian species is presented. 4 MATERIAL AND METHODS Specimens were obtained from collections of a number of studies. These include Graham Edgar's published and unpublished studies on algae from Tasmania and Queensland, Nicole Gallahar's unpublished study at Cape Banks, recent Australian Museum collections along the New South Wales coast, Museum of Tasmania