Invertebrate Fauna of Korea of Fauna Invertebrate

Number 40

Ministry of Environment Volume 21, 21, Volume National of Biological Institute Resources Flora and Fauna of Korea and Fauna Flora Arthropoda: Maxillopoda: Copepoda: Cyclopoida Maxillopoda: Arthropoda: Cyclopoid Copepods Associated with Cyclopoid Cnidarians, Polychaetes and Crustaceans Polychaetes Cnidarians, Invertebrate Fauna of Korea of Fauna Invertebrate

Cyclopoid Copepods Associated with Invertebrate Fauna of Korea Vol. 21, No. 40 NIBR Cnidarians, Polychaetes and Crustaceans

Invertebrate Fauna of Korea

Volume 21, Number 40 Arthropoda: Maxillopoda: Copepoda: Cyclopoida Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

2015

National Institute of Biological Resources Ministry of Environment

Invertebrate Fauna of Korea

Volume 21, Number 40 Arthropoda: Maxillopoda: Copepoda: Cyclopoida Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Il-Hoi Kim Gangnenug-Wonju National University Invertebrate Fauna of Korea Volume 21, Number 40 Arthropoda: Maxillopoda: Copepoda: Cyclopoida Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Published by the National Institute of Biological Resources Environmental Research Complex, Hwangyeong-ro 42, 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 : 9788968112102-96470 Government Publications Registration Number 11-1480592-000993-010819-01

Printed by Junghaengsa, Inc. in Korea on acid-free paper

Publisher : Kim, Sang-Bae Author : Il-Hoi Kim Project Staff : Joo-Lae Cho, Jumin Jun and Jin Han Kim

Published on November 30, 2015

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. The book cover and the logo were designed by Jee-Yeon Koo. Chlorococcales: 1

Preface

The biological resources include all the composition of organisms and genetic resources which possess the practical and potential values essential to human live. Biological resources will be firmed competition of the nation because they will be used as fundamental sources to make highly valued products such as new lines or varieties, new material, and drugs. As the Nagoya Protocol th was adopted in 2010 and entered into force in the 12 Conference of Parties of the Convention on

Biological Diversity (CBD) in 2014, it is expected that the competition to get biological resources will be much intensive under the rapidly changed circumstance on the access and benefic sharing of the genetic resources (ABS). Therefore, each nation 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 Resources of the Ministry of Environment has been publish- ing the ‘Flora and Fauna of Korea’ since 2006 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 related experts of taxonomy examined systematically a total of 12,631 species for the past eight years to publish 151 volumes in both Korean and English versions, and two volumes of World Monograph covering 216 species. This year, 11 volumes of the Flora and Fauna of Korea in both Korean and English versions including 517 species of invertebrates, insects and algae are additionally published. Flora and Fauna of Korea were 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. Furthermore, publication of flora and fauna through identification of native species and investigation of national biota would be helpful to declare sovereignty rights over our native biological resources, be used as positive proof, and be utilized to provide the basic information of biological resources for indus- trial application. The National Institute of Biological Resources of the Ministry of Environment will continue to accelerate the project of the publication of the ‘Flora and Fauna of Korea’. Personally I would like to express my sincere appreciation for Professor Il-Hoi Kim of Gangneung-Wonju National Univer- sity who has continuously made a lot of efforts to publish an excellent version of Korean fauna.

Kim, Sang-Bae President National Institute of Biological Resources

Chlorococcales: 1

Contents

List of Taxa 3 Introduction 5 Materials and Methods 7 Taxonomic Notes 9

1. Foliomolgus cucullus Kim 9 2. Hemadona clavicrura Ho and Kim 12 3. Hemicyclops ctenidis Ho and Kim 16 4. Hemicyclops gomsoensis Ho and Kim 20 5. Hemicyclops japonicus Itoh and Nishida 25 6. Hemicyclops membranatus Moon and Kim 29 7. Hemicyclops nasutus Moon and Kim 34 8. Hemicyclops parilis Moon and Kim 37 9. Hemicyclops saxatilis Ho and Kim 42 10. Hemicyclops spinulosus Itoh and Nishida 45 11. Hemicyclops tanakai Itoh and Nishida 51 12. Hemicyclops ventriplanus Kim 55

13. Pontoclausia antiqua (Kim) 58

14. Pontoclausia lobata (Kim) 61 15. Clausia parva Kim 64 16. Indoclausia bipartita Kim 69 17. Likroclausia namhaensis Ho and Kim 74 18. Maxilliclausia propria Kim 77 19. Rhodinicola laticauda Ho and Kim 81 20. Sabellacheres illgi Dudley 84 21. Kelleria grandisetiger Kim 89 22. Kelleria portiviva Kim 91 23. Kelleria undecidentata Kim 93 24. Kelleria vaga Kim 95 25. Enalcyonium digitigerum Ho 98 26. Macrochiron anomalum Kim 101 27. Pseudomacrochiron urostenum Kim 103 28. Polyankylis orientalis Ho and Kim 107 29. Polyankylis ovilaxa Kim 111 30. Terebelliphilus simplex Kim 116 31. Sewelloya plana Kim 121 32. Pseudanthessius excertus Kim 126 33. Critomolgus anthopleurus Kim 131 34. Critomolgus malmizalus Kim 135 35. Critomolgus vicinus Kim 139 36. Critomolgus nudus Kim 143 2 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

37. Lutumidomus panikkari (Gnanamuthu) 146 38. Lutumidomus parvus Kim 151

39. Lutumidomus tertius (Kim) 154 40. Paramolgus nudipes Kim 157 41. Paramolgus orientalis Choe and Kim 161 42. Verutipes scutatus Kim 165 43. Zamolgus cavernularius Kim 170 44. Eupolymniphilus orientalis Kim 175 45. Myxomolgus invulgus Kim 178 46. Nasomolgus firmus Humes and Ho 182 47. Thamnomolgus robustus Humes 187 Literature Cited 192 Index to Korean Names 193 Index to Korean Names as Pronounced 195 Index to Scientific Name 197 Chlorococcales: 3

List of Taxa

Order Cyclopoida Burmeister, 1835 Family Clausidiidae Embleton, 1901 Genus Foliomolgus Kim, 2001 Foliomolgus cucullus Kim, 2001 Genus Hemadona Ho and Kim, 2004 Hemadona clavicrura Ho and Kim, 2004 Genus Hemicyclops Boeck, 1873 Hemicyclops ctenidis Ho and Kim, 1990 Hemicyclops gomsoensis Ho and Kim, 1991 Hemicyclops japonicus Itoh and Nishida, 1993 Hemicyclops membranatus Moon and Kim, 2010 Hemicyclops nasutus Moon and Kim, 2010 Hemicyclops parilis Moon and Kim, 2010 Hemicyclops saxatilis Ho and Kim, 1991 Hemicyclops spinulosus Itoh and Nishida, 1998 Hemicyclops tanakai Itoh and Nishida, 2002 Hemicyclops ventriplanus Kim, 2000 Family Clausiidae Giesbrecht, 1895 Genus Pontoclausia Bacescu and Por, 1959

Pontoclausia antiqua (Kim, 2001)

Pontoclausia lobata (Kim, 2000) Genus Clausia Claparède, 1863 Clausia parva Kim, 2014 Genus Indoclausia Sebastian and Pillai, 1974 Indoclausia bipartita Kim, 2014 Genus Likroclausia Ho and Kim, 2003 Likroclausia namhaensis Ho and Kim, 2003 Genus Maxilliclausia Kim, 2014 Maxilliclausia propria Kim, 2014 Genus Rhodinicola Levinsen, 1878 Rhodinicola laticauda Ho and Kim, 2003 Family Gastrodelphyidae List, 1889 Genus Sabellacheres M. Sars, 1861 Sabellacheres illgi Dudley, 1964 Family Kelleridae Humes and Boxshall, 1996 Genus Kelleria Gurney, 1927 Kelleria grandisetiger Kim, 2006 Kelleria portiviva Kim, 2006 Kelleria undecidentata Kim, 2006 Kelleria vaga Kim, 2000 Family Lamippidae Joliet, 1882 Genus Enalcyonium Olsson, 1868 Enalcyonium digitigerum Ho, 1984 4 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Family Macrochironidae Humes and Boxshall, 1996 Genus Macrochiron Brady, 1872 Macrochiron anomalum Kim, 2000 Genus Pseudomacrochiron Reddiah, 1969 Pseudomacrochiron urostenum Kim, 2000 Family Polyankyliidae Ho and Kim, 1997 Genus Polyankylis Ho and Kim, 1997 Polyankylis orientalis Ho and Kim, 1997 Polyankylis ovilaxa Kim, 2014 Genus Terebelliphilus Kim, 2000 Terebelliphilus simplex Kim, 2001 Genus Sewelloya Kim, 2014 Sewelloya plana Kim, 2014 Family Pseudanthessiidae Humes and Stock, 1972 Genus Pseudanthessius Claus, 1889 Pseudanthessius excertus Kim, 2014 Family Rhynchomolgidae Humes and Stock, 1972 Genus Critomolgus Humes and Stock, 1983 Critomolgus anthopleurus Kim, 1996 Critomolgus malmizalus Kim, 1996 Critomolgus vicinus Kim, 1996 Critomolgus nudus Kim, 2000 Genus Lutumidomus Kim, 2006

Lutumidomus panikkari (Gnanamuthu, 1955) Lutumidomus parvus Kim, 2006

Lutumidomus tertius (Kim, 2000) Genus Paramolgus Humes and Stock, 1972 Paramolgus nudipes Kim, 2006 Paramolgus orientalis Choe and Kim, 2010 Genus Verutipes Humes, 1982 Verutipes scutatus Kim, 2006 Genus Zamolgus Humes and Stock, 1972, Zamolgus cavernularius Kim, 2000 Family Sabelliphilidae Gurney, 1927 Genus Eupolymniphilus Humes and Boxshall, 1996 Eupolymniphilus orientalis Kim, 2006 Genus Myxomolgus Humes and Stock, 1972 Myxomolgus invulgus Kim, 2001 Genus Nasomolgus Sewell, 1949 Nasomolgus firmus Humes and Ho, 1967 Family Thamnomolgidae Humes and Boxshall, 1996 Genus Thamnomolgus Humes, 1969 Thamnomolgus robustus Humes, 1969 Chlorococcales: 5

Introduction

Copepods, as small crustaceans containing more than 12,500 described species throughout the world’s aquatic environments, outnumber every other group of multicellular animals on earth (Huys, 2014). They consist of eight or nine orders, of which the order Cyclopoida contains many copepods associated with marine invertebrates.

In the Illustrated Encyclopedia of Fauna & Flora of Korea (Kim, 1998) 161 species of symbiotic copepods were reported, of which 10 species of cyclopoids were recorded as associates of cnidarians

(phylum Cnidaria), polychaetes (phylum Annelida), and crustaceans (phylum Arthropoda) as follows: Family Clausidiidae Embleton, 1901 Hemicyclops ctenidis Ho and Kim, 1990 Hemicyclops gomsoensis Ho and Kim, 1991 Hemicyclops saxatilis Ho and Kim, 1991 Family Lamippidae Joliet, 1882 Enalcyonium digitigerum Ho, 1984 Family Polyankyliidae Ho and Kim, 1997 Polyankylis orientalis Ho and Kim, 1997 Family Rhynchomolgidae Humes and Stock, 1972 Critomolgus anthopleurus Kim, 1996 Critomolgus malmizalus Kim, 1996 Critomolgus vicinus Kim, 1996 Family Sabelliphilidae Gurney, 1927 Nasomolgus firmus Humes and Ho, 1967 Family Thamnomolgidae Humes and Boxshall, 1996 Thamnomolgus robustus Humes, 1969 Subsequent records, described cyclopoid species, and their hosts of cnidarians, polychaetes and crustaceans are as follows (scientific names of copepods without author names were recorded as new species). The scientifirc names conform to WoRMS (2014).

Kim (2000a): Hemicyclops ventriplanus from the mud shrimp Upogebia major (De Haan); Clausia lobata from the polychaete Marphysa sanguinea (Montagu); Kelleria vaga from stagnant water; Macro- chiron anomalum from stagnant water; Pseudomacrochiron urostenum from stagnant water; Critomol- gus nudus from the pennatulacean Virgularia gustaviana (Herklots); Lutumidomus tertius (= Notoxynus tertius) from the sea anemone Cerianthus filiformis Carlgren.

Kim (2000b): Zamolgus cavernularius from the pennatulacean Cavernularia obesa Valenciennes in Milne Edwards and Halime.

Kim (2001a): Terebelliphilus simplex from the polychaete Terebella ehrenbergi Grube; Myxomolgus invulgus from Myxicola sp.

Kim (2001b): Clausia antiqua from the polychaete Arenicola brasilliensis Nonata.

Kim (2001c): Foliomolgus cucullus from the polychaete Marphysa sanguinea (Montagu)

Ho and Kim (2003): Likroclausia namhaensis from the polychaete Dasybranchus caudatus Grube.

Ho and Kim (2004): Hemadona clavicrura from the polychaete Dasybranchus caudatus Grube

Kim (2006a): Eupolymniphilus orientalis from burrows of polychaetes and decapods.

Kim (2006b): Kelleria grandisetiger, K. undecidentata and K. portiviva from burrows of polychaetes 6 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans and crustaceans.

Kim (2006c): Lutumidomus panikkari (Gnanamuth, 1955) from the sea anemone Paracondylactis hertwigi (Wassilieff); L. parvus and Paramolgus nudipes from the sea anemone Flosmaris mutsuensis (Uchida); Verutipes scutatus from an unidentified species of sea anemone.

Choe and Kim (2010): Paramolgus orientalis from the antipatharian Antipathes japonicus Brook.

Moon and Kim (2010): Hemicyclops parilis from crustacean burrows; H. nasutus from un unidentified polychaete;H. membranatus from an unidentified polychaete.

Kim (2014): Indoclausia bipartite from a polychaete; Clausia parva from the polychaete Hydroides ezoensis Okuda; Maxilliclausia propria from a polychaete; Pseudanthessius excertus from a polychaete; Polyankylis ovilaxa from the polychaete Thelepus japonicus Marenzeller; Sewelloya plana from the polychaete Loimia medusa (Savigny). The present report deals with the 47 species of copepods known from Korean waters as associates of the cnidarians, polychaetes and crustaceans. This report mostly comprises a compilation of previous descriptions made by the present author or with co-authors, but the following four species are recorded for the first time in Korean waters: Hemicyclops japonicus Itoh and Nishida, Hemicyclops spinulosus Itoh and Nishida, Hemicyclops tanakai Itoh and Nishida, and Sabellacheres illgi Dudley. Chlorococcales: 7

Materials and Methods

The present report includes a compilation of previous descriptions and figures made by the author or with his coauthors. Copepods studied here are associates of cnidarians, polychaetes and crustaceans from Korean waters. Although four species of the family Kelleridae and one species (Eupolymniphilus orientalis Kim, 2006) of the family Sabelliphilidae were found unknown of their hosts, they are also included in the present report, since they were collected mainly from burrows of crustaceans and polychaetes. To collect the copepods, stagnant water in the invertebrate burrows was sucked up with a large plastic pipette and filtered through a dip net. In the laboratory the filtrates were sorted out under a dissecting microscope. Some copepods were collected from the washings of hosts. These hosts were collected from the intertidal and sublittoral water or obtained from fishing boats operating in waters around Korea. Finally, all collected copepods were preserved in 80% ethanol. For microscopic observation, the copepods were immersed in lactic acid for at least 10 minutes and dissected. Dissected appendages were observed using the reverse slide method of Humes and

Gooding (1964). Drawings were made with the aid of a microscope equipped with a drawing ap- paratus. In the description section of each species, armature formula is given as: P1-P4 represent legs 1-4, Roman numerals indicate spines, and Arabic numerals indicate setae. The abbreviations “exp.” and “enp.” represents “exopod” and “endopod”, respectively. Lengths of copepod specimens and measurements of appendages are mostly based on a single dissected specimen of each species. In the description of species, males are described mainly on the basis of its differences from females. Collection data for each species include all specimens observed previously by the author.

Morphological terminology for the caudal setae follows Huys and Boxshall (1991) and for the maxilla Humes and Boxshall (1996) for the maxilla. The collector of the copepod material studied in this report is the author, unless otherwise mentioned.

Taxonomic Notes

Order Cyclopoida Burmeister, 1835

Geom-mul-byeo-ruk-mok (검물벼룩목)

Family Clausidiidae Embleton, 1901

Geom-mul-byeo-ruk-but-i-gwa (검물벼룩붙이과)

Body cyclopiform. Urosome 5-segmented in female and 6-segmented in male. Antennule 6-or 7-segmented, with 4 or 5 setae on first segment. Antenna 4-segmented; armature formula 1, 1, 4, and 7; third segment with 4 setae or 3 setae plus 1 claw. Mandible with 3 or 4 distal elements. Maxillule with 5+3 setae. Legs 1-4 with 3-segmented exopod nand endopod. Leg 1 with inner distal spine on basis in female. Leg 5 exopod with 4 elements.

Genus Foliomolgus Kim, 2001

Ip-geom-mul-byeo-ruk-sok (잎검물벼룩속)

As features of the family, except the followings: antenna with 1 hook-like process and 3 setae on third segment. Maxilla resembling that of the Clausiidae; proximal segment with 1 seta; distal segment reduced and blunt distally, with 3 small setae. Maxilliped absent in female. Associated with polychaetes.

1. Foliomolgus cucullus Kim, 2001 (Fig. 1)

Ip-geom-mul-byeo-ruk (잎검물벼룩)

Foliomolgus cucullus Kim, 2001c, p. 661, figs. 1-3.

Female: Body (Fig. 1A) cyclopiform, 2.67 mm long. Greatest width 1.02 mm. Prosome composed of cephalothorax and 3 pedigerous somites. Rostral area of cephalosome produced anteriorly. Prosomal somites with well developed epimera. Epimera of second and third pedigerous somites with pointed posterior corners. Epimera of fourth pedigerous somite with rounded corners.

Urosome (Fig. 1B) slender and 5-segmented. Fifth pedigerous somite 405 μm wide, characteristically with dorsal hood (487 μm wide) covering most of dorsal surface of fifth pedigerous somite and anterior part of genital double-somite. Genital double-somite 335×295 μm, widest at anterior third, then tapering posteriorly. Genital aperture located laterally, invisible from dorsal and ventral 10 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans views. Three abdominal somites 210×165, 162×142, and 115×115 μm, respectively. Anal somite with spinules along posteroventral border. Caudal ramus slender, divergent, 221×40 μm (5.53 : 1), with parallel lateral margins and 6 setae. Posteroventral border of caudal ramus armed with several denticles. All caudal setae naked. One of terminal setae distinctly larger than others, 1.05 mm long. Rostrum wider than long, rounded posterior margin. Antennule 7-segmented, 731 μm long, with armature formula 5, 15, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Antenna 4-segmented, elongate, with armature formula 1, 1, 3+1 claw, and 7. First and second segments slender, each about 3 times as long as wide. Seta on these segments relatively small and naked. Third segment armed with spinules on inner margin. Claw on this segment prominent, one of 3 setae very small. Fourth segment originating from proximal part of outer margin of third segment, wider than long, armed terminally with 4 spiniform setae and 3 simple setae, outermost seta plumose.

Labrum reduced, distinctly shorter than wide, covering only part of oral appendages. Mandible (Fig. 1C) armed with 1 terminal fleshy, atrophied element tipped with a small claw-like process, and 3 long, very thin, foliaceous subterminal elements covered with minute spinules on all surfaces. Paragnath with 2 hairy lobes on medial side. Maxillule tapering, proximally with several setules, and terminally bilobed, with 5 setae (3 lateral ons larger) on anterior lobe and 3 setae on posterior lobe. Maxilla (Fig. 1D) 2-segmented. Basal segment greatly expanded proximally, roughly triangular, with 1 terminal seta. Distal segment terminally truncated and sclerotized, and armed with 3 unequal lateral setae; terminal portion of distal segment covered with numerous, minute spinules. Maxilliped absent.

Leg 1 (Fig. 1E) and legs 2-4 with 3-segmented rami. All these legs spiniferous. All legs with 1 large inner coxal seta; those of legs 1-3 plumose on 1 side and weakly spinulated on the other side. Inner spine on basis of leg 1 shorter than first endopodal segment of same leg. Posteromedian part of basis of legs 1-4 ornamented with spinules. Each leg with endopod distinctly longer than exopod. Armature formula as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0; I-1; I, 7; enp. 0-1; 0-1; I, 5 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 7; enp. 0-1; 0-2; III, 3 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; I, 7; enp. 0-1; 0-2; III, 2 Leg 5 2-segmented. Protopod articulated from fifth pedigerous somite, distally armed with 1 small seta and spinules. Exopod elongate, tapering, armed with 1 outer lateral and 3 terminal setae; all setae naked, and as long as or shorter than distal segment. Leg 6 not seen. Male: Body similar to that of female. Length 2.20 mm. Urosome 6-segmented. Genital somite 175×317 μm, distinctly narrower than fifth pedigerous somite, without dorsal hood. Genital operculum with numerous spinules near inner margin. Four abdominal somites 154×187, 175×155,

150×135, and 104×113 μm, respectively. Caudal ramus 198×33 μm (6.0 : 1), more slender than that of female. Antennule with 1 additional seta on distal portion of third segment. Antenna, labrum, paragnath, and maxillule similar to those of female. Mandible with more prominent claw on tip of terminal element. Maxilla with first segment not expanded, but distal segment as in female. Maxilliped 4-segmented. First segment with 1 long distal seta. Second segment triangular, greatly expanded proximally, strongly tapering, with a flat, thin protrusion at inner proximal corner, and armed with

2 rows of thin, truncate spinules (1 row shorter), 2 rows of epicuticular extensions and 2 small setae (1 in the middle, the other smaller 1 in distal part) on inner margin. Third segment short and unarmed. Terminal segment forming a claw bearing proximally 2 setae and 1 distal membranous process. Cyclopoida: Clausidiidae: Foliomolgus 11

B A

Fig. 1. Foliomolgus cucullus, female. A. habitus, dorsal; B. urosome, ventral; C. mandible; D. maxilla; E. leg 1. Scales: A=0.5 mm, B=0.2 mm, C=0.05 mm, D, E=0.1 mm.

Legs 1-4 as in female. Protopod of leg 5 completely fused with fifth pedigerous somite, leaving only 1 distal seta. Free exopod broader and shorter than that of female. Leg 6 represented by 1 stiff seta on terminal corner of genital operculum.

Distribution: Japan, Korea. Korea: GN, JJ, JB, CN.

Specimens examined: 1♀, 2♂♂ (type specimens), on the external surface of the polychaete Mar- 12 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

physa sanguinea (Montagu), intertidal coral sands at Sinhung-ri on the northern shore of Jeju Is- land (approximately 33°33′N, 126°39′E), 3.vi.2000; 2♀♀, 4♂♂ from polychaetes, intertidal rocky shore, Gyeokpo, Buan, Jeollabuk-do, 8.iv.2001; 4♀♀, 1♂, from polychaetes on tidal flat, Geum- song-ri, Namhae I., Gyeongsangnam-do, 7.vii.2001; 2♀♀, 2♂♂, from polychaetes on tidal flat, Geumsong-ri, Namhae I., Gyeongsangnam-do, 22.vii.2001; 1♀, from polychaetes, intertidal rocky shore, Choonjangdae, Choongcheongnam-do, 28.ii.2002; 2♀♀, from polychaetes, intertidal rocky shore, Sapsi I., Choongcheongnam-do, 24.viii.2005; 9♀♀, 2♂♂, from polychaetes, intertidal, Seog- wipo port, Jeju I., 19.viii.2005. Remarks: This species is also distributed in Japan. The author could collect 1 female from mud- flat on the Matsushima coast east of Sendai, northeast of Japan, on October 4, 2008.

Genus Hemadona Ho and Kim, 2004

Nam-hae-geom-mul-byeo-ruk-sok (남해검물벼룩속)

General features as those of the family, except for the followings: Body elongate. Antennule 6-segmented, but second and third segment incompletely defined. Antenna with with 3 setae and 1 large, hook-like process on third segment. Maxilliped of female 3-segmented; with armature formula 2, 2, and 3; terminal segment strongly reduced. Maxilliped of male with large inner process on first segment. Associated with polychaets.

2. Hemadona clavicrura Ho and Kim, 2004 (Fig. 2)

Nam-hae-geom-mul-byeo-ruk (남해검물벼룩)

Hemadona clavicrura Ho and Kim, 2004, p. 97, figs. 1-3.

Female: Body (Fig. 2A) elongate, 6.34 mm long. Cephalothorax semicircular and containing first pedigerous somite. Second pedigerous somite widest of body, 1.04 mm; width of third and fourth pedigerous somites decreasing only slightly from that of second pedigerous somite. Urosome 5-segmented, 2.37 times longer than prosome. Genital double-somite longer than wide, 877×693 μm, with aliform dorsolateral protrusion in anterior half of somite covering area of egg sac attachment. Abdomen 3-segmented, with all segments longer than wide, 833×553 μm, 798×508 μm, and 880×430 μm. Caudal ramus 4.44 times longer than wide (720×162 μm), armed with 1 short, outer seta at about midlength of lateral margin, 1 short, medial, subterminal seta, and 2 short and

2 long terminal setae; longest terminal seta (830 μm) 1.15 times as long as ramus. Egg sac greatly elongated (7.05 mm), longer than body and cylindrical. Rostrum subquadrate in dorsal view, produced forward, and well demarcated from cephalothorax. Antennule short and robust, 6-segmented; formula of armature: 5, 16, 10, 4, 2+aesthetasc, and

7 +aesthetasc. Antenna (Fig. 2B) 4-segmented; first segment (coxobasis) longer than wide, with long outer-distal seta; second segment (first endopodal segment) shorter than proximal segment, with small subterminal seta; third segment (second endopodal segments) drawn out into a large uncinate claw, with basal patch of spinules on outer surface and 2 unequal setae plus 1 blunt tip, Cyclopoida: Clausidiidae: Hemadona 13

B A

F E

Fig. 2. Hemadona clavicrura, female. A. habitus, dorsal; B. antenna; C. mandible; D. maxilla; E. maxilliped; F. leg 1. Scales: A=0.5 mm, B-E=0.05 mm, F=0.1 mm. 14 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans bent, spiniform seta bearing terminal row of spinules on medial margin; terminal segment 2.75 times longer than wide, tipped with 3 unequal setae and 4 spiniform setae structured as that 1 on third segment. Labrum well-developed, with submarginal, inner, central process, and 2 disjunct, marginal rows of spinules on either side of this process. Mandible (Fig. 2C) armed terminally with 1 stout, pinnate element, 1 stout, spinulose element, and 1 pinnate and 1 naked setae. Paragnath an obtuse lobe fringed with spinules on distal margin. Maxillule bilobate, small outer lobe tipped with 1 long and 2 short setae and larger inner lobe with 5 unequal setae. Maxilla (Fig. 2D) 2-segmented; robust proximal segment (syncoxa) armed with 1 large spiniform and 1 small pinnate setae; distal segment (allobasis) tipped with 2 spiniform elements bearing spinules on 1 side and 2 pinnate setae. Maxilliped 3-segmented; proximal segment (syncoxa) with 2 unequal medial setae; middle segment (basis) greatly expanded laterally and carrying 2 unequal medial setae; terminal segment (endopod) tiny, bearing 1 spiniform and 2 setiform elements.

Leg 1 (Fig. 2E) and legs 2-4 biramous, with 3-segmented rami. Outer surface of all segments on rami fringed with spinules. Outer spines on all legs club-shaped, with swollen tip covered with fine denticles. Armature formula of ;egs 1-4 as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; II, 4 Leg 2: coxa 0-I; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; II, I, 3 Leg 3: coxa 0-I; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; II, II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp.0 -1; 0-2; II, II, 1 Leg 5 2-segmented; proximal segment small, carrying simple, outer seta; distal segment elongate, about 4 times longer than wide (750×187 μm), armed with 3 club-like spines and 1 thin, simple seta.

Male: Body elongate as in female, 3.40 mm long (excluding setae on caudal rami). Cephalotho- raxs hat-shaped, containing first pedigerous somite. Second pedigerous somite widest of body, 532 μm wide; width of third and fourth pedigerous somites decreasing only slightly from that of second pedigerous somite. Urosome 6-segmented, 1.51 times longer than prosome. Ventrally, protopod of leg 5 indistinctly separated from its somite. Genital somite slightly longer than wide, 310×300 μm; genital operculum small. Abdomen 4-segmented, with following measurements (proceeding from anterior to posterior): 295 ×282 μm, 366×275 μm, 317×254 μm, and 423×246 μm. Caudal ramus 4.08 times longer than wide (408×100 μm) and armed as in female.

Maxilliped (Fig. 2F) 4-segmented; proximal segment (syncoxa) with large, medial protrusion tipped with 3 sharp tines; second segment (basis) largest, armed with small patch of subterminal denticles on lateral surface, 1 seta in distomedial corner followed by 1 row of spinules on medial margin; third segment (first endopodal segment) smallest and naked; distal segment (second endopodal segment) drawn out in to 1 long claw with accessory tine and 2 simple setae on medial surface of basal region.

Distribution: Korea. Korea: GN.

Specimens examined: 3♀♀, 7♂♂ (type specimens), from washings of the polychaete Dasybran- chus caudatus Grube collected from intertidal mud flat on Namhae-do Island (34°49′N, 128°03′E) in Korea Strait, 22.vii.2001; 1♀, from polychaetes, tidal flat, Dapo-ri, Geoje I., 24.iv.2008; 2♀♀, from polychaetes, type locality, 21.vi.2012. Remarks: The body of this species is characteristically elongate and very large, growing up to 6 mm long in the female. Cyclopoida: Clausidiidae: Hemicyclops 15

Genus Hemicyclops Boeck, 1873

Geom-mul-byeo-ruk-but-i-sok (검물벼룩붙이속)

Body cyclopiform. Antennule 7-segmented (6-segmented in Hemicyclops ctenidis). Antenna 4-segmented, with armature formula of 1, 1, 4, and 7; one of setae on third segment may be transformed into claw. Mandible with 2 spiniform elements and 1 or 2 setae distally. Maxillule distally bilobed, with 3 and 5 setae, respectively on ech lobe. Maxilla 2-segmented; proximal segment with 2 setae; distal segment with 1 spine, 2 setae and 1 thick, spiniform process in female. Maxilliped 4-segmented; armature formula 2, 2, 0, and 5 in female. Legs 1-4 with 3-segmented exopod and endopod. Leg 4 exopod with 3 spines and 1 seta.

Key to the species of Hemicyclops

1. Antennule 6-segmented; second endopodal segment of leg 4 with 1 inner seta ·············· H. ctenidis - Antennule 7-segmented; second endopodal segment of leg 4 with 1 inner seta ······························ 2 2. Female genital double-somite trapezoidal; third exopodal segment of leg 1 with 9 elements ········· ·················································································································································· H. spinulosus - Female genital double-somite not trapezoidal; third exopodal segment of leg 1 with 8 elements ······································································································································································· 3 3. Caudal ramus of female less than twice as long as wide ····································································· 4 - Caudal ramus of female more than twice as long as wide ·································································· 5 4. ‌Third exopodal segment of leg 2 with 8 elements; lateral margins of genital double-somite of female strongly constricted ······································································································ H. japonicus - ‌Third exopodal segment of leg 2 with 9 elements; lateral margins of genital double-somite of female not constricted, but with anterior expansion ······························································ H. saxatilis 5. Genital double-somite of female complicated, with 2 pairs of anterolateral flaps ··························· 6 - Genital double-somite of female without anterolateral flaps ······························································ 7 6. Genital double-somite of female with 2 pairs of dorsolateral flaps, posterior 1 of which is blunt; caudal ramus of female 3.1-3.4 times as long as wide ··················································· H. gomsoensis - Genital double-somite of female with 2 pairs of dorsolateral flaps, posterior 1 of which is pointed; caudal ramus of female 2.4-2.9 times as long as wide ···················································· H. parilis 7. Caudal ramus of female more than 4 times as long as wide; third segment of antenna with 3 seta and 1 claw ·························································································································· H. membranatus - Caudal ramus of female less than 3 times as long as wide; third segment of antenna with 4 setae ······································································································································································· 8 8. First segment of antennule with 5 setae ················································································· H. nasutus - ‌First segment of antennule with 4 setae ································································································· 9 9. Genital double-somite of female with broad anterior 4/5 and narrower posterior 1/5; male genital somite with 2 posterolateral spine as leg 6 ········································································ H. tanakai - Genital double-somite of female with broad anterior 1/3 and narrower posterior 2/3; male genital somite with 1 posterolateral spine as leg 6 ······························································· H. ventriplanus 16 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

3. Hemicyclops ctenidis Ho and Kim, 1990 (Figs. 3-5)

Bit-geom-mul-byeo-ruk-but-i (빗검물벼룩붙이)

Hemicyclops ctenidis Ho and Kim, 1990, p. 231, figs. 1-5.

Female: Body (Fig. 3A) with widely separated tergal plates in prosome. Body length 1.01 mm and greatest width of prosome 0.39 mm. Urosome (Fig. 3B) 5-segmented. Fifth pedigerous somite 133×78 μm. Anterior part of this somite usually concealed by preceding somite in dorsal view. Genital double-somite quadrate, 101×102 μm, anteriorly with small, pointed lateral swelling. Ven- trally this somite divided by incomplete transverse suture at anterior two-fifths. Genital aperture situated antero-laterally. Three postgenital somites 63 ×88, 53×79, and 60×69 μm, respectively. Anal somite distally with spinules on each side. Caudal ramus gradually narrower distally, 51×25 μm, with numerous hairs on dorsal surface. Outer margin at proximal fourth with 1 minute spinule. Lateral seta plumose distally. Innermost seta shortest. Outer long terminal seta with similar spinules on outer margin in distal 3-fourths. Distal end of ramus with row of minute spinules near bases of long terminal setae. Egg sac 457 ×157 μm, reaching beyond caudal ramus. Spermato- phore (Fig. 3C) glove-shaped.

Rostrum triangular and weakly pointed ventrally. Antennule (Fig. 3D) about 260 μm long and 6-segmented; first segment with minute spinules on anterior margin and a number of setules on dorsal surface. Armature formula 4, 14, 9, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Plumose setae: 1 each on second and third, 2 on fourth, 1 on fifth and 4 on terminal segment. An- tenna (Fig. 3E) 4-segmented; armature formula 1, 1, 4, and 7. Third segment spinulose on anterior margin; anterodistal corner protruded; 2 spines on this corner unequal in size, both with a row of spinules on anterior margin and a subterminal setule; seta next to these spines longest among 4 elements, naked and curved near distal 1-third; fourth element spinulose. Distal segment 1.5 times as long as wide, with 4 long, annulated setae, 1 barbed seta, and 2 setae with spinules. Labrum ventrally with transverse row of hairs on either side and several subterminal hairs. Dor- sal surface with transverse row and a subsidiary row of denticles on each side. Posterior border armed with fine spinules arranged irregularly. Mandible (Fig. 3F) with 2 stout elements and 2 equal-sized, spinulose spines. Basal segment with 2 groups of ventral hairs. Paragnath (Fig. 3H) elongate, with rows of hairs posteriorly and dense tift of hairs apically. Maxillule (Fig. 3G) bilobate with 3 setae on smaller lobe and 5 setae on larger lobe. Maxilla (Fig. 3I) with 2 large, barbed spines and minute, naked spine on basal segment; second segment with 4 elements, 1 element thick and bifurcate distally. Maxilliped (Fig. 4A) with armature 2, 2, 0, 5. First segment with numerous spinules on outer distal corner and 3 groups of hairs on median surface. Legs 1-4 with endopods longer than exopods. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0; I-1; II, 6; enp. 0-1; 0-1; I, 5 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, 6; enp. 0-1; 0-2; III, 3 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 6; enp. 0-1; 0-1; III, 2

Spines of first endopodal segment of legs 1-4 typically with pectinate terminal part. Leg 1 (Fig. 4B) with blunt, barbed spines on inner surface of basis. This area of basis bearing row of spinules in legs 2 (Fig. 4C) and but naked in leg 4 (Fig. 4D). Ventral margin of intercoxal plate in leg 1 with hairs, but replaced with rows of spinules in legs 2-4. All outer spines on leg 1 exopod with subterminal setule, but in legs 2-4 such setule present only on distal spine of third segment.

Leg 5 (Fig. 4E) 2-segmented. Protopod with seta and spinules along outer distal corner. exopod 1.5 Cyclopoida: Clausidiidae: Hemicyclops 17

A B D

E G

Fig. 3. Hemicyclops ctenidis, female. A. habitus, dorsal; B. urosome, ventral; C. spermatophore; D. antennule; E. antenna; F. mandible; G. maxillule; H. paragnath; I. maxilla. Scales: A=0.1 mm, B-D, F=0.05 mm, E, G-I=0.02 mm. 18 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

C B

F D

A E

Fig. 4. Hemicyclops ctenidis. female: A, maxilliped; B. leg 1; C. leg 2; D. leg 4; E. left side of first two urosomal somites, dorsal. male: F. leg 5. Scales: 0.02 mm. Cyclopoida: Clausidiidae: Hemicyclops 19

D F

Fig. 5. Hemicyclops ctenidis, male. A. third segment of antennule; B. maxilla; C. maxilliped; D. terminal claw of maxilliped; E. leg 1; F. leg 2. Scales: A, B, D=0.02 mm, C, E, F=0.05 mm. 20 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans times as long as wide, with spinules on both outer and inner margins; terminally bearing 3 spines and 1 seta. Leg 6 represented by 3 minute spinules in genital operculum. Male: Body larger than that of female. Length 1.12 mm and greatest width 0.41 mm. Prosome flatter and broader than that of female. Genital somite as long as wide, 172×175 μm. Four abdominal somites each wider than long, 72×114, 67×103, 56×89, and 63×81 μm, respectively. Rostrum more conspicuous than in female. Antennule same as in female except for number of setae of second segment (Fig. 5A) bearing 15 instead of 14 as in female. Maxilla (Fig. 5B) as in female, but basal segment additionally with patch of spinules on ventral surface. Maxilliped (Fig. 5C, D) with 2 long setae on first segment; second segment with triangular process on inner dorsal area at proximal third, 3 rows of denticles on inner edge, 1 row of spinules on distal half, and 2 medial setae. Third segment very short.claw 0.8 times as long as second segment, with 2 setae and spinifor process near base.

Armature formula of legs as in female, but legs 1 (Fig. 5E) and 2 (Fig. 5F) showing significant sexual dimorphism. Third segment of leg 1 endopod elongate, with 2 groups of spines: 1 group on distal border and another on inner border. Leg 2 endopod strong and long, about 1.6 times as long as exopod; its third segment with 3 smooth, large spines, of which terminal 1 very long, distinctly longer than segment; of 3 setae on segment distal two somewhat transformed. Legs 3 and 4 nearly as those of female. Leg 5 (Fig. 4F) slender. Exopod 2.6 times as long as wide, with naked lateral side; elements on distal end comparatively longer than those in female. Leg 6 represented by unarmed seta on ventro-lateral corner of genital somite.

Distribution: Japan, Korea. Korea: GW, GN, JB. Specimens examined: All copepod specimens were collected from external washings of the polychaete Neanthes japonica (Okuda). 210♀♀, 110♂♂ (type specimens), Namdaecheon, Gangneung,

18.vi.1988; 4♀♀, Hwajinpo, Goseong, Gangwon-do, 19.x.1990; 1♂, Moonpo (Dongjin River),

Dongjin-myeon, Buan, Jeollabuk-do, 28.x.1990; 3717 specimens (including copepodids), Hyang- ho, Jumunjin, Gangwon-do, 16.vi.1991; 2♀♀, 1♂, Geumsong-ri, Namhae I, Gyeongsangnam-do, 22.vii.2001; 20 specimens, tidal flat, Dapo-ri, Geoje I.,.vii.2003; about 150 specimens, type locality, 14.iv.2012; about 200 specimens, type locality, 10.vii.2013. Remarks: This species is host-specific, living in association with the polychaete Neanthes japonicus (Izuka). It is also distributed in Japan (Itoh, 2003).

4. Hemicyclops gomsoensis Ho and Kim, 1991 (Figs. 6-8)

Gom-so-geom-mul-byeo-ruk-but-i (곰소검물벼룩붙이)

Hemicyclops gomsoensis Ho and Kim, 1991, p. 289, figs. 1-3; Itoh and Nishida, 2007, p. 134, figs. 1-10.

Female: Body (Fig. 6A) 1.45 mm long. Greatest width 0.66 mm. Prosome broad, 1.2 times longer than wide. Last prosomal somite with almost parallel lateral margins, its posterior corners pointed. Urosome (Fig. 6B) 5-segmented. Fifth pedigerous somite fringed with long hairs along posterior border. Genital double-somite complicated, constricted at middle, with 2 kinds of flaps at area Cyclopoida: Clausidiidae: Hemicyclops 21 of constriction, and fringed with fine hairs on posteroventral border. Three free abdominal somites wider than long, gradually shortened and narrowed from anterior to posterior. Anal somite with row of spinules along posterior border. Caudal ramus 2.7 times longer than wide, with fine hairs along inner lateral margin. All setae plumose, but outer lateral seta plumose only on outer edge. Both outer lateral and outermost setae with thorn-like process on outer edge. Dorsal seta articulated at base. Egg sac longer than urosome 683×256 μm, narrower and curved inward anteriorly; eggs about 72 μm in diameter.

Antennule (Fig. 6C) 7-segmented; armature formula of 4, 15, 6, 3, 4 +aesthetasc, 2+aesthetasc, and 7+aesthetasc. Plumose setae: 1 on each of second, fifth and sixth segments, and 3 on terminal segment. Antenna (Fig. 6D) 4-segmented, with armature 1, 1, 4, 7. First segment with a few spinules on outer and inner margins. Second segment with longer spinules on proximal two-thirds of outer margin and minute spinules on distal 1-third of outer margin. Third segment with large spinules on outer mrgin and small spinules near inner distal corner; outer distal corner projected, projection as long as its basal width, with 2 smooth setae, 1 barbed seta and 1 thin, smooth seta.

Fourth segment slightly wider than long, ratio 1 : 1.15.

Labrum (Fig. 6E) dorsal with transverse row of denticles on each side. Posterior border armed with 1 row of denticles and 1 row of fine spinules. Mandible (Fig. 6F) with 2 stout elements and 2 setae. Basal segment with manu hairs on ventral margin and several hairs on dorsal margin.

