Invertebrate Fauna of Korea

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Inver tebrate Fauna of Korea

Volume 21, Number 46 Crabs and Zoeas V Vol. 21, No. 46 Arthropoda: Malacostraca: Decapoda: Brachyura: Dromioidea, Calappoidea, Orithyioidea, Leucosioidea, Pinnotheroidea Crabs and Zoeas V

Flora and Fauna of Korea NIBR National Institute of Biological Resources Ministry of Environment

Invertebrate Fauna of Korea

Volume 21, Number 46

Crabs and Zoeas V

Arthropoda: Malacostraca: Decapoda: Brachyura: Dromioidea, Calappoidea, Orithyioidea, Leucosioidea, Pinnotheroidea

2018

National Institute of Biological Resources Ministry of Environment Invertebrate Fauna of Korea Volume 21, Number 46 Crabs and Zoeas V Arthropoda: Malacostraca: Decapoda: Brachyura: Dromioidea, Calappoidea, Orithyioidea, Leucosioidea, Pinnotheroidea

Copyright © 2018 by the National Institute of Biological Resources Published by the National Institute of Biological Resources Environmental Research Complex, 42, Hwangyeong-ro, Seo-gu Incheon 22689, Republic of Korea www.nibr.go.kr All rights reserved. No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording, or otherwise, without the prior permission of the National Institute of Biological Resources. ISBN: 978-89-6811-354-3(96470), 978-89-94555-00-3(Set) Government Publications Registration Number: 11-1480592-001394-01 Printed by Junghaengsa, Inc. in Korea on acid-free paper Publisher: National Institute of Biological Resources Authors: Hyun-Sook Ko, Seok-hyun Lee (Shilla University) Project Staff: Jung Sun Yoo, Hyun Jong Kil, Eun Jung Nam, Hyeonggeun Kim, Kwang Soo Kim Published on August 31, 2018 Invertebrate Fauna of Korea

Volume 21, Number 46

Crabs and Zoeas V

Arthropoda: Malacostraca: Decapoda: Brachyura: Dromioidea, Calappoidea, Orithyioidea, Leucosioidea, Pinnotheroidea

Hyun-Sook Ko and Seok-hyun Lee Shilla University The Flora and Fauna of Korea logo was designed to represent six major target groups of the project including vertebrates, invertebrates, insects, algae, fungi, and bacteria. PREFACE

Biological resources include all organisms and their genetic characteristics. Utilization and conserva tion of these resources have the capacity to improve human life and to enhance the world we live in. It is therefore imperative that the practical and potential value of these organisms is conserved and used wisely. The first step towards this goal is to document our diversity and to study it in detail. Biological resources increase the competitiveness of a nation through their use as fundamental resources for making highly valu ed products, such as new lines of medicines, new materials, and new drugs. Since the Nagoya Protocol was adopted in 2010 and entered into force during the 12th Conference of the Parties of the Convention on Bio logical Diversity (CBD) in 2014, it has been expected that competition for biological resources will become more intensive under the rapidly changing circumstance on access to and sharing of genetic resources (ABS). To cope with a new international paradigm on issues related to biological resources, the Ministry of the Environment of Korea enacted a law entitled ‘An act on access and benefit sharing of genetic resources’ on August 17, 2017. Each nation in the world is investigating and clearing information of native species within its territory in order to secure its sovereignty rights over biological resources. The National Institute of Biological Reso urces (NIBR) of the Ministry of Environment has published the ‘Flora and Fauna of Korea’ since 2009 to manage biological resources in comprehensive ways and to enhance national competitiveness by building up the foundation for the sovereignty over biological resources. Professional research groups, consisting of professors and other taxonomic experts, have systematically examined 15,154 species of vascular plants, animals and other organisms over the past 10 years and have published their findings in 184 volumes in Korean and 189 volumes in English, and two volumes of worldwide monographs covering 216 species of invertebrates. This year, 14 volumes of the Flora and Fauna of Korea in both Korean and English versions including 391 species of invertebrates and insects are additionally published. Flora and Fauna of Korea are the first professional records to describe all the species of the nation in a comprehensive way, and they would contribute to level up the taxonomic capacity. The NIBR will continue to publish flora and fauna of Korea that will contribute conservation and appli cation of biological resources for successful implementation of the ABS protocol. Finally, I would like to express my sincere appreciation to authors who spared no effort to publish the Flora and Fauna of Korea.

President National Institute of Biological Resources

CONTENTS

List of Taxa 3 Introduction 6 Materials and Methods 8 Taxonomic Notes 11 1. Cryptodromia tumida Stimpson 13 2. Lauridromia dehaani (Rathbun) 14 3. Lewindromia unidentata (Rüppell) 15 4. Metadromia wilsoni (Fulton and Grant) 16 5. Paradromia japonica (Henderson) 18 6. Petalomera granulata Stimpson 19 7. Calappa clypeata Borradaile 21 8. Calappa hepatica (Linnaeus) 22 9. Calappa lophos (Herbst) 23 10. Calappa philargius (Linnaeus) 24 11. Cycloes granulosa De Haan 25 12. Mursia armata De Haan 27 13. Mursia trispinosa Parisi 27 14. Ashtoret lunaris (Forskål) 29 15. Matuta planipes Fabricius 31 16. Matuta victor (Fabricius) 32 17. Orithyia sinica (Linnaeus) 35 18. Arcania globata Stimpson 38 19. Arcania undecimspinosa De Haan 39 20. Hiplyra platycheir De Haan 40 21. Lyphira heterograna Ortmann 41 22. Merocryptus lambriformis A. Milne-Edwards 42 23. Myra celeris Galil 43 24. Nursia rhomboidalis (Miers) 44 25. Philyra kanekoi T. Sakai 45 26. Philyra syndactyla Ortmann 46 27. Pyrhila carinata (Bell) 47 28. Pyrhila pisum (De Haan) 48

1 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

29. Euclosia obtusifrons (De Haan) 49 30. Leucosia anatum (Herbst) 50 31. Urnalana haematosticta (Adams and White) 52 32. Pinnixa tumida Stimpson 54 33. Arcotheres sinensis (Shen) 55 34. Pinnaxodes major Ortmann 57 35. Pinnotheres cyclinus Shen 58 36. Pinnotheres pholadis De Haan 59 37. Pinnotheres tsingtaoensis Shen 60 38. Sakaina koreensis Kim and T. Sakai 61 Literatures Cited 62 Plates 73 Index to Korean Names 87 Index to Scientific Names 89

2 LIST OF TAXA

LIST OF TAXA

Subphylum Crustacea Brünnich, 1772 Class Malacostraca Latreille, 1802 Subclass Eumalacostraca Grobben, 1892 Order Decapoda Latreille, 1802 Infraorder Brachyura Latreille, 1802 Superfamily Dromioidea De Haan, 1833 Family Dromiidae De Haan, 1833 Genus Cryptodromia Stimpson, 1858 Cryptodromia tumida Stimpson, 1858 Genus Lauridromia McLay, 1993 Lauridromia dehaani (Rathbun, 1923) Genus Lewindromia Guinot and Tavares, 2003 Lewindromia unidentata (Rüppell, 1830) Genus Metadromia McLay, 2009 Metadromia wilsoni (Fulton and Grant, 1902) Genus Paradromia Balss, 1921 Paradromia japonica (Henderson, 1888) Genus Petalomera Stimpson, 1858 Petalomera granulata Stimpson, 1858 Superfamily Calappoidea De Haan, 1833 Family Calappidae De Haan, 1833 Genus Calappa Weber, 1795 Calappa clypeata Borradaile, 1903 Calappa hepatica (Linnaeus, 1758) Calappa lophos (Herbst, 1782) Calappa philargius (Linnaeus, 1758) Genus Cycloes De Haan, 1837 Cycloes granulosa De Haan, 1837 Genus Mursia Desmarest, 1823 Mursia armata De Haan, 1837 Mursia trispinosa Parisi, 1914 Family Matutidae De Haan, 1835

3 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Genus Ashtoret Galil and Clark, 1994 Ashtoret lunaris (Forskål, 1775) Genus Matuta Weber, 1795 Matuta planipes Fabricius, 1798 Matuta victor (Fabricius, 1781) Superfamily Orithyioidea Dana, 1852 Family Orithyiidae Dana, 1852 Genus Orithyia Fabricius, 1798 Orithyia sinica (Linnaeus, 1771) Superfamily Leucosioidea Samouelle, 1819 Family Leucosiidae Samouelle, 1819 Subfamily Ebaliinae Stimpson, 1871 Genus Arcania Leach, 1817 Arcania globata Stimpson, 1858 Arcania undecimspinosa De Haan, 1841 Genus Hiplyra Galil, 2009 Hiplyra platycheir De Haan, 1841 Genus Lyphira Galil, 2009 Lyphira heterograna Ortmann, 1892 Genus Merocryptus A. Milne-Edwards, 1873 Merocryptus lambriformis A. Milne-Edwards, 1873 Genus Myra Leach, 1817 Myra celeris Galil, 2001 Genus Nursia Leach, 1817 Nursia rhomboidalis (Miers, 1879) Genus Philyra Leach, 1817 Philyra kanekoi T. Sakai, 1934 Philyra syndactyla Ortmann, 1892 Genus Pyrhila Galil, 2009 Pyrhila carinata (Bell, 1855) Pyrhila pisum (De Haan, 1841) Subfamily Leucosiinae Samouelle, 1819 Genus Euclosia Galil, 2003 Euclosia obtusifrons (De Haan, 1841) Genus Leucosia Weber, 1795 Leucosia anatum (Herbst, 1783)

4 LIST OF TAXA

Genus Urnalana Galil, 2005 Urnalana haematosticta (Adams and White, 1849) Superfamily Pinnotheroidea De Haan, 1833 Family Pinnotheridae De Haan, 1833 Subfamily Pinnothereliinae Alcock, 1900 Genus Pinnixa White, 1846 Pinnixa tumida Stimpson, 1858 Subfamily Pinnotherinae De Haan, 1833 Genus Arcotheres Manning, 1993 Arcotheres sinensis (Shen, 1932) Genus Pinnaxodes Heller, 1865 Pinnaxodes major Ortmann, 1894 Genus Pinnotheres Bosc, 1802 Pinnotheres cyclinus Shen, 1932 Pinnotheres pholadis De Haan, 1835 Pinnotheres tsingtaoensis Shen, 1932 Genus Sakaina Seréne, 1964 Sakaina koreensis Kim and T. Sakai, 1972

5 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

INTRODUCTION

Crabs are covered with a thick exoskeleton, armed wih a pair of chelipeds, and walking with four pairs of ambulatory legs. They are found in all of the world’s oceans, whereas some crabs live in freshwater, terrestrial or semi-terrestrial regions. They vary in size from the pinnotheroid crab with a few mm of carapace width, to the Japanese majoid crab with 4 m length of leg. Over 6,700 species are recognized in the checklist of extant brachyuran crabs of the world (Ng et al., 2008). The species of the five superfamilies of the Dromioidea, Calappoidea, Orithyioidea, Leucosioidea, and Pinnotheroidea can be often recognized by their different morphologies, behaviors, habitats, and distribu- tions of adults. The dromioid species typically carry pieces of camouflage over their backs by using of a subchelate dactylus of the last ambulatory leg. In the calappoid species a large proximal tooth on the dactylus of cheliped is present to open clams in the Calappidae or the paddle-like dactyli of all ambulatory legs exhibit in the Matutidae. The orithyioid species is only one species of Orithyia sinica distributed from north China to Yellow Sea of Korea. The leucosioid species having a triangular merus of the third maxilliped are small and usually globose. The pinnotheroid species are small and covered with a soft exoskeleton. They in- habit the mantle cavities of the clams and live in the tubes or burrows of other invertebrates. In his brachyuran monograph of Korea, Kim (1973) recorded 32 species of these five superfamilies. They were six dromioid species [Cryptodromia tumida, Petalomara wilsoni (= Dromia wilsoni), Dromia dehaani (= Lauridromia dehaani), Dromidia unidentata (= Lewindromia unidentata), Paradromia japonica, and Petalomera granulata], eight calappoid species [Calappa terraereginae (= Calappa clypeata), Calappa hepatica, Calappa lophos, Calappa philargius, Mursia armata, Mursia curtispina trispinosa (=Mursia trispinosa), Matuta lunaris (= Ashtoret lunaris), and Matuta planipes], an orithyioid species of Orithyia sinica, 12 leucosioid species [Arcania globata, Arcania undecimspinosa, Philyra heterograna (= Lyphira heterograna), Merocryptus lambriformis, Myra fugax (= Myra celeries), Philyra kanekoi, Philyra syndactyla, Philyra yangmataoensis (= Pyrhila carinata), Philyra pisum (= Pyrhila pisum), Leucosia longifrons (= Leucosia anatum), Leucosia obtusifrons (= Euclosia obtusifrons), and Leucosia haematosticta (= Urnalana haematosticta)], and five pinnotheroid species [Pinnixa tumida, Pinnotheres sinensis (= Arcotheres sinensis), Pinnaxodes major, Pinno- theres cyclinus, and Pinnotheres pholadis]. Later, a pinnotheroid species of Sakaina koreensis was included in the list of The Korean Society of Systematic Zoology (1997). Recently, five additional species (two calappoid species of Cycloes granulosa and Matuta victor, two leucosioid species of Hiplyra platycheir and Nursia rhomboidalis, and a pinnotheroid species of Pinnotheres tsingtaoensis) were reported in the Nation- al List of Species of Korea (Invertebrates-VII) (Ko, 2015). Currently, 38 species of these superfamilies (six dromioid, ten calappoid, an orithyioid, 14 leucosioid, and seven pinnotheroid species) are known from Ko- rean waters and these are reported upon in this present study.

6 INTRODUCTION

Brachyuran female crabs brood their eggs on their pleopods. Spawning usually takes place between late Spring to early Autumn. The eggs hatch into the first zoeas bearing little resemblance to their parents. At each molt, the zoeas grow in size and some appendage setae increase in number. For example, the four na- tatory setae on the distal segment of the first and second maxillipeds acquire two additional setae during the subsequent moult from first to second stage zoea. The number of zoeal stage can be different depending on the familiar species. It tends to be somewhat consistent, for example, two or four in the dromioid, five in the calappoid, three in the leucosioid and orithyioid, and three or five in the pinnotheroid species. The terminal zoeal stage metamorphoses into the megalopal phase which is transitional between a planktonic and benthic life style. The abdomen remains in the horizontal position and is still functional for swimming via the use of the pleopods. The megalop in turn metamorphoses into the first crab stage in which the abdomen is finally tucked under the sternum and is no longer function for swimming with the pleopods eventually developing further after a number of juvenile molts. In females the pleopods hold eggs and in males the first two pleopods only are retained and modified to be used to transfer sperm into the female. The color images of living zoeas are helpful for the identification of zoeas because the patterns of chro- matophores are slightly different at the species level. In this volume these images of representative species of zoeas belonging to the five superfamilies are provided and provisional keys are included for the identification of zoeas of species.

7 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

MATERIALS AND METHODS

The adults examined for the present study were collected from the mainland of South Korea associated islands. Adults were examined under a Leica EZ40 microscope and digital photographs of crabs taken using an Olympus E-30 camera. The following abbreviations are used in this book: CL (carapace length) from the tip of rostrum to the posterior dorsal margin of the carapace and CW (carapace width) across the widest point of the carapace excluding length of carapace spines. Measurements were made by using digital vernier caliper (Mitutojo, CD-15APX). The terminology used for carapace regions and spines generally follows that of Poore (2004). Most of zoeas were hatched from ovigerous female crabs in the laboratory and dissected under a Leitz zoom stereomicroscope. Zoeal appendages were examined under a Leitz Laborlux S microscope. Chromato- phores of zoeas were recorded by using a Leitz camera and zoom stereomicroscope. These images were then processed in Photoshop. All specimens were preserved in 95% ethyl alcohol. In zoeas, CL is measured from the anterior margin of the eye to the most posterior dorsal margin of the carapace and RDL (rostral spine to dorsal spine length) from the tip of rostral spine to the tip of dorsal spine. Descriptions of zoeas are based on the malacostracan somite plan, from anterior to posterior. Setal armature of the appendages is described from proximal to distal segments (Clark et al., 1998). The classification generally follows those of Ng et al. (2008) and WoRMS (World Register of Marine Spe- cies, 2015). Most of specimens examined were deposited at the National Institute of Biological Resources, Korea and Department of Biological Science, Silla University, Busan, Korea.

Morphology of crabs and zoeas

The dorsal surface of the carapace is divided into several regions (Fig. 1A). The five pairs of pereiopods consist of chelipeds and four pairs of ambulatory legs (Fig. 1B). The antennules and antennae are sensory or- gans in crabs. The buccal (mouth) cavity is covered by the third maxillipeds (Fig. 1B). The abdomen of male is with two pairs of pleopods (gonopods), compared with the female which is broader and with four pairs of pleopods. The zoeal carapace is globose and usually armed with three spines (rostral, dorsal and lateral). These spines may function as buoyancy devices. It may be difficult for a predator to swallow. Three pairs of mouthpart appendages i.e., the mandible, maxillule, maxilla, are used for feeding and the first and second maxillipeds are functional for swimming. The abdomen is flexible for rapid locomotion. The telson is usually forked and sometimes triangular or broadened with a median lobe (Fig. 2).

8 MATERIALS AND METHODS

Abdomen Basis

Fig. 1. External‌ morphology of crabs. A, dorsal view of Matuta victor; B, ventral view of Calappa lophos.

9 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Abdomen B C A

E F

G H I

Fig. 2. Diagrammatic external morphology of brachyuran zoea Matuta victor. A, lateral view; B, antennule; C, antenna; D, mandibles; E, maxillule; F, maxilla; G, maxilliped 1; H, maxilliped 2; I, pleon and telson.

10 TAXONOMIC NOTES

TAXONOMIC NOTES

Subphylum Crustacea Brünnich, 1772‌ Gab-gak-a-mun (갑각아문)

Class Malacostraca Latreille, 1802‌ Yeon-gab-gang (연갑강)

Subclass Eumalacostraca Grobben, 1892‌ Jin-yeon-gab-a-gang (진연갑아강)

Order Decapoda Latreille, 1802‌ Sib-gak-mok (십각목)

Infraorder Brachyura Latreille, 1802‌ Dan-mi-ha-mok (단미하목)

Superfamily Dromioidea De Haan, 1833‌ Hae-meon-chi-rae-sang-gwa (해면치레상과)

Most species cryptic, carrying pieces of sponges or ascidians over their backs using ambulatory legs 3, 4. Merus of maxilliped 3 subquadrate (Pl. 3). Genital opening of female on coxa of ambulatory leg 2, that of male on leg 3. Abdomen of both sexes with 6 somites. Small uropod plate present (Pl. 4). Two families (Dromiidae De Haan, 1833 and Dynomenidae Ortmann, 1892) and more than 130 species exist. Only one family Dromiidae known in the country.

11 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Family Dromiidae De Haan, 1833‌ Hae-meon-chi-rae-gwa (해면치레과)

Carapace subcircular, subhexagonal, or heptagonal. Ambulatory legs 3, 4 reduced, similar in size, subchelate, formed by distal propodal spine and datylus (Pl. 8). Spermatothecae present on female thoracic sternal sutures 7/8. Small uropod plate present between edges of abdominal somite 6 and telson. Male with 2 pairs of pleopods, first pair stout as gonopod 1, second pair styliform as gonopod 2 (Pl. 9).

Genera 39 (6 in Korea), species over 110 (6 in Korea).

Distribution: Worldwide.

Key to the genera of family Dromiidae

1. Dorsal surface of carapace entirely covered with dense setae or granules...... 2 - Dorsal surface of carapace relatively smooth...... Cryptodromia 2. Dorsal surface of carapace covered with dense setae...... 3 - Dorsal surface of carapace covered with granules...... Petalomera 3. Carapace broader than long...... 4 - Carapace as broad as long...... 5 4. Carapace covered with stiff setae...... Lauridromia - Carapace covered with soft setae...... Metadromia 5. Anterolateral margin of carapace without teeth...... Lewindromia - Anterolateral margin of carapace with teeth...... Paradromia

Key to the zoeas of family Dromiidae

General zoeal morphology of dromiid species similar to that of anomurans. Rostral carapace spine present, shorter than CL. Antenna with 4 plumose setae on endopod, scaphocerite with 10 or 11 plumose setae, protopod without spine at base of endopod. Endopod of maxillule with 2, 2+4 setae. Endopod of maxilla with 3+2+4 or 3, 2+4 setae. Basis and endopod of maxilliped 1 each with 2+3+3+3 and 3, 3, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+2 and 3, 3, 2, 5 setae, respectively. Telson fan-shaped posteriorly, posterior margin with 6+6 setae, outermost spine present.

