Parasitic Copepods of Mackerel - and Tuna-Like Fishes (Scombridae) of the World

Parasitic Copepods of Mackerel - and Tuna-like Fishes (Scombridae) of the World

ROGER CRESSEY and HILLARY BOYLE CRESSEY m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 311 SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION

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S. Dillon Ripley Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 311

Parasitic Copepods of Mackerel and Tuna-like Fishes (Scombridae) of the World

Roger Cressey and Hillary Boyle Cressey

SMITHSONIAN INSTITUTION PRESS City of Washington 1980 ABSTRACT

Cressey, Roger, and Hillary Boyle Cressey. Parasitic Copepods of Mackerel- and Tuna-like Fishes (Scombridae] of the World. Smithsonian Contributions to Zoology, number 311, 186 pages, 139 figures, 1980.—Forty-six species of parasitic copepods (one new genus, 8 new species) are reported from all known 46 species of Scombridae subfamily Scombrinae from the world. Literature records are included only where host and/or copepod identifications could be verified. Copepod families include: Bomolochidae; Shiinoidae; Caligidae; Euryphoridae; Tuxophoridae; Pseudocycnidae; Lernanthropidae; Lerneopodidae. The new bomolochid genus is Unicolax. The new species are U. collaterals, U. ciliatus, U. reductus, Holobomolochus divaricatus, H. nudiusculus, and H. asperatus (Bomolochidae) ; Caligus omissus (Caligidae) ; and Tuxophorus collettei (Tuxophoridae). The genus Pseudocynopsis Yamaguti has been placed in synonymy with Pseudocycnoides Yamaguti. Lists are included of all species of the Scombrinae and their geographic distribution, copepods parasitic on each scombrid species with the relative abundance of each copepod species on each host species, and the scombrid hosts for each copepod species indicating host preferences.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN : The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data Cressey Roger F 1930- Parasitic copepods of mackerel- and tuna-like fishes (Scombridae) of the world. (Smithsonian contributions to zoology ; no. 311) Bibliography: p. Supt. of Docs, no.: SI 1.27:311 1. Copepoda. 2. Parasites—Fishes. 3. Scombridae—Diseases. 4. Fishes—Diseases. I. Cres- sey, Hillary Boyle, 1949- joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology ; no. 311. QL1.S54 no. 311 [QL444.C7] 591'.08s [595'.34'04524] 79-20297 Contents

Page Introduction 1 Known Host Species with Distribution 2 Specimens Examined 3 Acknowledgments 3 Holobomolochus Vervoort, 1969 4 Holobomolochus divaricatus, new species 4 Holobomolochus nudiusculus, new species 6 Holobomolochus asperatus, new species 7 Discussion 8 Unicolax, new genus 8 Unicolax collateralis, new species 8 Unicolax anonymous (Vervoort, 1965), new combination 11 Unicolax mycterobius (Vervoort, 1965), new combination 13 Unicolax ciliatus, new species 15 Unicolax reductus, new species 17 Ceratacolax Vervoort, 1965 18 Ceratacolax euthynni Vervoort, 1965 18 Nothobomolochus Vervoort, 1962 19 Nothobomolochus kanagurta (Pillai, 1965), new combination 19 Orbitacolax Shen, 1957 20 Orbitacolax aculeatus (Pillai, 1962), new combination 20 Pumiliopes Shen, 1957 20 Pumiliopes capitulatus Cressey and Boyle, 1973 20 Shiinoa Kabata, 1968 21 Shiinoa inauris Cressey, 1975 21 Shiinoa occlusa Kabata, 1968 22 Caligus Miiller, 1785 22 Caligus coryphaenae Steenstrup and Liitken, 1861 22 Caligus regalis Leigh-Sharpe, 1930 23 Caligus productus Dana, 1852 24 Caligus asymmetricus Kabata, 1965 25 Caligus bonito Wilson, 1905 26 Caligus mutabilis Wilson, 1905 27 Caligus omissus, new species 28 Caligus biseriodentatus Shen, 1957 29 Discussion 31 Caligus cybii Bassett-Smith, 1898 31 Caligus pelamydis Kroyer, 1863 32 Caligus infestans Heller, 1868 33 Caligus diaphanus Nordmann, 1832 34

iii iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page Caligus savala Gnanamuthu, 1948 35 Caligus macarovi Gussev, 1951 35 Caligus amblygenitalis Pillai, 1961 36 Caligus pseudokalumai Lewis, 1968 36 Elytrophora Gerstaecker, 1853 36 Elytrophora brachyptera Gerstaecker, 1853 36 Elytrophora indica Shiino, 1958 37 Gloiopotes Steenstrup and Liitken, 1861 38 Gloiopotes hygomianus Steenstrup and Liitken, 1861 38 Caligulus Heegaard, 1962 38 Caligulus longispinosus Heegaard, 1962 38 Tuxophorus Wilson, 1908 39 Tuxophorus cybii Nunes-Ruivo and Fourmanoir, 1956 39 Tuxophorus cervicornis Heegaard, 1962 39 Tuxophorus collettei, new species 39 Pseudocycnus Heller, 1868 41 Pseudocycnus appendiculatus Heller, 1968 41 Pseudocycnoides Yamaguti, 1963 42 Pseudocycnoides armatus (Bassett-Smith, 1898) 42 Pseudocycnoides scomberomori (Yamaguti, 1939) 43 Pseudocycnoides buccata (Wilson, 1922), new combination 43 Lernanthropus Blainville, 1822 44 Lernanthropus kanagurta Tripathi, 1962 44 Brachiella Cuvier, 1830 45 Brachiella thynni Cuvier, 1830 45 Brachiella magna Kabata, 1968 45 Clavellisa Wilson, 1915 45 Clavellisa scombri (Kurz, 1877) 45 Clavellopsis Wilson, 1915 45 Clavellopsis saba Yamaguti, 1939 45 Pennella Oken, 1816 46 Pennella species 46 Summary 46 Literature Cited 51 Figures 54 Parasitic Copepods of Mackerel- and Tuna-like Fishes (Scombridae) of the World

Roger Cressey and Hillary Boyle Cressey

Introduction here, resolving problems of synonymy with several copepod species, and working closely with scombrid taxonomists (R. H. Gibbs and B. B. Collette) to assure There are 47 currently recognized species of the proper host identifications, we feel we have reached family Scombridae; plus an additional new species of that objective. We do not feel that we have produced Scomberomorus being described by Collette and Russo. a definitive work, but hopefully, a solid foundation to One species, Gasterochisma melampus Richardson, is which information in future collections can be added. placed in a separate subfamily and has not been con- Previous works by Lewis, Kabata, Shiino, and Pillai sidered in this study. For approximately the last 20 concerned with scombrid copepods are excellent and years the first author has been collecting and receiving we have included some of their information where it collections of the copepods parasitic on these fishes. fills gaps in our own collection data. For reasons stated Of the 46 species of parasitic copepods reported here, above, however, we have not considered most of the 40 species are represented in our collections. These earlier works in our host-parasite data. 40 species have been collected from 2422 individual Because most species of scombrid fishes are commer- fish from all 46 species of Scombrinae. As a result of cially important, various aspects of their biology have this exhaustive collection we have presented a com- been investigated for several years. One result of this prehensive account of the copepods parasitic on tunas interest has been the many reports of copepods para- and tuna-related fishes. sitic on them since the last century. The major failing Our primary objective has been to produce a base- in this, however, has been the general lack of coopera- line study of the copepods parasitic on scombrid fishes. tive effort between parasitologists and ichthyologists. Because of the unreliability of most past literature it Too often, the copepod worker has been content to has been impossible to assess the host-parasite relation- repeat unverified host identifications or unreliable ship, especially the degree of host specificity. By com- common names. This casual approach to that aspect piling a comprehensive collection such as we report of the record has resulted in an account too confusing to be of much value in determining true relationships Roger Cressey and Hillary Boyle Cressey, Department of between the copepod species and their hosts. Except Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. for a literature compilation by Silas and Ummerkutty SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY in 1967, no attempt has been made to critically Guinea being described by Collette and Russo. The examine the scombrid parasites in a comprehensive following list is adapted from information in Collette way to determine the taxonomic status of most of the andChao (1975) and Klawe (1977). described species. For example, Silas and Ummerkutty list 29 species of Caligus as reported parasites of the Tribe SCOMBRINI Bonaparte (Mackerels) Scombrinae. As a result of our examinations of type- specimens of a number of these Caligus species, and Rastrelliger brachysoma (Bleeker) : South China Sea east the data based on our own collections, we have been to Fiji able to place almost half of these in synonymy and Rastrelliger faughni Matsui: South China Sea to Indonesia Rastrelliger kanagurta (Cuvier): Indian and South Central reduce the number of Caligus species from these hosts Pacific oceans to 15. Scomber australasicus (Cuvier) : Western & Central Pacific We have included literature records in our data and Socorro Island (off Mexico) only in those cases where we could verify the host Scomber japonicus Houttuyn: circumglobal antitropical and/or the copepod identification. To include unveri- Scomber scombrus Linnaeus: North Atlantic Ocean and Mediterranean fied collections would serve no useful purpose and only cloud the host-parasite relationship. It was with this in mind that we waited until our collections were as Tribe SCOMBEROMORINI Starks (Seerfishes) complete as reasonably possible to publish our results. We have, with each species, given a skeleton synonymy Acanthocybium solandri (Cuvier): circutntropical and sub- tropical to provide a brief historical account of each. A com- Granimatorcynus bicarinatus (Quoy and Gaimard): Red Sea plete synopsis of the species of Caligus was recently and Western Pacific Ocean published by Margolis, Kabata, and Parker (1975) Scomberomorus brasiliensis Collette, Russo, and Zavala- but a complete revision of the genus is badly needed. Camin: Western Atlantic (Yucatan to south Brazil) There are slightly over 200 currently recognized spe- Scomberomorus cavalla (Cuvier): Tropical western Atlantic cies of Caligus. Reducing the 29 species cited by Silas Ocean and Ummerkutty to 15 illustrates the extent of the Scomberomorus commerson (Lacepede): Indian and West- ern Pacific oceans problem. Extrapolating from that reduction of 29 Scomberomorus concolor (Lockington): Gulf of California species to 15, it would not be unreasonable to expect and Monterey Bay that a generic revision might result in only 100 or so Scomberomorus guttatus (Bloch and Schneider): Indian and Caligus species as valid. western Pacific oceans The first author and B. B. Collette are planning a Scomberomorus koreanus (Kishinouye): Indian and western Pacific oceans joint paper summarizing the host-parasite relationship Scomberomorus lineolatus (Cuvier): India to Indonesia from both ecological and evolutionary standpoints. Scomberomorus maculatus (Mitchill): Western Atlantic We feel that this aspect can best be done separately (Yucatan to Massachusetts) and in conjunction with a scombrid specialist Scomberomorus multiradiatus Munro: Gulf of Papua (Collette). In addition to collections made by us, Scomberomorus niphonius (Cuvier): Northwestern Pacific Ocean (China to Japan) most of the other collections (about one-third of the Scomberomorus plurilineatus Fourmanoir: Mozambique total) were made by Collette in the course of his Channel (coastal) and South Africa studies on scombrids. Scomberomorus queenslandicus Munro: Northern Australia KNOWN HOST SPECIES WITH DISTRIBUTION.—Be- Scomberomorus regalis (Bloch): Western Atlantic (espe- low is a list of the known species of the scombrid sub- cially West Indies) Scomberomorus semifasciatus (Macleay): Northern Aus- family Scombrinae with their general geographic dis- tralia to New Guinea tributions. Although there are 2 recognized species of Scomberomorus sierra Jordan and Starks: Eastern Pacific Auxis, we were advised (B. B. Collette, pers. com.) Ocean that, for purposes of this study, we should refer to Scomberomorus sinensis (Lacepede): Northwestern Pacific Ocean them as "Auxis species" until the genus has undergone Scomberomorus tritor (Cuvier): Gulf of Guinea to Medi- revision. References to "Scomberomorus species" per- terranean tain to a new species from Northern Australia to New Scomberomorus species: Northern Australia to New Guinea NUMBER 3 1 1

Tribe SARDINI Jordan and Evermann (Bonitos) Museo di Storia Naturale, Genoa Museo de La Specola, Universita di Firenze, Florence Allothunnus fallai Serventy: circumglobal in southern hemi- Naturhistorisches Museum, Vienna sphere National Museum of Natural Sciences, Ottawa Cybiosarda elegans (Whitley): Northern Australia to New Northwest Fisheries Center Auke Bay Laboratory, National Guinea Marine Fisheries Service, NOAA, Auke Bay, Alaska Gymnosarda unicolor (Riippell) : Indian to central Pacific Queensland Museum, Brisbane oceans Rijksmuseum van Naturlijke Histoire, Leiden Orcynopsis unicolor (Geoffroy St. Hilaire): Gulf of Guinea J.L.B. Smith Institute of Ichthyology, Rhodes University, to Mediterraean Grahamstown Sarda australis (Macleay): southeast coast of Australia South African Museum, Cape Town Sarda chiliensis (Cuvier), 2 subspecies: Northeastern (S. c. Sea Fisheries Research Station, Haifa, Israel lineolata) and southeastern Pacific Ocean (S. c. chilien- Scripps Institution of Oceanography, La Jolla, Calif. sis) Senckenberg Museum, Frankfurt-am-Main Sarda orientalis (Temminck and Schlegel): Indian and Southeast Fisheries Center, National Marine Fisheries Serv- Pacific oceans (coastal) ice, NOAA (formerly Tropical Atlantic Biological Labo- Sarda sarda (Bloch): Atlantic Ocean and Mediterranean ratory), Miami, Fla. Institute of Fisheries, University of British Columbia, Van- couver Tribe THUNNINI Starks (Tunas) University of California, Los Angeles Rosenstiel School of Marine and Atmospheric Science, Auxis rochei (Risso) : circumtropical Miami, Fla. Auxis thazard Lacepede: circumtropical University of Michigan Museum of Zoology, Ann Arbor Euthynnus affinis (Cantor): Indian and Pacific oceans Smithsonian Institution, Washington, D.C. (USNM, former Euthynnus alletteratus (Rafinesque): Atlantic Ocean United States National Museum, collections in the Na- Euthynnus lineatus Kishinouye: Eastern Pacific Ocean tional Museum of Natural History, Smithsonian Insti- Katsuwonus pelamis (Linnaeus): circumtropical tution) Thunnus alalunga (Bonnaterre): circumtemperate and Western Australia Museum, Perth tropical Woods Hole Oceanographic Institution, Woods Hole, Mass. Thunnus albacares (Bonnaterre): circumsubtropical and Zoological Museum, Copenhagen tropical Zoological Museum, Oslo Thunnus atlanticus (Lesson): Western Atlantic Ocean Thunnus maccoyii (Castelnau): circumglobal in southern Representative collections of the following species hemisphere Thunnus obesus (Lowe): circumsubtropical and tropical will be deposited in the British Museum (Natural His- Thunnus thynnus (Linnaeus): North Pacific, North and tory, London), Museum National d'Histoire Naturelle South Atlantic (non-tropical) (Paris), Australian Museum (Sydney), and California Thunnus tonggol (Bleeker): Indian and Western Pacific Academy of Sciences (San Francisco): Holobomolo- chus divaricatus, H. nudiusculus, Unicolax collateral!*, SPECIMENS EXAMINED.—Scombrids housed in or U. reductus, U. ciliatus, Ceratacolax euthynni, Shiinoa borrowed from the following places were examined inauris, S. occlusa, Caligus coryphaenae, C. omissus, for copepods by us or Collette. C. bonito, C. productus, C. cybii, Elytrophora bra- chyptera, E. indica, Pseudocycnus appendiculatus, Australian Museum, Sydney (AMS) Pseudocycnoides armata, and P. buccata. All type Academy of Natural Sciences, Philadelphia, Pa. material and remaining collections are deposited in British Museum (Natural History), London Bernice P. Bishop Museum, Honolulu, Hawaii the Smithsonian Institution. California Academy of Sciences, San Francisco ACKNOWLEDGMENTS.—This work began with a Chesapeake Biological Laboratory, Solomons, Md. small collection of copepods parasitic on western CSIRO Marine Biological Laboratory, Cronulla, N.S.W., Atlantic pelagic fishes given to one of us (R. Cressey) Australia Dominion Museum, Wellington, New Zealand by Arthur Humes as a basis for a graduate research Field Museum of Natural History, Chicago, 111. project in 1960. The initial collection was made by Hebrew University, Jerusalem Robert Gibbs and Bruce Collette during various Los Angeles County Museum of Natural History, Los cruises of the R. V. Delaware, consisting primarily of Angeles, Calif. copepods from sharks and scombrids. The shark para- Museo Argentina de Ciencias Naturales, Buenos Aires Museum of Comparative Zoology, Harvard (MCZ) sites formed the basis for R. Cressey's doctoral dis- Muslum National d'Histoire Naturelle, Paris sertation revising the family Pandaridae (under the SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY direction of A. Humes). The remaining collection Georgia, Florida, Mississippi, Alabama, Louisiana, stimulated a continuing interest in scombrid copepods Texas, Panama (Atlantic), Surinam; 11 collections by the first author and later by the second. containing 28 9 20 $ and 3 immatures from S. regalis We thank the scientists and staffs of the many from Florida, Cuba, Haiti, Puerto Rico, Colombia institutions cited above for making fish specimens (Atlantic), Venezuela and Surinam. All copepods available for examination for parasitic copepods. were collected from the nasal sinuses of the host fish. Through the years many people have sent us mate- FEMALE.—Body form as in Figure la. Total length rial, for which we are grateful. Outstanding among 2.62 mm, greatest width (widest part of cephalon) these are: Bruce B. Collette (National Marine Fish- 1.20 mm. Genital segment (Figure \b) wider than eries Service, Washington), Robert H. Gibbs (Smith- long (348 X 483 jxm) with 3 dorsal setae at area of sonian Institution, Washington), Arthur G. Humes egg sac attachment. Abdomen 3-segmented, segments (Boston University), Richard Shomora (National measuring (1 X w) 224 X 318 /*m, 153 X 265 ^m, Marine Fisheries Service, Honolulu), and Vladimir 147 X 230 /*m respectively; ventral surface of last Walters (National Marine Fisheries Service, Miami). segment with 2 patches of prominent spinules (see Robert Gibbs was our principle consultant concern- figure \c). Caudal rami (Figure \c) longer than wide ing problems in scombrid taxonomy during the early (129 X 94 /*m) with six setae; longest seta 590 /*m. years of the work. Later, Bruce Collette began an First antenna (Figure Id) 5-segmented, second seg- intensive study of scombrid taxonomy and during the ment incompletely divided; first two segments with 15 course of his work spent many hours gathering cope- prominent plumose setae; segments 4 and 5 with one pods for us. We extend a special thanks to him for aesthete each. Second antenna (Figure 2a) of usual his enthusiastic and continued support bomolochid type; terminal segment with numerous We were assisted in the illustrations by Nancy Zacks rows of spinules, 2 processes with spinules, 3 terminal who rendered Figures 75-79 and 95 and by Michelle setae and 4 elongate spines. Mandible, paragnath, Wilcox who did the illustrations for Figures 82-84. first maxilla and second maxilla as in Figure 2b. Cynthia Hemmings assisted in the final preparation Maxilliped (Figure 2c) terminal segment with promi- of the manuscript. The staff of the SEM laboratory nent curved claw with slight projection on outer curve, all participated at various times in the preparation of and 3 plumose setae; basal segment with 1 plumose the SEM photographs (Walter Brown, Mary Jacque seta. Mann, and Susann Braden). We thank all of them Leg 1 (Figure 2d) exopod with stout, heavily plu- for their assistance. mose seta on outer corner; basipod with 2 patches of The manuscript was reviewed by Bruce Collette and hairs; exopod 2-segmented, first segment with promi- Brian Kensley and we are grateful for their many nent rugose spine on outer distal corner, second seg- helpful suggestions. ment with small spine at outer proximal comer, minute spine at outer distal corner and 2 stout flagel- Holobomolochus Vervoort, 1969 lated spines best viewed dorsally, terminal to inner edge of segment with 6 plumose setae; endopod 3-segmented, first segment with large patch of hairs Holobomolochus divaricatus, new species and inner seta, second segment with smaller patch of hairs and inner seta, third segment with 5 terminal FIGURES 1-4, ba-e, 96, 109-113 setae, all setae heavily plumose, outer edge of endopod MATERIAL EXAMINED. — Holotype 9 (USNM segments haired. Legs 2-4 biramous, rami 3-segmented. 172244), allotype $ (USNM 172245) paratypes 11 9 Leg 2 (Figure 2e) coxopod with short row of spinules (USNM 172246) from the nasal sinuses of Scorn- at outer distal corner; basipod with dorsal seta; exopod beromorus brasiliensis (USNM 188424) from Brazil. first segment with hairs on outer edge and spine on In addition, there are 13 collections containing 61 $ outer distal corner, second segment with inner seta 2$ from S. brasiliensis from Panama (Atlantic), and outer spine, slender spinules at base of spine, third Colombia (Atlantic), Brazil and Argentina; 25 collec- segment with 6 inner to terminal setae and 3 outer tions containing 70 $ 40 $ from S. maculatus from spines; exopod spines broad at base, tapering distally Cape Cod, New Jersey, Maryland, South Carolina, with small terminal flagellum, fine hairs on both NUMBER 3 1 1 edges of spines; endopod first segment with inner nent spinules forming a proximal transverse band and seta and row of fine hairs along outer half of distal 2 distal patches of spinules. Caudal rami (Figure 46) border, second segment similar except with 2 inner slightly longer than wide (41 X 35 /an) with 6 setae; setae, third segment with 3 inner to terminal seta and longest seta 572 /tm. 2 outer spines similar to, but smaller than, exopod First antenna (Figure 4c) 5-segmented, second seg- spines, outer edges of endopod segments haired; inter- ment incompletely divided, antenna similar to female, podal plate with long slender spinules. Leg 3 (Figure but segments not as elongate. Second antenna, mandi- 3a) coxopod with row of small spinules at outer distal ble, paragnath, first maxilla, second maxilla similar to, corner; basipod with dorsal seta; exopod segments but proportionately smaller than, female. Maxilliped heavily sclerotized; first segment with outer spine, (Figure 4d*) second segment with 2 short setae near second segment with outer spine and inner seta, third inner mid-margin, inner surface of segment with patch segment with 2 outer spines and 6 terminal to inner of elongate, blunt-tipped spinules and patch of smaller, setae, exopod spines heavily sclerotized, spines on rounder spinules; terminal claw with 2 setae and second and third segments of about equal length, numerous closely spaced "teeth" along entire inner spine on first segment slightly shorter than others; edge. Legs 1-4 biramous, rami 3-segmented except endopod similar to leg 2 except third segment with leg 4 endopod. only 2 setae; interpodal plate with short, stout spinules. Leg 1 (Figure Ae) coxopod heavily haired along Leg 4 (Figure 3b) coxopod and basipod similar to outer distal edge; basipod with dorsal seta and ventral leg 3; exopod first segment with spinules along ventro- ornamentation as indicated in figure; exopod first seg- lateral edge, hairs on inner edge and spine on outer ment with row of elongate spinules near base of spine distal corner, second segment with inner seta and on outer distal corner, second segment similar to first outer spine, slender spinules at base of seta, shorter, with the addition of an inner seta, third segment with stouter spinules at base of spine, third segment with 4 inner to terminal setae and 3 outer spines, proximal 6 inner to terminal setae, 2 outer spines, slender 2 spines similar to those of first and second segments, spinules at bases of 2 innermost setae, stouter spinules finely haired on edges, and with terminal flagellum at bases of spines, spines slender, elongate with fine and with row of spinules near base, distal spine elon- hair on edges and fine terminal flagellum; endopod gate with row of very fine hairs along outer edge; first segment with patch of slender spinules near outer endopod first segment enlarged, covering inner portion distal corner and inner seta spinulose along distal two- of exopod, stout hairs along distal edge and an inner thirds, second segment similar to first, third segment seta, second segment only about half as wide as first elongate, broadened distally with 2 terminal patches segment with inner seta, small spinule on outer distal of stout spinules and 2 spines flanking terminal seta, corner and row of fine hairs along outer half of distal spines elongate, of about equal length and finely haired edge, third segment with 4 terminal setae and small on edges, seta spinulose along distal half; interpodal outer spinule, outer half of segment with 2 converging plate with short, stout spinules. Leg 5 (Figure 3c) rows of small, rectangular, platelike spinules, outer basal segment with dorsal seta and small patch of edges of endopod segments heavily haired. Leg 2 slender spinules; free segment with 3 patches of coxopod with short row of small spinules on outer slender spinules, one spine at about mid-point of outer distal corner; basipod with dorsal seta; exopod first margin and 2 spines flanking terminal, naked seta, segment with irregular rows of small bumplike spinules spines elongate, simple with finely haired edges. Leg along outer edge, short row of slender spinules at base 6 (see Figure 16) represented by 3 naked setae at of spine on outer distal corner, second segment with area of egg sac attachment. inner seta and row of spinules at base of outer spine, MALE.—Body form as in Figure 3d. Total length third segment with 5 inner to terminal setae and 3 1.18mm, greatest width 0.50mm (measured at widest outer to terminal spines, proximal 2 spines similar to part of cephalon). Genital segment (Figure 4a) longer spines of first and second segments, finely haired on than wide (236 X 182 pm). Abdomen (Figure 4a) edges with terminal flagellum and short row of spi- 2-segmented, segments measure (length X width) nules near base, terminal spine elongate with row of 88 X 118 /un, 82 X 106 /un, respectively; ventral very fine hairs along outer two-thirds; endopod first surface of second segment with 2-3 rows of promi- segment with outer seta and row of hairs along outer SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY distal edge, second segment similar to first but with Holobomolochus nudiusculus, new species two inner setae, third segment with 3 inner to terminal FIGURES 5d-e, 6, la-b, 96 setae, short row of spinules at base of 2 outer spines, inner spine about twice as long as outer, outer edges MATERIAL EXAMINED. — Holotype 9 (USNM of endopod spines heavily haired. Leg 3 (Figure 5a) 172247), allotype $ (USNM 172248), paratypes similar to leg 2 except endopod third segment with 33 5 IS (USNM 172249) from the nasal sinuses of 2 rather than 3 setae. Leg 4 (Figure 56) coxopod Scomberomorus sierra from Buenaventura, Colombia and basipod similar to leg 3; exopod similar to leg 3 (USNM 218565). In addition 13 collections contain- with the following exceptions: spines more slender; ing 30 9 1