Paragnath (Fig. 6G) with 1 row of spinules on ventral margin and fine hairs on anterior and apical areas. Maxillule (Fig. 6H) ramified distally, with 8 setae (5+3); 1 seta on smaller ramification enlarged. Maxilla (Fig. 6I) with 2 large, barbed spines and accompanied, small seta; second segment with 3 setae and 1 large, ramified element. Maxilliped (Fig. 7A) with armature formula of 2, 2, 0, 6; first segment with 2 setae and 2 semi-circular rows of spinules; fourth segment with 6 setae, but one of them iriginating from base of second largest seta.

Legs 1-4 with endopods longer than exopods. Leg 1 (Fig. 7B) with inner spine on basis of 73 μm long. Intercoxal plate of leg 1 with row of hairs on ventral margin, but in legs 2-3, replaced by spinules. All spines on first exopod with terminal flagellum. Median terminal seta on third segment of first exopod naked. Outer terminal seta on this segment weakly plumose on outer edge. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0; I-1; I, 7; enp. 0-1; 0-1; I, 5 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 7; enp. 0-1; 0-2; III, 3 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; I, 7; enp. 0-1; 0-2; III, 2 Leg 5 2-segmented. Protopod about as long as wide, with plumose seta on elevated base; outer distal margin spinulated. Exopod (Fig. 7E) 1.7 times longer than wide, with 4 elements on distal margin; both outer margin and distal half of inner margin armed with spinules.

Male: Body (Fig. 7F) almost same as that of female, but with narrower prosome. Length 1.37 mm, and greatest width 0.54 mm. Urosome (Fig. 7G) 6-segmented. Genital somite wider than long, ratio 0.9 : 1. Spermatophore long and curved. First abdominal somite slightly longer than suc- ceeding somite. Third abdominal somite shortest. Caudal ramus as in female. Outer lateral seta plumose on both edges, without proximal spiniform process.

Antennule, antenna, labrum, mandible, paragnath, maxillule as in female. Maxilla (Fig. 8A) different from that of female in having inner seta on second segment replaced by stout process. Max- illiped (Fig. 8B) with 1 long seta on first segment; second segment widely expanded at base, with 1 smaller smooth seta, 1 longer plumose seta, and 3 rows of spines; third segment short and unarmed. Claw slightly longer than second segment, with proximally 2 setae and 1 process.

Leg 1 (Fig. 8C) same as that of female, but without inner spine on basis. Legs 2-5 same as those 22 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B C

D H

Fig. 6. Hemicyclops gomsoensis, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. paragnath; H. maxillule; I. maxilla. Scales: A=0.2 mm, B, C=0.1 mm, D, E, H, I=0.05 mm, F, G=0.02 mm. Cyclopoida: Clausidiidae: Hemicyclops 23

B C

F A

Fig. 7. Hemicyclops gomsoensis. female: A, maxilliped; B. leg 1; C. leg 2; D. leg 4; E. leg 5 exopod. male: F. habitus, dorsal; G. urosome, ventral. Scales: A-E=0.05 mm, F=0.2, G=0.1 mm. 24 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B C

Fig. 8. Hemicyclops gomsoensis, male. A. maxilla; B. maxilliped; C. leg 1. Scales: 0.05 mm.

of female. Leg 6 represented by 1 outer smooth seta and 1 shorter, barbed inner seta on posteriolat- eral corner of genital somite.

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

Specimens examined: The hosts are all Upogebia major de Haan. 5♀♀, 5♂♂ (type specimens) from burrows on mud flat, Gomso, Buan-gun, Jeollabuk-do, 22.x.1990; 1♀, tidal flat, Wando I.,

.iv.1991; 25 specimens, tidal flat, Namhae I. (west side), 7.ii.1993; 2♀♀, 2♂♂, Mangdeok (mouth of Seomjin River), 6.ii.1993; 10♀♀, 5♂♂, tidal flat, Hampyeong Bay, Jeollanam-do, 12.vii.1998; numerous specimens, tidal flat, Janghang, Seocheon-gun, Choongcheongnam-do, 14.viii.1998; numerous specimens from burrows of Upogebia major, tidal flat, Moochangpo, Choongcheongnam-do, 6.v.2000; 20♀♀, 10♂♂, tidal flat, Yoolpo, Geoje I., Gyeongsangnam-do, 30.vii.2003; about 150 specimens from burrows on tidal flat, Geumsong-ri, Namhae I, 21.x.2010; 65♀♀, 45♂♂ from burrows of Upogebia major, Daebu I., Ansan, Gyeonggi-do, 17.v.2012.

Remarks: This species closely resembles H. parilis Moon and Kim, 2010 (Moon and Kim, 2010).

Both are almost siblings and have 2 pairs of dorsolateral flaps (anterior and posterior) on the genital double-somite in the female. The anterior flap is a device holding the girdle of spermatophore, positioning just lateral to the copulatory pore. In both species the anterior flap is weakly bilobed, with the outer and inner lobes. This anterior flap is not so different between the two species. The posterior flap is, however, markedly differs between the two; in H. gomsoensis it is positioned dorsolaterally, with blunt posterior apex, whereas, that of H. parilis is positioned laterally, with a pointed posterior apex. The shapes of the spermatophores of two species are invariable within a species and clearly different between the two species. The spermatophore of H. gomsoensis is domino-shaped (Fig. 19I) and, thus, contrasted to that of H. parilis (Fig. 19E), which is L-shaped, with a prominent process near the opening. Ho and Kim (1991) recorded the ratio of the length to width of caudal ramus in Cyclopoida: Clausidiidae: Hemicyclops 25

the female of H. gomsoensis as 2.7 : 1. But in female specimens examined by Moon and Kim (2010), it was measured as 3.11-3.43 : 1, whereas that of H. parilis was measured as 2.44-2.93 : 1. This species was originally discovered from the burrows of the crab Macrophthalmus japonicus De Haan in the Yellow Sea. But this discovery from the burrows of that crab turned out to be accidental and its true host being Upogebia major (De Haan). Thousands of specimens can be collected from burrows at one site of mud flat where U. major lives. According to Itoh and Nishida (2007), H. gomsoensis is the most common species of the genus in Japan.

5. Hemicyclops japonicus Itoh and Nishida, 1993 (Figs. 9-11)

Tteo-dol-i-geom-mul-byeo-ruk-but-i (떠돌이검물벼룩붙이)

Hemicyclops japonicus Itoh and Nishida, 1993, p. 149, figs. 1-4.

Female: Body (Fig. 9A) 1.31 mm long. Prosome 765 μm long, with greatest width 610 μm. Sec- ond and third pedigerous somite fringed with membrane along posterior margin. Fourth pedigerous somite with uneven posterolateral areas. Urosome (Fig. 9B) 5-segmented. Fifth pedigerous somite 218 μm wide, tapering anteriorly, with several setules on each lateral side of posterodorsal margin. Genital double-somite 213×227 μm, with deep lateral incision at anterior third of somite length; posterior 0.7 part of somite rhomboidal, expanded in middle, with tooth-like point at lateral apex; genital aperture small, located dorsolaterally at anterior region. Three free abdominal somites 80×138, 65×120, and 60×108 μm, respectively. Anal somite with row of fine spinules along posteroventral margin. Caudal ramus (Fig. 9C) 79×44 μm (ratio 1.80 : 1), with setules on posterior half of medial margin and 6 setae; lateral and outermost terminal setae naked and tipped with setule. Dorsal seta naked; other 3 setae pinnate.

Antennule (Fig. 9D) 7-segmented and 354 μm long; armature formula 4, 14, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; setae naked, except for following plumose setae: 1 each on fifth and sixth and 4 on last segments. Antenna (Fig. 9E) 4-segmented, with armature formula 1, 1, 4, and 7; first segment the longest; third segment densely spinulose on medial margin, with distinctly produced mediodistal angle; last segment 24×21 μm, slightly longer than wide.

Labrum (Fig. 9F) broad, with spinulose posterior margin. Mandible (Fig. 9G) distally with 2 plate-like elements and 2 setae. Paragnath (Fig. 9H) densly setulose and articulated distally. Max- illule (Fig. 10A) distally bilobed, with 3 setae on small lobe and 5 setae on larger lobe. Maxilla (Fig. 9I) 2-segmented; basal segment with 2 large medial, spiniform seta and 1 small subsidiary seta inserted at base of one of larger setae; distal segment with 2 spinulose setae, 1 spinulose spine, and terminaled by large spinose process. Maxilliped (Fig. 10B) 4-segmented; first segment with 2 large setae on medial margin; second segment produced medially at proximal two-thirds , 2 large spinulose setae on produced region and longitudinal row of fine setules distal to setae; third segment small and unarmed; fourth segment also small bu armed with 2 spinulose spines (terminal one of them with 2 small setae at base) and 3 naked setae. Legs 1-4 with 3-segmented rami. Exopods shorter than endopods. Intercosal plate of leg 1 bearing setules on distal margin, but in leg 2-4 these setules replaced by spinules. Basis of leg 1 with pointed inner distal corner and inner distal spine of 69 μm long. Two medial setae on third endopodal segment of leg 4 spiniform. Armature formula of legs 1-4 as follows: 26 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

H D

Fig. 9. Hemicyclops japonicus, female. A. habitus, dorsal; B. urosome, dorsal; C. distal part of abdomen, ventral; D. antennule; E. antenna; F. labrum; G. mandible; H. paragnath; I. maxilla. Scales: A=0.2 mm, B=0.1 mm, C-F, I=0.05 mm, G, H=0.02 mm. Cyclopoida: Clausidiidae: Hemicyclops 27

A B C

E D

Fig. 10. Hemicyclops japonicus. female: A. maxillule; B. maxilliped; C. leg 1; D. leg 2; E. leg 4; F. leg 5. male: G. maxilla. Scales: A, B=0.02 mm, C-G=0.05 mm. 28 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

Fig. 11. Hemicyclops japonicus, male. A. habitus, dorsal; B. urosome, ventral; C. maxilliped; D. leg 1 endopod. Scales: A=0.2 mm, B=0.1 mm, C, D=0.05 mm.

Leg 1: coxa 0-1; basis 1-I; exp. I-0; I-1; I, 3, 4; enp. 0-1; 0-1; I, 2, 3 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 2, 5; enp. 0-1; 0-2; I, II, 2 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; I, 2, 5; enp. 0-1; 0-2; I, II, 2

Leg 5 (Fig. 10F) 2-segmented; protopod not articulated from somite, with 1 outer distal seta; exopod 128×54 μm (ratio 2.37 : 1), with spinules on outer and lateral margins, and armed distally with 3 spines and 1 seta. Leg 6 not discernible.

Male: Body (Fig. 11A) 1.18 mm long. Prosome narrower than that of female. Third pedigerous somite with angular posterolateral corners. Urosome (Fig. 11B) 6-segmented. Fifth pedigerous somite 225 μm wide. Genital somite nearly circular, 125×160 μm. Four abdominal somites 98×122,

85×108, 50×100, and 45×94 μm, respectively. Caudal ramus 73×38 μm (ratio 1.92 : 1) and armed as in female.

Antennule with 2 additional setae, 1 each on third and fourth segments (thus armature formula 7 and 4 on these segments). Antenna as in female. Labrum, mandible, and maxillule also as in female. Maxilla (Fig. 10G) on distal segment with 2 setae, 1 ramified spine, and 1 stout, claw-like Cyclopoida: Clausidiidae: Hemicyclops 29

process bearing distally with granules and 1 proximal tubercle on medial side. Maxilliped (Fig. 11C) consisting of 3 segments and terminal claw; first segment with 1 pinnate seta; second segment strongly expanded proximally, with 2 small setae and 2 rows of spinules on medial side; third segment small and unarmed; terminal claw elongate, gently curved, proximally with 2 small setae and 1 small spine.

Leg 1 lacking inner distal spine on basis (Fig. 11D). Legs 2-4 as in female. Leg 5 consisting 1 seta on somite and exopod (protopod completely fused with somite); exopod 127×66 μm. Leg 6 represented by 1 spine genital operculum (Fig. 11B).

Distribution: Japan, Korea. Korea: JB, GG. Specimens examined: 4♀♀, 3♂♂ from invertebrate burrows, tidal flat, Jakyak I., Incheon, 7.x.1998; 1♀, invertebrate burrows on tidal flat, Gyehwa I., Buan-gun, Jeollabuk-do, 5.viii.2000. Remarks: This species is unknown of its host, but a crustacean or a polychaete is presumed to be the host, as in general of other species of the genus. A strong lateral constriction on the genital double-somite is a distinguishing feature of this species.

6. Hemicyclops membranatus Moon and Kim, 2010 (Figs. 12-14)

Mak-geom-mul-byeo-ruk-but-i (막검물벼룩붙이)

Hemicyclops membranatus Moon and Kim, 2010, p. 288, figs. 7-9.

Female: Body (Fig. 12A) with flat prosome and cylindrical urosome. Body length 1.83 mm (1.90- 1.69 mm), based on 4 specimens. Dissected specimen 1.88 mm long. Prosome 1.09 mm long, with Greatest width 812 μm. Prosomal somites becoming distinctly narrowed from anterior to posterior, with rounded anterior and posterior corners. Cephalothorax 560 μm long, much wider than long, with membranous fringe along anterior and anterolateral margins. Urosome (Fig. 12B) 5-segmented. Fifth pedigerous somite 273 μm wide. Genital double-somite 269×307 μm, with distinct anterior expansion and narrower posterior half (179 μm across this area); genital aperture located apically on anterior expansion. Three abdominal somites 127×157, 96×142, and 77×131 μm, respectively. Anal somite (Fig. 12C) with transverse row of small spinules along posteroventral margin. Caudal rami slightly divergent and widely separated from each other; each ramus 109×45 μm

(ratio 2.42 : 1), gradually narrowed posteriorly, with small denticles on posteroventral margin and 7 setae including small outer proximal seta. Inner distal seta pinnate along inner margin. Inner dorsal seta weakly pinnate. Other 4 setae naked.

Rostrum (Fig. 13A) hardly discernible, with faint posterior margin. Antennule (Fig. 12D) 425 μm long and 7-segmented, with armature formula of 5, 15, 7, 4, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. First 2 segments distinctly wider than remaining 5. Aesthetascs on fifth to seventh segments very thin. Antenna (Fig. 12E) 4-segmented. First segment with 1 large seta. Second segment with smaller seta not reaching distal border of segment. Third segment with spinules along inner margin and 4 elements consisting of 1 small subdistal seta, 1 spiniform seta tipped by setule, 1 strong claw, and 1 ordinary seta. Terminal segment 41×18 μm, with setules on outer margin and 7 setae, inner 4 of latters claw-like. 30 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Labrum (Fig. 13B) with spinules along posterior margin. Mandible (Fig. 13C) with tubercle on ventral side, membranous fringe on dorsal margin, and 4 distal elements consisting of 2 spinulated spines and 2 pinnate setae. Paragnath (Fig. 13D) elongated, with numerous setules. Maxillule (Fig. 13E) distally bilobed and armed with 5 setae on broader lobe and 3 setae on narrower lobe, middle one of latters distinctly larger than others. Maxilla (Fig. 13F) 2-segmented. Proximal segment large, distally with 2 large and 1 small setae. Distal segment armed with 1 spiniform process, 1 spiniferous spine, and 2 spiniferous setae. Maxilliped (Fig. 13G) 4-segmented. First segment with 2 unequal, pinnate setae. Second segment with convex inner margin, 2 inner setae (1 naked seta and 1 smaller, unilaterally pinnate seta) and 2 longitudinal rows of long setules on inner side. Third segment small and unarmed. Terminal segment armed with 5 elements consisting of 3 naked setae, 1 small spine and 1 enlarged, pinnate spine.

Leg 1 (Fig. 13H), leg 2 (Fig. 13I), leg 3 and leg 4 (Fig. 14A) with 3-segmented rami. Posterior margin of intercoxal sclerite with setules in leg 1, but with spinules in legs 2-4. Some of outer spines on exopod tipped by setule. Outer seta on basis of legs 1-4 pinnate and large. Inner spine on basis of leg 1 thick but short. Outer spine on first exopodal segment of leg 1 setiform. Two inner setae on third endopodal segment of leg 4 pinnate proximally but spinulated distally. Legs 2 and 3 identical in armature and ornamentation. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-I; exp.I-0; I-1; II, 2, 4; enp. 0-1; 0-1; I, 2, 3 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; II, I, 3 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-2; II, I, 2

Leg 5 (Fig. 14B) consisting of protopod and exopod. Protopod longer than wide and armed with

1 distal seta. Exopod slightly curved, 125×61 μm (ratio 2.05 : 1), armed with 3 spines and 1 pinnate seta, and ornamented with large setules on proximal half of outer margin and distal part of inner margin and spinules near base of spines. Leg 6 not seen.

Male: Body (Fig. 14C) 1.73 μm long. Greatest width 632 μm. Prosome spatulate and 959 μm long. Cephalothorax fringed with membrane along anterior and anterolateral margins, as in female. Urosome (Fig. 14D) 6-segmented. Genital somite 128×203 μm, narrower than fifth pedigerous somite. Four abdominal somites 124×156, 99×136, 62×124, and 64×119 μm, respectively.

Caudal ramus 104×40 μm (ratio 2.60 : 1). Rostrum, Antennule and antenna not different from those of female. Labrum not observed.

Mandible, paragnath and maxillule as in female. Maxilla (Fig. 14E) different from that of female; its distal segment transformed to strong claw bearing 3 subdistal setae (one of them small and hardly visible). Maxilliped (Fig. 14F) consisting of 3 segments and terminal claw. First segment with 1 large pinnate seta. Second segment greatly expanded proximally with 3 longitudinal rows of spinules and 2 inner setae (distal one of latters unilaterally pinnate, proximal 1 very small and hardly visible). Small third segment unarmed. Terminal claw weakly curved and accompanied by 2 small proximal setae. Legs 1-4 not different from those of female in shape and armature. Leg 5 consisting of protopod and exopod; protopod fused to fifth pedigerous somite. Exopod different in shape from that of female, not curved, 104×58 μm (ratio 1.79 : 1), with pointed process near base of first and second outer spines. Leg 6 represented by 1 spine on genital operculum.

Distribution: Korea. Korea: CN. Specimens examined: 4♀♀, 1♂ from washing of an unidentified polychaete from intertidal sands at Sinjin Island, Taean, Chooncheongnam-do (36°41′00″N, 126°08′18″E), 09.x.2010, collected Cyclopoida: Clausidiidae: Hemicyclops 31

A B

Fig. 12. Hemicyclops membranatus, female. A. habitus, dorsal; B. urosome, dorsal; C. posterior part of urosome, ventral; D. antennule; E. antenna. Scales: A=0.2 mm, B=0.1 mm, C, D=0.05 mm, E=0.02 mm. 32 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A C

B F E

Fig. 13. Hemicyclops membranatus, female. A. rostral area ventral; B. labrum; C. mandible; D. paragnath; E. maxillule; F. maxilla; G. maxilliped; H. leg 1; I. leg 2. Scales: A=0.1 mm, B, F, H, I=0.05 mm, C-E, G=0.02 mm. Cyclopoida: Clausidiidae: Hemicyclops 33

B G

C E

Fig. 14. Hemicyclops membranatus. female: A. leg 4; B. leg 5. male: C. habitus, dorsal; D. urosome, ventral; E. maxilla; F. maxilliped; G. leg 5 exopod. Scales: A, B, E-G=0.05 mm, C=0.2 mm, D=0.1 mm. 34 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans by S.Y. Moon. Remarks: This species may be typified by 2 outstanding features: 1) the presence of the membranous fringe along the margins of cephalothorax and 2) a claw on the third segment of the antenna. The first character has not been reported in the genus Hemicyclops and is, therefore, a unique feature of H. membranatus. In the presence of claw on the third antennal segment, This species is com- parable to H. cylindraceus Pelseneer, 1929 associated with a bivalve in West Europe and H. livingsto- ni (T. Scott, 1894) known from a plankton sample in West Africa (Moon and Kim, 2010).

7. Hemicyclops nasutus Moon and Kim, 2010 (Figs. 15, 16)

Ko-geom-mul-byeo-ruk-but-i (코검물벼룩붙이)

Hemicyclops nasutus Moon and Kim, 2010, p. 285, figs. 5, 6.

Female: Body (Fig. 15A) moderately narrow and 1.63 mm long. Greatest width 611 μm measured across cephalothorax. Prosome 850 μm long, and cephalothorax 483 μm long. Cephalotho- rax and 2 following somites fringed with membrane along posterodorsal margin. Urosome (Fig. 15B) 5-segmented. Fifth pedigerous somite 233 μm wide, with small transparent posterolateral process on both sides. Genital double-somite (Fig. 15C) 256 μm long, with prominent anterolateral expansions (244 μm wide across this area), narrow posterior half (147 μm wide across this area), and dorsal stepwise curvature seen evidently in lateral view. Three postgenital somites 111×136, 83×133, and 75×118 μm, respectively. Anal somite with minute spines along dorsodistal margin and several larger spinules along ventrodistal margin. Caudal ramus (Fig. 15D) 183×43 μm (ratio

4.26 : 1) narrowest in middle and broader in extremities, with setules distal part of inner margin, 1 small outer proximal setules, and 6 setae. Outer subdistal and distal setae spiniform and tipped by naked setule. Dorsal seta naked. Other 3 distal setae pinnate.

Rostrum hemicircular. Antennule (Fig. 15E) 383 μm long and 7-segmented, with armature formula of 5, 15, 6, 4, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. One of setae pinnate on first, fifth, sixth, and seventh segments. Aesthetascs on 3 distal segments recurved. Antenna (Fig. 15F) 4-segmented with armature formula of 1, 1, 4, and 7. First segment with several setules on outer margin. Second segment with long setules proximally and spinules distally on outer margin. Third segment spinules of various sizes on outer margin and row of small spinules on inner margin. Fourth segment 42×33 μm, 1.27 times as long as wide.

Labrum (Fig. 15G) with spinules of various sizes posteriorly. Mandible (Fig. 15H) with smooth margins and 4 distal elements: claw-like element, spiniferous plate, and 2 pinnate setae. Parag- nath (Fig. 15I) lobate, with spinules and setules. Maxillule (Fig. 16A) distally ramified, with 5 setae on broader ramus and 3 setae on narrower ramus. Maxilla (Fig. 16B) 2-segmented. Proximal segment row of small setules proximally and 2 large setae distally, one of these 2 setae accompanied by 1 setule proximally; distal segment with 2 setae, 1 spine and 1 spiniform branched process; one of branches bearing 1 barb. Maxilliped (Fig. 16C) 4-segmented. First segment with 2 large inner setae. Second segment with 2 inner setae of unequal sizes and longitudinal row of large setules along distal part of inner side. Third segment small and unarmed. Fourth segment armed with 2 spiniform elements and 3 naked setae. Legs 1-4 with 3-segmented rami, with exopods distinctly longer than endopods. Armature of Cyclopoida: Clausidiidae: Hemicyclops 35

C E

F H

I G

Fig. 15. Hemicyclops nasutus, female. A. habitus, dorsal; B. urosome, dorsal; C. genital double somite, left; D. left caudal ramus, dorsal; E. antennule; F. antenna; G. labrum; H. mandible; I. paragnath. Scales: A=0.2 mm, B, C=0.1 mm, D-I=0.05 mm. 36 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B D

C A

E F

Fig. 16. Hemicyclops nasutus, female. A. maxillule; B. maxilla; C. maxilliped; D. leg 1; E. leg 2; F. leg 4; G. leg 5 exopod. Scales: 0.05 mm. Cyclopoida: Clausidiidae: Hemicyclops 37 these legs as follows: Leg 1: coxa 0-1; basis 1-I; exopod I-0; I-1; II, 2, 4; endopod 0-1; 0-1; I, 2, 3 Legs 2 and 3: coxa 0-1; basis 1-0; exopod I-0; I-1; II, II, 5; endopod 0-1; 0-2; I, II, 3 Leg 4: coxa 0-1; basis 1-0; exopod I-0; I-1; I, II, 5; endopod 0-1; 0-2; I, II, 2 Basis of leg 1 with pointed process on inn side of posterior margin near inner spine; inner 42 μm long. Four outer spines on leg 1 exopod and distal spine on exopod of legs 2-4 tipped by setule-like element. Intercoxal plate of leg 1 with setules along posterior margin but those of legs 2-4 replaced by spinules. Two inner setae on terminal segment of leg 4 endopod spiniform, proximally pinnate but spiniferous distally. Leg 5 2-segmented. Protopod segment articulated from fifth pedigerous somite, with 1 pinnate dorsodistal seta. Exopod (Fig. 16G) 120×77 μm (1.56 : 1), with spinules on both outer and inner margin, and 3 spines and 1 seta on distal margin. Leg 6 not seen. Male: Unknown.

Distribution: Korea. Korea: CN.

Specimens examined: 1♀ (holotype) from an unidentified polychaete at the Manripo Beach (36°47′08″N, 126°08′24″E), low intertidal, 12.viii.2010, collected by S.Y. Moon. Remarks: This species is easily distinguishable from other species of the genus from Korea by its elongate caudal rami which are more than 4 times as long as wide, in contrast to less than 3 times as long as wide in other species.

8. Hemicyclops parilis Moon and Kim, 2010 (Figs. 17-19)

Ssok-beol-re (쏙벌레)

Hemicyclops parilis Moon and Kim, 2010, p. 279, figs. 1-3, 4A, B.

Female: Body (Fig. 17A) 1.64 mm long, excluding setae of caudal rami. Greatest width 757 μm across second pedigerous somite. Prosome 942 μm long. Cephalothorax 474×751 μm, with pointed posterolateral corners. Fourth pedigerous somite with notch at posterolateral corners.

Urosome (Fig. 17B) 5-segmented. Fifth pedigerous somite 254 μm wide, with row of long setules along posterodorsal margin. Genital double-somite (Fig. 17C) flat, 250 (measured along mid-dorsal axis)×308 μm, with paired dorsal and lateral flaps and several setules on dorsal surface; dorsal flap branched, bearing minute setules along margin of lateral branch and 1 setule near tip of posterior branch; lateral flap flesh and ended with sharp point. Three postgenital somites 98×204, 77×162, and 62×137 μm. Genital double and first 2 postgenital somites fringed with membrane along posteroventral margin. Anal somite with transverse row of spinules along posteroventral border.

Caudal rami (Fig. 17D) divergent; each ramus 113×42 μm (ratio 2.69 : 1), with setules along inner margin, 1 proximolateral setule and 7 setae; outer subdistal and distal setae spiniform proximally but setiform distally; dorsal seta naked but other 5 setae pinnate. Egg sac (Fig. 17E) 492×223 μm, slightly curved inward, and narrowed anteriorly.

Rostrum hemicircular. Antennule (Fig. 17F) 7-segmented and 433 μm long, with armature formula of 4, 15, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; first segment with minute spinules 38 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

on ventral surface. One of setae on second, fifth and sixth segments pinnate. Antenna (Fig, 17G) 4-segmented, with armatrue formula of 1, 1, 4, and 7. First segment with setules on outer and inner margins. Second segment with setules and minute spinules on outer margin. Third segment with large and smaller spiunules on outer margin and row of minute setules on inner margin; outer distal corner projected. Fourth segment 256×282 μm, slightly wider than long, with 2 rows of minute setules near inner margin.

Labrum (Fig. 18A) with transverse row of denticles on dorsal surface and along posterior margin.

Mandible (Fig. 18B) with setules on distal half of posterior margin and 4 distal elements consisting of 1 stout claw-like element, 1 spiniferous plate and 2 pinnate setae. Paragnath (Fig. 17H) bi-articulated, with larger proximal article bearing spinules and minute setules and small distal article bearing minute spinules. Maxillule (Fig. 18C) distally ramified distally, with 5 setae on broader ramus and 3 setae on narrower ramus. Maxilla (Fig. 18D) 2-segmented. Proximal segment row of small setules proximally and 2 large setae distally, one of these 2 setae accompanied by 1 setule proximally; distal segment with 2 setae and 2 spines; one of spines ramified, each ramus bearing 1 and 2 barbs. Maxilliped (Fig. 18E) 4-segmented. First segment with 2 large inner setae. Second segment with 2 large inner setae and longitudinal row of minute setules along inner side. Third segment small and unarmed. Fourth segment armed with 2 spiniferous spiniform elements and 3 naked setae; shorter one of spiniform elements accompanied by proximal accessory seta.

Legs 1-4 (Figs. 18F-H, 19A) with 3-segmented rami, with exopods distinctly longer than endopods. Armature of these legs as follows: Leg 1: coxa 0-1; basis 1-I; exopod I-0; I-1; I, 3, 4; endopod 0-1; 0-1; I, 2, 3 Leg 2: coxa 0-1; basis 1-0; exopod I-0; I-1; II, 2, 5; endopod 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exopod I-0; I-1; II, II, 5; endopod 0-1; 0-2; I, II, 3 Leg 4: coxa 0-1; basis 1-0; exopod I-0; I-1; I, II, 5; endopod 0-1; 0-2; I, II, 2 Inner spine on basis of leg 1 67 μm long. Outer seta on basis of leg 1 distinctly larger than those of legs 2-4. Three outer spines on leg 1 exopod tipped by setule-like element. Intercoxal plate of leg 1 with setules along posterior margin but those of legs 2-4 replaced by spinules. Two inner setae on terminal segment of leg 4 endopod spiniform, proximally pinnate but spiniferous distally. Leg 5 2-segmented. Protopod articulated from fifth pedigerous somite, with 1 pinnate dorsodistal seta. Exopod (Fig. 19B) 160×81 μm (1.98 : 1), with spinules on both outer and inner margin, and 3 spines and 1 seta on distal margin. Leg 6 not seen.

Male: Body (Fig. 19C) resembling in form that of female and 1.56 mm long. Greatest width 640 μm across second pedigerous somite. Prosome 862 μm long. Cephalothorax 492×634 μm. Uro- some (Fig. 19D) 6-segmented. Fifth pedigerous somite 265 μm wide. Genital somite nearly circular, 252×274 μm, slightly wider than long. Four abdominal somites 100×150, 95×129, 43×118, and

65×111 μm, respectively. Caudal ramus 100×43 μm (2.33 : 1), slightly shorter than that of female.

Spermatophore (Fig. 19E) L-shaped, 288 μm long along middle axis, with projection proximally near opening. Rostrum as in female. Antennule with 1 additional seta on third and fourth segments, at places of dots in Fig. 17F. Antenna as in female.

Labrum, mandible, paragnath, and maxillule similar to those of female. Maxilla (Fig. 19F) with one of spines on distal segment transformed to stout claw. Maxilliped (Fig. 19G) composed of 3 segments and terminal claw. First segment with 1 large seta. Second segment greatly expanded proximally and gradually narrowed distally, with 2 setae near middle of inner side, and 1 row of minute spinules and 2 rows of larger spinules along inner margin. Third segment small and unarmed. Terminal claw nearly as long as 2 distal segments, proximally with 1 seta, 1 spiniform pro- Cyclopoida: Clausidiidae: Hemicyclops 39

A F

D C

G CP

Fig. 17. Hemicyclops parilis, female. A. habitus, dorsal; B. anal somite and caudal rami, ventral; C. egg sac; D. antennule; E. antenna; F. labrum; G. mandible; H. paragnath. Scales: A=0.2 mm, B, E-H=0.05 mm, C, D=0.1 mm. 40 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A F

C E D

H G

Fig. 18. Hemicyclops parilis, female. A. labrum; B. mandible; C. maxillule; D. maxilla; E. maxilliped; F. leg 1; G. leg 2; H. leg 4. Scales: 0.05 mm. Cyclopoida: Clausidiidae: Hemicyclops 41

B A

C H

Fig. 19. Hemicyclops parilis. female: A. leg 4; B. leg 5 exopod. male: C. habitus, dorsal; D. urosome, ventral; E. spermatophore; F. maxilla; G. maxilliped; H. leg 5 exopod; I. spermatophore of Hemicy- clops gomsoensis. Scales: A, B, E-I=0.05 mm, C=0.2 mm, D=0.1 mm. 42 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans cess, and 1 minute setule. Leg 1 lacking inner distal spine on basis, otherwise similar to that of female. Legs 2-4 as in female. Leg 5 with proximal segment completely fused to fifth pedigerous somite, leaving only 1 dorsal seta. Exopod 120×57 μm (2.11 : 1), narrower than distal segment female leg 5. Leg 6 represented by 2 spines on genital operculum.

Distribution: Korea. Korea: GN. Specimens examined: 6♀♀, 6♂♂ from crustacean burrows on intertidal mud flat, Namhae Is- land (34°21′56″N, 126°43′57″E), 21.x.2010, collected by S.Y. Moon. Remarks: This species is very similar to H. gomsoensis Ho and Kim, 1991. The morphological differences between two species are mentioned in the remarks of H. gomsoensis.

9. Hemicyclops saxatilis Ho and Kim, 1991 (Figs. 20, 21)

Dol-geom-mul-byeo-ruk-but-i (돌검물벼룩붙이)

Hemicyclops saxatilis Ho and Kim, 1991, p. 294, figs. 4-6.

Female: Body (Fig. 20A) with moderately broad prosome. Body length 2.04 mm and greatest width 0.79 mm. second to fifth pedigerous somite fringed with transparent membrane. Urosome (Fig. 20B) 5-segmented. Genital double-somite 1.25 times longer tha wide, with 2 recurved, leaf- like extensions on both anterior lateral sides. Free abdominal somites equal in width, but decrease in length toward posterior somite. Genital double-somite and first 2 abdominal somites fringed with transparent membrane on posterior border. Anal somite 0.52 times longer than wide, with transverse row of fine spinules on posteroventral and posterolateral borders. Caudal ramus short, 1.27 times longer than wide, with hairs on inner lateral margin and minute spinules on posteroventral margin. All setae plumose except for dorsal seta which naked and articulate at base. Outer- most terminal seta with proximal process on outer edge.

Antennule (Fig. 20C) with armature formula of 4, 14, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Plumose setae: 1 each on second and fourth to sixth segments, and 2 on seventh segment. Antenna (Fig. 20D) 4-segmented. First segment with 1 outer distal seta and spinules on both outer and inner sides. Second segment with 1 barbed seta on outer median side, longer spinules on outer proximal area and rows of short spinules on inner side; outer distal area naked. Third segment with short but broad outer distal corner, bearing 1 barbed, 1 weakly plumose and 2 naked setae; outer margin armed with longer spinules on distal half and short spinules on proximal half; posterior area with 1 row of minute spinules. Fourth segment shorter than wide, ratio 0.9 : 1, with 7 setae, one of which is recumbent and barbed.

Labrum (Fig. 20E) with dorsal row of denticles on both sides. Posterior margin fringed with 1 row of denticles and 1 row of small spinules. Mandible (Fig. 20F) with 2 stout elements and 2 setae. Paragnath (Fig. 20G) spinulose, not articulated, with narrow distal region. Maxillule (Fig. 20H) distally bilobed, with 5 setae on leger lobe and 3 setae on smaller lobe. Maxilla (Fig. 20I) with first segment bearing 2 setae and 1 small, accompanied seta; second segment with 4 unequal elements.

Maxilliped (Fig. 21A) with first segment bearing 2 setae; second segment with 2 setae and 1 row of Cyclopoida: Clausidiidae: Hemicyclops 43

A B

I G

Fig. 20. Hemicyclops saxatilis, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. paragnath; H. maxillule; I. maxilla. Scales: A=0.5 mm, B=0.2 mm, C, D=0.1 mm, E, F, H, I=0.05 mm, G=0.02 mm. 44 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B C

D F E

Fig. 21. Hemicyclops saxatilis. female: A. maxilliped; B. leg 1; C. leg 2; D. leg 4; E. leg 5. male: F. habitus, dorsal; G. maxilliped; H. leg 6. Scales: A, G, H=0.05 mm, B-E=0.1 mm, F=0.5 mm. Cyclopoida: Clausidiidae: Hemicyclops 45 long spinules on inner distal side; third segment very short and unarmed; fourth segment with 1 strong, claw-like seta, 1 thick basally ramified seta.

Leg 1 (Fig. 21B) with inner spine of 86 μm long on basis. Intercoxal plate on ventral edge and outer ventral margin of basis fringed with hairs. In legs 2-4 these areas bearing spinules (Figs 21C, D). All spines on exopod with flagellum. Legs -1 4 with armature as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0: I-1; II, 6; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; IV, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-2; IV, 1

Leg 5 (Fig. 21E) 2-segmented. Protopod as long as wide, with hairs on outer margin and spinules on outer distal angle. Dorsal seta plumose and shorter than second segment. Exopod 1.55 times longer than wide, with distall 3 broad spines and 1 seta; inner spine about twice longer than ther other 2; both outer and inner margin with spinules.

Male: Body (fig. 21F) 1.66 mm long. Greatest width 0.66 mm. Genital somite 1.33 times wider than long. Abdominal somites and caudal ramus like those of female. Antennule, antenna, labrum, mandible, and maxillule as in female. Maxilla without sexual dimorphsm. Maxilliped with 2 long setae on first segment; second segment broad at base, with 3 row of spines and 2 long plumose setae, of which proximal 1 curved upward; third segment short and unarmed; claw tapering and sharply pointed at tip, with 1 proximal seta. Legs 1-4 as in female. Leg 5 exopod 1.70 times longer than wide, slender than that of female. In- ner margin of second segment slightly concave and outer margin convex. Leg 6 (Fig. 21H) consisting of 1 smooth spine inserted on protrusion at posterodistal corners of genital somite.

Distribution: Korea. Korea: GW, GB, JJ, JB.

Specimens examined: 8♀♀, 2♂♂ (type specimens) from crab burrows of Pilumnopeus ma- kianus (Rathbum) on rocky shore in low intertidal zone at Gyeokpo, Buan-gun, Jeollabuk-do,

26.vii.1990; 3♀♀ from washings of the sponge Cliona celata (as a bycatch of fisheries), Sacheon Port, Gangneung, Gangwon-do, 10.iv.1993; 4♂♂, invertebrate burrows on tidal flat, Jakyak I., Incheon, 29.ix.1996; 1♀, type locality, invertebrate burrows on rocks, type locality, 28.x.2000; 1♀, 2♂♂ from invertebrate burrows, intertidal, Seogwipo Port, 19.viii.2005; 3♀♀, 1♂ from invertebrates epibenthic on walls of Yangpo Port, Pohang, Gyeongsangbuk-do, 24.viii.2007; 7♀♀, 4♂♂, invertebrate burrows on rocks, Choonjangdae, Choongcheongnam-do, 24.vi.2012; 4♀♀ from epibenthic invertebrates on walls of Yangpo Port, Pohang, Gyeongsangbuk-do, 22.viii.2012. Remarks: This species was originally discovered on the coast of the Yellow Sea. Subsequently it has been collected from all other coasts of Korea. Although the type specimens were collected in burrows occupied by small crabs, its true host is uncertain.

10. Hemicyclops spinulosus Itoh and Nishida, 1998 (Figs. 22-25)

Sil-cham-gaet-ji-reong-i-beol-re (실참갯지렁이벌레)

Hemicyclops spinulosus Itoh and Nishida, 1998, p. 85, figs. 1-4; 2008, p. 189, figs. 1-13.

Female: Body (Fig. 22A) moderately broad. Mean body length 2.11 mm (1.78-2.70 mm), based 46 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans on 55 specimens. Dissected specimen 2.67 mm long. Greatest width 1.05 mm. Prosome 1.29 mm long, consisting of cephalothorax and 3 pedigerous somites. Prosomal somites with pointed posterolateral corners. Urosome (Fig. 22B) 5-segmented. Fifth pedigerous somite 465 μm wide. Geni- tal double-somite characteristically trapezoid, 418×552 μm; genital apertures located ventrally (Fig. 22C). Three free abdominal somites 276×247, 171×198, and 125×198 μm, respectively. Anal somite (Fig. 22D) with row of spinules along posteroventral border. Caudal ramus (Fig. 22D) weakly tapering, 187×72 μm (ratio 2.60 : 1), with 7 setae, including rudimentary outer seta (seta I); longest inner one of 2 median terminal setae 1.04 mm long; distal outer seta (seta II) located distal third of ramus; dorsal seta (seta VII) pinnate, but other setae naked or finely spinulose. Egg sac (Fig. 22E) slightly curved, variable in size, and slightly broadened distally. Spermatophore (fig. 22F) about 372×132 μm, nearly haricot-shaped.

Rostrum as sclerotized rudiment. Antennule (Fig. 22G) 7-segmented and gradually narrowing distally; armature formula of 5, 15, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; pinnate setae 1 on second, fifth, and sixth segments each, and 4 on terminal segment; first segment with fine spinules. Antenna (Fig. 22H) 4-segmented; armature formula of 1, 1, 3, and 7; third segment with produced inner distal corner; terminal segment about 32×30 μm, slightly longer than wide; 4 large setae on terminal segment claw-like.

Labrum (Fig. 23A) as in general in the genus. Mandible (Fig. 23B) with 2 thick elements and 2 setae distally. Paragnath (Fig. 23C) lobate, without any articulation. Maxillule (Fig. 23D) distally bilobed, with 5 setae on broader lobe and 3 setae on narrower lobe. Maxilla (Fig. 23E) 2-segmented; basal segment with 2 setae, one of them bearing subsidiary setule proximally; distal segment with 2 thick spines and 2 setae. Maxilliped (Fig. 23F) 4-segmented; first segment spinulose, with 2 large setae on inner margin; second segment with 2 spiniform setae on inner margin; small third segment unarmed; terminal segment with 2 spines and 4 setae, one of latters inserted to proximal region of one of spines.

Legs 1-4 with 3-segmented rami. Leg 1 (Fig. 23G) basis with strong inner distal spine (103 μm long). Outer seta on basis of swimming legs very long (Figs. 23G, H, 24A). Legs 2 and 3 identical in armature formula. Some of spines on third exopodal segment of swimming legs setiform. Inner seta on first endopodal segment of legs -1 4 distally pectinate. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0: I-1; II, 6; enp. 0-1; 0-1; I, 5 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, 1, 5; enp. 0-1; 0-2; I, II, 3 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 1, 5; enp. 0-1; 0-2; I, II, 2

Leg 5 consisting of free protopod and exopod; protopod with 1 dorsodistal seta; exopod (Fig.

24B) 258×162 μm (ratio 1.59 : 1) spinulose along all margins, armed with 3 spines and 1 naked seta; lengths of these elements 134, 133, 275, and 229 μm, respectively, from outer to inner. Leg 6 represented by 2 small setule in genital aperture (Fig. 22C).