1. Carapace without posterolateral spines...... 2 - Carapace with posterolateral spines...... Lauridromia dehaani 2. Carapace with dorsal spine...... Metadromia wilsoni - Carapace without dorsal spine...... Paradromia japonica

12 DROMIIDAE: Cryptodromia

Genus Cryptodromia Stimpson, 1858‌ Gaet-ga-hae-meon-ch-rae-sok (갯가해면치레속)

Carapace broader than or as broad as long; dorsal surface smooth, convex; regions indistinct. Front tridentate. Ambulatory legs 1, 2 lobed, nodular, or tuberculate; legs 3, 4 reduced.

Species 22 (1 in Korea).

1. Cryptodromia tumida Stimpson, 1858 (Pl. 1) Gaet-ga-hae-meon-chi-rae (갯가해면치레)

Cryptodromia tumida Stimpson, 1858c, p. 240; T. Sakai, 1965, p. 8, pl. 3, fig. 3; Kim, 1973, p. 276, fig. 78, pl. 74, fig. 50; Kim and Chang, 1985, p. 44; McLay, 1993, p. 199 (key); Poore, 2004, p. 304. Cryptodromia tumida typica T. Sakai, 1936b, p. 18, fig. 4a, b, pl. 7, fig. 1; 1976, p. 13, fig. 4a-c; McLay, 2001, p. 88 (key).

Adult: Carapace subcircular, broader than long; dorsal surface smooth, convex, with minute setae; front bluntly tridentate, median tooth blunt, directed ventrally; anterolateral margin with 3 subequal teeth. Subhe- patic tubercle subequal size to first tooth. Chelipeds scattering of few tubercles; each carpus with 3 tubercles on outer surface; each propodus with 4 blunt tubercles on upper margin, outer surface bearing lines of small granules; fingers gaping in basal half. Ambulatory legs 1, 2 similar in size; each carpus with tubercle on distal margin; each propodus lobed on distal margin; leg 3 slightly shorter than leg 4, dactylus opposed by propodal spine. Abdomen of female with 6 somites. Telson broader than long. Color in live specimen yellowish. CL 6.8 mm, CW 8.2 mm.

Habitat: Intertidal-5 m.

Distribution: Korea, Japan, Malay Peninsula, Australia.

Korea: JJ.

Specimen examined: JJ: 1♀(Munseom: 19.v.2014, J.H. Park). Remarks: Larvae are unknown.

13 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Genus Lauridromia McLay, 1993‌ Hae-myeon-chi-rae-sok (해면치레속)

Carapace slightly broader than long; dorsal surface covered with stiff short setae; regions distinct. Front tridentate. Ambulatory legs 1, 2 smooth; legs 3, 4 reduced.

Species 2 (1 in Korea).

2. Lauridromia dehaani (Rathbun, 1923) (Pls. 2-4) Hae-myeon-chi-rae (해면치레)

Dromia rumphii (not Weber, 1795) H. Milne-Edwards, 1837, p. 174; Stimpson, 1858c, p. 240; Alcock, 1901, p. 44, pl. A, fig. 3, pl. 2, fig. 4; Borradaile, 1903b, p. 576, pl. 33, fig. 1. Dromia dormia (not Linnaeus, 1763) Rathbun, 1902, p. 32; Shen, 1931, p. 96, figs. 3a, b, 4a, b, pl. 4; Ch- hapgar, 1957, p. 3, pl. 1, fig. a. Dromia dehaani Rathbun, 1923b, p. 68; T. Sakai, 1936b, p. 8, pl. 5, fig. 1; Kamita, 1941, p. 22, fig. 2; Kim, 1973, pp. 273, 607, fig. 76, pl. 9, fig. 48; T. Sakai, 1976, p. 8, pl. 2, fig. 1. Miyake, 1983, p. 5, pl. 2, fig. 3; Dai and Yang, 1991, p. 17, fig. 4 (1), pl. 1 (1). Lauridromia dehaani McLay, 1993, p. 146 (key); Chan et al., 2009, p. 49, figs. 22-24. Dromia dehaani: Terada, 1983, p. 364, figs. 1-3 (A-I) (zoeal stages).

Adult: Carapace subcircular, slightly broader than long; dorsal surface strongly convex, covered with stiff short setae; cardiac and branchial grooves distinct; front tridentate, median tooth extremely small, deflexed, lateral teeth slightly larger than median one; anterolateral margin with 3 bluntly triangular teeth, tips acute. Subhepatic region with broadly triangular tooth between postorbital and first anterolateral carapace tooth. Chelipeds large, stout, covered with dense short setae; each merus with 6-8 minute tubercles on upper margin; each carpus with 2 large blunt tubercles on outer surface; each propodus with 3 blunt tubercles on upper surface; fingers not gaping when closed, tips white. Ambulatory legs 1, 2 shorter than chelipeds, covered with dense short setae; each merus, carpus, and propodus smooth; each dactylus with 9-12 small spines on posterior margin; legs 3, 4 reduced, similar in size, each dactylus opposed by propodal spine. Abdomen with 6 somites. Uropod plate present. Telson rounded on distal margin. Gonopod 1 stout tube with dense distal setae; gonopod 2 long, slender, styliform. CL 61.6 mm, CW 67.6 mm. Zoea I: Rostral carapace spine ca. 1CL; dorsal and lateral carapace spines absent; posterolateral carapace spines short. Antenna with 4 plumose setae on endopod, scaphocerite with 10 plumose setae, protopod without spine at base of endopod. Endopod of maxillule with 2, 2+4 setae. Endopod of maxilla with 3+2+4

14 DROMIIDAE: Lewindromia

setae. Basis and endopod of maxilliped 1 each with 2+3+3+3 and 3, 3, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1 +1 +2 and 3, 3, 2, 5 setae, respectively. Abdomen without pleopod bud. Telson fan-shaped posteriorly, posterior margin with 6+6 setae, outermost spine present. CL 1.20 mm (Terada, 1983).

Habitat: Mud; 30-100 m.

Distribution: Korea, Japan, China, Taiwan, India, Indonesia, Gulf of Aden.

Korea: GN, JJ.

Specimen examined: GN: 1♂(Dadaepo: 22.xii.2014, S.H. Lee).

Remarks: This species is the largest dromiid crab from Korean waters. It has four zoeal stages (Terada, 1983).

Genus Lewindromia Guinot and Tavares, 2003‌ Teol-bo-hae-myeon-chi-rae-sok (털보해면치레속)

Carapace as broad as long; dorsal surface covered with dense long setae; regions indistinct. Front tridentate. Anterolateral margin of carapace without teeth. Ambulatory legs 1, 2 smooth; legs 3, 4 reduced, each propodus with 2 spines on outer margin.

Species 1 (1 in Korea).

3. Lewindromia unidentata (Rüppell, 1830) Teol-bo-hae-myeon-chi-rae (털보해면치레)

Dromia unidentata Rüppell, 1830, p. 16, pl. 4, figs. 2, 2a, pl. 6, fig. 9; Alcock, 1901, p. 47, pl. 2, fig. 6. Dromidia unidentata: Kossmann, 1880, p. 67; T. Sakai, 1936b, p. 13, fig. 2, pl. 6, fig. 2; Kim, 1973, p. 275, fig. 77, pl. 74, fig. 49; T. Sakai, 1976, p. 11, fig. 2a, b, pl. 2, fig. 2; Miyake, 1983, p. 6, pl. 2, fig. 4. Dromidia unidentata hawaiiensis Edmondson, 1922, p. 34, pl. 2D, fig. 1a-j. Dromidia unidentata unidentata Garth, 1957, p. 316. Cryptodromia unilobata Campbell and Stephenson, 1970, p. 240, fig. 2A-I. Cryptodromiopsis unidentata: McLay, 1993, p. 188 (key), 192, figs. 7a-k, 18a; 2001, p. 843, fig. 8. Lewindromia unidentata: Guinot and Tavares, 2003, p. 74.

15 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Adult: Carapace subcircular, as broad as long; dorsal surface convex, covered with dense long setae; regions indistinct, only branchial groove marked; front tridentate, median tooth extremely small, lateral teeth continuous with supraorbital teeth, deflexed ventrally; anterolateral margin without teeth. Chelipeds slender, covered with dense long setae; cutting margins of fixed and movable fingers each with 5 acute and 7 small teeth, respectively. Ambulatory legs 1, 2 similar in size; legs 3, 4 reduced, each propodus with 2 unequal spines on outer margin, each dactylus opposed by stout propodal spine. CL 18.2 mm, CW 19.2 mm (Kim, 1973).

Habitat: Under rocks; 30-50 m.

Distribution: Korea, Japan, Thailand, Philippines, Singapore, India, Indonesia, Mozambique, Red Sea, Persian Gulf, Pakistan, Sri Lanka, Andaman Islands, Australia, Chesterfield Islands, New Caledonia, Ker- madec Islands, Mergui Archipelago, Easter Island, Hawaiian Islands.

Korea: JJ.

Specimen examined: None.

Remarks: Larvae are unknown.

Genus Metadromia McLay, 2009‌ Som-teol-mut-hi-sok (솜털묻히속)

Carapace much broader than long; dorsal surface sculptured, covered with dense soft setae; regions distinct. Front tridentate. Ambulatory legs 1, 2 smooth, not nodular or lobed; legs 3, 4 reduced.

Species 8 (1 in Korea).

4. Metadromia wilsoni (Fulton and Grant, 1902) (Pls. 5-9) Som-teol-mut-hi (솜털묻히)

Cryptodromia wilsoni Fulton and Grant, 1902, p. 61, pl. 9. Dromia pseudogibbosa Parisi, 1915, p. 102, pl. 2, figs. 1, 2; Balss, 1922, p. 106; Yokoya, 1933, p. 97. Petalomera wilsoni Rathbun, 1923a, p. 154, pl. 42, fig. 1; T. Sakai, 1936b, p. 34, fig. 9, pl. 1, fig. 4; 1965, p. 9, pl. 4, fig. 2; 1976, p. 24, pl. 6, fig. 1; Kim, 1973, p. 278, fig. 80, pl. 10, fig. 51; Miyake, 1983, p. 6, pl. 3, fig. 1; McLay, 1991, p. 470, pl. 1B, figs. 6a-d, 7a-c, 8a-c. Dromia wilsoni McLay, 1993, p. 151 (key), 156, fig. 16e; Chan et al., 2009, p. 43, figs. 18, 19. Metadromia wilsoni: McLay, 2009, pp. 15-19, fig. 4a, b.

16 DROMIIDAE: Metadromia

Petalomera wilsoni: Terada, 1983, p. 364, figs. 1-3 (A”-I”) (zoeal stages).

Adult: Carapace subhexagonal, 1.4 times broader than long; dorsal surface slightly convex, sculptured, covered with dense soft setae; cardiac and branchial grooves distinct; front tridentate, median tooth small, directed ventrally, lateral teeth continuous with supraorbital margin; anterolateral margin with 3 strong teeth, tips acute. Subhepatic region with broadly triangular tooth between postorbital and first anterolateral carapace tooth. Chelipeds large, stout; each merus smooth; each carpus with 2 blunt distal tubercles on outer surface, acute tooth on inner angle; each propodus with 4 proximal granules on upper margin, outer surface smooth; fingers gaping when closed. Ambulatory legs 1, 2 shorter than chelipeds, each merus smooth, each carpus and propodus with distal tubercle on anterior margin, each dactylus with 4 spines on posterior margin; legs 3, 4 reduced, similar in size, each dactylus opposed by propodal spine. Abdomen with 6 somites. Uropod plate present. Telson subtriangular. Gonopod 1 stout tube with distal dense setae; gonopod 2 slender, styliform. CL 28.7 mm, CW 40.1 mm. Zoea I: Rostral carapace spine shorter than CL; dorsal and lateral carapace spines short; posterolateral carapace spines absent. Antenna with 4 plumose setae on endopod, scaphocerite with 10 plumose setae, protopod without spine at base of endopod. Endopod of maxillule with 2, 2+4 setae. Endopod of maxilla with 3, 2+4 setae. Basis and endopod of maxilliped 1 each with 2+3+3+3 and 3, 3, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+2 and 3, 3, 2, 5 setae, respectively. Abdomen with pleopod bud. Telson fan-shaped posteriorly, posterior margin with 6+6 setae, outermost spine present. CL 1.33 mm.

Habitat: Subtidal-520 m.

Distribution: Korea, Japan, Taiwan, Philippines, Australia, New Zealand, South Africa.

Korea: GW, GB, GN.

Specimen examined: GW: 1♂(Samcheok: 22.xii.2011, S.H. Lee); GN: zoeas (Busan: 14.i.2011, S.H. Lee). Remarks: This species is one of the most widespread dromiid crabs distributed from the South Atlantic Ocean to the Indo-West Pacific. It has two zoeal stages.

Genus Paradromia Balss, 1921‌ Gaet-som-mut-hi-sok (갯솜묻히속)

Carapace as broad as long; dorsal surface with dense short setae; regions distinct. Front tridentate. Ambu- latory legs 1, 2 lobed; legs 3, 4 reduced.

Species 2 (1 in Korea).

17 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

5. Paradromia japonica (Henderson, 1888) (Pls. 10, 11) Gaet-som-mut-hi (갯솜묻히)

Cryptodromia japonica Henderson, 1888, p. 6, fig. 2. Cryptodromia asiatica Parisi, 1915, p. 105, pl. 2, fig. 3. Paradromia japonica: Balss, 1922, p. 108, figs. 3, 4; T. Sakai, 1936a, p. 32, figs. 1, 2; McLay, 1993, p. 164 (key). Petalomera japonica: T. Sakai, 1936b, p. 36, pl. 8, fig. 1; 1976, p. 22, fig. 12; Kamita, 1941, p. 23, fig. 3; Kim, 1973, p. 280, fig. 81, pl. 10, fig. 52; Miyake, 1983, p. 7; Dai and Yang, 1991, p. 27, pl. 2 (5). Petalomera japonica: Hong and Williamson, 1986, p. 1259, figs. 1-9 (larval stages).

Adult: Carapace subcircular, as broad as long; dorsal surface strongly convex, covered with dense short setae, scattering of granules; regions distinct; front tridentate, median tooth small, blunt, deflexed ventrally, lateral teeth blunt, slightly larger than median one; anterolateral margin with 3 blunt teeth, second one largest. Chelipeds large, stout, covered with dense soft setae; each merus smooth; each carpus scattering of granules, with 3 large tubercles on outer surface; each propodus scattering of granules, with 2 large tubercles on upper surface; fingers not gaping when closed. Ambulatory legs 1, 2 shorter than chelipeds, covered with dense soft setae; each merus smooth; each carpus and propodus lobed on distal margin; each dactylus with 4-5 small spines on posterior margin; leg 3 reduced, shorter than leg 4, dactylus opposed by propodal spine; propodus of leg 4 with small spine on outer margin, dactylus opposed by propodal spine. Abdomen with 6 somites. Uropod plate present. Telson trapezoid. Gonopod 1 stout tube with dense distal setae; gonopod 2 slender, styliform. CL 17.3 mm, CW 18.3 mm. Zoea I: Rostral carapace spine shorter than CL; dorsal and lateral carapace spines absent; posterolateral carapace spines absent. Antenna with 4 plumose setae on endopod, scaphocerite with 11 plumose setae, protopod without spine at base of endopod. Endopod of maxillule with 2, 2+4 setae. Endopod of maxilla with 3, 2+4 setae. Basis and endopod of maxilliped 1 each with 2+3+3+3 and 3, 3, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+2 and 3, 3, 2, 5 setae, respectively. Abdomen with pleopod bud. Telson fan-shaped posteriorly, posterior margin with 6+6 setae, outermost spine present. CL 1.46 mm (Hong and Williamson, 1986).

Habitat: Mud or sand; 10-100 m.

Distribution: Korea Japan, China, Indonesia, Australia.

Korea: GW, GN, JN, JJ, GG.

Specimen examined: GN: 1♂(Busan: 2.vi.2015, S. H. Lee).

Remarks: It has two zoeal stages (Hong and Williamson, 1986).

18 DROMIIDAE: Petalomera

Genus Petalomera Stimpson, 1858‌ So-reum-mut-hi-sok (소름묻히속)

Carapace as broad as long; dorsal surface entirely covered with granules; regions distinct. Front tridentate. Ambulatory legs 1, 2 covered with granules; legs 3, 4 reduced.

Species 3 (1 in Korea).

6. Petalomera granulata Stimpson, 1858 So-reum-mut-hi (소름묻히)

Petalomera granulata Stimpson, 1858c, p. 240; T. Sakai, 1936b, p. 37, fig. 10a-c, pl. 1, fig. 3; 1976, p. 25, pl. 4, fig. 4; Kim, 1973, p. 281, fig. 82, pl. 75, fig. 53; Miyake, 1983, p. 7, pl. 3, fig. 3; Dai and Yang, 1991, p. 25, fig. 7 (1-3), pl. 2 (3); McLay, 1993, p. 165 (key); Chan et al., 2009, p. 53, figs. 28, 29. Petalomera granulata var. indica Alcock, 1900a, p. 148. Petalomera indica Alcock, 1901, p. 55, pl. 3, figs. 14, 14a.

Adult: Carapace heptagonal, as broad as long; dorsal surface slightly convex, entirely covered with granules; cervical, urogastric, and branchiocardiac grooves well marked; front tridentate, small median tooth deflexed ventrally, lateral teeth larger than median one, with acute tip, serrated along margin continuous with supraorbital teeth; anterolateral margin with 2 distinct teeth, serrated along margin; posterolateral teeth absent. Subhepatic spine indistinct on dorsal view. Chelipeds large, stout, covered with acute granules; each merus smooth on inner surface, upper margin petaloid; each carpus with 2 acute tubercles on anterodistal margin; each propodus scattering of granules, with distal tubercle on upper surface. Ambulatory legs 1, 2 similar in size, covered with acute granules; leg 3 smallest, covered with acute granules, dactylus opposed by propodal spine; leg 4 smooth, slender, propodus with small spine on outer margin, dactylus opposed by propodal spine. Abdomen of male with 6 somites, scattering of granules on external surface. Uropod plate present. Telson rounded on distal margin. Gonopod 1 stout tube with dense distal setae; gonopod 2 slender, styliform. CL 25.0 mm, CW 24.6 mm (Kim, 1973).

Habitat: Mud; 50-150 m.

Distribution: Korea, Japan, China, Taiwan, Sri Lanka, Andaman Islands.

Korea: JJ.

Specimen examined: None.

Remarks: Lavae are unknown.

19 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Superfamily Calappoidea De Haan, 1833‌ Man-du-ge-sang-gwa (만두게상과)

Carapace subcircular or transversely ovate; posterolateral margin often with clypeiform expansion or lateral margin often with strong spine. Chelipeds massive; each propodus with denticulate crest on upper surface. Ambulatory legs with styliform or paddle-like dactyli. Genital opening of female on thoracic sternum; that of male on coxa of last ambulatory leg. Abdominal somites 3-5 fused in male, not fused in female. Two families (Calappidae De Haan, 1833 and Matutidae De Haan, 1835) exist. In Calappidae posterolateral margin of carapace usually with clypeiform expansion (Pl. 14), ambulatory legs with styliform dactyli; in Matu- tidae clypeiform expansion absent, ambulatory legs with paddle-like dactyli.

Family Calappidae De Haan, 1833‌ Man-du-ge-gwa (만두게과)

Dactylus of larger cheliped with large proximal tooth (Pl. 17) fitting into molariform depression. Merus of maxilliped 3 subquadrate or rarely subcircular. Ambulatory legs not flattened, with styliform dactyli. Gono- pod 1 of male stout; gonopod 2 slender.

Genera 8 (5 in Korea), species over 70 (10 in Korea).

Distribution: Worldwide.

Key to the genera of family Calappidae

1. Carapace with clypeiform expansion on postlateral margin...... Calappa - Carapace without clypeiform expansion on postlateral margin...... 2 2. Carapace with single strong spine on lateral margin...... Mursia - Carapace without strong spine on lateral margin...... Cycloes

Key to the zoeas of family Calappidae

Rostral carapace spine as long as or shorter than CL; dorsal carapace spine ca. CL; lateral carapace spine short. Antenna with long and spinous protopod, exopod short with 2 (1 long, 1 smaller) spines, endopod absent. Endopod of maxillule with 0, 2+4 setae. Endopod of maxilla with 2+5 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 and 2, 2, 0, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 1, 1, 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Fork of telson with 2 outer spines.

20 CALAPPIDAE: Calappa

1. Rostral carapace spine as long as dorsal carapace spine...... Calappa lophos - Rostral carapace spine less than 1/2 length of dorsal carapace spine...... Calappa philargius

Genus Calappa Weber, 1795‌ Man-du-ge-sok (만두게속)

Carapace transversely ovate, broader than long; regions indistinct, only grooves of cardiac region distinct; postlateral margin with clypeiform expansion, partially or completely covering ambulatory legs. Front triangular. Merus of maxilliped 3 subquadrate.

Species 41 (4 in Korea).