ramus (see Figure 76). Caudal rami (Figure 7 b) 7e) longer than wide (82 X 53 /un), each ramus with longer than wide (47 X 29 /un), each with small six setae and a patch of hairlike spinules on its ventral ventral patch of slender spinules and 6 setae (longest surface; longest seta 768 /*m. 531 /an). First antenna (Figure 7/) 7-segmented with an Cephalic appendages and legs 1-5 as in H. divari- aesthete on segments 6 and 7. Second antenna, mandi- catus male. ble, paragnath, first maxilla, second maxilla, and ETYMOLOGY.—The Latin nudiusculus ("somewhat maxilliped similar to corresponding appendages in H. naked") refers to the small patches of minute hairs divaricatus. on the ventral surface of the last abdominal segment Leg 1 similar to that of H. divaricatus except endo- and the naked surface of the female caudal rami. pod and basipod lack hairs found in H. divaricatus. REMARKS.—Females of this species can be distin- Leg 2 (Figure 86) similar to that of H. divaricatus guished from H. divaricatus and H. asperatus by the except exopod second segment lacks spinules at base ornamentation of the ventral surface of the last ab- of outer spine; endopod first and second segments each dominal segment. Holobomolochus nudiusculus has 2 have a minute patch of small hairs on outer distal small patches of minute hairs on the last abdominal corner, outer spines on third segment are small. Leg 3 segment; H. divaricatus has 2 large patches of stout (Figure 8

on the rami of H. divaricatus or H. nudiusculus) and ETYMOLOGY.—The Latin, uni-colax (with "colax"), the lack of prominent patches of spinules on leg 4 the suffix "colax" is common in other genera of the (present in H. divaricatus and H. nudiusculus). family and the name alludes to the new genus "with" Holobomolochus asperatus can be distinguished from others of the family. all other members of the genus by the nature of the DISCUSSION.—Unicolax can be separated from all leg 3 exopod spines (see Discussion). the other bomolochid genera except Bomolochus by This species has been collected only from Scombero- the modified fourth seta on the basal segment of the morus cava.Ua from Georgia to northern Brazil. first antenna. Holobomolochus, Acanthocolax, Pseudo- Although its range overlaps that of H. divaricatus, the eucanthus, Orbitacolax, Pseudorbitacolax, Pumiliopes, 2 parasites have never been collected from a common and Pumiliopsis have no modified setae or cuticular host species. process on the first antenna; Boylea has the fifth seta modified; in Nothobomolochus the third, fourth, and Discussion fifth setae are modified; in Dicrobomolochus the sec- ond and third setae are modified; Ceratacolax and The 3 species of Holobomolochus described here all Tegobomolochus have a cuticular process in addition have highly developed spines on the exopod of leg 3 to the usual 15 plumose setae on the first antenna. and the exopod segments themselves are heavily Bomolochus is characterized by having the fourth sclerotized. This character separates these 3 species seta of the first antenna modified, but in Bomolochus from all previously described Holobomolochus. this seta is sharply curved and lightly sclerotized, Although we do not feel that this difference alone whereas in Unicolax it is straight or nearly so, and warrants assigning these new species to a new genus, heavily sclerotized. Unicolax can further be separated we feel that an improved understanding of the genus from Bomolochus as the rami of the first leg of Uni- might result in a better basis for doing so. No other colax males are flattened and broad as in the females; species of Holobomolochus has been collected from the first leg of Bomolochus males does not resemble the nasal sinuses of the hosts. This unusual habitat that of Bomolochus females. may account for the unique nature of these 3 species. Unicolax has, so far, only been collected from the nasal sinuses of its hosts. Vnicolax, new genus

DIAGNOSIS.—Bomolochidae. Body form typical of Vnicolax collateralis, new species family. Thoracic segments bearing legs 3-5 free in FIGURES 9-13, 97, 114-116* female, thoracic segments bearing legs 2-5 free in male. Abdomen 3-segmented in female, 2-segmented MATERIAL EXAMINED. — Holotype 9 (USNM in male. Caudal rami in both sexes with 6 setae, 2 172256), allotype $ (USNM 172257) paratypes 12 9 much longer than other 4. First antenna 6- or 7-seg- 23 d (USNM 172258) from the nasal sinus of mented in female, fourth seta on basal segment modi- Euthynnus alletteratus (USNM 203838) from St. fied to form heavily sclerotized straight or nearly George Bay, Lebanon. The following collections from straight spine; first antenna in male 6-segmented with Auxis species: 2 9 from Woods Hole, Mass.; 2 S no modified setae. Second antenna and oral append- from St. George Bay, Lebanon; 2 9 3$ from ages typical of family. Maxilliped hook in female Ghardaqa, Egypt; 2 collections containing 7 9 3$ without accessory process; maxilliped basal segment from Hong Kong; 2 collections containing 2 9 1 $ in male with numerous small spinules. Legs 1-4 from Japan; 7 collections containing 12 9 4$ from biramous in both sexes; rami of legs 1-3 and leg 4 the Philippines; 2 9 from Chusan, China; 1 9 from exopod 3-segmented, leg 4 endopod 2- or 3-segmented. Hawaii. One collection containing 5 9 from Cybio- Female leg 2 endopod second segment with 2 inner sarda elegans from Brisbane, Australia. The following setae, leg 3 endopod second segment with 1 inner seta. collections from Euthynnus affinis: 2 9 2 3 from Elat, Rami of leg 1 in both sexes broad and flattened. Israel; 2 9 from Mozambique; 2 9 from Madagascar; TYPE SPECIES.—Unicolax collateralis new genus, 1 S from the Seychelles; 4 9 from the Arabian Sea new species. 24°N, 67°E; 2 9 from Java; 16 collections containing NUMBER 311

29 9 9 $ from the Gulf of Thailand; 2 9 from For- sory blade smaller. Paragnath with distal and lateral mosa; 2 2 from Hong Kong; 4 collections containing hairs; also distal edge with fine teeth. First maxilla 8 9 1 $ from the Philippines; 1 2 from Palau; 1 9 with 3 long, plumose setae and one short, naked seta. from Okinawa; 1 $ from Tokyo; 4 2 7 $ from Bris- Second maxilla with 2 blades, each with uniform bane, Australia. The following collections from spinules along edges. Maxilliped (Figure 106) with Euthynnus alletteratus: 1 9 from the Caribbean 4 plumose setae; hook slightly curved, without acces- (9°11'N, 77°50'W); 2 9 from Brazil; 4 collections sory process. containing 29 9 16 $ from St. George Bay, Lebanon. Legs 1-4 biramous, each ramus 3-segmented except The following collections from Euthynnus lineatus: 1 leg 1 exopod. Leg 1 (Figure 10c) coxopod with patch 9 from Galapagos; 1 9 from Lower California; 1 9 of stout hairs on outer corner and broad inner seta; from Mexico (Pacific); 2 9 from Costa Rica (Pacific). basipod with outer seta, row of hairs above insertion One collection containing 1 9 from Orcynopsis uni- of endopod, and inner blunt process, exopod first seg- color from St. George Bay, Lebanon. Two collections ment with stout outer spine terminating in short containing 5 9 3 <$ from Sarda australis from New flagellum and second segment incompletely divided, South Wales, Australia. Two collections from Sarda with 2 short, outer flagellate spines, and 6 terminal orientalis containing 3 9 from Pearl Islands, Panama, to inner setae, endopod first segment with inner seta and Durban, South Africa. All collections from nasal and short row of spinules (hairs?) along outer, lower sinuses of hosts. edge, second segment similar to first, third segment FEMALE.—Body form as in Figure 9a. Total length with 5 terminal setae, outer edges of endopod seg- 1.55 mm, greatest width 0.91 mm; length of cephalon ments heavily haired, all setae heavily plumose. Leg 2 0.30 mm. Genital segment (Figure 96) wider than (Figure 10a") coxopod with patch of slender spinules long (200 X 312 fitn). Abdomen (see Figure 96) on outer corner, row of short hairs along mid lower 3-segmented, segments measuring (1 X w) 153 X edge, basipod with outer dorsal seta and triangular 206 pin, 100 X 171 /on, 94 X 141 /urn respectively. patch of minute spinules on midventral surface; exo- First 2 segments without ornamentation, last seg- pod first segment with long, slender spinules on outer ment with 2 large ventral patches of fine hairs (see edge, serrate spine with terminal flagellum on outer Figure 9c). Caudal rami (Figure 9c) longer than distal corner, inner edge with small patch of hairs, wide (88 X 53 /(in), each ramus with ventral patch second segment with inner seta and outer spine similar of fine hairs and 6 setae as indicated in Figure 9c; to that on first segment, third segment with 5 inner longest seta 413 fan. to terminal setae and 4 terminal to outer spines, First antenna (Figure 9d) 7-segmented. Fourth seta proximal spine with short hairs on both edges and on first segment modified to form a heavily sclerotized terminal flagellum, next 2 spines similar to those on spine. Ornamentation of segments as follows. Segment first and second segments, distal spine elongate with 1: 3 plumose setae; 1 spine; 1 plumose seta. Segment broad outer serrations and short inner hairs; all exo- 2: 4 antero-dorsal naked setae; 7 antero-ventral plu- pod setae long, slender and plumose; endopod first mose setae; 3 ventral plumose setae; 2 ventral naked segment with inner seta and row of fine hairs along setae. Segment 3: 1 dorsal naked seta; 3 antero-ventral distal edge, second segment enlarged with 2 inner plumose setae; 1 ventral naked seta. Segment 4: no setae and row of fine hairs on outer half of distal edge, ornamentation. Segment 5: 1 dorsal naked seta; 1 third segment narrower than second with 3 inner to antero-ventral naked seta; 2 ventral naked setae. terminal setae and 2 short terminal spines with short Segment 6: 2 terminal ventral naked setae; 1 aesthete. lateral hairs and terminal flagella; endopod setae stout Segment 7: 6 terminal naked setae; 1 subterminal basally and heavily plumose; outer edges of endopod naked seta; 1 aesthete. Second antenna (Figure 9e) segments haired. Leg 3 (Figure lla) coxopod with 3-segmented; second segment with one plumose seta row of short spinules on outer distal corner, small and one short, naked seta; third segment armed with central patch of blunt spinules and row of short hairs several rows of short spinules, and with 4 hooklike on mid-lower edge; basipod with outer dorsal seta; terminal spines and 3 setae. Mouthparts as in Figure exopod first segment with outer distal spine, dorso- 10a. Labrum with 2 large patches of hairs, hairs longer lateral surface of segment with several rows of short along anterior edge. Mandible blade slender, acces- spinules giving bumpy appearance, second segment 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY with inner seta and outer spine, third segment with 5 large patch of small, rounded spinules on second seg- inner to terminal setae and 3 terminal to outer spines, ment; last segment with 2 setae near base and row exopod spines similar to corresponding spines of leg 2, of teeth along inner edge. except leg 3 lacks haired spine present on third seg- Legs 1-4 biramous. Leg 1 (Figure 12*/) coxopod ment of leg 2, all setae plumose; endopod first seg- with inner seta and spinules as indicated in Figure ment with inner seta and row of short hairs along I2d; basipod with outer seta, blunt process between distal edge, second segment similar to first, third seg- insertion of rami, and inner process sclerotized at ment with 2 inner to terminal setae and 2 terminal to base with terminal, fringed membrane, spinules near outer spines, each with serrate edges and terminal base of this process; exopod 3-segmented, first segment flagellum, all setae plumose, outer edges of endopod with outer spine, second segment with outer spine and segments haired. Leg 4 (Figure life) coxopod with inner seta, third segment with 2 outer spines and 5 central patch of small spinules and row of short hairs setae; endopod first segment with inner seta, distal along mid-lower edge; basipod lacking ornamentation; outer edge of segment with row of hairs, second seg- exopod similar to leg 3 except first segment lacks dorso- ment similar to first, third segment with 5 setae lateral spinules; endopod first segment with inner seta terminally and one outer spine, minute spinules near and row of fine hairs along distal edge, seta shorter bases of setae; all setae heavily plumose. Leg 2 (Fig- than corresponding seta of leg 2, constricted slightly ure 12tf) coxopod with short row of spinules on outer at midpoint, plumose distally finely serrate proximally, distal corner and row of hairs on mid-distal edge; hairs on distal edge of segment slightly longer than basipod with naked seta dorsally near outer edge; corresponding hairs of leg 2, second segment similar exopod first segment with outer distal serrate spine to first, third segment with row of fine hairs distally with terminal flagellum, row of fine spinules near and 2 spines flanking terminal seta, spines broadly base of spine, dorso-lateral surface of segment with serrate and with terminal flagellum, seta finely serrate several rows of short, rounded spinules, second seg- on distal half, outer edges of endopod segments haired. ment with inner seta and outer spine similar to that Leg 5 (Figure lie) basal segment with outer dorsal on first segment, third segment with 5 inner to ter- seta and dense patch of short spinules on outer distal minal setae and 3 terminal to outer spines, proximal corner; free segment with 3 dense patches of short 2 spines similar to those on first and second segments, spinules, one spine on mid-outer margin and 2 spines distal spine elongate with broad outer serrations and flanking one terminal naked seta, spines with serrate short inner hairs; endopod first segment with inner edges and terminal flagellum. Leg 6 (see Figure 9b) seta and short row of hairs along distal edge, second represented by 2 long and one shorter seta on genital segment similar, third segment with 3 inner to ter- segment. minal setae and 2 outer spines, each finely haired on MALE.—Body form as in Figure lid. Total length edges and with terminal flagellum; outer edge of 1.09 mm, greatest width 0.44 mm; length of cephalon endopod segments heavily haired. Leg 3 (Figure 13a) 0.20 mm. Genital segment (Figure 11*) longer than similar to leg 2, with following exceptions; exopod wide (236X218 fan). Abdomen (see Figure lie) second segment lacks outer spine and fine spinules; 2-segmented, segments measuring 94 X 112 /*m and endopod third segment with only 2 setae, inner spine 70 X 88 /*m respectively. Last abdominal segment about twice as long as outer, ornamentation on edges ornamented ventrally with anterior row of fine spinules of each spine serrate rather than haired as in leg 2. and 2 large patches of fine hairs (see Figure 12a). Leg 4 (Figure 13ft) coxopod, basipod, and exopod Caudal rami (Figure 12a) longer than wide (53 X similar to leg 3 except exopod third segment with 35 fan) ; each ramus with ventral patch of fine hairs only 4 setae; endopod 2-segmented, first segment with and 6 setae; longest seta 295 /un. inner seta, second segment elongate with row of fine First antenna (Figure 12b) 6-segmented. First seg- hairs distally and 2 spines flanking terminal seta, ment with 5 plumose setae and no indication of spines finely serrate with terminal flagella, inner spine modified spine present on female; fifth and sixth seg- about one-third longer than outer, seta finely serrate ments each with one aesthete. Second antenna, man- along distal half. Leg 5 (Figure 13c) basal segment dible, paragnath, first and second maxillae similar to with outer dorsal seta and dense patch of spinules on female. Maxilliped (Figure 12c) with one seta and outer distal corner; free segment with terminal outer NUMBER 3 1 1 11 seta extending almost to end of genital segment, and Unicolax anonymous (Vervoort, 1965), new inner spine broadly serrate on edges and with terminal combination flagellum; segment with dense patch of spinules on FIGURES 14-16, 98, 116fr—/, 117 distal half. ETYMOLOGY.—The Latin collateredis ("standing Parabomolochus anonymous Vervoort, 1965:3. side by side") alludes to its occurrence in collections MATERIAL EXAMINED.—Two collections containing with U. mycterobius. 319 2 S from the nasal rosettes of Euthynnus allet- REMARKS.—Females of this species can be dis- teratus from Ghana and the Gulf of Mexico. tinguished from U. mycterobius, U. ciliatus, and FEMALE.—Body form as in Figure 14a. Total length U. reductus principally by the nature of the spines of cephalon 0.37 mm. Genital segment wider than long (177 X 247 /*m). Abdomen 3-segmented, seg- on the exopod third segment of leg 2 as the proximal ments measure 82 X 153 /*m, 53 X 141 /*m, 70 X spine of U. collateralis is finely haired on both edges 123 /urn length X width respectively. First 2 segments and the next two spines have broad serrations on both without ornamentation, last segment with 2 patches side. This is unlike the ornamentation of correspond- of fine hairs ventrally. Gaudal rami each with patch of ing spines on any of the other above species. Unicolax fine hairs and 6 setae; longest 171 ^m. collateralis can be distinguished from U. anonymous, First antenna (Figure 146) 7-segmented. Fourth which it most closely resembles, by having 3 spines, seta on first segment modified to form heavily sclero- 5 setae on the exopod third segment of leg 4; U. tized spine; one aesthete on each of last 2 segments. anonymous has 3 spines, 4 setae on the corresponding Second antenna (Figure 14c) 3-segmented, last seg- segment. The ventral surface of the endopod segments ment with several rows of hooklike spinules, 4 setae, of legs 2-4 are densely haired in U. anonymous, and 4 hooklike terminal spines. Mandible, paragnath, whereas U. collateralis has only a single row of hairs first maxilla, second maxilla as in Figure 14a\ Parag- along the distal edge of some of its endopod segments. nath with outer row of teethlike spinules and long hairs; first maxilla with 3 long plumose setae and Males of this species can be distinguished from those one short, naked seta. Maxilliped (Figure 14e) with of U. anonymous by the ornamentation of leg 5. Uni- one plumose seta on basal segment, 3 plumose setae colax collateralis has a patch of stout spinules that on last segment; hook slightly curved, without acces- extends from about the middle of the free segment sory process. distally and around the entire distal edge; in U. Legs 1-4 biramous, each ramus 3-segmented except anonymous the spinules are distinctly elongate and leg 1 exopod. Leg 1 (Figure 15a) coxopod with the patch extends distally from above the middle of broad inner seta; basipod with outer seta, one small the segment and does not continue around to the blunt spinule near lower inner edge and two large outer distal edge. Males can be distinguished from patches of hairs; exopod first segment with outer U. mycterobius, U. ciliatus, and U. reductus by the spine, second segment with 2 short outer spines and ornamentation of the ventral surface of the last ab- 6 terminal to inner setae; endopod first and second dominal segment. In U. collateralis this segment has segments each with inner seta and patch of hairs on a single, even row of spinules near the anterior lower to outer edge, third segment with 5 setae, outer border, and 2 large patches of hairs. Unicolax myc- edges of endopod segments heavily haired. Leg 2 terobius has similar ornamentation, but the anterior (Figure 156) coxopod with inner seta, outer distal row of spinules is irregular and the spinules are corner with short curved spinules; basipod with outer seta and patch of small spinules between insertion of minute. rami; exopod first segment with long, slender spinules This copepod is circumglobal and occurs in the on outer edge and stout broadly serrate spine with nasal sinuses of a variety of scombrid hosts (species terminal flagellum on outer distal corner, second seg- of Auxis, Cybiosarda, Euthynnus, Orcynopsis, and ment with inner seta and outer spine similar to that Sarda). It is often collected with U. mycterobius, of first segment, third segment with 5 inner to termi- which parasitizes some of the same hosts (species of nal setae and 4 terminal to outer spines, proximal Auxis and Euthynnus). spine with short hairs along both sides and terminal 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

flagellum, next 2 spines similar to those of first and 0.81 mm, greatest width 0.36 mm; length of cephalon second segments, distal spine elongate with broad 0.20 mm. Genital segment as long as wide (177 X outer serrations and short inner hairs; endopod first 177 /an). Abdomen 2-segmented, segments measuring segment with inner seta and large patch of short (length X width) 35 X 94 /un and 59 X 70 /un. hairs distally, second segment enlarged, with 2 inner Caudal rami slightly longer than wide (35 X 29 /*m) ; setae and patch of short hairs on outer distal corner, longest seta 177 pin. third segment narrower than second with 3 inner to All appendages and ornamentation similar to (al- terminal setae and 2 short terminal spines each with though proportionately smaller than) U. collaterals hairs on edges and terminal flagellum, outer edges except leg 5. of endopod segments haired. Leg 3 (Figure 16a) Leg 5 (Figure 16«) similar to U. collaterals except coxopod and basipod similar to leg 2; exopod first patch of spinules on free segment covers lower two- segment with outer distal spine, dorso-lateral surface thirds of inner half of segment and does not extend of segment with several uneven rows of short spinules around the entire distal edge; also, spinules on both giving bumpy appearance, second segment with inner basal and free segments ar*» longer and more slender seta and outer spine, third segment with 5 inner to than those of U. collateralis. terminal setae and 3 terminal to outer spines, exopod REMARKS.—Unicolax anonymous was described by spines similar to corresponding spines of leg 2, except Vervoort (1965) as Parabomolochus anonymous from leg 3 lacks haired spine present on third segment of the nasal sinus of Euthynnus alletteratus from the leg 2; endopod first segment with inner seta and Gulf of Guinea. patch of hairs covering distal half of segment, second We believe the copepods above to be the same segment with inner seta and patch of hairs on outer species as Vervoort's even though our descriptions distal corner, third segment with 2 inner to terminal differ on some points. Vervoort (1965:6) reported setae and 2 terminal to outer spines, inner spine "no spinules on the abdominal somites" of the female. longer, each with serrate edges and terminal flagel- We, however, found 2 patches of hairs on the ventral lum, outer edges of endopod segments haired. Leg 4 surface of the last abdominal segment as well as a (Figure 166) coxopod with row of short hairs on ventral patch of hairs on each caudal ramus. Ver- mid-distal edge; basipod with outer seta; exopod voort (1965:11) further reported that "the external first segment with spine on outer distal corner, second margin of the first exopod segment [of legs 2-4] is segment with inner seta and outer spine, third seg- strongly haired." We found this true only of leg 2; ment with 4 inner to terminal setae and 3 terminal on legs 3 and 4 the first exopod segment was orna- to outer spines, exopod spines similar to correspond- mented by several irregular rows of bumplike spinules. ing spines of leg 3; endopod first segment with inner In spite of these differences we feel we are dealing seta, proximal half plumose, distal portion edged with with the same species for the following reasons: the short bristles, segment with large patch of short hairs copepods reported here are from the same host species distally, second segment with inner seta similar to and the same locality (and the Gulf of Mexico as that of first segment, and patch of short hairs on outer well) as those described by Vervoort; Vervoort re- distal corner, third segment with 2 spines flanking ported collecting Ceratocolax euthynni from the same terminal seta, spines broadly serrate distally, seta with individual hosts as U. anonymous, we also found short bristles on lower half, distal edge of segment C. euthynni in both our collections with U. anony- with short spinules, outer edges of endopod segments mous; apart from the differences mentioned above, haired. Leg 5 (Figure 16c) basal segment with outer our descriptions are in agreement. seta and dense patch of short spinules on outer distal Females of U. anonymous can be distinguished corner; free segment with 3 dense patches of short from U. mycterobius, U. ciliatus, and U. reductus spinules, one spine on mid-outer margin, and 2 spines by the nature of the spines on the exopod third seg- flanking one naked seta distally, spines each with fine ment of leg 2. The proximal spine in U. anonymous serrations along distal half and terminal flagellum. is finely haired on both sides while the remaining Leg 6 (see Figure 14a) represented by 3 setae on spines are broadly serrate; this ornamentation is un- genital segment. like that of the corresponding spines of any other of MALE.—Body form as in Figure 16d. Total length the above species. Unicolax anonymous can be dis- NUMBER 3 1 1 13 tinguished from U. collateralis, which it most closely Cephalic appendages and leg 1 similar to Unicolax resembles, by the ornamentation of the exopod third collateralis. segment of leg 4; U. anonymous has 3 spines, 4 setae; Leg 2 (Figure 176) coxopod with inner seta and U. collateralis has 3 spines, 5 setae. Further, the row of fine spinules on outer distal corner; basipod endopod segments of legs 2-4 in U. anonymous each with outer seta and 2 small patches of fine hairs; have a dense patch of hairs; U. collateralis has only exopod first segment with large outer patch of fine a single row of hairs near the distal border of some hairs and spine on outer distal corner, second segment of its endopod segments. with inner seta and outer spine, third segment with Unicolax anonymous males have, on the free seg- 5 inner to terminal setae and 4 terminal to outer ment of leg 5, a patch of elongate spinules that ex- spines, exopod spines squat with short hairs along tends from above the middle of the segment to the margins and terminal flagellum except distal spine on inner distal border; the spinules do not continue third segment elongate and tapered with fine hairs on around the entire distal border of the segment. This outer edge, widely spaced hairs on inner edge; endo- ornamentation is unlike that of other species of Uni- pod first segment with stout inner seta, outer distal colax males. portion of segment with patch of short hairs, second This copepod has, so far, only been reported from segment enlarged with 2 stout inner setae and patch the nasal sinuses of Euthynnus alletteratus from the of short hairs on outer distal edge, third segment Gulf of Guinea and the Gulf of Mexico. It is a rela- narrower than second, with 3 stout inner to terminal tively small copepod and may, as Vervoort noted, setae and 2 terminal spines similar to those on exopod, frequently be overlooked. outer edges of endopod segments heavily haired. Leg 3 (Figure \7c) coxopod and basipod similar to leg 2; exopod first segment with patch of short hairs on mid- Unicolax mycterobius (Vervoort, 1965), outer margin and spine on outer distal corner, second new combination segment with inner seta and outer spine, third seg- ment with 5 inner to terminal setae and 3 terminal FIGURES 17-19, 99, 118, 119a-