Male: Body (Fig. 24C) resembling that of female. Urosome 6-segmented. Genital somite subcircular. Antennule with 2 additional setae each on third and fourth segments at places indicated by dark spots in Fig. 22G. Antenna, labrum, mandible, maxillule as in female. Maxilla (Fig. 24D) with rows of spinules on proximal region of ventral surface of basal segment. Maxilliped consisting of 3 segments and large terminal claw; first and second segments with 2 setae each; small third segment unarmed; terminal claw with 2 proximal and 2 subproximal setae. Males in 3 types, showing differences, as follows:

Small male: Body length 1.90 mm (1.60-2.18 mm), based on 37 specimens. Caudal ramus 2.64 times as long as wide. Maxilliped (Fig. 24E) with straight inner and proximal margins. Leg 1 (Fig. Cyclopoida: Clausidiidae: Hemicyclops 47

E G D

F H

Fig. 22. Hemicyclops spinulosus, female. A. habitus, dorsal; B. urosome, dorsal; C. genital double-somite, ventral; D. distal part of abdomen, ventral; E. egg sac; F. spermatophore; G. antennule; H. antenna. Scales: A, G, H=0.05 mm, B=0.2 mm, C-F=0.1 mm. 48 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

F A

B C

H G

Fig. 23. Hemicyclops spinulosus, female. A. labrum; B. mandible; C. paragnath; D. maxillule; E. maxilla; F. maxilliped; G. leg 1; H. leg 2. Scales: A, B, D-F=0.05 mm, C= 0.02 mm, G, H=0.1 mm. Cyclopoida: Clausidiidae: Hemicyclops 49

B C

E F

G H

Fig. 24. Hemicyclops spinulosus. female: A. leg 4; B. leg 5 exopod. male: C. habitus, dorsal; D. maxilla; E. maxilliped of small male; F. maxilliped of medium-sized male; G, H. maxillipeds of large male. Scales: A, B, E, F=0.1 mm, C=0.5 mm, D=0.05 mm, G, H=0.2 mm. 50 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A D

C E

Fig. 25. Hemicyclops spinulosus, male. A. leg 1 of small male; B. C, third endopodal segment of leg 1 of small male; D. leg 1 endopod of leg 1 of medium-sized male; E. third endopodal segment of leg 1 of medium-sized male; F. leg 1 endopod of large mall; G. third endopodal segment of leg 1 of large male. Scales: A, E, G=0.05 mm, B, C=0.02 mm, D, F=0.1 mm.

25A) with stout endopod; inner distal spine on basis 86 μm long; endopod with several membranous flaps: 1 on second and 3 or 4 on third segments (Fig. 25B, C); outer spine on third segment 40 μm long and straight. Leg 5 exopod 1.92 times as long as wide.

Medium-sized male: Body length 2.40 mm (2.27-2.76 mm), based on 16 specimens, Caudal ramus

2.43 times as long as wide. Maxilliped (Fig. 24F) with curved inner and proximal margin. Leg 1 endopod (Fig. 25D) narrower and longer than that of small male; inner distal spine on basis 109 μm long, extending beyond distal margin of first endopodal segment; membranous flaps 1 on second and 3 on third segment (Fig. 25E); outer spine on third segment 53-60 μm long and slightly curved. Leg 5 exopod 1.93 times as long as wide.

Large male: Body length 3.08 mm (2.68-3.59 mm), based on 10 specimens. Caudal ramus 2.65 times as long as wide. Maxilliped (Fig. 24G, H) with 1 large, digitiform process on proximal region Cyclopoida: Clausidiidae: Hemicyclops 51

of inner margin; terminal claw strongly curved. Leg 1 endopod (Fig. 25F) elongate, much longer than those of other types of males; inner seta on coxa small. Inner distal spine on basis 154 μm long, extending to distal margin of first endopodal segment; membranous flaps short, 1 on second and 2 on third segments; outer spine on third segment straight, 65 μm long. Leg 5 exopod 2.30 times as long as wide, narrower than those of other types of males.

Distribution: Japan, Korea. Korea: GW.

Specimens examined: 64♀♀, 70♂♂ (37 small ♂♂, 16 medium ♂♂, 17 large ♂♂) from the external surface and burrows of the polychaete Tyllorhynchus heterohaetus, Maeho (brackish coastal lagoon), Yanynag-gun, Gangwon-do, 8.x.2007; more than 100 specimens, Maeho, 17.vi.2012; about 300 specimens, Maeho, 10.vii.2013.

Remarks: Itoh and Nishida (2008) studied the life cycle of this species and reported that the re- production of this species occurs mainly from early summer to autumn. They also grouped the males into 3 types, as small, medium, and large. The biological aspects of these male types may be a subject of research in future. This species is very common in Maeho which is one of brackish la- goons distributed along the east coast of Gangwon-do district of Korea.

11. Hemicyclops tanakai Itoh and Nishida, 2002 (Figs. 26-28)

Ssok-but-i-beol-re (쏙붙이벌레)

Hemicyclops tanakai Itoh and Nishida, 2002, p. 139, figs. 1-4.

Female: Body (Fig. 26A) moderately broad. Body length 1.42 mm. Urosome (Fig. 26B) 5-segmented. Fifth pedigerous somite 232 μm wide, with setules on posterolateral corners. Genital double-somite 164×212 μm (ratio 1.25 : 1) with laterally expanded anterior third and narrower posterior two-thirds; genital aperture small, located laterally in anterior region of anterior expansion. Genital double-somite and first and second free abdominal somites with membranous fringe along posterodorsal border. Three abdominal somites 102×124, 70×118, and 47×109 μm, respectively.

Anal somite with row spinules along posteroventral border (Fig. 26C). Caudal rami (Fig. 26C) divergent; each ramus weakly tapering, 115×47 μm (ratio 2.45 : 1), with 6 setae; outer lateral seta positioned distal 1-fifth of outer margin.

Rostrum hemicircular, with rounded posterior apex. Antennule (Fig. 26D) 384 μm long and 7-segmented; armature formula of 4, 15, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. An- tenna (Fig. 26E) 4-segmented; armature formula 1, 1, 4, and 7; inner distal corner of third segment produced; terminal segment 25×25 μm, as long as wide.

Labrum (Fig. 26F) as usual in the genus. Mandible (Fig. 26G) distally with 2 thick elements and 2 pinnate setae. Paragnath (Fig. 26H) spinulose and distally articulated. Maxillule (Fig. 26I) distally bilobed, with 5+3 setae. Maxilla (Fig. 26J) with 2 spiniform setae and 1 small setule on inner margin of basal segment; distal segment with 2 spines and 2 setae. Maxilliped (Fig. 27A) 4-segmented; first and second segments each with 2 large setae on inner margin; third segment unarmed; terminal segment with 3 setae and 1 spine, and terminated in spiniform process bearing 1 seta proximally (these spine and seta not articulated at base). 52 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

G A B

C I

Fig. 26. Hemicyclops tanakai, female. A. habitus, dorsal; B. urosome, dorsal; C. distal part of urosome, ventral; D. antennule; E. antenna; F. labrum; G. mandible; H. paragnath; I. maxillule; J. maxilla. Scales: A=0.2 mm, B-D=0.1 mm, E-J=0.05 mm. Cyclopoida: Clausidiidae: Hemicyclops 53

A B

D C

Fig. 27. Hemicyclops tanakai, female. A. maxilliped; B. leg 1; C. leg 2; D. leg 4; E. leg 5. Scales: 0.05 mm. 54 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B D A

Fig. 28. Hemicyclops tanakai, male. A. habitus, dorsal; B. urosome, ventral; C. maxilla; D. maxilliped. Scales: A=0.2 mm, B=0.1 mm, C, D=0.05 mm.

Legs 1-4 with 3-segmented rami (Fig. 27B-D). Intercoxal plate of leg 1 with setules along distal margin but those of legs 2-4 with spinules. Inner distal spine on leg 1 basis slender and extending over distal border of first endopodal segment. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0: I-1; I, 3, 4; enp. 0-1; 0-1; I, 5 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 2, 5; enp. 0-1; 0-2; I, II, 3 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; I, 2, 5; enp. 0-1; 0-2; I, II, 2

Leg 5 (Fig. 27E) composed of protopod and exopod; protopod with 1 dorsodistal seta; exopod

123×75 μm (ratio 1.65 : 1), with 3 spines and 1 seta. Leg 6 not discernible.

Male: Body (Fig. 28A) similar to that of female. Body length of dissected specimen 1.45 mm.

Greatest width 676 μm. Urosome (Fig. 28B) 6-segmented. Genital somite 214×277 μm, increasing- ly broader distally. Four abdominal somites 97×137, 89×124, 55×117, and 47×110 μm, respectively. Caudal ramus 114×46 μm (ratio 2.48 : 1).

Rostrum as in female. Antennule with setation as in female, but fourth segment with 4 setae (not 3 setae as in female). Antenna as in female.

Labrum, mandible and maxillae as in female. Maxilla (Fig. 28C), second segment with 1 bifur- Cyclopoida: Clausidiidae: Hemicyclops 55

cate spine, 2 setae and 1 stout process. Maxilliped (Fig. 28D) consisting of 3 segments and terminal claw; first segment with 1 long seta; second segment strongly tapering, with 2 small inner seta; third segment small and unarmed; terminal claw arched, with 2 small seta proximally. Leg 1 without inner distal element on basis. Legs 2-4 as in female. Leg 5 consisting of dorsolateral seta on fifth pedigerous somite and free exopod (protopod completely fused with somite); exopod 1.68 times as long as wide. Leg 6 represented by 1 spiniform seta on genital operculum (Fig. 28B).

Distribution: Japan, Korea. Korea: GN, JJ, CN.

Specimens examined: Hosts are Callianassa japonica (Ortmann). 30♀♀, 20♂♂ from washings of, Sinheung-ri, Jocheon, Jeju I., 29.ix.1999; 8♀♀, 1♂, Sinheung-ri, 2.vi.2000; 2♀♀, tidal flat, Geumsong-ri, Namhae I., Gyeongsangnam-do, 21.x.2010; 14♀♀, 2♂♂, Choonjangdae beach, Chooncheongnam-do, 15.x.2011; 4♀♀, Daecheon Beach, Choongcheongnam-do, 24.vi.2012; 6♀ ♀, 5♂♂, Sangju Beach, Namhae I., Gyeongsangnam-do, 5.vii.2012; 16♀♀, Choonjangdae Beach, 20.v.2014.

Remarks: Callianassa japonica (Ortmann), the host of this species, lives in sands or mudy sands of the intertidal zone and shallow water.

12. Hemicyclops ventriplanus Kim, 2000 (Fig. 29)

Nap-jak-geom-mul-byeo-ruk-but-i (납작검물벼룩붙이)

Hemicyclops ventriplanus Kim, 2000a, p. 369, figs. 1-3.

Female: Body (Fig. 29A) of dissected specimen 1.39 mm long. Greatest width 0.55 mm. Prosome composed of cephalothorax and 3 pedigerous somites. Rostral area of cephalosome produced anteriorly. Prosomal somites with well developed epimerae. Second and third pedigerous somites with posterior margin of tergite fringed with membrane. Fourth pedigerous somite with pointed posterior corners. Urosome (Fig. 29B) 5-segmented. Fifth pedigerous somite 194 μm wide, with round lateral margins. Genital double-somite ventrally flat, dorsally complex as in Figure 1B,

212×188 μm (1.13 : 1). Genital aperture located laterally near anterior portion of somite at area of greatest width. Three abdominal somites 103 ×117, 74×109, and 62×101 μm, respectively. Pos- terodorsal border of second abdominal somite convex, fringed with membrane. Anal somite with spinules on medial side of posteroventral border. Caudal ramus 102×40 μm (2.55 : 1), with nearly parallel lateral margins and six setae. Outermost seta divided into two parts: thick basal and plumose distal parts. Longest inner median terminal seta 653 μm. Next longest, outer median terminal seta 359 μm. Egg sac oval (Fig. 1A), 441×223 μm. Rostrum incorporated with ventral surface of cephalothorax. Antennule 7-segmented, 363 μm long, with armature formula of 4, 15, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Anten- na 4-segmented, with armature formula 1, 1, 4, 7. Third segment with distinct inner distal protuberance and spinules on inner margin. Terminal segment slightly wider than long. Mandible armed terminally with 4 elements: 1 dorsal, plate-like element bearing denticles, another plate-like element bearing spinules on dorsal side, and 2 ventral plumose setae. Paragnath a 56 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B A D

Fig. 29. Hemicyclops ventriplanus, female. A. habitus, dorsal; B. urosome, dorsal; C. maxilla; D. maxilliped. Scales: A=0.2 mm, B=0.1 mm, C=0.05 mm, D=0.02 mm.

lobe bearing distal process and a posterior row of spinules. Maxillule distally bilobed, with five setae on larger lobe and 3 setae on smaller lobe. Maxilla (Fig. 29C) 2-segmented. Basal segment with row of setules basally and 2 distal, barbed setae, one of them accompanied basally with 1 setule.

Maxilliped (Fig. 29D) 4-segmented, with formula of armature: 2, 2, 0, 3+II. First and second segments each with row of spinules in addition to setae. Inner margin of second segment produced near base of setae. Terminal spine on terminal segment with 1 basal, terminally-forked setae and 6 or 7 setules on outer side of proximal half. Inner spine enlarged, with several stiff spinules on both margins. Legs 1-4 with 3-segmented rami. Formula of armature of these legs as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0; I-1; II, 6; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 7; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 7; enp. 0-1; 0-2; III, 3 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; I, 7; enp. 0-1; 0-2; III, 2 Cyclopoida: Clausiidae 57

Median portion of posterior margin of basis of legs 1-4 spinulated. Posterior margin of intercoxal plate of leg 1 with hairs, but those of legs 2-4 with spinules. Outer margin of basis in leg 1 with setae, but those in legs 2-4 with spinules. Inner spine of basis of leg 1 slender and longer than first endopodal segment. Outer margin of endopod of all biramous legs with hairs. Outer margin of first exopodal segment smooth but those of legs 2-4 with spinules.

Leg 5 2-segmented. Protopod with 1 seta, and spinules near base of seta. Exopod 103×51 μm

(ratio 2.02 : 1), with 3 spines, 1 seta, and spinules on outer margin and on distal half of inner margin. Leg 6 not observable, concealed from dorsal and ventral views of body. Spermatophore 120×62 μm, a curved sac. Male: Body resembling that of female. Length 1.38 mm. Urosome 6-segmented. Fifth pedigerous somite 212 μm wide, narrowed posteriorly. Genital somite nearly hexagonal, 188×218 μm. Four abdominal somites 88 ×126, 91×106, 62×100, and 52×95 μm, respectively. Caudal ramus

91×38 μm (2.39 : 1). Antennule, antenna, labrum, mandible, paragnaths, and maxillule identical to those of female. Maxilla sexually dimorphic; second segment terminated in a claw-like, blunt process armed distally with minute spinules. Maxilliped composed of 3 segments and terminal claw. First segment with 1 long plumose seta. Second segment greatly expanded proximally and narrowed distally, with 1 row of minute spinules, 2 rows of larger spinules and distantly separated, 2 small setae on inner surface. Third segment unarmed. Claw weakly curved, proximally with 1 seta and 1 spinule. Leg 1 without inner spine on posterior margin of basis. Legs 2-4 as in female. Protopod of leg 5 completely incorporated into fifth pedigerous somite. Leg 5 exopod similar in shape to that of female. Leg 6 represented by 2 small spines on genital operculum.

Distribution: Japan, Korea. Korea: GN, GG.

Specimens examined: 52♀♀, 33♂♂ (type specimens) from Upogebia sp., tidal flat, Jakyak I., Incheon, 28.ix.1996; 3♀♀, 3♂♂ from invertebrate burrows, collected along with Hemicyclops japonicus, type locality, 7.x.1998; numerous specimens, invertebrate burrows, type locality, 7.x.1998;

1♀ from washings of the gobiid fish Istigobius campbeli (Jordan and Snyder) (accidental association?), Cheokpo Port, Tongyeong, Gyengsangnam-do, 1.vi.2012, S.Y. Moon;

Remarks: The original record reported Upogebia major (De Haan) as the host of this species. However, the true host seems to be some other species of Upogebia.

Family Clausiidae Giesbrecht, 1895

Gin-bae-geom-mul-byeo-ruk-gwa (긴배검물벼룩과)

Body cyclopiform. Urosome 3-6- (generally 6-) segmented in both sexes. Antennule 4-6-segmented (7-segmented in Boreoclausia). Antenna 3- or 4- segmented; setation variously reduced. Mandible with 1 or 2 distal elements. Maxillule with at most 6 setae. Maxilla 2-segmented; first segment usually unarmed; distal segment with 1 or 2 elements. Maxilliped 2-4-segmented (absent in Maxilliclausia). Legs 1-4 variously reduced in segmentation and setation. Inner distal spine on basis of leg 1 usually absent (present in Pontoclausia and Likroclausia). Free exopod of leg 5 present 58 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans or absent. Associated with polychaetes.

Key to the genera of family Clausiidae (based on female)

1. Second to fourth pedigerous somite with spinulose, long processes; distal ·endopodal segment of leg 1 modified to a powerful hook ························································································ Likroclausia - Prosomal somites smooth; leg 1 endopod normal ················································································ 2 2. ‌Prosome not segmented; mandible with 1 elongate spine distally; distal segment of maxilla transformed to a sucker ······································································································· Maxilliclausia - ‌Prosome segmented; mandible with 2 distal elements; distal segment of maxilla not transformed to a sucker ··················································································································································· 3 3. Legs 1-3 with 3-segmented rami; leg 4 with 2- or 3-segmented rami ··············································· 4 - Legs 1 and 2 with 2-segmented rami, but legs 3 and 4 rudimentary or absent ································ 5 4. Leg 1 basis with inner distal spine ······················································································· Pontoclausia - Leg 1 basis without inner distal spine ··················································································· Rhodinicola 5. Leg 4 as a lobe bearing seta ····································································································· Indoclausia - Leg 4 absent ······································································································································ Clausia

Genus Pontoclausia Bacescu and Por, 1959

Jang-da-ri-geom-mul-byeo-ruk-sok (장다리검물벼룩속)

Urosome 6-segmented in female. Legs 1-3 with 3-segmented exopod and endopod. Leg 4 with 2- or 3-segmented exopod and endopod. Leg 1 basis with inner distal spine. Other characters as those of the family.

13. Pontoclausia antiqua (Kim, 2001) (Fig. 30)

Se-ma-di-geom-mul-byeo-ruk (세마디검물벼룩)

Clausia antiqua Kim, 2001b, p. 218, figs. 1-3.

Female: Body (Fig. 30A) large. Body length 3.20 mm. Greatest width 1.51 mm, measured across second pedigerous somite. Prosome broad and flat. Prosomal somites with well developed, rig- id tergites. Prosome of young adult with less developed tergites and faint suture lines between somites. Cephalosome and first pedigerous somite completely fused to form cephalothorax.

Mid-posterior margin of fourth pedigerous somite conspicuously protruded posteriorly. Urosome (Fig. 30B) 6-segmented. Fifth pedigerous somite about 618 μm wide. Genital somite 265×538 μm, anterior portion expanded laterally, with steeply oblique lateral margins. Paired genital fields located dorsolaterally. Abdomen gradually narrowing. Four abdominal somites 235×305, 229×265, 164×229 and 212×206 μm, respectively. Genital and first 3 abdominal somites ornamented with minute spinules on ventral surface. Anal somite with minute spinules near posteroventral border.

Caudal ramus weakly tapering, 173 ×72 μm (2.37 : 1), with minute spinules near distal border of Cyclopoida: Clausiidae: Pontoclausia 59

ventral surface. Caudal setae 7 in number, including proximal lateral seta (seta I); all setae naked, largest 1 741 μm long. Rostrum nearly quadrangular, constricted laterally, and wider than long. Antennule 6-segmented, gradually tapering, 490 μm long, with armature formula of 4, 12, 8, 4 +aesthetasc, 2+aesthetasc and 7+aesthetasc. All aesthetascs setiform, hardly distinguishable from setae. All setae naked. Antenna (Fig. 30C) 3-segmented. All segments with minute spinules on inner surface. First and second segments each armed with 1 distal seta. Third segment subdivided by an incomplete suture line into proximal and distal portions (original third and fourth segments). Proximal portion with 2 small and 1 large, lanceolate setae; the latter rimmed with membrane along inner margin. Distal portion reduced, with 3 plain and 4 spiniform setae; the latter 4 setae each tipped with small spinulated pad.

Labrum reduced, tapering posteriorly, and not covering mouthparts. Mandible (Fig. 30D) with 2 extremely unequal, terminal and subterminal spines; terminal spine large, rod-shaped, directed posteriorly, crenate on tip and along outer (posterior) margin; subterminal spine distinctly smaller than terminal, claw-like, with spinules on both sides. Paragnath naked lobe. Maxillule (Fig. 30E) lobate bearing anterior protrusion, armed with 6 distal setae; posteriormost seta bent basally; next posteriormost seta thick but shorter. Maxilla (Fig. 30F) largest mouthpart, 2-segmented: first segment broad and unarmed; second segment blunt, wider than long, terminally sclerotized, forming cutting edge, with 1 small seta at subterminal part of posterior margin. Maxilliped rudimentary, 3-segmented and unarmed: terminal segment very small, with pointed tip. Legs 1-4 with 3-segmented rami. Inner spine on basis of leg 1 shorter than first endopodal segment (Fig. 30G), with bilateral marginal membrane. Most spines on rami of legs setiform. Leg 4 (Fig. 30H) without inner coxal seta. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-I; exp. I-0; I-1; III, I, 3; enp. 0-1; 0-1; I, 4 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 4; enp. 0-1; 0-1; III, 3 Leg 4: coxa 0-0; basis 1-0; exp. I-0; I-1; II, I, 4; enp. 0-1; 0-1; III, 2

Leg 5 (Fig. 30B) 2-segmented. Protopod with 1 outer seta and spinules on outer margin. Exo- pod roughly oval, 233×140 μm (1.66 : 1) with 2 outer and 2 terminal setae. Both outer and inner margins ornamented with spinules. All setae shorter than exopod. Leg 6 represented by 2 minute spinules in genital field. Male: Body 3.61 mm long. Prosomal somites with faint suture lines between somites. Second pedigerous somite as wide as cephalothorax. Third pedigerous somite narrower than second, with epimera well separated from those of second and fourth pedigerous somites. Fourth pedigerous somite almost fused with fifth pedigerous somite, much narrower than third, its epimera directed posterolaterally. Urosome 6-segmented. Genital somite nearly quadrangular, 606×629 μm, slightly wider than fifth pedigerous somite. Four abdominal somites 235×329, 200×288, 135×247 and

200×232 μm, respectively. Caudal ramus 147×75 μm (1.96 : 1).

Antennule with additional seta on third segment (9 setae) than in female. Antenna with more conspicuous spinules on inner margins of segments, otherwise as in female. Labrum, mandible and maxillule as in female. Maxilla with large patch of spinules on ventral surface of first segment. Maxilliped well-developed, 4-segmented. First segment gradually broadening distally, with 1 small seta on inner margin. Second segment with minute spinules on inner surface and pair of small setae in middle of inner margin. Third segment short and unarmed. Fourth segment forming long, slender, distally curved claw, basally with 2 setae on opposite sides. Leg 1 sexually dimorphic: endopod 2-segmented with armature formula 0-1; I,5. Inner spine on basis longer than first endopodal segment. Legs 2-4 as in female Leg 5 exopod with narrower 60 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B C

D F

Fig. 30. Pontoclausia antiqua, female. A. habitus, dorsal; B. urosome, dorsal; C. antenna; D. mandible; E. maxillule; F. maxilla; G. leg 1; H. leg 4. Scales: A=0.5 mm, B=0.2 mm, C, E, F=0.05 mm, D=0.02 mm, G, H=0.1 mm. Cyclopoida: Clausiidae: Pontoclausia 61

distal half, 173×73 μm (2.37 : 1). Distalmost seta longest, 210 μm, longer than exopod. Leg 6 represented by 1 seta on posterior corner of genital operculum.

Distribution: Korea. Korea: GN, JJ, CN.

Specimens examined: 7♀♀, 4♂♂ (type specimens) from washings of the polychaete Arenico- la brasilliensis Nonata, in the intertidal sands at Sinhung-ri, Jocheon, Jeju I., 29.ix.1999; 1♂ from A. brasilliensis in the intertidal mud flat at Sapsi I., Choongcheongnam-do, 26.x.1999; ♀1 , 2♂♂ from A. brasilliensis, Geumsong-ri, Namhae I., Gyeongsangnam-do, 7.vii.2001. Remarks: The large size of body with strongly expanded prosome is a typical feature of this species. The polychaete host Arenicola brasiliensis of this copepod is widely distributed in the world. Therefore, P. antiqua is thought to be distributed widely as well.

14. Pontoclausia lobata (Kim, 2000) (Fig. 31)

Du-ma-di-geom-mul-byeo-ruk (두마디검물벼룩)

Clausia lobata Kim, 2000a, p. 375, figs. 4-6.

Female: Body (Fig. 31A) small, flat, 650 μm long. Greatest width 265 μm. Ventrolateral sides of cephalothorax with many wavy adhesion ridges. Rostral area of cephalothorax projected, broad, truncate anteriorly, and demarcated by faint line from remaining part of cephalothorax. Third pedigerous somite narrower but distinctly longer than second pedigerous somite. Fourth pedigerous somite with characteristic lobate protuberance on each lateral side. Urosome (Fig. 31B) 6-segmented. Fifth pedigerous somite 103 μm wide, distinctly wider than genital somite. Genital somite 49×83 μm, or 1.69 times wider than long, nearly quadrangular, wider posteriorly. Demarcation between genital and first abdominal somites distinct dorsally but obscure ventrally. Abdomen slightly tapering. Four abdominal somites 43×53, 38×45, 28×40, 25×39 μm, respectively. Anal somite with 2 pairs of membranous flaps, each proximally and distally on ventral surface. Caudal rami divergent, 32×16 μm (2.0 : 1). Outer lateral seta located two-thirds length of ramus. Two mid-terminal setae weakly pinnate in distal half. Other 4 setae naked. Egg sac elongate, longer than urosome, 323×105 μm, containing relatively large eggs. Rostrum completely incorporated into cephalothorax, leaving only faint. Antennule 6-segmented and 142 μm long, with armature formula of 4, 9, 9, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Plumose setae are 1 on both fourth and fifth segments and 3 on terminal segment. First segment distinctly broader than long. Antenna (Fig. 31C) 3-segmented, with armature formula of 1, 0, and 7. Second segment with 3 membranous flaps. Third segment slightly broadened distally, with 1 membranous flap; outermost seta on terminal segment plumose, and longest. Innermost seta broadened, spiniform, with membrane on inner margin and tipped with minute spinule. Other five setae naked.

Labrum (Fig. 31D) reduced, not covering mouth parts, with narrow posterior margin. Mandi- ble (Fig. 31D) small, armed terminally with 2 spiniform elements of different sizes. Maxillule (Fig. 31D) with 3 setae terminally and protuberance on inner side; outer one of these setae plumose on outer side, but other 2 setae naked. Maxilla (Fig. 31D) largest among mouth parts, 1-segmented, 62 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B C

Fig. 31. Pontoclausia lobata, female. A. habitus, dorsal; B. urosome, dorsal; C. antenna; D. mouthparts; E. leg 1; F. leg 4. Scales: A=0.1 mm, B=0.05 mm, C-F=0.02 mm. Cyclopoida: Clausiidae: Pontoclausia 63 with 3 membranous flaps posteroventrally and 2 sclerotized blades distally, without seta or spine.

Maxilliped (Fig. 31D) 3-segmented, blunt, and unarmed. Third segment short, distally truncate, blade-like and indistinctly marked from second segment.

Leg 1 (Fig. 31E) and legs 2 and 3 with 3-segmented rami. Leg 4 (Fig. 31F) with 2-segmented rami. All spines on rami of these legs setiform. Inner spine on posterior margin of leg 1 thick but short. Membranous flaps seen on intercoxal plate, coxa, basis and segments of rami of legs -1 4. Outer seta on basis of leg 3 slender compared with that of leg 2, and that of leg 4 naked. Legs 3 and 4 without inner seta of coxa. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis1-I; exp. I-0; I-1; II, 4; enp. 0-1; 0-1; 2, 2 Leg 2: coxa 0-1; basis 1-0; exp. I-0; 0-1; I, 5; enp. 0-1; 0-1; I, 3 Leg 3: coxa 0-0; basis 1-0; exp. I-0; 0-1; I, 5; enp. 0-1; 0-1; I, 3 Leg 4: coxa 0-0; basis 1-0; exp. I-0; 5; enp. 0-1; 3

Leg 5 (Fig. 31B) 2-segmented. Each segment with 3 membranous flaps. Protopod well defined from fifth pedigerous somite, with 1 outer distal seta mounted on digitiform elevation. Exopod nearly quadrate, 25×20 μm, with 4 long, naked terminal setae. Leg 6 represented by 1 spinule in genital aperture. Male: Body distinctly tapering, very small. Length 0.37 mm. Greatest width 177 μm. Urosome 6-segmented. Fifth pedigerous somite wider than genital somite. Genital somite 30×52 μm, dorsally well marked from, but ventrally fused with, fifth pedigerous somite. Genital operculum not seen. Four abdominal somites 26 ×42, 21×35, 18×30, and 18×28 μm, respectively. Anal somite with membranous flap on each side of ventral surface. Caudal ramus 18×11 μm (1.64 : 1). Maxilliped 3-segmented. First segment very short and unarmed. Second segment with seven small tubercles on inner surface. Third segment the longest, tapering and claw-like, with 1 tubercle proximally and 1 curved row of small bead-like spinules. Other mouthparts as those of female. Leg 1 endopod 2-segmented. First segment with 1 inner seta. Second segment with 4 setae. Legs 2-4 with armature formula identical to that of female. Leg 5 protopod completely incorporated into fifth pedigerous somite, leaving 1 long seta near base of exopod. Exopod small, lobate, 7.7×4.6 μm (1.67 : 1), with 4 setae. Leg 6 not observed.

Distribution: Korea. Korea: JN, JB, GG.

Specimens examined: 36♀♀, 3♂♂ (type specimens) from body surface of the polychaete Mar- physa sanguinea (Montagu), tidal flat, Jakyak I., Incheon, 1.ix.1996; ♀♀2 from washings of invertebrates, Gosapo, Buan-gun, Jeollabuk-do, 26.ix.1999; 4 pairs in amplexus from Marphysa sanguinea, Moochangpo, Choongcheongnam-do, 6.v.2000; 10♀♀, 10♂♂ from M. sanguinea, tidal flat, Yongyu I., Incheon, 8.x.2006; 9♀♀, 8♂♂ from polychaetes, tidal flat, Weoldoo, Mooan-gun, Jeollanam-do, 17.x.2012. Remarks: The body of this copepod is small and easily unnoticed under the dissecting microscope. 64 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Genus Clausia Claparède, 1863

Gin-bae-geom-mul-byeo-ruk-sok (긴배검물벼룩속)

Urosome 6-segmented. Antenna 3-segmented. Manddible with 2 spines distally. Legs 1 and 2 with 2-segmented exopod and endopod. Leg 3 as lobe bearing 1 or 2 setae. Leg 4 absent. Leg 5 consisting of 1 seta on fifth pedigerous somite and free exopod bearing 4 elements. Other features as those of the family.

15. Clausia parva Kim, 2014 (Figs. 32-34)

Jag-eun-gin-bae-geom-mul-byeo-ruk (작은긴배검물벼룩)

Clausia parva Kim, 2014, p. 100, figs. 4-6.

Female: Body (Fig. 32A) elongate, cylindrical, and relatively small. Body length of dissected specimen 1.49 mm. Prosome (Fig. 32A, B) indistinctly segmented, but 4 prosomal somites well-defined from 1 another by constrictions and thin exoskeletons between them. Greatest width of prosome 278 μm across second pedigerous somite. All prosomal somites with rounded lateral margins. Cephalothorax 275 μm wide and as long as wide, with truncated rostral protuberance. Second to fourth pedigerous somites similar in length and width, with rudimentary dorsal tergite. Urosome slightly shorter than prosome and 6-segmented. Fifth pedigerous somite 272 μm wide, as wide as prosomal somites. Genital somite 135 ×219 μm; genital apertures locating dorsolaterally in subdistal region of somite. Abdomen distinctly narrower than preceding somites, 96×126, 87×125, 67×110, and 108×102 μm, respectively; all abdominal somites smooth, without ornamentation. Caudal rami (Fig. 32C) small and widely separated from each other; each ramus 51×27 μm

(ratio 1.89 : 1) and weakly tapering in distal half, with 6 naked setae; largerst terminal seta (seta V) 250 μm long, markedly larger than other 5 setae; outer lateral seta located at midlength of ramus.

Egg sac (Fig. 32D) longer than urosome, containing several uniserial eggs; each egg about 200 μm in diameter.

Rostrum as anterior protuberance of cephalothorax. Antennule (Fig. 32E) short, 142 μm long and 5-segmented, with armature formula of 5, 23, 4, 2+aesthetasc, and 7+aesthetasc; all setae naked; first segment with proximal patch of minute spinules on anterior surface; second segment the longest, longer than combined distal 3 segments. Antenna (Fig. 32F) 3-segmented; first segment (basis) with 1 distal seta and patches of spinules on medial and outer surfaces; second segment (first endopodal segment) unarmed, with 2 patches of minute spinules; terminal segment (second endopodal segment) with 4 distal claws of unequal lengths, 2 small, subdistal claws (proximal one of them being setiform) on medial margin, 2 weakly pinnate setae on outer margin, and minute spinules on medial margin and middle of outer margin.

Labrum (Fig. 33A) unornamented, with tapering posterior lobes, and deep and broad posteromedian invagination. Mandible (Fig. 33A) with 2 very unequal, distal spines; larger anterior one of them with 5 teeth distally. Maxillule (Fig. 33A) as lobe bearing 3 outer setae, 1 inner seta, and on ventral surface patch of minute spinules. Maxilla (Fig. 33B) 2-segmented; proximal segment broad, unarmed but with 1 small patch of minute spinules on dorsal (anterior) surface; distal segment Cyclopoida: Clausiidae: Clausia 65

A E

D F

Fig. 32. Clausia parva, female. A. habitus, dorsal; B. anterior part of prosome, right; C. caudal rami, dorsal; D. egg sac; E. antennule; F. antenna. Scales: A, B=0.1 mm, C=0.05 mm, D=0.2 mm, E, F=0.02 mm. 66 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

C G

Fig. 33. Clausia parva, female. A. labrum, mandibles, and left maxillule; B. maxilla; C. maxilliped; D. leg 1; E. leg 2; F. leg 3; G. leg 5. Scales: A-F=0.02 mm, G=0.05 mm.

forming distally obliquely expanded spinulose pad, with 2 small setae (each on anterior and posterior surface) and scattered minute spinules. Maxilliped (Fig. 33C) 3-segmented; first segment broad but short and unarmed; second segment the longest, with 2 small medial setae at proximal third of medial margin; terminal segment short, stout, and distally forming spinulose pad, with 1 subdistal spine.

Legs 1 and 2 (Fig. 33D, E) with 2-segemented rami; inner seta on coxa small, blunt and naked; basis with patch of spinules on posterior region near base of endopod; outer seta on basis thick and stiff; outer surfaces of rami covered with spinules; medial surface of first endopodal segment of leg Cyclopoida: Clausiidae: Clausia 67

A D

Fig. 34. Clausia parva, male. A. habitus, dorsal; B. urosome, ventral; C. maxilliped; D. leg 1; E. leg 2. Scales: A, B=0.1 mm, C-E=0.02 mm. 68 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

1 also covered with spinules; outer spines on leg 2 exopod with subdistal setule. Leg 3 (Fig. 33F) as small lobe bearing 2 setae. Leg 4 absent. Armature formula of legs 1-3 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; II, I, 0; enp. 0-0; 0, II, 2 Leg 2: coxa 0-1; basis 1-0; exp. I-0; III, I, 1; enp. 0-0; 0, I, 2

Leg 3: coxa 2 (on lobe)

Leg 5 (Fig. 33G) consisting of protopod and distal exopod; protopod 108×69 μm, articulated at base from somite, with 1 small outer seta. Exopod distinctly smaller than protopod, 51×32 μm (ratio 1.59 : 1); with 2 patches of minute spinules and 4 setae; these 4 setae smooth or weakly setulose, and 120, 172, 55, and 105 μm long, respectively, from outer to inner. Leg 6 represented by 1 small setule in genital aperture.

Male: Body (Fig. 34A) cylindrical as in female. Body length 0.89 mm in dissected paratype. Pro- some well-segmented. Cephalothorax 221×190 μm, longer than wide. Three metasomites distinctly shorter than cephalothorax and 72×165, 51×151, and 44×146 μm, respectively. Urosome (Fig. 34B) 6-segmented. Fifth pedigerous somite 72×149 μm. Genital somite 94×140 μm, with weakly convex lateral margins. Four abdominal somites 62×102, 65×94, 58×79, and 87×81 μm, respectively. Caudal ramus 48×25 μm (ratio 1.92 : 1), not different from that of female in shape.

Rostrum, antennule and antenna as in female. Maxilliped (Fig. 34C) 3-segmented as in female; first segment small and unarmed; second segment strongly protuberant medially, with 2 naked setae on tapering medial protuberance; terminal segment forming strong, smooth, distally curved claw. Other mouthparts as in female.

Legs 1 and 2 (Fig. 34D, E) with 2-segmented rami as in female but with different armature formula; setae on coxa and rami well-developod and pinnate, unlike those of female. Terminal spine on distal exopodal segment in legs 1 and 2 larger than that of female, with subterminal setule. Second endopodal segment of leg 1 with 2 spinulose bulges on medial side posterior to base of medial seta. Leg 3 as lobe bearing 2 setae, as in female. Leg 4 absent. Armature formula of legs 1-3 as follows:

Leg 1: coxa 0-1; basis 1-0; exp. I-0; III, I, 3 (or II, I, 2); enp. 0-0; I, 1, 1 Leg 2: coxa 0-1; basis 1-0; exp. I-0; II, I, 3; enp. 0-0; 0, I, 3

Leg 3: coxa 2 (on lobe)

Leg 5 similar to that of female; exopod 48×31 μm (ratio 1.55 : 1). Leg 6 represented by 1 rudimentary seta on genital operculum.

Distribution: Korea. Korea: GW, GN.

Specimens examined: 15♀♀, 3♂♂ (type specimens) from the polychaete Hydroides ezoensis

Okuda epizoic on the shell of the oyster Crassostrea gigas (Thunberg), Gangneung Port (37°46′15″N, 128°57′08″E), depth about 50 cm, 21.xi.2012; 2♀♀ from washings of H. ezoensis epizoic on the shell of Crassostrea gigas, Yangpo Port (35°15′54″N, 129°31′28″E), 31.vii.2007. Remarks: Currently, the genus Clausia contains two species, C. lubbockii Claparède, 1863 and this species. This species differs from C. lubbockii in the following differences: The antennule is 5-segmented in this species, but 6-segmented in C. lubbockii. The armature formulae of legs 1 and 2 of this species are I-0; II, I, 0 (vs. I-0; II, II, 1 in C. lubbocki) for leg 1 exopod; 0-0; 0, II, 2 (vs. 0-1; 0, I, 1 in

C. lubbockii) for leg 1 endopod; I-0; III, I, 1 (vs. I-0; II, I, 1 in C. lubbocki) for leg 2 exopod; and 0-0; 0, I,

2 (vs. 0-1; II, I, 0 in C. lubbocki) for leg 2 endopod (Kim, 2014). Cyclopoida: Clausiidae: Indoclausia 69

Genus Indoclausia Sebastian and Pillai, 1974

In-do-geom-mul-byeo-ruk-sok (인도검물벼룩속)

Characters as those of the genus Clausia, except that leg 4 present as a lobe bearing 1 or 2 setae.

16. Indoclausia bipartita Kim, 2014 (Figs. 35-37)

Du-ma-di-in-do-geom-mul-byeo-ruk (두마디인도검물벼룩)

Indoclausia bipartita Kim, 2014, p. 96, figs. 1-3.

Female: Body (Fig. 35A) elongate and cylindrical. Body length 2.87 mm (2.48-3.47 mm) based on 8 specimens. Dissected specimen 3.30 mm long. Prosome-urosome division indistinct. Prosome longer than urosome, unsegmented, but each somite clearly defined by constrictions between somites. Cephalothorax nearly circular, 653 ×693 μm, with slightly produced and truncated anteri- or apex. Second to fourth pedigerous somites 346×673, 426×574, and 495×564 μm, respectively, with rounded lateral margins. Urosome (Fig. 35B) 4-segmented, consisting of fifth pedigerous somite, genital double-somite, and 2 free abdominal somites. Fifth pedigerous somite about 465×594 μm and nearly hexagonal. Genital double-somite 238×520 μm, much wider than long; genital aperture located dorsally. Two free abdominal somites 158×350 and 218×304 μm, respectively. Anal somite (second free abdominal somite) distinctly longer than first free abdominal somite, without ornamentation on ventral surface. Caudal rami parallel and widely separated from each other; each ramus gradually narrowing distally, 264×105 μm (ratio 2.51 : 1), with 6 naked setae; outer lateral seta locating at midlength of ramus; largest terminal seta (seta V) 541 μm long; next largest seta (seta IV) 198 μm long; other 4 setae much smaller, shorter than ramus width.

Rostrum (Fig. 36A) distinct, highly sclerotized and much broader than long, with convex posteri- or margin. Antennule (Fig. 35C) 383 μm long and 6-segmented, with armature formula of 5 (or 6), 18, 14, 4, 2+aesthetasc, and 7+aesthetasc; first segment short but widest among segments; remaining segments grandually narrowing; suture between second and third segments indistinct; setae dense on second and third segments; all setae naked and thin. Antenna (Fig. 35D, E) 3-segmented; first segment (basis) longer and wider than 2 distal segments, with 1 seta medio-distally; second segment (first endopodal segment) unarmed but ornamented with large patch of fine spinules on medio-distal surface; terminal segment (second endopodal segment) shorter than second segment and armed with 4 distal claws of unequal sizes, 2 outer subdistal setae, 2 medial subdistal claws, and 1 minute medial seta proximal to 2 subdistal claws and ornamented with 2 patches of fine spinules, one of latters on medial surface and the other on posterior surface.