7. Calappa clypeata Borradaile, 1903 Top-man-du-ge (톱만두게)

Calappa pustulosa var. clypeata Borradaile, 1903a, p. 436. Calappa terraereginae Ward, 1936, p. 11, pl. 3, figs. 9-11; Kim, 1973, p. 319, fig. 108, pl. 78, fig. 76; T. Sakai, 1976, p. 130, fig. 72a, b; Miyake, 1983, p. 199; Kim and Chang, 1985, p. 44; Chen, 1993, p. 682, fig. 4a. Calappa terrae-reginae: T. Sakai, 1937, p. 92, figs. 6b, 7, pl. 18, fig. 1; Kamita, 1941, p. 32; Dai and Yang, 1991, p. 103, fig. 50 (1), pl. 11 (5). Calappa clypeata Galil, 1997, p. 283, figs. 1d, 4d, 6, 34, 35d.

Adult: Carapace transversely ovate; dorsal surface smooth; margins of front and orbit not denticulate; anterolateral margin with small blunt teeth, clypeiform expansion with marginal blunt teeth. Propodus of each cheliped bearing more and larger tubercles on outer surface than in C. lophos, upper margin crested with 7 blunt teeth. Telson of female much longer than in C. lophos. CL 36 mm, CW 47 mm (Kim, 1973).

Habitat: 6-160 m.

Distribution: Korea, Japan, China, Thailand, Philippines, Indonesia, west Malay Peninsula, Zanzibar, South China Sea, Madagascar, Seychelles, Maldives, Australia, New Caledonia.

Korea: JJ.

Specimen examined: None.

Remarks: This is a rare species from Korean waters, therefore, it has been only reported by T. Sakai (1937)

21 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

in Jejudo Island (JJ). Larvae are unknown.

8. Calappa hepatica (Linnaeus, 1758) (Pls. 12, 13) Gan-man-du-ge (간만두게)

Cancer hepaticus Linnaeus, 1758, p. 630. Calappa tuberculata Weber, 1795, p. 92; Stimpson, 1858a, p. 162; Brocchi, 1875, p. 101, pl. 18, figs. 160, 161; Serène, 1968, p. 41. Calappa (Calappa) hepatica: De Haan, 1833-1849 (1837), p. 70, pl. E. Calappa spinosissima H. Milne-Edwards, 1837, p. 106. Calappa hepatica: White, 1847, p. 44; Borradaile, 1903a, p. 436, pl. 22, fig. 6; Gravely, 1927, p. 142, pl. 22, fig. 27; Ward, 1928, p. 243, pl. 27; T. Sakai, 1936a, p. 44, fig. 7; 1976, p. 128, pl. 38, figs. 1, 3; Ka- mita, 1941, p. 245; Forest and Guinot, 1961, p. 11, figs. 1a, b, 2; Kim, 1973, p. 315, fig. 105, pl. 78, fig. 73; Takeda, 1982, p. 105, fig. 307; Miyake, 1983, p. 20, pl. 7, fig. 6; Dai and Yang, 1991, p. 102, fig. 49 (1), pl. 11 (3); Chen, 1993, p. 677; Galil, 1997, p. 296, figs. 10e, f, 13e, f, 14, 31; Poore, 2004, p. 327, figs. 96a, 97c. Calappa spinosissimus: Stebbing, 1917, p. 19. Calappa hepatica var. spinosissima: Buitendijk, 1939, p. 231.

Adult: Carapace transversely ovate, 1.5 times broader than long; dorsal surface slightly convex, tuber- culated, 2 gastric and 2 cardiac tubercles distinct in midline, longitudinal groove distinct on each side of mesogastric and intestinal regions; anterolateral margin denticulated; clypeiform expansion with 4 triangular teeth, completely covering ambulatory legs; posterior margin smooth, convex medially. Front small, triangular, with minute median incision. Merus of maxilliped 3 subquadrate. Chelipeds massive; each merus smooth on outer surface, with crest on anterodistal margin; each carpus with tubercles on surface; each propodus broadened, subtriangular, with small granules and tubercles on surface, upper margin crested with 8 teeth, lower margin granulate; larger dactylus with large proximal tooth fitting into molariform depression. Ambulatory legs smooth, slightly compressed, with styliform dactyli. Abdomen of female with 6 somites, somite 6 longest. Telson triangular. CL 31.9 mm, CW 46.8 mm.

Habitat: Intertidal-100 m.

Distribution: Korea, Japan, China, Taiwan, Thailand, Philippines, India, Indonesia, South Africa, Mada- gascar, Kenya, Tanzania, Seychelles, Red Sea, Gulf of Aden, Maldives, Sri Lanka, Andaman Islands, Papua New Guinea, Australia, New Zealand, New Caledonia, Fiji, Hawaiian Islands.

Korea: JJ.

22 CALAPPIDAE: Calappa

Specimen examined: None.

Remarks: This is a rare species from Korean waters, therefore, it has been only reported by T. Sakai (1937) in Jejudo Island (JJ). Larvae are unknown.

9. Calappa lophos (Herbst, 1782) (Pls. 14-18) Beom-mu-nui-man-du-ge (범무늬만두게)

Cancer lophos Herbst, 1782, p. 201, pl. 13, fig. 77. Calappa lophos: Fabricius, 1798, p. 346; T. Sakai, 1937, p. 90, fig. 6a, pl. 12, fig. 1; 1976, p. 129, pl. 37, fig. 1, pl. 38, fig. 2; Kamita, 1941, p. 30, figs. 7a, 8; Kim, 1973, p. 318, fig. 107, pl. 16, fig. 75; Takeda, 1982, p. 105, fig. 309; Miyake, 1983, p. 19, pl. 7, fig. 3; Dai and Yang, 1991, p. 103, fig. 49 (2), pl. 11 (4); Chen, 1993, p. 680, fig. 3; Galil, 1997, p. 275 (key), 302, figs. 16, 17a, 20a, 32, 35c; Poore, 2004, p. 330, figs. 96e, 97e. Calappa (Lophos) lophos: De Haan, 1833-1849 (1837), p.72, pl. 20, fig. 1, pl. E. Calappa guerini Brito-Capello, 1871, pp. 128, 133, pl. 2, fig. 2. Calappa quadrimaculata Takeda and Shikatani, 1990, p. 479, figs. 1-4. Calappa lophos: Terada, 1987 (zoea I).

Adult: Carapace transversely ovate, 1.5 times broader than long; dorsal surface with 13 tubercles on gastric region, smooth posteriorly, longitudinal ridges on branchial region, longitudinal groove distinct on each side of mesogastiric and intestinal regions; anterolateral margin slightly denticulated; clypeiform expansion with 4 triangular teeth, partially covering ambulatory legs; posterior margin granulate, with 6 broad lobes, which surfaces with granules. Front triangular, deflexed ventrally. Chelipeds massive; each merus smooth on outer surface, with crest on anterodistal margin; each carpus smooth on surface; each propodus broadened, subtriangular, relatively smooth on outer surface, upper margin crested with 8 teeth, lower surface with row of rough tubercles increasing in size distally, granulate on margin; larger dactylus with large proximal tooth fitting into molariform depression. Ambulatory legs smooth, slightly compressed, with styliform dactyli. Abdominal somites 3-5 fused in male. Telson triangular. Gonopod 1 stout, curved outwards; gonopod 2 slender, filamentose, shorter than gonopod 1. CL 76.5 mm, CW 118.5 mm. Zoea I: Rostral and dorsal carapace spines ca. 1CL; lateral carapace spine short. Antenna with long and spinous protopod, exopod short with 2 (1 long, 1 shorter) spines, endopod absent. Endopod of maxillule with 0, 2+4 setae. Endopod of maxilla with 2+5 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 and 2, 2, 0, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 1, 1, 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Fork of telson with 2 outer spines. CL 0.35 mm, RDL 0.95 mm (Terada, 1987).

23 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Habitat: Sand, mud, gravel; 5-140 m.

Distribution: Korea, Japan, China, Philippines, Thailand, Indonesia, South Africa, Madagascar, Tanzania, Seychelles, Persian Gulf, India, Sri Lanka, Australia, New Caledonia.

Korea: GN, JJ.

Specimen examined: JJ: 1♂(Seoguipo: 6.vii.2013, S.H. Lee).

Remarks: Terada (1987) described only the first zoeal stage of this species.

10. Calappa philargius (Linnaeus, 1758) (Pl. 19) An-geong-man-du-ge (안경만두게)

Cancer philargius Linnaeus, 1758, p. 626. Cancer inconspectus Herbst, 1794, p. 162, pl. 40, fig. 3. Calappa cristata Fabricius, 1798, p. 346; Stimpson, 1858, p. 162; Sokolowsky, 1945, p. 71, pl. 2, figs. 1-3. Calappa inconspecta Bosc, 1801, p. 185. Calappa (Lophos) philargius: De Haan, 1833-1849 (1837), p. 71 (part), pl. 19, fig. 1. Calappa philargius: Herklots, 1861, p. 25; Balss, 1922, p. 122; T. Sakai, 1936a, p. 45, pl. 8, fig. 2; 1976, p. 130, pl. 37, fig. 2; Kamita, 1941, p. 33, fig. 9; Sokolowsky, 1945, p. 72, pl. 2, figs. 4-6; Guinot, 1962, p. 26, figs. 11-15, 17a, b, 18a, b, pl. 1, fig. 1, pl. 2; Kim, 1973, p. 317, fig. 106, pl. 78, fig. 74a, b; Takeda, 1982, p. 106, fig. 310; Miyake, 1983, p. 20, pl. 7, fig. 4; Dai and Yang, 1991, p. 104, fig. 50 (2), pl. 11 (6); Chen, 1993, p. 679, fig. 2; Galil, 1997, p. 307, figs. 17d, 20d, 21, 33; Poore, 2004, p. 330, fig. 97a, f, pl. 18f. Calappa philargius: Terada, 1987 (zoea I).

Adult: Carapace transversely ovate, 1.4 times broader than long; dorsal surface with low tubercles on gastric region, smooth posteriorly; longitudinal ridges on branchial region, longitudinal groove distinct on each side of gastric, cardiac, and intestinal regions; anterolateral margin slightly denticulated; clypeiform expansion with 4 acute triangular teeth, completely covering ambulatory legs; posterior margin with 5 acute teeth, which surfaces with minute granules. Front with 2 triangular teeth. Chelipeds massive; each merus smooth on outer surface, with crest on anterodistal margin; each carpus with low tubercles on outer surface; each propodus broadened, subtriangular, relatively smooth on outer surface, upper margin crested with 7 teeth, lower surface with row of rounded tubercles increasing in size distally, granulate on margin; larger dactylus with large proximal tooth fitting into molariform depression. Ambulatory legs smooth, slightly compressed, with styliform dactyli. Abdominal somites 3-5 fused in male. Telson triangular. Gonopod 1 stout, curved outwards; gonopod 2 slender, filamentose, shorter than gonopod 1. CL 45.5 mm, CW 64.8 mm.

24 CALAPPIDAE: Cycloes

Zoea I: Rostral carapace spine ca. 1/2 CL; dorsal carapace spine ca. 1CL; lateral carapace spine short. Antenna with long and spinous protopod, exopod short with 2 (1 long, 1 shorter) spines, endopod absent. Endopod of maxillule with 0, 2+4 setae. Endopod of maxilla with 2+5 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 and 2, 2, 0, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 1, 1, 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Fork of telson with 2 outer spines. CL 0.37 mm, RDL 0.93 mm (Terada, 1987).

Habitat: Sand, muddy sand; 8-120 m.

Distribution: Korea, Japan, China, South China Sea, Taiwan, Philippines, Thailand, Indonesia, Red Sea, Sri Lanka, New Guinea, Australia, New Caledonia.

Korea: GN.

Specimen examined: GN: 1♂(Busan: 13.ix.2012, S.H. Lee).

Remarks: Terada (1987) described only the first zoeal stage of this species.

Genus Cycloes De Haan, 1837‌ Jeju-man-du-ge-sok (제주만두게속)

Carapace subcircular; regions indistinct; branchial region with 3 longitudinal ridges; lateral margin without strong spine, postlateral margin without clypeiform expansion. Front with 2 triangular teeth. Merus of maxilliped 3 subcircular. Chelipeds massive; each propodus with denticulate crest; larger dactylus with proximal tooth fitting into molariform depression. Ambulatory legs slightly compressed, with styliform dactyli. Gonopod 1 of male stout; gonopod 2 slender, filamentose.

Species 2 (1 in Korea).

11. Cycloes granulosa De Haan, 1837 (Pls. 20-22) Je-ju-man-du-ge (제주만두게)

Cycloës granulosa De Haan, 1837, p. 71, pl. 19, fig. 3, pl. E. Cycloes granulosa: T. Sakai, 1976, p. 139, pl. 43, fig. 3; Miyake, 1983, p. 199; Dai and Yang, 1991, p. 108, fig. 54, pl. 12 (3); Yamaguchi and Baba, 1993, p. 313, fig. 97; Galil and Clark, 1996, p. 194, figs. 9B, 10A-C, 11A, B; Minemizu, 2000, p. 195; Yang et al., 2015, p. 203, figs. 1A-D, 2. Cryptosoma granulosum: Lucas, 1844, p. 438; Miers, 1886, p. 293; T. Sakai, 1936a, p. 49, pl. 7, fig. 2;

25 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Takeda, 1982, p. 109, fig. 319. Not Cryptosoma granulosum: Alcock and Anderson, 1894, p. 198 (list), 203; Borradaile, 1903a, p. 436 (= Cycloes marisrubri Galil and Clark, 1996).

Adult: Carapace subcircular, slightly longer than borad; dorsal surface densely granulate, with longitudinal rows of low tubercles anteriorly; anterolateral margin granulate, lateral margin without strong spine, posterolateral margin minutely granulate; longitudinal groove distinct on each side of gastric, cardiac, and intestinal regions. Front with 2 triangular teeth. Merus of maxilliped 3 subcircular. Chelipeds massive; each merus smooth, with lanceolate tooth distally; each carpus with 3 teeth on upper margin; each propodus broadened, subtriangular, densely granulate on outer surface, upper margin crested with 9 teeth, lower margin with 2 rows of granules; larger dactylus with large proximal tooth fitting into molariform depression. Ambulatory legs smooth, slightly compressed, with styliform dactyli. Abdomen of female with 6 somites. Telson triangular. CL 36.6 mm, CW 35.9 mm.

Habitat: Sand; 5-100 m.

Distribution: Vietnam, Singapore, Taiwan, China, Japan, Korea.

Korea: JJ. Specimen examined: JJ 1♀ [Hanrim: 28.x.2007, K.C. Yang). Remarks: Larvae are unknown.

Genus Mursia Desmarest, 1823‌ Geum-ge-bu-chi-sok (금게붙이속)

Carapace subcircular; dorsal surface covered with granules and tubercles; lateral margin with strong spine, postlateral margin without clypeiform expansion. Front with 3 lobes. Merus of maxilliped 3 subquadrate. Chelipeds massive; each propodus with denticulate crest; larger dactylus with proximal tooth fitting into cup-like depression. Ambulatory legs compressed, with long styliform dactyli. Gonopod 1 of male stout; gonopod 2 slender, filamentose.

Species 25 (2 in Korea).

26 CALAPPIDAE: Mursia

12. Mursia armata De Haan, 1837 Geum-ge-but-i (금게붙이)

Mursia armata De Haan, 1833-1849 (1837), p. 73, pl. 19, fig. 2, pl. E; T. Sakai, 1937, p. 85, pl. 11, fig. 3; 1976, p. 135, pl. 43, fig. 2; Kim, 1973, p. 311, fig. 103, pl. 15, fig. 71; Miyake, 1983, pp. 23, 199, pl. 8, fig. 3; Dai and Yang, 1991, p. 107, fig. 53, pl. 12 (2); Chen, 1993, p. 689, fig. 9; Galil, 1993, p. 352 (part), figs. 1b, 2c, d, 3c, d. Thealia acanthophora Lucas, 1839, p. 579, pl. 21, figs. 1-3. Mursia armata typica Doflein, 1904, p. 40, pl. 17, fig. 1, pl. 18, fig. 2; T. Sakai, 1936a, p. 47, pl. 7, fig. 1. Not Mursia armata: Barnard, 1950, p. 356, fig. 67g; Kensley, 1981, p. 38 (= Mursia africana Galil, 1993).

Adult: Carapace subcircular, broader than long; dorsal surface covered with minute granules, 5 rows of longitudinal tubercles present; longitudinal groove distinct on each side of gastric, cardiac, and intestinal regions; anterolateral margin with 7-10 larger granules, lateral margin with strong medial spine of 1/2 CL, posterolateral margin minutely granulate, posterior margin with 2 leaf-like tubercles. Front narrow, with 3 small lobes, of which median lobe slightly protruding. Chelipeds massive; each merus prismatic, outer margin with 2 teeth (1 long distal, 1 shorter subdistal); each propodus broadened, upper margin crested with 8 teeth, dorsal surface with 12 tubercles forming 3 transverse rows, lower margin with minute teeth; movable finger much slender than fixed one. Ambulatory legs long, slender; each carpus with 1 acute distal spine; each dactylus slightly curved, much longer than propodus, with styliform tip. Male with abdominal somites 3-5 fused, female with 6 somites. Telson triangular. Gonopod 1 of male stout, curved outwards; gonopod 2 long, slender, with loop-shaped tip. Male CL 36 mm, CW 47 mm (Kim, 1973).

Habitat: Mud or muddy sand; 50-60 m.

Distribution: Vietnam, Philippines, South China Sea, China, Japan, Korea.

Korea: JJ.

Specimen examined: None.

Remarks: Larvae are unknown.

13. Mursia trispinosa Parisi, 1914 (Pl. 23) Sae-ga-si-geum-ge-but-i (세가시금게붙이)

Mursia armata trispinosa Parisi, 1914, p. 290, pl. 12. Mursia armata curtispina: Balss, 1922, p. 124 (part); T. Sakai, 1936a, p. 48, pl. 7, fig. 3. Mursia curtispina trispinosa: T. Sakai, 1965, p. 53, figs. 8b, b’, pl. 21, fig. 1; Kim, 1973, p. 313, fig. 104, pl.

27 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

15, fig. 72. Mursia trispinosa: T. Sakai, 1976, p. 137, fig. 74b, b’, pl. 43, fig. 4; Miyake, 1983, p. 24, pl. 8, fig. 6; Chen, 1993, p. 690, fig. 10; Galil, 1993, p, 370, figs. 7c, 9e, f, 10h, I.

Adult: Carapace subcircular, broader than long; dorsal surface covered with minute granules and rows of longitudinal tubercles; longitudinal groove distinct on each side of gastric, cardiac, and intestinal regions; anterolateral margin with 9 larger granules, lateral margin with strong medial spine, which length (12.9 mm) less than 1/3 CL, posterior margin with 3 (2 acute lateral, 1 small medial) tubercles. Front narrow, with 3 small lobes, of which median lobe protruding. Chelipeds massive; each merus prismatic, outer margin with long distal and 2 shorter subdistal teeth; each carpus scattering of tubercles on dorsal surface; each propodus broadened, upper margin crested with 8 teeth, dorsal surface scattering of 9 tubercles, lower surface with 3 acute tubercles. Ambulatory legs broader than those of M. armata; each carpus with acute distal spine. Ab- dominal somites 3-5 fused in male. Telson triangular. Gonopod 1 stout, curved outwards; gonopod 2 long, slender, with hook-shaped tip. CL 45.3 mm, CW 53.9 mm.

Distribution: Philippines, South China Sea, Japan, New Caledonia, Korea.

Habitat: Mud or muddy sand; 50-70 m.

Korea: JJ.

Specimen examined: JJ: 1♂(33°48ʹN, 128°13ʹE: 20.viii.2012, J.H. Choi). Remarks: Larvae are unknown. The male specimen is deposited at National Marine Biodiversity Institute of Korea.

Family Matutidae De Haan, 1835‌ Geum-ge-gwa (금게과)

Carapace subcircular; regions indistinct; lateral margin with strong spine; posterolateral margin sharply convergent. Merus of maxilliped 3 subtriangular (Pl. 25). Chelipeds massive; each propodus with 3 teeth on upper surface, ridges on outer median surface; cutting margin of dactylus irregularly dentate. Ambulatory legs flattened, with paddle-like dactyli. Abdominal somites 3-5 fused in male; somites 2, 3 carinate. Gono- pod 1 stout, straight; gonopod 2 slender, filamentose.

Genera 3 (2 in Korea), species over 10 (3 in Korea).

Distribution: From east African coast to Indo-West Pacific.

28 MATUTIDAE: Ashtoret

Key to the genera of family Matutidae

1. Propodus of cheliped with parallel ridge on outer median surface, dactylus of male smooth on outer distal‌ ...... Ashtoret - Propodus‌ of cheliped with oblique ridge on outer median surface, dactylus of male with strong milled ridge on outer distal surface...... Matuta

Key to the zoeas of family Matutidae

Rostral and dorsal carapace spines shorter than CL; lateral carapace spine absent. Antennal protopod long, spinous, exopod absent. Endopod of maxillule with 1, 3 (4) setae. Endopod of maxilla with 2+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 and 3, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1 +1+1+1 and 1, 1, 5 (6) setae, respectively. Lateral processes on abdominal somites 2, 3; posterolateral process on abdominal somite 4 long, with blunt tip. Fork of telson with 3 outer spines.