This species is found from the western Pacific to outer spine with fine serrations and terminal flagellum, the Gulf of Guinea and is, so far, the only species of second segment incompletely divided, with one outer Unicolax found on species of Scomberomorus. flagellated spine and 6 terminal to inner plumose setae; endopod first segment with inner plumose seta Vnicolax reductus, new species and small hairs on distal and outer portion of segment, second segment with inner plumose seta, third seg- FIGURES 23-26, 99, \\9d-f, 12O-122a,6 ment with 5 terminal plumose setae. Leg 2 (Figure 24c) coxopod with few, widely spaced hairs along MATERIAL EXAMINED. — Holotype 9 (USNM distal edge; basipod with outer dorsal seta and patch 172259), allotype $ (USNM 172260), paratypes of fine spinules on distal edge near insertion of endo- 15 5 (USNM 172261) from Katsuwonus pelamis pod; exopod first segment with patch of fine spinules (USNM 176974) nasal sinus from New South Wales. on lower, outer portion of segment, row of several, Additional material from the same host: 24 9 2$ stout spinules near base of spine on outer distal corner, from Tahiti and 25 9 1 & from Japan. All copepods spine with few, broad serrations and fine terminal collected from the nasal sinuses of the hosts. flagellum, second segment with short, inner, naked FEMALE.—Body form as in Figure 23a. Total length seta, row of stout spinules near base of outer spine 2.94 mm, greatest width 1.50 mm; length of cephalon similar to that on first segment, third segment with 0.94 mm. Genital segment (Figure 236) wider than 4 inner to terminal, sparcely plumose setae, and 3 long (289 X 501 /*m). Abdomen (see Figure 236) outer to terminal spines, proximal 2 spines each with 3-segmented, segments measuring 206 X 247 /*m, row of stout spinules near base and similar to those of 188 X 230 Aim, 230 X 200 /un respectively; segments first and second segments, terminal spine elongate without ornamentation. Caudal rami (Figure 23c) with few, broad outer serrations; endopod first seg- longer than wide (206 X 88 /an) each ramus bearing ment with stout, inner, plumose seta and patch of 6 setae and no other ornamentation, longest seta fine hairs on lower, outer edge of segment, second 401 fim. segment enlarged, with 2 inner, plumose setae and First antenna (Figure 23d) 6-segmented. Ornamen- patch of fine hairs on outer, distal corner, third seg- tation of segments as follows. Segment 1: 3 plumose ment with 3 inner to terminal plumose setae and 2 setae; 1 seta modified to form a sclerotized spine; small, outer to terminal spines, each with finely ser- 1 plumose seta. Segment 2: 5 antero-dorsal naked rate edges, outer edges of endopod segments haired. setae; 10 long plumose setae in a diagonal line across Leg 3 (Figure 24d) coxopod without ornamentation; ventral surface of segment; 4 short, ventral plumose basipod with outer, dorsal seta and small patch of setae; 1 ventral naked seta. Segment 3: one antero- fine spinules near insertion of endopod; exopod similar ventral naked seta; one postero-ventral sparcely plu- to leg 2 except spinules on lower, outer portion of first mose seta. Segment 4: one ventral naked seta; one segment slightly larger and more sparcely placed than dorsal naked seta. Segment 5: 2 terminal ventral naked corresponding spinules of leg 2; endopod first segment setae; one ventral aesthete. Segment 6: 6 terminal with inner, sparcely plumose seta and patch of fine naked setae; one sub terminal naked seta; one terminal spinules on outer half of distal edge, second segment aesthete. Second antenna (Figure 23*) similar to similar to first, third segment with 2 inner to terminal U. collaterals and ciliatus. Labrum similar to U. col- sparcely plumose setae, 2 outer to terminal spines each lateralis. Remaining mouthparts as in Figure 23/. with broadly serrate edges and terminal flagellum Maxilliped (Figure 24a) basal segment with one plu- (terminal spine longer than outer spine) and small mose seta; second segment with 3 plumose setae; hook patch of spinules near base of spines, outer edges of bent, with elbowlike shape, and without accessory endopod segments with short hairs. Leg 4 (Figure process. 25a) coxopod and basipod similar to leg 3; exopod Legs 1-4 biramous, each ramus 3-segmented except similar to leg 3 with following exceptions: first seg- leg 1 exopod and leg 4 endopod. Leg 1 (Figure 246) ment with very few spinules on outer portion of seg- coxopod with broad inner seta, patch of small hairs ment; third segment with only 3 rather than 4 and a blunt process near inner distal edge; basipod with sparcely plumose setae; also spines slightly longer than outer patch of small hairs; exopod first segment with corresponding spines of leg 3; endopod first segment 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY with inner seta finely serrate distally and patch of similar to female except exopod setae comparatively spinules along outer distal edge, few spinules on outer longer; endopod setae longer than and not as robust edge, second segment incompletely divided with tri- as in female and second segment not as inflated. Legs angular patch of spinules proximal to division indicat- 3 and 4 similar to female. Leg 5 (Figure 26d) basal ing probable fusion of second and third segments, segment with dorsal naked seta; free segment orna- uneven row of stout spinules distally near bases of mented similarly to female, but segment itself not as terminal seta flanked by 2 spines, seta finely serrate robust. distally, outer spine similar to those on exopod, inner ETYMOLOGY.—The Latin reductus ("reduced") spine with fine serrations, outer edge of segment with alludes to the reduced number of setae on exopods two patches of short hairs. Leg 5 (Figure 25b) basal of legs 2-4 and the reduced number of segments of segment with naked, dorsal seta, patch of spinules on the endopod of leg 4. outer, distal corner; free segment with patch of spi- REMARKS.—Females of U. reductus can be distin- nules on inner edge and a row of stout spinules near guished from other Unicolax by the following char- base of each of the following: one, terminal, naked acters: the ventral surface of the last abdominal seg- seta; 3 spines, one on mid-outer edge, one on each ment has' no ornamentation; the caudal rami have side of terminal seta, all spines stout with broad serra- only six setae each, with no ventral hairs or spinules tions distally and fine, terminal flagellurn. Leg 6 (see (other species have a ventral patch of hairs on each Figure 23b) represented by 3 short setae, of about ramus); leg 2 exopod third segment has 3 spines, equal length, on genital segment. 4 setae (other species have 4 spines, 5 setae); leg 3 MALE.—Body form as in Figure 25c. Total length exopod third segment has 4 setae (other species have 1.78 mm, greatest width 0.78 mm; length of cephalon 5) ; leg 4 exopod third segment has 3 setae (U. col- 0.39 mm. Genital segment (Figure 25d) slightly lateralis, U. mycterobius, and U. ciliatus have 5, U. wider than long (277 X 300 /mi), abdomen 2-seg- anonymous has 4) ; leg 4 endopod is 2-segmented (in mented, segments measuring (length X width) 159 all other species it is 3-segmented). X 177 /an, and 194 X 147 /on; second segment with Males of U. reductus differ from others of the genus two ventral rows of minute spinules (see Figure 25^). by the following characters: the ventral surface of Caudal rami (Figure 25«) longer than wide (141 X the last abdominal segment is ornamented by only 2 73 fixn) with no ornamentation other than 6 setae distal rows of minute spinules; the caudal rami have (longest 371 pin). no ornamentation on the ventral surface (other males First antenna similar to that of U. collateralis male. have a patch of hairs on the ventral surface of each Remaining cephalic appendages (except rnaxilliped) ramus). Legs 3, 4, and 5 are strikingly similar to those similar to those of female. Maxilliped (Figure 26a) of the female, while the legs of males of others in second segment inflated with 2 naked setae and nu- the genus differ in several ways from their female merous irregular rows of low, rounded spinules; ter- conspecifics. minal segment claw-like with 2 naked setae and row of Unicolax reductus is the only member of the genus teethlike spinules along inner edge to apex. found on Katsuwonus pelamis and is probably re- Legs 1-4 biramous. Leg 1 (Figure 266) basipod stricted to that host species. with stout outer seta, patch of hairs near insertion of Because of the number and nature of its unique rami, inner spine sclerotized at base with terminal characteristics, it is the most easily identified member fringed membrane, patch of spinules near base of of the genus. spine extending midway up segment; exopod 3-seg- mented, first segment with outer spine with stout Ceratacolax Vervoort, 1965 terminal flagellum, second segment with inner seta and outer spine similar to that on first segment, third Ceratacolax enthynni Vervoort, 1965 segment with 5 setae and 2 outer spines, proximal FIGURES 27-31, 100, 122c-/, 123-125 spine similar to previous 2, distal spine simple, not heavily sclerotized; endopod 3-segmented, similar to Ceratacolax euthynni Vervoort, 1965:26. U. mycterobius male except patches of hairs on first MATERIAL EXAMINED.—34 collections containing and second segments smaller. Leg 2 (Figure 26c) 94 9 and 63 S from the following hosts and localities: NUMBER 3 1 1 19

Euthynnus alletteratus from Massachusetts, New to the spinulose setae of the female. Perhaps the most Jersey, Florida (west coast), and Ghana; Sarda sarda unique feature of the male is the presence of long from Massachusetts, Rhode Island, Chesapeake Bay, slender spinules on the base of one seta on each Florida (east and west coasts), Mississippi, Texas, endopod last segment of legs 1-3. Venezuela, Brazil, Spain (Cadiz), Gulf of Guinea, In both sexes there is some individual variation in and South Africa (Port Elizabeth). All collections the pattern of spinules of patches on the last abdomi- from the nasal sinuses of the hosts. nal segment and caudal rami. In the female, particu- Vervoort provided a good discription of both the larly, spinules on the caudal rami may appear as a male and female of this species. We have figured both single longitudinal row or as a patch of 2-3 rows. sexes completely and will comment only on those We could not correlate this variation to host or geo- characteristics not included in, or which differ from, graphic distribution. The variation seems to occur Vervoort's description. randomly with occasional variation on the same in- FEMALE.—Ventral surface of last abdominal seg- dividual from right to left ramus. ment and caudal rami (Figure 27d) with patches of prominent spinules (omitted by Vervoort). Leg 1 Nothobomolochus Vervoort, 1962 (Figure 286) exopod 2-segmented (Vervoort indicated 1), second segment incompletely divided; first seg- Nothobomolochus kanagurta (Pillai, 1965), ment with one crenate spine on outer distal corner, new combination second segment with 3 outer spines (omitted by Ver- voort) and 6 inner to terminal setae. FIGURES 32, 33, 100 MALE.—Ventral surface of last abdominal segment Bomolochus kanagurta Pillai, 1965:51. and caudal rami with patches of spinules as indicated in Figures 27c and d (Vervoort omitted spinules on MATERIAL EXAMINED.—5 collections containing 9 9 caudal rami). Legs 1-3 (Figures 30c, 30rf, 31a) endo- from the gills of Rastrelliger kanagurta from China, pod third segment, second from innermost seta with India (Madras), Red Sea; R. faughni from the row of 4-5 long, slender spinules along proximal outer Philippines. edge (spinules omitted by Vervoort). FEMALE.—Body form as in Figure 32a. Total length REMARKS.—This bomolochid can be easily sepa- 1.50 mm. Abdomen 3-segmented, segments measure rated from other known members of the family on the 118 X 206 /an, 95 X 195 /an, 95 X 177 pin, length basis of the following characteristics. The structure of by width respectively. Last abdominal segment (Fig- the first antenna is unique by the presence of a long, ure 32b) with 2 oblique patches of spinules. Caudal curved, heavily sclerotized dorsal hook situated at the rami (see Figure 326) about twice as long as wide junction of the first and second segments; this hook (94 X 55 jan); each ramus with a lateral seta, 2 sub- is in addition to the usual row of 15 plumose setae on terminal and 2 terminal setae, one of which is much the basal segments and apparently does not represent larger, its base nearly as wide as distal end of ramus. a modified seta or setae as in some other bomolochid First antenna (Figure 32c) with 3 modified setae on genera. The genital segment includes 3 flaplike struc- basal segment, outer 2 longer than more heavily tures at the area of egg sac attachment; within these sclerotized middle seta; remaining setae of usual structures are 3 spinulose setae (not easily visible with- bomolochid type; an aesthete on each of last 2 seg- out dissection) representing leg 6. Leg 2 exopod and ments. Second antenna with rows of hooklets on sec- both rami of legs 3 and 4 are characterized by patches ond segment and armed with terminal setae typical of stout spinules along outer edges, and by setae that of the genus. Oral appendages typical bomolochid. are spinulose rather than plumose. Leg 4 endopod is Maxilliped (Figure 32d) with recurved hook bearing conspicuously elongate, especially the terminal seg- a short, blunt-tipped accessory process, base armed ment. with 3 prominent plumose setae. Males are characterized by a first leg that is neither Legs 1-4 biramous, each ramus 3-segmented. Leg 1 flattened nor modified as in the female. Legs 2 and 3 endopod with broad segments as in other bomolochids. endopod second segment both have 2 inner setae. Leg 2 (Figure 33a) coxopod with a group of long The setae of the male legs are all plumose in contrast spinules on outer distal corner and a row of short 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY spinules at articulation with basipod; basipod with a ing from between bases of first antennae and bearing cluster of long spinules arranged in a circle near inner a pair of ventral hooks (see Figure 346). margin and a long, stout seta at outer distal corner; Abdomen 3-segmented, ventral surface covered with exopod first segment with a patch of long spinules spinules. Caudal rami about twice as long as wide, along outer margin and a heavy serrated spine at each with one lateral, 2 subterminal, and 3 terminal outer distal corner, second segment with serrated spine setae (innermost much stouter and longer than at outer distal corner and an inner seta, third seg- others). First antenna (Figure 346) bearing 15 ment with 3 serrated spines, a fringed terminal spine, plumose and 6 naked setae along outer edge and 1 and 5 terminal to inner setae (last 5 spines armed on plumose and 3 naked setae directed posteriorly on 1 margin only) ; endopod first 2 segments each with first 3 segments, fourth and fifth segments with 3 setae a row of spinules on distal margin in addition to usual each, last segment with 7 short and 1 long terminal inner setae, last segment with 2 short, outer, lightly setae. Second antenna (Figure 34c) with 6 rows of fringed spines and 3 terminal setae. Leg 3 (Figure small hooks along outer edge of second segment and 336) coxopod with a row of stout spinules at outer 6 setae at tip (Pillai indicates 5). Labrum with 2 distal corner; basipod with 3 patches of spinules (outer dense patches of spatulate spinules, posterior corners 2 fine, inner patch heavier) and a seta on outer distal produced as spinulose knobs. Other oral appendages corner; exopod similar to leg 2 except no spinules on as in Figure 34d. Maxilliped (Figure 34e) hook with first segment and 1 less spine on last segment; endopod accessory process, posterior edge of basal segment with as in leg 2 except 1 less seta on last segment. Leg 4 spinules (not indicated by Pillai). (Figure 33c) as in leg 3 except 1 less patch of fine We will not repeat a full description of legs 1-5 but spinules on basipod; 1 less spine and seta on exopod rather restrict our discussion to those points where our last segment; endopod patches of spinules larger, material differs with the description of Pillai's. Pillai setae with short spinules rather than usual plumosities, illustrated the basipod of leg 1 with 1 large patch of last segment with only 1 long seta. Leg 5 (Figure 33d) spinules; in our specimens there are 2 (see Figure 34/). with 2 prominent patches of spinules in distal third, Leg 2 as described by Pillai except endopod second innermost with longer spinules; 1 lateral, naked seta segment with 2 setae rather than one. Leg 3 exopod and 3 terminal setae, outer 2 finely spinulose (all setae similar to that of leg 2; endopod first and second seg- of about equal length). ments each with one inner seta, third segment with 2 MALE.—We did not collect males of this species naked subequal outer spines and 2 inner setae. Pillai but Pillai noted (1965:53) that the second segment shows no setae on first and second segments and only of the maxilliped bears 2 rows of "tubercles" on inner one outer spine on third segment. Legs 4 and 5 (Fig- margin. ure 35a,6) as described by Pillai. REMARKS.—This species has been collected only REMARKS.—In spite of the differences between our from species of Rastrelliger from the Indo-West material and the description given by Pillai we feel Pacific. certain that we are dealing with the same species as the same genus of host is involved in both cases. So far this copepod has been collected only from Rastrel- Orbitacolax Shen, 1957 liger from India and the Philippines. Orbitacolax aculeatus (Pillai, 1962), new combination Pumiliopes Shen, 1957

FIGURES 34, 35a-b, 100 Pumiliopes capitulatus Cressey and Boyle, 1973 Bomolochus aculeatus Pillai, 1962a: 610. FIGURES 35c-e, 36, 37, 100, 126a-c MATERIAL EXAMINED.—2 collections containing Pumiliopes capitulatus Cressey and Boyle, 1973:1. 15 9 from the nasal sinuses of 2 Rastrelliger faughni from Manila and Lingayan Gulf, Philippines. MATERIAL EXAMINED.—23 collections containing FEMALE.—Body form as in Figure 34a. Total length 35 5 from the orbits of the following hosts and locali- 1.71 mm. Greatest width 0.73 mm. Rostrum protrud- ties: Rastrelliger kanagurta the Red Sea, Sri Lanka, NUMBER 311 21

India (Madras), western Indian Ocean, Philippines, panodon punctatus from Hong Kong. The specimens Java; R. faughni from Philippines; Scomber japonicus reported here vary somewhat from the original de- from Gulf of Guinea, Mauritania, Zanzibar; S. aus- scription. The most obvious difference is the greater tralasicus from Philippines. number of scalelike spinules in the patches on the FEMALE.—Body form as in Figure 35c. Total length caudal rami and abdomen of the scombrid parasites. of holotype 1.53 mm, greatest width 0.75 mm. Cepha- Also, the scombrid copepods have only 1 setule (in- lon about as long as wide. Genital segment (Figure appropriately referred to as setae in our original de- 35d) wider than long. Abdomen (see Figure 35

as in Figure 38c. Maxilliped absent. When Kabata described this species he stated that Leg 1 (Figure 38rf) biramose; exopod 2-segmented, his description was based on an immature female. The first segment with a short toothed spine on outer distal first author examined that specimen during the course corner, second segment with 4 toothed spines and 3 of another study and compared some of the present weak setae along inner margin; endopod 3-segmented, material with it. It was concluded that Kabata's speci- first segment unarmed, a non-articulated spine on men was a nonovigerous female adult. We have found outer distal corner of second segment, last segment no differences between the specimens reported here with 2 non-articulated terminal spines and a small and Kabata's description except for the presence of seta on inner margin. Leg 2 (Figure 39a) as in leg 1 egg sacs on some of our specimens. We will point out except one less spine and seta on exopod last segment the differences between this species and S. inauris and an additional seta on endopod last segment. Leg 3 rather than repeat a full description here. We have, (Figure 39b) 1-segmented, bearing a spine on inner however, provided several SEM photographs of a distal corner and 2 short terminal setae. Legs 4, 5, and female S. occlusa (from Gymnosarda unicolor). Fe- 6 absent. males of S. occlusa can be distinguished from S. in- Egg sacs typically cyclopoid. auris by the following: the abdomen of S. occlusa is MALE.—Body form as in Figure 38a. All males about one-sixth of the total body length (one-third in collected were attached to females. Body segmentation S. inauris) ; the exopods of legs 1 and 2 of S. occlusa well defined. Abdomen 4-segmented. Caudal rami as are 3-segmented (2-segmented in S. inauris). Males in female. of the 2 species can be separated by the following First antenna 8-segmented each with short setae. characters: distal segment of the second antenna of Second antenna (Figure 39c) 3-segmented; first seg- S. occlusa with 3 clawlike terminal spines (a bifid ment with 2 short setae on inner margin, second seg- claw in S. inauris); 3-segmented exopods of legs 1 ment with a seta on outer distal comer and 3 setae on and 2 of S. occlusa (2-segmented in S. inauris); 2 a sclerotized ridge at inner distal corner, third segment stout, but unequal, spines at tip of leg 2 exopod of in form of a bifurcate claw with an accessory spine S. occlusa (1 stout spine in S. inauris). near base. Legs 1-3 similar to female except last REMARKS.—This species is very similar to S. inauris exopod segment of leg 2 with a long, heavily sclero- but easily separated by the characters cited above. The tized spine (Figure 39d). species so far has been collected from the Western REMARKS.—This species so far has been collected Pacific, Indian Ocean, and Eastern Atlantic. It should only from species of Scomberomorus from the western be pointed out that a third species (S. elagata Cressey, Atlantic. For a more complete description the reader 1976) has been described from the carangid Elagatus should consult the original description (Cressey, 1975). bipinnulata (Quoy and Gaimard) from the Western Pacific. Shiinoa occlusa Rabat a, 1968 Caligus Muller, 1785 FIGURES 39e-h, 101, \2U-f, 127-128a,fc Shiinoa occlusa Kabata, 1968a:497. Caligus coryphaenae Steenstrup and Liitken, 1861

MATERIAL EXAMINED.—21 collections containing FIGURES 40, 41a-6, 102; 128c-/, 129, 130 41 9 11 $ from the nasal lamellae of the following hosts and localities: Grammatorcynus bicarinatus from Caligus coryphaenae Steenstrup and Liitken, 1861:352. Caligus scutatus Milne-Edwards, 1840:453. North Celebes, Solomon Islands, Palau, Caroline Is- Caligus thymni Dana 1852:56. lands; Gymnosarda unicolor from Solomon Islands; Caligus bengoensis Scott, 1894:130. Scomberomorus commerson from Mozambique, Paki- Caligus aliuncus Wilson, 1905:576. stan, Gulf of Thailand, Solomon Islands, Philippines, Caligus elongatus Heegaard, 1943:11. Caligus tesserifer Shiino, 1952:89. Palau; S. guttatus from China; S. niphonius from Japan; S. queenslandicus from Papua; S. tritor from MATERIAL EXAMINED.—138 collections containing Canary Islands; Acanthocybium solandri from Kapin- 316 9 and 281 $ from the body surface and gills of garmarangi Atoll. the following hosts and localities: Acanthocybium NUMBER 3 1 1 23 solandri from northeast coast of Malagasy Republic, spinules, patches of longer hairs, a short plumose seta Auxis species from Gulf of Guinea; Euthynnus allet- on inner margin, a long, very plumose, seta at outer teratus from Puerto Rico (Atlantic) ; Katsuwonus distal corner; exopod 3 distal spines each with promi- pelamis from northwest Malagasy Republic, Christmas nent fringes, inner 2 with accessory process; endopod Island (Pacific), Peru, Ecuador, east coast of United reduced to a short, sclerotized process. Leg 2 endopod States (several localities), Puerto Rico (Atlantic), (Figure 40e) with patches of long spinules on outer Venezuela, Brazil (north coast), Cape Verde Islands, margin of each segment. Leg 3 exopod (Figure 40/) Gulf of Guinea; Thunnus alalunga from New Jersey; with prominent thumblike spine on outer distal corner Thunnus albacares from Christmas Island (Pacific), of first segment, each of last 2 segments with long east coast of United States (several localities), Brazil, setules on outer margin in addition to usual spines and Gulf of Guinea; Thunnus atlanticus from Puerto Rico setae. Leg 4 exopod (Figure 41a) 3-segmented, first (Atlantic); Thunnus obesus from Christmas Island and second segment each with a prominent fringed (Pacific), east coast of United States (several locali- spine on outer distal corner; last segment with 3 ties), Surinam, Brazil (north coast), Cape Verde Is- fringed spines, distalmost longest, all spines with a lands; Thunnus thynnus from east coast of United fringe at base; spines measure 366, 236, 153, 206, and States (several localities). 247 /un distalmost to proximalmost respectively. Shiino (1959a), Pillai (1962b), and Lewis (1967) MALE.—Body form as in Figure 416. Total length have provided good descriptions and figures for this 5.3 mm. Cephalon comprises more than half total well known species. We will restrict our consideration length. Appendages as in female except second an- of this species to those characters that serve to dis- tenna with accessory process on claw. Maxilliped with tinguish it from the other Caligus species found on small sclerotized area opposite tip of claw. Sternal scombrid hosts. furca similar to female except tines not quite as diver- FEMALE.—Body form as in Figure 40a. Lewis, et al. gent. Thoracic appendages as in female. Legs 5 and 6 (1969:423) provided morphometric data for 36 represented by setae at the posterior corners of the specimens from 4 host species from the Indian Ocean. genital segment. The average total length for his material was 5.57 REMARKS.—This species has been reported many mm. Our material from the Indian Ocean and Pacific times since its original description in 1861. It is com- areas did not differ significantly from this mean. The mon on the body surface of scombrids of the tribe Atlantic specimens, however, tended to be larger (5.8- Thunnini. The single collection from Acanthocybium 6.5 mm based on 10 specimens from various localities). solandri (Scomberomorini) is the only exception re- The cephalon comprises about one-half of the total ported here of all scombrids examined. It is also com- length with the genital segment and abdomen each mon on species of Coryphaena and is found in all ex- comprising about one-fourth. cept polar oceans (see Figure 102). Margolis, Kabata, Frontal lunules widely spaced; space between lun- and Parker, 1975:25 provide a complete synonymy to ules (1.31 mm) more than twice greatest diameter of this species. either lunule (0.58 mm). Further discussion of this species follows the descrip- Genital segment slightly longer than wide (1.5 X tion of C. regedis. 1.4 mm) with posterior outer corners produced some- what beyond origin of abdomen. Abdomen 3-seg- mented; 1st segment constricted in posterior two-thirds Caligus regalis Leigh-Sharpe, 1930 giving the appearance of 2 segments, second segment FIGURES *\c-g, 102 shortest, third segment with 2 distal lobes separating caudal rami; segments measure 0.9, 0.3, and 0.37 mm Caligus regalis Leigh-Sharpe, 1930:5. respectively. Caudal ramus as in Figure 406. Caligus euthynus Kurian, 1961:63. Caligus alveolaris Heegaard, 1962:156. Postantennal spine lacking. Postoral spine wide at base, triangular. Sternal furca (Figure 40c) with MATERIAL EXAMINED.—5 collections containing 149 short, widely divergent tines; furca with an accessory 5, 113 $ from the body surface of Euthynnus sclerotized cuticular process on each side. affinis collected off Nosy Be, Malagasy Republic. Leg 1 (Figure 40rf) basipod with a patch of short Types of C. alveolaris (AMS P. 16408) from the 24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY same host species, reported as E. allitteratus [sic] We have placed Heegaard's C. alveolaris and from north Queensland, Australia. Kurian's C. euthynus in synonymy with C. regalis of FEMALE.—Body form as in Figure 41c. Total length Leigh-Sharpe. Although Leigh-Sharpe's figures are 4.62 mm (4.27-4.73 mm) based on an average of 5 poor there is little doubt that they illustrate the cope- specimens. Greatest width 2.1 mm (measured at pods discussed here, especially considering that he widest part of cephalon). Cephalon comprises about collected his material from the same host-species (as one-half total body length. Genital segment somewhat E. yaito). longer than wide (1.77 X 1.55 mm), posterior corners produced to form rounded lobes. Abdomen 2-seg- Caligus productus Dana, 1852 mented, segments measure 1.02 and 0.43 mm long respectively, comprising about one-third total body FIGURES 42-44, 103 length. Caudal rami as in Caligus coryphaenae except outer terminal seta much stouter in C. regalis. Caligus productus Dana, 1852:56. Caligus alalongae Kroyer, 1863:129. Frontal lunules widely spaced, each about 400 /*m Caligus monacanthi Kroyer, 1863:133. wide, rostral space between each lunule about 950 /xrn. Caligus lobatus Wilson, 1935:1. Cephalic appendages as in C. coryphaenae except Caligus katuwo Yamaguti, 1936:6. sternal furca not as divergent in C. regalis (Figure Caligus dentatus Heegaard, 1962:160. Caligus microdontus Heegaard, 1964:139.