Labrum (Fig. 36A) unornamented, with deep and broad posteromedian invagination. Mandible (Fig. 35F) elongate, with 2 unequal distal elements: larger element 49 μm long, with 6 or 7 teeth in distal region; smaller element as flame-shaped broad seta. Paragnath not discernible. Maxillule (Fig. 36B) as a blunt lobe bearing 3 larger outer setae, 2 smaller inner setae and on posterior surface

1 large patch of minute spinules. Maxilla (Fig. 36C) 2-segmented; proximal segment much wider than long, with 1 large distal protuberance covered entirely with minute spinules; distal segment blade-like and bluntly ended, with 2 small setae (1 on anterior and 1 on posterior surfaces) and 70 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B A

D E

Fig. 35. Indoclausia bipartita, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. distal part of antenna; F. mandible. Scales: A=0.5 mm, B=0.2 mm, C, D=0.05 mm, E, F=0.02 mm. Cyclopoida: Clausiidae: Indoclausia 71

B A

Fig. 36. Indoclausia bipartita, female. A. cephalic area ventral; B. maxillule; C. maxilla; D. maxilliped; E. leg 1; F. leg 2; G. leg 3; H. leg 4. Scales: A=0.1 mm, B=0.02 mm, C-H=0.05 mm. 72 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B A

Fig. 37. Indoclausia bipartita. female: A. leg 5. male: B. habitus, dorsal; C. maxilliped; D. left genital operculum, ventral. Scales: A=0.1 mm, B=0.2 mm, C, D=0.05 mm.

transparent flange on cutting edge. Maxilliped (Fig. 36D) 4-segmented; first segment widest but very short and unarmed; second segment with 1 patch of minute spinules on medial surface; small third segment unarmed; terminal segment gradually narrowed distally, terminating in blunt tip covered with minute spinules, with 2 small setae.

Legs 1 and 2 (Fig. 36E, F) with 2-segmented rami and patches of spinules on coxa, basis and segments of rami. Distal spines on second exopodal segment of these legs with minute spinules along outer margin. Setae on these legs small and naked. Leg 3 (Fig. 36G) lobate, with 3 naked setae dis- Cyclopoida: Clausiidae: Likroclausia 73

tally. Leg 4 (Fig. 36H) also lobate, tipped with 1 seta, and with distal patch of minute spinules near base of seta. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-0; basis 1-0; exp. 1-0; 2, II, 1; enp. 0-0; 0, 1, 0 Leg 2: coxa 0-0; basis1-0; exp. 1-0; 1, II, 1; enp. 0-0; 0, 1, 0

Leg 3: coxa 3 (on lobe)

Leg 4: coxa 1 (on lobe)

Leg 5 (Fig. 37A) 2-segmented, consisting of protopod and 1-segmented exopod; protopod with

1 naked outer seta; exopod 222×67 μm (ratio 3.31 : 1), distally with several patches of minute spinules and 4 unequal, stiff setae; lengths of these setae 228, 433, 87, and 166 μm, respectively, from outer to inner. Leg 6 probably represented by 1 small seta in genital aperture.

Male: Body (Fig. 37B) similar to that of female. Mean body length 1.79 mm (1.25-2.07 mm), based on 10 specimens. Prosomal somites gradually narrowed from anterior to posterior. Cepha- lothorax 438×404 μm. Second to fourth pedigerous somites 154×342, 145×273, and 158×258 μm, respectively. Urosome 4-segmented as in female, and slightly shorter than prosome. Fifth pedigerous somite 175×283 μm. Genital somite 154×233 μm. Two free abdominal somites 125×204 and 175×175 μm, respectively. Anal somite with minute spinules on ventral surface. Caudal rami slightly divergent; each ramus 156×62 μm (ratio 2.52 : 1), with 6 caudal setae, and spinules on ventral surface; largest terminal seta 620 μm long, other 5 setae small. Rostrum, antennule, and antenna not different from those of female. Mouthparts, except for maxilliped, as those of female. Maxilliped (Fig. 37C) massive and 4-segmented; first segment short and unarmed; second segment laterally expanded, with 1 large claw-like process and 1 spinules-covered bulge on medial margin and 1 small patch of minute spinules distally; third segment small and unarmed; terminal segment as strong claw, forming pincers along with medial process on second segment, and bearing 2 small setae proximally.

Legs 1-4 as in female. Leg 5 as in female, but exopod more slender than that of female. Leg 6 (Fig. 37D) represented by 1 small seta on genital operculum.

Distribution: Korea. Korea: CN.

Specimens examined: 9♀♀, 12♂♂ (type specimens) from washings of several species of polychaetes epizoic on shells of the oyster Ostrea denselamellosa Lischke, bought at a fish market at Dae- cheon (36°19′44″N, 126°30′18″E) on the coast of the Yellow Sea, 15.viii.2004. Remarks: This species has a 4-segmented abdomen in both sexes. This character differentiates it from its only congener, I. bacescui Sebastian and Pillai, 1974 and also most of other species of the related genera (Kim, 2014).

Genus Likroclausia Ho and Kim, 2003

Ppul-geom-mul-byeo-ruk-sok (뿔검물벼룩속)

Second to fourth pedigerous somites with elongate posterolateral processes. Legs 2-4 with 3-segmented exopod and endopod. Leg 1 with 3-segmented exopod and 2-segmented endopod (endopod 3-segmented in male); distal segment of endopod transformed to a powerful hook; basis with inner distal spine. Maxilliped of female 2-segmented. Other characters as those of the family. 74 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

17. Likroclausia namhaensis Ho and Kim, 2003 (Figs. 38, 39)

Ppul-geom-mul-byeo-ruk (뿔검물벼룩)

Likroclausia namhaensis Ho and Kim, 2003, p. 569, figs. 1-4.

Female: Body (Fig. 38A) cyclopiform, 3.18 mm long. Cephalothorax semicircular, with 6 short sensila on lateral surface and 1 long sensilum at tip of protruded posterolateral corner. Lateral area of tergum of second and third pedigerous somites produced posterolaterally into antler-like large process, bearing 2 short sensila on lateral surface, 2 at terminal end, and 1 longer 1 at tip of inner basal protrusion. Antler-like process on fourth pedigerous somite with 2 short sensila at terminal end and 1 long sensilum on each short protrusion located on inner and outer surface. Urosome (Fig. 38B) 6-segmented. Genital double-somite wider than long, 300×615 μm, with 6 dorsal conical protrusion near area of egg sac attachment. Abdomen 4-segmented, reducing size as progress- ing posteriorly and bearing a transverse row of setules on ventral surface of first to third somites and 2 patches of denticles on either side of ventral surface of anal somite. Caudal ramus 4.84 times longer than wide (310×64 μm), distal edge on ventral surface fringed with spinules; ramus prop- er armed with 1 median dorsal seta, and 4 short and 1 long terminal seta; longest terminal seta 4 times as long as ramus. Rostum truncate anteriorly and oval posteroventrally. Antennule slender and 6-segmented; armature formula 4, 14, 10, 4+aesthetasc, 2+asthetasc, and 7+aesthetasc. Antenna (Fig. 38C) slender as in antennule and 3-segmented; first segment with middle region of outer margin fringed with a row of spinules and armed with 1 distal-inner seta; second segment with longitudinal patch of spinules; third segment 6.67 times as long as wide (240×36 μm) and armed with spinules on 2/3 of proximal inner margin and setules on proximal half of outer margin; distally with 2 setae and 1 spiniform element on inner margin, 2 pectinate setae on outer margin, 1 small subterminal knob, and 4 unequal, terminal, spines with bent tip. Labrum small, covering only tips of mandible; with 2 spiniform, marginal processes in central region. Mandible (Fig. 38D) small; gnathobase armed terminally with smooth but sharply pointed process. Maxillule bilobate, small outer lobe tipped with long seta and longer inner lobe with spinulation at tip and 2 unequal setae at base. Maxilla (Fig. 38E) 2-segmented; robust proximal segment armed with long patch of spinules on posteroventral surface, sigmoid distal segment armed with 2 setae, row of setules between these 2 setae, and patch of spinules at tip. Maxilliped 2-segmented; proximal segment with long medial-terminal seta; distal segment slightly bipartite distally, with fine setules in middle region of inner surface and coarse setules at tip of inner lobe.

Legs 1-4 biramous, with 3-segmented rami except 2-segmented endopod of leg 1 (Fig. 38F). Out- er surface of all leg segments covered with spinules. Coxa of leg 1 with loose row of spinules on ventral surface, but basis bearing a long and dense patch of spinules. Coxa of legs 2, 3, and 4 with short row of spinules at outer-distal corner; while basis of legs 2 and 3 without spinules that of leg 4 bearing long patch of marginal spinules between bases of two rami. Distal segment of endopod of leg 1 modified into a strong, curved, sharp process. Armature formula of legs -1 4 as follows: Leg 1: coxa 0-0; basis 1-1; exp. I-0; 1-1; 3, 4; enp. 0-0; 0-1 Legs 2 and 3: coxa 0-0; basis 1-0: exp. I-0; 0-1; 3, 5; enp. 0-1; 0-2; I, 3 Leg 4 coxa 0-0; basis 1-0; exp. I-0; 0-1; II, 1, 5; enp. 0-1; 0-2; I, 2 Leg 5 2-segmented; armature formula I-0; 2, 2. Leg 6 represented by 2 setae in egg sac attachment area. Cyclopoida: Clausiidae: Likroclausia 75

Fig. 38. Likroclausia namhaensis, female. A. habitus, dorsal; B. urosome, dorsal; C. antenna; D. mandible; E. maxilliped; F. leg 1. Scales: A=0.5 mm, B=0.2 mm, C, F=0.1 mm, D=0.02 mm, E=0.05 mm. 76 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B A

Fig. 39. Likroclausia namhaensis, male. A. habitus, dorsal; B. urosome, dorsal; C. maxilliped. Scales: A=0.5 mm, B=0.2 mm, C=0.05 mm.

Male: Body (Fig. 39A) similar to that of female. Urosome (Fig. 39B) 6-segmented. Antennule, antenna, and mouth organs, except for maxilliped, as in female. Maxilliped (Fig. 39C) consisting of 3 segments and terminal claw; first segment with stout process and 1 seta in inner distal region; second segment tapering distally, with 2 inner setae (1 naked and 1 pinnate) and numerous spinules on inner surface; small third segment unarmed; terminal clal rather stout, with 2 large setae (1 proximal and 1 at 0.4 region)

Legs 1-5 as in female. Leg 6 (Fig. 39B) represented by 1 naked seta on genital operculum.

Distribution: Korea. Korea: GN.

Specimens examined: 8♀♀ (type specimens) collected from washings of Dasybranchus caudatus

Grube collected from intertidal mud flat in Namhae Island (34°49′N 128°03′E), Gyeongsangnam-do, 22.vii.2001; 3♀♀, host and locality same as the types, 31.vii.2003; 2♀♀, from polychaetes, tidal Cyclopoida: Clausiidae: Maxilliclausia 77 flat, Dapo-ri, Geoje I., Gyeongsangnam-do, 23.iv.2008; 1♂ from invertebrate burrows, type locality, 21.x.2010.

Remarks: In the original description of this species, Ho and Kim (2003) mentioned that the most remarkable feature of this species is the possession of a pair of antler-like lateral processes on the second to fourth pedigerous somites. The 2-segmented maxilliped and the modified leg 1 endopod are also unique. This is the first record on the male of this species.

Genus Maxilliclausia Kim, 2014

Ppal-pan-geom-mul-byeo-ruk-sok (빨판검물벼룩속)

Female: Body cylindrical, consisting of cephalothorax, second to fifth pedigerous somites, genital somite, and 4 abdominal somites. Caudal ramus with 6 setae. Egg sac containing mutiserial eggs. Antennule 5- or 6-segmented; first segment with 4 setae. Antenna 4-segmented, non-prehensile, and consisting of coxa, basis, and 2-segmented endopod; terminal segment (second endopodal segment) blunt at tip, with setae, and lacking claw or spine. Labrum with elongate, digitiform posteromedian process. Mandible with extremely elongate distal spine. Maxillule lobate, with adhesion pad and setae. Maxilla, as an attachment organ, large and 2-segmented; second segment transformed to sucking disk. Maxilliped absent. Legs 1 and 2 with 2-segmented rami; coxa without inner element. Leg 3 lobate, bearing 3 setae. Leg 4 absent. Leg 5 represented by 2 setae (Kim, 2014). Associated with polychaetes.

18. Maxilliclausia propria Kim, 2014 (Figs. 40, 41)

Ppal-pan-geom-mul-byeo-ruk (빨판검물벼룩)

Maxilliclausia propria Kim, 2014, p. 105, figs. 7, 8.

Female: Body (Fig. 40A) cylindrical, consisting of cephalothorax, second to fifth pedigerous somite, genital somite, and 4 abdominal somites. Body length 2.75 mm in dissected paratype. So- mites from cephalothorax to genital somite not articulated from 1 another but well defined by deep constrictions between them and with convex lateral margins. Sizes of cephalothorax to fifth pedigerous somite 523×685, 277×454, 277×492, 292×531, and 431×568 μm, respectively. Cephalotho- rax nearly trapezoid, broadening distally, with rounded posterolateral corners and nearly circular dorsal sclerotization band. Fifth pedigerous somite subcircular, longer and broader than preceding metasomal somites. Urosome (Fig. 40B) 5-segmented and consisting of genital somite and 4 abdominal somites. Genital somite 217 ×433 μm, much wider than long; lateral margins projected laterally and tapering; genital aperture large and locating dorsolaterally. Abdomen distinctly narrower than anterior part of body, smooth, and well-segmented; four abdominal somites 192×192, 196×175, 150×158, and 175×154 μm, respectively. Anal somite lacking anal operculum. Caudal ramus (Fig. 40C) 102×50 μm (ratio 2.04 : 1), with 6 naked setae; 1 (seta V) of 2 median terminal setae much larger than other 5 setae, about twice as long as second largest nearby seta (seta IV). Egg sac (Fig. 40D) 1.33×0.36 mm, containing multiserial eggs; each egg about 190 μm in diameter. 78 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Rostrum as broad apical protuberance of cephalothorax, truncated apically, and not articulated at base. Antennule (Fig. 40E) distinctly narrowing distally, 307 μm long, and 6-segmented, but suture line between second and third segment indistinct and existing only on dorsal surface; armature formula of 4, 11, 7, 4, 2+aesthetasc, and 7+aesthetasc; all setae naked; aesthetascs on 2 distal segments shorter than their accompanying setae. Antenna (Fig. 40F) 4-segmented, consisting of coxa, basis, and 2-segmented endopod; coxa short, much broader than long, and unarmed; basis with large outer protrusion ornamented apically with about 15 foliaceous, tapering processes, and with 1 small medio-distal seta; first endopodal segment about 94×32 μm, with large, corrugated adhesion pad on medial surface and 1 outer subdistal seta; second endopodal segment 88×32 μm, broadest at proximal 2/5 region, blunt distally, with large, corrugated adhesion pad along medial surface, and 5 (3 large and 2 small) setae on outer side.

Labrum (Fig. 41A) characteristically with large, digitiform posteromedian process extending to posterior region maxilla. Mandible (Fig. 41B) slender, 53 μm long (excluding distal spine), and armed distally with 1 elongate, rod-shaped spine of 83 μm long; this distal spine blunt at tip, with about 16 small teeth along distal 2/5 of outer margin. Paragnath absent. Maxillule (Fig. 40H) lobate, armed with 3 unequal outer setae and 1 small medial seta; distal part of maxillule expanded and forming large adhesion pad. Maxilla (Figs. 40G, 41C) 2-segmented; proximal segment elongated, unarmed but with large patch of spinules on posterior surface; distal segment stout and forming an sucking disc bearing 1 small seta anterodistally and corrugated adhesion rim along dorso- disal margin of disk. Maxilliped absent.

Legs 1 and 2 (Fig. 41D, E) biramous, with 2-segmented rami; coxa lacking inner seta, but with 1 row of minute spines on ventral (anterior) surface near distal margin; both sides of intercoxal sclerite roundly projected, with several rows of minute setules. Basis of leg 1 with large pinnate outer seta and 3 transverse rows of minute setules on ventral surface. Basis of leg 2 with smaller, naked outer seta and 2 transverse rows of minute setules on ventral surface. Rami of legs 1 and 2 with several longitudinal rows of minute setules along outer surface. Outer setae on exopod of legs 1 and 2 small. Leg 3 (Fig. 41F) as broad lobe tipped with 3 naked setae, medial one of which being thick. Leg 4 absent. Armature formula of legs 1 and 2 as follows: Leg 1: coxa 0-0; basis 1-0; exp. 1-0; 3, 2, 2; enp. 0-1; 0, 1, 1 Leg 2: coxa 0-0; basis 1-0; exp. 1-0; 2, 2, 3; enp. 0-1; 0, 1, 1

Leg 5 (Fig. 41G) represented by 2 naked setae (1 thick and 1 slender). Leg 6 (Fig. 41H) represented by 1 minute seta in genital aperture. Male: Unknown.

Distribution: Korea. Korea: GN, JN.

Specimens examined: 2♀♀ (type specimens) from washings of polychaetes (mixing of several species) collected from a tidal flat (34°49′54″N, 128°02′08″E), Namhae I., Gyeongsangnam-do, 21.vi.2012; 1♀ from washings of polychaetes, tidal flat, Weoldoo, Muan-gun, Jeollanamdo, 4.vii. 2012, collected by H.S. Hwang.

Remarks: This species is the only species of the genus. Kim (2014) defined the genus Maxilliclau- sia by the following diagnostic features: 1) the antenna is of an unusual form, non-prehensile, without any claw on distal segments, but with large adhesion pad on the 2 endopodal segments; 2) the labrum has a large, digitiform posteromedian process; 3) the mandible is armed with a very elongate distal spine; 4) the distal segment of the maxilla is transformed into a sucking disk; and 5) the maxilliped is absent. Cyclopoida: Clausiidae: Maxilliclausia 79

B A D E

G H

Fig. 40. Maxilliclausia propria, female. A. habitus, dorsal; B. urosome, dorsal; C. right caudal ramus, dorsal; D. egg sac; E. antennule; F. antenna; G. mouthparts; H. maxillule. Scales: A, D=5 mm, B=0.1 mm, C, E-G=0.05 mm, H=0.02 mm. 80 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A C

B H F

Fig. 41. Maxilliclausia propria, female. A. labrum; B. mandible; C. maxilla; D. leg 1; E. leg 2; F. leg 3; G. leg 5; H. genital aperture, dorsal. Scales: A, C-H=0.05 mm, B=0.02 mm. Cyclopoida: Clausiidae: Rhodinicola 81

Genus Rhodinicola Levinsen, 1878

Neolb-jeok-bae-geom-mul-byeo-ruk-sok (넓적배검물벼룩속)

Body cylindrical. Urosome 6-segmented (5-segmented in R. gibbosus Bresciani, 1964). Antennule 4-6-segmented. Antenna 3- or 4-segmented. Mandible with 1 or 2 distal elements. Maxilla 2-segmented; proximal segment unarmed, distal segment unarmed or with 1 seta. Maxilliped 2-4-segmented. Legs 1 and 2 with 2- or 3-segmented exopod and endopod. Legs 3 and 4 with 2- or 3-segmented exopod and 1-3 segmented endopod. Leg 1 basis without inner distal element. Other characters as those of the family.

19. Rhodinicola laticauda Ho and Kim, 2003 (Fig. 42)

Neolb-jeok-bae-geom-mul-byeo-ruk (넓적배검물벼룩)

Rhodinicola laticauda Ho and Kim, 2003, p. 574, figs. 5-7.

Female: Body (Fig. 42A) elongate and broad through out, 1.22 mm long (excluding setae on caudal rami) and 357 μm wide (across second pedigerous somite). Cephalothorax shield like, 288×350 μm, and with protruded rostral area. Metasomal somites short and wide, at least 4 times wider than long. Urosome 6-segmented, except for anal somite (135×238 μm), all other somites more than twice wider than long. Caudal ramus longer than wide, 58×38 μm, and carrying 5 (2 outer, 1 dorsal, and 2 terminal) short setae and 1 long terminal seta, which is 4.31 times longer than the ramus. Rostrum with protruded corners posteroventrally. Antennule 6-segmented; armature formula 4,

13, 9, 4, 2+aesthetasc, and 7+aesthetasc. Antenna (Fig. 42B) stocky, 3-segmented, and profusely armed with spinules; proximal segment with bilaterally pectinate spiniform seta on distal-inner corner; middle segment without seta; terminal segment armed with 4 unequal, terminal, hooked spines, 2 subterminal, bipectinate, long, outer setae and 2 subtermnal, short, inner setae.

Labrum small, covering only tips of mandible. Mandible (Fig. 42C) small, gnathobase armed sub- terminally 1 short, smooth, spiniform process and terminally 1 long process with spinules on distal inner margin. Maxillule as thumb-like process covered with denticles and bearing 3 setae on outer surface and 1 seta at base of inner surface. Maxilla (Fig. 42D) 2-segmented, largest oral appendage, greatly inflated basal segment covered with a large patch of denticles on ventral surface; terminal segment subquadrate, with patch of spinules on dorsal margin, seta on posterior surface, fine denticles on distal surface. Maxilliped (Fig. 42E) reduced and 2-segmented; proximal segment with 2 setae on medial margin and row of coarse spiniform elements on anterior margin; distal segment produced into a spiniform process armed with 2 unequal setae at base.

Leg 1 (Fig. 42F), legs 2 and 3, and leg 4 (Fig. 42F) biramous, with 3-segmented rami. Outer surface of all leg segments covered with spinules. Coxa of all legs with row of spinules on outer-distal margin. Basis of all legs with row of spinules on posterior margin, that of leg 1 with additional row on ventral surface. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-0; basis 1-0; exp. I-0; 1-1; III, 1, 3; enp. 0-1; 0-1; I, 2 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 4; enp. 0-1; 0-2; II, 3 82 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B H

D C E

F G

Fig. 42. Rhodinicola laticauda, female. A. habitus, dorsal; B. antenna; C. mandible; D. maxilla; E. maxilliped; F. leg 1; G. leg 4; H. leg 5. Scales: A=0.1 mm, B-E=0.02 mm, F-H=0.05 mm. Cyclopoida: Gastrodelphyidae: Sabellacheres 83

Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-2; I, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, 1, 4; enp. 0-1; 0-1; I, 2

Leg 5 (Fig. 42H) 2-segmented; protopod (100×93 μm) enlarged laterally and without ornamentation; exopod (78×45 μm) smaller, with hooked inner-distal corner and rows of spinules on outer margin, formula of armature 1-0; 4. Leg 6 represented by seta in egg sac attachment area. Male: Unknown.

Distribution: Korea. Korea: JB.

Specimens examined: 1♀ (holotype) from washings of polychaetes living in crevices of intertidal sandst1 at Gyeokpo, Buan-gun, Jeollabuk-do, 8.iv.2001. Remarks: Only a single female, the holotye, of this species was recorded and no further specimen has been collected in spite of a number of field works were carried out at environments similar to that of the type locality.

Family Gastrodelphyidae List, 1889

Keun-bae-geom-mul-byeo-ruk-gwa (큰배검물벼룩과)

Body somite obscure in female but well demarcated in male. Female of most species with internal brood pouch. Urosome of female greatly reduced. Antennule 5-segmented; first and second segments expanded. Antenna 3- or 4-segmented, with 1 bifid hook and 1 simple hook on terminal segment. Mandible small, distally with 3 spiniform elements. Maxillule with 3 distal elements. Maxilla 2-segmented; basal segment unarmed; distal segment slender and spiniform. Maxilliped absent in female, but consisting of 4 segments and terminal claw in male. Legs 1-3 varying with species. Leg 4 as seta-bearing lobe. Leg 5 absent. Associated with sabellid polychaetes. Contain- ing 2 genera: Gastrodelphys (antenna 4-segmented) Sabellacheres (antenna 3-segmented).

Genus Sabellacheres M. Sars, 1861

Kkot-gaet-ji-reong-i-beol-re-sok (꽃갯지렁이벌레속)

Antenna 3-segmented. Brood pouch internal. Legs 1-3 with 3-segmented exopod and endopod. Leg 4 formed as lobe tipped with seta. Other characters as those of the family. 84 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

20. Sabellacheres illgi Dudley, 1964 (Figs. 43-45)

Kkot-gaet-ji-reong-i-beol-re (꽃갯지렁이벌레)

Sabellacheres illgi Dudley, 1964, p. 23, figs. 8-16.

Female: Body (Fig. 43A, B) consisting of large, cylindrical prosome and small urosome. Prosome 3.12 mm long, unsegmented, consisting of narrower anterior part corresponding to cephalosome to third pedigerous somite and longer and broader posterior part corresponding to fourth pedigerous somite. Cephalosome and first to third pedigerous somites discernible by their dorsal tergites. Cephalosome 330×350 μm in dorsal view. Fourth pedigerous somite greatly expanded posteriorly, with eggs inside. Urosome (Fig. 43C) perpendicular to axis of prosome, 4-segmented, and consisting of genital double-somite and 3 free abdominal somites. Fifth pedigerous not defined, completely fused to fourth pedigerous somite. Genital double-somite rectangular, 146 ×223 μm, much wider than long, with weak lateral constriction; genital aperture located dorsally. Three free abdominal somites 56×117, 54×90, and 52×75 μm, respectively. Caudal ramus 48×29 μm (ratio

1.66 : 1), with 5 setae; outer medial distal seta (seta IV) large and other setae small; inner median distal seta (seta V) minute and inserted to base of seta IV. Mating girdle and spermatophore as in Fig. 44E.

Rostrum (Fig. 43D) promienent, clearly defined from cephalosome, plate-like, shaped as Fig. 43D, and directed posteroventrally. Antennule short, 5-segmented; first and second segments strongly expanded; armature formula of 4, 15+spine, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. An- tenna (Fig. 43F, G) 3-segmented, inserted on segment-like elevation of body surface; first and second segments with 1 distal seta each; terminal segment with 1 patch of minute spinules on inner margin, and 2 small setae and 3 unequal claws (large, medium and small) distally; small claw fused to base of large claw.

Labrum (Fig. 44A) consisting of 3 parts: tapering lateral lobes and plate-like median extension.

Mandible (Fig. 44B) with 3 distal elements: small, thick spine, long, spinulose straight spine, and more longer, basally strongly curved element, latter element dentate along one margin and spinulose along other margin, and expanded in distal third. Maxillule (Fig. 44C) curved and armed with

1 subdistal, leaf-like seta and 2 distal, blunt setae. Maxilla (Fig. 44D) 2-segmented; basal segment broad and unarmed; distal segement narrow, with suture in distal third, and ramified distally; one of distal rami (posterior ramus) fringed with finely serrate membrane. Maxilliped absent.

Legs 1-3 (Figs. 44F, G, 45A) with 3-segmented rami; inner coxal seta absent in these legs. Leg 4 (Fig. 45B) rudimentary and represented by small lobe tipped by 1 curved seta. Armature formula of legs 1-3 as follows: Leg 1: coxa 0-0; basis 1-0; exp. I-0; I-1; II, I, 4; enp. 0-1; 0-1; I, 4 Leg 2: coxa 0-0; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-2; II, 3 Leg 3: coxa 0-0; basis 1-0; exp. I-0; I-1; I, I, 4; enp. 0-1; 0-2; I, 2

Leg 5 absent. Leg 6 represented by 2 setae at genital aperture (Fig. 43C).

Male: Body (Fig. 45C, D) cyclopiform, narrow, very different from that of female, and well-segmented. Body length of dissected specimen 1.25 mm. Prosome-urosome division unclear. Cepha- losome triangular, 310×350 μm. Genital somite 104×195 μm, with distal half narrowing posteriorly. Four abdominal somites 48×92, 63×85, 54×72, and 48×68 μm, respectively. First abdominal somite with triangular, plate-like extension on ventral surface (Fig. 45E). Caudal ramus 47×27 μm.

Rostrum, antennule, antenna, and mouthparts, except from maxilliped, as in female. Maxilliped (Fig. 45F) consisting of 4 segments and terminal claw; first and third segment short and unarmed; Cyclopoida: Gastrodelphyidae: Sabellacheres 85

B A

D F

Fig. 43. Sabellacheres illgi, female. A. habitus, dorsal; B. habitus, right; C. urosome, dorsal; D. rostrum; E. antennule; F. antenna; G. distal part of antenna. Scales: A, B=0.5 mm, C=0.1 mm, D-G=0.05 mm. 86 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A C

B D

G F

Fig. 44. Sabellacheres illgi, female. A. labrum; B. mandible; C. maxillule; D. maxilla; E. mating girdle and spermatophores; F. leg 1; G. leg 2. Scales: A-D=0.02 mm, E=0.1 mm, F, G=0.05 mm. Cyclopoida: Gastrodelphyidae: Sabellacheres 87

B A

Fig. 45. Sabellacheres illgi. female: A. leg 3; B. leg 4. male: C. habitus; dorsal; D. habitus; left; E. posterior half of body, ventral; F. maxilliped. Scales: A, F=0.05 mm, B=0.02 mm, C, D=0.2 mm, E=0.1 mm. 88 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans second segment with 1 short seta on inner margin; fourth segment with 1 subdistal seta; terminal claw long, with 2 conical processes proximally. Legs 1-4 as in female. Leg 5 absent as in female. Leg 6 as a lobe bearing 1 distal spine and 1 subdistal seta on genital operculum (Fig. 45E).

Distribution: North Pacific (Pacific coast of North America, and Korea). Korea: GN, JN, JB, CN, GG. Specimens examined: All of examined specimens were collected from the polychaete Pseudopota- milla myriop Marenzeller. 7♀♀, 3♂♂, tidal flat, Sapsi I., Choongcheongnam-do, 26.x.1999; 7♀♀, 3♂♂, Sangrok Beach, Buan-gun, Jeollabuk-do, 16.v.2003; 3 ♀♀, tidal flat, Dapo-ri, Geoje I., Gyeo- ngsangnam-do, 8.v.2008; 3♀♀, tidal flat, Weoldoo, Muan-gun, Jeollanam-do, 17.x.2012; 3♀♀, tidal flat, Daebu I., Gyeonggi-do, 2.i.2014.

Remarks: This is the second record of the species. Dudley (1964) recorded 3 hosts of this copepod from the Pacific coast of North America: Parasabella rugosa (Moore) (as Distylidia rugosa), Pseu- dopotamilla occelata Moore and Megalomma splendida? (Moore) (as Branchiomma burrardum). The female adults of the copepod were found attached to the skin near the mouth of the hosts, but males were found from washings of the hosts. Known hosts belong to the family Sabellidae.

Family Kelleridae Humes and Boxshall, 1996

Gu-meong-beol-re-gwa (구멍벌레과)

Body cyclopiform. Urosome 5-segmented in female and 6-segmented in male. Antennule 7-segmented. Antenna 4-segmented; armature formula 1, 1, 2+claw, and 5+2 claws; claws setiform. Labrum deeply incised. Mandible with tapering lash, with spine row on oblique inner margin. Maxillule with 4 elements. Maxilla with unarmed proximal segment; distal segment with row of teeth along distal margin; inner spine strong and spinulose. Maxilliped 3-segmented; armature formula 0, 2, and 4. Male maxilliped 4-segmented; armature formula 0, 2, 0, 2+claw. Legs 1-3 with 3-segmented exopod and endopod. Leg 4 with 3-segmented exopod and 1-segmented endopod. Leg 5 exopod with 2 setae or 1 spine+1 seta. Containing a single genus, Kelleria.

Genus Kelleria Gurney, 1927

Gu-meong-beol-re-sok (구멍벌레속)

Generic characters as those of the family. Containing 18 known species.

Key to the species of genus Kelleria (based on female)

1. Caudal ramus more than 4 times as long as wide ················································································· 2 - Caudal ramus 3 times as long as wide or less ······················································································· 3 2. Distal segment of maxilla with 9 teeth on distal margin; convex side of mandible with tuft of spinules ············································································································································· K. vaga Cyclopoida: Kelleridae: Kelleria 89

- Distal segment of maxilla with 10 teeth on distal margin; covex side of mandible with row of about 5 spinules ······················································································································· K. portiviva 3. Distal segment of maxilla with 7 teeth on distal margin; leg 5 exopod about 2.6 times as long as wide, with 2 distal setae ··································································································· K. grandisetiger - Distal segment of maxilla with 11 teeth on distal margin; leg 5 exopod about 4.6 times as long as wide, with 1 spine and 1 seta distally ··········································································· K. undecidentata

21. Kelleria grandisetiger Kim, 2006 (Fig. 46)

Keun-teol-gu-meong-beol-re (큰털구멍벌레)

Kelleria grandisetiger Kim, 2006b, p. 1, figs. 1, 2.

Female: Body (Fig. 46A) moderately broad, 1.40 mm long. Greatest width of body 450 μm Pro- some 787 μm long. Cephalothorax with faint dorsal suture line between cephalosome and first pedigerous somite. Urosome (Fig. 46B) 5-segmented. Fifth pedigerous somite with distinct, thumb-like process near base of exopod of leg 5. Genital double-somite 173×163 μm, gradually narrowing posteriorly, with 1 large spinulose seta on lateral side posterior to genital aperture. Three free abdominal somites 54×90, 40×83, and 66×81 μm, respectively. Caudal ramus 95×32

μm (ratio 2.97 : 1), with 6 setae and strongly tapering posterior margin; outer lateral and dorsal setae naed, other setae pinnate. Rostrum with rounded posterior margin. Antennule 7-segmented; armature formula of 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all setae naked; asethetascs thin, setiform. Anten- na 4-segmented; armature formula of 1, 1, 2+claw, and 5+2 claws; terminal segment 75×20 μm; spines on third and terminal segments narrow, setiform.

Labrum with round posterior lobes. Mandible (Fig. 46C) with 10 spines on inner margin, a tuft of about 9 spinues on convex side, more than 10 thick teeth on convex margin; distal lash elongate, with spinules on both margins. Maxillule with 1 subdistal, setiform element and 3 apical setae.

Maxilla (Fig. 46D) with unarmed proximal segment; distal segment with 7 teeth of unequal sizes on distal margin followed by terminal spine; terminal spine demarcated from segment by suture line; anterior seta naked; inner spine strong, extending beyond terminal spine of segment, armed with

4 or 5 spinules on distal margin and 4 spinules on proximal margin. Maxilliped (Fig. 46E) 3-segmented. First segment longest but unarmed. Second segment with angular process proximally on outer margin and armed with 2 large inner setae; proximal seta with large spinules along distal margin and no setules or spinules on proximal margin; distal seta with fine hairs on both margins, those of proximal margin larger. Terminal segment with 4 setae, largest one of them naked and not articulated at base.

Legs 1-3 with 3-segmented rami. Leg 4 (Fig. 46F) with 3-segmented exopod and 1-segmented endopod. Leg 4 endopod 86×28 μm, with small pointed process on outer margin and patch of minute spinules on outer surface; 2 terminal spines 55 and 43 μm, respectively. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 2

Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5 (or III, I, 5); enp. II, 1 90 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A F

G E

C D

Fig. 46. Kelleria grandisetiger, female. A. habitus, dorsal; B. urosome, dorsal; C. mandible; D. maxilla; E. maxilliped; F. leg 4; G. right leg 5 and genital aperture, dorsal. Scales: A=0.2 mm, B-G=0.05 mm. Cyclopoida: Kelleridae: Kelleria 91

Leg 5 (Fig. 46G) exopod 87×55 μm (ratio 2.64 : 1), slightly swollen proximally in lateral view, with small spinules on surface; 2 terminal setae 78 μm (inner) and 54 μm (outer). Leg 6 (Fig. 46G) represented by 2 setae on genital operculum. Male: Unknown.

Distribution: Korea. Korea: JJ.

Specimens examined: 3♀♀ (type specimens) from invertebrate burrows (mostly of polychaetes and decapods crustaceans) on intertidal zone in Seogwipo Port, Jeju Island, 17.ix.2005. Remarks: The most important characteristic features of this species is the presence of a large seta on the lateral side of the genital double-somite and the thumb-like process on the fifth pedigerous somite near the base of the exopod of leg 5 (Kim, 2006).

22. Kelleria portiviva Kim, 2006 (Fig. 47)

Hang-gu-gu-meong-beol-re (항구구멍벌레)

Kelleria portiviva Kim, 2006b, p. 8, figs. 5, 6.

Female: Body (Fig. 46A) rather narrow, 1.23 mm long. Greatest width 0.41 mm. Prosome 0.73 mm long. Cephalothorax with dorsal suture line delimiting cephalosome and first pedigerous somite. Urosome (Fig. 47B) 5-segmented. Genital double-somite 175×158 μm, with distinct anterior expansion and 1 lateral seta on each side of posterior part of anterior expansion. Three free abdominal somites 52×73, 36×67, and 60×67 μm, respectively. Caudal ramus 120 ×29 μm (ratio 4.14: 1), with 6 setae and oblique posterior margin; outer lateral seta naked, other setae pinnate. Rostrum elongate and tapergin, with rounded posterior apes. Antennule 7-segmented; armature formula 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; several setae on distal segments pinnate distally; aesthetascs thin, setiform. Antenna 4-segmented; armature formula 1, 1, 2+claw, and 5+2 claws; outer margin of second and terminal segments spinulose; claws of third and terminal segments setiform; terminal segment 78×21 μm.

Labrum with prominent posterior lobes and deep median incision. Mandible (Fig. 47C) with narrow hyaline membrane along outer margin; inner margin with 12 spines; convex outer corner with tuft of small spinules; distal margin with thick teeth; distal lash elongate, with spinulose margins. Maxillule armed with 1 inner setiform process, 3 distal setae, and narrow membrane along distal outer margin. Maxilla (Fig. 47D) with unarmed basal segment; distal segment with 10 or 11 teeth of unequal sizes on distal margin; anterior seta naked; inner spine strong, extending slightly beyond tip of terminal spine of segment, armed on both margins with.many spinules which are increas- ingly larger distally. Maxilliped (Fig. 47E) 3-segmented; first segment unarmed; second segment expanded medially and armed with 2 large inner setae: proximal seta with large spinules on proximal part of distal margin, 2 spinules on proximal part of proximal margin, and fine hairs along distal three-fourths of distal margin; distal seta with fine hairs on both margins; third segment with 4 setae, one of them being large, without articulation at base.

Legs 1-4 with segmentation and setation as in preceding species, Leg 4 (Fig. 47F) endopod 64×19 μm, with small pointed process on outer margin and row of minute spinules on distal part of outer 92 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B E

Fig. 47. Kelleria portiviva, female. A. habitus, dorsal; B. urosome, dorsal; C. mandible; D. maxilla; E. maxilliped; F. leg 4; G. left leg 5 and genital aperture, dorsal. Scales: A=0.2 mm, B, E, F=0.05 mm, C, D, G=0.02 mm. Cyclopoida: Kelleridae: Kelleria 93 margin; 2 terminal spines 72 and 50 μm long, respectively.

Leg 5 exopod 79×17 μm (ratio 3.57 : 1), with parallel lateral margins, spinules on distal half of outer side, and distally with 1 spine (31 μm long) and 1 seta (54 μm long). Leg 6 (Fig. 47G) represented by 1 spiniform seta and 1 blunt process on genital operculum. Male: Unknown.

Distribution: Korea. Korea: JJ.

Specimens examined: 3♀♀ (type specimens) from invertebrate burrows (mostly of polychaetes and decapods crustaceans) on intertidal zone in Seogwipo Port, Jeju Island, 17.ix.2005. Remarks: In this species, the exopod of leg 5 has parallel lateral margins without proximal swelling or inner process and is armed distally with 1 short compound spine and 1 seta, as distinguishing features. It is very similar to K. vaga Kim, 2000. Differences between the two are: 1) the caudal ramus of this species is 4.14 times as long as wide (vs. 4.55 times in K. vaga); 2) leg 5 exopod of K. portiviva is 3.57 times as long as wide (vs. 3.30 times in K. vaga); 3) the distal segment of the maxilla of this species has 10 or 11 teeth (vs. 9 teeth in K. vaga); and 4) the genital aperture of this species is widely exposed (vs. closed in K. vaga).

23. Kelleria undecidentata Kim, 2006 (Fig. 48)

Yeol-han-ga-si-gu-meong-beol-re (열한가시구멍벌레)

Kelleria undecidentata Kim, 2006b, p. 4, figs. 3, 4.

Female: Body (Fig. 48A) with relatively broad prosome and narrow urosome. Body length 1.04 mm. greatest width 0.42 mm. Prosome 0.65 mm long. Urosome (Fig. 48B) 5-segmented. Fifth pedigerous somite 97 μm wide. Genital double-somite 148×121 μm, with expanded anterior and narrower posterior parts, and with 1 small seta laterally. Three free abdominal somites 50×58,

33×53, and 52×63 μm, respectively. Caudal ramus 85×29 μm (ratio 2.93 : 1), with 6 setae and strongly tapering posterior margin; outer lateral and dorsal setae naked, other setae pinnate; 2 median terminal setae expanded, with hyaline membrane on both sides. Rostrum strongly tapering, with pointed posterior apex. Antennule 7-segmented; armature formula 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all setae naked; aesthetascs thin, setiform. Antenna 4-segmented;armature formula 1, 1, 2+claw, and 5+2 claws; claws on third and terminal segments setiform; terminal segment 65×21 μm.

Labrum with prominent posterior lobes, and deep and wide median incision. Mandible (Fig. 48C) with about 13 spines on inner margin, row of about 5 minute spinules on outer corner, and 15 thick teeth on convex marginl distal lash elongate, with spinulose margins. Maxillule with 1 subdistal setiform process, 3 apical setae, and hyaline membrane on distal region of outer margin.

Maxilla (Fig. 48D) 2-segmented; proximal segment unarmed but with 1 blunt process on outer side; distal segment with 11 unequal teeth on distal margin, 2 of which located on terminal spine; terminal spine demarcated from segment by suture line; anterior seta naked; inner spine strong, extending to tip of terminal spine of segment, armed on both margins with many spinules becoming larger from proximal to distal ones. Maxilliped 3-segmented. First segment unarmed. Second seg- 94 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

F B A

C E

Fig. 48. Kelleria undecidentata, female. A. habitus, dorsal; B. urosome, dorsal; C. mandible; D. maxilla; E. maxilliped; F. leg 4; G. right leg 5 and genital aperture, dorsal. Scales: A=0.1 mm, B, E, F=0.05 mm, C, D, G=0.02 mm. Cyclopoida: Kelleridae: Kelleria 95 ment expanded medially and with 2 large inner setae; proximal seta with large spinules on proximal part of both margins and fine hairs along distal tho-thirds of distal margin; distal seta with fine hairs on both margins. Terminal segment with 4 setae, largest one of them lacking suture at base.

Legs 1-3 with 3-segmented rami. Leg 4 (Fig. 48F) with 3-segmented exopod and 1-segmented endopod. Thrid endopodal segment of legs 2 and 3 with prominent terminal process. Leg 4 endopod 64×19 μm, with small pointed process on outer margin and row of minute spinules on distal part of outer marginl 2 terminal spines 57 and 38 μm, respectively. Armature formula of legs 1-4 as in preceding species.

Leg 5 (Fig. 48G) exopod widest across procimal third in lateral view, 79×17 μm (ratio 4.66 : 1), terminally with 1 spine (43 μm long) and 1 seta (30 μm long). Leg 6 (Fig. 48G) as 1 spiniform seta and 1 pointed process on genital operculum. Male: Unknown.