1. Endopod of maxillule with 1, 3 setae, endopod of maxilliped 2 with 1, 1, 5 setae...... Matuta planipes - Endopod of maxillule with 1, 4 setae, endopod of maxilliped 2 with 1, 1, 6 setae...... 2 2. Dorsal carapace spine slightly curved...... Ashtoret lunaris - Dorsal carapace spine straight...... Matuta victor

Genus Ashtoret Galil and Clark, 1994‌ Geum-ge-sok (금게속)

Propodus of cheliped with parallel ridge on outer median margin; dactylus of male smooth on outer distal surface (Pl. 25). Carpus of ambulatory leg 3 bicarinate on upper surface. Abdominal somite 2 with transversely tuberculate carina.

Species 8 (1 in Korea).

14. Ashtoret lunaris (Forskål, 1775) (Pls. 24, 25) Geum-ge (금게)

Cancer lunaris Forskål, 1775, p. 91 (part). Matuta banksii Leach, 1817, p. 14; Barnard, 1950, p. 359, fig. 67h-k; Miyake, 1983, p. 200 (list); Dai and

29 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Yang, 1991, p. 111, fig. 56 (1), pl. 12 (6). Matuta victor (nec Fabricius, 1781) Desmarest, 1825, p. 101, pl. 7 (2). Matuta lessueri Rüppell, 1830, p. 7 (part). Matuta lunaris: White, 1847, 46 (part); Yamaguchi and Baba, 1993, p. 317, fig. 100. Matuta banksi: Ortmann, 1892, p. 573; T. Sakai, 1976, p. 141 (part), pl. 44, fig. 3; Takeda, 1982, p. 110, fig. 322. Ashtoret lunaris: Galil and Clark, 1994, p. 5, fig. 1a, b, pl. 1a, b; Poore, 2004, p. 345, fig. 102e. Not Matuta lunaris: Miers, 1877, p. 247, pl. 40, figs. 10, 11; Henderson, 1887, p. 66, fig. 6; Kim, 1973, p. 321, fig. 110, pl. 79, fig. 77 [= Matuta victor (Fabricius, 1781)]. Matuta lunaris: Terada, 1987 (zoeal stages).

Adult: Carapace subcircular; mesogastric, cardiac and branchial regions minutely granulate, each region with 1 indistinct tubercle; anterolateral margin with 5-7 small and 3 larger tubercles; lateral margin with 1 strong medial spine of 1/6 CW; posterolateral margin convergent, minutely granulate; posterior margin narrow, straight. Front with 3 (2 truncate lateral, 1 protruding medial) lobes, of which medial lobe with V-shaped notch. Chelipeds massive; each carpus with 3 low tubercles on outer surface; each propodus with 3 teeth on upper margin decreasing in size distally, of which proximal tooth tuberculate, upper outer distal surface with 2 rows of granulate tubercles, outer median surface with sharp spine proximally and parallel ridge extending to base of fixed finger, lower proximal angle with acute spine, lower margin with row of tubercles; each dactylus smooth on outer surface. Ambulatory legs 1-3 each with triangular tooth on posterior margin of propodus, propodus of leg 4 extended posteriorly; carpus of leg 3 with bicarinate on upper surface. Abdominal somites 3-5 of male fused; somite 2 with transversely tuberculate carina, somite 3 with transversely low tuberculate carina. Gonopod 1 stout, straight; gonopod 2 slender, filamentose. Telson subtriangular. Carapace covered with red spots and rings. CL 23.9 mm, CW 25.1 mm. Zoea I: Rostral carapace spine shorter than CL; dorsal carapace spine slightly curved, shorter than CL; lateral carapace spine absent. Antenna with spinous protopod, exopod absent. Endopod of maxillule with 1, 4 setae. Endopod of maxilla with 2+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 and 3, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 1, 1, 6 setae, respectively. Lateral processes on abdominal somites 2, 3; posterolateral process on abdominal somite 4 long, with blunt tip. Fork of telson with 3 outer spines. CL 0.32 mm, RDL 0.66 mm (Terada, 1987).

Distribution: South Africa, Mozambique, Red Sea, Indian Ocean, Andaman, Malaysia, Singapore, Bor- neo, Palau, Indonesia, Philippines, Taiwan, China, Japan, Australia, New Caledonia, Korea.

Habitat: Sand; shallow waters.

Korea: GB, GN, JN, JJ.

30 MATUTIDAE: Matuta

Specimen examined: GN: 1♂(Busan: 27.x.2012, S.H. Lee).

Remarks: This species has five zoeal stages (Terada, 1987).

Genus Matuta Weber, 1795‌ Gue-mul-mu-nei-geum-ge-sok (그물무늬금게속)

Propodus of cheliped with oblique ridge on outer median surface, dactylus of male with strong milled ridge on outer surface (Pl. 28). Carpus of ambulatory leg 3 unicarinate on upper surface. Abdominal somite 3 with transversely tuberculate carina.

Species 4 (2 in Korea).

15. Matuta planipes Fabricius, 1798 (Pl. 26) Gue-mul-mu-nei-geum-ge (그물무늬금게)

Matuta planipes Fabricius, 1798, p. 369; Balss, 1922, p. 125; T. Sakai, 1936a, p. 50, pl. 8, fig. 1; 1976, p. 141, pl. 44, fig. 2; Kamita, 1941, p. 36, fig. 12; Chhapgar, 1957, p. 7, pl. A, fig. 4, pl. 2d-f; Romimohtarto, 1972, p. 11, figs. 1-3, 5, 8, 15-20, pls. 1a, 3a; Kim, 1973, p. 323, fig. 111, pl. 16, fig. 78; Takeda, 1982, p. 109, fig. 321; Dai and Yang, 1991, p. 109, fig. 55 (1), pl. 12 (4); Tirmizi and Kazmi, 1991, p. 64, fig. 18E- H; Galil and Clark, 1994, p. 35, fig. 6c, d, pl. 12a, b; K. Sakai, 1999, p. 21, pl. 9B; Poore, 2004, p. 346, fig. 102d. Cancer lunaris (not Rumphius, 1741): Herbst, 1799, p. 43, pl. 48, fig. 6. Matuta victor (not Fabricius, 1781): Latreille, 1818, p. 273, figs. 3, 4. Matuta appendiculata Bosc, 1830, p. 222. Matuta victor (not Fabricius, 1781): De Haan, 1833-1849 (1841), p. 127 (part, var. IV). Matuta lunaris (non Rumphius, 1741): White, 1847, p. 46 (part); Miers, 1877, p. 247, pl. 40, figs. 10, 11; Henderson, 1887, p. 66, fig. 6. Matuta rubrolineata Miers, 1877b, p. 244, pl. 39, figs. 5, 6. Matuta lineifera Miers, 1877, p. 245, pl. 39, fig. 7. Matuta laevidactyla Miers, 1880, p. 316; De Man, 1898, p. 699 (list), pl. 32, fig. 45. Matuta flagra Shen, 1936, p. 64, fig. 1. Matuta planipes: Hashmi, 1969 (zoea I).

31 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Adult: Carapace subcircular; mesogastric and branchial regions minutely granulate; protogastric, mesogastric, cardiac, and branchial regions each with 1 distinct tubercle; anterolateral margin with 11-12 tubercles; lateral margin with 1 strong medial spine of 1/7 CW; posterolateral margin convergent, minutely granulate; posterior margin narrow, straight. Front with 3 (2 truncate lateral, 1 protruding medial) lobes, of which medial lobe with V-shaped notch. Chelipeds massive; each carpus with 3 low tubercles on outer surface; each propodus with 3 teeth on upper margin decreasing in size distally, of which proximal tooth tuberculate, outer distal surface with 2 rows of tubercles, outer median surface with sharp spine proximally and oblique ridge extending to tip of fixed finger, lower proximal angle with tubercle, lower margin with row of tubercles; each dactylus with strong milled ridge on outer distal surface. Ambulatory legs 1-3 each with triangular tooth on posterior margin of propodus, propodus of leg 4 extended posteriorly; carpus of leg 3 unicarinate on upper surface. Abdominal somites 3-5 of male fused; somite 2 with transversely low carina, somite 3 with transversely tuberculate carina. Gonopod 1 stout, straight; gonopod 2 slender, filamentose, with submedial sharp tubercle, distal half more slender than proximal half. Telson subtriangular. Carapace covered with reticulating brown lines. CL 30.4 mm, CW 31.9 mm. Zoea I: Rostral and dorsal carapace spines slightly shorter than1 CL; lateral carapace spine absent. Anten- na with spinous protopod, exopod absent. Endopod of maxillule with 1, 3 setae. Endopod of maxilla with 2+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 and 3, 2, 1, 2, 5 setae, respectively. Basis and endopod maxilliped 2 each with 1+1+1+1 and 1, 1, 5 setae, respectively. Lateral processes on abdominal somites 2, 3; posterolateral process on abdominal somite 4 long, with blunt tip. Fork of telson with 3 outer spines. RDL 0.76 mm (Hashmi, 1969).

Habitat: Sand; shallow waters.

Distribution: Persian Gulf, Indian Ocean, Pakistan, India, Sri Lanka, Gulf of Thailand, Vietnam, Malaysia, Singapore, Indonesia, Taiwan, South China Sea, China, Japan, New Guinea, Australia, Korea.

Korea: GN, JN, JJ, JB, CN, GG.

Specimen examined: JB: 1♂(Gochang: 11.x.2011, S. H. Lee).

Remarks: Hashmi (1969) described only the first zoeal stage of this species.

16. Matuta victor (Fabricius, 1781) (Pls. 27-31) Jan-jeom-mu-nui-geum-ge (잔점무늬금게)

Cancer victor Fabricius, 1781, p. 502. Cancer lunaris: Herbst, 1783, p. 140, pl. 6, fig. 44. Matuta victor: Fabricius, 1798, p. 369; H. Milne-Edwards, 1837, p. 115, pl. 20, figs. 3-6; Hilgendorf, 1869, p. 93, pl. 3, fig. 2; Hoffmann, 1874, p. 27, pl. 6, figs. 45-48; Gravely, 1927, p. 142, pl. 22, fig. 28; Galil and

32 MATUTIDAE: Matuta

Clark, 1994, p. 39, fig. 7a, b, pl. 13a, b; Ng, 1998, p. 1095, fig. 10; K. Sakai, 1999, p. 21, pl. 9C. Matuta peronii Leach, 1817, p. 13, pl. 127, figs. 1, 2. Matuta lesuerii Leach, 1817, p. 14. Matuta victrix Miers, 1877, p. 243, pl. 39, figs. 1-3. Matuta lunaris: Rathbun, 1902, p. 30; T. Sakai, 1936a, p. 49, pl. 8, fig. 3; Kim, 1973, p. 321, fig. 110, pl. 79, fig. 77; Takeda, 1982, p. 109, fig. 320; Miyake, 1983, p. 24, pl. 8, fig. 7; Dai and Yang, 1991, p. 110, fig. 55 (2), pl. 12 (5). Matuta victrix var. crebrepunctata Miers, 1877, p. 244, pl. 39, fig. 4. Matuta crebrepunctata Ortmann, 1892, p. 572; Ward, 1941, p. 1.

Adult: Carapace subcircular; mesogastic, cardiac and branchial regions minutely granulate; protogastric, mesogastric, cardiac, and branchial regions each with 1 tubercle; anterolateral margin with 5 small and 3 larger tubercles; lateral margin with 1 strong medial spine of 1/5 CW; posterolateral margin convergent, minutely granulate; posterior margin narrow, straight. Front with 3 (2 rounded lateral, 1 protruding medial) lobes, of which medial lobe with V-shaped notch. Chelipeds massive; each carpus with 3 tubercles on outer surface; each propodus with 3 teeth on upper margin decreasing in size distally, of which proximal tooth tuberculate, outer distal surface with 2 rows of tubercles, outer median surface with sharp spine proximally and oblique ridge extending to tip of fixed finger, lower proximal angle with acute spine, lower margin with row of tubercles; each dactylus with strong milled ridge on outer distal surface. Ambulatory legs 1-3 each with triangular tooth on posterior margin of propodus, propodus of leg 4 extended posteriorly; carpus of leg 3 unicarinate on upper surface. Abdominal somites 3-5 of male fused; somite 2 with transversely low carina, somite 3 with transversely tuberculate carina. Gonopod 1 stout, almost straight; gonopod 2 slender, filamentose, with submedial sharp tubercle, distal half more slender than proximal half. Telson subtriangular. Carapace entirely covered with minute red dots. CL 43.8 mm, CW 46.2 mm. Zoea I: Rostral carapace spine shorter than 1 CL; dorsal carapace spine straight, shorter than 1 CL; lateral carapace spine absent. Antenna with spinous protopod, exopod absent. Endopod of maxillule with 1, 4 setae. Endopod of maxilla with 2+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 and 3, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 1, 1, 6 setae, respectively. Lateral processes on abdominal somites 2, 3; posterolateral process on abdominal somite 4 long, with blunt tip, that on somite 5 shorter, with blunt tip. Fork of telson with 3 outer spines. CL 0.45 mm, RDL 0.85 mm.

Habitat: Sand; shallow waters.

Distribution: South Madagascar, Mozambique, Tanzania, Somalia, Red Sea, Indian Ocean, Pakistan, In- dia, Sri Lanka, Andaman Islands, Vietnam, Singapore, Indonesia, Taiwan, China, Japan, Australia, Palau, New Caledonia, Korea.

33 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Korea: GN, JJ.

Specimen examined: GN: 1♂(Busan: 11.x.2013, S.H. Lee); JJ: zoeas (Hamdeok: 21.viii.2016, H.S. Ko). Remarks: The first zoeal stage of this species is reported for the first time in the volume. It shows similar- lity more to Ashtoret lunaris than to Matuta planipes in the characteristics of the mouthpart appendages.

Superfamily Orithyoidea Dana, 1852‌ Beom-ge-sang-gwa (범게상과)

Carapace subcircular. Merus of maxilliped 3 subtriangular, narrowing anteriorly. Chelipeds massive; each propodus without crest on upper surface. Only ambulatory leg 4 with paddle-like dactylus. Genital opening of female on thoracic sternum, not covered by abdomen; that of male on coxa of last ambulatory leg. Ab- dominal somites not fused in both sexes. Only one family, Orithyiidae Dana, 1852 exists.

Family Orithyiidae Dana, 1852‌ Beom-ge-gwa (범게과)

Carapace longer than broad; dorsal surface granulate. Chelipeds short, unequal. Ambulatory legs 1-3 with styliform dactyli, leg 4 with paddle-like dactylus. Abdomen with 6 somites in both sexes. Gonopod 1 of male broadened proximally, stout distally.

Genus 1 (1 in Korea), species 1 (1 in Korea).

Distribution: China, Korea.

Genus Orithyia Fabricius, 1798‌ Beom-ge-sok (범게속)

Carapace with 2 spines on lateral margin; dorsal surface granulate with tubercles. Right cheliped larger than left one.

34 ORITHYIIDAE: Orithyia

17. Orithyia sinica (Linnaeus, 1771) (Pls. 32-35) Beom-ge (범게)

Cancer sinicus Linnaeus, 1771, p. 541. Orithyia mammilarlis Fabricius, 1798, p. 363. Orithyia sinica: Urita, 1926, p. 436; Kamita, 1941, p. 38, pl. 2; Kim, 1973, p. 325, pl. 17, fig. 79; T. Sakai, 1976, p. 143, fig. 75; Takeda, 1982, p. 110, fig. 324; Dai and Yang, 1991, p. 113, pl. 12 (8), fig. 57. Orithyia sinica: Hong, 1976 (zoeal stages).

Adult: Carapace slightly longer than broad; dorsal surface convex, covered with granules; protogastric, hepatic, cardiac, and intestinal regions each with tubercle, mesogastric and branchial regions each with 2 tubercles; margins granulate, anterolateral margin with 2 tubercles, lateral margin with large medial spine; posterolateral margin with 2 smaller spines. Front with 3 spines, of which medial one protruding. Orbit margin with 3 (2 small inner, 1 larger outer) spines. Maxilliped 3 with short setae on margin; ischium with longitudinal groove on surface; merus subtriangular, 1.3 times longer than ischium. Chelipeds short, stout, as long as CL; unequal, right one larger than left; each merus prismatic, upper margin with distal spine, lower margin with 2 (1 medial, 1 distal) spines; each carpus with numerous setae on posterior margin, with 2 (1 small subdistal, 1 larger distal) spines on upper margin, smallest spine on outer distal margin; each propodus convex, granulate on outer surface, upper margin with 3 spines; fingers longer than propodus. Ambulatory legs stout; each carpus and propodus of legs 3, 4 carinate on upper margin; dactyli of legs 1-3 styliform, in leg 4 paddle-like. Abdomen of male relatively broad, with 6 somites; somite 1 with small spine; somites 3 with 3 tubercles transversely. Telson tongue-shaped. Gonopod 1 broadened in proximal part, numerous setae on outer distal margin; distal part stout, slightly curved outerwards; tip rounded. In fresh specimen, carapace with large black spot on each side of branchial region; ambulatory legs with black bands. CL 76.4 mm, CW 65.4 mm. Zoea I: Carapace with extremely long rostral and dorsal spines; rostral spine ca. 5.5 CL, dorsal spine ca. 3.5 CL, lateral spine short. Protopod of antenna with spinous tip; exopod with bifid tip, slightly longer than protopod; endopod present. Endopod of maxillule with 1, 1+4 setae. Endopod of maxilla with 3+4 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 (10) setae and 2, 2, 1, 2, 5 setae, respectively. Ba- sis and endopod of maxilliped 2 each with 1+1+1 (3) setae and 0, 1, 4 setae, respectively. Lateral processes on abdominal somite 2. Telson slender; fork long, with 2 small outer spines. CL 0.86 mm, RDL 8.64 mm.

Habitat: Sandy mud; shallow waters.

Distribution: China, Korea.

Korea: GG, CN, JB.

35 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Specimen examined: JN: 1♂(Gochang: 27.iii.2010, S.H. Lee); zoeas (Gochang: 9.x.2010, S.H. Lee).

Remarks: This species has three zoeal stages (Hong, 1976).

Superfamily Leucosioidea Samouelle, 1819‌ Bam-ge-sang-gwa (밤게상과)

Carapace variable in shape. Front narrow. Buccal frame narrowing anteriorly. Merus of maxilliped 3 triangular, narrowing anteriorly (Pl. 45). Branchial channel running along side of buccal cavity, covered with maxilliped 3. Genital opening of female on thoracic sternum; that of male on coxa of last ambulatory leg or thoracic sternum. Two families (Leucosiidae Samouelle, 1819 and Iphiculidae Alcock, 1896) and more than 480 species exist. In Leucosiidae some abdominal somites of both sexes fused, in Iphiculidae all abdominal somites not fused (Ng et al., 2008). Only one family Leucosiidae known in Korea.

Family Leucosiidae Samouelle, 1819‌ Bam-ge-gwa (밤게과)

Carapace subcircular, subtriangular, subpentagonal or rhomboidal; dorsal surface smooth, granular, sometimes with spines. Chelipeds short, or sometimes long, slender. Ambulatory legs with styliform dactyli. Sev- eral abdominal somites fused in both sexes. Gonopod 1 of male straight (Pl. 46), contorted or coiled (Pl. 64). Two subfamilies (Ebaliinae and Leucosiinae) known in the country.

Genera 62 (11 in Korea), species over 470 (11 in Korea).

Distribution: Worldwide.

Key to the genera of family Leucosiidae

1. Gonopod 1 of male straight...... 2 (Subfamily Ebaliinae) - Gonopod 1 of male contorted or coiled...... 9 (Subfamily Leucosiinae) 2. Carapace subcircular...... 3 - Carapace not subcircular...... 8 3. Carapace with spine...... 4 - Carapace without spine...... 5 4. Carapace only with posterior spines...... Myra - Carapace with marginal or dorsal spines...... Arcania

36 LEUCOSIIDAE

5. Abdominal surface with subdistal tubercle in male...... 6 - Abdominal surface without subdistal tubercle in male...... 7 6. Apical process of gonopod 1 short...... Lyphira - Apical process of gonopod 1 long or absent...... Philyra 7. Apical process of gonopod 1 absent...... Hiplyra - Apical process of gonopod 1 present...... Pyrhila 8. Carapace subtriangular...... Merocryptus - Carapace pentagonal...... Nursia 9. Ventral margin of thoracic sinus without granules...... Urnalana - Ventral margin of thoracic sinus with granules...... 10 10. Thoracic sinus with anterior loop-shaped margin...... Euclosia - Thoracic sinus without anterior loop-shaped margin...... Leucosia

Key to the zoeas of family Leucosiidae

Rostral carapace spine present. Antenna uniramous process, without endopod and exopod. Endopods of maxillule and maxilla each with 2+2 and 2+1 (rarely 2+2) setae on distal segment, respectively. Basis and endopod of maxilliped 1 each with 2+2+2+2 and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1 +1+1+1 and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate (Pl. 58), with 3 or 4 (rarely 1) outer spines, fork absent.