Legs 1-4 as in C. coryphaenae; the stout spine on This species has been recorded many times from the first exopod segment of leg 3 (Figure 41«) is scombrids and occasionally non-scombrid hosts (see curved inwards rather than straight as in C. cory- Lewis, 1967:116). Shiino, 1959b and Lewis, 1967 phaenae. The apical seta on the exopod of leg 4 have provided good descriptions and we do not feel (Figure 41/) is only slightly longer than the adjacent a need to repeat another here. We have, however, seta, setae measure 189, 177, 177, 177, and 183 /*m provided pertinent illustrations and a consideration distal to proximal respectively. of certain characters to enable the reader to identify MALE.—Body form as C. coryphaenae. Total length this species without resorting to other literature. A 3.51 mm (3.37-3.53 mm) based on an average of 5 complete synopsis of the literature has been provided specimens. Greatest width 1.73 mm (measured at by Margolis, Kabata, and Parker (1975:64). widest part of cephalon). Genital segment (Figure MATERIAL EXAMINED.—207 collections containing 41g) wider than long (914 X 711 /un). Caudal rami 2286 9 , 657 <$ from the mouth, gills, and body sur- with outermost terminal seta stout and about as wide face of the following hosts and localities: Acanthyocy- at base as other 3 (much smaller in C. coryphaenae). bium solandri from the northeast Malagasy Republic, Legs 5 and 6 at posterior outer corners. Abdomen 2- Seychelles, Line Islands, Campeche Bay; Scombero- segmented, segments measure 247 and 479 /tin re- morus cavalla from east coast United States (several spectively. localities); Scomberomorus regalis from Dominican REMARKS.—Pillai, 1962b, discussed the relationship Republic, Puerto Rico, Virgin Islands; Scombero- between this species and C. coryphaenae and provided morus species from Australia (east coast); Scombero- a comprehensive description of C. regalis (as C. morus tritor from Nigeria; Sarda sarda from Massa- euthynus). All collection records to date indicate that chusetts; Sarda orientalis from Panama (Pacific); C. regalis is specific to Euthynnus affinis and C. Sarda chiliensis lineolatus from Baja California; Auxis coryphaenae is primarily found on other species of species from Panama (Pacific), Mexico (Pacific); scombrids of the tribe Thunnini. Euthynnus affinis from Hawaii; Euthynnus allettera- These 2 species of Caligus differ from other mem- tus from Bermuda, Anagada Passage (W. Indies); bers of the genus in lacking a postantennal process Katsuwonus pelamis from northwest Malagasy Re- (Lewis 1967:105, considered the group of sensillae public, Tahiti, Line Islands, Suitcases Bank (E. at the usual site of the postantennal process as rep- Pacific), Peru, Ecuador, Gulf of Mexico (Alabama), resenting the process). When the genus Caligus has New Jersey, Dominican Republic, Venezuela, Brazil, undergone a much needed revision it may be that Gulf of Guinea; Thunnus alalunga from North Caro- these 2 species should be removed from the genus. lina; Thunnus albacares from northwest Malagasy NUMBER 3 1 1 25

Republic, Mozambique Channel, Chagos Islands, tainly our own records, based in large part on the Somalia, Palau, Line Islands, Brazil (north coast), examination of preserved hosts, would be biased in Dominican Republic (Atlantic), Bermuda, east coast that regard. United States (several localities) ; Thunnus atlanticus Although we collected Caligus productus from 14 from West Indies (several localities), Nicaragua; species of scombrid fishes it is common (found on Thunnus thynnus from east coast United States more than 25% of the specimens examined) on only (several localities), Gulf of Mexico, Japan. 4 species; Katsuwonus petamis 52%, Thunnus alba- FEMALE.—Body form as in Figure 42a. Range of cares 45%, T. atlanticus 92%, and T. thynnus 28%. total length (including data of Lewis 1967, 1968) It is curious that, although this parasite was found 3.85-5.45 mm. We did not find any significant differ- on 8 of the 13 species of Thunnini, we did not ences in total length between our measurements of collect it from any of the 40 specimens of T. obesus Atlantic specimens and those of ours and Lewis' from examined. the Indian and Pacific oceans. Examination of speci- This species was not found on any species of the mens from several different hosts from widely sepa- tribe Scombrini (Rastrelliger and Scomber). Leigh- rated geographic areas revealed no significant varia- Sharpe (1926:384) reported this species from Scomber tions in form. It appears that this is a very stable scombrus and placed the previously described Caligus species, well established as a parasite of scombrids scomberi Bassett-Smith in synonymy with it. Later except members of the tribe Scombrini {Scomber and writers (see Margolis, Kabata, and Parker 1975:74) Rastrelliger). and our examination of the type of Caligus scomberi Caligus productus females can be easily separated agree that it is a synonym of Caligus pelamydis from other species of the genus by a combination of Kroyer, 1863 and the records of C. productus from the following characters: the outer distal corners of Scomber scombrus discounted. the genital segment are rounded and extend well beyond the insertion of the abdomen; the ventral surface of the abdomen bears a medial subterminal Caligus asymmetricus Kabata, 1965 patch of minute spinules and a terminal transverse FIGURES 45-47, 104 patch of fine hairs (see Figure 426); the last exopod segment of leg 1 lacks the usual 3 lateral setae; the Caligus asymmetricus Kabata, 1965:109. Caligus thynni Pillai, 1963:89. outer distal corner of the first endopod segment of leg 2 bears a row of long spinules, the second seg- In addition to Pillai's original description, Lewis ment bears a double row of stout spines (7-9 per row) (1967:131) redescribed this species and presented an along its outer edge; leg 4 exopod is 2-segmented, the account of its rather confusing history. In view of last segment bearing one lateral and 3 terminal setae these recent works we will not present another full (outermost seta slightly longer than others). description of C. asymmetricus, but, as with some MALE.—Cephalothoracic appendages as in female other Caligus species from scombrids, discuss only the except second antenna (Figure 44«) with short claw salient features and provide full illustrations to enable and 2 prominent bossed areas on basal segment. Geni- the reader to identify this species without consulting tal segment and abdomen as in Figure 440*. the earlier works. REMARKS.—This species is common on scombrid MATERIALS EXAMINED.—22 collections containing fishes throughout the circumtropical and subtropical 719 and 25 $ from the gills, gill arches, and roof of area. We have recorded it here from 15 species of the mouth (1 collection) of the following hosts and hosts, infesting the buccal area, gills and body surface. localities: Grammatorcynus bicarinatus from Palau, Data presented by Lewis, et al, 1969, and our own Marshall Islands, Kapingamarangi Atoll; Scombero- collection data indicate the parasite is most commonly morus commerson from New South Wales, East found in the buccal area, next in the gill area, and Indies; Sarda australis from New South Wales; Sarda occasionally on the body surface. The fewer records orientalis from Seychelles, Hawaii; Cybiosarda elegans from the body surface could result from the more from western Australia; Auxis sp. from New South exposed site and consequently the copepods more Wales; Euthynnus affinis from New South Wales, easily drop off prior to examination of the host. Cer- northwest Malagasy Republic; Katsuwonus pelamis 26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY from northwest Malagasy Republic; Thunnus alba- our specimens from Sarda orientalis from Hawaii are cares from Queensland. usually distorted, including one specimen with 1 cen- FEMALE.—Body form as in Figure 45a. Range of tral tine rather than the usual 2. In certain other total length 3.00-4.20 mm (3.49 mm average of 22 collections the furcae were normally developed. specimens). Those from Cybiosarda elegans (western Australia) averaged somewhat longer (3.78 mm of 9 Caligus bonito Wilson, 1905 specimens) than those from other hosts from other areas of the Pacific and Indian oceans. Those meas- FIGURES 48-50, 105 ured from other areas and other hosts did not vary significantly. Caligus bonito Wilson, 1905:589. Caligus asymmetricus can be separated from other Caligus sarda Pearse, 1952:17. Caligus species found on scombrids by the combina- Caligus kuroshio Shiino, 1959b: 51. tion of the following characters: abdomen short This species has been reported many times (see (about 10 percent of total length); sternal furca Margolis, Kabata, and Parker, 1975:18 for complete narrow, tines often asymmetrical or otherwise dis- literature history) and has been recently redescribed torted; leg 2 endopod first segment with long spinules by Pillai, 1971:161 and Lewis, 1967:124. Because at outer distal corner, second segment with a row of of these recent good descriptions it is unnecessary to 6-8 stout spines; leg 4 exopod 2-segmented, outermost completely redescribe this species again. We discuss seta longest, other 4 exopod setae all about equal in only certain taxonomic features that we consider im- length to each other. portant in distinguishing this species from other MALE.—Characters described above apply to males Caligus found on scombrid fishes. We have, however, except abdomen comprises about 15 percent of total provided illustrations so the reader can identify speci- body length. mens without consulting other work. REMARKS.—Caligus asymmetricus was reported by MATERIAL EXAMINED.—76 collections containing Pillai (1963:89) from Euthynnus affinis from India, 155 9 and 58 $ from the mouth and gills of the fol- by Kabata (1965:110) from the same host (reported lowing hosts and localities: Euthynnus affinis from as E. alletteratus) from Queensland, Australia, and New South Wales; E. alletteratus from Florida (Gulf by Lewis (1967:131) from the same host (reported coast), Dominican Republic (Atlantic); E. lineatus as E. yaito) from Hawaii. We record this species from from Galapagos Islands, Panama (Pacific), Baja Cali- 9 scombrid species within the Indo-West Pacific. Al- fornia (Pacific) ; Thunnus thynnus from Florida (At- though the records are scattered throughout 3 of the lantic) ; Scomberomorus regalis from Antigua; Sarda 4 tribes of Scombrinae, our collections indicate that sarda from western North Atlantic (several localities Indo-West Pacific species of the tribe Sardini may be Massachusetts to Brazil); Florida (north Gulf coast), preferred hosts (17 percent infestation rate on 3 host Rio de Janeiro, Norway, Tunisia, Gulf of Guinea, species as opposed to 4 percent infestation rate on 2 Angola, South Africa (Port Elizabeth) ; Sarda austra- species of Scomberomorini, 2 percent infestation rate lis from New South Wales; Sarda chiliensis chiliensis on 4 species of Thunnini and no records from the from Peru; Sarda chiliensis lineolatus from Cedros 5 Indo-West Pacific species of Scombrini). Islands, California; Sarda orientalis from Red Sea Kabata (1965:110) notes that both his and Pillai's (Elat), South Africa (Port Elizabeth), India specimens were collected with specimens of Caligus (Cochin), China, Galapagos; Gymnosarda unicolor bonito (reported as C. kuroshio). Of the 22 collections from Bikini. reported here only 2 were accompanied by C. bonito, FEMALE.—Body form as in Figure 48fl. Total length nor did we find any other species common in collec- 6.45 mm. (average of 37 specimens 5.10-8.10 mm tions of C. asymmetricus. range). Measurements taken from collections from We found that the peculiar asymmetry or distortion Sarda sarda, S. australis, and S. chiliensis. No signifi- of the tines of the sternal furca reported previously in cant differences could be seen between C. bonito from the collections from India, Australia, and Hawaii did different host species or geographic areas, except that not occur in all collections but rather seems to occur specimens from warmer water tend to be smaller than randomly within populations. The sternal furcae of those from colder. NUMBER 3 1 1 27

Caligus bonito females can be easily separated from Pearse (USNM 92667) we have placed it in synonymy other species of the genus except Caligus omissus new with Caligus bonito Wilson. species by a combination of the following characters: cephalon, genital segment, and abdomen each com- Caligus mutabilis Wilson, 1905 prising about one-third of the body length (abdomen widest anteriorly), endopod first and second segments FIGURES 51, 52, 53a-b, 105 of leg 2 with a group of prominent spinules at outer Caligus mutabilis Wilson, 1905:573. distal corner and a double row of stouter spinules along outer edge respectively (see Figures 49e,f), tines MATERIAL EXAMINED.—8 collections containing 8 2 of sternal furca (Figure 49c) nearly parallel and blunt from the gills or gill chambers of Scomberomorus tipped, lateral setae of leg 1 exopod last segment with brasiliensis from Brazil and Costa Rica; S. cavalla stout spinules on basal fourth (see Figure 49d), leg from Surinam; S. maculatus from west coast of 4 exopod 2-segmented with terminalmost seta about Florida; Scomber japonicus from Campeche. twice as long as others. See discussion of characters FEMALE.—Body form as in Figure 51a. Total length separating C. bonito from C. omissus new species in 2.78 mm. Greatest width 1.12 mm. Frontal lunules the description of the new species. large, greatest width of lunule more than least distance MALE.—Body form as in Figure 50c. Total lengths between each lunule (307 vs 254 fan). Genital seg- of males are approximately 80 percent that of the ment (Figure 516) longer than wide (986 X 725 /*m), females in any collection. The ranges would be pro- widest posteriorly, posterior corners rounded and only portionately the same as given for the females. Male slightly produced. Abdomen (see Figure 516) incom- as in female except in the following characters: pletely divided into 2 segments, about twice as long abdomen 2-segmented; second antenna (Figure 50d) as wide (681 X 362 fin), posterior ventral surface second segment with 2 rugose areas, claw with 2 short with patches of spinules as in Figure 51c. Caudal processes at tip and a seta on inner margin; maxilliped rami (Figure 51c) longer than wide (159 X 106 /an) (Figure 50«) basal segment with sclerotized bifid knob bearing 6 plumose setae as indicated in figure. on margin opposite tip of claw. Oral area as in Figure 52c. First antenna (Figure DISCUSSION.—We have recorded this species here 5ld) first segment with 27 setae (all but 2 are plu- from 11 scombrid species. From our data Caligus mose), last segment with 13 naked setae. Second bonito seems to be primarily a parasite of scombrids antenna (Figure 51/) with spatulate posterior process, of the tribe Sardini (collected from 7 species) ; to a otherwise of usual form of genus. Maxillae and maxil- lesser extent a parasite of species of Thunnini (from liped as in Caligus bonito. Sternal furca (Figure 52b) 4 species) ; rarely found on members of the Scom- tines slightly divergent, blunt tipped. boromorini (1 species) ; not found on the tribe Leg 1 (Figure 52d) basipod with large patch of Scombrini (Rastrelliger and Scomber). It is re- fine spinules and 2 plumose setae; exopod first seg- corded from all except polar oceans and previous ment with row of spinules on inner edge and short records are cited by Lewis (1967:125) (including spine at outer distal corner, second segment with 3 nonscombrid hosts). terminal spines (2 with accessory processes), a ter- Lewis (1967:131) discussed the difficulties dis- minal seta plumose on outer edge and 3 lateral setae, tinguishing this species from Caligus productus and each with outer edge armed with stout spinules at C. quadratus Shiino. Although these 3 species super- base becoming finer toward tip and short plumosities ficially resemble each other, the nature of the armature along inner edge. Leg 2 (Figure 52«) exopod similar on the first, second, and fourth legs should easily to other species of genus, endopod first segment with separate them. Caligus quadratus is easily separated a patch of long spinules at outer distal corner, second from C. productus by the normally developed setae segment with a dense patch of long spinules along on the leg 1 exopod of C. quadratus (absent in C. outer edge, last segment with 3-4 shorter spinules at productus) and from C. bonito by the stout spinules proximal outer edge, all segments with plumose setae present on these same setae of C. bonito (normal as in figure. Leg 3 (Figure 53a) similar to other plumosities on C. quadratus). species of genus, thumblike spine at outer distal corner After examining the type-specimens of Caligus sarda of exopod first segment only slightly recurved. Leg 4 28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(Figure 536) exopod 2-segmented, first segment with paratypes (USNM 172241) from the gills of S. sierra prominent spine at outer distal corner, second segment from San Juan Lagoon, Mexico (USNM 219623). with 3 terminal and 1 lateral spine, distalmost longest FEMALE.—Body form as in Figure 53c. Total length (setae measure 177, 112, 83, 83, 124 fim distal to 2.96 mm (2.78-308 mm) based on an average of 6 proximal respectively). Leg 5 represented by 3 short specimens from the type series. Greatest width 1.15 setae near posterior corner of genital segment. mm (1.12-1.19). Cephalon comprises about 40 per- MALE.—None collected. cent of total length. Frontal lunules prominent, REMARKS.—This species is apparently common on diameter of lunule (266 /*m) slightly more than a number of host species in the western Atlantic and least distance (218 /*m) between lunules (interlunular Gulf of Mexico (scombrid and non-scombrid). It space). Genital segment (Figure 53d) slightly longer has also been reported from the Gulf of California than wide (1.16 X 1-09 mm), posterior corners and Pacific coast of Mexico from a number of host rounded, only slightly produced, comprising about 30 species by Causey (1960:329) and Wilson (1937:27). percent of total length. Abdomen (see Figure 53d) We describe here a new, closely related, species of 1-segmented, nearly 3 times as long as wide (0.96 X Caligus from eastern Pacific Scomberomorus and it 0.36 mm), usually slightly wider in posterior third seems likely that at least some of the specimens re- but never noticeably wider in anterior half; distal ported by these 2 authors may have been the new ventral surface with patches of spinules as in Figure species. None of that material has been deposited in 53*. Caudal rami (see Figure 53e) longer than wide the Smithsonian collections so we have not been able (142 X 94 fixn) armed with 6 plumose setae as in- to verify the identifications. dicated in figure, longest seta 307 /*m. Examination of the type-specimens (USNM 6155) Oral area as in Figure 54a. First antenna (Figure from Woods Hole, Mass, indicate variation from our 546) first segment with 27 setae, all plumose except specimens on at least 2 points. The outermost of the 2 in middle anterior group; last segment with 13 4 terminal setae on the caudal rami of the type speci- naked setae. Second antenna (Figure 54rf) with mens is less than half the length of the outer edge of prominent hook, basal segment with conspicuous the ramus, whereas in our specimens that seta is pointed posterior process. Post antennal process (Fig- almost as long as the outer edge of the ramus. The ure 54c) a prominent recurved hook. Postoral process longest seta on the exopod of leg 4 of the type speci- (Figure 54e) with slight outward curve, extending mens is only slightly longer than the adjacent seta well beyond tip of mouth tube. Maxilliped (Figure (218 and 177 /*m) whereas in our material this seta 54/) with a strongly sclerotized, recurved claw. is much longer than the adjacent seta (177 in 112 Sternal furca (Figure 54g) with widely divergent /*m). We do not consider these differences important tines, each with a blunt tip. enough to warrant describing a new species but rather Leg 1 (Figure 55a) basipod with a patch of may reflect ecological differences between the 2 groups prominent spinules covering two-thirds of ventral of material. surface; exopod first segment with row of spinules along inner edge and short, naked spine on outer distal corner; exopod with 3 terminal spines (2 with Caligus omissus, new species accessory spines), 1 terminal seta (plumose on outer edge), and 3 lateral setae with stout spinules on base FIGURES 53c-e, 54-56, 105 of outer edge of each followed by short plumosities; MATERIAL EXAMINED.—58 collections containing endopod reduced to a short process with 2 small distal 147 9 41 S from the gills, gill arches, gill rakers, inner spines. Leg 2 (Figure 556) as in other species of the operculum, and roof of the mouth from 41 specimens genus except exopod spine on outer distal corner of (114 examined) of Scomberomorus sierra from Peru, last segment very small; endopod first segment with Colombia, Panama, Costa Rica, Pacific coast of Mex- 3-4 stout spinules on outer distal corner, second seg- ico, and Baja California; from 7 specimens (47 ex- ment with a double row of stout spinules along outer amined) of Scomberomorus concolor from off Sonora, edge, a patch of 4-5 short spinules on proximal outer Mexico and Gulf of California. Holotype (USNM edge of last segment, all segments with plumose setae 172239), allotype (USNM 172240) and 40$ 3* as in figure. Leg 3 (Figure 55c) basipod with con- NUMBER 3 1 1 29 spicuous spinules on outer quarter and a patch of other 2 species as well as all other species of Caligus finer spinules at inner, proximal corner; endopod first found on scombrids by a combination of the follow- segment with stout, recurved claw on outer distal ing characters: abdomen narrowest anteriorly, widely corner, a short spine on outer distal corner of second divergent tines on the sternal furca, distal process segment, 3 short spines on outer edge of last segment on the base of the second antenna pointed, basipod and each segment with plumose setae as in figure; of leg 1 with a large patch of prominent spinules endopod with plumose setae as indicated in figure. (fine spinules in C. mutabilis and C. bonito), the Leg 4 (Figure 55d) exopod 2-segmented, first segment spinules on the outer basal margin of the lateral setae with a prominent spine on outer distal corner, second of leg 1 are fingerlike (short and stout in C. mutabilis segment with one lateral spine and 3 terminal spines, and C. bonito). distalmost spine longest, spines measure 201, 124, 106, Wilson (1937:27) reported C. mutabilis from 100, and 130 /tun distal to proximal spine respectively. Scomberomorus maculatus (=5. sierra) from Mexico Leg 5 represented by 3 short plumose setae near (Pacific). We suspect that this material was actually distal corner of genital segment. the new species reported here. Causey (1960:329) MALE.—Body form as in Figure 56a. Total length reported C. mutabilis from the Gulf of California and 3.15 mm (3.08-3.23 mm), based on an average of Pacific coast of Mexico from several species of fishes 2 specimens. Greatest width 1.27 mm. Cephalon com- including Scomberomorus sierra. It seems likely to us prising about one-half total length. Genital segment that at least the specimens from S. sierra were actually (Figure 566) longer-than wide (884 X 580 /an). C. omissus and some or all of those from the non- Abdomen (Figure 566) 2-segmented, first segment scombrid hosts may have been as well. nearly square (348 X 348 /*m), second segment longer Caligus omissus was collected from 36 percent (41 than wide (449 X 334 /*m) with ventral patches of of 114) of the specimens of S. sierra and 15 percent spinules as in female. Caudal ramus as in female. (7 of 47) of the S. concolor examined. This would in- Appendages as in female except as follows. Post- dicate that these species may well be the preferred antennal process (Figure 56d) a prominent, sharply hosts for this species. Examination of collections from recurved, hook. Second antenna (Figure 56c) with non-scombrid hosts from the Eastern Pacific may well 2 rugose areas on basal segment and a short bifid claw. turn up additional material of this new species. Maxilliped (Figure 56/) basal segment with a pair of sclerotized processes on inner margin opposite tip of Caligus biseriodentatus Shen, 1957 claw. Sternal furca (Figure 56g) with tines less divergent than female. FIGURES 57-59, 105 Legs 1-4 as in female. Legs 5 and 6 each repre- Caligus biseriodentatus Shen, 1957:352. sented by 2 short plumose setae on the genital segment Caligus proboscidatus Heegaard, 1962:161. as in Figure 566. Caligus obovatus Heegaard, 1962:166. ETYMOLOGY.—The Latin omissus ("neglected") Caligus auxisi Pillai, 1963:85. alludes to its common association with eastern Pacific MATERIAL EXAMINED.—1 $ from the inner surface Scomberomorus species but not collected or, if so, of the operculum of Auxis thazard from Trivandrum, not recognized as new. India (a specimen of Caligus auxisi donated to the REMARKS.—This new species seems to be closely Smithsonian by Dr. Pillai) ; holotype (AMS P16418) related to Caligus mutabilis and C. bonito to a lesser of C. obovatus Heegaard from Scomberomorus extent. These 3 Caligus share a number of characters: queenslandicus from Queensland. Holotype (AMS the similar pattern of fine spinule patches on the PI6420) of C. proboscidatus Heegaard from Scom- distal-ventral surface of the abdomen, a prominent beromorus queenslandicus; from Queensland; 237 posterior process arising from the base of the second immature 9 and $ from mouth, gill arches, and antennae, inner lateral setae of leg 1 with spinules inner operculum of the following hosts and localities: along the basal part of the outer edge, prominent, Scomberomorous commerson from northwest Mala- stout spinules along the outer edges of the leg 2 gasy Republic, Somalia, Ceylon, East Indies, Philip- endopod segments, and a 2-segmented exopod of pines, China; Scomberomorus guttatus from Arabian leg 4. The new species can be separated from the Sea, Ceylon, Thailand, East Indies (various locali- 30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY ties), Hong Kong; Scomberomorus queenslandicus MALE.—According to Pillai (1963:87) the second from Onslow (W. Australia), Ambon (Moluccas); antenna basal segment with a bossed pad opposite tip Scomberomorus plurilineatus from Zanzibar. of claw. The postantennal process (first maxilla of FEMALE.—Body form as in Figure 57a. Total length Pillai) a long "sickle-shaped" claw. Maxilliped base 3.15 mm. Greatest width 1.50 mm (measured at with 4 low projections on inner border. Total length widest part of cephalon) ; cephalon comprising about 2.70 mm. 40 percent of total length. Genital segment somewhat IMMATURE FEMALE.—Body form as in Figure 58d. longer than wide (0.90 X 0.75 mm). Abdomen about Total length 2.90 mm (3.00-2.70 mm) based on an twice as long as wide (0.68 X 0.30 mm), 1-seg- average of 5 specimens. Frontal lunules greatest mented; dorsal surface distal half with a large patch width (236 fim) less than least space (interlunular of spinules, ventral surface distal half with groups of space) between them (265 /*m). First antenna typical spinules as in Figure 57b. Caudal rami slightly longer of the genus. Second antenna with strong claw and than wide, each with a seta on middorsal surface and pointed posterior process on base. Postantennal process 6 subterminal to terminal setae. a short recurved hook. Postoral process a sharply Space between frontal lunules (254 fun) greater pointed process, tip not extending beyond tip of than diameter of lunule (207 /un). First antenna of mouth tube. Second maxilla typical of the genus. usual type of genus except last segment about 4 Maxilliped (Figure 58