Distribution: Korea. Korea: JJ.

Specimens examined: 4♀♀ (type specimens) from invertebrate burrows (mostly of polychaetes and decapods crustaceans) on intertidal zone in Seogwipo Port, Jeju Island, 17.ix.2005. Remarks: This species is similar to K. grandisetiger, but differentiated in having the smaller body size, a smaller seta on the genital double-somite, and the more slender exopod of leg 5 than in K. grandisetiger.

24. Kelleria vaga Kim, 2000 (Fig. 49)

Tteo-dol-i-gu-meong-beol-re (떠돌이구멍벌레)

Kelleria vaga Kim, 2000a, p. 402, figs. 20-22.

Female: Body (Fig. 49A) narrow, 1.01 mm long. Greatest width 331 μm. Prosome spindle-shaped and consisting of cephalosome and 4 metasomites. Urosome (Fig. 49B) 5-segmented. Fifth pedigerous somite 102 μm wide. Genital double-somite 125×160 μm, with distinctly expanded anteri- or two-thirds and narrower posterior part. Three abdominal somites 39×63, 27×58, 48×54 μm, respectively, without ornamentation. Caudal ramus elongate, 100×22 μm (4.55 : 1), with parallel margins. Lateral seta located slightly posterior to mid-point of caudal ramus. Outer lateral and inner dorsal setae naked. Other setae plumose. Outer posterior corner of caudal ramus with a small duct and terminal pore. Rostrum narrow and long, with round posterior tip. Antennule 249 μm long, 7-segmented, with armature formula of 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Plumose setae are

1 on fifth and sixth segment, and 4 on terminal segment. Antenna (Fig. 49C) slender, 4-segmented, with armature formula of 1, 1, 3+claw, and 5+2 claws. Claws on third and terminal segments slender and setiform. Second and terminal segments with minute spinules on outer margin, each segment about 2.4 and 3.3 times longer than wide, respectively. Labrum deeply incised, with round posterior corners; posterior margin of each lobe unornamented. Mandible (Fig. 49D) with tapering, moderately long lash; proximal notch shallow; inner margin oblique to lash, with rather large spinules; outer margin denticulate and proximally with a 96 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A C B

Fig. 49. Kelleria vaga, female. A. habitus, dorsal; B. urosome, dorsal C, antenna; D mandible; E. maxilla; F. maxilliped; G. leg 5 exopod. Scales: A, B=0.1 mm, C=0.05 mm, D-G=0.02 mm. Cyclopoida: Kelleridae: Kelleria 97 row of several small spinules. Maxillule with 3 terminal spinulated setae, 1 lateral seta, and serrate membrane on distal half of outer margin. Maxilla (Fig. 49E) 2-segmented; first segment unarmed; second segment with 1 slender seta and 1 large, terminally bifurcate, spinulated seta; distal portion of second segment curved perpendicularly to proximal portion, tapering and sharply pointed, with

9 or 10 large spines of unequal sizes. Maxilliped (Fig. 49F) 3-segmented. First segment unarmed and longest. Second segment with 2 large setae of similar size; proximal seta with long and thick setules both sides of proximal third, and small setules on distal margin of remaining two-thirds.

Third segment small, with 1 long (86 μm), naked seta, 1 small plumose seta, and 2 more smaller, naked setae. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 1-segmented endopod; inner seta on coxa minute; exopod with 3 or 2 spines on outer margin (formula III, I, 5 or II, I, 5); endopod with or without spinule-like small process on outer margin; median one of 2 terminal spines larger than the inner. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 2

Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5 (or III, I, 5); enp. II, 1

Leg 5 composed of 1 seta near corner of fifth pedigerous somite and exopod; exopod (Fig. 49G)

50×15 μm (3.3 : 1), margins nearly parallel, with small angular proximal protuberance on inner margin, 1 terminal spiniform process, and terminally 1 spine (31 μm) and 1 naked seta (45 μm). Leg 6 represented by 1 seta in genital aperture and distantly isolated, 1 spinule and 1 setule; the latter two located on dorsal surface of genital double-somite. Male: Body more slender than that of female, 877 μm long. Urosome 6-segmented. First urosomal somite 75 μm wide. Genital somite 131×130 μm, as long as wide, with round lateral margins. Four abdominal somites 35×56, 31×48, 23×44, and 35×46 μm, respectively. Caudal ramus 75×21

μm (3.57 : 1). Rostrum and antenna as in female. Antennule with 3 more aesthetascs than in female: 2 on second and 1 on fourth segment. All aesthetascs extending over end of antennule. Mouth parts as in female, except for maxilliped. Maxilliped consisting of 3 segments and terminal claw. First segment unarmed. Second segment with 2 setae at proximal third of inner margin and spinules on entire inner margin. Third segment unarmed. Claw long, 152 μm, evenly curved, proximally with 2 moderately long setae.

Leg 1 with 2 spines and 4 setae (II, 4) on terminal endopodal segment. Legs 2-4 as in female. Leg

5 exopod 23×10 μm (2.30 : 1), without proximal protuberance; terminal spine 21 μm, and seta 34 μm. Leg 6 represented by 2 setae and 1 spinule on genital operculum.

Distribution: Korea. Korea: GG.

Specimens examined: 3♀♀, 1♂ (type specimens) from stagnant water on the intertidal mud flat at Jakyak-do Island, Inchon, 01.ix.1996.

Remarks: This species resembles Kelleria camortensis Sewell, 1949 and K. pectinata (A. Scott, 1909), but K. vaga can be distinguished from these two congeners by having the long caudal rami of 4.76 times as long as wide and a short exopod of female leg 5. 98 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Family Lamippidae Joliet, 1882

San-ho-sok-sal-i-gwa (산호속살이과)

Body vermiform or stelliform, without segmentation. Prosome-urosome division absent. Caudal ramus with at most 5 setae or setiform processes. Antennule unsegmented, Antenna 1-4-segmented, with terminal claw. Mouthparts rudimentary, located in oral prominence. Legs 1 and 2 usually with 1- or 2-segmented exopod; endopod absent or rudimentary. Legs 3-5 absent. Genital aperture located ventrally. Internally associated with octocorals.

Genus Enalcyonium Olsson, 1868

Yeon-san-ho-sok-sal-i-sok (연산호속살이속)

Body cylindrical or fusiform. Caudal ramus not articulated from body, with 3-5 process-like setae. Antennule usually unsegmented, with at most 15 setae. Antenna 3-segmented, with terminal claw. Maxilliped absent. Legs 1 and 2 with 1- or 2-segmented exopod; endopods absent or rudimentary. Legs 3-5 absent. Containing about 35 known species associated with gorgonaceans, al- cyonaceans, and pennatulaceans.

25. Enalcyonium digitigerum Ho, 1984 (Fig. 50)

Gon-bong-ttal-gi-san-ho-sok-sal-i (곤봉딸기산호속살이)

Enalcyonium digitigerum Ho, 1984, p. 41, figs. 15, 16; Kim, 1998, p. 400, fig. 161.

Female: Body (Fig. 50A, B) vermiform, without segmentation, gradually broadened posteriorly. Body length 1.67 mm. Cephalic region with faint sclerotization sculptureon dorsal surface. Genital aperture located ventrally at 75% region of trunk. Caudal ramus root-like, with 5 setiform branches.

Rostrum absent. Antennule (Fig. 50C) unsegmented, tapering, with about 12 setae and terminating in long setiform process. Antenna (Fig. 50D) consisting of 3 segments and terminal claw; first segment longest; first and second segments unarmed; second and third segments nearly equal in length; third segment distinctly narrower than second segment, with 1 small seta proximally. Ter- minal claw about haf as long second segment, curved distally, with 1 small seta proximally.

Mouthparts reduced and forming oral prominence (Fig. 50E); mouth organs hardly visible.

Legs 1 and 2 (Fig. 50F, G) with 2-segmented exopod; endopods absent. Leg 1 positioned just posterior to oral prominence, and leg 2 positioned at 25% region of trunk. Leg 1 with 1 outer seta on protopod (fused coxa and basis); first exopodal segment also with 1 outer seta; second exopodal segment with 1 outer proximal seta and 2 large distal spines, outer one of latter with 1 long setiform process. Leg 2 with 1 large, bifurcate seta on ventral surface of protopod; first exopodal segment with 2 setae; second exopodal segment with 2 spines each with 1 distal, setiform process.

Male: Body (Fig. 50H) as in female in form, but smaller, 1.18 mm long. Genital operculum slit- Cyclopoida: Lamippidae: Enalcyonium 99

B A

H D

G F

Fig. 50. Enalcyonium digitigerum. female: A. habitus, dorsal; B. habitus, ventral; C. antennule; D. antenna; E. mouthparts; F. leg 1; G. leg 2. male: H. habitus, ventral. Scales: A, B, H=0.2 mm, C-G=0.02 mm.

shaped. Spermatophore kidney-shaped. Caudal rami, oral prominence, and legs 1 and 2 as those of female.

Distribution: Japan, Korea. Korea: GB, GN, JN. 100 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Specimens examined: Many specimens from the alcyonacean Bellonella rubra Brundin, Naegal-do, Sinan-gun, Jeollanam-do, 28.vi.2000; 6♀♀ from Bellonella rigida Putter, Yangpo Port, Pohang-shi, 24.vi.2011. Remarks: In Korea, this species is the only known species of the family Lamippidae, but a number of other species wait to be described.

Family Macrochironidae Humes and Boxshall, 1996

Keun-son-beol-re-gwa (큰손벌레과)

Body cyclopiform. Urosome 5-segmented in female and 6-segmented in male. Antennule 7-segmented. Antenna 3- or 4-segmented; third segment with 3 setae and fourth segment with 1 or 2 claws plus setae. Mandible with straight inner margin which is usually perpendicular to distal lash. Maxillule with 4 elements. Maxilla 2-segmented; distal segment terminating in lash. Max- illiped 3-segmented in female and 4-segmented in male. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 1-segmented (occasionally 2-segmented) endopod bearing 2 spines (2 spines and 1 seta in Macrochiron anomalum Kim, 2000). Leg 5 with free exopod bearing 2 distal elements. Containing 4 genera. Hosts are cnidarians, mainly medusae.

Key to the genera of family Macrochironidae (adapted from Humes and Boxshall, 1996)

1. Antenna 3-segmented ································································································································ 2 - Antenna 4-segmented ································································································································ 3 2. Antenna with 1 terminal claw ····························································································· Sewellochiron - Antenna with 2 pectinate claws, or with 1 pectinate claw and 1 claw-like seta ············ Macrochiron 3. Antenna with 1 terminal claw ························································································ Paramacrochiron

- Antenna with 2 terminal claws (except 1 claw in P. fusicolum) ······························ Pseudomacrochiron

Genus Macrochiron Brady, 1872

Keun-son-beol-re-sok (큰손벌레속)

Body cyclopiform. Antenna massive and 3-segmented, terminally with 2 pectinate claws, or with 1 pectinate claw and 1 claw-like seta. Other characters as those of the family. Containing 14 known species. Cyclopoida: Macrochironidae: Macrochiron 101

26. Macrochiron anomalum Kim, 2000 (Fig. 51)

Teol-keun-son-beol-re (털큰손벌레)

Macrochiron anomalum Kim, 2000a, p. 410, figs. 26-28.

Female: Body (Fig. 51A) relatively narrow. Body length 1.19 mm. Greatest width 0.42 mm. Pro- some occupying 61% of body length and five-segmented. Urosome (Fig. 51B) 5-segmented as well. Fifth pedigerous somite 159 μm wide. Genital double-somite 200×162 μm, with anterior two- thirds expanded broadly and roundly. Posterior third slightly tapering. Genital aperture located dorsally in the middle of anterior broad part. Genital double- and first 2 abdominal somites with serrate membranous fringe along posterodorsal and posteroventral margins. Three abdominal somites 63×72, 45×63, and 37×55 μm, respectively. Anal somite (Fig. 51C) with minute spinules along posterodorsal and posteroventral margins. Caudal ramus (Fig. 51C) 55×22 μm (2.50 : 1), terminating in a duct with terminal pore at tip. Rostrum short, wider than long, with angular posterior tip. Antennule 7-segmented, 313 μm long, with armature formula of 4, 13, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all setae naked.

Antenna (Fig. 51D) massive, 3-segmented. First and second segment each much wider than long, with 1 inner distal seta respectively. Third segment with ten setae; 3 setae on inner margin broad, proximal two of them with spinulated inner margin, and remaining 1 with serrate inner margin; of seven terminal setae, 1 large, curved with serrate concave margin; remaining six setae comprising 2 long, thin, simple setae, 1 small naked seta, and 3 moderately thick, spiniform setae; the latter 3 setae with minute spinules on concave margin.

Labrum with 2 round, unornamented posterior lobes. Mandible (Fig. 51E) with deep proximal notch; terminal lash long, thin, and well defined proximally; inner margin perpendicular to terminal lash, with onw row of spinules; outer margin with proximal row of spinules and continued to serrate margin. Maxillule tapering, with 3 terminal and 1 subterminal setae. Maxilla (Fig. 51F) 2-segmented. First segment unarmed. Second segment drawn out into moderately long lash bearing serrate convex margin, with 2 setae, one of the latter large and spinulated on both margins.

Maxilliped (Fig. 51G) 3-segmented. First segment longest, slightly tapering and unarmed. Second segment with spinules and 2 extremely unequal setae. Third segment slender and tapering, with 1 proximal and 1 distal setae.

Legs 1-3 with 3-segmented rami. Leg 4 (Fig. 51H) with 3-segmented exopod and 1-segmented endopod. Seta on outer margin of basis of legs 1-4 small. Inner posterior corner of basis of legs 3 and 4 pointed. Endopod of leg 4 103×31 μm (3.32 : 1); inner of terminal spines more than twice as long as outer 1; proximal seta on inner margin slightly longer than width of endopod; outer margin with small spinule at proximal one-third. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. II, 1

Leg 5 consisting of 1 small seta on fifth pedigerous somite and exopod. Exopod (Fig. 51I) 131×30

μm (4.37 : 1), tapering, with minute spinules all over surface and with 1 spine (61 μm) and 1 seta (42 μm) terminally. Leg 6 represented by 3 spinules in genital aperture. Male: Body distinctly narrower than that of female. Length 858 μm. Segmentation between cephalosome and first pedigerous somite incomplete. Urosome 6-segmented. Fifth pedigerous so- 102 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A G

I C

E H

Fig. 51. Macrochiron anomalum, female. A. habitus, dorsal; B. urosome, dorsal; C. anal somite and caudal rami, dorsal; D. antenna; E. mandible; F. maxilla; G. maxilliped; H. leg 4; I. leg 5 exopod. Scales: A, B=0.1 mm, C, E-G=0.02 mm, D, H, I=0.05 mm. Cyclopoida: Macrochironidae: Pseudomacrochiron 103 mite 79 μm wide. Genital somite distinctly longer than wide, 123×96 μm. Four abdominal somites 46×48, 38×44, 29×41, and 31×42 μm, respectively. All abdominal somites with denticulated posteroventral margin. Caudal ramus 54×21 μm (2.57 : 1). Outer one of 2 median terminal setae truncate. Antennule with 3 additional aesthetascs: 2 on second, and 1 on third. All aesthetascs extending beyond distal end of antennule. Antenna with 1 extra seta on inner margin of third segment (thus 10 setae). Maxilliped consisting of 3 segments and terminal claw. First segment distinctly tapering and unarmed. Second segment with 1 longitudinal row of spinules and 2 extremely unequal setae on inner side. Claw long, slender, evenly curved, proximally with 1 long seta and 1 minute spinule. Other mouth parts as in female.

Terminal endopodal segment of leg 1 armed with 2 spines and 4 setae (II, 4; instead of I, 5 as in female). Legs 2-4 as in female. Leg 5 exopod relatively short, 33×12 μm (2.75 : 1), with nearly parallel margins. Leg 6 represented by 2 setae on genital operculum.

Distribution: Korea. Korea: GG.

Specimens examined: 2♀♀, 5♂♂ (type specimens) from stagnant water on the intertidal mud flat at Jakyak-do I., Incheon, 29.ix.1996. Remarks: The presence of a seta on the inner margin of leg 4 endopod is the most striking feature of this species, because none of its congeners has been reported to have this setation.

Genus Pseudomacrochiron Reddiah, 1969

Heot-keun-son-beol-re-sok (헛큰손벌레속)

Body narrow. Antenna 4-segmented, with 2 terminal claws. Other characters as those of the family. Containing 8 known species.

27. Pseudomacrochiron urostenum Kim, 2000 (Fig. 52)

Heot-keun-son-beol-re (헛큰손벌레)

Pseudomacrochiron urostenum Kim, 2000a, p. 415, figs. 29-31.

Female: Body (Fig. 52A) slender, 1.03 mm long. Prosome 5-segmented. Segmentation between cephalosome and first pedigerous somite distinct. Lateral margins of second pedigerous somite almost straight, its posterior corners pointed. Lateral margins of third and fourth pedigerous somites with narrow membranous fringe. Urosome (Fig. 52B) slender and 5-segmented. Fifth pedigerous somite 97 μm wide. Genital double-somite 132×98 μm, with moderately expanded anterior part and narrower posterior part; genital aperture located at area of Greatest width in expanded anterior part. Three abdominal somites 33×56, 27×53, and 67×51 μm, respectively. Anal somite 104 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

E A B

Fig. 52. Pseudomacrochiron urostenum, female. A. habitus, dorsal; B. urosome, dorsal; C. rostrum; D. maxilla; E. leg 4. Scales: A=0.1 mm, B, C, E=0.05 mm, D=0.02 mm.

distinctly longer than wide, longer than combined preceding 2 abdominal somites, with minute spinules on posteroventral margin. Caudal ramus elongate, 120 ×19 μm (6.32 : 1), with 6 setae. Outer lateral seta located at 61% length of caudal ramus. Rostrum longer than wide, tapering, with angular posterior tip. Antennule 7-segmented, 259 μm long, with armature formula of 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. All setae naked. Antenna slender and 4-segmented, with armature formula of 1, 1, 2+1 claw, and 5+2 claws. Claws on third and fourth segments slender, setiform. Fourth segment about 55×18 μm (3.06

: 1). Labrum widely incised; posterior margins of both lobes fringed with narrow membrane. Mandi- ble with deep proximal notch; terminal lash long, slender, spinulated, and well defined proximally; inner margin with 1 row of spinules; outer margin with proximal row of spinules and continued to serrate margin. Maxillule tapering, with 3 terminal and 1 subterminal setae. Maxilla (Fig. 52D) Cyclopoida: Macrochironidae: Pseudomacrochiron 105

2-segmented. First segment unarmed. Second segment drawn out into relatively short lash bearing spines on convex margin and 1 spiniform process on concave margin, with 2 setae and 1 proximal setule; seta on concave margin small and plumose. Maxilliped 3-segmented. First segment longest and unarmed. Second segment with 2 identical setae on inner margin proximal to middle.

Third segment tapering and terminating in sharp point, with 2 proximal (1 being spiniform) and 2 distal setae.

Legs 1-3 with 3-segmented rami. Leg 4 (Fig. 52E) with 3-segmented exopod and 1-segmented endopod. Endopod of leg 4 shorter than combined first 2 segments of exopod, 52×19 μm, with small spiniform process on outer margin; both margins with hairs; two terminal spines subequal. Arma- ture formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. II Leg 5 consisting of 1 seta on fifth pedigerous somite and exopod. Exopod small, lobate, 26×16

μm (1.63 : 1), with several setules on convex inner margin and terminally with 1 long spine (65 μm),

1 seta (45 μm) and 1 spiniform process. Leg 6 represented by 1 seta and 2 spinules in genital aperture. Male: Body 0.67 μm long. Greatest width 0.20 mm. Lateral margins of prosome nearly parallel. Urosome 2-segmented. Fifth pedigerous somite 62 μm wide. Genital somite 107×93 μm. Four abdominal somites 21×44, 22×38, 19×36, 33×34 μm, respectively. Caudal ramus 63×22 μm (2.86 : 1). Outermost terminal seta transformed, basally thickened. Antennule with 3 additional aesthetascs: 2 on second and 1 on third. All aesthetascs large, extending beyond end of antennule. Antenna spinulated on inner surface of second segment, otherwise, identical to that of female. Maxilliped with unarmed first and third segments. Second segment proximally narrow and distally expanded, with 1 longitudinal row of spinules and 2 setae on inner side; the latter 2 located proximal to middle of segment, and one of them slightly expanded in proximal two-thirds. Claw long, weakly curved, as long as proximal 3 segments combined, with proximally 1 seta and 1 setule.

Leg 1 with terminal endopodal segment bearing 2 spines and 4 setae (II, 4; instead of I, 5 as in female). Legs 2-4 as in female. Leg 5 exopod ovoid, 19×12 μm (1.58 : 1), both margins convex. Ter- minal spine 32 μm, and seta 37 μm. Leg 6 represented by 2 setae and 1 spiniform process on genital operculum.

Distribution: Korea. Korea: GG.

Specimens examined: 7♀♀, 5♂♂ (type specimens) collected from stagnant water on the intertidal mud flat at Jakyak-do I., Inchon, 29.ix.1996. Remarks: This species has long caudal rami with a length to width ratio of 6.32 in contrast to less than 4 of its congeners. Its leg 5 exopod is very short, with the ratio of length to width being 1.63, while this ratio in the other known species is more than 3 (Kim, 2000). 106 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Family Polyankyliidae Ho and Kim, 1997

Bal-top-geom-mul-byeo-ruk-gwa (발톱검물벼룩과)

Body varuous (cyclopiform, vermiform, or cylindrical). Antennule at most 6-segmented; first segment with at most 3 setae. Antenna 4-segmented; third segment with 1 claw and 2 setae; fourth segment with 3 claws and 3 setae (4 claws and 3 setae in Stockia). Mandible with strongly tapering blade and curved row of spinules on inner margin. Maxillule with 4 elements (2 elements in Oc- tophiophora). Maxilla 2-segmented; distal segment with short, dentate distal lash and large inner spine. Maxilliped of female 3-segmented; second segment usually with 2 enlarged setae; third segment tapering or terminating in claw. Male maxilliped consisting of 3 segments and long terminal claw; 2 setae on second segment enlarged or transformed to stout spines. Legs 1 and 2 usually with 3-segmented rami. Leg 3 usually with with 3-segmented exopod and at most 2-segmented endopod. Leg 4 with 1-3-segmented exopod, without endoopod, Leg 5 with free exopod bearing 1 spine and 1 seta; sometimes exopod absent. Associated with polychaetes.

Key to the genera of family Polyankyliidae (based on females; Kim 2014)

1. Legs 1-3 with 2-segmented exopod ······························································································ Stockia - Legs 1-3 with 3-segmented exopod ········································································································ 2 2. Body vermiform; antennule 5-segmented; endopod of legs 1-3 rudimentary ········ Octophiophora - ‌Body not modified; antennule 6-segmented; endopod of legs 1-3 well-developed and 2-segmented ························································································································································· 3 3. Leg 4 with 3-segmented exopod; legs 1-3 without inner coxal seta; fourth pedigerous somite narrow, without epimera ···································································································· Terebelliphilus - Leg 4 with 1- or 2-segmented exopod; legs 1-3 with inner coxal seta; fourth pedigerous somite with epimera ·············································································································································· 4 4. Terminal segment of female maxilliped terminated in long spiniform claw or spine; leg 5 with free exopod bearing 2 elements ······························································································ Polyankylis - Terminal segment of female maxilliped simple and tapering, with 2 small setae; leg 5 represented by 3 setae ································································································································· Sewelloya

Genus Polyankylis Ho and Kim, 1997

Bal-top-geom-mul-byeo-ruk-sok (발톱검물벼룩속)

Body cyclopiform. Antennule 6-segmented with 2, 6, and 6 setae from first to third segments, respectively. Antenna with 1 spine plus 2 setae on third segment and 3 spines plus 2 or 3 setae on terminal segment. Mandible with 2 outer elements on convex side. Maxilliped with 2 spines on second segment. Legs 1-3 with 3-segmented exopod and 2-segmented endopod. Leg 4 with 1- or 2-segmented exopod; endopod lacking. Cyclopoida: Polyankyliidae: Polyankylis 107

28. Polyankylis orientalis Ho and Kim, 1997 (Figs. 53-55)

Dong-yang-bal-top-geom-mul-byeo-ruk (동양발톱검물벼룩)

Polyankylis orientalis Ho and Kim, 1997, p. 432, figs. 1-3.

Female: Body (Fig. 53A) 1.04 mm long, with truncated rostral area. Prosome dorsoventrally de- pressed. Rostrum broadly rounded posteroventrally. Urosome (Fig. 53B) 5-segmented. Genital double-somite 169 μm long and 142 μm wide. Caudal ramus 75×22 μm, ratio 3.4 : 1, with 6 setae. Proximalmost seta located dorsally at midlength of ramus. Inner median terminal seta elongated, 338 μm long.

Antennule (Fig. 53C) 6-segmented; armature formula 2, 6, 6, 4+aesthetasc, 2+aesthetasc, and

7+aesthetasc. Antenna (Fig. 53D) 4-segmented, with coxa and basis fused to form first segment (coxobasis) and carrying 3-segmented endopod; coxobasis bearing 1 seta and recurved hook at mediodistal corner; first endopodal segment with medial seta and row of outer spinules; second endopodal segment with setiform claw and 2 unequal setae; third endopodal segment tipped with 3 claws and 3 setae.

Labrum (Fig. 53E) deeply incised. Mandible (Fig. 53F) forming short and strongly tapering blade with row of strong teeth on convex edge and much finer teeth on concave edge; proximal notch as base not defined; inner margin with heavily denticulate process; outer scales forming 2 large but unequal tooth-like protuberance. Paragnath (Fig. 53G) as small spinulose lobe. Maxillue (Fig.

53H) a single lobe armed tyerminally with 1 setiform process and 3 barbed setae. Maxilla (Fig. 53I)

2-segmented; proximal segment (syncoxa) large and armed with strongly sclerotized, ponted process on ventral surface; basis drawn out into tapering blade, armed with 3 large teeth followed by a row of 6 small teeth; seta III missing, but both setae I and II well-developed and heavily armed.

Maxilliped (Fig. 54A) 3-segmented; first segment small and unarmed; second segment large, bearing 2 large, spiniform, and heavily armed setae; terminal segment drawn out into long tapering blade with small, pointed, basal protrusion; basal part of endopod carrying 1 naked and 1 barbed seta.

Legs 1-3 (Fig. 54B-D) with 3-segmented exopod and 2-segmented endopod. Leg 4 (Fig. 54E) with 1-segmented exopod, endopod absent. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 4; enp. 0-1; I, 6 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; III, 4 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; III, 3

Leg 4: coxa 0-0; basis 1-0; exp. I, I, 2; enp. (missing)

Leg 5 (Fig. 54F) consisting of 1 seta on fifth pedigerous somite and a small exopod mesuring

37×16 μm, carrying terminally 1 large naked seta (87 μm) and 1 short spine (26 μm). Leg 6 a long seta located in genital aperture.

Male: Body (Fig. 55A) 0.92 mm long, generally as in female. Urosome (Fig. 55B) 6-segmented. Genital somite 135×156 μm. Caudal ramus 67×20 μm, armature as in female.

Antennule (Fig. 55C) segmented and armed as in female, except for addition of 2 aesthetascs on second segment and 1 aesthetasc on third segment. Antenna, mandible, maxillule and maxilla as in female. Maxilliped (Fig. 55D, E) 4-segmented; first segment longest but unarmed; second segment armed on medial margin with 2 midified setae bearing spinulose know and on outer surface a row of spinules ending with 3 long setae distally; third segment smallest and unarmed; fourth segment forming a long, curve claw carrying proximally 1 short and 1 long setae. Leg armature as in female. Leg 5 exopod 62×20 μm. 108 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

G H

Fig. 53. Polyankylis orientalis, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. paragnath; H. maxillule; I. maxilla. Scales: A, B=0.1 mm, C=0.05 mm, D-G, I=0.02 mm, H=0.01 mm. Cyclopoida: Polyankyliidae: Polyankylis 109

A C

B D

F E

Fig. 54. Polyankylis orientalis, female. A. maxilliped; B. leg 1; C. leg 2; D. leg 3; E. leg 4; F. leg 5. Scales: 0.02 mm. 110 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B C

E D

Fig. 55. Polyankylis orientalis, male. A. habitus, dorsal; B. urosome, ventral; C. antennule; D. maxilliped; E. distal part of maxilliped. Scales: A=0.1 mm, B, C=0.05 mm, D, E=0.02 mm. Cyclopoida: Polyankyliidae: Polyankylis 111

Distribution: Korea. Korea: GG.

Specimens examined: 1♀, 1♂ (type specimens) from stagnant water on the tidal flat, Jakyak I., Incheon, 29.ix.1996. Remarks: This species was found from stagnant water on the mud flat at Jayakdo Island, off Incheon. Although it is not known of its host, a polychaete is assumed to be the host, as in other species of the Polyankyliidae.

29. Polyankylis ovilaxa Kim, 2014 (Figs. 56-58)

Jag-eun-bal-top-geom-mul-byeo-ruk (작은발톱검물벼룩)

Polyankylis ovilaxa Kim, 2014, p. 111, figs. 11-13.

Female: Body (Fig. 56A) 838 μm long, excluding caudal seta, in dissected paratype. Prosome 512 μm long and strongly tapering posteriorly. Cephalothorax 333×343 μm, divided by faint dorsal suture line into cephalosome and first pedigerous somite. Cephalosome with apical, rostral prominence and slight lateral constriction indicating division between oral and maxillipedal areas. Second to fourth pedigerous somites with distinct epimera. Fourth pedigerous somite small. Uro- some (Fig. 56B) 5-segmented. Fifth pedigerous somite 62 μm wide and distinctly narrower than genital double-somite. Genital double-somite (Fig. 56C) 92×77 μm, consisting of broader anterior 70% and narrower posterior 30%, ventrally divided by transverse line into anterior and posterior halves. Three free abdominal somites narrow, unornamented, 37×40, 35×38, and 46×40 μm, respectively. Anal region large. Caudal rami rather slender, slightly divergent, and widely separated from each other by ramus width; each ramus 66 ×15 μm (ratio 4.40 : 1), with 6 naked setae; outer lateral seta locating slightly proximal to midlength of ramus; largest caudal seta (seta V) 206 μm long, more than twice as long as next longest nearby seta (seta VI); all caudal setae naked. Egg sac containing loosely aggregated eggs; each egg about 40 μm in diameter.

Rostrum (Fig. 56D) tapering posteriorly, slightly wider than long, with round posterior apex. An- tennule (Fig. 56E) slender, 202 μm long, and 6-segmented; all setae naked; armature formula: 2, 6, 6, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; 2 proximal setae on third segment obscurely positioned between second and third segments, therefore armature formula of first 3 segments may be interpreted as 2, 7, and 5 (or 2, 8, and 4). Antenna (Fig. 57A) 4-segmented, consisting of coxobasis and 3-segmented endopod; coxobasis narrow proximally and broadened distally, with 1 distal seta medially; first endopodal segment the longest segment, gradually broadening distally, with 1 seta in middle of medial margin; second endopodal segment short, armed with 1 claw and 2 setae (proximal seta very small); third endopodal segment 2.5 times as long as wide and armed with 3 claws and 2 naked outer setae; 2 medial claws strongly curved and smaller than outer 1, the latter with subdistal spinule on medial margin.

Labrum (Fig. 57B) narrow and covering only part of mandible, with large posterior lobes, and deep and narrow median incision. Mandible (Fig. 57C) with short, strongly tapering blade and 2 spines (or spiniform scales) on convex side (proximal 1 distinctly longer than distal 1); blade terminating in blunt tip, with more than 20 teeth along convex outer margin and wavy row of small, densely arranged spinules on concave medial margin. Paragnath not seen. Maxillule (Fig. 57D) 112 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

lobate, slightly longer than wide, and armed with 4 setae (2 lateral and 2 distal); 2 middle ones of these 4 setae weakly pinnate unilaterally. Maxilla (Fig. 57E) 2-segmented; proximal segment ornamented proximally with patch of setules on posterior surface; distal segment drawn out into short, acutely tapering process bearing with 4 large and 1 small teeth on convex side; medial seta (seta I) large, longer than segment, with 1 row of large spines and 1 row of small spinules on distal (outer) margin and several setules on proximal (medial) margin; anterior seta (seta II) spiniform, unilaterally spinulose along medial margin; outer proximal seta (seta III) lacking. Maxilliped (Fig. 57F) 3-segmented. First segment unarmed. Second segment inflated medially, with 2 large, spiniform setae; proximal seta much longer than segment, with 4 long setules on medial margin and 2 shorter setules on outer margin; distal seta slightly shorter than segment, almost straight, with minute spinules along outer margin and minute setules along medial margin. Terminal segment forming distally slender, elongate claw and proximally with 1 spinulose spine, 1 small seta, and 1 spiniform process.

Legs 1-3 (Figs. 57G, 58A, B) with 3-segmented exopod and 2-segmented endopod. Leg 4 (Fig. 58C) uniramous, with 2-segmented exopod. Endopods of legs 1-3 longer than exopods; distal endopodal segment of these legs much longer than wide and with small, pointed process on outer margin. Three spines on outer and distal margins of exopod of leg 4 setiform, elongate and distally barbed. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, 4; enp. 0-1; I, 1, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, 5; enp. 0-1; II, I, 4 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, 5; enp. 0-1; II, I, 3

Leg 4: coxa 0-0; basis1-0; exp. I-0; II, I, 3; enp. (missing) Leg 5 consisting of 1 dorsolateral seta on fifth pedigerous somite and 1-segmented, free exopod; exopod (Fig. 58D) small, 19×10 μm (ratio 1.90 : 1), carrying terminally 1 setiform spine and 1 long seta, both naked. Leg 6 represented by large seta on ventral surface of genital double-somite.

Male: Body (Fig. 58E) similar to that of female. Body length 702 μm in dissected paratype. Uro- some (Fig. 58F) 6-segmented. Fifth pedigerous somite narrow, less than half width of genital somite. Genital somite longer than wide, 102×88 μm, with pronounced posterolateral corners. Four abdominal somites 26 ×36, 28×33, 22×30, and 33×33 μm, respectively. All abdominal somites smooth, without ornamentation. Caudal ramus 51×13 μm (ratio 3.92 : 1). Antennule segmented and armed as in female except for addition of 2 aesthetascs on second segment and 1 aesthetasc on third segment. Antenna ornamented with spinules on outer margin of second segment (basis).

Labrum, mandible, maxillule, and maxilla as in female. Maxilliped (Fig. 58G) 4-segmented. First segment with large, acutely pointed process distally. Second segment with 1 longitudinal row of setules (3 distal setules enlarged) on outer surface, and 2 expanded, heavily barbed, transformed setae; proximal one of latters longer than distal 1, gradually narrowing distally, with spinules on proximal half of proximal margin; distal seta blunt, short, with 1 setule proximally and spinules distally. Third segment small and unarmed. Terminal segment forming large, arched claw bearing proximally 1 large and 1 small setae.

Legs 1-4 not different from those of female. Leg 5 similar to that of female, but exopod 14×6 μm

(ratio 2.33 : 1), more slender, and armed with 2 unequal, distal setae.

Distribution: Korea. Korea: GN.

Specimens examined: 17♀♀, 11♂♂ (type specimens) from the terebellid polychaete Thelepus Cyclopoida: Polyankyliidae: Polyankylis 113

B A

Fig. 56. Polyankylis ovilaxa, female. A. habitus, dorsal; B. urosome, dorsal; C. first two urosomal somites, ventral; D. rostral area, ventral; E. antennule. Scales: A=0.1 mm, B, D, E=0.05 mm, C=0.02 mm. 114 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B D

A C

Fig. 57. Polyankylis ovilaxa, female. A. antenna; B. labrum; C. mandible; D. maxillule; E. maxilla; F. maxilliped; G. leg 1. Scales: A=0.05 mm, B-G=0.02 mm. Cyclopoida: Polyankyliidae: Polyankylis 115

A C B

F E

Fig. 58. Polyankylis ovilaxa. female: A. leg 2; B. leg 3 endopod; C. leg 4; D. leg 5 exopod. male: E. habitus, dorsal; F. urosome, ventral; G. maxilliped. Scales: A-D, G=0.02 mm, E=0.1 mm, F=0.05 mm. 116 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

japonicus Marenzeller collected from a tidal flat (34°49′54″N, 128°02′08″E), Namhae I., Gyeong- sangnam-do, 22.vii.2001. Remarks: This species differs from P. orientalis in the following features: 1) the caudal ramus is

4.4 times as long as wide (vs. 3.4 times in P. orientalis); 2) leg 5 exopod 1.90 times as long as wide (vs.

2.31 times P. orientalis); 3) the absence of a claw on the first segment of the antenna (vs. present in P. orientalis); and 4) leg 4 exopod is 2-segmented (vs. 1-segmented in P. orientalis).

Genus Terebelliphilus Kim, 2000

Yu-ryeong-gaet-ji-reong-i-beol-re-sok (유령갯지렁이벌레속)

Body narrow. Fourth pedigerous somite without epimera. Antennule 6-segmented, with 2, 6, and 6 setae from first to third segment, respectively. Antenna 4-segmented, with 1 spine plus 2 setae on third segment and 3 spines plus 3 setae on terminal segment. Legs 1-3 with 3-segmented exopod and 2-segmented endopod. Leg 3 without seta on first endopodal segment. Leg 4 with 3-segmented exopod; endopod lacking. Inner coxal seta absent in legs 1-4. Containing a single species.

30. Terebelliphilus simplex Kim, 2001 (Figs. 59-61)

Yu-ryeong-gaet-ji-reong-i-beol-re (유령갯지렁이벌레)

Terebelliphilus simplex Kim, 2001a, p. 2, figs. 1-3.

Female: Body (Fig. 59A) small, narrow, 743 μm long, with thin exoskeleton. Prosome consisting of cephalothorax and 3 pedigerous somites. Cephalothorax narrow, longer than wide, 288×238 μm, with round anterior margin, nearly parallel lateral margins, and weak, round lateral expansion at posterior parts of lateral margins. Second and third pedigerous somites with round epimera, but fourth pedigerous somite without epimera. Urosome (Fig. 59B) 5-segmented but suture lines faint and incomplete. Fifth pedigerous somite 44×110 μm. Genital double- and first 2 abdominal somites each armed with horizontal row of about 6 denticles near posteroventral border. Genital double-somite 88×99 μm, narrower than fifth pedigerous somite, and slightly expanded anteriorly and weakly tapering. Genital aperture characteristically located on ventral surface. Three abdominal somites 41×58, 29×54, and 24×45 μm, respectively. Caudal rami widely separated from each other, originated posteroventrally from anal somite, each 25×14 μm (1.79 : 1), with 6 setae, terminal one of them distinctly larger. All caudal setae naked.

Rostrum absent. Antennule (Fig. 59C) 6-segmented, 150 μm long, with armature formula: of 2, 6, 6, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Third segment subdivided proximally by a faint suture line. Antenna (Fig. 59D) 4-segmented, with armature formula of 1, 1, 2+claw, and 3+3 claws. Second segment about 4 times as long as wide, with many denticles on inner margin. One of setae on third segment spinulated along inner margin. Inner one of 3 terminal claws thicker than other two. Cyclopoida: Polyankyliidae: Terebelliphilus 117

Labrum (Fig. 59E) bilobed and deeply incised posteromedially. Mandible (Fig. 59F) with small process proximal to denticulated convex margin. Inner margin short, with row of spinules. Termi- nal lash very short, strongly tapering. Paragnath not observed. Maxillule (Fig. 59G) armed with 1 lateral setiform process and 3 setae, terminal one of them spiniform, barbed on both margins.

Maxilla (Fig. 59F) 2-segmented. First segment thick, with several oblique rows of spinules on ventral and posterior surfaces. Second segment with 2 setae. Inner (dorsal) seta extremely long, basally expanded, with setiferous outer margin. Anterior seta naked. Outer basal seta not observed.

Convex margin with 4 large teeth proximal to terminal lash. Terminal lash very short. Maxilliped (Fig. 60A) 3-segmented. First segment unarmed and tapering. Second segment expanded medially, with 2 small setae and strongly convex inner margin. Terminal segment strongly tapering, with 2 unequal setae near inner margin and row of several spinules on both sides near tip.

Legs 1-3 (Fig. 60B-D) with 3-segmented exopod and 2-segmented endopod. Leg 4 (Fig. 60E) with 3-segmented exopod, endopod lacking. Second endopodal segment of leg 1 with 2 cusps on outer margin. Second endopodal segment in legs 2 and 3 with 1 cusp on outer margin. Legs 1-4 without inner coxal seta. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-0; basis 1-0; exp. I-0; I-1; II, I, 4; enp. 0-1; I, 6 Leg 2: coxa 0-0; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; III, 4 Leg 3: coxa 0-0; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-0; III, 3

Leg 4: coxa 0-0; basis 1-0; exp. I-0; I-1; I. I, 4; enp (missing)

Leg 5 (Fig. 60F) 1-segmented, with no trace of protopod or seta, distally broadened, 23 ×8.7 μm

(2.64 : 1), terminally with a pointed process, 1 spine (30 μm) and 1 naked seta (36 μm). Leg 6 represented in genital aperture by 1 small denticle and 1 relatively large seta.

Male: Body (Fig. 60G) resembling that of female, 639 μm long. Urosome (Fig. 61A) 6-segmented. Genital and 4 abdominal somites 56 ×89, 37×65, 32×54, 24×45, and 21×39 μm, respectively. Cau- dal ramus 23×14 μm (1.64 : 1).

Antennule added by 2 aesthetascs, each proximal and distal, on second segment. Antenna (Fig. 61B) with 1 additional seta on third segment.

Labrum (Fig. 61C) armed with spinules on posterior margin of lobes. Maxilliped (Fig. 61D) 4-segmented. First segment unarmed, tapering. Second segment with 2 thick, short, spinulated setae and several rows of spinules. Third segment short and unarmed. Fourth segment forming long, curved claw, basally with 2 setae, proximal one of latters greatly enlarged, longer than claw. Other mouthparts as in female.

Legs 1-4 with same armature formula as those of female. Leg 5 24×7 μm (3.43 : 1). Terminal spine 35 μm. Leg 6 represented by 1 seta and 2 spinules on genital operculum.

Distribution: Korea. Korea: CN.

Specimens examined: 25♀♀, 9♂♂ (type specimens) collected from washings of the polychaete

Terebella ehrenbergi Grube in intertidal mud flat, at Sapsi-do Island (36°20′N, 126°21′E), Choongc- heongnam-do, 26.x.1999.