1. Dorsal carapace spine present...... 2 - Dorsal carapace spine absent...... 7 2. Lateral carapace spine present...... 3 - Lateral carapace spine absent...... Philyra syndactyla 3. Endopod of maxilla with 3 setae...... 4 - Endopod of maxilla with 4 setae...... 5 4. Lateral carapace spine with blunt tip...... Leucosia anatum - Lateral carapace spine with spinous tip...... Euclosia obtusifrons 5. Dorsal carapace spine longer than 1.5 CL...... Arcania undecimspinosa - Dorsal carapace spine shorter than 1.5 CL...... 6 6. Dorsal carapace spine ca. 1.2 CL...... Nursia rhomboidalis - Dorsal carapace spine ca. 1 CL...... Myra celeris 7. Antenna with rounded tip...... Hiplyra platycheir - Antenna with spinous tip...... 8 8. Base of dorsal carapace spine with chromatophor...... Pyrhila carinata, P. pisum - Base of dorsal carapace spine without chromatophore...... Philyra kanekoi

37 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Subfamily Ebaliinae Stimpson, 1871‌ Bam-ge-a-gwa (밤게아과)

Front of carapace not strongly produced. Propodi of chelipeds cylindrical, relatively swollen or slightly flattened. Gonopod 1 of male straight.

Genera 52 (8 in Korea), species over 370 (10 in Korea).

Distribution: Worldwide.

Genus Arcania Leach, 1817‌ Yeol-han-ga-si-bam-ge-sok (열한가시밤게속)

Carapace subcircular, longer than broad; regions indistinct; dorsal surface covered with granules or spines, marginal spines present. Front divided into 2 lobes. Chelipeds long, slender; propodi cylindrical; fingers long. Gonopod 1 of male long, straight.

Species 20 (2 in Korea).

18. Arcania globata Stimpson, 1858 (Pls. 36, 37) Ga-si-tu-seong-bam-ge (가시투성밤게)

Arcania globata Stimpson, 1858a, p. 160; Miers, 1879, p. 44; Yokoya, 1933, p. 134; Kim, 1973, p. 297, fig. 92, pl. 76, fig. 61; T. Sakai, 1976, p. 92, fig. 48; Galil, 2001, pp. 182-184, figs. 2B, 5C.

Adult: Carapace longer than broad; dorsal surface covered with small curved spines; margins with 10 larger spines; intestinal region with larger spine. Front produced, bilobed; each lobe covered with granules, tip acute. Ischium of maxilliped 3 twice longer than merus measured along inner margin. Chelipeds relatively stout; each merus covered with acute granules, 3 small spines on posterior margin; each carpus and propodus covered with minute granules; fingers as long as propodus, cutting edges denticulate. Ambulatory legs slender, each merus spinulate on anterior margin, each dactylus slightly shorter than propodus. Abdom- inal somites 1, 2 of female articulated, somites 3-6 fused, surface granulate; telson tongue-shaped. CL 12.1 mm, CW 11.0 mm.

Habitat: Muddy sand or sand; 30-100 m.

38 LEUCOSIIDAE: Arcania

Distribution: China, Japan, Korea.

Korea: JN.

Specimen examined: JN: 1♀(Jindo: 27.v.2015, S.H. Lee). Remarks: This species is rare. Larvae are unknown.

19. Arcania undecimspinosa De Haan, 1841 (Pls. 38-40) Yeol-han-ga-si-bam-ge (열한가시밤게)

Arcania undecimspinosa De Haan, 1833-1849 (1841), p. 135, pl. 33, fig. 8; Bell, 1855, p. 309; Kamita, 1941, p. 40, fig. 13; Kim, 1973, p. 295, fig. 91, pl. 11, fig. 60; T. Sakai, 1976, p. 91, pl. 28, fig. 1; Miyake, 1983, p. 60, pl. 20, fig. 6; Dai and Yang, 1991, p. 73, fig. 33 (1), pl. 8 (1); Galil, 2001a, p. 197 (part), figs. 3D, 7D (part); Poore, 2004, p. 340, fig. 99; Naruse, 2014, p. 302, figs. 1-6. Arcania undecimspinosa: Terada, 1984, p. 157, fig. 2A’-H’ (zoea I).

Adult: Carapace subcircular; dorsal surface covered with granules; regions indistinct, only branchiocardiac groove distinct; margins with total of 11 spines, of which 2 subhepatic and 2 anterolateral spines smallest. Front with 2 triangular lobes. In maxilliped 3 ischium 2.5 times longer than merus measured along inner margin. Chelipeds long, slender, ca. 2CL; each merus cylindrical, covered with granules; each carpus and propodus sparsely covered with minute granules; fingers slightly longer than propodus, denticulate on cutting margins. Ambulatory legs long, slender, covered with minute granules; each merus longer than other segments. Abdominal somites 3-5 of male fused; somite 6 long, lateral margin slightly convex. Telson subtriangular, twice longer than broad. Gonopod 1 long, straight; apical process slender. CL 23.4 mm, CW 20.8 mm. Zoea I: Carapace with long spines; rostral spine ca. 1.4 CL, lateral spine ca. 1.2 CL, dorsal spine ca. 1.6 CL. Protopod of antenna knob-like, without spinules. Endopod of maxillule with 4 setae. Endopod of maxilla with 2+2 (4) setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 1 outer spine; fork absent. CL 0.51 mm, RDL 1.80 mm.

Habitat: Muddy sand or sand; 20-30 m.

Distribution: China, Taiwan, Japan, Korea.

Korea: GB, GN, JN, CN.

Specimen examined: GN 1♂(Busan: 13.ix.2012, S.H. Lee); JN: zoeas (Yeosu: 29.vii.2012, S.H. Lee).

Remarks: Terada (1984) described only the first zoeal stage of this species.

39 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Genus Hiplyra Galil, 2009 Jeep-ge-teol-bam-ge-sok (집게털밤게속)

Carapace subcircular, longer than broad, without spine; regions indistinct; dorsal surface punctuate or granulate. Front slightly produced. Chelipeds long, flattened; each merus long in male; fingers with dense setae on cutting margins. Only abdominal somite 1 of male articulate; somites 2-6 without denticles. Gono- pod 1 of male long, straight; apical process absent.

Species 6 (1 in Korea).

20. Hiplyra platycheir De Haan, 1841 (Pls. 41, 42) Jeep-ge-teol-bam-ge (집게털밤게)

Philyra platycheira De Haan, 1841, p. 132, pl. 33, fig. 6; T. Sakai, 1976, p. 109, fig. 59a, pl. 32, fig. 4; Take- da, 1982, p. 100, fig. 294; Miyake, 1983, p. 65, pl. 22, fig. 4; Dai and Yang, 1991, p. 84, fig. 38 (1, 2), pl. 9 (5); Poore, 2004, p. 343, fig. 100d; Lee and Ko, 2007, pp. 1-4, figs. 1, 2. Hiplyra platycheir: Galil, 2009, p. 295, figs. 2F, 10. Philyra platychira: Ko, 2000 (larval stages).

Adult: Carapace subcircular, slightly longer than broad; dorsal surface sparsely covered with granules; regions indistinct, only branchiocardiac groove distinct; margins lined with granules. Front deflexed, with triangular tip medially. In maxilliped 3 ischium 1.2 times longer than merus measured along inner margin. Chelipeds long, flattened, ca. 2.2 CL; each merus covered with granules proximally; each carpus smooth; each propodus longer than fingers; fingers with dense setae on cutting margins. Ambulatory legs slender, smooth; each propodus longer than other segments. Abdominal somite 1 of male narrow, articulate; somites 2-6 fused, somites 4-6 smooth on surfaces. Telson triangular, twice longer than broad. Gonopod 1 long, straight; tip setose, apical process absent. CL 19.2 mm, CW 17.2 mm. Zoea I: Carapace globose, only with short rostral spine. Protopod of antenna short, tip rounded. Endo- pod of maxillule with 0, 4 setae. Endopod of maxilla with 3 setae. Basis and endopod of maxilliped 1 each with 2+2 +2 +2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1 +1 +1 +1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 3 outer spines; fork absent. CL 0.42 mm.

Habitat: Muddy sand or sand; 7-30 m.

Distribution: China, Taiwan, Japan, Korea.

40 LEUCOSIIDAE: Lyphira

Korea: GN, JJ.

Specimen examined: GN: 1♂(Busan: 25.vi.2009, H.S. Ko); JJ: zoeas (Mosulpo: 25.viii.1996, H.S. Ko).

Remarks: This species has three zoeal stages (Ko, 2000).

Genus Lyphira Galil, 2009‌ Ma-leum-mo-bam-ge-sok (마름모밤게속)

Carapace subcircular, longer than broad, without spine; regions indistinct; dorsal surface smooth, granular. Front slightly produced. Each merus of chelipeds short, subcylindrical; propodus swollen. Only abdominal somite 1 of male articulate; somites 2-6 heavily granulate, surface with subdistal tubercle. Gonopod 1 of male long, straight; apical process short.

Species 4 (1 in Korea).

21. Lyphira heterograna (Ortmann, 1892) (Pl. 43) Ma-reum-mo-bam-ge (마름모밤게)

Philyra heterograna Ortmann, 1892, p. 582, pl. 26, fig. 17; Kamita, 1941, p. 43, fig. 15; Kim, 1973, pp. 306, 613, fig. 99, pl. 77, fig. 67a, b; T. Sakai, 1976, p. 111, fig. 63a, pl. 32, fig. 3; Takeda, 1982, p. 101, fig. 297; Dai and Yang, 1991, p. 86, fig. 39 (2), pl. 9 (7); K. Sakai, 1999, p. 18, fig. 1b, pl. 7C. Lyhira heterograna: Galil, 2009, p. 300, figs. 12B, 14.

Adult: Carapace subcircular, slightly longer than broad; dorsal surface sparsely covered with granules; regions indistinct, only branchiocardiac groove distinct; margins lined with granules. Front truncate. In maxilliped 3 ischium as long as merus measured along inner margin; exopod broader than merus. Chelipeds subcylindrical, ca. CL; each merus short, covered with granules; each carpus relatively smooth; each propodus smooth; fingers longer than propodus. Ambulatory legs short, slender, smooth; each merus subcylindrical. Abdomen heavily granulate on surface; somites 1, 2 of female articulate; somites 3-6 fused. Telson tongue- shaped. CL 14.5 mm, CW 14.0 mm.

Habitat: Muddy sand; 5 m.

Distribution: China, Taiwan, Japan, Korea.

Korea: GN, JN.

Specimen examined: JN: 1♀(Jindo: 26.vii.2012, S.H. Lee). Remarks: Larvae are unknown.

41 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Genus Merocryptus A. Milne-Edwards, 1873‌ Sae-mo-bam-ge-sok (세모밤게속)

Carapace subtriangular; regions distinct; dorsal surface irregularly sculpted, with granules; posterolateral margin forming wing-like projection. Front slightly produced, divided into 2 lobes. Chelipeds short. Gono- pod 1 of male long, straight.

Species 4 (1 in Korea).

22. Merocryptus lambriformis A. Milne-Edwards, 1873 Sae-mo-bam-ge (세모밤게)

Merocryptus lambriformis A. Milne-Edwards, 1873, p. 261, pl. 13, figs. 1, 1a-c; Kim, 1973, p. 294, fig. 90, pl. 76, fig. 59; T. Sakai, 1976, p. 78, fig. 40c, pl. 25, fig. 4; Takeda, 1982, p. 95, fig. 279; Kim and Chang, 1985, p. 45; Dai and Yang, 1991, p. 64, fig. 29, pl. 6 (8). Ebalia rugosa Yokoya, 1933, p. 124, fig. 43; Poore, 2004, p. 336, fig. 98f, pl. 19e.

Adult: Carapace subtriangular, broader than long; dorsal surface with granules; hepatic and posterobran- chial regions concave; branchial region swollen, with wing-like projection; intestinal region protruding, with 2 large spines on posterior margin. Front slightly produced, divided into 2 triangular lobes. Chelipeds short, slightly longer than CL, with granules on anterior surfaces; each merus bearing 2 acute tubercles on posterior margin proximally. Ambulatory legs covered with granules. Abdominal somites 3-6 of male fused, somite 2 small. Gonopod 1 long, straight; apical process slender. CL 10.9 mm, CW 14.0 mm (Kim, 1973).

Habitat: Mud; 60 m.

Distribution: Japan, Australia, Korea.

Korea: JJ.

Specimen examined: None.

Remarks: Larvae are unknown.

Genus Myra Leach, 1817‌ Gin-son-bam-ge-sok (긴손밤게속)

Carapace subcircular, only with posterior spines; dorsal surface smooth or granulate. Front narrow, with shallow median notch. Chelipeds extremely long, slender; each propodus cylindrical; fingers long. Gonopod

42 LEUCOSIIDAE: Myra

1 of male long, straight.

Species 18 (1 in Korea).

23. Myra celeris Galil, 2001 (Pls. 44-46) Gin-son-bam-ge (긴손밤게)

Myra fugax De Haan, 1833-1849 (1841), p. 134, pl. 33, fig. 1; Kim, 1973, pp. 298, 612, pl. 12, fig. 62; T. Sakai, 1976, p. 101, pl. 27, fig. 4; Takeda, 1982, p. 99, fig. 289; Miyake, 1983, p. 62, pl. 2, fig. 6; Dai and Yang, 1991, p. 78, pl. 8 fig. 6, fig. 36 (3). Myra fugax var. coalita Ortmann, 1892, p. 582 (pro parte). Persephona fugax: Rathbun, 1902, p. 30. Myra affinis Stimpson, 1858a, p. 160. Myra fugax coalita: T. Sakai, 1937, p. 136, text-fig. 23. Myra coalita: T. Sakai, 1976, p. 101, text-fig. 55. Myra celeris Galil, 2001b, pp. 418-421, figs. 1d, 7. Myra fugax: Terada, 1979 (zoeal stages).

Adult: Carapace subcircular, longer tha broad; dorsal surface covered with granules; regions indistinct; margins lined with granules, posterior margin with 3 spines (1 large median, 2 smaller lateral). Front with shallow median notch. In maxilliped 3 ischium twice longer than merus measured along inner margin. Che- lipeds extremely long, slender, ca. 3 CL; each merus cylindrical, covered with granules proximally, 1.2 times longer than CL; each carpus smooth; each propodus smooth, flattened, 1.5 times longer than movable finger; fingers denticulate on cutting margins. Ambulatory legs smooth, slender; each merus longer than other segments. Abdomen of male long, somites 3-6 fused, surface smooth. Telson subtriangular, 1.5 times longer than broad. Gonopod 1 long, straight; apical process slender, curved. CL 30.6 mm, CW 27.3 mm. Zoea I: Rostral and lateral carapace spines shorter than 1 CL, dorsal carapace spine ca. 1 CL. Protopod of antenna rounded. Endopod of maxillule with 4 setae. Endopod of maxilla with 4 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 1 outer spine; fork absent. CL 0.50 mm (Terada, 1979).

Habitat: Muddy sand or sand; 20-30 m.

Distribution: Indonesia, China, Japan, New Guinea, Australia, New Caledonia, Korea.

Korea: GN, JJ.

43 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Specimen examined: GN: 1♂(Busan: 10.x.2009, S.H. Lee).

Remarks: This species has five zoeal stages (Terada, 1979).

Genus Nursia Leach, 1817 O-gak-bam-ge-sok (오각밤게속)

Carapace pentagonal; regions distinct; dorsal surface with transverse ridge; margins thin, expanded. Front slightly produced. Gonopod 1 of male straight.

Species 20 (1 in Korea).

24. Nursia rhomboidalis (Miers, 1879) (Pls. 47-49) O-gak-bam-ge (오각밤게)

Ebalia rhomboidalis Miers, 1879, p. 42. Nursia sinica Shen, 1937, p. 279, figs. 1, 2; Shen and Dai, 1964, p. 17. Nursia rhomboidalis: T. Sakai, 1965, p. 39, fig. 5a, b; 1976, p. 87, fig. 44a, b; Dai and Yang, 1991, p. 67, fig. 30 (1), pl. 7 (3); Lee et al., 2009, pp. 1-3, fig. 1. Nursia rhomboidalis: Lee and Ko, 2017, pp. 228-234, figs. 1a, 2 (zoea I).

Adult: Carapace pentagonal, broader than long; dorsal surface relatively smooth; regions indistinct, median transverse ridge prominent; cardiac and intestinal regions each with distinct tubercle; anterolateral margin slightly convex medially; epibranchial angle forming triangular lobe; posterolateral margin with 2 triangular lobes; posterior margin with 2 smaller triangular lobes projecting downwards. Front slightly produced, margin truncate. In maxilliped 3 ischium twice longer than merus measured along inner margin. Chelipeds smooth, flattened, shorter than 2 CL; each merus crested on outer margin; each propodus crested on outer margin, 1.5 times longer than movable finger; fingers denticulate on cutting margins. Ambulatory legs short; each merus and propodus crested on anterior and posterior margins; each carpus prismatic, anterior margin and upper surface crested. Abdominal somites 3-6 of male fused, surface smooth, with 3 (2 proximal, 1 subdistal) tubercles. Telson subtriangular, 1.5 times longer than broad. Gonopod 1 long, straight; apical process slender, curved. CL 13.6 mm, CW 22.0 mm. Zoea I: Rostral carapace spine ca. 0.9 CL, lateral carapace spine ca. 0.6 CL, dorsal carapace spine ca. 1.2 CL. Protopod of antenna knob-like. Endopod of maxillule with 4 setae. Endopod of maxilla with 2+2 (4) setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively.

44 LEUCOSIIDAE: Philyra

Basis and endopod of maxilliped 2 each with 1+1+1+1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 1 outer spine; fork absent. CL 0.46 mm, RDL 1.27 mm.

Habitat: Mud; 5-30 m.

Distribution: China, Japan, Korea.

Korea: JN, CN.

Specimen examined: JN: 1♂(Jindo: 26.vii.2012, S.H. Lee); zoeas (Jindo: 27.v.2015, S.H. Lee).

Remarks: Lee and Ko (2017) described only the first zoeal stage of the species.

Genus Philyra Leach, 1817‌ Du-due-reo-gi-bam-ge-sok (두드러기밤게속)

Carapace subcircular, as long as broad, without spine; regions indistinct; dorsal surface covered with granules. Front slightly produced. Chelipeds short, stout. Abdomen of male with subdistal tubercle on surface. Gonopod 1 straight; apical process long or absent.

Species 32 (2 in Korea).

25. Philyra kanekoi T. Sakai, 1934 (Pls. 50, 51) Du-deu-reo-gi-bam-ge (두드러기밤게)

Philyra kanekoi T. Sakai, 1934, p. 286, fig. 4; 1976, p. 112, fig. 62, pl. 32, fig. 2; Kamita, 1941, p. 45, fig. 16; Kim, 1958, p. 7, fig. 1; 1973, p. 307, fig. 100, pl. 77, fig. 68a-d; Kim and Chang, 1985, p. 45. Philyra nipponensis Yokoya, 1936, p. 140, fig. 7; T. Sakai, 1937, p. 162. Philyra kanekoi: Ko, 2001 (zoeal stages).

Adult: Carapace subcircular, as long as broad; dorsal surface covered with granules except branchiocardiac region; regions relatively distinct, branchial, cardiac and intestinal regions convex, branchiocardiac groove depressed. Front with shallow median notch. In maxilliped 3 ischium as long as merus measured along inner margin. Chelipeds short, flattened, slightly longer than 1 CL; surface covered with granules; each merus covered with granules proximally; each propodus longer than fingers. Ambulatory legs short. Abdominal somite 1 of male articulate; somites 2-6 fused, surface with sudistal tubercle. Telson triangular, twice longer than broad. Gonopod 1 long, straight; apical process long, slender. CL 9.1 mm, CW 9.1 mm.

45 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Zoea I: Carapace globose, only with short rostral spine. Protopod of antenna short, with spinous tip. En- dopod of maxillule with 0, 4 setae. Endopod of maxilla with 3 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1 +1 +1 +1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 3 outer spines; fork absent. CL 0.45 mm.

Distribution: Japan, Korea.

Habitat: Muddy sand or sand; intertidal region.

Korea: GN, JJ.

Specimen examined: GN: 1♂ (Namhae: 25.v.2012, H.S. Ko); JJ: zoeas (Seongsanpo: 25.vii.1999, H.S. Ko).

Remarks: This species has three zoeal stages (Ko, 2001).