Shiino later (1954:150) reported it (as C. fulvipur- antenna with posterior process on basal segment. pureus) from the body surface of several Cololabis Postantennal process (first maxilla of Pillai) long saira from Japan and again (Shiino, 1959b:277) and blunt. Sternal furca with widely divergent tines, from off Mexico (Pacific) as C. macarovi. This para- spread of tines wider than base. Leg 1 lateral setae site apparently is common on Pacific Cololabis (also with spinules on basal part of outer edge. Leg 2 reported from this host by Kazachenko, et al, 1972: endopod segments with a patch of narrow spinules 224). It is not surprising that a copepod common on on the outer edge of each, second spine on exopod a prey species of fish (Cololabis) would occasionally with strong teeth on outer edge. Leg 4 exopod 2- be found on its predator species. The female of this segmented, bearing 4 spines (no lateral spine on last species is characterized by the following. Cephalon segment which is usually present in other species when comprises about 40 percent of total length (see Fig- exopod is 2-segmented), distalmost spine about one- ure 73g). Genital segment about as wide as cephalon, third longer than adjacent spine. sides nearly parallel. Abdomen one-segmented, about twice as long as wide. Second antenna with usual claw, posterior process on basal segment. Postantennal Caligus pseudokalumai Lewis, 1968 spine short, angle of inner curve greater than 90° Caligns pseudokalumai Lewis, 1968:59. (see Figure 73h). Maxilliped with 2-segmented claw. Sternal furca (Figure 73*) tines not widely divergent, This species was described from a single female not sharply pointed. Leg 1 basipod with patch of collected from the body surface of a Gymnosarda spinules, 3 inner lateral setae of last segment with unicolor from Eniwetok Atoll. Lewis compared this spinules on outer basal part followed by fringe. Leg species with a previously described species, Caligus 2 endopod segments (Figure 74a) each with a dense kalumai Lewis, 1964, collected from Acanthurus patch of spinules on outer edge. Leg 4 (Figure 74fc) guttatus from Hawaii. The differences between the exopod 2-segmented, last segment without midlateral 2 species are slight and Lewis suggested that their spine, distalmost spine longer than adjacent spine. close relationship should be more closely examined Male as in female except in body form. Second an- when additional material is available. We feel that tenna with short claw and rugose areas on basal this species represents an accidental infestation and segment. Maxilliped with rugose area on basal seg- Gymnosarda unicolor is not its usual host. Lewis has ment opposite tip of claw. provided a good description with illustrations and since this is undoubtedly not a scombrid parasite we We did not collect this parasite during the course will not repeat a description here. of this study. This is undoubtedly due to its rarity on scombrids and its occurrence on the body surface of the host (not usually found on preserved hosts). Elytrophora Gerstaecker, 1853

Elytrophora brachyptera Gerstaecker, 1853 Caligns amblygenitalis Pill a i, 1961 FIGURES 75-77, 106 FIGURE l\c-g Elytrophora brachyptera Gerstaecker, 1853:58. Caligns amblygenitalis Pillai, 1961:98. Elytrophora hemiptera Wilson, 1921:4.

Pillai describted this species from a single female MATERIAL EXAMINED.—78 collections containing collected from the body surface of Euthynnus affinis 771 9 269 $ from the gill area of the following hosts from Vizhingom, India. It has not been recorded and localities: Thunnus alalunga from He Amster- since. dam, Chile, New Jersey, Brazil (north coast), North The female is distinguished by the following char- Atlantic (25°N, 35°W), Azores Islands; T. albacares acters. Frontal lunules widely separated, interlunular from Chagos, Australia (NSW), Christmas Island space about 3 times diameter of lunule (according to (Pacific), Hawaii, New Jersey, Brazil (north coast) ; Pillai's figure). Genital segment about as long as T. atlanticus from St. Thomas Island; T. maccoyii cephalon and with lobed posterior corners. Abdomen from Western Australia (Albany) ; T. obesus from 1-segmented, about twice as long as wide. Second He Amsterdam, Seychelles, Christmas Island (Pacific), NUMBER 3 1 1 37

Hawaii, Juan Fernandez Islands, New Jersey, Brazil part of cephalon). Genital segment (Figure 77«) (north coast), Cape Verde Islands, Canary Islands, longer than wide (1.74 X 1.53 mm). Abdomen 2- Azores Islands; T. thynnus from South Africa (west segmented, segments measure (1 X w) 0.56 X 0.72 coast), Eastern Pacific, Western North Atlantic, New mm, 0.52 X 0.69 mm respectively. Caudal rami simi- Jersey, Portugal (Lisbon); Allothunnus fallai from lar to female. First antenna 2-segmented, similar to New Zealand, California. female. Second antenna (Figure 77/) terminal seg- FEMALE.—Body form as in Figure 75a. Total length ment long, recurved claw with stout spine on inner 8.26 mm, greatest width 4.71 mm (measured at wid- margin; triangular process on basal segment not as est part of cephalon). Genital segment longer than prominent as in female. Remaining cephalic appen- wide (2.61 X 1-89 mm). Abdomen 2-segmented, seg- dages except maxilliped similar to female. Maxilliped ments measure (1 X w) 0.72 X 0.87 mm, 0.68 X modified as in Figure 77g. Sternal furca slender, tines 0.85 mm respectively; first segment partially hidden less divergent than in female. Legs 1-6 similar to in dorsal view by flaps of genital segment. Caudal female, except leg 4 endopod setae longer. rami (Figure 756) longer than wide (680 X 400 /*m), REMARKS.—Examination of Wilson's types of E. each with 6 setae (2 very small). First antenna (Fig- hemiptera indicate that they are actually E. brachyp- ure 75c) 2-segmented. Second antenna (Figure 75d) tera and we have placed his species in synonomy. basal segment with prominent triangular process, Hewitt (1968) made a tentative revision of the genus terminal segment clawlike with 1 short spine and 1 Elytrophora in which he considered E. brachyptera seta. Mouthtube, mandible, post oral process and first to be the only species with 2 additional subspecies maxilla as in Figure 75e; mandible with 12 teeth; {atlantica and indica). We feel that E. indica is a post oral process about as long as mouthtube; first valid species and have indicated it as such below. maxilla with 3 small setae. Second maxilla (Figure Elytrophora brachyptera is a circumglobal species 75/) terminal segment with 2 prominent pectinated found on several species of Thunnini. membranes at about mid-point, terminally with 2 We noticed that material collected from colder long curved, subequal fringed processes. Maxilliped waters (New Zealand and lie Amsterdam) show a (Figure 75g) claw distal half finely grooved. Sternal variation in the armature of leg 4. One of the 4 inner furca (Figure 76a) with stout, widely divergent tines. setae is strongly developed whereas in specimens from Leg 1 (Figure 76b) biramous, endopod first seg- more temperate waters all of these setae are reduced ment without ornamentation, second segment spinu- (compare Figures 77& and c). We did not feel this lose along outer edge, 3 inner setae; exopod one- variation warranted designating the colder water segmented with 3 stout, fringed outer spines, 1 material as a new species. These variant specimens terminal plumose seta, 3 inner plumose setae. Leg 2 were collected from the lie Amsterdam specimen of (Figure 76c) biramous, rami 3-segmented, endopod Thunnus obesus and the southernmost specimens of first and second segments each with spinulose outer Allothunnus. edges, third segment small, with 6 setae; exopod with 4 outer spinulose spines, last segment with 6 setae. Leg 3 (Figure 76d) endopod first segment with inner Elytrophora indica Shiino, 1958 seta, second segment with 2 inner setae, third segment FIGURES 78, 79a,b, 106 small, with 4 setae; exopod as in Figure 77a. Leg 4 (Figure 77b) endopod 2-segmented, first segment Elytrophora indica Shiino, 1958:107. with inner seta, second segment incompletely divided Elytrophora brachyptera indica Shiino.—Hewitt, 1968:124. with one short inner seta and 3 short terminal setae; MATERIAL EXAMINED.—9 collections containing exopod first segment with outer spinulose spine, 128 $ 27 $ from the gill area of Thunnus obesus from second segment similar, with inner seta, third segment He Amsterdam, Seychelles, Christmas Island (Pacific), with 3 outer to terminal spines and 4 short inner Juan Fernandez Islands. setae. Legs 5 and 6 represented by 1 and 3 setae FEMALE.—Body form as in Figure 78a. Total respectively on lateral genital segment. length 10.15 mm, greatest width 5.80 mm (measured MALE.—Body form as in Figure 77d\ Total length at widest part of cephalon). Genital segment (Figure 6.67 mm, greatest width 3.62 mm (measured at widest 78fc) longer than wide (2.97 X 2.20 mm). Abdomen 38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(see Figure 786) first segment crescent-shaped, pro- Islands, Malagasy Republic, He Amsterdam, Christ- duced at posterior corners to envelope anterior half mas Island (Pacific), Washington Island, Hawaii, of second segment; segments measure (1 X w) 1.01 X Socorro Island, Revillagigedo, Puerto Rico, Nicaragua, 1.50 mm, 1.02 X 1.10 mm respectively. Caudal rami North Atlantic (39°N, 41 °W), North Carolina, Bra- longer than wide (0.65 X 0-43 mm) similar to E. zil, Canary Islands, Cape Verde Islands, Azores brachyptera. First antenna (Figure 78billfishes (Istiophoridae). The genus Shiino's records of this copepod on the map (Figure Gloiopotes was revised by the first author (Cressey, 105). 1967).

Gloiopotes Steenstrup and Liitken, 1861 Caligulus Heegaard, 1962

Gloiopotes hygomianus Steenstrup and Liitken, Caligulus longispinosus Heegaard, 1962 1861 FIGURE 80 FIGURE 79*-* Caligulus longispinosus Heegaard, 1962:171. Gloiopotes hygomianus Steenstrup and Liitken, 1861:363. We did not collect this species but we have examined MATERIAL EXAMINED.—27 collections containing the type male deposited in the Australian Museum 77 9 56 $ from the body surface of Acanthocybium (AMS P. 16429). Heegaard stated that a female solandri from the following localities: Seychelles holotype and male allotype were deposited but only NUMBER 311 39 a partially dissected male was in the vial when we T. solandri are based on immature specimens. Each received it from the Australian Museum. Examina- description is based on a single specimen and neither tion of this specimen and Heegaard's figures of the Nunes-Ruivo and Fourmanior or Kurian show egg female lead us to believe that this species was erro- strings in the illustrations (Heegaard does). neously placed in the Euryphoridae by the author. One of our collections of T. cervicornis is from His basis for doing so was the presence of dorsal Pakistan, extending the range of that species to within plates covering the fourth thoracic segment. His fig- the known range of T. cybii. ures do not indicate these and the male specimen does Since we did not collect specimens of T. cybii or not have them. Unfortunately, lacking the female, we immature T. cervicornis we feel it would be premature could not resolve the problem of its true taxonomic to place T. cybii in synonymy. We trust that future position. We hope that additional material of this collection will bear out our suspicions regarding the interesting species will be collected eventually and its synonymy of these 2 species. taxonomic position made clear. We have refigured the male maxilliped (Figure 80c), sternal furca (Figure SOd), leg 1 (Figure 80*), Tuxophorus cervicornis Heegaard, 1962 and leg 2 (Figure 80f). FIGURE 81 b-g The original collection was made from the body surface of Euthynnus affinis (reported as E. alletter- Tuxophorus cervicornis Heegaard, 1962:172. atus) from Howick Island, north Queensland, MATERIAL EXAMINED.—2 collections containing 2 9 Australia. from the body surface of Scomberomorus commerson The total length of the male is 3.35 mm. from Coffs Harbor, New South Wales, Australia, and Karachi, Pakistan. Tuxophorus Wilson, 1908 FEMALE.—Body form as in Figure 81fe. Total length 6.80 mm. Greatest width 3.15 mm. Cephalon slightly Tuxophorus cybii Nunes-Ruivo and Founnanoir, longer than wide (3.10 X 3.15 mm). Thoracic seg- 1956 ment bearing leg 4 with dorso-lateral winglike plates. Genital segment slightly wider than long (2.10 X 2.25 FIGURE 81a mm, including posterior processes) ; outer posterior corners produced. Abdomen 3-segmented. Caudal Tuxophorus cybii Nunes-Ruivo and Fourmanoir, 1956:76. ramus about 6.5 times longer than wide (1.95 X 0.30 Tuxophorus solandri Kurian, 1961:72. mm). Oral appendages as in T. collettei except sternal We did not collect this species but we have repro- furca with pointed tines (Figure 81tf) rather than duced the figure of the female based on the illustra- spatulate. Leg 1 as in T. collettei except spinules on tion of Nunes-Ruivo and Fourmanoir. In 1961 Kurian terminal spines thicker in T. cervicornis. Legs 2-3 described a new species, T. solandri, based on a single as in T. collettei. Leg 4 (Figure 81g) exopod 3-seg- specimen from the body surface of Acanthocybium mented, terminalmost spine shorter (relative to ad- solandri from India. Kurian was apparently unaware jacent spine) than in T. collettei. of the description of T. cybii earlier as he made no MALE.—Unknown. mention of it. Comparisons of the 2 descriptions leave REMARKS.—This species has been reported from no doubt that they represent the same species and, Scomberomorus commerson from northern Australia consequently, we have placed Kurian's T. solandri and Pakistan. See the discussion following the descrip- in synonymy with T. cybii. tion of T. cybii for comparisons with that species. Furthermore, we suspect that T. cervicornis of Hee- gaard (described below) may also be a synonym of T. cybii. The only difference we find between Hee- Tuxophorus collettei, new species gaard's species and T. cybii and T. solandri is the FIGURES 82-84 highly developed processes on the outer distal corners of T. cervicornis (absent in the other 2). This could MATERIAL EX MINED.—Holotype $ (USNM be accounted for if the descriptions of T. cybii and 172250), allotype (USNM 172251), and 13 $ 1 $ 40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY paratypes (USNM 172252 collected from the body inner row of spinules, others bifid and with broad surface of Scomberomorus regalis caught off the Vir- serrations on inner and outer edges, each spine with gin Islands, 8 June 1967 during a cruise of the R.V. a fringe near its base, a short terminal seta and 3 Oregon II. Collected by C. A. Child. lateral plumose setae: endopod a small process near FEMALE.—Body form as in Figure 82a. Total length base of exopod bearing a row of hairs near tip. Leg (including caudal rami) 9.0 mm, greatest width 2 (Figure 83e) basipod unarmed except a prominent (measured at widest part of cephalon) 2.33 mm. fringe on both inner and outer margins; exopod 3- Cephalon somewhat longer than wide (2.55 X 2.33 segmented, first segment with a stout serrate spine, mm). Thoracic segments bearing legs 1-3 fused, in- inwardly directed, on outer distal corner and an inner corporated within posterior lateral lobes of cephalon. seta, second segment armed as first except spine smaller Thoracic segment bearing leg 4 with lateral winglike and more finely serrate, third segment with 2 outer plates, posterior margins of plates reaching to "shoul- spines (terminalmost finely fringe) and 6 plumose ders" of genital segment. Genital segment longer than setae; endopod first segment outer edge with a row wide (1.58 X 1-35 mm) with lateral margins parallel of long spinules and an inner seta, second segment giving segment a boxlike shape; distal corners rounded with 2 inner setae and spinules on outer margins and slightly produced, ventral surface with 2 lateral (smaller than those on first segment), third segment patches of stout spinules (Figure 82b) ; a dorsal, with spinules on outer edge and 6 plumose setae. heavily sclerotized process near each distal corner Third leg (Figure 84a) basipod with patch of spi- (Figure 82a). Abdomen about 6 times longer than nules on outer half and smaller patch of finer spinules wide (3.45 X 0-58 mm) narrowing near mid-margin near inner margin; exopod first segment with stout and near distal end. Caudal rami about 3.8 times spine on outer distal corner and fringe on outer longer than wide (490 X 129 fim), widest proximally; margin, second segment with spine on outer distal 2 short setae on each lateral margin at distal two- corner, inner seta, and patch of long spinules on outer thirds, 1 subterminal inner seta, 3 short, plumose, edge, last segment with 3 outer spines, 4 short termi- terminal setae. nal to inner setae, and long spinules on outer edge; First antenna (Figure 82*/) 2-segmented; first seg- endopod first segment with patch of spinules along ment with 13 short, stout, plumose setae, second seg- outer edge and inner seta, second segment with patch ment with 11 naked setae. Second antenna (Figure of spinules on outer edge and 6 plumose setae. Leg 4 82tf) robust with stout recurved claw, claw with short (Figure 84b) basipod unarmed; exopod 3 segmented, seta near base of longer seta near middle of outer first and second segments each with a fringed spine edge. Mandible of usual caligoid type with 12 teeth on outer distal corner, last segment with 3 fringed at tip. First maxilla represented by 3 short setae spines measuring 82, 112, and 236 fim respectively, lateral to mouthtube near base of post-oral process. each exopod spine with a fringe near its base. No Second maxilla (Figure 83a) long and slender, a patch evidence of legs 5 or 6 could be found. of long hairs (compound setae?) at junction of first Egg strings of usual caligoid type, reaching well and second segment, a palmate fringe near mid- beyond tip of abdomen. margin of second segment; 2 terminal fringed flagella, MALE.—Body form as in Figure 84c. Total length outermost about twice length of inner. Maxilliped (including caudal rami) 6.15 mm, greatest width (Figure 83b) basal segment with a short, stout seta (measured at widest part of cephalon) 2.68 mm. near inner mid-margin; second segment in form of a Cephalon about as wide as long (2.68 X 2.33 mm). recurved claw bearing a short seta near middle. Sternal General morphology of cephalon as in female. Genital furca (Figure 83c) with short, divergent, truncate segment (Figure 84a1) longer than wide (1.3 X 0.9 tines, tines with lateral flanges on basal half; accessory mm), ventral surface with spinules as indicated in processes lateral to furca. figure. Abdomen similar to that of female except only Legs 1-3 biramose. Leg 1 (Figure 83d) basipod 3.7 times as long as wide (2.68 X 0.73 mm), small with posterior seta and plumose seta on outer distal spinules on ventral surface. Caudal rami as in female corner; exopod first segment with row of hairs along (590 X 141 fim). Cephalic appendages as in female posterior border and short spine on outer distal corner; except second antenna (Figure 84e) with large ru- second segment with 3 terminal spines, outermost with gose areas on basal segments and maxilliped with NUMBER 3 1 1 41 small rugose knob on basal segment opposing tip of without setae; second segment with 4 setae, outer dis- claw. talmost stout (not as prominent as in Pseudocyc- Legs 1-4 as in female. Legs 5 and 6 absent. noides), other segments with setae as in the figure. REMARKS.—Of the 5 previously described species Second antenna (Figure 85c) with heavily sclerotized (not including T. tylosuri, which was replaced in the claw, 2 short, stout setae on inner margin. Mouthtube genus Caligus by Pillai, 1961) the new species can (Figure 85d) without ornamentation on labrum. Man- be separated from T. wilsoni Kirtisinghe and T. dible (Figure 85d) with 7 teeth at tip. First maxilla caligodes Wilson by the long, slender caudal ramus (Figure 85«) with 2 stout and 1 weak setae at tip. of the new species (nearly square in T. wilsoni and Second maxilla (Figure 85/) with irregular rows of T. caligodes). It can be separated from T. cybii Nunes- stout spinules along outer edge of distal 2 segments. Ruivo and Fourmanoir, T. solandri Kurian (synony- Maxilliped (Figure 86a) with wide basal articulation, mous with T. cybii), and T. cervicornis Heegaard by basal segment narrows proximally to articulate with the much longer abdomen, the spatulate tines of the claw; claw with bifid tip and stout accessory process sternal furca, and the relatively longer inner seta on near mid-margin. the end of leg 4 (twice as long as other leg 4 setae) Leg 1 (Figure 86b) biramous; basipod with 2 patch- of T. collettei. es of long spinules; exopod 1-segmented, bearing 3 ETYMOLOGY.—This species is named for Dr. Bruce short, stout spines at tip; endopod mostly obscured Collette whose enthusiastic support and collecting by fleshy tissuelike processes from basipod, bearing efforts made this paper considerably more compre- 2 spines and 2 setae at tip. Leg 2 (Figure 86c) basipod hensive than it would have been otherwise. with 3 patches of spinules and well-developed seta at outer distal corner; exopod with 4 stout spines at tip and 3 spinules along outer and distal margins; Pseudocycnus Heller, 1868 endopod with 3 stout, terminal spines, 2 groups of spinules as figured; both rami 1-segmented. Leg 3 Pseudocycnus appendiculatus Heller, 1868 (Figure 86d) basipod with a large patch of spinules; exopod fused with basipod and bearing 3 spines at tip. FIGURES 85-87, 131-136a Leg 4 represented by a single lateral seta at proximal Pseudocycnus appendiculatus Heller, 1868:218. part of genital segment. Egg strings long. Pseudocycnus spinosus Pearse, 1952:30. MALE.—Body form as in Figure 87a. Total length Pseudocycnus thynnus Brandes, 1955:190. 2.70 mm. Greatest width 0.75 mm. Genital segment MATERIAL EXAMINED.—43 collections containing less than half total body length (520 X 370 /xm 85 9 4 $ from the gills of the following hosts and length by width). Abdomen (Figure 87b) longer than localities: Thunnus tonggol from Pakistan, Gulf of wide (300 X 220 fim). Caudal rami (see Figure 81b) Thailand; T. albacares from Seychelles Island, Soma- somewhat longer than abdomen and about 4 times lia, Australia (Queensland), Philippines, Caroline as long as wide (450 X HO [tin) ventral surface with Islands, Kapingamarangi Atoll, Peru, Costa Rica 5 patches of sclerotized knobs. (Pacific), Juan Fernandez Islands, Gulf of Guinea; First antenna (Figure 87c) 6-segmented (fourth T. obesus from Juan Fernandez Islands, North At- segment of male is fused fourth and fifth segments lantic (40°N, 49°W); Katsuwonus pelamis from of female), segment with bulbous spine on outer distal Malagasy Republic, Surinam, Gulf of Guinea; Eu- corner, remaining segments with setae as in the figure. thynnus alletteratus from Florida (West Coast), Gulf Maxilliped (Figure 87d) base with patches of short of Mexico; E. affinis from Malagasy Republic, Gulf spinules on inner margin and a heavily sclerotized of Thailand. claw, a short seta at outer distal corner of penultimate FEMALE.—Body form as in Figure 85a. Thoracic segment. Other cephalic appendages as in female ex- segments bearing legs 2 and 3 free, without lateral cept first maxilla with 1 less seta and second maxilla processes. Abdomen short, incompletely separated from with more spinules at tip. genital segment. Caudal rami long, about one-half Leg 1 (Figure 81 e) basipod with 2 patches of stout length of genital segment, fused with abdomen. First spinules, a seta at outer distal corner, a prominent, antenna (Figure 85b) 7-segmented; first segment heavily sclerotized process arising from inner mid- 42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY margin, process extends to tip of exopod; exopod with (1.05-1.35 mm based on 5 specimens). Genital seg- 1 spine at tip and surface ornamentation as in the ment comprising well over half total body length. figure; endopod an elongate process modified at tip Abdomen and caudal rami fused; each ramus tapering giving it a tubelike appearance (see SEM photo). Leg to a point. First antenna (Figure 88b) 7-segmented, 4 (Figure 87/) a prominent lateral process with a first segment with no ornamentation, second segment single seta at tip. Other legs as in the female. with prominent sclerotized spine at anterodistal corner REMARKS.—This species has been recorded many and other setae as in the figure; all segments with times from scombrids and is apparently circumglobal membranelike covering. Second antenna (Figure 88c) in distribution. Kabata (1970:171) summarized the terminal segment clawlike with prominent medial known hosts and distribution to that time. spine and small proximal seta. Mouthtube (Figure 88a1) tapering posteriorly; labrum with 4 patches of Our collection indicates that this parasite is most bumplike spinules; mandible with 7 teeth. First common on scombrids of the tribe Thunnini and to a maxilla as in P. buccata Wilson. Second maxilla lesser extent Sardini. The 6 species of Thunnini in- (Figure 88^) basal segment with large patch of fested were parasitized at a rate of nearly 10 percent spinules; second segment elongate, curved distally, and the single species of Sardini about 2 percent. together with short third segment forming a claw with Females of this species show considerable variation rows of numerous stout spines along outer distal edge. in total length. Lewis, et al. (1969) have presented Maxilliped (Figure 89a) basal segment enlarged, data based on hosts and locality. Total lengths range visible from dorsal aspect; terminal segment forming from 11-18 mm with the longer specimens from stout, heavily sclerotized claw, darkened and rugose at colder waters. tip with small accessory spine on inner curve. Leg 1 (Figure 89fc) biramous, rami not distinctly Pseudocycnoid.es Yamaguti, 1963 separate from basipod; coxopod with small patch of slender spinules and triangular process on outer distal Pseudocycnoides armatus (Bassett-Smith, 1898) corner; exopod 1-segmented with 2 terminal spines; endopod 1-segmented with medial inner spine and 4 FIGURES 88-91a-fe, 107, 1366-/, 137 terminal spines, innermost shortest. Leg 2 (Figure Hellaria armata Bassett-Smith, 1898a: 10. 89c) basipod with patches of slender spinules and Cybicola armata.—Bassett-Smith, 1898b:371 stout outer spine; rami 1-segmented, exopod with 4 Pseudocycnus armatus.—Kirtisinghe, 1935:339. terminal spines, outermost stout, toothed along outer Paracycnus lobosus Heegaard, 1962:182. edge; endopod with patches of slender spinules and Pseudocycnoides armatus.—Yamaguti, 1963:172. 3 terminal spines. Leg 3 (Figure 89a1) uniramous, Pseudocycnoides rugosa Kensley and Grindley, 1973:104. ramus with slender spinules and 3 terminal spines, MATERIAL EXAMINED.—31 collections containing outermost spine stout, toothed along outer edge. Leg 4 149 5 11 <$ from the following hosts and localities: represented by a simple seta on anterolateral genital Scomberomorus commerson from Red Sea (Suez), segment. Egg strings usually as long as body. Malagasy Republic, India (Cochin), Pakistan, Sri MALE.—Body form as in Figure 89e. Total length Lanka, Gulf of Thailand, Australia (NSW and 2.39 mm. Greatest width 0.59 mm. Genital segment Queensland), Philippines, Hong Kong; S. guttatus more than half total body length. Abdomen (Figure from Borneo (Padang), Hong Kong; S. semifasciatus 90a) without ornamentation. Caudal rami (see Figure from New Guinea; S. lineotatus from India (Palk 90a) with numerous spinules near tips and 5 setae. Strait) ; S. koreanus from India (Bombay), Sumatra. First antenna (Figure 90fe), second antenna, mouth- China; S. queenslandicus from Papua New Guinea; tube, mandible, and first maxilla as in female. Second S. plurilineatus from South Africa (Natal). maxilla (Figure 90c) similar to female, but spines on FEMALE.—Body form (Figure 88a) elongate with last 2 segments more numerous and more closely winglike process on each side of second free thoracic spaced. Maxilliped (Figure 90^) basal segment with segment and anteriormost genital segment (fused with 3 patches of small spinules and process on inner edge; third thoracic segment). Total length 8.4 mm (7.8- distal 2 segments clawlike, each segment with 1 8.7 mm based on 5 specimens). Greatest width 1.2 mm accessory spine. NUMBER 3 1 1 43