Remarks: This species was originally placed in the family Sabelliphilidae, but Kim (2014) removed it to the family Polyankyliidae. The major diagnostic character of Terebelliphilus simplex is the absence of epimera on the fourth pedigerous somite, which is observable without dissection. 118 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

G D E

H F

Fig. 59. Terebelliphilus simplex, female. A. habitus, dorsal; B. urosome, ventral; C. antennule; D. antenna; E. labrum; F. mandible; G. maxillule; H. maxilla. Scales: A=0.1 mm, B=0.05 mm, C-H=0.02 mm. Cyclopoida: Polyankyliidae: Terebelliphilus 119

F G

Fig. 60. Terebelliphilus simplex. female: A. maxilliped; B. leg 1; C. leg 2; D. leg 3 endopod; E. leg 4; F. leg 5. male: G. habitus, dorsal. Scales: A-F=0.02 mm, G=0.1 mm. 120 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B D A

Fig. 61. Terebelliphilus simplex, male. A. urosome, ventral; B. distal part of antenna; C. labrum; D. maxilliped. Scales: A=0.05 mm, B-D=0.01 mm.

Genus Sewelloya Kim, 2014

Mu-jok-bal-top-geom-mul-byeo-ruk-sok (무족발톱검물벼룩속)

Body cyclopiform. Urosome 5-segmented in female and 6-segmented in male. Antennule 6-segmented, with 2, 6, and 7 setae respectively on first to third segments in female, but in male with 1 additional aesthetasc on second segment. Antenna 4-segmented; third segment with 1 claw and 2 setae, and terminal segment with 3 claws and 4 setae. Labrum with clearly defined posterior lobes. Mandible short, strongly tapering, with 1 claw-like element on convex side and curved row of spinules on inner margin. Maxillule with 4 setae. Maxilla 2-segmented; distal segment with 2 setae and short, curved distal process bearing several dentiform spines on convex margin. Female maxilliped 3-segmented; first segment unarmed; middle segment with 2 setae; distal segment small and tapering with 2 small setae and spinulose apex. Male maxilliped 4-segmented; first and third segment without seta; second segment with 2 stout spiniform setae; terminal segment as large claw bearing 2 setae proximally. Legs 1-3 with 3-segmented exopod and 2-segmented endopod. Leg 4 uniramous, with 1-segmented exopod. Leg 5 represented by 3 setae; exopod absent. Leg 6 represented by 1 seta (Kim, 2014). Cyclopoida: Polyankyliidae: Sewelloya 121

31. Sewelloya plana Kim, 2014 (Figs. 62-64)

Mu-jok-bal-top-geom-mul-byeo-ruk (무족발톱검물벼룩)

Sewelloya plana Kim, 2014, p. 116, figs. 14-16.

Female: Body (Fig. 62A) dorsoventrally flat, with laterally inflated prosome and cylindrical urosome. Body length 1.19 mm in dissected paratype (1.12 mm in holotype). Prosome 775 μm long and consisting of cephalothorax and second to fourth pedigerous somites. Greatest width of prosome 555 μm. Cephalothorax divided dorsally by faint suture line into cephalosome and first pedigerous somite, with broad apical prominence and rounded posterolateral corners. Fourth pedigerous somite small and much narrower than preceding somite. Urosome (Fig. 62B) 5-segmented.

Fifth pedigerous somite short but nearly as borad as genital double-somite, with 3 unequal setae

(representing leg 5) on both sides. Genital double-somite (Fig. 62C, D) consisting of expanded anterior part (67×165 μm) and narrower distal part (50×103 μm); these anterior and posterior parts divided dorsally by suture but not divided ventrally; genital aperture locating laterally. Genital double-somite and first and second free abdominal somites with membranous flange along posterodorsal margin (Fig. 14B) and spinules along posteroventral margin. Three free abdominal somites 58×97, 58×85, and 58×67 μm, respectively. Caudal rami slightly divergent; each ramus

92×26 μm (ratio 3.54 : 1), with 6 naked setae; seta I absent; seta II locating dorsally at halfway of ramus; seta V the largest, 396 μm long, more than 3 times as long as next longest seta. Egg sac unknown.

Rostral area (Fig. 63A) broad but rostrum absent. Antennule (Fig. 62E) 300 μm long and 6-segmented; first to third segments gradually narrowing distally; armature formula 2, 6, 7 (1 being minute), 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all setae slender and some of them weakly pinnate. Antenna (Fig. 62F) 4-segmented; coxobasis gradually broadened distally, with 1 small medio-distal seta and minute spinules in distal region; first endopodal segment the longest, gradually broadened distally, with 1 small seta on medial margin slightly distal to midlength of the margin and 1 transverse row of minute spinules near outer distal corner; second endopodal segment small, with short claw and 2 setae, proximal one of latters minute; third endopodal segment twice as long as wide, with 3 broad claws and 2 setae distally and 2 minute setae subdistally on outer side; medial one of 2 distal setae truncate and tipped by 1 spinule.

Labrum (Fig. 63B) small, with prominent distal lobes articulated at base. Mandible (Fig. 63C) short, strongly tapering, broad at base; convex outer side with 1 claw-like spine and several minute spinules; convex side of blade with row of fine teeth; concave side of blade with curved row of fine spinules; distal end of blade rather blunt, not flexible. Paragnath not seen. Maxillule (Fig.

63D) lobate, with 4 naked setae on oblique distal margin. Maxilla (Fig. 63E) 2-segmented; proximal segment with long membranous flange on outer (ventral) side, 1 tuft of setules on proximal region and 1 curved row of fine spinules on distal region; distal segment strongly recurved in distal region bearing 5 or 6 (usually 6) strong teeth along convex outer side and 1 row of minute spinules on anterior side, with 2 setae (setae I and II); seta I (medial seta) large, nearly as long as segment and spinulose along distal margin; seta II (anterior seta) stiff and spinulose along medial margin.

Maxilliped (Fig. 63F) 3-segmented; first segment (syncoxa) unarmed but with 3 transverse rows of fine spinules on medial side; middle segment (basis) with 2 spiniform setae: small, spinulose medial 1 and larger, unilaterally spinulose 1 terminating in thin, setiform tip; distal segment (endopod) small, tapering, with 2 minute setae near middle and row of fine spinules distally. 122 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

Fig. 62. Sewelloya plana, female. A. habitus, dorsal; B. urosome, dorsal; C. anterior part of urosome, right; D. anterior part of urosome, ventral; E. antennule; F. antenna. Scales: A=0.2 mm, B-E=0.05 mm, F=0.02 mm. Cyclopoida: Polyankyliidae: Sewelloya 123

G F

Fig. 63. Sewelloya plana, female. A. cephalic region, ventral; B. labrum; C. mandible; D. maxillule; E. maxilla; F. maxilliped; G. leg 1. Scales: A=0.05 mm, B-G=0.02 mm. 124 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

F D

Fig. 64. Sewelloya plana. female: A. leg 2; B. leg 3; C. leg 4. male: D. habitus, dorsal; E. first two urosomal somites, ventral; F. maxilliped. Scales: A, B, F=0.02 mm, C, E=0.05 mm, D=0.2 mm. Cyclopoida: Polyankyliidae: Sewelloya 125

Legs 1-3 (Figs. 63G, 64A, B) biramous, with 3-segmented exopod and 2-segmented endopod. Leg

4 (Fig. 64C) uniramous, consisting of unarmed coxa, 1 seta-bearing basis, and 1-segmented exopod; intercoxal sclerite with pair of pointed distal processes; 7 setae on exopod stiff and spiniform. Coxa of legs 1-3 with small, pinnate inner seta and spinules on outer distal corner. Basis of leg 1 with 1 medio-distal, membranous flange near base of endopod and 2 rows of minute spinules on ventral surface. Basis of legs 2 and 3 with 2 ventral rows of minute spinules, each row near bases of exopod and endopod. Endopods of legs 1-3 with transparent knob on outer distal corner of proximal segment and outer margin of distal segment. First exopodal segment of legs 1-3 with fine spinules on outer margin. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 4; enp. 0-1; I, 2, 4 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; II, I, 4 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; II, I, 3

Leg 4: coxa 0-0; basis 1-0; exp. 2, 2, 3; enp. (missing) Leg 5 represented by 3 unequal setae; lengths of these setae 112, 38, and 22 μm, repectively, from dorsal to medial; minute spinules present near base of setae. Leg 6 represented by 1 small, simple seta on genital operculum.

Male: Body (Fig. 64D) narrower than that of female. Body length 1.06 mm. Greatest width 450 μm measuring across cephalothorax. Urosome 6-segmented. Fifth pedigerous somite slightly narrower than genital somite. Genital somite (Fig. 64E) 133×175 μm, with convex lateral margins, distinct genital operculum tipped by 1 seta, and deep posteroventral incision. Four abdominal somites gradually narrowing from proximal to distal. Caudal ramus 60 ×23 μm (ratio 2.61 : 1), shorter than that of female. Rostrum absent, as in female. Antennule with 1 additional aesthetasc on second segment at opposite side of dot in Fig. 14E. Antenna as in female.

Labrum, mandible, maxillule, and maxilla as in female. Maxilliped (Fig. 64F) 4-segmented; first segment with 2 tapering processes at medio-distal corner; second segment with convex outer margin and 1 medial, longitudinal row of fine spinules and armed with 2 massive, transformed spines on medial margin; latter 2 spines (proximal 1 larger than distal 1) distally expanded and spinulose, and tipped by 1 setule; third segment small and unarmed; terminal segment forming long, strongly curved claw bearing 1 large and 1 minute setae proximally. Legs 1-5 not different from those of female. Leg 6 represented by 1 seta on genital operculum.

Distribution: Korea. Korea: CN.

Specimens examined: 2♀♀, 1♂ (type specimens) from the terebellid polychaete Loimia medusa

(Savigny) collected from the intertidal zone of Choonjangdae (36°09′40″N, 126°30′57″E), Choongc- heongnam-do, 03.i.2014; 1♀, 1♂ (both dissected) from the same host and locality as type material, 1.i.2002. Remarks: The genus Sewelloya is a monotypic genus. It is distinguished from the type genus,

Polyankylis, mainly in the female maxilliped and leg 5 (Kim, 2014). 126 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Family Pseudanthessiidae Humes and Stock, 1972

Min-da-ri-geom-mul-byeo-ruk-gwa (민다리검물벼룩과)

Body cyclopiform. Urosome 5-segmented in female and 6-segmented in male. Antennule 7-segmented. Antenna 3- or 4-segmented; third segment with 3 setae or 2 setae plus 1 claw; terminal segment with 7 elements. Labrum with deep incision and elongate posterior lobes. Mandible strongly tapering, with short distal lash. Maxilliped 3-segmented in female and 4-segmented in male. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 1-segmented endopod (endopod 2-segmented in Sipadania). Free exopod of leg 5 present in Sipadania Humes and Lane, 1993 and Tubiporicola Kim, 2009, but absent in other genera. Major hosts are echinoderms and polychaetes.

Genus Pseudanthessius Claus, 1889

Min-da-ri-geom-mul-byeo-ruk-sok (민다리검물벼룩속)

Antennule 7-segmented; first segment without enlarged setae (all setae shorter than half length of

Antennule). Antenna 4-segmented. Third exopodal segment of leg 3 with 3 spines and 2 setae (formula I, II, 2). Leg 5 without free exopod.

32. Pseudanthessius excertus Kim, 2014 (Figs. 65, 66)

Dol-gi-min-da-ri-geom-mul-byeo-ruk (돌기민다리검물벼룩)

Pseudanthessius excertus Kim, 2014, p. 108, figs. 9, 10.

Female: Body (Fig. 65A) rather small. Body length 945 μm in dissected and figured paratype (in other 2 measured specimens 868 and 968 μm). Prosome 584 μm long. Cephalothorax 403×372 μm, nearly globular, as dorsoventrally deep as wide, with weak dorsal suture delimiting cephalosome and first pedigerous somite. Second pedigerous somite broad, with prominent, tapering lateral projection on both side. Third and fourth pedigerous somite much narrower and shorter than second pedigerous somite, 240 and 154 μm wide, respectively, with round lateral margins. Urosome (Fig. 65B) 5-segmented. Fifth pedigerous somite 117 μm wide. Genital double-somite 140×132 μm, slightly longer than wide, with broad lateral expansion, widest at 0.4 region of somite length and gradually narrowing posteriorly from widest region; genital aperture large and locating dorsally. Three free abdominal somites 37×72, 26×60, and 51×60 μm, respectively. Anal somite twice as long as penultimate abdominal somite, with large anal region. Caudal ramus (Fig. 65C)

71×26 μm (ratio 2.73 : 1), slightly narrowing distally, with 6 setae; outer lateral seta (seta II) locating at 0.6 region of ramus length.

Rostrum (Fig. 65D) stout, as long as wide; proximal half with parallel lateral margins and distal half strongly tapering, with rounded posterior apex. Antennule (Fig. 65E) 183 μm long and 7-seg- Cyclopoida: Pseudanthessiidae: Pseudanthessius 127

B A

C G D

Fig. 65. Pseudanthessius excertus, female. A. habitus, dorsal; B. urosome, dorsal; C. right caudal ramus, dorsal; D. rostrum; E. antennule; F. antenna; G. labrum; H. mandible; I. maxillule. Scales: A=0.1 mm, B=0.05 mm, C-I=0.02 mm. 128 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

C A B

F D

Fig. 66. Pseudanthessius excertus, female. A. maxilla; B. maxilliped; C. leg 1; D. leg 3; E. leg 4; F. right side of first two urosomal somites, dorsal. Scales: 0.02 mm. Cyclopoida: Pseudanthessiidae: Pseudanthessius 129 mented; armature formula 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all setae naked. Antenna (Fig. 65F) 4-segmented; last segment 33×20 μm (ratio 1.65 : 1); armature formula 1, 1, 2+claw, and 3+4 claws; claws slender; outermost one of 4 terminal claws distinctly longer than other 3.

Labrum (Fig. 65G) with elongate posterior lobes bearing hemi-circular notch on medial margin and membranous flange along posterior margin and distal part of medial margin. Mandible (Fig. 65H) curved in a right angle, with 1 broad, transparent scale on convex outer corner at base of blade; medial margin of blade with about 10 spines near middle of margin and fine spinules distally; outer margin of blade smooth, only with delicate striations; distal part of blade blunt, with small point, without lash. Paragnath as small digitiform process (indicated by dotted line in Fig.

65G). Maxillule (Fig. 65I) lobate, with 1 lateral and 3 distal setae; 2 larger ons of latters unilaterally sinulose in distal half. Maxilla (Fig. 66A) 2-segmented; proximal segment expanded but unarmed; distal segment armed with 3 setae: medial margin seta (seta I) large, almost extending to distal end of distal lash of segment; anterior seta (seta II) stout, spiniform, and spinulose along medial margin; proximal seta (seta III) small and blunt at tip; distal lash of second segment relatively short, armed with 2 rows of thick teeth on convex side and with setules along distal region. Maxilliped (Fig. 66B) 3-segmented; first segment unarmed; middle segment slightly longer than wide, with 1 massive spine bearing 4 teeth, 1 small seta, and 1 longitudinal row of minute spinules; distal segment terminating in spiniform process bearing 4 or 5 teeth, with 1 stout spine bearing 5 teeth.

Legs 1 and 2 (Fig. 66C, D) and leg 3 with 3-segmented rami. Leg 4 (Fig. 66E) with 3-segmented exopod and 1-segmented endopod. Third exopodal segment of leg 3 characteristically armed with 3 spines and 5 setae. Leg 4 with minute inner seta on coxa; endopod 46×15 μm, about 3 times as long as wide, with several setules on outer margin but lacking notch or process on outer margin. Armature formula of legs 1-4 as follows: Leg 1: coxa 1-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 2, 3 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-2; I, II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. II

Leg 5 (Fig. 66F) represented by lobe bearing 1 dorsal seta proximally and 1 spine and 1 seta distally. Leg 6 represented 2 small setae (distal one of them proximally thick and distally thin) and 1 detiform process on genital operculum (Fig. 66F). Male: Unknown.

Distribution: Korea. Korea: GW.

Specimens examined: 6♀♀ (type specimens) from polychaetes (mainly Hydroides ezoensis

Okuda) epizoic on shells of the oyster Crassostrea gigas (Thunberg) attached to the wharf in Gang- neung Port (37°46′15″N, 128°57′08″E), depth about 50 cm, 21.xi.2012. Remarks: Although the genus Pseudanthessius is very speciose, this species can easily be differentiated from its congeners by the possession of a pair of prominent lateral processes on the second pedigerous somite, and 3 spines and 5 setae (armature formula II, I, 5, not III, I, 5) on the third exopodal segment of leg 3. 130 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Family Rhynchomolgidae Humes and Stock, 1972

San-ho-i-gwa (산호이과)

Body cyclopiform or swollen. Urosome 5-segmented in female and 6-segmented in male. Anten- nule 7-segmented. Antenna generally 4-segmented but sometimes 3-segmented. Mandible broad, with inner margin perpendicular to distal lash. Maxillule typically with 1 subdistal and 3 distal setae. Maxilla 2-segmented; basal segment unarmed; distal segment with long lash. Maxilliped 3-segmented in female and 4-segmented in male. Legs 1-3 with 3-segmented rami. Leg 1 without inner distal seta on basis. Leg 4 with 3-segmented exopod and 1- or 2-segmented endopod; sometimes endopod absent.

Key to the genera of family Rhynchomolgidae dealt with in the present report

1. Antenna 3-segmented ··············································································································· Lambanetes - Antenna 4-segmented ································································································································ 2 2. Terminal segment of antenna with 2 claws ···························································································· 3 - Terminal segment of antenna with 1 claw ······························································································ 4 3. Body swollen; leg 3 with 2-segmented endopod ···································································· Verutipes - Body not swollen; leg 3 with 3-segmented endopod ·························································· Critomolgus 4. Second endopodal segment of leg 4 with 2 spines and 3 setae ······································ Lutumidomus - Second endopodal segment of leg 4 with 2 spines ················································································ 5 5. Convex side of mandible greatly produced; third exopodal segment of leg 4 with 4 spines and 5 setae ··············································································································································· Zamolgus - Convex side of mandible not strongly produced; third endopodal segment of leg 4 with 3 spines and 5 setae ································································································································ Paramolgus

Genus Critomolgus Humes and Stock, 1983

Mal-mi-jal-i-sok (말미잘이속)

Remarks. Critomolgus anthoplureus, C. vicinus, and C. malmizalus those mentioned below are sibling species and a careful comparion is needed to distinguish them. The genus Critomolgus is closely related to the genus Dorodicola Leydig, 1954. Both genera are distinguished from each other only by a single morphological character: the third exopodal segment of leg 4 has 4 spines and 5 setae

(armature formula III, I, 5) in the former genus, but 3 spines and 5 setae (II, I, 5) in the latter.

Key to the species of genus Critomolgus

1. Genital double-somite of female wider than long; caudal ramus less than 1.5 times as long as wide; distal setae on leg 5 exopod longer than exopod ·························································· C. nudus - Genital double-somite of female longer than wide; caudal ramus more than 1.5 times as long as wide; distal setae on leg 5 exopod shorter than exopod ······································································ 2 2. Leg 5 exopod without medial swelling ············································································ C. malmizalus - Leg 5 exopod with medial swelling ········································································································ 3 Cyclopoida: Rhynchomolgidae: Critomolgus 131

3. Genital double-somite in female widest at halfway; body length of female usually over 1.9 mm; second segment male maxilliped with crest-like, medio-distal expandsion ············ C. anthoplureus - Genital double-somite in female widest across anterior part; body length of female less than 1.9 mm; second segment of male maxilliped without crest-like medio-distal expansion ······· C. vicinus

33. Critomolgus anthopleurus Kim, 1996 (Figs. 67-69)

Nok-saek-mal-mi-jal-sok-sal-i (녹색말미잘속살이)

Critomolgus anthopleurus Kim, 1996, p. 367, figs. 4-6.

Female: Body (Fig. 67A) moderately broad. Length 2.29 mm (1.88-2.70 mm) and greatest width

1.01 mm (0.85-1.13 mm), based on 10 specimens. First pedigerous somite separated from cephalosome by weak dorsal suture line. Epimeral area of posomal somites rounded posteriorly. Urosome (Fig. 67B) 5-segmented. Fifth pedigerous somite 325 μm wide. Genital double-somite 365 ×313

μm, ratio 1.17 : 1; convex in middle. Genital aperture located dorsolaterally at anterior 0.35 of somite length. Second free abdominal somite shortest among 3 free abdominal somites. Anal somite with posteroventral margin fringed with row of minute spinules. Caudal ramus 125×75 μm (ratio

1.67 : 1), with 6 setae; outer lateral and dorsal setae naked; other 4 distal setae plumose. Egg sac 845×420 μm and kidney-shaped.

Rostrum with hemicircularly rounded posterior margin. Antennule (Fig. 67C) 7-segmented, 533 μm long, decreasing in width distally, with armature formula of 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Several larger setae on first and second segments with minute setules.

Antenna (Fig. 67D) 4-segmented, with armature formula 1, 1, 3, and 5+2 claws. Fourth segment about 4 times as long as wide; all setae naked; 2 terminal claws almost equal in size.

Labrum (Fig. 67E) with 2 posteroventral lobes. Mandible (Fig. 67F) with convex side of base bearing row of small spines and followed by serrate fringe; concave side of base neyond indentation bearing setules of 2 different sizes; lash moderately long and barbed. Paragnath a small hairy lobe.

Maxillule (Fig. 67G) with 3 terminal seta and 1 subterminal setiform process. Maxilla (Fig. 68A) 2-segmented, with unornamented proximal segment; distal segment with naked posterior seta, inner barbed spine, and small outer proximal setule; distal lash long and spinose. Maxilliped (Fig. 68B) 3-segmented; first segment unarmed; second segment with 1 large distal and 1 small proximal setae; third segment with 2 naked setae of unequal sizes and terminating in blunt process.

Legs 1 and 2 (Fig. 68C, D) and leg 3 with 3-segmented rami. Outer seta on basis of all of these legs naked. Inner seta on coxa of leg 4 naked. Two terminal spines of leg 4 56 μm (outer) and 119

μm (inner). Leg 4 (Fig. 68F) with 3-segmented exopod and 2-segmented endopod. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. 1-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 2, 3 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; II

Leg 5 consisting of 1 small dorsolateral setae of fifth pedigerous somite and free exopod; exopod (Fig. 68G) 209 μm long, bearing distinct proximal inner expansion, and ornamented with small spinules along outer side and armed with 2 terminal setae of 67 μm (outer) and 119 μm (inner); 132 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

C A B

Fig. 67. Critomolgus anthopleurus, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. maxillule. Scales: A=0.2 mm, B-D=0.1 mm, E=0.05 mm, F, G=0.02 mm. Cyclopoida: Rhynchomolgidae: Critomolgus 133

C B A

E G

Fig. 68. Critomolgus anthopleurus, female. A. maxilla; B. maxilliped; C. leg 1; D. leg 2; E. terminal endopodal segment of leg 3; F. leg 4; G. leg 5 exopod. Scales: A, B, G=0.05 mm, C-F=0.1 mm.

greatest width measured across proximal expansion 44 μm; width in middle 35 μm. Leg 6 represented by 2 mall setae on genital operculum.

Male: Body (Fig. 69A) similar to that of female. Length 1.58 mm (1.48-1.58 mm) and breatest width 0.60 mm (0.57-0.62 mm), based on 7 specimens. Urosome (Fig. 69B) 6-segmented. Fifth 134 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B A

Fig. 69. Critomolgus anthopleurus, female. A. habitus, dorsal; B. urosome, dorsal; C. maxilliped; D. terminal endopodal segment of leg 2; E. leg 5. Scales: A=0.2 mm, B=0.1 mm, C, E=0.05 mm, D=0.02 mm.

pedigerous somite 153 μm wide. Genital somite 249-212 μm, longer than wide. Four abdominal somites 53×98, 5×93, 40×86, and 58×100 μm, respectively, Caudal ramus 74×42 μm, ratio 1.76 : 1.

Rostrum as in female. Antennule as in female, but with 3 additional aesthetascs (2 on second and 1 on fourth segments). Antenna as in female, but with more distinct spinulation on inner side of second segment.

Labrum, mandible, paragnath, maxillule, and maxilla as in female. Maxilliped (Fig. 69C) 4-segmented, including terminal claw; first segment unarmed; second segment with longitudinal row of spinules, 2 equal setae near midlength and serrate crest distally; third segment small and unarmed; terminal claw elongate, with small tubercle near proximal third on concave side and proximmaly 2 extremely unequal setae, the longer seta finely spinulated along both side of distal half.

Legs 1-4 segmented armed as in female, except for third endopodal segment (Fig. 69D) of leg 1 armed with 2 spines and 4 setae. Third endopodal segment of leg 2 with foliaceous lamella be- Cyclopoida: Rhynchomolgidae: Critomolgus 135 tween 2 terminal spines.

Leg 5 (Fig. 69E) with exopod of 108×25 μm (ratio 4.32 : 1), not expanded proximally, and armed with 1 spine (45 μm and serrate) and 1 setae (61 μm and naked). Leg 6 represented by 2 extremely unequal setae on genital operculum.

Distribution: Korea. Korea: GW. Specimens examined: Specimens were all collected from the gastrovascular cavity of the actiniarian Anthopleura midori Uchida and Muramatsu. 23♀♀, 9♂♂ (type specimens), intertidal, Anin, Gangneung, Gangwon-do, 21.vi.1992. Remarks: This species and C. vicinus are co-associated with the sea anemone Anthopleura midori, but the former species lives in the gastrovacular cavity of the host and the latter species on the body surface, mainly on the tentacles. Unlike C. vicinus, C. anthopleurus has a soft exoskeleton.

34. Critomolgus malmizalus Kim, 1996 (Figs. 70-72)

Gal-saek-mal-mi-jal-sok-sal-i (갈색말미잘속살이)

Critomolgus malmizalus Kim, 1996, p. 363, figs. 1-3.

Female: Body (Fig. 70A) moderately broad. Length 1.91 mm (1.82-2.03 mm) and greatest width

0.85 mm (0.80-0.88 mm), based on 10 specimens. First pedigerous somite separated from cephalosome by indistinct dorsal suture line. Urosome (Fig. 70B) 5-segmented. Fifth pedigerous somite 254 μm wide. Genital double-somite elongate, 208 ×204 μm, with 2 weakly expanded region at 0.17 and 0.55 of somite length; anterior half slightly broader than posterior half; genital apertures located dorsolaterally at 0.44 region of somite length. Three free abdominal somites 96×154, 69×146, and 96×154 μm, respectively. Anal somite with minute spinules on posteroventral margin. Cau- dal ramus 117×68 μm (ratio 1.72 : 1), with 6 setae. Outer lateral and dorsal setae naked. Other 4 setae plumose. Egg sac nearly ovoid, 500×370 μm.

Rostrum with hemicirculary rounded posterior margin. Antennule (Fig. 70C) 524 μm long; arm- ture formua 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Antenna (Fig. 70D) segmented and armed as in preceding species; second segment with minute spinules on medial surface. Terminal segment about 3.5 times as long as wide; two terminal claws almost equal in size.

Labrum (Fig. 70E), mandible (Fig. 70F), maxillule (Fig. 70G), and maxilla (Fig. 71A) not different from those preceding species. Maxilliped (Fig. 71B) 3-segmented; first segment unarmed; second segment with 1 large distal and 1 small proximal setae; third segment with 2 naked setae of unequal sizes and terminating in blunt process.

Legs 1 and 2 (Fig. 71C, D), leg 3, and leg 4 (Fig. 71E) segmented and armed as in preceding species. Leg 1 coxa with weak outer distal prominence. Outer seta on basis of legs 1-4 naked. Leg 4 with naked inner coxal seta; 2 terminal spines 58-65 μm (outer) and 105-109 μm (inner).

Leg 5 exopod (Fig. 71G) 188-216×33-34 μm, length: width ratio about 6.0 : 1, and ornamented with small spines along outer side; 2 terminal setae 79 and 125 μm, respectively. Leg 6 represented by small setae on genital operculum.

Male: Body (Fig. 72A) 1.44 mm long, and greatest width 0.64 mm. Urosome (Fig. 72B) 6-seg- 136 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B C

G F

Fig. 70. Critomolgus malmizalus, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. maxillule. Scales: A=0.2 mm, B-D=0.1 mm, E, F=0.05 mm, G=0.02 mm. Cyclopoida: Rhynchomolgidae: Critomolgus 137

A B

E G F

Fig. 71. Critomolgus malmizalus, female. A. maxilla; B. maxilliped; C. leg 1; D. leg 2; E. third endopodal segment of leg 3; F. leg 4; G. leg 5 exopod. Scales: A, B=0.02 mm, C-F=0.1 mm, G=0.05 mm. 138 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A F B D

Fig. 72. Critomolgus malmizalus, male. A. habitus, dorsal; B. urosome, ventral; C. maxilliped; D. third segment of maxilliped; E. leg 1; F. third endopodal segment of leg 2; G. leg 5. Scales: A=0.2 mm, B=0.1 mm, C-F=0.05 mm, G=0.02 mm. Cyclopoida: Rhynchomolgidae: Critomolgus 139 mented. Genital somite 250×243 μm, nearly as long as wide. Anal somite distinctly shorter than other 2 preceding somites. Caudal ramus 75×43 μm (ratio 1.74 : 1), shorter than that of female.

Antennule with 3 additional aestetascs (2 on second and 1 on fourth segments). Antennule with more distinct spinulation on second segment than in female.

Labrum, paragnath, maxillule, and maxilla as in female. Maxilliped (Fig. 72C, D) second segment with longitudinal row of spinules, 2 subequal setae near weak prominence at midlength and serrate crest distally.

Leg 1 (Fig. 72E) with 2 spines and 4 setae on third endopodal segment. Leg 2 with foliaceous distal lamella on third endopodal segment (Fig. 72F). Legs 3 and 4 as in female.

Leg 5 (Fig. 72G) exopod 81×24 μm (ratio 3.38 : 1), without proximal swelling; terminal spine serrate and 40 μm long, and terminal seta naked and 56 μm long. Leg 6 represented by 2 unequal naked setae on genital operculum.

Distribution: Korea. Korea: GW, GB, JN, JJ. Specimens examined: All specimens were collected from the external surface of the actiniarian

Anthopleura japonica Verrill. 13♀♀, 8♂♂ (type specimens), intertidal and shallow sublittoral, Anin, Gangneung, 24.v.1996; 10♀♀, 5♂♂, Yongmeori, Jeju I., 28.ix.1999; numerous specimens, Naegaldo I., Sinan-Gun, Jeollanam-do, 28.vi.2000; Nang-do I., Jeollanam-do, 10.x.2002; 3♀♀, 2♂♂, Dapo-ri, Geoje I., 24.iv.2008; 2 ♀♀, 3♂♂, Gyeongjeong-ri, Yeongdeok-gun, Gyeongsangbuk-do, 20.vii.2012. Remarks: Critomolgus malmizalus and its two sibling species can be distinguished by the differences listed in the Table 1.

35. Critomolgus vicinus Kim, 1996 (Figs. 73-75)

Nok-saek-mal-mi-jal-i (녹색말미잘이)

Critomolgus vicinus Kim, 1996, p. 370, figs. 7-9.

Female: Body (Fig. 73A) length 1.73 mm (1.54-1.89 mm) and breatest width 0.80 mm (0.68-0.91 mm), based on 10 specimens. Cephalothorax with weak dorsal suture line delimiting cephalosome and first pedigerous somite. Urosome (Fig. 73B) 5-segmented. Fifth pedigerous somite 229 μm wide. Genital double-somite 27×224 μm (ratio 1.21 : 1); greatest width measuring across distinctly eapanded anterior 0.25 region of somite length; remaining part of somite distal to expanded region gradually narrowed, with posterior third bearing parallel lateral margins; genital aperture located dorsolaterally at anterior 0.4 of length of somite. First free abdominal somite shorter than other free abdominal somites. Anal somite with row of minute spinules on posteroventral margin. Cau- dal ramus (Fig. 73C) 108×57 μm, length: width ratio 1.89 : 1. Egg sac 680×382 μm.

Rostrum as in 2 preceding species. Antennule (Fig. 73D) 525 μm long, with armature formula as in preceding species. Antenna (Fig. 73E) with terminal segment about 4 times as long as wide; two terminal claws almost equal in size.

Labrum (Fig. 73F), mandible (Fig. 73G), maxillule (Fig. 74A), maxilla (Fig. 74B), and maxilliped (Fig 74C) not different from those of preceding species. 140 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

C F

Fig. 73. Critomolgus vicinus, female. A. habitus, dorsal; B. urosome, dorsal; C. caudal ramus; D. antennule; E. antenna; F. labrum; G. mandible. Scales: A=0.2 mm, B, D, E=0.1 mm, C, F=0.05 mm, G=0.02 mm. Cyclopoida: Rhynchomolgidae: Critomolgus 141

C D

B A

F H

Fig. 74. Critomolgus vicinus, female. A. maxillule; B. maxilla; C. maxilliped; D. leg 1; E, leg 2; F. third endopodal segment of leg 3; G. leg 4; H. leg 5 exopod. Scales: A=0.02 mm, B, C, H=0.05 mm, D-G=0.1 mm. 142 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

F E C

Fig. 75. Critomolgus vicinus, male. A. habitus, dorsal; B. urosome, ventral; C. maxilliped; D. leg 1; E. third endopodal segment of leg 2; F. leg 5. Scales: A=0.2 mm, B=0.1 mm, C-F=0.05 mm.

Legs 1 and 22 (Fig. 74D, E), leg 3, and leg 4 (Fig. 74G) segmented and armed as in preceding species. Leg 4 with 2 terminal spines of 58 μm (outer) and 108 μm (medial), respectively.

Leg 5 slender; exopod (Fig. 74H) 185 μm long, with distinct proximal swelling (45 μm wide across this region) and small spinules on outer side, and armed with 2 terminal setae of 65 and 112 μm Cyclopoida: Rhynchomolgidae: Critomolgus 143 long, respectively; distally region 31 μm wide. Leg 6 represented by 2 small setae on genital operculum.

Male: Body (Fig. 75A) length 1.46 mm (1.21-1.74 mm) and greatest width 0.58 mm (0.51-0.64 mm), based on 8 specimens. Urosome (Fig. 75B) 6-segmented. Fifth pedigerous somite 167 μm wide. Genital somite 218×225 μm. Four abdominal somites 60×105, 57×98, 40×90, and 50×95

μm, respectively. Caudal ramus 65×45 μm, ratio 1.44 : 1.

Rostrum as in female. Antennule with 3 additional aesthetascs (2 on second and 1 on fourth segments). Antenna with spinulation on second segment more distinct than in female.

Maxilliped (Fig. 75C) second segment with longitudinal row of spinules, 2 identical setae ner midlength and patch of minute spinules near inner distal area. Other mouthparts as in female.

Leg 1 (Fig. 75D) with 2 spines and 4 setae on third endopodal segment. Leg 2 with distal, foliaceous lamella on third endopodal segment (Fig. 75E). Legs 3 and 4 as in female.

Leg 5 (Fig. 75F) exopod 105×22 μm, ratio 4.77 : 1, without proximal swelling; terminal spine 43 μm long and seta 52 μm long. Leg 6 represented by 2 unequal, naked setae on genital operculum.

Distribution: Korea. Korea: GW, JB, CN. Specimens examined: All specimens were collected from the external surface of the actiniarian

Anthopleura midori Uchida and Muramatsu. 16♀♀, 9♂♂ (type specimens), intertidal, Aninjin,

Gangneung, Gangwon-do, 21.vi.1992 (collected along with Critomolgus anthopleurus Kim); 23♀♀, 34♂♂, Gyeokpo, Buan-gun, Jeollabuk-do, 23.v.1993. Remarks: Although this species and C. anthopleurus are co-associates of A. midori, the former species is not distributed on the coast of the Yellow Sea.

36. Critomolgus nudus Kim, 2000 (Fig. 76)

Ba-da-jo-reum-i (바다조름이)

Critomolgus nudus Kim, 2000a, p. 423, figs. 35-37.

Female: Body (Fig. 76A) 2.00 mm long. Greatest width 1.00 mm. Prosome ovoid, broad and dorsoventrally flat, with weak suture line between cephalosome and first pedigerous somite. Uro- some (Fig. 76B) 5-segmented, with thick exoskeleton. Fifth pedigerous somite 290 μm wide. Gen- ital double-somite 293×305 μm, with greatest width across anterior third and weak lobate process just posterior to genital aperture. Genital aperture located dorsolaterally. Three abdominal somites 115 ×180, 85×163, and 110×162 μm, respectively. First two abdominal somites each broadened posteriorly. Caudal ramus short, 77×64 μm (1.20 : 1), with six setae. Outer lateral seta naked, located in the middle of, and about twice as long as, caudal ramus. Inner dorsal seta naked. Other 4 setae plumose. Rostrum weakly developed, its posterior margin obscure. Antennule slender, 592 μm long, with armature formula of 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Setae relatively short. Antenna (Fig. 76C) stocky, with armature formula of 1, 1, 2+claw, 5+2 claws. Setae on first two segments small. Terminal segment about twice as long as wide; two of setae near outer distal corner short and broad. Two terminal claws strong. 144 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Labrum deeply incised; lateral margins nearly parallel; posterior margin unornamented. Man- dible (Fig. 76D) with deep proximal notch; terminal lash narrow, spinulated and moderately long; inner margin perpendicular to lash, with row of fine spinules; convex side broadly produced, with row of spinules and 1 small, dentiform process. Paragnath (Fig. 35H) a lobe, distally hairy. Maxil- lule with 1 lateral seta (or process) and 3 broad terminal setae. Maxilla (Fig. 76E) 2-segmented; first segment with minute spinules on outer (ventral) surface; second segment with 1 minute outer seta,

1 long, curved seta (spinulated on inner margin) on inner margin, 1 small seta on anterior surface, and 1 small denticle near base of lash; lash shorter than seta on inner margin, with serrate convex margin. Maxilliped 3-segmented and stocky; first segment unarmed; second segment with 2 simple setae of unequal length; third segment terminating into blunt process, with 1 spine and 1 small seta.

Legs 1-3 with 3-segmented rami. Leg 4 (Fig. 76F) with 3-segmented exopod and 2-segmented endopod; inner seta on coxa small and naked; both terminal corners of second endopodal segment with pointed process; terminal spines unequal in length. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; II

Leg 5 consisting of 1 naked seta on fifth pedigerous somite and free exopod. Exopod (Fig. 76G)

109×38 μm (2.87 : 1), with convex inner margin and 2 subequal, naked setae; these 2 setae longer than exopod. Leg 6 represented by 2 spinules and 1 spinule-like process in genital aperture. Male: Body resembling that of female. Length 1.80 mm. Greatest width 0.80 mm. Urosome 6-segmented. Fifth pedigerous somite short, 230 μm wide. Genital somite nearly spherical, large, 370×450 μm. Four abdominal somites 75×138, 70×130, 50×120, 70×132 μm, respectively. Cau- dal ramus 60×52 μm (1.15 : 1). Antennule with 3 additional aesthetascs: 2 on second and 1 on fourth segments. Antenna as in female. Maxilliped consisting of 3 segments and claw. First segment unarmed and similar in length to second segment. Second segment with 1 row of spinules on inner side, and 2 equal, small setae at midlength of inner margin. Third segment unarmed. Claw strongly curved, proximally with 1 small and 1 thick seta. Terminal endopodal segment of leg 1 with 2 terminal processes more prominent than in female, but armature formula of legs 1-4 as in female. Leg 5 exopod rather varied in shape, 55×23 μm (2.39: 1). Leg 6 represented by 2 small setae on genital operculum.

Distribution: Korea. Korea: JB, GG.

Specimens examined: 7♀♀, 13♂♂ (type specimens) from the pennatulacean, Virgularia gustaviana (Herklots), dug from the intertidal mud flat at Jakyak-do I., Incheon, 13.xii.1996; 3♀♀ from V. gustaviana, Gosapo Beach, Buan-gun, Jeollabuk-do, 6.x.2006; 23♀♀, 35♂♂ from V. gustaviana, weoldoo (tidal flat), Muan-gun, Jeollanam-do, 18.x.2012, collected by H.S. Hwang. Remarks: Unlike other three species of the same genus from Korea, this species has a patch of minute spinules on the basal segment of maxilla, no sexual dimorphism on the endopod of leg 1, and the small exopod of leg 5, which is widest in the middle. Cyclopoida: Rhynchomolgidae: Critomolgus 145

B C

E D

Fig. 76. Critomolgus nudus, female. A. habitus, dorsal; B. urosome, dorsal; C. antenna; D. mandible; E. maxilla; F. leg 4; G. leg 5 exopod. Scales: A=0.5 mm, B, C, F=0.1 mm, D, E, G=0.05 mm. 146 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Genus Lutumidomus Kim, 2006

Gae-peol-mal-mi-jal-i-sok (개펄말미잘이속)

Body generally narrow. Antennule 7-segmented. Antenna 4-segmented; terminal segment armed with 1 claw and 1 elongated seta in addition to other smaller setae. Mandible narrow. Max- illule with 2 setae apically. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 2-segmented endopod. Third endopodal segment of leg 1 armed with 2 spines and 4 setae in both sexes. Third endopodal segment of leg 3 with 2 spines and 2 setae. Leg 4 endopod with 1 seta on the first segment and 2 spines plus 3 setae on the second segment. Containing 3 species associated with sea anemone.

Key to the species of genus Lutumidomus

1. ‌Legs 2-4 with second exopodal segment bearing outer spine, and with third exopodal segment bearing 4 spines ···························································································································· L. tertius - ‌Legs 2-4 with second exopodal segment bearing no outer spine, and with third exopodal segment bearing less than 4 spines ··············································································································· 2 2. Body length more than 2.0 mm; caudal ramus about 4 times as long as wide ··············· L. panikkari - Body length less than 1.5 mm; caudal ramus less than 3 times as long as wide ················ L. parvus

37. Lutumidomus panikkari (Gnanamuthu, 1955) (Figs. 77-79)

Keun-gae-peol-mal-mi-jal-i (큰개펄말미잘이)

Lichomolgus panikkari Gnanamuthu, 1955, p. 151, figs. 1-3. Indomolgus panikkari: Humes and Ho, 1966, p. 11; Humes and Stock, 1973, p. 185. Lutumidomus panikkari: Kim, 2006c, p. 146, figs. 1, 2.