26. Philyra syndactyla Ortmann, 1892 (Pl. 52) Nap-jak-son-bam-ge (납작손밤게)

Philyra syndactyla Ortmann, 1892, p. 583, pl. 26, fig. 18; T. Sakai, 1936a, p. 65, fig. 26, pl. 12, fig. 4; 1976, p. 109, fig. 59b, pl. 33, figs. 1, 2; Kim, 1973, p. 304, fig. 98, pl. 14, fig. 66. Philyra syndactyla: Terada, 1979 (zoeal stages).

Adult: Carapace subcircular, slightly longer than broad; dorsal surface smooth; regions indistinct. Front not produced, with shallow median notch. In maxilliped 3 ischium as long as merus measured along inner margin. Chelipeds long, slender, flattened, more than 2 CL; each merus with granules proximally; each carpus and propodus smooth; fingers slightly longer than propodus. Ambulatory legs long, slender, smooth. Abdominal somites 1, 2 of male articulate; somites 3-6 fused, surface with sudistal tubercle. Telson subtriangular, as long as broad. Gonopod 1 long, straight; apical process absent. CL 14.6 mm, CW 13.4 mm. Zoea I: Carapace with rostral and dorsal spines, which slightly shorter than 1 CL; lateral spine absent. Protopod of antenna short, with spinous tip. Endopod of maxillule with 0, 4 setae. Endopod of maxilla with 3 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 3 outer spines; fork absent. CL 0.42 mm (Terada, 1979).

Habitat: Sand; 2-30 m.

Distribution: Japan, Korea.

46 LEUCOSIIDAE: Pyrhila

Korea: GB, GN.

Specimen examined: GN: 1♂(Busan: 30.vii.2009, H.S. Ko).

Remarks: This species has three zoeal stages (Terada, 1979).

Genus Pyrhila Galil, 2009‌ Bam-ge-sok (밤게속)

Carapace subcircular, slightly longer than broad, without spine; dorsal surface with granules; regions indistinct. Front slightly produced. Chelipeds short, stout. Only abdominal somite 1 of male articulate; somites 2-6 fused, surface without subdistal tubercle. Gonopod 1 straight; apical process present.

Species 3 (2 in Korea).

27. Pyrhila carinata (Bell, 1855) (Pls. 53-55) Yang-ma-bam-ge (양마밤게)

Philyra carinata Bell, 1855, p. 302, pl. 33, fig. 4; T. Sakai, 1937, p. 162; 1976, p. 113; Dai and Yang, 1991, p. 85, pl. 9 (6), fig. 39 (1). Philyra yangmataoensis Shen, 1932, p. 27, fig. 15, pl. 1, fig. 3; Kamita, 1941, p. 52, fig. 20; Kim, 1973, p. 310, fig. 102, pl. 78, fig. 70. Pyrhila carinata: Galil, 2009, p. 306, figs. 12E, 19. Pyrhila carinata: Lee and Ko, 2017, pp. 228-234, figs. 1B, 3 (zoea I).

Adult: Carapace subcircular, slightly longer than broad; dorsal surface covered with granules except on branchiocardiac region, longitudinal row of granules prominent medially; regions indistinct, only branchiocardiac groove distinct; margins lined with granules. Front truncate, with extremely shallow median notch. In maxilliped 3 ischium slightly longer than merus measured along inner margin; exopod as broad as merus, with row of longitudinal granules medially. Chelipeds short, flattened, more than CL; surface covered with granules; each propodus as long as fingers. Ambulatory legs smooth; each merus subcylindrical. Abdomen smooth on surface, without subdistal tubercle; somite 1 of male articulate; somites 2-6 fused. Telson subtriangular, longer than broad. Gonopod 1 long, straight; apical process short. CL 16.5 mm, CW 15.9 mm. Zoea I: Carapace globose, only with short rostral spine. Protopod of antenna short, with spinous tip. En- dopod of maxillule with 0, 4 setae. Endopod of maxilla with 3 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with

47 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

1+1+1+1 (4) and 0, 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 3 outer spines; fork absent. CL 0.49 mm.

Habitat: Sandy mud; intertidal region.

Distribution: Borneo Island, China, Korea.

Korea: CN, GG, JN.

Specimen examined: JN: 1♂(Muan: 21.viii.2012, S.H. Lee); zoeas (Muan: 5.vi.2014, S.H. Lee).

Remarks: Lee and Ko (2017) described only the first zoeal stage of this species.

28. Pyrhila pisum (De Haan, 1841) (Pls. 56-58) Bam-ge (밤게)

Philyra pisum De Haan, 1833-1849 (1841), p.131, pl. 33, fig. 7; T. Sakai, 1936a, p. 63, pl. 12, fig. 2; Kim, 1973, p. 308, fig. 101, pl. 14, fig. 69; Dai and Yang, 1991, p. 88, fig. 41 (2), pl. 10 (1). Pyrhila pisum: Galil, 2009, p. 308, figs. 12D, 20. Philyra pisum: Ko, 1996 (larval stages).

Adult: Carapace subcircular, slightly longer than broad; dorsal surface with granules on branchial, gastric, and cardiac regions, smooth on branchiocardiac region; regions indistinct, only branchiocardiac groove distinct; margins lined with granules. Front with shallow median notch. In maxilliped 3 ischium slightly longer than merus measured along inner margin; exopod as broad as merus, with minute granules on surface. Che- lipeds short, stout, slightly flattened, slightly longer than CL, granules on anterior and posterior surfaces; each propodus slightly shorter than fingers. Ambulatory legs smooth, slender; each merus subcylindrical. Abdomen smooth on surface, without subdistal tubercle; somite 1 of male articulate; somites 2-6 fused. Tel- son subtriangular, twice longer than broad. Gonopod 1 long, straight; apical process long. CL 21.9 mm, CW 20.9 mm. Zoea I: Carapace globose, only with short rostral spine. Protopod of antenna short, with spinous tip. En- dopod of maxillule with 0, 4 setae. Endopod of maxilla with 3 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1 +1 +1 +1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 3 outer spines; fork absent. CL 0.51 mm.

Distribution: Taiwan, China, Japan, Korea.

Habitat: Mud; intertidal region.

Korea: GG, CN, JB, JN, GN.

48 LEUCOSIIDAE: Euclosia

Specimen examined: GN 1♂(Busan: 2.vi.2012, S.H. Lee); JN: zoeas (Jindo: 25.vii.1994, H.S. Ko). Remarks: This species has two zoeal stages. The general morphology of it coincides well with that of Pyrhila carinata, therefore, to distingusish them is impossible at present.

Subfamily Leucosiinae Samouelle, 1819‌ Gin-i-ma-bam-ge-a-gwa (긴이마밤게아과)

Carapace highly polished, thoracic sinus present laterally. Front strongly produced. Postorbital neck present. Gonopod 1 of male contorted or coiled.

Genera 6 (3 in Korea), species over 70 (3 in Korea).

Distribution: Worldwide.

Genus Euclosia Galil, 2003‌ Dung-geun-mu-ni-bam-ge-sok (둥근무늬밤게속)

Carapace subcircular; dorsal surface smooth; thoracic sinus (Pl. 61) bearing row of granules, anterior margin loop-shaped. Front produced. Chelipeds short, stout. Abdominal somite 2 of male minute; somites 3-5 fused. Gonopod 1 coiled 3 times; apical process absent.

Species 10 (1 in Korea).

29. Euclosia obtusifrons (De Haan, 1841) (Pls. 59-61) Dung-geun-mu-ni-bam-ge (둥근무늬밤게)

Leucosia obtusifrons De Haan, 1833-1849 (1841), p. 133, pl. 33, fig. 2; T. Sakai, 1936a, p. 60, fig. 21, pl. 11, fig. 2; 1976, p. 119, figs. 65d, 67a, pl. 34, fig. 5; Kim, 1973, p. 301, fig. 95, pl. 13, fig. 64; Miyake, 1983, p. 67, pl. 23, fig. 3; Kim and Chang, 1985, p. 45. Leucosia mimasensis T. Sakai, 1969, p. 249, fig. 1b, b’; 1976, p. 120, pl. 36, figs. 68a, b, 1. Euclosia obtusifrons: Galil, 2003, p. 337, figs. 1E, 6E. Leucosia obtusifrons: Terada, 1984 (larval stages).

Adult: Carapace slightly longer than broad; dorsal surface convex, smooth, punctuate anteriorly. Front

49 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

with rounded margin. Postocular neck short. Thoracic sinus bearing row of granules, anterior margin loop- shaped. In maxilliped 3 endopod and exopod punctuate on surface, granules on distal half surface. Cheli- peds short, stout, slightly longer than CL; each merus prismatic, with tubercles proximally; each carpus and propodus with row of granules on inner margin; fingers slightly shorter than propodus, tips white. Ambu- latory legs smooth, subcylindrical except in propodi and dactyli. Abdominal somites 3-5 of female fused. Three yellowish spots on each side of gastric region, larger spot on each side of cardiac region distinct. CL 27.6 mm, CW 25.9 mm. Zoea I: Carapace with long rostral and dorsal spines, which ca. 1.5 CL; lateral spine long, ca. 0.5 CL. Protopod of antenna short, with rounded tip. Endopod of maxillule with 0, 4 setae. Endopod of maxilla with 3 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 3 outer spines; fork absent. CL 0.7 mm, RDL 2.0 mm (Terada, 1984).

Habitat: Mud; 70 m.

Distribution: Taiwan, Japan, Korea.

Korea: JJ.

Specimen examined: 1♀(Korea; purchasing date, 25.iv.2013, collector unknown).

Remarks: The specimen is deposited at Marine Natural History Museum of Busan Metropolitan City. Kim (1973) reported the gonopod 1 of this species coiled three times. It has three zoeal stages (Terada, 1984).

Genus Leucosia Weber, 1795‌ Gin-i-ma-bam-ge-sok (긴이마밤게속)

Carapace subcircular; dorsal surface smooth or punctuate; regions indistinct; thoracic sinus deep, anterior margin without loop-shaped granules. Front strongly produced. Chelipeds short, stout. Abdominal somite 2 of male minute; somites 3-5 fused. Gonopod 1 coiled, with apical process.

Speceis 25 (1 in Korea).

30. Leucosia anatum (Herbst, 1783) (Pls. 62-64) Gin-i-ma-bam-ge (긴이마밤게)

Cancer anatum Herbst, 1783, p. 93.

50 LEUCOSIIDAE: Leucosia

Leucosia australiensis Miers, 1886, p. 322, pl. 27, figs. 1, 1a-c. Leucosia neocaledonica A. Milne-Edwards, 1874, p. 40, pl. 2, fig. 1. Leucosia ornata Miers, 1877, p. 236, pl. 38, figs. 7-9. Leucosia polita Hess, 1865, p. 155, pl. 6, fig. 14. Leucosia splendida Haswell, 1879, p. 47, pl. 5, fig. 1. Leucosia anatum: Leach, 1815, p. 334; Sakai, 1976, p. 116, figs. 64, 65a, pl. 34, figs. 1-3; Miyake, 1983, p. 67, pl. 23, fig. 1; Kim and Chang, 1985, p. 45; Dai and Yang, 1991, p. 93, fig. 43 (1-3), pl. 10 (4); Poore, 2004, p. 337, fig. 99a. Leucosia longifrons De Haan, 1833-1849 (1841), p. 132, pl. 33, fig. 4; T. Sakai, 1936a, p. 61, pl. 11, fig. 1; Kamita, 1941, p. 42, fig. 14; Kim, 1973, p. 300, fig. 94, pl. 12, fig. 63. Leucosia longifrons: Terada, 1979 (larval stages).

Adult: Carapace subcircular, longer than broad; dorsal surface smooth, convex; regions indistinct. Front strongly produced, with triangular margin. Postocular neck long. Thoracic sinus bearing row of granules, anterior margin without loop-shaped granules. Maxilliped 3 smooth on surface, ischium as long as merus measured along inner margin. Chelipeds short, stout, slightly longer than 1 CL; each merus slightly flattened, with strong tubercles on proximal part and margins; each carpus smooth; each propodus smooth on surface, with row of granules on anterior margin; fingers shorter than propodus. Ambulatory legs smooth, each merus and carpus subcylindrical, each propodus and dactylus flattened. Abdominal somite 2 of male minute, somites 3-5 fused. Telson tongue-shaped. Gonopod 1 long, coiled 3 times, with apical process. CL 28.1 mm, CW 24.4 mm. Zoea I: Carapace with long rostral and dorsal spines, which ca. 2.0 CL; lateral spine long, more than 0.5 CL. Protopod of antenna short, with rounded tip. Endopod of maxillule with 0, 4 setae. Endopod of maxilla with 3 setae. Basis and endopod of maxilliped 1 each with 2+2+2+2 (8) and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 (4) and 3 setae, respectively. Lateral processes on abdominal somites 2, 3. Telson triangular plate, with 4 outer spines; fork absent. CL 0.52 mm (Terada, 1979).

Habitat: Sandy mud; 30-100 m.

Distribution: Madagascar, Pakistan, Sri Lanka, Andaman Islands, Indonesia, Philippines, Taiwan, China, Japan, New Caledonia, Fiji, Korea.

Korea: GB, GN, JJ.

Specimen examined: GN: 1♂(Busan: 13.ix.2012, S.H. Lee).

Remarks: This species has three zoeal stages (Terada, 1979).

51 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Genus Urnalana Galil, 2005‌ Bulg-eun-jeom-bam-ge-sok (붉은점밤게속)

Carapace rhomboidal; dorsal surface smooth or punctuate; regions indistinct. Thoracic sinus deep, setose; ventral margin without granules. Front strongly produced. Chelipeds short, stout. Abdominal somite 2 of male small; somites 3-5 fused. Gonopod 1 short, contorted; apical process curved.

Species 19 (1 in Korea).

31. Urnalana haematosticta (Adams and White, 1849) Bulg-eun-jeom-bam-ge (붉은점밤게)

Leucosia haematosticta Adams and White, 1849, p. 54, pl. 12, fig. 2; T. Sakai, 1936a, p. 61, pl. 12, fig. 1; 1976, p. 121, fig. 65e, pl. 36, fig. 2; Kim, 1973, p. 302, fig. 96, pl. 13, fig. 65; Dai and Yang, 1991, p. 97, fig. 45 (1-2), pl. 10 (8); Poore, 2004, p. 337, fig. 99b.

Adult: Carapace rhomboidal, posterior margin rounded; dorsal surface strongly convex; regions indistinct, obtuse median ridge present between front and gastric region. Front with rounded margin. Postocular neck short. Thoracic sinus deep, setose; ventral margin without granules. Chelipeds short, stout, slightly longer than CL; each merus with granules on margins, slightly flattened distally; each carpus smooth, convex, granules on proximal part and anterior margin; each propodus convex, carinate on posterior margin, row of granules on anterior margin; movable finger longer than fixed finger. Each propodus of ambulatory legs flattened, carinate on margin; each dactylus straight, with acute tip. Abdominen of male with 4 somites. Go- nopod 1 short, contorted; apical process curved. CL 11.7 mm, CW 11.2 mm (Kim, 1973).

Habitat: Sandy mud; intertidal-100 m.

Distribution: India, Singapore, Indonesia, China, Japan, Australia, Korea.

Korea: GN, JN.

Specimen examined: None.

Remarks: Larvae are unknown.

52 PINNOTHERIDAE

Superfamily Pinnotheroidea De Haan, 1833‌ Sok-sal-i-ge-sang-gwa (속살이게상과)

Symbionts on other invertebrates in mantle cavity of bivalve, cloaca of holothurians, tubes of polychaete worms, and burrows of echiurans and other crustaceans. Carapace poorly calcified in commensal females; dorsal surface smooth. Merus of maxilliped 3 completely or incompletely fused with ischium. One family, Pinnotheridae De Haan, 1833 and more than 290 species exist.

Family Pinnotheridae De Haan, 1833‌ Sok-sal-i-ge-gwa (속살이게과)

Carapace subcircular, subhexagonal, or transversely ovate. Antennules and antenna extremely small. Eye reduced. Genital openings of both sexes on thoracic sterna. All abdominal somites articulated. Males usually smaller than females. Two subfamilies, Pinnothereliinae Alcock, 1900 and Pinnotherinae De Haan, 1833 exist.

Key to the genera of family Pinnotheridae

1. Carapace twice broader than long...... Pinnixa (Pinnothereliinae) - Carapace not twice broader than long...... 2 (Pinnotherinae) 2. Carapace transversely ovate, 1.8 times broader than long...... Sakaina - Carapace subcircular or subhexagonal...... 3 3. Carapace covered with soft setae...... Pinnaxodes - Carapace not covered with soft setae...... 4 4. Dactylus of maxilliped 3 reaching tip of propodus...... Pinnotheres - Dactylus of maxilliped 3 not reaching tip of propodus...... Arcotheres

Key to the zoeas of family Pinnotheridae

Dorsal carapace spine absent, except in Pinnaxodes major. Endopod of maxillule with 0, 2+2 setae. En- dopod of maxilla with 1+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 (2) and 2, 2, 1, 2, 5 and setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 0, 5 (4) setae, respectively; except in Pinnaxodes major. Telson (Pl. 74) broader than long, with 3 lobes (1 rounded median, 2 acute lateral).

53 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

1. Carapace without dorsal spine, telson broader than long...... 2 - Carapace with dorsal spine, telson longer than broad...... Pinnaxodes major 2. Carapace with rostral and lateral spines...... Pinnotheres pholadis - Carapace without rostral and lateral spines...... Arcotheres sinensis

Subfamily Pinnothereliinae Alcock, 1900‌ Hae-sam-sok-sal-i-ge-a-gwa (해삼속살이게아과)

Carapace transversely ovate, twice broader than long. Merus of maxilliped 3 incompletely fused with ischium.

Genera 8 (1 in Korea), species over 70 (1 in Korea).

Distribution: Worldwide.

Genus Pinnixa White, 1846‌ Huin-hae-sam-sok-sal-i-ge-sok (흰해삼속살이게속)

Carapace twice broader than long. Front with median groove. Merus of maxilliped 3 incompletely fused with ischium. Ambulatory leg 3 longer, stouter than other legs.

Species 56 (1 in Korea).

32. Pinnixa tumida Stimpson, 1858 (Pl. 65) Huin-hae-sam-sok-sal-i-ge (흰해삼속살이게)

Pinnixa tumida Stimpson, 1858b, p. 108; Shen, 1932, p. 127, figs. 75-77, pl. 5, fig. 3; T. Sakai, 1936a, p. 201, pl. 57, fig. 5; 1976, p. 585, pl. 202, fig. 2; Kim, 1973, p. 422, fig. 171, pl. 88, fig. 132a-c; Dai and Yang, 1991, p. 437, fig. 220.

Adult: Carapace transversely ovate, ca. twice broader than long, fully calcified; dorsal surface smooth, convex; pair of grooves between gastric and cardiac regions; anterolateral margin with long setae ventrally; posterior margin slightly concave. Front narrow, divided into 2 small lobes by median groove. Maxilliped 3 with long setae on inner margin; ischium incompletely fused with merus; dactylus broad, spoon-shaped, exceeding tip of propodus. Chelipeds stout; each merus with short setae on inner margin; each carpus and

54 PINNOTHERIDAE: Arcotheres

propodus smooth, convex; fingers shorter than propodus, distal part of fixed finger and medial part of movable finger each with large triangular tooth on cutting margin. Ambulatory leg 1 shorter than leg 2; leg 3 longest, stoutest; leg 4 shortest. Abdomen of female broad, with 6 somites. CL 6.7 mm, CW 12.6 mm.

Habitat: Sandy mud; low tidal region.

Distribution: China, Japan, Korea.

Korea: CN.

Specimen examined: CN: 1♀(Taean: 26.ix.2007, H.S. Ko).

Remarks: This species inhabits the cloaca of holothurian, Paracaudina chilensis (see Kim, 1973). It has 5 zoeal stages (Konishi et al., 1988). The zoeal morphology of it is not described in the paper of Konishi et al. (1988) reported as an abstract.

Subfamily Pinnotherinae De Haan, 1833‌ Jo-gae-sok-sal-i-ge-a-gwa (조개속살이게아과)

Carapace subcircular, subhexagonal or transversely ovate, not twice broader than long. Merus of maxilliped 3 completely fused with ischium.

Genera 41 (4 in Korea), species over 220 (6 in Korea).

Distribution: Worldwide.

Genus Arcotheres Manning, 1993‌ Gul-sok-sal-i-ge-sok (굴속살이게속)

Carapace subhexagonal, smooth, poorly calcified. Merus of maxilliped 3 completely fused with ischium, dactylus not reaching tip of propodus. Dactyli of ambulatory legs 3, 4 longer than those of legs 1, 2.

Species 20 (1 in Korea).