Leg 1 (Figure 90*) biramous, each ramus 1-seg- and heavily spinulose (longest spine on endopod of mented and not distinctly separate from basipod; P. armatus much shorter and with few spinules). Leg basipod with patch of slender spinules, triangular 3 (Figure 91 g) outer spine not as robust as in P. process, and several lobate processes; exopod with armata. dense outer patch of spinules, smaller spinules in semi- MALE.—Unknown. circle and 2 medial lobes, 2 spines and 3 lobed REMARKS.—This species is apparently restricted to processes distally; endopod produced as a slender, Scomberomorus niphonius, which is endemic to waters elongate, heavily sclerotized spine with several small of North China and Japan. Yamaguti described this distal processes, spine with membranelike covering. parasite from S. chinensis (probably S. niphonius) Leg 2 (Figure 91a) biramous, each ramus 1-seg- from the Inland Sea of Japan. mented; basipod with dense inner patch of spinules, Pseudocycnoides scomberomori is closely related to 2 lighter patches of spinules and long, slender outer P. armatus but easily separated from it by the char- spine; exopod with irregular distal row of short acters cited above. spinules, 4 terminal spines, outermost spine enlarged, toothed along outer edge, row of hairs near bases of 3 smaller, inner spines; endopod with irregular patches Pseudocycnoides buccata (Wilson, 1922), of spinules, 1-2 rows of hairs at bases of 3 terminal new combination spines, innermost spine elongate, densely spinulose FIGURES 92-94a-c, 107, 138, 139 around entire distal three-fourths, tapering gradually. Leg 3 (Figure 916) similar to female. Legs 4 and 5 Pseudocycnus buccata Wilson, 1922:79. Cybicola elongate Pearse, 1951:365. represented by simple spines on lateral genital segment. Pseudocycnopsis buccata (Wilson).—Yamaguti, 1963:172. REMARKS.—This species is common on species of Scomberomorus throughout the Indo-West Pacific MATERIAL EXAMINED.—125 collections containing (except S. niphonius). We examined the type material 667 5 10 <$ from the gill filaments of following hosts of P. rugosa Kensley and Grindley and concluded that and localities: Scomberomorus sierra from Peru, Co- their material was actually P. armatus. Their conclu- lombia (Pacific), Panama (Pacific), Costa Rica (Pa- sion that P. rugosa represented a new species was cific), Gulf of California, Baja California; S. concolor based on comparisons with poor literature descriptions from Gulf of California, California; S. regalis from of P. armatus (B. Kensley, pers. comm.). We place Brazil (north coast), Surinam, Gulf of Venezuela, P. rugosa in synonymy with P. armatus. Virgin Islands, Haiti, Cuba, Florida (Atlantic) ; S. maculatus from Mexico (Vera Cruz), Florida (west coast), Alabama, United States east coast (several Pseudocycnoides scomberotnori (Yamaguti, 1939) localities from Florida to Cape Cod); S. brasiliensis FIGURES 91 c-g from Brazil (Rio de Janeiro to north coast), Trinidad, Panama (Atlantic), Costa Rica, British Honduras, Pseudocycnus scomberomori Yamaguti, 1939:457. Mexico (Mazatlan) ; S. cavalla from Brazil, Trinidad, Pseudocycnoides scomberomori.—Yamaguti, 1963:172. Cuba, Gulf of Mexico, Cape Cod. MATERIAL EXAMINED.—7 collections containing 22 9 FEMALE.—Body from as in Figure 92a. Total from the gills of Scomberomorus niphonius from length 5.14 mm. Greatest width 1.20 mm. Thoracic Korea, North China, Japan. segments bearing legs 2 and 3 free. Genital segment FEMALE.—Body form as in Figure 9It. Total length comprising about 60 percent total body length; ventral 4.8 mm. Greatest width 0.9 mm. Abdomen and caudal surface with a patch of spinules between bases of rami of about equal length (0.6 mm) ; abdomen much leg 3, 3 patches of spinules between leg 4 setae, and shorter in P. armatus. Appendages as in P. armatus another patch distal to leg 4 setae. Considerable vari- except as described below. Maxilliped (Figure 91 d) ation of total length occurs in this species and this claw longer and tip more recurved. Leg 1 (Figure 9\e) will be considered in a separate study by the first basipod with large patch of spinules (spinules on outer author and B. B. Collette. Abdomen (Figure 92a) edge only in P. armatus). Leg 2 (Figure 91/) endopod somewhat wider than long (652 X 493 /Am). Caudal longest spine extending beyond tip of exopod spines rami about twice as long as wide (362 X 188 /mi), 44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY bearing 2 short terminal setae. Entire body covered patches of spinules and spinulose lobes in distal third, with dense, fine hairs. First antenna (Figure 92c) 2 short spines at tip; endopod modified as a curved, 7-segmented, no ornamentation on first segment, re- heavily sclerotized process, lobed at tip and most of maining segments with short, naked setae as indicated its length enclosed in a membrane (compare with in the figure. Second antenna (Figure 92d) a simple P. armatus). Leg 2 similar to P. buccata except claw with 2 short, stout setae on inner edge of claw. exopod inner 3 spines smaller and distal half with Mouth tube (Figure 92e) without surface ornamenta- widely spaced spinules; endopod with 2 distinct tion. Mandible with 6-7 teeth. First maxilla (Figure patches of spinules. Leg 3 (Figure 94d) reduced to a 92/) bearing 3 setae at tip. Second maxilla (Figure spinulose process with a short spine at tip. 92g) similar to that of P. armatus except no patch REMARKS.—Yamaguti (1963:172) removed this of spinules on basal segment. Maxilliped (Figure species from Pseudocycnus in which it had been placed 93a) with a short, heavily sclerotized claw (Figure by Wilson. He designated the new genus Pseudo- 93b), basal segment with a thumblike process at inner cynopsis for this species and another new genus Pseu- proximal corner as in P. armatus. docycnoides to include P. armatus and also P. Legs 1-3 rudimentary. Leg 1 (Figure 93c) rami scomberomori (Yamaguti), both originally described 1-segmented; exopod with 5 short terminal spines; as species of Pseudocycnus. We agree with Yamaguti endopod bearing a short terminal spine; both rami that these 3 species should be removed from Pseudo- covered with dense hairs and difficult to study. Leg 2 cycnus but we do not feel that placing P. buccata in (Figure 93c) rami 1-segmented; basipod with a well- a genus separate from P. armatus and P. scombero- developed seta at outer distal corner; exopod bearing mori is justified. Consequently we have placed this 4 terminal spines; endopod with 1 easily visible spine, species in the genus Pseudocycnoides. The original 2 very short spines, and spinules as in the figure. Leg 3 description of P. buccata was superficial and inaccu- (Figure 93c) reduced to a short knob bearing a short rate and was undoubtedly the basis for Yamaguti's spine. Leg 4 represented by a lateral seta on anterior conclusions. Our studies of these 3 species indicate genital segment. them to be much more closely related than previously Egg string long. known. These conclusions are based on the compara- MALE.—Body form as in Figure 93d. Total length tive morphology, not only of the female, but first 1.88 mm. Greatest width 0.55 mm. Thoracic segments descriptions of males of P. buccata and P. armatus as bearing legs 2 and 3 incompletely separated. Genital well. It should be pointed out that the first pair of segment about twice as long as wide (1.04 X 0.53 legs in all 3 of these species is very small, between mm) ; abdomen (Figure 93e) nearly twice as wide as the bases of the maxillipeds, and not easily seen. long (259 X 147 /xm) with a ventral patch of spinules This species is found on all American species of near each outer distal corner. Caudal rami (see Fig- Scomberomorus (western Atlantic and eastern Pacific) ure 93e) about twice as long as wide (123 X 64 fim), and is relatively common on all its hosts. bearing 2 prominent lateral setae, 2 small lateral setae Examination of the type-specimen of Cybicola and a terminal spine, ventral surface covered with elongata Pearse (USNM 88538) leaves no doubt that spinules. First antenna as in female. Second antenna it is the same as Wilson's species. The host for Pearse's (Figure 93/) similar to female except claw longer and material was Scomberomorus maculatus. more pointed. Mouth tube (Figure 94a) with 3 The genus Pseudocycnopsis Yamaguti should be patches of spinules on lab rum. First maxilla (Figure considered a synonym of Pseudocycnoides Yamaguti. 94a) with 2 stout setae at tip, setae relatively longer than in female. Second maxilla as in S. armatus except Lernanthropus Blainville, 1822 no spinules on basal segment. Maxilliped (Figure 94fc) with 2-segmented claw and spinules on proximal inner Lernanthropus kanagurta Tripathi, 1962 corner of base. FIGURE 94«-g Leg 1 (Figure 94c) biramous, each ramus 1-seg- mented and not distinctly articulated with basipod; Lernanthropus kanagurta Tripathi, 1962:194. basipod with a patch of spinules and a triangular MATERIAL EXAMINED.—1 collection containing 1 9 process and a seta at outer distal corner; exopod with from the gills of Rastrelliger brachysoma from Borneo. NUMBER 311 45

FEMALE.—Body form as in Figure 94e. Total length commerson from Australia (New South Wales) and 3.23 mm. Greatest width 1.35 mm. Cephalon about S. sinensis (gills) from Hong Kong. one-fourth total body length (not including posterior The original description was based on a collection processes) and only about one-half greatest width. The from the gills of S. commerson from off Queensland, single specimen was somewhat mutilated so we were Australia. Considering the number of specimens of the not able to examine all of the appendages. The form 2 hosts examined (121) it is apparently uncommon on of the second antenna and leg 1 are as in Figures 94/ and known only from west Pacific Scomberomorus. and 94g. Kabata has provided a good description with illus- REMARKS.—When Tripathi described this copepod trations. he reported it from Megalaspis cordyla (Linnaeus) from India. Since it seems to be rare on species of Rastrelliger it may be that Megalaspis (Carangidae) Clavellisa Wilson, 1915 or some other host is preferred. Clavellisa scombri (Kurz, 1877)

Brachiella Cuvier, 1830 FIGURES 95C, 108 Anchorella scombri Kurz, 1877:403. Brachiella thynni Cuvier, 1830 Clavella scombri.—Brian, 1906:116. Clavellisa scombri.—Wilson, 1915:694. FIGURES 95a, 108 Brachiella thynni Cuvier, 1830:257. MATERIAL EXAMINED.—10 collections containing Thynnicola ziegleri Miculicich, 1904:47. 17 9 from the following hosts and localities: Scomber japonicus from Philippines, Peru, Gulf of Mexico MATERIAL EXAMINED.—58 collections containing (Campeche and Alabama), Sierra Leone, Liberia; 114 9 from the axil of the pectoral fin of the follow- Scomber australasicus from Taiwan, Australia. ing hosts and localities: Acanthocybium solandri from This species was originally described from material Malagasy Republic, He Amsterdam, Seychelles, Palau collected from the gills of Scomber scombrus from Island, Yap Island, Kapingamarangi Atoll; Raroia Trieste. Our material was collected from the other 2 Island, Line Islands, California, Nicaragua (Atlantic), species of Scomber. Since we examined 97 specimens Campeche, Brazil (north coast), Surinam, Azores; of the type host without recovering this species we Thunnus albacares from Christmas Island (Pacific), wonder if perhaps the earlier hosts were misidentified. Juan Fernandez Islands, off New Jersey, Azores; Whatever the case, it seems apparent that this species Thunnus obesus from Seychelles, Christmas Island is restricted to species of Scomber and is found (Pacific), Hawaii, mid-North Atlantic (39°N, 44°W), wherever its hosts are. Later records by Yamaguti Azores; Thunnus thynnus from east coast of United (1939) and Shiino (1959c) cite Scomber japonicus as States, Portugal; Scomberomorus regalis from Lee- the host species. Yamaguti (1939) redescribed this ward Islands. species and provided good illustrations. He inadver- This species has been reported many times from tently omitted this species in his 1963 synoptic survey scombrids and other hosts. It is usually attached to its of fish copepods. host in the axil of the pectoral fin. This copepod is easily recognized and has been well described (both sexes) by Shiino (1956:238) and we will not repeat Clavellopsis Wilson, 1915 a description here. Clavellopsis saba Yamaguti, 1939

Brachiella magna Kabata, 1968 FIGURES 95d, 108

FIGURES 95b, 108 Clavellopsis saba Yamaguti, 1939:558. Brachiella magna Kabata, 1968b;508, fig. 2a-j This species was described from Japan from MATERIAL EXAMINED.—3 collections containing 3 9 Scomber japonicus and reported again by Shiino from the gills and nasal lamellae of Scomberomorus (1959c: 367) from the same host from the Sea of 46 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Japan (Tsunodayama). During our studies we ex- copepods are listed with each host in descending order amined 11 specimens of S. japonicus from Japan but of frequency. did not recover this copepod. Shiino (1959c) has provided a good description SCOMBRINI with illustrations. Rastrelliger kanagurta (108) Pumiliopes capitulates (11) Pennella Oken, 1816 Nothobomolochus kanagurta (4) Rastrelliger brachysoma (33) Pennella species Lernanthropus kanagurta (1)

Members of this genus have been reported many Rastrelliger faughni (14) Pumiliopes capitulatus (2) times from large marine pelagic animals. It is easily Nothobomolochus kanagurta (2) recognized attached to the body of the host; the head Orbitacolax aculeatus (2) buried in the muscle tissue and the long wormlike Scomber scombrus (97) body trailing free. Specimens have been recorded Caligus pelamydis (1) several inches long. Scomber japonicus (500) This genus is in need of revision and species names Pumiliopes capitulatus (8) should be used with caution. At least 6 species have Clavellisa scombri (9) been reported from whales, 7 from flyingfishes, and 4 Caligus pelamydis (4) from swordfish. A revision would undoubtedly reduce Caligus mutabilis (1) these numbers of Pennella species. Clavelopsis saba We have collected Pennella from Acanthocybium Scomber australasicus (17) solandri and Thunnus thynnus. Because of the con- Clavellisa scombri (1) fusion surrounding the taxonomy of this genus we Pumiliopes capitulatus (1) have not assigned species names to our specimens. SCOMBEROMORINI

Summary Acanthocybium solandri (61) Brachiella thynni (37) Our data, relative to infestation rates, are subject Gloiopotes hygomianus (26) to considerable bias because of variables in collecting Caligus productus (9) Caligus coryphaenae (1) methods throughout the course of the study. Much of Shiinoa occlusa (1) the material was collecetd from preserved fish in Pennella species (1) museum collections. In these cases copepods usually Grammatorcynus bicarinatus (37) found on the body surfaces of the hosts are lost. Con- Shiinoa occlusa (5) sequently, rate of infestation data for each species, Caligus asymmetricus (4) based on data from the entire collection, is biased. Scomberomorus brasiliensis (62) Nevertheless, we have included infestation rates in Pseudocycnoides buccata (39) spite of this, as we feel that they do reflect, in a gen- Holobomolochus divaricatus (14) eral way, host preferences and relative common-to- Caligus mutabilis (4) rare occurrences of the parasite species on their hosts. Shiinoa inauris (3) Selected collections will be considered in more detail Scomberomorus cavalla (36) in the joint work with Collette. Pseudocycnoides buccata (18) The following is a list of scombrid fishes with the Holobomolochus asperatus (10) Caligus mutabilis (2) parasitic copepods collected from them during the Caligus productus (1) course of this study. The number in parentheses after the host name indicates the number of individuals ex- Scomberomorus commerson (113) Pseudocycnoides armatus (20) amined. The number in parentheses after the copepod Unicolax ciliatus (20) name indicates the number of fish infested. The Shiinoa occlusa (11) NUMBER 3 1 1 47

Caligus biseriodentatus (11) Scomberomorus sinensis (8) Caligus cybii (10) Caligus cybii (2) Caligus infestans (5) Brachiella magna (1) Caligus asymmetricus (2) Brachiella magna (2) Scomberomorus tritor (21) Tuxophorus cervicornis (2) Unicolax ciliatus (4) Shiinoa occlusa (1) Scomberomorus concolor (47) Caligus productus (1) Pseudocycnoid.es buccata (14) Caligus diaphanus (1) Holobomolochus nudiusculus (13) Caligus omissus (7) Scomberomorus queenslandicus (19) Caligus biseriodentatus (11) Scomberomorus guttatus (55) Pseudocycnoides armatus (2) Caligus biseriodentatus (16) Unicolax ciliatus (1) Unicolax ciliatus (13) Shiinoa occlusa (1) Pseudocycnoides armatus (2) Shiinoa occlusa (1) Scomberomorus species (3) Unicolax ciliatus (1) Scomberomorus koreanus (17) Caligus productus (1) Caligus cybii (11) Caligus cybii (1) Pseudocycnoides armatus (3)

Scomberomorus lineolatus (10) Pseudocycnoides armatus (1) SARDINI Unicolax ciliatus (1) Caligus biseriodentatus (1) Sardasarda (104) Caligus bonito (23) Scomberomorus maculatus (76) Ceratacolax euthynni (21) Pseudocycnoides buccata (27) Caligus pelamydis (7) Holobomolochus divaricatus (24) Caligus productus (1) Shiinoa inauris (7) Caligus mutabilis (1) Sar

Scomberomorus semifasciatus (17) Cybiosarda elegans (8) Pseudocycnoides armatus (2) Caligus asymmetricus (2) Unicolax ciliatus (2) Unicolax collateralis (1) Caligus cybii (1) Orcynopsis unicolor (6) Scomberomorus sierra (116) Unicolax collateralis (1) Pseudocycnoides buccata (48) Caligus omissus (39) Allothunnus fallot (3) Holobomolochus nudiusculus (28) Elytrophora brachyptera (3) 48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

THUNNINI Thunnus maccoyii (2) Elytrophora brachyptera (2) Auxis species (65) Unicolax collaterals (19) Thunnus obesus (40) Unicolax mycterobius (9) Elytrophora brachyptera (20) Caligus productus (2) Caligus coryphaenae (18) Caligus asymmetricus (1) Brachiella thynni (10) Caligus coryphaenae (1) Elytrophora indica (9) Caligus pelamydis (1) Pseudocycnus appendiculatus (3)

Euthynnus affinis (71) Thunnus thynnus (57) Unicolax collateralis (29) Caligus coryphaenae (16) Caligus asymmetricus (5) Caligus productus (16) Caligus re gaits (5) Elytrophora brachyptera (11) Pseudocycnus appendiculatus (4) Pennella species (3) Unicolax mycterobius (3) Brachiella thynni (2) Caligus pelamydis (2) Caligus bonito (1) Caligus productus (1) Pseudocycnus appendiculatus (1) Caligus bonito (1) Thunnus tonggol (26) Euthynnus alletteratus (64) Pseudocycnus appendiculatus (7) Caligus coryphaenae (9) Unicolax collateralis (8) Below is a list of the copepods and the fishes on Ceratacolax euthynni (7) which they were found. The number in parentheses Caligus productus (5) after the host name indicates the infestation rate Caligus bonito (4) (number of infested fish/number of fish examined). Pseudocycnus appendiculatus (3) We remind the reader of the bias in the infestation Unicolax mycterobius (3) Unicolax anonymous (2) rates discussed above. No number after a fish name Caligus pelamydis (1) indicates the record is from the literature.