Female: Body (Fig. 77A) large and narrow, with weak exoskeleton. Body length 2.83 mm. Pro- some 1.54 mm long. Greatest width of prosome 811 μm. Cephalothorax divided by dorsal suture line into cephalosome and first pedigerous somite. Urosome (Fig. 77B) 5-segmented. Fifth pedigerous somite 429 μm wide. Genital double-somite 313×370 μm, with laterally expanded anterior part and narrower posterior part; genital aperture large, positioned dorsally at posterior part of anterior expansion. Three abdominal somites 125×204, 104×183, and 125×208 μm, respectively. All of these somites smooth, without ornamentation. Caudal rami divergent; each ramus 445×100 μm

(ratio 4.45 : 1) with 6 naked setae; outer lateral seta located near middle of lateral margin; all of setae shorter than ramus. Rostrum faint and strongly tapering; posterior apex not discernible. Antennule 7-segmented; armature formula 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all of setae naked and thin; all of aethetascs setiform and confluent with setae. Antenna (Fig. 77D) 4-segmented, with armature formula of 1, 1, 3, and 6+claw; second segment slightly inflated, with spinule row on outer margin; terminal segment 1.81 times as long as wide; claw strongly curved, rather slender, slightly longer than terminal segment; one of terminal setae distinctly larger than other seta and longer than claw. Cyclopoida: Rhynchomolgidae: Lutumidomus 147

Labrum (Fig. 77E) with elongated and divergent posterior lobes. Mandible (Fig. 77F) narrow; inner proximal notch shallow; inner margin with row of spinules; convex outer margin with 1 scale followed by serrate margin distally; distal lash moderately long and spinulose. Maxillule

(Fig. 77G) elongate and terminally with 2 large, naked setae. Maxilla (Fig. 78A) 2-segmented; first segment unarmed; second segment with simple anterior seta, spiniform inner seta bearing several spinules on distal margin; outer proximal seta (seta III) not seen. Distal lash arched, with large spinules along outer convex margin. Maxilliped (Fig. 77H) 3-segmented; first segment unarmed; second segment with 2 spinulose inner setae; terminal segment tapering, terminal with conical process, with 2 setae.

Legs 1-3 (Fig. 78B-D) with 3-segmented exopod and endopod. Leg 4 (Fig. 78E) with 3-segmented exopod and 2-segmented endopod. Third exopodal segment of legs 2-4 swollen, without spine on outer margin. All spines of legs 2-4 minimized and digitiform. All setae and spines on third exopodal segment of legs 3 and 4 characteristically located on inner margin. Outer seta on basis of legs 1-4 naked and small. Inner coxal seta of leg 4 naked, but that of legs 1-3 pinnate. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; IV, 4; enp. 0-1; 0-1; I, I, 4 Leg 2: coxa 0-1; basis 1-0; exp. I-0; 0-1; III, 5; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; 0-1; II, 5; enp. 0-1; 0-2; II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; 0-1; II, 5; enp. 0-1; II, 3

Leg 5 consisting of exopod and 1 seta on fifth pedigerous somite. Exopod (Fig. 78F) 140×53 μm

(ratio 2.64 : 1), tapering, blunt at tip, with 2 stiff subdistal setae. Leg 6 represented by 2 minute setae on genital operculum.

Male: Body (Fig. 79A) similar to that of female. Body length 2.68 mm. Prosome 1.46 mm long.

Greatest width of prosome 702 μm. Urosome (Fig. 79B) 6-segmented. Fifth pedigerous somite 333 μm wide. Genital somite nearly circular in dorsal view, 351×400 μm. Suture line obscure between fifth pedigerous and genital somites. Four abdominal somites 143×184, 123×167, 96×147, and

123×167 μm. Caudal ramus 324×79 μm (ratio 4.10 : 1). Outer lateral seta located slightly anterior to middle of lateral margin. Two median terminal setae slightly longer than caudal ramus, other 2 terminal setae shorter. Rostrum longer than wide, with round posterior apex. Antennule 537 μm long and 7-segmented; armature formula not different from that of female. Antenna as in female; distalmost seta on third segment distinctly smaller than 2 proximal setae; terminal segment 99×48 μm (ratio 2.06 : 1).

Labrum, mandible, maxillule and maxilla as in female. Maxilliped (Fig. 79C) consisting of 3 segments and terminal claw; first segment largest among segments, with 1 minute spinule at inner distal corner; second segment with 2 inner setae of similar size and small spinules on proximal half of inner side; third segment small and unarmed; terminal claw with 2 proximal setae and membranous flange along concave margin.

Legs 1-4 as in female. All spines and setae on third exopodal segment (Fig. 79D) of leg 3 positioned on inner margin. All setae and spines on third exopodal segment of legs 3 and 4 characteristically located on inner margin (Fig. 79D).

Leg 5 exopod 110×32 μm (ratio 3.44 : 1). Leg 6 represented by 2 thin setae on posterior margin genital operculum. Spermatophore sac-like, 364×175 μm.

Distribution: India and Korea. Korea: JN, GG. 148 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

D E

H F

Fig. 77. Lutumicomus panikkari, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. maxillule; H. maxilliped. Scales: A=0.5 mm, B, C=0.1 mm, D, E, H=0.05 mm, F, G=0.02 mm. Cyclopoida: Rhynchomolgidae: Lutumidomus 149

B C

A F

Fig. 78. Lutumicomus panikkari, female. A. maxilla; B. leg 1; C. leg 2; D. leg 3; E. leg 4; F. exopod of leg 5. Scales: A, F=0.05 mm, B-E=0.05 mm. 150 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A C B

Fig. 79. Lutumicomus panikkari, male. A. habitus, dorsal; B. urosome, dorsal; C. maxilliped; D. exopod of leg 3. Scales: A=0.5 mm, B=0.2 mm, C, D=0.1 mm.

Specimens examined: 1♂ from the sea anemone Paracondylactis hertwigi (Wassilieff, 1908), tidal flat, Yongyu I., Incheon, 01.iii.2006; ♀1 from washings of an unidentified sea anemone, intertidal mud flat. Waon, Sooncheon Bay, Jeollanam-do, 22.vi.2009.

Remarks: This species was originally described as Lichomolgus panikkari from India (Gnanamuthu, 1955). Subsequently, it was removed to the genus Indomolgus Humes and Ho, 1966 by Humes and

Ho (1966) and then to the genus Lutumidomus Kim, 2006 by Kim (2006). Cyclopoida: Rhynchomolgidae: Lutumidomus 151

38. Lutumidomus parvus Kim, 2006 (Figs. 80, 81)

Wae-gae-peol-mal-mi-jal-i (왜개펄말미잘이)

Lutumidomus parvus Kim, 2006c, p. 149, figs. 3-5.

Female: Body (Fig. 80A) narrow, nearly cylindrical, with weak exoskeleton. Body length 1.41 mm, excluding caudal setae. Greatest width of prosome 420 μm. Cephalothorax divided by dorsal suture line into cephalosome and first pedigerous somite. cephalosome 365 μm long. Urosome (Fig. 80B) 5-segmented, gradually narrowed posteriorly. Genital double-somite 212×195 μm, with slightly expanded anterior part and narrower, weakly tapering posterior part. Genital aperture located dorsolaterally at posterior area of slightly expanded anterior part. Suture line faint between fifth pedigerous and genital double-somites. Genital double-somite, abdominal somites and caudal rami with numerous fine hairs on surface. Three abdominal somites 62 ×120, 43×110, and

88×115 μm. Caudal ramus (Fig. 80C) slightly tapering, 100×43 μm or 2.33 times as long as wide, with 6 naked setae. Outer lateral seta located slightly posterior to middle of lateral margin. One of terminal setae as long as caudal ramus, but other setae shorter.

Rostrum conically projected to ventral direction. Antennule (Fig. 80D) 258 μm long and 7-segmented, with armature formula of 4, 13, 5, 3, 5, 2+aesthetasc, and 7+aesthetasc; all setae thin and naked; aethetascs setiform and confluent with setae. Antenna (Fig. 80E) 4-segmented, with armature formula of 1, 1, 3, and 5+claw; second segment similar to first in length, with setules on outer margin; distal one of 3 setae on third segment distinctly smaller than other two; fourth segment approximately 46×20 μm (ratio 2.30 : 1); terminal claw strongly curved. One of setae on 4th segment enlarged, straight, and more than twice as long as claw.

Labrum not observed. Mandible (Fig. 80F) slender and strongly curved. Inner proximal notch obscure. Inner margin with row of spinules. Convex outer margin with 1 longer and 1 shorter rows of spinules. Distal lash long, with several spinules on each side. Maxillule (Fig. 80G) slender and terminally with 2 naked setae. Maxilla (Fig. 80H) 2-segmented. First segment unarmed. Second segment with outer proximal seta, simple anterior seta, and smooth, spiniform inner seta. Distal lash terminally bifurcate, with 6 large spines along outer convex margin and 5 spinules on inner concave margin. Maxilliped (Fig. 80I) 3-segmented. Largest first segment unarmed. Second segment with 2 small, unequal setae on inner side. Third segment (or third and fourth segments) divided by faint suture line into unarmed proximal part and strongly tapering distal part bearing pointed tip and 2 small seta.

Legs 1-3 (Fig. 81A-C) with 3-segmented exopod and endopod. Leg 4 (Fig. 81D) with 3-segmented exopod and 2-segmented endopod. All spines on legs 2-4 setiform. Outer seta on basis of legs 1-4 naked. Inner coxal seta of these legs plumous. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, I, 4 Leg 2: coxa 0-1; basis 1-0; exp. I-0; 0-1; II, I, 5; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; 0-1; II, 5; enp. 0-1; 0-2; II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; 0-1; II, 5; enp. 0-1; II, 3

Leg 5 consisting of free exopod and 1 seta near base exopod. Exopod (Fig. 81E) 87×18 μm, or 4.83 times as long as wide, smooth, slightly tapering distally, with 2 thin and long distal setae. Leg 6 represented by 1 small seta and 1 small spine in genital aperture (Fig. 81F).

Male: Body (Fig. 81G) similar to that of female. Body length 1.35 mm. Greatest width of body

330 μm. Urosome 6-segmented. Genital somite (Fig. 81H) nearly quadrate in dorsal view, 185×190 152 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

I G

Fig. 80. Lutumidomus parvus, female. A. habitus, dorsal; B. urosome, dorsal; C. caudal ramus, ventral; D. antennule; E. antenna; F. mandible; G. maxillule; H. maxilla; I. maxilliped. Scales: A=0.2 mm, B=0.1 mm, C-E=0.05 mm, F-I=0.02 mm. Cyclopoida: Rhynchomolgidae: Lutumidomus 153

A B G

C D

E I

H F

Fig. 81. Lutumidomus parvus. female: A. leg 1; B. leg 2; C. leg 3; D. leg 4; E. leg 5 exopod; F. genital aperture. male: G. habitus, dorsal; H. anterior part of urosome, ventral; I. maxilliped. Scales: A-D, H=0.05 mm, G=0.2 mm, E, F, I=0.02 mm. 154 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

μm. Four abdominal somites 63×112, 48×103, 33×98, and 67×98 μm.

Rostrum, antennule, antenna, mandible, maxillule, and maxilla as in female. Maxilliped (Fig. 81I) consisting of 3 segments and terminal claw. First segment unarmed. Second segment with 1 row of spinules and 2 similar setae. Small third segment unarmed. Terminal claw large, almost as long as 3 proximal segments combined and strongly curved, with 2 unequal proximal setae.

Legs 1-4 with armature formula identical to that of female. Leg 5 exopod 70×15 μm (ratio 4.67 : 1). Leg 6 represented by 2 thin setae and 1 minute denticle on posterior part of genital operculum.

Distribution: Korea. Korea: JB.

Specimens examined: 3♀♀, 1♂ (type specimens) collected from the gastrovascular cavity of the sea anemone Flosmaris mutsuensis (Uchida, 1938) at Gosapo Beach (35°39′28″N, 126°30′30″S), Yellow Sea, 29.x.2000. Remarks: This species can be distinguished from L. panikkari in the absence of scale on the convex side of mandible, the usual form of the third exopodal segment of legs 2-4, and the smaller caudal rami; from Lutumidomus tertius (Kim, 2000) in the absence of the outer spine on the second exopodal segment of legs 2-4. The body length of this species is 1.41 mm in the female, it is markedly smaller than other two species: 2.39 mm in L. tertius (see Kim, 2000) and 2.7-3.1 mm in L. panikkari (see Gnanamuthu, 1955).

39. Lutumidomus tertius (Kim, 2000) (Fig. 82)

Gae-peol-mal-mi-jal-i (개펄말미잘이)

Notoxynus tertius Kim, 2000a, p. 428, figs. 38, 39. Lutumidomus tertius: Kim, 2006c, p. 145.

Female: Body (Fig. 82A) narrow, almost cylindrical, stocky, 2.39 mm long, with very thin exoskeleton. Greatest width 0.61 mm. Prosome 5-segmented, with faint suture line between cephalosome and first pedigerous somite. Epimera of prosomal somites weakly developed. Urosome (Fig. 82B) 5-segmented, tapering, with weak segmentation. Fifth pedigerous somite distinctly wider than genital double-somite, 358 μm wide. Genital double-somite 375×296 μm (1.27 : 1). Genital aperture small, located dorsolaterally at anterior third of lateral margin. Three abdominal somites 129×212, 79×188, and 96×179 μm, respectively. Posteroventral border of anal somite with minute spinules. Caudal ramus 137×71 μm (1.93 : 1), with distinctly convex inner margin and several minute spinules on terminal margin. Caudal setae relatively short and naked. Outer lateral seta located at midlength of outer margin of caudal ramus. Rostrum semicircular and much wider than long. Antennule 487 μm long and 7-segmented, with armature formula of 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Setae thin and naked. Antenna (Fig. 82C) 4-segmented, with armature formula of 1, 1, 3, 6+1 claw. Third segment very short. Terminal segment about 77×34 μm (2.26 : 1), weakly curved. Terminal claw strongly curved. One of terminal setae very large, distinctly longer than terminal segment.

Labrum deeply incised, with unornamented, weakly tapering lobes. Mandible (Fig. 82D) with broad proximal notch. Terminal lash long and spinulated. Inner margin oblique to lash, with a row of setules. Maxillule (Fig. 82E) a slender lobe, with 2 terminal, subequal setae (in copepodid Cyclopoida: Rhynchomolgidae: Lutumidomus 155

F A B

D E

H I G J

Fig. 82. Lutumidomus tertius, female. A. habitus, dorsal; B. urosome, dorsal; C. antenna; D. mandible; E. maxillule; F. maxilla; G. leg 1; H. third endopodal segment of leg 2; I. leg 4; J. leg 5. Scales: A=0.5 mm, B=0.1 mm, C, G-J=0.05 mm, D-F=0.02 mm. 156 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

V, number of these setae being 3). Maxilla (Fig. 82F) 2-segmented. First segment unarmed. Sec- ond segment terminating in moderately long lash bearing spinules on convex margin, with 1 minute outer seta, 1 naked seta on anterior surface, and 1 relatively small, distally spinulated seta on concave margin. Maxilliped 3-segmented. First segment unarmed and slightly longer than wide. Second segment with 2 simple, unequal setae on inner margin. Terminal segment blunt, tapering, with small, blunt terminal process, 1 small spine and 1 small seta.

Leg 1 (Fig. 82G) and legs 2 and 3 with 3-segmented rami. Leg 4 (Fig. 82I) with 3-segmented exopod and 2-segmented endopod. Hairs on setae and segments stiff and relatively thick. Second endopodal segment of leg 4 tapering, with 2 minute spiniform processes on outer margin; terminal 2 spines very unequal, inner one of which about twice as long as inner 1. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; II, 4 Leg 2: coxa 0-1; basis 1-0; exp. I-1; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-1; I-1; III, I, 5; enp. 0-1; 0-2; II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-1; I-1; III, I, 5; enp. 0-1; II, 3

Leg 5 (Fig. 82J) composed of 1 naked seta on fifth pedigerous and free exopod. Exopod distinctly tapering, 142×54 μm (2.63 : 1), with minute spinules on surface and 2 naked terminal setae (outer one 110 μm, and inner one 83 μm). Leg 6 represented by 2 spinules in genital aperture. Male: Body resembling that of female. Length 1.96 mm. Greatest width 258 μm. Urosome 6-segmented, with incomplete segmentation between fifth pedigerous somite and genital somite. Genital somite distally broadened, 292×296 μm. Four abdominal somites 120×296, 96×171,

54×156, and 71×154 μm, respectively. Caudal ramus 113×56 μm (2.02 : 1). Antennule with armature formula identical to that of female. Antenna with spinules on outer margin of second segment. Maxilliped consisting of 3 segments and terminal claw. First segment unarmed and longest. Second segment with longitudinal row of spinules and 2 identical setae on inner side. Third segment unarmed. Claw evenly curved, relatively large, proximally with 1 setule and 1 seta bearing spinules distally. Legs 1-4 with armature formula identical to that of female. Leg 5 identical in shape to that of female, 92×47 μm (1.96 : 1). Leg 6 represented by 2 thin setae and 1 minute spinule on genital operculum; the latter located at base of inner seta.

Distribution: Korea. Korea: GN, JB, GG.

Specimens examined: 4♀♀, 3♂♂ (type specimens) from the gastrovascular cavity of the mud-inhabiting sea anemone, Cerianthus filiformis Carlgren, collected in the intertidal mud flat at Jakyak-do I., Incheon, 28.ix.1996; 3♀♀, 2♂♂ from a sea anemone, on tidal flat, Sanglok Beach, Buan-gun, Jeollabuk-do, 28.x.2000; 3♀♀, 1♂ from C. filiformis, Geumsong-ri, Namhae I., Gyeo- ngsangnam-do, 1.viii.2003; 1♂ from Paracondylactis hertwigi (Wassilieff, 1908), tidal flat, Yongyu I., Incheon, 01.iii.2006. Remarks: The distinguished features of this species and differences with its congeners are mentioned in the remarks of the preceding species. Cyclopoida: Rhynchomolgidae: Paramolgus 157

Genus Paramolgus Humes and Stock, 1972

Ppul-san-ho-beol-re-sok (뿔산호벌레속)

Antenna 4-segmented, with 1 terminal claw. Mandible without any process or element on convex side. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 2-segmented endopod. Third exopodal segment of leg 4 with 3 spines and 5 setae. Armature of leg 4 endopod 0-1; II. Other characters as those of the family. Containing about 35 species associated with various cnidarians.

40. Paramolgus nudipes Kim, 2006 (Figs. 83-85)

Min-da-ri-ppul-san-ho-beol-re (민다리뿔산호벌레)

Paramolgus nudipes Kim, 2006c, p. 153, figs. 6-8.

Female: Body (Fig. 83A) stout. Body length of dissected specimen 2.02 mm, excluding setae on caudal rami. Greatest width 792 μm. Prosome 1.24 mm long. Cephalothorax divided by dorsal suture line into cephalosome and first pedigerous somite. First to fourth pedigerous somites nearly equal in length. Urosome (Fig. 83B) 5-segmented, all somites much wider than long. Fifth pedigerous somite short, 350 μm wide. Genital double-somite 269×373 μm, anteriorly expanded laterally. Genital apertures located dorsally at anterior 1-third. Three abdominal somites 108×246, 77×215, and 115×215 μm. Caudal ramus (Fig. 83C) tapering, 130 ×57 μm or 2.28 times as long as wide, with 6 naked setae. Outer lateral seta located near posterior 1-third of lateral margin. One of terminal setae enlarged, more than twice as long as caudal ramus, but other setae shorter than ramus.

Rostrum (Fig. 83D) strongly tapering posteriorly and continued as ridge to labrum. Antennule (Fig. 83E) 294 μm long and 7-segmented, with armature formula of 3, 11, 6, 3, 4 +aesthetasc, 2+aesthetasc, and 7+aesthetasc. All setae short and naked. Aethetascs small. Antenna (Fig. 83F) 4-segmented, with armature formula of 1, 1, 2, and 4+claw. First segment wider than long, with 1 small inner distal seta. Second segment distinctly longer than first segment, with 1 small seta on inner margin. Fourth segment slightly curved and approximately 3 times as long as wide. Terminal claw strongly curved in middle. Outer distal seta strongly curved to lateral direction.

Labrum (Fig. 83G) with broad posterior lobes, each lobe with small point and minute spinules on posterior margin. Mandible (Fig. 83H) with deep and broad inner proximal notch; inner margin bilobed and expanded. Outer margin distally with several small spinules. Distal lash elongated, denticulated along whole outer margin and along distal half of inner margin. Maxillule (Fig. 83I) with protuberant inner margin and armed with 3 distal setae (inner one of them distinctly smaller) and 1 subdistal setiform element, Maxilla (Fig. 84A) 2-segmented. First segment unarmed. Second segment with spiniform anterior and inner setae. Inner seta with sense spinules along distal margin. Outer proximal seta not seen. Distal lash arched, with spinules along outer convex margin.

Maxilliped (Fig. 84B) 3-segmented. First segment broad and unarmed. Second segment with 1 large, spiniform seta and 1 small seta. Third segment tapering, distally with 3 spines and blunt tip.

Legs 1-3 (Fig. 84C-E) with 3-segmented exopod and endopod. Leg 4 (Fig. 84F) with 3-segmented exopod and 2-segmented endopod. All setae naked and small. Armature formula of legs 1-4 as 158 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

F C

Fig. 83. Paramolgus nudipes, female. A. habitus, dorsal; B. urosome, dorsal; C. left caudal ramus, dorsal; D. rostral area, ventral; E. antennule; F. antenna; G. labrum; H. mandible; I. maxillule. Scales: A=0.5 mm, B, D=0.1 mm, C, E, F=0.05 mm, G-I=0.02 mm. Cyclopoida: Rhynchomolgidae: Paramolgus 159

D I E

Fig. 84. Paramolgus nudipes. female: A. maxilla; B. maxilliped; C. leg 1; D. leg 2; E. leg 3 endopod; F. leg 4; G. leg 5; H. right genital aperture. male: I. habitus, dorsal. Scales: A, B=0.02 mm, C-H=0.05 mm, I=0.2 mm. 160 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B C

Fig. 85. Paramolgus nudipes, male. A. anterior part of urosome, ventral; B. maxilliped; C. leg 1. Scales: A=0.1 mm, B=0.02 mm, C=0.05 mm.

follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, I, 4 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; II

Leg 5 (Fig. 84G) consisting of exopod and 1 seta near base exopod. Exopod 50×25 μm (ratio 2.0 : 1), subquadrate, with 2 naked distal setae. Leg 6 represented by 2 small spines in genital aperture.

Male: Body (Fig. 84I) narrower than that of female. Body length 1.45 mm. Greatest width of body 444 μm. Urosome 6-segmented. Fifth pedigerous somite 187 μm wide. Genital somite (Fig. 85A) 219×223 μm, with round anterior and posterior corners. Four abdominal somites 65 ×121, 63×106, 42×96, and 67×104 μm. All somites unornamented. Rostrum as in female. Antennule with 3 additional aeathetascs: 2 on second and 1 on fourth segments. These 3 aesthetascs longer than antennule. Antenna, mandible, maxillule, and maxilla as in female. Maxilliped (Fig. 85B) consisting of 3 segments and terminal claw. First segment unarmed. Second segment with 2 small spiniform setae of similar sizes. Small third segment unarmed. Ter- minal claw large, almost as long as 3 proximal segments combined and strongly curved near middle, with 2 unequal proximal setae.

Leg 1 (Fig. 85C) and legs 2-4 with armature formula identical to that of female. Setae of these legs distinctly more developed and plumous. Leg 5 exopod about twice as long as wide, distally with 1 spine and 1 small seta.

Distribution: Korea. Korea: JB.

Specimens examined: 19♀♀, 1♂ (type specimens) from the gastrovascular cavity of the sea Cyclopoida: Rhynchomolgidae: Paramolgus 161

anemone Flosmaris mutsuensis (Uchida), Gosapo Beach, Buan-gun, Jeollabuk-do, 20.x.2000. Remarks: This species has the following combination of diagnostic characters that serve to dis- tinghuish it from other species of the genus: the caudal ramus of the female is tapering and about 2.3 times as long as wide, the third endopodal segment of leg 1 is armed with 2 spine and 4 setae in both sexes, the female genital double-somite is distinctly wider than long, and the exopod of female leg 5 lacks a proximal swelling.

41. Paramolgus orientalis Choe and Kim, 2010 (Figs. 86-88)

Dong-yang-ppul-san-ho-beol-re (동양뿔산호벌레)

Paramolgus orientalis Choe and Kim, 2010, p. 49, figs. 1-3.

Female: Body (Fig. 86A) moderately narrow. Body length of dissected paratype 1.64 mm. Great- est width 600 μm. Prosome 951 μm long. Cephalothorax 610×600 μm, divided by dorsal suture line into cephalosome and first pedigerous somite. Second pedigerous somite posterolaterally produced. Urosome 5-segmented. Fifth pedigerous somite 183 μm wide. Genital double-somite (Fig. 86B) 225×197 μm, divided by distinct lateral incision into roundly expanded anterior 2/3 and narrower posterior 1/3; narrower posterior part 128 μm wide; genital apertures located dorsally in midway of anterior part. Three free abdominal somites 70×100, 52×84, and 85×110 μm, respectively; second abdominal somite narrower than preceding and following somites. Anal somite trapezoid, with fine spinules on posterolateral border both dorsally and ventrally. Caudal rami (Fig. 86B) well isolated from each other; each ramus 190×45 μm, 4.22 times as long as wide, with 6 setae; outer lateral seta located subdistally.

Rostrum (Fig. 86C) large and hemicircular. Antennule (Fig. 86D) 7-segmented; numbers of seta on first to last segments 4, 13, 6, 3, 4, 2, and 7; distal 3 segments each with 1 aesthetasc, in addition to setae; all setae smooth. Antenna (Fig. 86E) 4-segmented, with 1 terminal claw; first and second segments each armed with 1 inner seta; short third segment with 3 inner setae; fourth segment 83×25 μm, 3.32 times as long as wide, distally with 6 small setae; terminal claw 75 μm long, slightly curved.

Labrum (Fig. 86F) with broad posterior incision; both lobes fringed with broad membrane on posterior margin. Mandible (Fig. 86G) with broad proximal notch; inner margin with setules proximally and spinules distally; convex margin with row of spinules; distal lash elongate with serrate margins. Maxillule (Fig. 86H) with 1 small lateral seta and 3 larger distal setae. Maxilla (Fig. 87A) 2-segmented; proximal segment unarmed; distal segment with small proximal seta; spiniform anterior seta, and large inner seta bearing 3 rows of spinules on distal margin; distal lash as long as inner seta, with 1 row of spinules and additional row of larger spinules on proximal half of distal margin. Maxilliped (Fig. 87B) 3-segmented; first segment unarmed; second segment nearly as long as first; with 2 smooth similar setae; terminal segment terminated by nipple shaped process and armed with 2 setae subdistally, one of them spinule-bearing and spiniform.

Legs 1-3 (Fig. 87C-E) with 3-segmented rami. Leg 4 (Fig. 87F) with 3-segmented exopod and 2-segmented endopod. Legs 1-4 with smooth outer seta on basis. Leg 4 with second endopodal segment of 83×23 μm; its 2 distal spines 32 (outer) and 65 μm long. Armature formula of legs 1-4 as follows: 162 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A H D

G F

Fig. 86. Paramolgus orientalis, female. A. habitusm dorsal; B. genital double-somite and abdomen, dorsal; C. rostral area, ventral; D. antennule; E. antenna; F. labrum; G. mandible; H. maxillule. Scales: A=0.2 mm, B, C=0.1 mm, D-F=0.05 mm, G, H=0.02 mm. Cyclopoida: Rhynchomolgidae: Paramolgus 163

A C

Fig. 87. Paramolgus orientalis, female. A. maxilla; B. maxilliped; C. leg 1; D. leg 2; E. leg 3 endopod; F. leg 4; G. leg 4 exopod. Scales: A, B=0.2 mm, C-G=0.05 mm. 164 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

B E C

Fig. 88. Paramolgus orientalis, male. A. urosome, ventral; B. antenna; C. maxilliped; D. leg 1 endopod; E. leg 5 exopod. Scales: A=0.1 mm, B-D=0.05 mm, E=0.02 mm.

Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 1, 4 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; II

Leg 5 consisting of small dorsolateral seta on fifth pedigerous somite and free exopod; exopod (Fig. 87G) slightly expanded proximally and gradually narrowed distally, with many scattered minute spinules (or scales) and 2 terminal setae of 100 μm (outer) and 40 μm (inner). Leg 6 represented by 2 small setae in genital aperture.

Male: Body narrower than that of female. Body length of dissected specimen 1.38 μm. Urosome (Fig. 88A) 628 μm long and 6-segmented. Fifth pedigerous somite narrow, 115 μm wide. Geni- tal somite 252×207 μm, longer than wide. Four abdominal somites 52×75, 43×73, 30×68, and 53×83 μm. Caudal ramus 130×35 μm, 3.71 times as long as wide. Rostrum as in female. Antennule with 1 additional aesthetasc on second and 4th segments at places indicated by dots in Fig. 86D. Antenna (Fig. 88B) additionally with rows of scales (spinules) on proximal half of inner margin of second segment; fourth segment more longer than that of female, 90×21 μm, 4.29 times as long as wide.

Labrum, mandible, maxillule, and maxilla as in female. Maxilliped (Fig. 88C) consisting of 3 segments and terminal claw; first segment longest but unarmed; second segment with 1 longitudinal row of spinules, 2 unequal setae (one of them broad, with serrate margins), and 1 pointed distal process on expanded inner margin; small third segment unarmed; terminal claw strong, proximal- Cyclopoida: Rhynchomolgidae: Verutipes 165 ly with 2 extremely unequal setae.

Leg 1 with third endopodal segment (Fig. 88D) bearing 2 spines and 4 setae (armature formula I,

I, 4). Legs 2-4 as in female. Leg 5 exopod (Fig. 88E) rectangular, 57×13 μm, 4.38 times as long as wide, with few spinules on outer margin; 2 terminal setae 87 (outer) and 26 μm (inner). Leg 6 represented by 2 subequal setae on genital operculum (Fig. 88A).

Distribution: Korea. Korea: JJ.

Specimens examined: 2♀♀, 4♂♂ (type specimens) from washings of the antipatharian Antipa- thes japonicus Brook, off Sagye-ri (SCUBA), Andeok-myeon, south of Jeju Island, 22.ix.2009, M.-K. Choe. Remarks: The genus Paramolgus consists of more than 30 known species, of which 4 are associated with antipatharians. This species can be distinguished from its congeners by having 4 setae on the maxillule and the long caudal ramus of 4.22 times as long as wide (Choe and Kim, 2010).

Genus Verutipes Humes, 1982

Wang-ga-seum-mal-mi-jal-i-sok (왕가슴말미잘이속)

Body with swollen prosome and small urosome. Antenna 4-segmented, with 2 terminal claws. Mandible without any process or element on convex side. Legs 1-3 with 3-segmented rami and reduced setation. Leg 4 with 3-segmented exopod and 2-segmented endopod. Leg 4 endopod with armature 0-0; I. Containing 2 species associated with actiniarians.

42. Verutipes scutatus Kim, 2006 (Figs. 89-91)

Wang-ga-seum-mal-mi-jal-i (왕가슴말미잘이)

Verutipes scutatus Kim, 2006c, p. 157, figs. 9-11.

Female: Body (Fig. 89A) with greatly expanded prosome and narrow urosome. Body length of dissected largest specimen 1.69 mm excluding caudal setae. Prosome shield-shaped, unsegmented, but with 3 dorsal lines as vestigial segmentations. Anterior part of prosome broadened, with 2 pairs of small lateral lobes, posterior pair larger. Greatest length of prosome 1.38 mm, and greatest width 1.08 mm. Urosome (Fig. 89B) 5-segmented, but fifth pedigerous somite obscurely delimited. Genital double-somite much wider than long, anteriorly expanded, about 225 μm long, 270 μm wide, and obscurely delimited from fifth pedigerous somite, with rudimentary dorsal suture line between expanded anterior and narrower posterior parts. Three abdominal somites 85×150, 50×143, and 87×140 μm. Anal somite and caudal rami with fine hairs on surfaces, Caudal ramus distinctly tapering, 60×50 (ratio 1.20 : 1) μm, with 6 naked setae.

Rostrum lacking. Antennule (Fig. 89C) 410 μm long and 7-segmented, with armature formula of 4, 12, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Plumous setae: 1 on each fifth and 166 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

sixth segments and 4 on last segments. Antenna (Fig. 89D) 4-segmented, gradually narrowed from proximal to distal, with armature formula of 1, 1, 3, and 5+2 claws. Setae on these segments very small. Fourth segment approximately 93×48 μm. Two terminal claws recurved.

Labrum (Fig. 89E) with divergent posterior lobes. Mandible (Fig. 89F) with moderately distinct inner proximal notch. Inner margin linear, with fine spinules. Outer margin slightly expanded roundly, with row of minute spinules proximally and serration distally. Distal lash thin, elongate, and weakly serrate along both margins. Paragnath (Fig. 89G) lobate, with thick and dense setules distally. Maxillule (Fig. 90A) gradually narrowed distally, curved at distal 1-third, proximally with small point, and distally with 3 setae, inner one of them spiniform. Maxilla (Fig. 90B) 2-segmented. First segment expanded and unarmed. Second segment with small outer proximal seta, short, scal- pel-like anterior seta and spiniform inner seta bearing serration along distal margin. Distal lash long, with serrated convex margin and minute spinules along distal half of concave margin. Max- illiped (Fig. 90C) 3-segmented. First segment unarmed. Second segment longer than first, with 2 inner setae. Third segment with 1 spine, 1 seta, and spiniform process terminally.

Legs 1 and 2 (Fig. 90D, E) with 3-segmented exopod and endopod. Legs 3 and 4 (Figs. 90F, 91A) with 3-segmented exopod and 2-segmented endopod. First exopodal segment of legs 1-4 distally expanded, with rod-shaped outer spine. Other segments of these legs narrow. Setation of legs 1-4 variable. Setae naked or weakly plumous. Inner coxal seta lacking in legs 1-4. Armature formula of legs 1-4 as follows:

Leg 1: coxa 0-0; basis 1-0; exp. I-0; I-1; III, I, 3; enp. 0-1; 0-1 (or 0-0); II, 4

Leg 2: coxa 0-0; basis 1-0; exp. I-0; I-1 (or I-0); III, I, 3; enp. 0-1; 0-2 (or 0-0); II, I, 2

Leg 3: coxa 0-0; basis 1-0; exp. I-0; I-1; III, I, 1; enp. 0-0; I. 2 (or I)

Leg 4: coxa 0-0; basis 1-0; exp. I-0; I-1 (or I-0); III, I, 1; enp. 0-0; I

Leg 5 (Fig. 91B) consisting of free exopod and nearby seta. Exopod 28×14 μm, with convex outer margin; its 2 terminal setae 55 μm and 35 μm long. Leg 6 represented by 2 thick spines in genital aperture (Fig. 91C).

Male: Body (Fig. 91D) similar to that of female. Body length of dissected specimen 1.29 mm.

Greatest width 740 μm. Greatest length of prosome 800 μm. Urosome (Fig. 91E) 6-segmented. Fifth pedigerous somite fused with genital somite, with faint suture line between these somites. Combined fifth pedigerous and genital somites 213×353 μm. Genital somite with lobate posterolateral corners. Four abdominal somites gradually narrowed from proximal to distal, 73×197, 60×170, 33×140, and 60×117 μm. First to third abdominal somites with roundly convex lateral margins. Caudal ramus 40×40 μm. Rostrum absent as in female. Antennule with additional aesthetasc on distal area of second segment as indicated by dot in Fig. 89C. Antenna as in female.

Labrum, mandible, paragnath, maxillule, and maxilla as in female. Maxilliped (Fig. 91F) consisting of 3 segments and terminal claw. First segment unarmed. Second segment with strongly projected and tapering, 1 pointed and 1 blunt setae, and row of spinules on inner margin. Small third segment unarmed. Terminal claw stout, distinctly shorter than second segment, proximally with 1 small and 1 large setae.

Legs 1-4 basically as those of female, with variable setation. Leg 5 exopod (Fig. 91G) small, 16×14 μm, but its distal setae larger than in female, each 90 and 48 μm long.

Distribution: Korea. Korea: JJ.

Specimens examined: 4♀♀, 4♂♂ (type specimens) from the gastrovascular cavity of a sea Cyclopoida: Rhynchomolgidae: Verutipes 167

F G

Fig. 89. Verutipes scutatus, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. paragnath. Scales: A=0.2 mm, B=0.1 mm, C-E=0.05 mm, F, G=0.02 mm. 168 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Fig. 90. Verutipes scutatus, female. A. maxillule; B. maxilla; C. maxilliped; D. leg 1; E. leg 2; F. leg 3. Scales: A-C=0.02 mm, D-F=0.05 mm. Cyclopoida: Rhynchomolgidae: Verutipes 169

C E

Fig. 91. Verutipes scutatus. female: A. leg 4; B. leg 5; C. right genital aperture. male: D. habitus, dorsal; E. urosome, dorsal; F. maxilliped; G. leg 5. Scales: A, F=0.05 mm, B, C, G=0.02 mm, D=0.2 mm, E=0.1 mm. 170 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

anemone (green color), Moseulpo Port, depth about 3 m, 4.vi.2004. Remarks: The genus Verutipes consists of only rwo species, this species and V. laticeps Humes, 1982. Verutipes laticeps is known from the New Caledonia and has the following morphological features, as differences from this species: 1) the body size is much smaller, only 0.73-1.03 mm in the female and 0.61-0.95 mm in the male (vs. 1.69 and 1.29 mm in the female and male, respectively, in this species); 2) the prosome is distinctly T-shaped and clearly segmented (vs. not T-shaped in this species); 3) the caudal ramus is 2.01 times as long as wide in the female (vs. 1.20 times in this species); 4) the inner element on the second maxillary segment is setiform (vs. spiniform in this species) and spinulated on both margins; and 5) the outer spine on the first exopodal segment of legs

1-4 is tapering and pointed as usual (vs. rod-shaped, with rounded tip, in this species).

Genus Zamolgus Humes and Stock, 1972

Tteok-dol-gi-beol-re-sok (턱돌기벌레속)

Antenna 4-segmented, with 1 terminal claw. Mandible with large process on convex side. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 2-segmented endopod. Leg 1 endopod sexually dimorphic. Third exopodal segment of leg 4 with 4 spines and 5 setae. Leg 4 endopod with armature 0-1; II. Other characters as those of the family.

43. Zamolgus cavernularius Kim, 2000 (Figs. 92-94)

Tteok-dol-gi-beol-re (턱돌기벌레)

Zamolgus cavernularius Kim, 2000b, p. 251, figs. 1-3.

Female: Body (Fig. 92A) 1.63 mm long and greatest width 0.90 mm. Prosome broad, dorsoventrally flat, and composed of cephalosome and 4 pedigerous somites. Suture line between cephalosome and first pedigerous somite indistinct. Urosome (Fig. 92B) 5-segmented. Fifth pedigerous somite 255 μm wide; its latral margins strongly oblique. Genital double-somite greatly expanded laterally, much wider than fifth pedigerous somite, 153×330 μm, more than twice wider than long. Genital aperture located dorsally. Three abdominal somites 53×128, 56×119, and 83×130 μm, respectively. Caudal ramus 92×56 μm (ratio 1.64 : 1), with 6 setae. Egg sac (Fig. 92C) 438×276 μm, containing about 10 eggs.

Rostrum (Fig. 92D) broad, with rounded posterior apex. Antennule (Fig. 92E) slender, 7-segmented; armature formula 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all setae naked.

Antenna (Fig. 92F, G) 4-segmented; armature formula 1, 1, 2, and 6+claw; terminal segment about twice as long as wide; terminal claw strong.

Labrum (Fig. 92H) broad, with broad median incision; posterolateral part of each lobe produced inwardly. Mandible (Fig. 92I) with large, tapering process on convex side of blade; inner margin straight and spinulose; distal lash thin, elongate, and well defined from inner margin. Paragnath

(Fig. 92J) a spinulose lobe. Maxillule (Fig. 92K) with 3 setae distally. Maxilla (Fig. 93A) 2-segmented; basal segment unarmed; distal segment with terminated in rather short lash bearing serrate Cyclopoida: Rhynchomolgidae: Zamolgus 171

A E

H K J

Fig. 92. Zamolgus cavernularius, female. A. habitus, dorsal; B. urosome, dorsal; C. egg sac; D. rostral area, ventral; E. antennule; F. antenna; G. distal part of antenna; H. labrum; I. mandible; J. paragnath; K. maxillule. Scales: A, C, D=0.2 mm, B=0.1 mm, E-H=0.05 mm, I-K=0.02 mm. 172 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A C

F B H

Fig. 93. Zamolgus cavernularius, female. A. maxilla; B. maxilliped; C. distal part of maxilliped; D. leg 1; E. leg 2; F. leg 4; G. leg 5; H. leg 6. Scales: A, C, G, H=0.02 mm, B, D-F=0.05 mm. Cyclopoida: Rhynchomolgidae: Zamolgus 173

A D

Fig. 94. Zamolgus cavernularius, male. A. habitus, dorsal; B. urosome, ventral; C. maxilliped; D. leg 1 endopod; E. leg 5. Scales: A=0.2 mm, B=0.1 mm, C-E=0.05 mm.

outer margin; inner seta large, much longer than lash, with spinules along outer (distal) margin; an- teterior seta naked; proximal setaminute. Maxilliped (Fig. 93B, C) 3-segmented; first segment unarmed; second segment with 2 unequal setae; terminal segment bifurcate, distally forming 2 claws.

Legs 1 and 2 (Fig. 93D, E) and leg 3 with 3-segmented rami. Leg 4 (Fig. 93F) with 3-segmented exopod and 2-segmented endopod. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 3; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 3; enp. 0-1; 0-2; I, II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; II

Leg 5 consisting of 1 dorsolateral seta on fifth pedigerous somite and free exopod; exopod (Fig. 93G) consisting of broader proximal half and narrower distal half, with 2 naked setae and 1 conical process distally. Leg 6 represented by 2 small, spiniform setae on genital operculum (Fig. 93H).

Male: Body (Fig. 94A) similar to that of female. Body length 1.25 mm. Urosome (Fig. 94B) 6-segmented. Genital somite subcircular, 237×297 μm. Four abdominal somites 50×105, 48×97, 174 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

40×90, and 65×95 μm, respectively. Caudal ramus 73×41 μm (ratio 1.78 : 1). Rostrum as in female. Antennule with 3 additional aesthetascs: 2 on second and 1 on fourth segments. Antenna with minute spinules on inner margin of second segment.

Labrum, mandible, maxillule, maxilla as in female. Maxilliped (Fig. 94C) 4-segmented; first segment unarmed; second segment with 2 setae of similar lengths, 3 rows of denticles proximal to setae; short third segment unarmed; terminal segment as long , evenly curved claw bearing 2 unequal setae proximally. Leg 1 endopod (Fig. 94D) with 2 spines and 4 setae on third segment. Legs 2-4 as in female.