33. Arcotheres sinensis (Shen, 1932) Gul-sok-sal-i-ge (굴속살이게)

Pinnotheres sinensis Shen, 1932, p. 131, figs. 78, 79, pl. 6, figs. 3, 4; T. Sakai, 1939, p. 584, fig. 70a-d; 1976, p. 569, fig. 309a-a’, pl. 200, figs. 1, 2; Kamita, 1941, p. 143, fig. 77; Kim, 1973, p. 415, fig. 167,

55 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

pl. 86, fig. 128a-d; Dai and Yang, 1991, p. 421, fig. 206 (1-3). Pinnotheres alcocki (not Rathbun, 1909) T. Sakai, 1936a, p. 197, pl. 56, fig. 4. Pinnotheres gordoni (not Shen, 1932) T. Sakai, 1939, p. 592, fig. 79. Arcotheres sinensis (Shen, 1932): Ng et al., 2008, p. 248. Pinnotheres sinensis: Konishi, 1983 (zoeal stages); Ko, 1991 (zoea I).

Adult: Carapace subhexagonal, poorly calcified, ca. 1.7 times broader than long; dorsal surface slightly smooth, convex; posteroior margin concave medially. Front narrow, deflexed ventrally. Merus of maxilliped 3 completely fused with ischium; dactylus digitiform, not reaching tip of propodus. Chelipeds smooth, slender; each merus smooth, cylindrical; each carpus and propodus smooth; fingers shorter than propodus, fixed finger with soft setae on lower margin, 2 small teeth on cutting margin, movable finger with larger tooth on cutting margin proximally; fingers curved, tips acute. Ambulatory legs smooth, slender; leg 1 shortest, leg 3 longest; dactyli of legs 1, 2 short; dactylus of leg 3 slightly shorter than propodus; dactylus of leg 4 longer than propodus, covered with short setae. Abdomen of female broad, with 6 somites. CL 7.5 mm, CW 10.5 mm. Zoea I: Carapace without spines. Antenna reduced as small seta. Endopod of maxillule with 0, 2+2 setae. Endopod of maxilla with 1+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 0, 4 setae, respectively. Abdomen broadened posteriorly, lateral processes on somites 2, 3. Telson broader than long; rounded median lobe protruding postriorly; 3 setae present between median and acute lateral lobe. CL 0.44 mm (Ko, 1991).

Habitat: In bivalve molluscs, Ostrea, Mytilus.

Distribution: China, Japan, Korea.

Korea: GN, JN, GG.

Specimen examined: GN: 1♀(Busan: 26.viii.1991, H.S. Ko).

Remarks: This species has three zoeal stages (Konishi, 1983).

Genus Pinnaxodes Heller, 1865‌ Yung-dan-sok-sal-i-ge-sok (융단속살이게속)

Carapace subcircular, fully calcified, covered with soft setae. Merus of maxilliped 3 completely fused with ischium, broad dactylus exceeding tip of propodus. Dactyli of ambulatory legs subequal in size.

Species 5 (1 in Korea).

56 PINNOTHERIDAE: Pinnaxodes

34. Pinnaxodes major Ortmann, 1894 (Pls. 66-68) Yung-dan-sok-sal-i-ge (융단속살이게)

Pinnaxodes major Ortmann, 1894, p. 697, pl. 23, figs. 10, 10h, i, k, z; T. Sakai, 1936a, p. 200, fig. 104a-c; 1965, p. 178, fig. 24a, b, pl. 87, fig. 3; 1976, p. 578, fig. 317a, pl. 202, figs. 4, 5; Kim, 1973, p. 421, fig. 170, pl. 87, fig. 131a, b. Pinnotheres major: Sakai, 1933, pp. 978, 980, 981, fig. 4a, b. Pinnaxodes major: Hong, 1974 (larval stages).

Adult: Carapace subcircular, ca. 1.2 times broader than long, fully calcified; dorsal surface slightly convex, covered with soft short setae; posterolateral margin slightly convex. Front transverse in dorsal view, triangular tip deflexed ventrally. Merus of maxilliped 3 completely fused with ischium; broad dactylus spoon- shaped, attached to propodus medially, exceeding tip of propodus. Chelipeds covered with soft short setae; each merus short, long setae on inner margin; each carpus convex on outer surface, long setae on inner angle; each propodus twice longer than broad; fingers shorter than propodus, fixed and movable fingers each with 4 (3 small, larger) teeth on cutting margin proximally; tips of fingers acute, curved. Ambulatory legs covered with soft short setae; leg 4 shortest, legs 2, 3 longest; all dactyli short, acute, subequal in size. Ab- domen of female broad, with 6 somites. CL 10.7 mm, CW 12.6 mm. Abdomen of male broadened proximally, narrowing to telson; distal half of telson expanded, posterior margin rounded. Gonopod 1 reaching to telson, proximal part straight, distal part strongly curved outwards; tip angled at right angle in relation to body of gonopod. CL 8.8 mm, CW 10.3 mm. Zoea I: Carapace with long spines, tips blunt; rostral spine ca. 1.2 CL, dorsal spine ca. 1.5 CL, lateral spine ca. 1.0 CL. Antenna with well developed protopod bearing 2 rows of spinules, exopod reduced as small seta. Endopod of maxillule with 0, 2+2 setae. Endopod of maxilla with 1+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1 +1 +1 and 0, 5 setae, respectively. Abdomen not broadened posteriorly, lateral processes on somites 2, 3. Telson longer than broad, without rounded median lobe; fork with minute lateral spine. CL 0.48 mm.

Habitat: In pen shell, Atrina pectinata.

Distribution: Sakhalin, Kurile Islands, Japan, Korea.

Korea: GN, JN, CN.

Specimen examined: GN: 1♀, 1♂(Busan: 5.v.2009, S.H. Lee); zoeas (Busan: 12.vi.2012, S.H. Lee).

Remarks: This species has five zoeal stages (Hong, 1974). Its zoeas are remarkably different from those of the genera Arcotheres and Pinnotheres (subfamily Pinnotherinae) by having a well developed antennal protopod (vs. reduced as a seta) and a slender telson with forks (vs. broad telson with a medial lobe). Such

57 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

zoeal characteristics agree well with those of Japanese species Pinnixa rathbuni (subfamily Pinnothereli- inae) described by Konishi (1983). Furthermore, this female species significantly differs by having a broad and spoon-shaped dactylus of the third maxilliped (vs. digitiform or styliform in other Pinnotherinae species). Such type of the third maxiiliped coincides with that of Pinnixa species described by Zmarzly (1992) and T. Sakai (1976). Therefore, the subfamily name of Pinnaxodes major should be altered to the subfamily Pinnothereliinae.

Genus Pinnotheres Bosc, 1802‌ Sok-sal-i-ge-sok (속살이게속)

Carapace subcircular or subhexagonal, poorly calcified, not covered with setae. Merus of maxilliped 3 completely fused with ischium, dactylus reaching tip of propodus. Dactyli of ambulatory legs subequally short or those of legs 3, 4 longer than others.

Species 71 (3 in Korea).

35. Pinnotheres cyclinus Shen, 1932 (Pl. 69) Dae-hap-sok-sal-i-ge (대합속살이게)

Pinnotheres cyclinus Shen, 1932, p. 135, figs. 80, 81, pl. 6, fig. 5; T. Sakai, 1939, p. 587, fig. 73; 1976, p. 570, fig. 310b; Kamita, 1941, p. 144, fig. 78; Kim, 1973, p. 417, fig. 168, pl. 86, fig. 129a, b; Dai and Yang, 1991, p. 423, fig. 207 (1, 2).

Adult: Carapace subhexagonal, poorly calcified, ca. 1.2 times broader than long; dorsal surface slightly convex anteriorly; lateral margin convex; posterior margin slightly concave medially. Front slightly protruding on dorsal view, triangular tip deflexed ventrally. Merus of maxilliped 3 completely fused with ischium; styliform dactylus attached to propodus medially, reaching tip of propodus. Chelipeds smooth, slender; each merus and carpus subcylindrical; each propodus subcylindrical, longer than merus, twice longer than broad; fingers shorter than propodus, fixed and movable fingers each with 3 (2 small, 1 larger) teeth on cutting margin proximally; fingers curved, tips acute. Ambulatory legs smooth, slender; leg 1 shortest, leg 2 longer than leg 4, leg 3 longest; dactylus of each legs curved, short in legs 1, 2, longer in leg 3, longest in leg 4, setae on posterior margins in legs 3, 4. Abdomen of female broad, with 6 somites. CL 12.6 mm, CW 15.5 mm.

Habitat: In bivalve shell, Saxidomus purpurata.

58 PINNOTHERIDAE: Pinnotheres

Distribution: China, Japan, Korea.

Korea: GN, JN, CN, GG.

Specimen examined: GN: 1 ovigerous♀(Saryangdo: 9.vi.2013, S.H. Lee). Remarks: Larvae are unknown.

36. Pinnotheres pholadis De Haan, 1835 (Pls. 70-73) Seob-sok-sal-i-ge (섭속살이게)

Pinnotheres pholadis De Haan, 1833-1849 (1835), p. 63; (1837), pl. 16, fig. 7. Pinnotheres pholadis: T. Sakai, 1936a, p. 198, pl. 56, fig. 2; 1939, p. 590, fig. 76, pl. 69, fig. 2; 1976, p. 571, pl. 199, figs. 2, 3; Kamita, 1941, p. 147, fig. 81; Kim, 1973, p. 419, fig. 169, pl. 87, fig. 130a-d; Miyake, 1983, p. 155, pl. 52, fig. 2; Dai and Yang, 1991, p. 428, fig. 213 (1, 2). Pinnotheres pisoides Ortmann, 1894a, p. 698, pl. 23, fig. 11. Pinnotheres pholades Lenz, 1901, p. 468. Pinnotheres affinis (not Bürger, 1895) Shen, 1932, p. 138, figs. 82, 83, pl. 6, fig. 7. Pinnotheres cardii (not Bürger, 1865) T. Sakai, 1933, pp. 978, 980, fig. 3; 1936, p. 198, pl. 56, fig. 3; Kami- ta, 1941, p. 146, fig. 79a, b. Pinnotheres pholadis: Konishi, 1983 (zoeal stages I, III).

Adult: Carapace subcircular, moderately calcified, ca. 1.2 times broader than long; dorsal surface smooth, slightly convex; posterior margin transverse. Front slightly deflexed ventrally, with triangular tip. Merus of maxilliped 3 completely fused with ischium; digitiform dactylus exceeding tip of propodus. Chelipeds smooth, slender; fingers shorter than propodus, fixed finger with 7 minute teeth on proximal half of cutting margin, movable finger with large tooth on cutting margin proximally; fingers slightly curved, tips not acute. Ambulatory legs slender, scattering of long setae; leg 2 longest, leg 4 shortest; each dactylus slightly curved, much shorter than propodus. Abdomen of female broad, with 6 somites. CL 8.7 mm, CW 10.2 mm. Zoea I: Carapace without dorsal spine; rostral spine ca. 0.6 CL, lateral spine ca. 0.5 CL, tips blunt. An- tenna reduced as small seta. Endopod of maxillule with 0, 2+2 setae. Endopod of maxilla with 1+2 setae. Basis and endopod of maxilliped 1 each with 2+2+3+3 and 2, 2, 1, 2, 5 setae, respectively. Basis and endopod of maxilliped 2 each with 1+1+1+1 and 0, 5 setae, respectively. Abdomen broadened posteriorly, lateral processes on somites 2, 3. Telson broader than long; rounded median lobe protruding postriorly; 3 setae present between median and acute lateral lobe. CL 0.45 mm.

Habitat: In pectinid, Chlamys farreri.

Distribution: Japan, Korea.

59 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Korea: GN, JB, GG.

Specimen examined: GN: 1♀(Busan: 24.x.2014, S.H. Lee); zoeas (Geoje: 6.vi. 2012, S.H. Lee).

Remarks: Number of zoeal stages is not known in the species because of Konishi (1983)’s zoeas collected from plankton materials.

37. Pinnotheres tsingtaoensis Shen, 1932 Gae-ryang-jo-gae-sok-sal-i-ge (개량조개속살이게)

Pinnotheres tsingtaoensis Shen, 1932, p. 149, figs. 92-94, pl. 6, figs. 10, 11; Dai and Yang, 1991, p. 433, fig. 217 (1-3); Kim, 1985, p. 85, fig. 4C, D.

Adult: Carapace subcircular, ca. 1.1 times broader than long. Front transverse on dorsal view. Merus of maxilliped 3 completely fused with ischium; propodus narrowed distally; styliform dactylus attached to propodus proximally, exceeding tip of propodus. Chelipeds scattering of setae; lower surface of propodus with series of setae, which extending to tip of fixed finger; fixed finger with 5-6 obtuse teeth on proximal half of cutting margin, movable finger with large tooth on cutting margin proximally; tips of fingers curved. Ambulatory leg 2 longest, leg 4 shortest; anterior marginal setae longer than posterior marginal setae in each merus; in legs 2, 3 each carpus with line of setae on upper surface, each propodus with anterior ridge and long marginal setae; each dactylus acute. CL 7.6 mm, CW 8.4 mm (Dai and Yang, 1991).

Habitat: In bivalve mollusc, Mactra chinensis.

Distribution: China, Taiwan, Korea.

Korea: JB.

Specimen examined: None.

Remarks: Larvae are unknown.

Genus Sakaina Serène, 1964‌ Im-po-yeop-gil-ge-sok (임포옆길게속)

Carapace transversely ovate, ca. 1.8 times broader than long. Merus of maxilliped 3 completely fused with ischium; dactylus attached to tip of propodus. Ambulatory legs decreasing in length from first to last.

Species 5 (1 in Korea).

60 PINNOTHERIDAE: Sakaina

38. Sakaina koreensis Kim and Sakai, 1972 Im-po-yeop-gil-ge (임포옆길게)

Sakaina koreensis Kim and T. Sakai, 1972, p. 29, figs. 1, 2; T. Sakai, 1976, p. 582, fig. 321.

Adult: Carapace transversely ovate, ca. 1.8 times broader than long; dorsal surface smooth, glabrous, anterior and posterior parts slightly deflexed; anterolateral margin rounded, rim with dense short setae; posterior margin straight. Front narrow, divided into 2 small lobes by median groove. Eyes and frontal lobes defined by rim of short setae. Each merus of cheliped short, covered with short setae; each carpus with short setae on upper inner angle; each propodus with 4 granules on upper inner margin; fingers with 8 denticles on cutting margins. Ambulatory legs covered with short setae; leg 1 subequal to length of leg 2, with triangular merus; legs 2, 3 each with depressed merus; leg 4 shortest; each dactylus strongly curved, black tip acute. Abdomen of female with 6 somites, lateral margins of segments 4-6 convergent distally. CL 2.9 mm, CW 5.3 mm (Kim and T. Sakai, 1972).

Habitat: With seaweeds.

Distribution: Korea.

Korea: JN.

Specimen examined: None.

Remarks: This species is a Korean endemic. Larvae are unknown.

61 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

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Lee, K.H. and Ko, H.S., 2007. Redescription of a leucosiid crab, Philyra platycheira (Crustacea, Brachyura) from Korea. J. Nat. Sci. Silla Univ., 16: 1-4. Lee, S.H. and Ko, H.S., 2017. First zoeas of Nursia rhomboidalis and Pyrhila carinata (Crustacea: Decapo- da: Leucosiidae) with a key to the known zoeas of ten leucosiid species from Korean Water. Anim. Syst. Evol., 33(4): 228-234. Lee, S.K., Song, S.J. and Kim, W., 2009. New record of leucosiid crab Nursia rhomboidalis (Crustacea: Decapoda: Brachyura) from Korea. Korean J. Syst. Zool., 25(3): 287-289. Lenz, H., 1901. Crustaceen. Ergebnisse einer Reise nach dem Pacific (Schauinsland 1896 1897). Zool. Jb., (Syst.), 14(5): 429-482, pl. 32. Linnaeus, C., 1758. Systema Naturae per Regna tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. (ed. 10), 1: 1-823. Linnaeus, C., 1763. Centuria Insectorum, Quam, Praesidae D.D. Car. von Linné, Proposuit Boas Johansson, Calmariensis. In: Linnaeus, C., Amoenitates Academicae; seu Dissertations variae, physicae, medicae, botanicae, Antehac seorsim editae, nunc collectae and auctae, 6: 384-415. Linnaeus, C., 1771. Mantissa Plantarum, Regni Animalis Appendix Insecta, pp. 521-552. Lucas, H., 1839. Observations sur un nouveau genre de Crustacés de l’Ordre des Décapodes brachyures. Ann. Soc. Entomol. Fr., 8: 573-581, pl. 21. Lucas, H., 1844. Cryptosome. In: C. d’Orbigny, Dictionnaire universal d’Histoire Naturelle résumant et complétant tous les faits présentés par les Encyclopédies, les anciens Dictionnaires, les oeuvres com- plètes de Buffon, et les meilleurs Traités spéciaux sur les diverses branches des sciences naturelles: - Donnant la description des êtres et des divers phénomênes de la nature, l’ètymologie et la définition des noms scientifiques, les principales applications des corps organiques et inorganiques, à l’agriculture à la médecine, aux arts industrials, etc.: ouvrage utile aux Médecins, aux Pharmaciens, aux Agriculteurs, aux Industriels, et généralement ê tous les hommes désireux de s’initier aux merveilles de la nature. Paris: Renard, Martinet et Cie, 4(1849): 1-752. Man, J.G. de, 1898. Bericht über die von Herrn Schiffscapitän Storm zu Atjeh, an den westlichen Küsten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Decapoden und Stomatopoden. Theil 6. Zool. Jahrb., Abt. Syst. Geogr. Biol. Thiere, 10: 677-708, pls. 28-38. [1898/II] McLay, C.L., 1991. A small collection of deep water sponge crabs (Brachyura: Dromiidae) from French Polynesia, including a new species of Sphaerodromia Alcock, 1899. Bull. Mus. natn Hist. nat., Paris, (4) 13, sect. A, (3-4): 457-481. McLay, C.L., 1993. Crustacea Decapoda: The Sponge Crabs (Dromiidae) of New Caledonia and the Philip- pines with a review of the genera. In: A. Crosnier (ed.), Résultats des Campagnes MUSORSTOM, Vol- ume 10. Mém. Mus. natn. Hist. nat., Paris, 156: 111, 251, figs. 1-19. McLay, C.L., 2001. Dynomenidae and Dromiidae (Decapoda, Brachyura) from Guam, Philippine Islands,Tonga and Samoa. Zoosystema, 23(4): 807-856. McLay, C.L., 2009. New records of crabs (Decapoda: Brachyura) from the New Zealand region, including a new species of Rochinia A. Milne-Edwards, 1875 (Majidae), and a revision of the genus Dromia Weber, 1795 (Dromiidae). Zootaxa, 2011: 1-66. Miers, E.J., 1877. Notes upon the Oxystomatous Crustacea. Trans. Linn. Soc. London, (Zool.), (2) 1(5):

67 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

235-249, pls. 38-40. Miers, E.J., 1879. On a collection of Crustacea made by Capt. H.C. St. John R.N. in the Corean and Japa- nese Seas. Part I. Podophthalmia. With an Appendix by Capt. H.C. St. John. Proc. Zool. Soc. London, 1879: 18-61, pls. 1-3. Miers, E.J., 1880. On a collection of Crustacea from the Malaysian Region. Part II. Thelphusidae, Catome- topa, and Oxystomata. Ann. Mag. Nat. Hist., 5(5): 304-317, pl. 14. Miers, E.J., 1886. Report on the Brachyura collected by H.M.S. Challenger during the years 1873-1876. In: C.W. Thompson & J. Murray, Report on the Scientific Results of the exploring Voyage of H.M.S. Chal- lenger during the years 1873-1876, under the command of Captain George S. Nares, R.N., F.R.S. and the Late Captain Frank Tourle Thomson, R.N., Zoology, 17: 1-362, pls. 1-29. Milne-Edwards, A., 1873. Description de quelques Crustacés nouveaux ou peu connus provenant du Musée de M.C. Godeffroy. J. Mus. Godeffroy, 1: 253-264, pls. 12-13. Milne-Edwards, A., 1874. Recherches sur la faune carcinologique de la Nouvelle-Calédonie. Part III. Nouv. Arch. Mus. Nat. Hist. nat. Paris, 10: 39-58, pls. 2-3. Milne-Edwards, H., 1837. Histoire naturelle des Crustacés, comprenant l’anatomie, la physiologie et la classification de ces animaux. Libraire Encyclopédique de Roret, Paris, Vol. 2: 1-532, pls. 1-2, 7-8, 10, 14, 18-19, 21, 24. Minemizu R, 2000. Marine Decapods and Stomatopod Crustaceans mainly from Japan. Buninchi Sogo Pub- lishing Co., Tokyo, pp. 1-344. Miyake, S., 1983. Japanese Crustacean Decapods and Stomatopods in color. Vol. II. Brachyura (Crabs), Hoikusha, Osaka, pp. 1-277, pls. 1-64. Naruse, T., 2014. Description of two new species of Arcania Leach, 1817, from the western Pacific Ocean and redescriptions of A. undecimspinosa De Haan, 1841, and A. elongata Yokoya, 1933 (Crustacea: Brachyura: Leucosiidae). Zootaxa, 3814(3): 301-332. Ng, P.K.L., 1998. Crabs. In: K.E. Carpenter & N. Volker (eds.), FAO Species Identification Guide for fish- ery purposes. The living marine Resources of the Western Central Pacific. Volume 1. Food and Agricul- ture Organisation, Rome, pp. 1046-1155. Ng, P.K.L., Guinot, D. and Davie, P.J.F., 2008. Systema Brachyurorum: Part 1. An annotated checklist of extant brachyuran crabs of the world. Raffles Bull. Zool., 17: 1-286. Ortmann, A.E., 1892. Die Decapoden-Krebse des Strassburger Museums, mit besonderer Berücksichtigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und z.Z. im Strassburg- er Museum aufbewahrten Formen. Theil V. Die Abtheilungen Hippidea, Dromiidea und Oxystomata. Zool. Jb. (Syst.), 6: 532-588, pl. 26. Ortmann, A.E., 1894. Die Decapoden-Krebse des Strassburger Museums mit besonderer Berücksichtigung der von Herrn Dr. Döderlein bei Japan und bei den Liu-Kiu-Inseln gesammelten und zur Zeit im Strass- burger Museum aufbewahrten Formen. Theil VIII. Abtheilung: Brachyura (Brachyura genuina Boas), III. Unterabtheilung: Cancroidea. 2. Section: Cancrinea, 2. Gruppe: Catametopa. Zool. Jb., (Syst.), 7: 683- 772, pl. 23. Parisi, B., 1914. Decapodi giapponesi del Museo di Milano. I. Oxystomata. Atti Soc. Ital. Sci., 53: 280-312, figs. 1-5, pls. 11-13.