Euthynnus lineatus (15) Unicolax collateralis (4) BOMOLOCHIDAE Caligus bonito (3) Ceratacolax euthynni (2) Holobomolochus divaricatus Unicolax mycterobius (1) Scomberomorus maculatus (32) Scomberomorus regalis (29) Katsuwonus pelamis (132) Scomberomorus brasiliensis (23) Caligus coryphaenae (51) Caligus productus (51) Holobomolochus nudiusculus Pseudocycnus appendiculatus (7) Scomberomorus concolor (28) Unicolax reductus (3) Scomberomorus sierra (24) Caligus asymmetricus (1) Holobomolochus asperatus Thunnus alalunga (12) Scomberomorus cavalla (28) Elytrophora brachyptera (7) Caligus coryphaenae (1) Unicolax collateralis Caligus productus (1) Euthynnus affinis (41) Pseudocycnus appendiculatus (1) ^4uxu species (29) Euthynnus lineatus (27) Thunnus albacares (110) Sarda orientalis (21) Caligus productus (49) Orcynopsis unicolor (17) Elytrophora brachyptera (39) Cybiosarda elegans (13) Caligus coryphaenae (32) Euthynnus alletteratus (13) Pseudocycnus appendiculatus (21) Brachiella thynni (8) Unicolax anonymous Caligus asymmetricus (1) Euthynnus alletteratus (3)

Thunnus atlanticus (76) Unicolax mycterobius Caligus productus (70) Auxis species (14) Caligus coryphaenae (9) Euthynnus alletteratus (5) Elytrophora brachyptera (1) Euthynnus affinis (4) NUMBER 311 49

Unicolax ciliatus Caligus productus Scomberomorus gut talus (24) Thunnus atlanticus (92) Scomberomorus tritor (19) Thunnus albacares (45) Scomberomorus commerson (18) Katsuwonus pelamis (39) Scomberomorus semifasciatus (12) Thunnus thynnus (28) Scomberomorus niphonius (11) Acanthocybium solandri (15) Scomberomorus lineolatus (10) Sarda orient alls (13) Euthynnus alletteratus (8) Unicolax reductus Thunnus alalunga (8) Katsuwonus pelamis (2) Scomberomorus regalis (8) Ceratacolax euthynni Scomberomorus tritor (5) Sarda sarda (20) Scomberomorus cavalla (3) Euthynnus alletteratus (14) Auxis species (3) Sarda chiliensis lineolatus (2) Nothobomolochus kanagurta Euthynnus affinis (1) Rastrelliger faughni (14) Sarda sarda (1) Rastrelliger kanagurta (4) Scomberomorus species Rastrelliger species Caligus asymmetricus Orbitacolax aculeatus Cybiosarda elegans (25) Rastrelliger faughni Sarda orientalis (21) Grammatorcynus bicarinatus (11) Pumiliopes capitulatus Sarda australis (10) Rastrelliger faughni (14) Euthynnus affinis (7) Rastrelliger kanagurta (10) ^4uxu species (2) Scomber japonicus (2) Scomberomorus commerson (2) Scomber scombrus (1) Katsuwonus pelamis (1) Thunnus albacares (1)

SHIINOIDAE Caligus bonito Sarda australis (65) Shiinoa inauris Sarda chiliensis chiliensis (54) Scomberomorus regalis (13) Sarda chiliensis lineolatus (35) Scomberomorus maculatus (9) Sarda orientalis (39) Scomberomorus brasiliensis (5) Euthynnus lineatus (20) Gymnosarda unicolor (14) Shiinoa occlusa Euthynnus alletteratus (6) Grammatorcynus bicarinatus (14) Scomberomorus regalis (3) Gymnosarda unicolor (14) Thunnus thynnus (2) Scomberomorus commerson (10) Euthynnus affinis (1) Scomberomorus niphonius (6) Scomberomorus queenslandicus (5) Caligus mutabilis Scomberomorus guttatus (1) Scomberomorus brasiliensis (6) Scomberomorus tritor (1) Scomberomorus cavalla (6) Acanthocybium solandri (1) Scomberomorus maculatus (1) Scomber japonicus (1) Caligus omissus CALIGIDAE Scomberomorus sierra (34) Scomberomorus concolor (15) Caligus coryphaenae Thunnus obesus (45) Caligus biseriodentatus Katsuwonus pelamis (38) Scomberomorus queenslandicus (58) Thunnus albacares (29) Scomberomorus guttatus (29) Thunnus thynnus (28) Scomberomorus commerson (10) Euthynnus alletteratus (14) Scomberomorus lineolatus (10) Thunnus atlanticus (12) species Thunnus alalunga (8) Auxis species (1) Caligus cybii Scomberomorus koreanus (65) Caligus regalis Scomberomorus sinensis (25) Euthynnus affinis (70) Scomberomorus species (20) 50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Scomberomorus commerson (9) Tuxophorus collettei Scomberomorus semifasciatus (6) Scomberomorus regalis (3)

Caligus pelamydis Sarda australis (55) PSEUDOCYCNIDAE Scomberomorus niphonius (11) Sarda chiliensis lineolatus (10) Pseudocycnus appendiculatus Sarda sarda (7) Thunnus tonggol (27) Euthynnus affinis (3) Thunnus albacares (19) Euthynnus alletteratus (2) Thunnus obesus (8) Auxis species (2) Euthynnus affinis (6) Scomber japonicus (1) Euthynnus alletteratus (5) Scomber scombrus (1) Katsuwonus pelamis (5)

Caligus infestans Pseudocycnoides armatus Scomberomorus commerson (4) Scomberomorus plurilineatus (100) Scomberomorus koreanus (18) Caligus diaphanus* Scomberomorus commerson (18) Scomberomorus tritor (5) Scomberomorus semifasciatus (12) Scomberomorus queenslandicus (11) Caligus savala* Scomberomorus lineolatus (10) Euthynnus affinis Scomberomorus guttatus (4)

Caligus macarovi* Pseudocycnoides scomberomori Auxis species Scomberomorus niphonius (33)

Caligus amblygenitalis* Pseudocycnoides buccata Euthynnus affinis Scomberomorus brasiliensis (63) Scomberomorus cavalla (50) Caligus pseudokalumai Scomberomorus sierra (41) Gymnosarda unicolor Scomberomorus maculatus (36) Scomberomorus regalis (32) Scomberomorus concolor (30) EURYPHORIDAE

Elytrophora brachyptera LERNANTHROPIDAE Allothunnus fallal (100) Thunnus maccoyii (100) Lernanthropus kanagurta* Thunnus alalunga (58) Rastrelliger brachysoma (3) Thunnus obesus (50) Thunnus albacares (35) Thunnus thynnus (19) LERNEOPODIDAE Thunnus atlanticus (1) Brachiella thynni Elytrophora indica Acanthocybium solandri (61) Thunnus obesus (23) Thunnus obesus (25) Thunnus albacares (7) Gloiopotes hygomianus Thunnus thynnus (4) Acanthocybium solandri (43) Scomberomorus regalis (3) Caligulus longispinosus Brachiella magna Euthynnus affinis Scomberomorus sinensis (13) Scomberomorus commerson (2)

TUXOPHORIDAE Clavellisa scombri Scomber australasicus (6) Tuxophorus cybii Scomber japonicus (2) Scomberomorus commerson Clavellopsis saba Acanthocybium solandri Scomber japonicus Tuxophorus cervicornis Scomberomorus commerson (2) * Probably not usually scombrid parasites. Literature Cited

Bassett-Smith, P. W. Gerstaecker, A. 1896. Notes on the Parasitic Copepoda of Fish Obtained 1853. Ueber eine neue und eine weniger gekannte at Plymouth, with Descriptions of New Species. Siphonostomen-Gattung. Archiv fur naturges- Annals and Magazine of Natural History, 1(6): chichte, 19:58-70. 8-16. Gnanamuthu, C. P. 1898a. Some New Parasitic Copepods Found on Fish at 1948. Notes on the Anatomy and Physiology of Caligus Bombay. Annals and Magazine of Natural His- savala n. sp., a Parasitic Copepod from Madras tory, 1(7):1-17. Plankton. Proceedings of the Zoological Society, 1898b. Some New or Rare Parasitic Copepods Found on 118(3):591-606. Fish in the Indo-Tropic Region. Annals and Gussev, A. B. Magazine of Natural History, 11(7):359-372. 1951. Parasitic Copepoda of Some Marine Fishes. Col- Brandes, C. H. lected Papers on Parasitology from Zoological 1955. Uber eine neue Art der Parasitischen Copepoden: Institute, Academy of Sciences SSSR, 13:394-463. Pseudocycnus thynnus n. sp. Veroffentlichungen Heegaard, P. des Instituts fur Meeresforschung in Bremerhaven, 1943. Parasitic Copepods Mainly from Tropical and 3:190-198. Antarctic Seas. Arkiv for Zoologi, 34A(18): 1-37. Brian, A. 1962. Parasitic Copepoda from Australian Waters. Rec- 1906. Copepodi Parassiti dei Pesci a"Italia. 187 pages. ords of the Australian Museum, 25(9): 149-233. Genoa, Italy. 1964. New Names for Caligus dentatus and Hatschekia Causey, D. elongata. Crustaceana, 6:319-320. 1960. Parasitic Copepoda from Mexican Coastal Fishes. Heller, C. Bulletin of Marine Science of the Gulf and Car- 1868. Crustaceen Reise der Osterreichischen Fregatte ibbean, 10:323-337. Novara um die erde in den Jahren 1857, 1858, Collette, B. B., and L. Chao 1859. Zoologischer Theil, 2(3): 1-280. Hewitt, G. C. 1975. Systematics and Morphology of the Bonitos 1968. Elytrophora brachyptera Gerstaecker (Euryphori- (Sarda) and Their Relatives (Scombridae, Sar- dae, Caligoida) from New Zealand Waters, with dini). Fishery Bulletin, 73(3) :516-625. a Tentative Revision of the Genus. Transactions Cressey, R. F. of the Royal Society of New Zealand, Zoology, 1967. Genus Gloiopotes and a New Species with Notes 10(12):117-126. on Host Specificity and Intraspecific Variation Kabata, Z. (Copepoda: Caligoida). Proceedings of the United 1965. Copepoda Parasitic on Australian Fishes, IV: States National Museum, 122(3600): 1-22. Genus Caligus (Caligidae). Annals and Magazine 1975. A New Family of Parasitic Copepods (Cyclopoida, of Natural History, 8(13): 109-126. Shiinoidae). Crustaceana, 28(2):211-219. 1968a. Copepoda Parasitic on Australian Fishes, VII: 1976. Shiinoa elagata, a New Species of Parasitic Shiinoa occlusa gen et sp. nov. Journal of Natural Copepod (Cyclopoida) from Elagatus (Carangi- History, 2:497-504. dae). Proceedings of the Biological Society of 1968b. Copepoda Parasitic on Australian Fishes, VIII; Washington, 88(40): 433-438. Families Lernaeopodidae and Naobranchiidae. Cressey, R. F., and H. Boyle Journal of Natural History, 2:505-523. 1973. Five New Bomolochid Copepods Parasitic on 1970. Copepoda Parasitic on Australian Fishes, X: Indo-Pacific Clupeid Fishes. Smithsonian Contri- Families Eudactylinidae and Pseudocycnidae. butions to Zoology, 161:1-25. Journal of Natural History, 4:159-173. Cuvier, G. Kabata, Z., and A. V. Gussev 1830. Le Regne Animal distribui d'apres son Organiza- 1966. Parasitic Copepoda of Fishes from the Collection tion. Volume 3, pages 255-258. of the Zoological Institute in Leningrad. Journal Dana, J. D. of the Linnean Society (Zoology), 46(309):155- 1852. Conspectus crustaceorum quae in orbis terrarum 207. circumnavigatione Carolo Wilkes e classe reipubli- Kazachenko, V. N., D. Korotaeva, and Yu. V. Kurochkin cae faederatae duce, Pars II. Proceedings of the 1972. Parasitic Copepods of Some Fishes from the American Academy of Arts and Sciences, 2:9-61. Pacific Ocean. Izvestiya Tikhookeanskogo Nauchno-

51 52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Issledovanniyu Institute. Rybnogo Khozyaistva i Milne-Edwards, M. H. Okeanographii, 81:224-238. 1840. Histoire naturelle des Crustads. Volume 3, 638 Kensley, B., and J. R. Grindley pages. 1973. South African Parasitic Copepoda. Annals of the Nordmann, A. V. South African Museum, 62(3):69-130. 1832. Mikrographische Beitrdge zur Naturgeschichte der Kirtisinghe, P. wirbellosen Thiere. 150 pages. 1935. Parasitic Copepods of Fish from Ceylon. Parasit- Nunes-Ruivo, L., and P. Fourmanoir ology, 27(3):332-344. 1956. Copepodes Parasites de Poissons de Madagascar. 1964. A Review of the Parasitic Gopepods of Fish Re- Memoires de I'Institut Scientifique de Madagascar, corded from Ceylon with Descriptions of Addi- 10(A):69-80. tional Forms. Bulletin of the Fisheries Research Pearse, A. S. Station, Ceylon, 17(1) :45-132. 1951. Parasitic Crustacea from Bimini, Bahamas. Pro- Klawe, W. L. ceedings of the United States National Museum, 1977. What Is a Tuna? Marine Fisheries Review, 39 101(3280):341-372. (11):1-5. 1952. Parasitic Crustacea from the Texas Coast. Publica- Kroyer, H. tions of the Institute of Marine Science, The 1838. Om Snyltekrebsene isaer med Hensyn til den University of Texas, 2(2): 5-42. Danske Fauna. Naturhistorisk Tidsskrift, 2:8-52. Pillai, N. K. 1863. Bidrag til Kundskab om Snyltekrebsene. Natur- 1961. Copepods Parasitic on South Indian Fishes, 1: historisk Tidsskrift, 2(3):75-320. Caligidae. The Bulletin of the Central Research Kurian, C. V. Institute, University of Kerala, Trivandrum, 1961. Parasitic Copepods of Fishes from Kerala—No. 3. 8:87-130. Bulletin of the Central Research Institute, Trivan- 1962a. On a New Species of Bomolochus (Copepoda) drum, 8 (series C):63-77. with Remarks on Orbitacolax Shen. The Journal Kurz, W. of Parasitology, 48(4):610-612. 1877. Studien Uber de Familie der Lernaeopodiden. 1962b. Observations on the Synonymy of Caligus Zeitschrift fur Wissenschaftliche Zoologie, 29:380- coryphaenae Stp-Lutk. Annals and Magazine of 428. Natural History, series 5, volume 13:513-522. Leigh-Sharpe, W. H. 1963. Copepods Parasitic on South Indian Fishes— 1926. A List of Parasitic Copepoda Found at Plymouth Family Caligidae. Journal of the Marine Biologi- with a Note on the Bulla of Clavella devastatrix. cal Association of India, 5(1): 68—96. Parasitology, 18:384-386. 1965. Copepods Parasitic on South Indian Fishes: Fam- 1930. Parasitic Copepoda. In V. van Straelen, editor, ily Bomolochidae, 3. Journal of the Bombay Nat- Resultats Scientifiques du voyage aux Indes ural History Society, 62 (1): 38-55. Orientales Neerlandaises de LL. AA. RR. le Prince 1971. Notes on Some Copepod Parasites in the Collec- et la Princesse Leopold de Belgique. Memoirs tion of the British Museum (N.H.), London. de Musee Royal d'Histoire Naturelle de Belgique, Journal of the Marine Biological Association of 3(2):1-11. India, 11:149-174. Lewis, A. G. Scott, T. 1964. Caligoid Copepods (Crustacea) of the Hawaiian 1894. Report on Entomostraca from the Gulf of Guinea, Islands: Parasitic on Fishes of the Family Collected by John Rattray, B.Sc. Transactions of Acanthuridae. Proceedings of the United States the Linnean Society, London, 6(2): 1-161. National Museum, 115(3482) : 137-244. 1901. Notes on Some Parasites of Fishes. Report of the 1967. Copepod Crustaceans Parasitic on Teleost Fishes Fishery Board for Scotland, 19(3) : 120-153. of the Hawaiian Islands. Proceedings of the United Shen, C. States National Museum, 121(3574): 1-204. 1957. Parasitic Copepods from Fishes of China, Part II: 1968. Copepod Crustaceans Parasitic on Fishes of Caligoida, Caligidae (1). Ada Zoologica Sinica, Eniwetok Atoll. Proceedings of the United States 9(4):351-377. National Museum, 125(3656): 1-78. Shiino, S. M. Lewis, A. G., J. Dean, and E. Gilfillan III 1952. Copepods Parasitic on Japanese Fishes, 1: On the 1969. Taxonomy and Host Associations of Some Parasitic Species of Caligus and Lepeophtheirus. Report Copepods (Crustacea) from Pelagic Teleost Fishes. of the Faculty of Fisheries, Prefectural University Pacific Science, 23(4): 414-437. of Mie, 1:79-113. Margolis, L., Z. Kabata, and R. Parker 1954. A New Fish-Louse Found on the Mackerel-Pike. 1975. Catalogue and Synopsis of Caligus. Bulletin of the Annotationes Zoological Japonenses, 27(3):150- Fisheries Research Board of Canada, 192:1-117. 153. Miculicich, M. 1956. Copepods Parasitic on Japanese Fishes, 12: Family 1904. Ein neuer Lernaepodidae. Zoologischer Anzeiger, Lernaeopodidae. Report of the Faculty of Fisher- 28:47-52. ies, Prefectural University of Mie, 2(2):269-311. NUMBER 3 1 1 53

1958. Parasitic Copepods from Fishes Collected in the ceedings of the First All-India Zoological Congress, Indian Ocean. Annual Report of the Prefectural 2:191-217. University of Mie, section 2 (Natural Science), Vervoort, W. 2(3):98-113. 1965. Three New Species of Bomolochidae (Copepoda, 1959a. Revision der auf Goldmakrele, Coryphaena hip- Cyclopoida) from Tropical Atlantic Tunnies. purus L. schmarotzenden Caligidenarten. Annual Zoologische Verhandelingen, 76:3—40. Report of the Prefectural University of Mie, sec- Wilson, C. B. tion 2 (Natural Science), 3(1): 1-34. 1905. North American Parasitic Copepods Belonging to 1959b. Ostpazifische Parasitierende Copepoden. Report the Family Caligidae, 1: The Caliginae. Pro- of the Faculty of Fisheries, Prefectural University ceedings of the United States National Museum, of Mie, 3(2):267-333. 28:479-672. 1959c. Sammlung der parasitischen Copepoden in der 1915. North American Parasitic Copepods Belonging to Prafturuniversitat von Mie. Report of the Faculty the Lernaeopodidae, with a Revision of the En- of Fisheries, Prefectural University of Mie, 3(2): tire Family. Proceedings of the United States Na- 334-374. tional Museum, 47(2063): 565-729. 1960. Parasitic Copepods of Fishes from the Eastern 1921. New Species and a New Genus of Parasitic Pacific. Report of the Faculty of Fisheries, Pre- Copepods. Proceedings of the United States Na- fectural University of Mie, 3(3) :527-541. tional Museum, 59(2354): 1-17. 1965. Parasitic Copepods of the Eastern Pacific Fishes, 1922. North American Parasitic Copepods Belonging to 5: Caligus. Report of the Faculty of Fisheries, the Family Dichelesthiidae. Proceedings of the Prefectural University of Mie, 5:391-420. United States National Museum, 60(2400): 1-100. 1935. New Parasitic Copepods. Smithsonian Miscellane- Silas, E. G., and A.N.P. Ummerkutty. ous Collections, 91 (19): 1-9. 1967. Parasites of Scombroid Fishes, Part II: Parasitic 1937. Parasitic Copepods Taken During the Third Han- Copepoda. Symposium on Scombroid Fishes, cock Expedition to the Galapagos Islands. Allan 3:876-993. Hancock Pacific Expedition, 2(4): 23-30. Steenstrup, J., and C. Liitken. Yamaguti, S. 1861. Bidrag til Kundskab om det aabne Havs Synltekrebs 1936. Parasitic Copepods from Fishes of Japan, part 2: og Lernaeer samt om nogle andre nye eller hiltil Caligoida, I. 22 pages, [private publication.] kun ufuldstaen digt kjendte parasitiske Copepoder, 1939. Parasitic Copepods from Fishes of Japan, part 5: 5:341-432. Caligoida, III. Volumen Jubilare Pro Prof. Sadao Tripathi, Y. R. Yoshida 3:443-487. 1962. Parasitic Copepods from Indian Fishes, III: 1963. Parasitic Copepoda and Branchiura of Fishes. Family Anthosomatidae and Dichelesthiidae. Pro- 1104 pages. New York: Interscience Publishers. Figures

FIGURE 1.—Holobomolochus divaricatus, new species, female: a, dorsal; b, genital segment and abdomen, ventral; c, last abdominal segment and caudal rami, ventral; d, first antenna.

54 NUMBER 3 1 1 55

E D

FIGURE 2.—Holobomolochus divaricatus, new species, female: a, second antenna; b, mandible, paragnath, first maxilla, second maxilla; c, maxilliped; d, leg 1; e, leg 2. 56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Holobomolochus divaricatus, new species, female: a, leg 3; b, leg 4; c, leg 5; male: d, dorsal. NUMBER 3 1 1 57

E

D

FIGURE 4.—Holobomolochus divaricatus, new species, male: a, genital segment and abdomen, ventral; b, last abdominal segment and caudal rami, ventral; c, first antenna; d, maxilliped; e, leg 1. 58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 5.—Holobomolochus divaricatus, new species, male: a, leg 3; b, leg 4; c, leg 5. Holobomolochus nudiusculus, new species, female: d, dorsal; e, genital segment and abdomen, ventral. NUMBER 3 1 1 59

FIGURE 6.—Holobomolochus nudiusculus, new species, female: a, last abdominal segment and caudal rami, ventral; b, leg 2; c, leg 3; d, leg 5; male: e, dorsal. 60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7.—Holobomolochus nudiusculus, new species, male: a, genital segment and abdomen, ventral; b, last abdominal segment and caudal rami, ventral. Holobomolochus asperatus, new species, female: c, dorsal; d, genital segment and abdomen, dorsal; e, last abdominal segment and caudal rami, ventral; /, first antenna; g, second antenna. NUMBER 3 1 1 61

FIGURE 8.—Holobomolochus asperatus, new species, female; a, mandible, paragnath, first maxilla, second maxilla; b, leg 2; c, leg 3; d, leg 4; e, leg 5. 62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 9.—Unicolax collateralis, new genus, new species, female: a, dorsal; b, genital segment and abdomen, dorsal; c, last abdominal segment and caudal rami, ventral; d, first antenna; e, second antenna. NUMBER 311 63

FIGURE 10.—Unicolax collateralis, new genus, new species, female: a, labrum, mandible, paragnath, first maxilla, second maxilla; b, maxilliped; c, leg 1; d, leg 2. 64 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 11.—Unicolax collaterals, new genus, new species, female: a, leg 3; b, leg 4; c, leg 5; male: d, dorsal; e, genital segment and abdomen, dorsal. NUMBER 3 1 1 65

FIGURE 12.—Unicolax collateralis, new genus, new species, male: a, last abdominal segment and caudal rami, ventral; b, first antenna; c, maxilliped; d, leg 1; e, leg 2. 66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Unicolax collaterals, new genus, new species, male: a, leg 3; b, leg 4; c, leg 5. NUMBER 3 1 1 67

E

B

D

FIGURE 14.—Unicolax anonymous (Vervoort), new genus, female: a, dorsal; b, first antenna; c, second antenna; d, mandible, paragnath, first maxilla, second maxilla; e, maxilliped. 68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 15.—Unicolax anonymous (Vervoort), new genus, female: a, leg 1; b, leg 2. NUMBER 3 1 1 69

E

FIGURE 16.—Unicolax anonymous (Vervoort), new genus, female: a, leg 3; b, leg 4; c, leg 5; male: d, dorsal; e, leg 5. 70 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 17.—Unicolax mycterobius (Vervoort), new genus, female: a, dorsal; b, leg 2; c, leg 3. NUMBER 3 1 1 71

FIGURE 18.—Unicolax mycterobius (Vervoort), new genus, female: a, leg 4; b, leg 5; male: c, dorsal; d, last abdominal segment and caudal raini, ventral; e, leg 1; /, leg 1 exopod. 72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Unicolax mycterobius (Vervoort), new genus, male: a, leg 2; b, leg 3; c, leg 4; d, leg 4 last 2 endopod segments. NUMBER 3 1 1 73

B

FIGURE 20.—Unicolax ciliatus, new genus, new species, female: a, dorsal; b, first antenna; e, leg 1 exopod; d, leg 2 exopod. 74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 21.—Unicolax ciliatus, new genus, new species, female: a, leg 3; b, leg 4; c, leg 5; male: d, dorsal; e, last abdominal segment and caudal rami, ventral; /, leg 1 endopod. NUMBER 311 75

B

FIGURE 22.—Unicolax ciliatus, new genus, new species, male: a, leg 2; b, leg 3; c, leg 4; d, leg 5. 76 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 23.—Unicolax reductus new genus, new species, female: a, dorsal; b, genital segment and abdomen, dorsal; c, last abdominal segment and caudal rami, ventral; d, first antenna; e, second antenna; /, mandible, paragnath, first maxilla, second maxilla. NUMBER 3 1 1 77

B

FIGURE 24.—Unicolax reductus, new genus, new species, female: a, maxilliped; b, leg 1; c, leg 2 \d, leg 3. 78 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

FIGURE 25.—Unicolax reductus, new genus, new species, female: a, leg 4; b, leg 5; male: c, dorsal; d, genital segment and abdomen, ventral; e, last abdominal segment and caudal rami, ventral. NUMBER 3 1 1 79

FIGURE 26.—Unicolax reductus, new genus, new species, male: a, maxilliped; b, leg 1; c, leg 2; d, leg 5. 80 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 27.—Ceratacolax euthynni Vervoort, female: a, dorsal; b, lateral; c, genital segment and abdomen, ventral; d, last abdominal segment and caudal rami; e, first antenna; /, second antenna; g, mandible, paragnath, first and second maxilla. NUMBER 3 1 1 81

D \

FIGURE 28.—Ceratacolax euthynni Vervoort, female: a, maxilliped; b, leg 1; c, leg 2; d, leg 3. 82 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