Leg 5 exopod 63×22 μm (ratio 2.86 : 1), with parallel lateral margins and 2 distal setae. Leg 6 as 2 setae on genital operculum.

Distribution: Korea. Korea: CN, GG.

Specimens examined: 60♀♀, 20♂♂ (type specimens), from external surface of the pennatulacean Cavernularia obesa Valenciennes in Milne Edwards and Halime, tidal flat, Sapsi I., Choong- cheongnam-do, 26.x.1999; 20♀♀, 15♂♂ from C. obesa, tidal flat, Daebu-do, Ansan, Gyeonggi-do, 2.vi.2012; about 300 specimens from C. obesa, tidal flat, Daebu-do, Ansan, Gyeonggi-do, 17.x.2012, H.S. Hwang. Remarks: This species was found from a pennatulacean, unlike other three species of the genus which are known as associates of gorgonaceans. The laterally strongly expanded female genital double-somite and the 2 large, claw-like terminal processes on the female maxilliped are the outstanding features of Z. cavernularius.

Family Sabelliphilidae Gurney, 1927

Gaet-ji-reong-i-sal-i-gwa (갯지렁이살이과)

Body cyclopiform. Urosome 5-segmented in female and 6-segmented in male. Antennule 7-segmented. Antenna 4-segmented; third segment usually with 1 claw and 3 setae (sometimes 3 setae only or 1 claw plus 2 setae); terminal segment with 2-4 claws. Mandible short and strongly tapering; terminal lash not developed. Maxilla 2-segmented; distal segment terminating in lash bearing large teeth; inner seta developed. Maxilliped 3-segmented in female and 4-segmented in male. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and 1-3 segmented endopod. Leg 5 with free exopod bearing 2 elements. Associated with tubicolous polychaetes and Phoroni- da.

Key to the genera of family Sabelliphilidae (based on females)

1. Leg 4 endopod 3-segmented ···················································································································· 2 - Leg 4 endopod 1- or 2-segmented ··········································································································· 6 2. Third endopodal segment of leg 4 with 5 spines ························································ Eupolymniphilus - Third endopodal segment of leg 4 with 1 or 2 elements ······································································ 3 3. Antenna with 4 terminal claws ························································································ Myxomolgoides - Antenna with 3 terminal claws ················································································································ 4 4. Rostrum bifid; first and second segments of antennule expanded· ································ Sabelliphilus Cyclopoida: Sabelliphilidae: Eupolymniphilus 175

- Rostrum simple; first and second segments of antennule not expanded ·········································· 5 5. ‌Third segment of maxilliped elongate and claw-like; first and second segments of antennule without spines ························································································································ Serpuliphilus - ‌Third segment of maxilliped not elongate nor claw-like; first and second segments of antennule with spines ······························································································································ Myxomolgus 6. Leg 4 endopod 2-segmented ···················································································································· 7 - Leg 4 endopod 1-segmented ···················································································································· 8 7. Distal segment of maxilliped elongate, whip-like ······························································· Nasomolgus - Distal segment maxilliped not elongate or whip-like ···················································· Acaenomolgus 8. Leg 4 endopod with 1 spine ···································································································· Phoronicola - Leg 4 endopod with 2 spines ································································································ Tubiporincola

Genus Eupolymniphilus Humes and Boxshall, 1996

O-ga-si-gaet-ji-reong-i-sal-i-sok (오가시갯지렁이살이속)

Rostrum simple. Antenna 4-segmented, with 1 claw plus 3 setae on third segment and 3 or 4 spines plus setae on terminal segment. Leg 4 with 3-segmented rami, with 5 spines on third endopodal segment. Containing 7 known species.

44. Eupolymniphilus orientalis Kim, 2006 (Fig. 95)

Dong-yang-o-ga-si-gaet-ji-reong-i-sal-i (동양오가시갯지렁이살이)

Eupolymniphilus orientalis Kim, 2006a, p. 574, figs. 1-3.

Female: Body (Fig. 95A) rather slender. Body length of dissected specimen 1.01 mm, greatest width 413 μm. Prosome 688 μm long. Cephalothorax divided by dorsal suture line into cephalosome and first pedigerous somite. Lateral margin of epimera of third pedigerous somite bordered by narrow membrane. Urosome (Fig. 95B) 5-segmented, small, 317 μm long, less than half length of prosome. Fifth pedigerous somite 155 μm wide, with sleeve-like, large posterior extension on both sides near base of exopod of leg 5. Genital double-somite 155×125 μm, expanded near middle, 70 μm wide across narrowed posterior part. Genital apertures located laterally. Three abdominal somites 25×65, 25×65, and 17×63 μm, from anterior to posterior. Posteroventral margin of anal somite ornamented with fine spinules. Caudal ramus nearly rectangular, 42×28 μm (ratio

1.50 : 1), slightly shorter than anal somite; posteroventral margin equipped with fine spinules. Two median terminal setae out of 6 caudal setae weakly plumose, other 4 setae naked; proximal caudal seta located dorsally at midlength of caudal ramus and characteristically swollen proximally. Rostrum distinct and strongly tapering. Antennule 293 μm long and 7-segmented, with armature formula of 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 4+aesthetasc; all setae naked. First and second segments distinctly wider than remaining ons. Two of 4 setae on first segment distinctly larger than other 2 setae on same segment. Terminal segment shorter than wide. Antenna (Fig.

95C) 4-segmented. First segment (coxobasis) with 1 inner distal seta. Second segment with 1 inner 176 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B C

D F

Fig. 95. Eupolymniphilus orientalis, female. A. habitus, dorsal; B. urosome, dorsal; C. antenna; D. mandible; E. maxillule; F. leg 4. Scales: A=0.1 mm, B, C, F=0.05 mm, D, E=0.02 mm.

seta and pointed outer distal corner. Third segment with 1 setiform claw and 3 setae. Fourth segment 50×20 μm, with 3 setiform claws and 4 setae.

Labrum posteriorly elongated, with deep and narrow posteromedian incision. Mandible (Fig. 95D) curved at right angle near base of blade, with 1 scale on convex side, a row of spines, and 1 Cyclopoida: Sabelliphilidae: Eupolymniphilus 177 isolated subterminal spine on concave margin of blade. Blade moderately elongate and evenly tapering. Convex margin of lash with numerous minute denticles. Maxillule (Fig. 95E) armed with 3 terminal naked setae and 1 lateral setiform element. Maxilla with unarmed first segment. Second segment with small proximal seta, naked anterior seta, elongate, spinulated inner setae, and spines along inner margin distal to inner seta. Distal lash slender and basally well demarcated from second segment. Maxilliped with unarmed first segment. Second segment with protruded outer margin in proximal half, with 1 small tubercle proximally near inner margin and 2 naked setae, one of these characteristically recurved. Third segment divided by incomplete proximal suture line into 2 parts, distally tapering, with 1 seta on distal part.

Legs 1-3 and leg 4 (Fig. 95F) with 3-segmented exopod and endopod. Third exopodal segment of legs 1-3 with bifid terminal process. Third endopodal segment of legs 3 and 4 with dentiform process on distal part of inner margin. Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; I, II, I+1 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-1; I, II, II Leg 5 consisting of free exopod and 1 plumose dorsolateral seta on fifth pedigerous somite. Ex- opod 36×18 μm, terminally serrated and armed with 1 long spine (112 μm) and 1 naked seta (74 μm). Leg 6 represented in genital aperture by plumose, proximally thickened anterior seta and naked posterior seta bearing rod-shaped base. Male: Body similar to that of female. Body length 858 μm. Greatest width 290 μm. Urosome 6-segmented. Fifth pedigerous somite 105 μm wide. Genital somite 150×137 μm. Four abdominal somites 28×52, 23×51, 13×50, and 25×50 μm, respectively. Caudal ramus 45×22 μm, ratio of length to width 2.05 : 1. Rostrum as in female. Antennule with 3 additional aesthetascs: 2 on second and 1 on fourth segments. Antenna with a few additional spinules on proximal half of inner margin of second segment. Mouthparts as in female except for maxilliped. Maxilliped 4-segmented including terminal claw. First segment unarmed. Second segment expanded in middle, with 1 longitudinal row of spinules and 2 inner setae. Third segment small and unarmed. Terminal claw arched, as long as 3 proximal segments combined, proximally with 1 large transformed seta and 1 small seta. Legs 1-4 as in female. Third endopodal segment of leg 1 armed as in female. Leg 5 exopod with parallel lateral margins. Leg 6 represented by 1 longer and 1 shorter seta on genital operculum.

Distribution: Korea. Korea: JJ.

Specimens examined: 6♀♀ and 1♂ from invertebrate burrows (mainly of polychaetes and decapods) in intertidal muds at Seogwipo Port (33°14′11″N 126°33′44″E), Jeju Island, 19.viii.2005.

Remarks: The genus Eupolymniphilus contains 7 known species. Recently et al. (2013) described E. mediterraneus as a new species and discussed that this Mediterranean species is closely related to this species, E. brecaudatus Kim, 2009 and E. occidentalis Kim, 2009. They could differentiate them by different proportional ratios of the length to width of caudal ramus of these species: 3.0 : 1 in E. mediterraneus, 1.50 : 1 in this species, 1 : 1 in E. brevicaudatus, and 2.69 : 1 in E. occidentalis. 178 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Genus Myxomolgus Humes and Stock, 1972

Gaet-ji-reong-i-jib-beol-re-sok (갯지렁이집벌레속)

Rostrum simple. Antennule with spines on first and second segments. Antenna with 1 spine plus 3 setae on third segment and 3 spines plus setae on terminal segment. Maxillule with 3 setae apically. Containing 4 known species associated with polychaetes.

45. Myxomolgus invulgus Kim, 2001 (Figs. 96-98)

Gaet-ji-reong-i-jib-beol-re (갯지렁이집벌레)

Myxomolgus invulgus Kim, 2001a, p. 5, figs. 4-6.

Female: Body (Fig. 96A) relatively slender. Body length, excluding rostrum, 1.27 mm. Greatest width 491 μm. Prosome consisting of cephalosome and 4 pedigerous somites. Cephalosome anteriorly tapering. Suture line between cephalosome and first pedigerous somite relatively distinct.

Urosome (Fig. 96B) 5-segmented. Fifth pedigerous somite 167 μm wide. Genital double-somite 145×181 μm, with anterior part greatly expanded laterally, and lateral margins of narrower posterior part nearly parallel. Three abdominal somites 44×87, 35×79, and 56×75 μm, respectively. Posteroventral margins of genital double- and first 2 abdominal somites rimmed with crenated membrane. Posteroventral margin of anal somite (Fig. 96C) with spinules. Caudal ramus (Fig.

96C) 64×28 μm (2.29 : 1), with slightly convex inner margin and 6 caudal setae, of which outer terminal and 2 mid-terminal ons plumose.

Rostrum (Fig. 96D) characteristically directed anteriorly, triangular, extending over anterior margin of cephalosome with pointed apex. Antennule (Fig. 96E) short, 214 μm, 7-segmented, short and stocky, characteristically bearing 2 small aesthetascs on second segment, with armature formula of

4, 13+2 aesthetascs, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc. Antenna (Fig. 96F) 4-segmented, with armature formula of 1, 1, 3+claw, and 4+3 claws. Two of 4 setae on fourth segment located on outer margin, their tip weakly bifurcated. Outer one of 3 claws on terminal segment slightly thicker than inner two. These 3 claws equal in length.

Labrum (Fig. 97A) with 2 narrow posterior lobes, with deep posteromedian incision and several minute spinules on inner margin of lobes. Mandible (Fig. 97B) short, with strongly tapering blade, without proximal notch. Convex margin with 1 conspicuous, claw-like outer scale and followed by numerous denticles. Inner margin of blade spinulated. Paragnath not observed. Maxillule (Fig. 97C) an elongate lobe with 1 small seta on lateral margin and 2 equal naked seta at apex. Maxil- la (Fig. 97D) 2-segmented. First segment unarmed. Second segment drawn out into moderately long lash bearing 5-6 strong spines on convex margin, with 3 seta. Anterior seta specialized, thick, distally ramified into 3 or 4 spiniform processes. Seta on inner margin spinulated all over surface, slightly shorter than terminal lash. Maxilliped (Fig. 97E) 3-segmented. First segment unarmed, as long as second segment. Second segment with 1 tubercle on proximal part of inner surface and 2 small setae, and scattered minute spinules. Third segment tapering, distally forming curved claw, proximally with 2 small setae.

Legs 1 and 2 (Fig. 97F, G) and leg 3 with 3-segmented rami. Two terminal spines on endopod of Cyclopoida: Sabelliphilidae: Myxomolgus 179

F D

Fig. 96. Myxomolgus invulgus, female. A. habitus, dorsal; B. urosome, dorsal; C. anal somite and caudal rami, ventral; D. anterior part of cephalosome, ventral; E. antennule; F. antenna. Scales: A=0.2 mm, B, D=0.1 mm, C=0.02 mm, E, F=0.05 mm. 180 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B C

D E F

Fig. 97. Myxomolgus invulgus, female. A. labrum; B. mandible; C. maxillule; D. maxilla; E. maxilliped; F. leg 1; G. leg 2; H. leg 4; I. leg 5 exopod. Scales: A-E=0.02 mm, F-I=0.05 mm. Cyclopoida: Sabelliphilidae: Myxomolgus 181

A B

Fig. 98. Myxomolgus invulgus, male. A. habitus, dorsal; B. urosome, ventral; C. maxilliped; D. third endopodal segment of leg 2. Scales: A=0.2 mm, B=0.05 mm, C, D=0.02 mm.

Leg 4 (Fig. 97H) 112 μm (inner) and 53 μm (outer). Armature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Legs 2 and 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 4: coxa 0-1; basis 1-0; exp I-0; I-1; II, I, 5; enp. 0-1; 0-1; II

Leg 5 consisting of 1 dorsal seta on fifth pedigerous somite and free exopod. Exopod (Fig. 97I)

70×45 μm (1.56 : 1), slightly widened medially, terminally with 1 denticle, 1 spine (90 μm) and 1 seta (122 μm). Leg 6 represented by 1 spinule and 2 setae in genital aperture.

Male: Body (Fig. 98A) more slender than that of female. Urosome (Fig. 98B) 6-segmented. Fifth pedigerous somite 95 μm wide, distinctly narrower than genital somite. Genital somite expanded, 117 ×150 μm, roughly quadrangular. Lateral margins of abdomen nearly parallel. Four abdominal somites 28×52, 27×47, 17×43, and 32×47 μm, respectively. Caudal ramus 39×18 μm (2.17 : 1). Rostrum as in female. Antennule added by 1 aesthetasc on fourth segment. Asethetascs on second and fourth segments large, longer than whole antennule. Antenna as in female.

Maxilliped (Fig. 98C) 4-segmented. First segment unarmed. Second segment expanded proximally, with 2 setae and rows of spinules on inner margin. Third segment short and unarmed. 182 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Fourth segment with 2 proximal setae and distally forming large claw bearing ramified tip and membrane on concave margin. Other mmouthparts as in female.

Terminal endopod segment of leg 2 (Fig. 98D) sexually dimorphic: terminal process enlarged, and 2 terminal setae transformed respectively to small setule and irregular process. Legs 1, 3-5 as in female. Leg 6 represented by 2 setae and 1 denticle on posterior part of genital operculum.

Distribution: Korea. Korea: CN.

Specimens examined: 22♀♀, 20♂♂ (type specimens) from the mucous wall of the tube of a polychaete of Myxicola sp. in the intertidal sand, at Daechon Beach (36°18′N, 126°31′E), Choongc- heongnam-do, 28.x.1999. Remarks: This species lives in a peculiar habitat, the mucous wall of the polychaete tube. The mucous wall was inhabited by numerous, minute green plants. It is uncertain whether the copepods feed on the mucus material or the green plants.

Genus Nasomolgus Sewell, 1949

Gaet-ji-reong-i-sal-i-sok (갯지렁이살이속)

Rostrum simple. Antenna 4-segmented, with 2 terminal claws. Labrum with long lateral process on both sides. Female maxilliped with elongate, whip-like terminal claw. Leg 4 endopod 2-segmented, with 2 distal setae on distal segment. Other characters as those of the family. Containing 5 known species.

46. Nasomolgus firmus Humes and Ho, 1967 (Figs. 99-101)

Kkeun-kkeun-i-gaet-ji-reong-i-sal-i (끈끈이갯지렁이살이)

Nasomolgus firmus Humes and Ho, 1967, p. 386, figs. 60-86; Humes and Stock, 1973, p. 250, fig. 139; Izawa, 1976, p. 94, figs. 20-42; Kim, 1998, p. 521, figs. 231, 232.

Female: Body (Fig. 99A) with globular prosome and narrow urosome. Body length 1.12 mm. Geatest width 0.49 mm. Prosome 5-segmented. First pedigerous somite distinctly broader than cephalosome. Second to fourth pedigerous somite distinctly narrower than second pedigerous somite. Urosome (Fig. 99B) 5-segmented. Fifth pedigerous somite narrow, 92 μm wide. Genital double-somite 138×131 μm; anterior part strongly expanded laterally, with greatest width at 35% region of somite; genital aperture broad, located posterior region expansion. Free abdominal somites

54×56, 46×51, and 54×50 μm, respectively. Caudal ramus 52×21 μm (2.48 : 1); all caudal setae naked. Egg sac elongate.

Rostrum nearly pentagonal, with tapering posterior half. Antennule (Fig. 99C) 7-segmented; armature formula 4, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; aesthetascs small. An- tenna (Fig. 99D) 4-segmented and stout; armature formula 1, 1, 3, and 4 +3 claws; first segment with 1 claw-like, conical process in middle; second to terminal segments each much shorter than Cyclopoida: Sabelliphilidae: Nasomolgus 183 wide. terminal segment with 3 claws, one of which slender and remaining 2 very thick and subequal in length.

Labrum (Fig. 99E) characteristically with 1 pair of proximo-lateral, setiform processes. Mandible (Fig. 99F) simple and short; convex margin of blade serrate; middle region of blasé with 2 denticles; concave margin with row of spinules. Maxillule (Fig. 99G) with 2 setae and 2 setiform processes.

Maxilla (Fig. 99H) with 3 setae on second segment; distal part of second segment with 8 or 9 teeth on convex margin. Maxilliped (Fig. I) consisting of 2 segments and elongate terminal, lash-like claw; first segment naked; second segment with 2 equal inner setae in middle of inner margin; terminal claw 141 μm long, much longer than combined basal segments, proximally with rudiment of suture and 1 small seta and spinules on both margins of distal haf.

Legs 1-3 (Fig. 100A-C) with 3-segmented rami. Leg 4 (Fig. 100D) with 3-segmented exopod and 2-segmented endopod. Endopod of leg 5 slender; its distal segment 55×11 μm, 5 times as long as wide, its 2 distal setae spiniform, 68 μm long (inner seta) and 43 μm long (outer seta). All spines on legs small. Posterior margin of basis of legs 1-4 with conical process between bases of rami. Ar- mature formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-2; II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; II Leg 5 represented by 1 dorsolateral seta on fifth pedigerous somite and free exopod;. Exopod very small, shorter than long, with 2 distal setae. Leg 6 represented by 2 small setae and 1 denticle in genital aperture (Fig. 99J).

Male: Body (Fig. 101A) distinctly narrower than that of female and 754 μm long. Prosome 427 μm, with greatest width of 237 μm. Dorsal suture line between cephalosome and first pedigerous somite distinct. Urosome (Fig. 101B) 6-segmented. Genital somite 123×95 μm, longer than wide, with rounded anterior and posterior corners. Four abdominal somites 32×44, 32×42, 26×34, and

28×35 μm, respectively. Caudal ramus 31×16 μm (ratio 1.91 : 1). Rostrum, Antennule and antenna as in female. Maxilla similar to that of female, except for lacking of conical process on proximal segment. Maxilliped (Fig. 101C) consisting of 3 segments and terminal claw; first and third segment naked; second segment with 2 similar setae on inner margin and 1 elongate circle of small spinules on inner side; terminal claw large, longer than combined 2 distal segments, gently curved, and tipped with hyaline, semicircular membrane, with 2 proximal, unequal setae (larger 1 spinulose) and numerous denticles along distal two-thirds of concave margin. Other mouthparts as in female. Legs 1-5 as in female. Leg 6 represented by 2 naked setae on genital operculum.

Distribution: Madagascar, Japan, and Korea. Korea: JJ, CN.

Specimens examined: 2♀♀ from the polychaete Sabellastarte spectabilis (Grube), Soheuksan I. (SCUBA), Jeollanam-do, 17.vi.1993; 15♀♀, 1♂ from Sabellastarte sp., Moonseom, Seogwipo, Jeju I., 9.xi.2012, H.S. Hwang.

Remarks: Known hosts of this copepod are Sabellastarte magnifica (Shaw) (type host) in Madagas- car and S. indica in Japan and Korea. 184 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

D G

F H

E J

Fig. 99. Nasomolgus firmus, female. A. habitus, dorsal; B. urosome, dorsal; C. antennule; D. antenna; E. labrum; F. mandible; G. maxillule; H. maxilla; I. maxilliped; J. right genital aperture. Scales: A=0.2 mm, B, C=0.05 mm, D-J=0.02 mm. Cyclopoida: Sabelliphilidae: Nasomolgus 185

A B

D C

Fig. 100. Nasomolgus firmus, female. A. leg 1; B. leg 2; C. leg 3; D. leg 4. Scales: 0.05 mm. 186 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

A B

Fig. 101. Nasomolgus firmus, male. A. habitus, dorsal; B. urosome, ventral; C. maxilliped. Scales: A=0.1 mm, B=0.05 mm, C=0.02 mm.

Family Thamnomolgidae Humes and Boxshall, 1996

Hae-song-but-ge-gwa (해송붙게과)

Body cyclopiform. Urosome 5-segmented in female and 6-segmented in male. Rostrum broad, with truncate posterior apex. Antennule 7-segmented. Antenna 4-segmented; third segment with 3 setae; fourth segment with 1, well-developed terminal claw. Labrum broad median incision. Mandible slender, elongate, unarmed, but with spinules along concave margin. Maxillule with 2 apical setae. Maxilla 2-segmented; distal segment spiniform. Maxilliped 3-segmented in female and 4-segmented in male. Legs 1-3 with 3-segmented rami. Leg 4 with 3-segmented exopod and

2- or 3-segmented endopod. Leg 5 with free exopod bearing 2 setae. Associated with cnidarians (antipatharians amd gorgonaceans). Cyclopoida: Thamnomolgidae: Thamnomolgus 187

Genus Thamnomolgus Humes, 1969

Hae-song-but-ge-sok (해송붙게속)

Characters as those of the family, but leg 4 endopod 2-segmented. Containing 2 species associated with antipatharians.

47. Thamnomolgus robustus Humes, 1969 (Figs. 102-104)

Hae-song-but-ge (해송붙게)

Thamnomolgus robustus Humes, 1969, p. 19, figs. 64-67; Humes and Stock, 1973, p. 81; Humes, 1990, p. 591; Kim, 1998, p. 548, figs. 247-249.

Female: Body (Fig. 102A) stout. Body length 1.39 mm. Greatest width of body 0.59 mm. Pro- some 4-segmented, consisting of cephalothorax and 3 pedigerous somites. Cephalothorax nearly spherical. Metasomites with relatively developed epimera. Urosome (Fig. 102B) 5-segmented. Fifth pedigerous somite 220 μm wide, wider than next somite. Genital double-somite 213×193 μm and laterally expanded. Genital aperture located at midway of expansion. First to third free abdominal somites 43×113, 35×110, and 63×132 μm, respectively. Anal somite with minute spinules along posteroventral margin. Caudal ramus (Fig. 102C) oval, 100×66 μm (1.52 : 1), with short caudal setae, dorsal one of which simple but other 5 setae with inflated proximal part, and 2 median terminal setae fringed with membrane along both margins.

Rostrum (Fig. 102D) broad and distally truncated. Antennule (Fig. 102E) 7-segmented; armature formula 3, 13, 6, 3, 4+aesthetasc, 2+aesthetasc, and 7+aesthetasc; all setae short and naked. An- tenna (Fig. 102F) 4-segmented; first segment naked; second segment with 1 inner seta proximally; third segment with 3 setae at inner distal corner; terminal segment with 1 inner distal seta; termi- mal claw long and slender, as long as terminal segment.

Labrum (Fig. 102D) with broad and shallow median incision. Mandible (Fig. 102G) digitiform, slightly curved, and pectinate along concave margin. Maxillule (Fig. 102H) slender and tapering, with 2 equal, naked setae. Maxilla (Fig. 103A) 2-segmented; basal segment large and naked; distal segment claw-like, slender and elongate, with 2 proximal and row of several denticles on distal half. Maxilliped (Fig. 103B) 3-segmented; first segment short and broad; second segment with 2 inner setae; third segment serrate distally, with small process and 1 large, process-like seta on inner side.

Legs 1-4 (Fig. 103C-F) with 3-segmented rami. Leg 4 endopod very slender and short. Arma- ture formula of legs 1-4 as follows: Leg 1: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 4; enp. 0-1; 0-1; I, 5 Leg 2: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; III, 3 Leg 3: coxa 0-1; basis 1-0; exp. I-0; I-1; III, I, 5; enp. 0-1; 0-2; II, 2 Leg 4: coxa 0-1; basis 1-0; exp. I-0; I-1; II, I, 5; enp. 0-1; 0-1; II

Leg 5 (Fig. 103G) exopod oval 40×22 μm (1.82 : 1), with 2 long, naked setae distally.

Male: Body (Fig. 104A) as that of female in form. Body length 1.24 mm. Urosome (Fig. 104B) 6-segmented. Genital somite nearly rectangular and 205×230 μm, with strongly produced postero- 188 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

C B A

D F

H G

Fig. 102. Thamnomolgus robustus, female. A. habitus, dorsal; B. urosome, dorsal; C. caudal ramus; D. frontal part of cephalosome, ventral; E. antennule; F. antenna; G. mandible; H. maxillule. Scales: A=0.2 mm, B, D=0.1 mm, C, E, F=0.05 mm, G, H=0.02 mm. Cyclopoida: Thamnomolgidae: Thamnomolgus 189

C D B

E F

Fig. 103. Thamnomolgus robustus, female. A. maxilla; B. maxilliped; C. leg 1; D. leg 2; E. leg 3; F. leg 4; G. leg 5. Scales: A, B, G=0.02 mm, C-F=0.05 mm. 190 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

E D

Fig. 104. Thamnomolgus robustus, male. A. habitus, dorsal; B. urosome, dorsal; C. maxilliped; D. leg 1; E. leg 1 endopod. Scales: A=0.2 mm, B=0.1 mm, C, D=0.05 mm, E=0.02 mm.

ventral corers. Abdominal somites broadened from anterior to posterior. First to third abdominal somites short. Caudal ramus 94×72 μm (1.31 : 1). Antennule with 2 additional aesthatascs on second segment and 1 additional aesthetasc on fourth segment. Antenna as in female. Cyclopoida: Thamnomolgidae: Thamnomolgus 191

Maxilliped (Fig. 104C) consisting of 3 segments and terminal claw; first segment narrow and naked; second segment with broad outer expansion, 2 unequal, inner setae, row of denticles on proximal two-thirds of inner margin, and patch of minute spinules at inner distal region; third segment short and unarmed; terminal claw rather stout, as long as combined 2 distal segments, proximally with 1 ovoid process, 1 seta and 1 setule. Other mouthparts as in female.

Legs 1-4 with same armature formula as that of female, but leg 1 endopod (Fig. 104D, E) sexually dimorphic in having distal part third segment highly sclerotized and covered dark material, and spinulose distal tubercle; outer spine on this segment slender and elongated; outer distal seta enlarged. Leg 5 exopod small, immovable, almost fused with somite, with suture line at bese. Leg 6 represented by 2 setae on posterior margin of genital operculum.

Distribution: Madagascar, the Philippines, Korea. Korea: JJ. Specimens examined: All specimens were collected from the external surface of Antipathes japonica Brook. 40♀♀, 25♂♂, Moonseom (SCUBA), Seogwipo, Jeju I., 14.xi.1997, J.G. Je; 15♀♀,

12♂♂, near Biyang-do I. (SCUBA), Jeju-do, 19.iv.2003; 31♀♀, 15♂♂, off Sagye-ri (SCUBA), An- deok-myeon, Jeju-do, 22.ix.2009, M.-K. Choe; 32♀♀, 22♂♂, off Sagye-ri (SCUBA), Andeok-myeon, Jeju-do, 22.ix.2009, M.-K. Choe. Remarks: This species is distinguishable from its only congener, T. nodulus, in the ratio of the length to width of caudal ramus about 1.5 : 1 in the female (vs. 2.1 : 1 in T. nodulus; see Humes, 1990) and the ratio of the length to width of leg 5 exopod of the female about 1.8 : 1 (vs. 2.48 : 1 in T. nodulus; see Humes, 1990). 192 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Literature Cited

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area in Tokyo Bay, Japan. Hydrobiologia, 254: 149-157. Itoh, H. and S. Nishida, 1998. A new species of Hemicyclops (Copepoda, Poecilostomatoida) rom burrows of the ocypodid crab Macrophthalmus japonicus in an estuarine mud-flat in Tokyo Bay, Japan. Hydrobiolo- gia, 379: 85-92. Itoh, H. and S. Nishida, 2002. A new species of Hemicyclops (Copepoda, Poecilostomatoida) from burrows of the mud shrimp Upogebia major in an estuarine mud-flat in Tokyo Bay, Japan. Hydrobiologia, 474: 139- 146. Itoh, H. and S. Nishida, 2007. Life history of the copepod Hemicyclops gomsoensis (Poecilostomatoida, Clausi- diidae) associated with decapod burrows in the Tama-River estuary, central Japan. Plankton Benthos Res., 2(3): 134-146. Itoh, H. and S. Nishida, 2008. Life history of the copepod Hemicyclops spinulosus (Poecilostomatoida, Clausi- diidae) associated with crab burrows with notes on male polymorphism and precopulatory mate guarding. Plankton Benthos Res., 3(4): 189-201. Izawa, K., 1976. Two semi-parasitic copepods of marine invertebrates from Japan (Cyclopoida: Lichomolgi- dae). Publ. Seta Mar. Biol. Lab., 23(1/2): 89-98. Kim, I.-H., 1996. Three new species of Critomolgus (Copepoda, Poecilostomatoida, Rhynchomolgidae) associated with sea anemons in Korea. Korean J. Zool., 39: 362-377. Kim, I.-H., 1998. Illustrated Encyclopedia of Fauna & Flora of Korea. Vol. 38. Cirripedia, Symbiotic Copepo- da, Pycnogonida. Ministry of Education, Korea, 1038 pp. Kim, I.-H., 2000a. Poecilostomatoid copepods from an intertidal mud flat in the Yellow Sea. J. Nat. Hist., 34(3): 367-432. Kim, I.-H., 2000b. Zamolgus cavernularius n. sp. (Copepoda, Poecilostomatoida, Rhynchomolgidae) associated with a pennatulacean in the Yellow Sea. Korean J. Biol. Sci., 4(3): 251-255. Kim, I,-H., 2001a. A new genus and two new species of Copepoda (Poecilostomatoida: Sabelliphilidae) associated with the tubicolous polychaetes in the Yellow Sea. Korean J. Biol. Sci., 5(1): 1-9. Kim, I.-H., 2001b. A new species of Clausia (Copepoda, Poecilostomatoida, Causiidae) associated with the polychaete Arenicola brasilliensis Nonata in Korea. Hydrobiologia, 452(1/3): 217-223. Kim, I.-H., 2001c. Foliomolgus cucullus, a new genus and species of Clausidiidae (Copepoda, Poecilostomatoi- da) associated with a polychaete in Korea. Proc. Biol. Soc. Wash., 114(3): 660-666. Kim, I.-H., 2006a. Eupolymniphilus orientalis n. sp. (Copepoda, Cyclopoida, Sabelliphilidae) from intertidal burrows in Korea, with the recognition of four species in the genus. Crustaceana, 79(5): 573-582. Kim, I.-H., 2006b. Three new species of Kelleria (Copepoda: Poecilostomatoida: Kelleridae) from Korea. Kore- an J. Syst. Zool., 22(1): 1-11. Kim, I.-H., 2006c. Poecilostomatoid copepods (Rhynchomolgidae) associated with sea anemons (Actiniaria) from Korea. Integrative Biosciences, 10(3): 145-161. Kim, I.-H., 2014. Six new species of Copepoda (Clausiidae, Pseudanthessiidae, Polyankyliidae) associated with polychaetes from Korea. Journal of Species Research, 3(2): 95-122. Moon, S.Y. and I.-H. Kim, 2010. Three new species of Hemicyclops (Copepoda, Cyclopoida, Clausidiidae) from Korea. Korean J. Syst. Zool., 26(3): 279-293. WoRMS, 2014. Accessed through: World Register of Marine Species at http://www.marinespecies.org/ aphia.php?p=taxdetails&id=1066 on 2014-11-10. 194 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Index to Korean Names

ㄱ 두마디인도검물벼룩 69 떠돌이검물벼룩붙이 25 갈색말미잘속살이 135 떠돌이구멍벌레 95 개펄말미잘이 154 개펄말미잘이속 146 갯지렁이살이과 174 ㅁ 갯지렁이집벌레 178 갯지렁이집벌레속 178 막검물벼룩붙이 29 검물벼룩목 9 말미잘이속 130 검물벼룩붙이과 9 무족발톱검물벼룩 121 검물벼룩붙이속 15 무족발톱검물벼룩속 120 곤봉딸기산호속살이 98 민다리검물벼룩과 126 곰소검물벼룩붙이 20 민다리검물벼룩속 126 구멍벌레과 88 민다리뿔산호벌레 157 구멍벌레속 88 긴배검물벼룩과 57 긴배검물벼룩속 64 ㅂ 꽃갯지렁이벌레 83 꽃갯지렁이벌레속 83 바다조름이 143 끈끈이갯지렁이살이 182 발톱검물벼룩과 106 갯지렁이살이속 182 발톱검물벼룩속 106 빗검물벼룩붙이 16 빨판검물벼룩 77 ㄴ 빨판검물벼룩속 77 뿔검물벼룩 74 남해검물벼룩 12 뿔검물벼룩속 73 남해검물벼룩속 12 뿔산호벌레속 157 납작검물벼룩붙이 55 넓적배검물벼룩 81 넓적배검물벼룩속 81 ㅅ 녹색말미잘속살이 131 녹색말미잘이 139 산호속살이과 98 산호이과 130 세마디검물벼룩 58 ㄷ 실참갯지렁이벌레 45 쏙벌레 37 돌검물벼룩붙이 42 쏙붙이벌레 51 돌기민다리검물벼룩 126 동양발톱검물벼룩 107 동양뿔산호벌레 161 ㅇ 동양오가시갯지렁이살이 175 두마디검물벼룩 61 연산호속살이속 98 Index to Korean Names 195

열한가시구멍벌레 93 큰배검물벼룩과 83 오가시갯지렁이살이속 175 큰손벌레과 100 왕가슴말미잘이 165 큰손벌레속 100 왕가슴말미잘이속 165 큰털구멍벌레 89 왜개펄말미잘이 151 유령갯지렁이벌레 116 유령갯지렁이벌레속 116 ㅌ 인도검물벼룩속 69 잎검물벼룩 9 턱돌기벌레 170 잎검물벼룩속 9 턱돌기벌레속 170 털큰손벌레 101

ㅈ ㅎ 작은긴배검물벼룩 64 작은발톱검물벼룩 111 항구구멍벌레 91 장다리검물벼룩속 58 해송붙게 187 해송붙게과 186 해송붙게속 187 ㅋ 헛큰손벌레 103 헛큰손벌레속 103 코검물벼룩붙이 34 큰개펄말미잘이 146 196 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Index to Korean Name as Pronounced

B Hae-song-but-ge-gwa 186 Hae-song-but-ge-sok 187 Ba-da-jo-reum-i 143 Hang-gu-gu-meong-beol-re 91 Bal-top-geom-mul-byeo-ruk-gwa 106 Heot-keun-son-beol-re 103 Bal-top-geom-mul-byeo-ruk-sok 106 Heot-keun-son-beol-re-sok 103 Bit-geom-mul-byeo-ruk-but-i 16

I D In-do-geom-mul-byeo-ruk-sok 69 Dol-geom-mul-byeo-ruk-but-i 42 Ip-geom-mul-byeo-ruk 9 Dol-gi-min-da-ri-geom-mul-byeo-ruk 126 Ip-geom-mul-byeo-ruk-sok 9 Dong-yang-bal-top-geom-mul-byeo-ruk 107 Dong-yang-o-ga-si-gaet-ji-reong-i-sal-i 175 Dong-yang-ppul-san-ho-beol-re 161 J Du-ma-di-geom-mul-byeo-ruk 61 Du-ma-di-in-do-geom-mul-byeo-ruk 69 Jag-eun-gin-bae-geom-mul-byeo-ruk 64 Jag-eun-bal-top-geom-mul-byeo-ruk 111 Jang-da-ri-geom-mul-byeo-ruk-sok 58 G

Gae-peol-mal-mi-jal-i 154 K Gae-peol-mal-mi-jal-i-sok 146 Gaet-ji-reong-i-jip-beol-re 178 Keun-bae-geom-mul-byeo-ruk-gwa 83 Gaet-ji-reong-i-jip-beol-re-sok 178 Keun-gae-peol-mal-mi-jal-i 146 Gaet-ji-reong-i-sal-i-gwa 174 Keun-son-beol-re-gwa 100 Gaet-ji-reong-i-sal-i-sok 182 Keun-son-beol-re-sok 100 Gal-saek-mal-mi-jal-sok-sal-i 135 Keun-teol-gu-meong-beol-re 89 Geom-mul-byeo-ruk-but-i-gwa 9 Kkeun-kkeun-i-gaet-ji-reong-i-sal-i 182 Geom-mul-byeo-ruk-but-i-sok 15 Kkot-gaet-ji-reong-i-beol-re 83 Geom-mul-byeo-ruk-mok 9 Kkot-gaet-ji-reong-i-beol-re-sok 83 Gin-bae-geom-mul-byeo-ruk-gwa 57 Ko-geom-mul-byeo-ruk-but-i 34 Gin-bae-geom-mul-byeo-ruk-sok 64 Gom-so-geom-mul-byeo-ruk-but-i 20 Gon-bong-ttal-gi-san-ho-sok-sal-i 98 M Gu-meong-beol-re-gwa 88 Gu-meong-beol-re-sok 88 Mak-geom-mul-byeo-ruk-but-i 29 Mal-mi-jal-i-sok 130 Min-da-ri-geom-mul-byeo-ruk-gwa 126 H Min-da-ri-geom-mul-byeo-ruk-sok 126 Min-da-ri-ppul-san-ho-beol-re 157 Hae-song-but-ge 187 Mu-jok-bal-top-geom-mul-byeo-ruk 121 Index to Korean Name as Pronounced 197

Mu-jok-bal-top-geom-mul-byeo-ruk-sok 120 San-ho-sok-sal-i-gwa 98 Se-ma-di-geom-mul-byeo-ruk 58 Sil-cham-gaet-ji-reong-i-beol-re 45 N Ssok-beol-re 37 Ssok-but-i-beol-re 51 Nam-hae-geom-mul-byeo-ruk 12 Nam-hae-geom-mul-byeo-ruk-sok 12 Nap-jak-geom-mul-byeo-ruk-but-i 55 T Neolb-jeok-bae-geom-mul-byeo-ruk 81 Neolb-jeok-bae-geom-mul-byeo-ruk-sok 81 Teol-keun-son-beol-re 101 Nok-saek-mal-mi-jal-i 139 Tteo-dol-i-geom-mul-byeo-ruk-but-i 25 Nok-saek-mal-mi-jal-sok-sal-i 131 Tteo-dol-i-gu-meong-beol-re 95 Tteok-dol-gi-beol-re 170 Tteok-dol-gi-beol-re-sok 170 O

O-ga-si-gaet-ji-reong-i-sal-i-sok 175 W

Wae-gae-peol-mal-mi-jal-i 151 P Wang-ga-seum-mal-mi-jal-i 165 Wang-ga-seum-mal-mi-jal-i-sok 165 Ppal-pan-geom-mul-byeo-ruk 77 Ppal-pan-geom-mul-byeo-ruk-sok 77 Ppul-geom-mul-byeo-ruk 74 Y Ppul-geom-mul-byeo-ruk-sok 73 Ppul-san-ho-beol-re-sok 157 Yeol-han-ga-si-gu-meong-beol-re 93 Yeon-san-ho-sok-sal-i-sok 98 Yu-ryeong-gaet-ji-reong-i-beol-re 116 S Yu-ryeong-gaet-ji-reong-i-beol-re-sok 116

San-ho-i-gwa 130 198 Invertebrate Fauna of Korea·Cyclopoid Copepods Associated with Cnidarians, Polychaetes and Crustaceans

Index to Scientific Names

C nasutus 34 parilis 37 Clausia 64 saxatilis 42 parva 64 spinulosus 45 Clausidiidae 9 tanakai 51 Clausiidae 57 ventriplanus 55 Critomolgus 130 anthopleurus 131 malmizalus 135 I nudus 143 vicinus 139 Indoclausia 69 Cyclopoida 9 bipartita 69

E K

Enalcyonium 98 Kelleria 88 digitigerum 98 grandisetiger 89 Eupolymniphilus 175 portiviva 91 orientalis 175 undecidentata 93 vaga 95 Kelleridae 88 F

Foliomolgus 9 L cucullus 9 Lamippidae 98 Likroclausia 73 G namhaensis 74 Lutumidomus 146 Gastrodelphyidae 83 panikkari 146 parvus 151 tertius 154 H

Hemadona 12 M clavicrura 12 Hemicyclops 15 Macrochiron 100 ctenidis 16 anomalum 101 gomsoensis 20 Macrochironidae 100 japonicus 25 Maxilliclausia 77 membranatus 29 propria 77 Index to Scientific Names 199

Myxomolgus 178 laticauda 81 invulgus 178 Rhynchomolgidae 130

N S

Nasomolgus 182 Sabellacheres 83 firmus 182 illgi 83 Sabelliphilidae 174 Sewelloya 120 P plana 121

Paramolgus 157 nudipes 157 T orientalis 161 Polyankyliidae 106 Terebelliphilus 116 Polyankylis 106 simplex 116 orientalis 107 Thamnomolgidae 186 ovilaxa 111 Thamnomolgus 187 Pontoclausia 58 robustus 187 antiqua 58 lobata 61 Pseudanthessiidae 126 V Pseudanthessius 126 excertus 126 Verutipes 165 Pseudomacrochiron 103 scutatus 165 urostenum 103

Z R Zamolgus 170 Rhodinicola 81 cavernularius 170 Number 2

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