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Parisi, B., 1915. I Decapodi giapponesi del Museo di Milano. II. Dromiacea. Atti Soc. Ital. Sci. nat., Milano, 54: 102-116, figs. 1-2, pls. 2-3. Poore, G., 2004. Marine decapod crustacea of Southern Australia: a guide to identification. CSIRO publishing, Australia, pp. 1-574. Rathbun, M.J., 1902. Japanese stalk-eyed Crustaceans. Proc. U.S. nat. Mus., 26(1307): 23-55, figs. 1-24. Rathbun, M.J., 1909. New crabs from the Gulf of Siam. Proc. Biol. Washington, 22: 107-114. Rathbun, M.J., 1923a. Report on the crabs obtained by the F.I.S. Endeavour on the coasts of Queensland, New South Wales, Victoria, South Australia, and Tasmania. 3. Biological results of the fishing experi- ments carried on by the F.I.S. Endeavour 1909-14. Australian Dept. Trade & Customs, Fisheries, Syd- ney, 5(3): 95-156, figs. 1-3, pls. 16- 42. Rathbun, M.J., 1923b. An analysis of Dromia dormia (Linnaeus). Proc. Boil. Soc. Wash., 36: 65-70. Romimohtarto, K., 1972. Five species of Matuta (Calappidae, Brachyura, Decapode) from Indonesia. Mar. Res. Indonesia, 12: 1-23, figs. 1-41, pls. 1-3. Rumphius, G.E., 1705-1766. D’Amboinsche rariteitkamer, behelzende eene beschryvinge van allerhande zoo weeke als harde schaalvisschen, te weeten raare krabben, kreeften, en diergelyke zeedieren, als mede allerhande Hoorntjes en schulpen, die men in d’Amboinsche zee vindt: daar beneven zommige Mener- aalen, gesteenten en soorten van aarde, die in d’Amboinsche, en zommige omleggende eilanden gevon- den worden, 1: 1-340, frontispiece, pls. 1-60. Rüppell, E.S., 1830. Beschreibungen und Abbildungen von 24 Arten kurzschwänzigen krabben, als Beiträge zur Naturgeschichte des Rothen Meeres. H.L. Brönner, Frankfurt am Main, pp. 1-28, pls. 1-6. Sakai, K., 1999. J.F.W. Herbst - collection of Decapod Crustacea of the Berlin Zoologica Museum, with remarks on certain species. Naturalists, Publ. Tokushima Biol. Lab. Shikoku Univ., 6: 1-45, pls. 1-21. Sakai, T., 1933. Botany and Zoology, Tokyo, 1(2): 977-983. Sakai, T., 1934. Brachyura from the coast of Kyushu, Japan. Sci. Rep. Tokyo Bunrika Daigaku, (B) 1 (Suppl. no. 25): 281-330. Sakai, T., 1936a. Crabs of Japan: 66 plates in life colours with descriptions. Sanseido, Tokyo, pp. 1-239, figs. 1-122, pls. 1-66. Sakai, T., 1936b. Studies on the crabs of Japan. I. Dromiacea. Sci. Rep. Tokyo Bunrika Daigaku, (B) 3 (Suppl. no. 1): 1-66, figs. 1-13, pls. 1-9. Sakai, T., 1937. Studies on the Crabs of Japan. II. Oxystomata. Sci. Rep. Tokyo Bunrika Daigaku, (B) 3: 67-192, figs. 1-45, pls. 10-19. Sakai, T., 1939. Studies on the crabs of Japan. IV. Brachygnatha, Brachyrhyncha. Yokendo Co., Tokyo, pp. 365-741, figs. 1-129, pls. 42-111. Sakai, T., 1965. The Crabs of Sagami Bay, collected by His Majesty the Emperor of Japan, Maruzen Co., Tokyo, 1-206 (English text), figs. 1-27, pls. 1-100: 1-92 (Japanese text): 1-26 (references and index in English): 27-32. Sakai, T., 1969. Two new genera and twenty-two new species of crabs from Japan. Proc. Biol. Soc. Wash- ington, 82: 243-280, figs. 1-20, pls. 1-2. Sakai, T., 1976. Crabs of Japan and the Adjacent Seas. Kodansha Ltd, Tokyo, pp. 1-773, figs. 1-379, pls.

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1-251. Serène, R., 1968. The Brachyura of the Indo-West Pacific region. In: Prodromus for a check list of the non-planctonic marine fauna of South East Asia. UNESCO Singapore National Academy of Sciences, Special publication No. 1, Fauna IIIC3: 33-112. Shen, C.J., 1931. The Crabs of Hong Kong. Part 1. Hong Kong Naturalist, 2(2): 92-110, 11 figs, pls. 4-10. Shen, C.J., 1932. The Brachyuran Crustacea of North China. Zool. Sinica, Peiping, (A) 9(1): 1-320, figs. 1-171, pls. 1-10. Shen, C.J., 1936. On a collection of Brachyuran Decapoda from Hainan Island, with description of three new species. Chinese J. Zool., 2: 63-80, figs. 1-4. Shen, C.J., 1937. Second Addition to the Fauna of Brachyuran Crustacea of North China, with a check list of the species recorded in this particular region. Contr. Inst. Zool. natn. Acad. Peiping, 3(6): 277-313, 11 figs. Shen, C.J. and Dai, A.Y., 1964. Illustrations of Animals in China (Crustacea part II), Peking, pp. 1-172, 277 figs. Sokolowsky, P.A., 1945. Biologisch-morphologische Betrachtung einiger Calappinae, Ortmann. Nebst Bes- chreibung einer anscheinend neuen Art aus dem Oestlichen Sued Amerika. Ann. Mus. Civ. Stor. Nat. Genova, 62: 62-75, pls. 1-2. Stebbing, T.R.R., 1917. The Malacostraca of Natal. I. Ann. Durban Mus., 2(1): 1-33, pls. 1-6. Stimpson, W., 1858a. Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et descripsit W. Stimpson. Pars VI. Crustacea Oxystomata. Proc. Acad. Nat. Sci. Philadelphia, 10: 159-164. Stimpson, W., 1858b. Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et descripsit. Pars V. Crustacea Ocypodoidea. Proc. Acad. Nat. Sci. Philadelphia, 10: 93-110. Stimpson, W., 1858c. Prodromus descriptionis animalium evertebratorum, quae in expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers Ducibus, observavit et descripsit, Pars VII. Crustacea Anomoura. Proc. Acad. Nat. Sci. Philadelphia, l0(4): 225-252. Takeda, M., 1982. Keys to the Japanese and foreign crustaceans fully illustrated in colors. Hokuryukan, Tokyo, pp. 1-285, figs. 1-779. Takeda, M. and Shikatani, N., 1990. Crabs of the genus Calappa from the Ryukyu Islands, with description of a new species. Zool. Sci., Tokyo, 7(3): 477-484, figs. 1-4. Terada, M., 1979. On the zoeal development of five species of the subfamilies Iliinae and Leucosiinae (Family Leucosiidae). Res. Crust., 9: 27-42. Terada, M., 1983. Zoea larvae of three crabs in the family Dromiidae. Zool. Mag., 92: 361-370. Terada, M., 1984. Larval forms of eight species of crabs (Philyrinae and Leucosiinae). Res. Crust., 13, 14: 153-164.

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Terada, M. 1987. Zoeal forms of 14 species of crabs from the Enshunada. Res. Crust., 16: 93-120. The Korean Society of Systematic Zoology, 1997. Lists of animals in Korea (excluding insects). Academy Publishing Co., Seoul, pp. 1-489. Tirmizi, N.M. and Kazmi, Q.B., 1991. Marine fauna of Pakistan: 4. Crustacea: Brachyura (Dromiacea, Archaeobrachyura, Oxystomata, Oxyrhyncha). Univ. Karachi BCCI (Bank Credit Commer. Int.) Founda- tion Chair, Publication No. 1 (1988): 1-244, figs. 1-65, pls. 1-4. Urita, T., 1926. On macrurous and brachyurous crustaceans from Tsingtao. Zool. Mag., 38: 421-438. Urita, T., 1926. On macrurous and brachyurous crustaceans from Tsingtao. Zoological Magazine, 38: 421- 438. Ward, M., 1928. The Crustacea of the Capricorn and Bunker Groups, Queensland. Austral. Zoologist, 5(3): 241-246, 1 fig., pls. 27-29. Ward, M., 1936. Crustacea Brachyura from the Coasts of Queensland. Mem. Queensl. Mus., 11(pt. 1): 1-13, pls. 1-3. Ward, M., 1941. New Brachyura from the Gulf of Davao, Mindanao, Philippine Islands. Am. Mus. Novit., New York, 1104: 1-15, figs. 1-30. Weber, F., 1795. Nomenclator entomologicus secundum Entomologiam systematicam ill. Fabricii adjectis speciebus recens detectis et varietatibus. Chilonii and Hamburgi, I -viii, pp. 1-172. White, A., 1847. List of the specimens of Decapoda Crustacea in the collection of the British Museum. Lon- don, Trustees of British Museum, pp. 1-143. World Register of Marine Species (WoRMS), 2015. [Internet]. WoRMS, Accessed 15 Aug 2015, . Yamaguchi, T. and Baba, K., 1993. Crustacean specimens collected in Japan by Ph. F. Von Siebold and H. Bürger and held by the Nationaal Natuurhistorisch Museum in Leiden and Other Museums. In: T. Yama- guchi (ed.), Ph. von Siebold and Natural History of Japan. Crustacea. Carcinological Society of Japan, pp. 145-570, figs. 1-200. Yang, K.C., Lee, S.H. and Ko. H.S., 2015. New report of two species of crabs, Cycloes granulosa and Pugettia vulgaris (Crustacea: Decapoda) collected from Korea. Anim. Syst. Evol. Divers., 31: 201-207. Yokoya, Y., 1933. On the distribution of decapod crustaceans inhabiting the continental Shelf around Japan, chiefly based upon the materials collected by S.S. Sôyô-Maru, during the years 1923-1930. J. Coll. Agric. Tokyo Imp. Univ., 12(1): 1-226, figs. 1-71. Yokoya, Y., 1936. Some rare and new species of decapod crustaceans found in the vicinity of the Misaki Marine Biological Station, Japan. Jap. J. Zool., 7(1): 129-146, figs. 1-10. Zmarzly, D.L., 1992. Taxonomic review of pea crabs in the genus Pinnixa (Decapoda: Brachyura: Pinnothe- ridae) occurring on the California Shelf, with descriptions of two new species. J. Crust. Biol., 12(4): 677-713.

도판 LITERATURES CITED

PLATES

73 109 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

PLATES

1. Cryptodromia tumida, dorsal view of female. 2. Lauridromia dehaani, dorsal view of male. 3. Lauridromia dehaani, ambulatory legs 3, 4 of male (↑ = subchelate tip). 4. Lauridromia dehaani, gonopods of male. 5. Metadromia wilsoni, dorsal view of male. 6. Metadromia wilsoni, anterior ventral view of male (↑ = merus of maxilliped 3). 7. Metadromia wilsoni, ventral view of abdomen and telson in male (↑ = uropod plate). 8. Metadromia wilsoni, gonopods of male. 9. Metadromia wilsoni, first stage zoeas. 10. Paradromia japonica, dorsal view of male. 11. Paradromia japonica, male carrying sponges. 12. Calappa hepatica, dorsal view of female. 13. Calappa hepatica, frontal view of female. 14. Calappa lophos, dorsal view of male (↑ = clypeiform expansion of carapace). 15. Calappa lophos, frontal view of male. 16. Calappa lophos, ventral view of male. 17. Calappa lophos, anterior ventral view of male (↑ = proximal tooth of cheliped). 18. Calappa lophos, gonopods of male. 19. Calappa philargius, dorsal view of male. 20. Cycloes granulosa, dorsal view of female. 21. Cycloes granulosa, frontal view of female. 22. Cycloes granulosa, ventral view of female. 23. Mursia trispinosa, dorsal view of male. 24. Ashtoret lunaris, dorsal view of male. 25. Ashtoret lunaris, anterior ventral view of male. 26. Matuta planipes, dorsal view of male. 27. Matuta victor, dorsal view of male. 28. Matuta victor, ventral view of male (↑ = milled ridge).. 29. Matuta victor, anterior ventral view of male (↑ = merus of maxilliped 3). 30. Matuta victor, gonopods of male. 31. Matuta victor, first stage zoeas. 32. Orithyia sinica, dorsal view of male.

74 PLATES

33. Orithyia sinica, ventral view of male. 34. Orithyia sinica, dorsal view of female. 35. Orithyia sinica, first stage zoea. 36. Arcania globata, dorsal view of female. 37. Arcania globata, ventral view of female. 38. Arcania undecimspinosa, dorsal view of male. 39. Arcania undecimspinosa, dorsal view of female. 40. Arcania undecimspinosa, first stage zoeas. 41. Hiplyra platycheir, dorsal view of male. 42. Hiplyra platycheir, dorsal view of female. 43. Lyphira heterograna, dorsal view of female. 44. Myra celeries, dorsal view of male. 45. Myra celeries, anterior ventral view of male (↑ = merus of maxilliped 3). 46. Myra celeries, gonopod of male. 47. Nursia rhomboidalis, dorsal view of male. 48. Nursia rhomboidalis, ventral view of male. 49. Nursia rhomboidalis, first stage zoeas. 50. Philyra kanekoi, dorsal view of male. 51. Philyra kanekoi, dorsal view of male (white color). 52. Philyra syndactyla, dorsal views of male and female. 53. Pyrhila carinata, dorsal view of female. 54. Pyrhila carinata, ventral view of female. 55. Pyrhila carinata, first stage zoeas. 56. Pyrhila pisum, dorsal view of male. 57. Pyrhila pisum, first stage zoeas. 58. Pyrhila pisum, abdomen and telson of first stage zoea. 59. Euclosia obtusifrons, dorsal view of female. 60. Euclosia obtusifrons, ventral view of female. 61. Euclosia obtusifrons, left thoracis sinus of female. 62. Leucosia anatum, dorsal view of male. 63. Leucosia anatum, anterior ventral view of male. 64. Leucosia anatum, gonopod of male. 65. Pinnixa tumida, dorsal view of female. 66. Pinnaxodes major, dorsal view of male. 67. Pinnaxodes major, first stage zoea.

75 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

68. Pinnaxodes major, first stage zoeas. 69. Pinnotheres cyclinus, frontal view of female (↑ = merus fused with ischium in maxilliped 3). 70. Pinnotheres pholadis, dorsal view of female. 71. Pinnotheres pholadis, dorsal view of male. 72. Pinnotheres pholadis, first stage zoeas. 73. Pinnotheres pholadis, abdomen and telson of first stage zoea.

76 PLATES

1 2

3 4

5 6

7 8

77 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

9 10

11 12

13 14

15 16

78 PLATES

17 18

19 20

21 22

23 24

79 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

25 26

27 28

29 30

31 32

80 PLATES

33 34

35 36

37 38

39 40

81 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

41 42

43 44

45 46

47 48

82 PLATES

49 50

51 52

53 54

55 56

83 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

57 58

59 60

61 62

63 64

84 PLATES

65 66

67 68

69 70

71 72

85 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

86 INDEX TO KOREAN NAMES

INDEX TO KOREAN NAMES

ㄱ 대합속살이게 58 두드러기밤게 45 가시투성밤게 38 두드러기밤게속 45 간만두게 22 둥근무늬밤게 49 갑각아문 11 둥근무늬밤게속 49 개량조개속살이게 60 갯가해면치레 13 ㅁ 갯가해면치레속 13 갯솜묻히 18 마름모밤게 41 갯솜묻히속 17 마름모밤게속 41 굴속살이게 55 만두게과 20 굴속살이게속 55 만두게상과 20 그물무늬금게 31 만두게속 21 그물무늬금게속 31 금게 29 ㅂ 금게과 28 금게붙이 27 밤게 48 금게붙이속 26 밤게과 36 금게속 29 밤게상과 36 긴손밤게 43 밤게속 47 긴손밤게속 42 밤게아과 38 긴이마밤게 50 범게 35 긴이마밤게속 50 범게과 34 긴이마밤게아과 49 범게상과 34 범게속 34 범무늬만두게 23 ㄴ 붉은점밤게 52 납작손밤게 46 붉은점밤게속 52

ㄷ ㅅ

단미하목 11 섭속살이게 59

87 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

세가시금게붙이 27 ㅈ 세모밤게 42 세모밤게속 42 잔점무늬금게 32 소름묻히 19 제주만두게 25 소름묻히속 19 제주만두게속 25 속살이게과 53 조개속살이게아과 55 속살이게상과 53 진연갑아강 11 속살이게속 58 집게털밤게 40 솜털묻히 16 집게털밤게속 40 솜털묻히속 16 십각목 11 ㅌ

ㅇ 털보해면치레 15 털보해면치레속 15 안경만두게 24 톱만두게 21 양마밤게 47 연갑강 11 열한가시밤게 39 ㅎ 열한가시밤게속 38 오각밤게 44 해면치레 14 오각밤게속 44 해면치레과 12 융단속살이게 57 해면치레상과 11 융단속살이게속 56 해면치레속 14 임포옆길게 61 해삼속살이게아과 54 임포옆길게속 60 흰해삼속살이게 54 흰해삼속살이게속 54

88 INDEX TO SCIENTIFIC NAMES

INDEX TO SCIENTIFIC NAMES

A D

Arcania 38 Decapoda 11 globata 38 Dromiidae 12 undecimspinosa 39 Dromioidea 11 Arcotheres 55 sinensis 55 Ashtoret 29 E lunaris 29 Ebaliinae 38 Euclosia 49 B obtusifrons 49 Eumalacostraca 11 Brachyura 11

H C Hiplyra 40 Calappa 21 platycheir 40 clypeata 21 hepatica 22 lophos 23 L philargius 24 Calappidae 20 Lauridromia 14 Calappoidea 20 dehaani 14 Crustacea 11 Leucosia 50 Cryptodromia 13 anatum 50 tumida 13 Leucosiidae 36 Cycloes 25 Leucosiinae 49 granulosa 25 Leucosioidea 36 Lewindromia 15 unidentata 15

89 INVERTEBRATE FAUNA OF KOREA Crabs and Zoeas V

Lyphira 41 P heterograna 41 Paradromia 17 japonica 18 M Petalomera 19 granulata 19 Malacostraca 11 Philyra 45 Matuta 31 kanekoi 45 planipes 31 syndactyla 46 victor 32 Pinnaxodes 56 Matutidae 28 major 57 Merocryptus 42 Pinnixa 54 lambriformis 42 tumida 54 Metadromia 16 Pinnothereliinae 54 wilsoni 16 Pinnotheres 58 Mursia 26 cyclinus 58 armata 27 pholadis 59 trispinosa 27 tsingtaoensis 60 Myra 42 Pinnotheridae 53 celeris 43 Pinnotherinae 55 Pinnotheroidea 53 Pyrhila 47 N carinata 47 pisum 48 Nursia 44 rhomboidalis 44 S

O Sakaina 60 koreensis 61 Orithyia 34 sinica 35 Orithyiidae 34 U Orithyioidea 34 Urnalana 52 haematosticta 52

90 Invertebrate Fauna of Korea Vol. 21, No. 46 Crabs and Zoeas V NIBR ) set ( 3 3 00- 354- 96470 94555- 6811- 89- 89- 113543 Not for sale / No charge Not for sale 978- 978-

BN BN IS IS 788968 9