E

FIGURE 29.—Ceratacolax euthynni Vervoort, female: a, leg 4; b, leg 5; male: c, dorsal; d, genital segment and abdomen, ventral; e, last abdominal segment and caudal rami, ventral; /, first antenna. NUMBER 311 83

B

D

FIGURE 30.—Ceratacolax euthynni Vervoort, male: a, second antenna; b, maxilliped; c, leg 1; d, leg 2. 84 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

s

FIGURE 31.—Ceratacolax euthynni Vervoort, male: a, leg 3; b, leg 4; c, leg 5. NUMBER 3 1 1 85

B

FIGURE 32.—Nothobomolochus kanagurta Pillai, female: a, dorsal; b, last abdominal segment and abdomen, ventral; c, first antenna; d, maxilliped. 86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 33.—Nothobomolochus kanagurta Pillai, female: a, leg 2; b. leg 3; c, leg 4; d, leg 5. NUMBER 311 87

D

FIGURE 34.—Orbitacolax aculeatus (Pillai), female: a, dorsal; b, first antenna; c, second an- tenna; d, oral area; e, maxilliped; /, leg 1. 88 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

E

FIGURE 35.—Orbitacolax aculeatus (Pillai), female: a, leg 2; b, leg 5. Pumiliopes capitulatus Cressey and Boyle, female: c, dorsal; d, genital segment, abdomen, caudal rami, ventral; e, last abdominal segment and caudal rarni, ventral. NUMBER 3 1 1 89

B

D

FIGURE 36.—Pumiliopes capitulatus Cressey and Boyle, female: a, first antenna; b, second an- tenna ; c, oral area; d, maxilliped; e, leg 1. 90 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

B

FIGURE 37.—Pumiliopes capitulatus Cressey and Boyle, female: a, leg 2; b, leg 3; c, leg 4; d, leg 5. NUMBER 3 1 1 91

D

FIGURE 38.—Shiinoa inauris Cressey, female: a, lateral with attached male; b, rostral area, lateral; c, oral area; d, leg 1. 92 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 39.—Shiinoa inauris Cressey, female: a, leg 2; b, leg 3; male: c, second antenna; d, exopod of leg 2. Shiinoa occlusa Kabata, female: e, lateral; male: /, genital segment and abdomen; g, caudal ramus; h, second antenna. NUMBER 3 1 1 93

B

E

FIGURE 40.—Caligus coryphaenae Steenstrup and Liitken, female: a, dorsal; b, caudal ramus, ventral; c, sternal furea; d, leg 1; e, leg 2 endopod; /, leg 3 exopod. 94 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 41.—Caligus coryphaenae Steenstrup and Lutken, female: a, leg 4 exopod; male: b, dorsal. Caligus regalis Leigh-Sharpe, female: c, dorsal; d, sternal furca; e, leg 3 exopod; /, leg 4 exopod: male: g, genital segment and abdomen, ventral. NUMBER 3 1 1 95

B

FIGURE 42.—Caligus productus Miiller, female: a, dorsal; b, last abdominal segment and caudal rami, ventral; c, oral area; d, first antenna; e, second antenna. 96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 43.—Caligus productus Miiller, female: a, second maxilla; b, maxilliped; c, sternal furca; d, leg 1; e, leg 2. NUMBER 3 1 1 97

E

B

FIGURE 44.—Caligus productus Muller, female: a, leg 3; b, leg 4; male: c, dorsal; ,d, genital segment and abdomen; e, second antenna. 98 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 45.—Caligns asymmetricus Kabata, female: a, dorsal; b, abdomen and caudal rami, ventral; c, oral area; d, first antenna; e, second antenna. NUMBER 3 1 1 99

E D

FIGURE 46.—Caligus asymmetricus Kabata, female: a, second maxilla; b, maxilliped; c, sternal furca; d, leg 1; e, leg 2. 100 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 47.—Caligus asymmetricus Kabata, female: a, leg 3; b, leg 4; male: c, dorsal; d, geni- tal segment and abdomen, ventral; e, second antenna. NUMBER 311 101

FIGURE 48.—Caligus bonito Wilson, female: a, dorsal; b, posterior portion of abdomen and caudal rami, ventral; c, oral area; d, first antenna; e, second antenna. 102 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 49.—Caligns bonito Wilson, female: a, second maxilla; b, maxilliped; c, sternal furca; d, leg 1; e, leg 2; /, detail of outer edge of leg 2 endopod second segment. NUMBER 311 103

D

FIGURE 50.—Caligus bonito Wilson, female: a, leg 3; b, leg 4; male: c, dorsal; d, second antenna; e, distal portion of maxilliped. 104 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

D

FIGURE 51.—Caligus mutabilis Wilson, female: a, dorsal; b, genital segment and abdomen, ventral; c, last abdominal segment and caudal rami, ventral; d, first antenna; «, postantennal process; /, second antenna. NUMBER 3 1 1 105

FIGURE 52.—Caligus mutabilis Wilson, female: a, postoral process; b, sternal furca; c, oral area; d, leg 1; e, leg 2. 106 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

c

FIGURE 53.—Caligus mutabilis Wilson, female: a, leg 3; b, leg 4. Caligns omissus, new species; female: c, dorsal; d, genital segment and abdomen, ventral; e, last abdominal segment and caudal ranii, ventral. NUMBER 3 1 1 107

FIGURE 54.—Caligus omissus, new species, female: a, oral area; b, first antenna; c, postantennal process; d, second antenna; e, postoral process; /, maxilliped; g, sternal furca. 108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 55.—Caligus omissus, new species, female: a, leg 1; b, leg 2; c, leg o; d, leg 4. NUMBER 3 1 1 109

FIGURE 56.—Caligus omissus, new species, male: a, dorsal; b, genital segment and abdomen, ventral; c, second antenna; d, postantennal process; e, postoral process; /, maxilliped; g, sternal furca. 110 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 57.—Caligus biseriodentatus Shen, female: a, dorsal; b, abdomen and caudal rami, ventral; c, cephalon, ventral; d, leg 1; e, leg 2 endopod. NUMBER 3 1 1 111

FIGURE 58.—Caligus biseriodentatus Shen, female: a, leg 3 exopod; b, leg 4; male: c, dorsal; immature female: d, dorsal; e, maxilliped; /, sternal furca; g, leg 1 last segment. 112 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

E

FIGURE 59.—Caligus biseriodentatus Shen, immature female: a, leg 2; b, leg 4; immature male: c, dorsal; d, tip of abdomen and caudal ramus, ventral; e, postantennal process; /, maxilliped. NUMBER 311 113

B

FIGURE 60.—Caligus cybii Bassett-Smith, female: a, dorsal; b, genital segment and abdomen, ventral; c, last abdominal segment and caudal rami, ventral; d, oral area; e, first antenna. 114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 61.—Caligus cybii Bassett-Smith, female: a, postantennal process; b, second antenna; c, postoral process; d, second maxilla; e, maxilliped; /, sternal furca; g, leg 1. NUMBER 3 1 1 115

FIGURE 62.—Caligus cybii Bassett-Smith, female: a, leg 2; 6, leg 3; c, leg 4; male: d, dorsal. 116 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

FIGURE 63.—Caligus cybii Bassett-Smith, male: a, genital segment and abdomen, ventral; b, second antenna; c, postoral process; d, maxilliped. NUMBER 3 1 1 117

B

FIGURE 64.—Caligus pelamydis Kroyer, female: a, dorsal; b, genital segment and abdomen, ventral; e, oral area; d, first antenna; e, second antenna. 118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

E

FIGURE 65.—Caligns pelamydis Kroyer, female: a, second maxilla; b, maxilliped; c, sternal furca; d, leg 1; e, leg 2. NUMBER 3 1 1 119

FIGURE 66.—Caligns pelamydis Kroyer, female: a, leg 3; b, leg 4; male: e, dorsal; d, genital segment and abdomen, ventral; e, postoral process; /, maxilliped. 120 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 67.—Caligus infestans Heller, female: a, dorsal; b, genital segment and abdomen, ventral; c, last abdominal segment and caudal rami, ventral; d, oral area; e, first antenna. NUMBER 3 1 1 121

F

B

FIGURE 68.—Caligus infestans Heller, female: a, second antenna; b, tip of second maxilla; c, maxilliped: d, sternal furca; e, leg 1; /, leg 2. 122 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 69.—Caligus infestans Heller, female: a, leg 3; b, leg 4; male: c, dorsal; d, genital segment and abdomen, ventral; e, second antenna. NUMBER 3 1 1 123

FIGURE 70.—Caligus diaphanus Nordmann, female: a, dorsal; b, genital segment and abdomen, ventral; c, distal end of abdomen and caudal rami, ventral; d, first antenna; e, postantennal process; /, second antenna; g, postoral process. 124 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 71.—Caligns diaphanus Nordmann, female: a, second maxilla; b, maxilliped; c, sternal furca; d, oral area; e, leg 1; /, leg 2. NUMBER 311 125

B

FIGURE 72.—Caligus diaphanus Nordmann, female: a, leg 3; b, leg 4; male: c, dorsal; d, second antenna; e, postoral process; /, maxilliped. 126 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 73.—Caligus savala Gnanamuthu, female: a, dorsal; b, second antenna; c, postantennal process; d, sternal furca; e, leg 2 endopod; /, leg 4 exopod last segment. Caligus macarovi Gussev, female: g, dorsal; h, second antenna, postantennal process, postoral process; i, sternal furca. (C macarovi after Shiino 1959.) NUMBER 3 1 1 127

FIGURE 74.—Caligus macarovi Gussev, female: a, leg 2 endopod; b, leg 4. Caligus ambly- genitalis Pillai, female: c, dorsal; d, sternal furca; e, leg 1; /, leg 2 endopod edge; g, leg 2 exopod; h, leg 4. (C. macarovi after Shiino 1959, C. amblygenitalis after Pillai 1961.) 128 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 75.-—Elytrophora brachyptera Gerstaeckcr, female: a, dorsal; b, caudal ramus; c, first antenna; d, second antenna; e, mouth tube and postoral process; /, second maxilla; g, maxil- liped. NUMBER 3 1 1 129

D

FIGURE 76.—Elytrophora brachyptera Gerstaecker, female: a, sternal furca; b, leg 1; c, leg 2; d, leg 3; e, leg 3 endopod. 130 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 77.—Elytrophora brachyptera Gerstaecker, female: a, leg 3 exopod; b, leg 4 (cold water; c, leg 4 (warm water); male: d, dorsal; e, genital segment and abdomen, ventral; /, second antenna; g, maxilliped. NUMBER 3 1 1 131

E

FIGURE 78.—Elytrophora indica Shiino, female: a, dorsal; b, genital segment and abdomen, ventral; c, first antenna; d, second antenna; e, mouth tube and postoral process; /, leg 4; g, leg 6. 132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 79.—Elytrophora indica Shiino, male: a, dorsal; b, second antenna. Gloiopotes hygomianus Steenstrup and Lutken, female: c, dorsal; d, sternal furca; e, leg 1; f, tip of exopod of leg 1; g, endopod of leg 1; h, leg 4; male: i, dorsal. NUMBER 3 1 1 133

FIGURE 80.—Caligulus longispinosus Heegaard, female: a, dorsal; male: b, dorsal; c, maxil- liped; d, sternal furca; e, leg 1; /, leg 2 (female after Heegaard, 1962). 134 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

D

FIGURE 81.—Tuxophorus cybii Nunes-Ruivo and Fourmanoir, female: a, dorsal. Tuxophorus cervicornis Heegaard, female: b, dorsal; c, second antenna; d, postantennal process; e, sternal furca; /, leg 1; g, leg 4. (7\ cybii after N.-R and Fourmanoir, 1956.) NUMBER 3 1 1 135

A

FIGURE 82.—Tuxophorus collettei, new species, female: a, dorsal; b, genital segment and abdomen, ventral; c, caudal rami, ventral; d, first antenna; e, second antenna and postantennal process; /, postoral process. 136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

FIGURE 83.—Tuxophorus collettei, new species, female: a, second maxilla; b, maxilliped; c, sternal furca; d, leg 1; e, leg 2. NUMBER 3 1 1 137

FIGURE 84.—Tuxophorus collettei, new species, female: a, leg 3; b, leg 4; male: c, dorsal; d, genital segment and abdomen, ventral; e, second antenna; /, maxilliped. 138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 85.—Pseudocycnus appendiculatus Heller, female: a, dorsal; b, first antenna; c, second antenna; d, mouth tube and mandible; e, first maxilla; /, second maxilla. NUMBER 3 1 1 139

D

FIGURE 86.—Pseudocycnus appendiculatus Heller, female: a, maxilliped; b, leg 1; c, leg 2; d, leg 3. 140 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D B

FIGURE 87.—Pseudocycnus appendiculatus Heller, male: a, dorsal; b, abdomen and caudal rami, ventral; c, first antenna; d, maxilliped; e, leg 1; /, leg 4. NUMBER 3 1 1 141

D

FIGURE 88.—Pseudocycnoides armatus (Bassett-Smith), female: a, dorsal; b, first antenna; c, first and second antennae; d, mouth tube; e, second maxilla. 142 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 89.—Pseudocycnoides armatus (Bassett-Smith), female: a, maxilliped; b, leg 1; c, leg 2; d, leg 3; male: e, dorsal. NUMBER 3 1 1 143

E

FIGURE 90.—Pseudocycnoides armatus (Bassett-Smith), male: a, abdomen and caudal rami; b, first antenna; c, second maxilla; d, maxilliped; e, leg 1. 144 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 91.—Pseudocycnoides armatus (Bassett-Smith), male: a, leg 2; b, leg 3. Pseudo- cycnoides scomberomori (Yamaguti), female: c, dorsal; d, maxilliped; e, leg 1; /, leg 2; g. ^g 3. NUMBER 311 145

FIGURE 92.—Pseudocycnoides buccata (Wilson), female: a, dorsal; b, abdomen and caudal rarni, young female, ventral; c, first antenna; d, second antenna; e, mouth tube; /, first maxilla; g, second maxilla. 146 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 93.—Pseudocycnoides buccata (Wilson), female: a, maxilliped; b, claw of maxilliped; c, legs 1-3; male: d, dorsal; e, abdomen and caudal ramus, ventral; /, second antenna. NUMBER 3 1 1 147

E

FIGURE 94.—Pseudocycnoid.es buccata (Wilson), male: a, first maxilla, mandible and mouth tube; b, maxilliped; c, leg 1; d, leg 3. Lernanthropus kanagurta Tripathi, female: e, dorsal; /, second antenna; g, leg 1. 148 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 95.—Brachiella thynni Cuvier, female: a, ventral. Brachiella magna Kabata: b, dorsal. Clavellisa scombri (Kurz), female: c, ventral. Clavelliopsis saba Yamaguti, female: d, lateral. (5. thynni after Shiino, 1956; C. saba after Shiino, 1959c.) NUMBER 3 11 149

o divaricatus • nudiusculus

FIGURE 96.—Distribution of Holobomolochus divaricatus, new species, H. nudiusculus, new species, and H. asperatus, new species.

FIGURE 97.—Distribution of Unicolax collaterals, new species. 150 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

• anonymous • ciliatus

FIGURE 98.—Distribution of Unicolox anonymous (Vervoort) and U. ciliatus, new species.

o mycterobius r • reductus

FIGURE 99.—Distribution of Unicolax mycterobius (Vervoort) and U. reductus, new species. NUMBER 3 1 1 151

o C. euthynni * P. capitulatus • N. kanagurta kO.aculeatus

FIGURE 100.—Distribution of Ceratacolax euthynni Vervoort, Pumiliopes capitulatus Cressey and Boyle, Nothobomolochus kanagurta Pillai, and Orbitacolax aculeatus (Pillai).

o inauns • occlusa

FIGURE 101.—Distribution of Shiino inauris Cressey and S. occlusa Kabata. 152 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

• coryphaenae • regalis

FIGURE 102.—Distribution of Caligus coryphaenae Steenstrup and Liitken, and C. regain Leigh-Sharpe. (Includes some records of Lewis, et al. 1969; Shiino 1959a, 1973; Kabata and Gussev, 1966; Pillai, 1963.)

FIGURE 103.—Distribution of Caligus productus Muller. NUMBER 3 1 1 153

o pelamydis * asymmetricus

FIGURE 104.—Distribution of Caligns pelamydis Kroyer, C. asymmetricus Kabata, C. infestans Heller, and C. cybii Bassett-Smith.

o bonito * omissus • mutabilis \! • biseriodentatus ~|

FIGURE 105.—Distribution of Caligus bonito Wilson, C. omissus, new species, C. mutabilis Wilson, and C. biseriodentatus Shen. 154 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

o indica • brachyptera

FIGURE 106.—Distribution of Elytrophora indica Shiino, and E. brachyptera Gerstaecker. (Includes Shiino, 1958 and 1965 E. indica.)

o buccata * armatus • scomberomori -

FIGURE 107.—Distribution of Pseudocycnoides buccata (Wilson), P. armatus (Bassett-Smith), and P. scomberomori (Yamaguti). NUMBER 3 1 1 155

"f 'itf mf 1

* B. thynni • C. scombri m B.magna oC.saba

FIGURE 108.—Distribution of Brachiella thynni Cuvier, Clavellisa scombri (Kurz), B. magna Kabata, and Clavellopsis sab a Yamaguti. (Includes Kabata, 1968b B. magna.) 156 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 109.—Holobomolochus divaricatus, new species, female: a, rostral area (1775X) b, abdomen and caudal rami, ventral (540x); c, abdomen, outer distal corner (295OX); d, labrum (1000X ); e, paragnath (3145X ); /, leg 2 exopod (500X)• NUMBER 3 1 1 157

FIGURE 110.—Holobomolochus divaricatus, new species, female: a, leg 2 exopod spines (2850); b, leg 2 endopod first segment (1500X): c, leg 2 armature of interpodal plate (3750X); d, leg 2 (above) and leg 3 (below) (200x); e, leg 3 exopod first segment outer spine (1400X); /, same, lateral view (1315X). 158 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 111.—Holobomolochus divaricatus, new species, female: a, leg 3 exopod first segment base of outer spine (1400X ); b, leg 4 tip of endopod (2000X ) ; male: c, abdomen and caudal ramus, ventral (500x); d, caudal ramus outer distal corner (5000X); e, second antenna (350X); /, hooks on second antenna (3000X). NUMBER 311 159

FIGURE 112.—Holobomolochus divaricatus, new species, male: a, inner edge of maxilliped (1250X); b, same (3750X); c, leg 1 interpodal plate (2000X); d, leg 1 basipod (1500X); e, leg 1 exopod third segment (1500X); f, same, outer edge (3000X). 160 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 113.—Holobomolochus divaricatus, new species, male: a, leg 1 exopod third segment outer edge (5000x). Unicolax collateralis, new species, female: b, cephalon. anterior view (300x); c, same (900X); d, abdomen and caudal rami, ventral (440X); e, tip of caudal ramus, ventral (2000X); /, modified seta of first antenna (600X). NUMBER 311 161

FIGURE 114.—Unicolax collaterally new species, female: a, second antenna (750x); b, hooks on second antenna (2400x); c, same (2600X); d, labrum (600X); e, oral area (1500X); /, tip of mandible (10,000x)- 162 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 115.—Unicolax collaterals, new species, female: a, tip of paragnath (lO.OOOx); b, leg 2 exopod (lOOOx); c, spines on leg 2 exopod (2000x); d, hyaline fringe of leg 2 exopod outer edge (8000X); e, spine on leg 2 exopod (25OOX); /, leg 4 endopod last seg- ment (1000X). NUMBER 3 1 1 163

FIGURE 116.—Unicolax collateralis, new species, female: a, tip of leg 4 endopod (2000x)- Unicolax anonymous (Vervoort), female: b, cephalon, anterior view (600x); c, first and second antenna, anterior view (700X); d, modified seta of first antenna (2300X); t> hooks on second antenna (75OOX); /, oral area (750X)- 164 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 117.—Unicolax anonymous (Vervoort), female: a, leg 2 exopod (1650X); b, leg 3 exopod outer edge (1750X)- Unicolax mycterobius (Vervoort), female: c, cephalon, anterior view (200x ) ; d, surface of cephalon dorsal to insertion of first antennae (1400X ) ; «, cepha- lon, anteroventral view (200X ) ; /, modified seta of first antenna (400X ). NUMBER 3 1 1 165

FIGURE 118.—Unicolax mycterobius (Vervoort), female: a, second antenna (750x); b, tip of second antenna (1400X); c, oral area (1050x); d, tip of paragnath (5250X); e, leg 1 exopod (500X); /, leg 1 basipod (1000X). 166 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 119.—Unicolax mycterobius (Vervoort), female: a, leg 1 exopod first segment (700X ); b, leg 1 basipod and exopod insertion (1050X); c, leg 4 endopod tip (75Ox)- Unicolax reductus, new species, female: d, distal end of caudal rarnus (1500X); e, modified seta of first antenna (500x); f, second antenna (lOOOx). NUMBER 311 167

FIGURE 120.—Unicolax reductus, new species, female: a, oral area (700x); b, paragnath (3400X); c, leg 2 (260x); d, leg 2 exopod first segment outer corner (2000X); e, leg 2 exopod last segment tip (1200X ); /, leg 2 interpodal plate (1400X). 168 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 121.—Unicolax reductus, new species, female: a, leg 3 (260x); 6, leg 4 endopod first segment spinules (2000X); c, leg 3 exopod first segment (1000X); d, leg 3 exopod first segment spinules (5000X ); e, leg 4 (240X ); /, leg 4 exopod (650x )• NUMBER 311 169

FIGURE 122.—Unicolax reductus, new species, female: a, leg 4 exopod second segment outer corner (1200X); b, leg 4 endopod tip (1600X). Ceratacolax euthynni Vervoort, female: c, cephalon, anterolateral view (230X); d. same, oblique view (185X); e, same, anterior view (200X); /, rostrum (600X). 170 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 123.—Ceratacolax euthynni Vervoort, female: a, rostrum detail (2280x)- b posterior corner of genital segment (260X ) ; c, same, lateral view (370X); d, abdomen and caudal rami, ventral (300X ) ; e, caudal ramus, ventral (500X); /, caudal ramus posterior corner ventral (5000X). NUMBER 311 171

FIGURE 124.—Ceratacolax euthynni Vervoort, female: a, base of modified seta on first antenna, rostrum on left (420 X ) ; b, second antenna, distal half (1000X ) ; c, hooks on second antenna (5000X); d, terminal seta of second antenna (4000X): e, oral area (400X ); /, oral area, detail (650X). 172 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 125.—Ceratacolax euthynni Vervoort, female: a, tip of mandible and paragnath (1350X ) ; b, detail of second maxilla terminal flagellum (2000X ) ; c, maxilliped claw (900X ) : d, fringe of leg 1 endopod (1900X ) ; e, leg 2 exopod outer edge (800X); f, leg 5 lateral spine (750X). NUMBER 311 173

FIGURE 126.—Pumiliopes capitulatus Cressey and Boyle, female: a, spinules on abdomen, ventral (2800 X ) ; b, oral area (300 X ) ; c, flagellum of second maxilla and labium below (2000X)- Shiinoa occlusa Kabata, female: d, rostral area, anterior view (100x); e, rostrum above and second antenna below, lateral view (75 X ) ; /, detail of pores on rostrum and second antenna, lateral view (280X)- 174 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 127.—Shiinoa occlusa Kabata, female: a, tip of second antenna and first antenna (200X); b, base of first antenna (500x); c, oral area and legs 1 and 2 (90x); d, leg 1 (660x); e, leg 2 (440X); /, legs 3 (180x). NUMBER 311 175

FIGURE 128.—Shiinoa occlusa Kabata, female: a, tip of genital segment, abdomen, and caudal rami, lateral view (70x); b, detail of genital area (240x)- Caligus coryphaenae Steenstrup and Liitken, female: c, cephalon, ventral (30x ) ; d, frontal lunule (200X ) ; e, second antenna (200X); f, a happy mouth tube (280X ). 176 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 129.—Caligus coryphaenae Steenstrup and Liitken, female: a, tip of mouth tube, labrum above (550 X ); b, leg 1 (150 X ) ; c, tip of leg 1 exopod (500x) ; d, midterminal spine on leg 1 exopod (1300X); e, fringe at base of midterminal spine on leg 1 exopod (2800x); /, plumosities on leg 1 lateral seta (1300x)- NUMBER 3 1 1 177

FIGURE 130.—Caligus coryphaenae Steenstrup and Liitken, female: a, leg 2 exopod (26OX); b, spine on leg 2 first segment (650X ) ; c, spinules on outer edge of leg 2 endopod (1000X ) ; d, tip of leg 4 (290 X ) ; e, spine on leg 4 second segment (600 X ); /, fringe at base of terminal seta on leg 4 (1200 X ). 178 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 131.—Pseudocycnus appendiculatus Heller, female: a, cephalon, anterior view (55x); b, same, between bases of first antennae (300X ) ; c, anal opening (240X ) ; d, same (373x) ; *, second antenna (105 X ); /, second maxilla (500X ) • NUMBER 311 179

FIGURE 132.—Pseudocycnus appendiculatus Heller, female: a, maxilliped (200X); b, tip of maxilliped (500X ) ; c, leg 1 (420X ) ; d, leg 1 exopod base of terminal spine (1800X ); ', leg 1 endopod (75OX ) ; /, same, lateral view (800x)- 180 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1p^ - ' £

D •

i K i *——

FIGURE 133.—Pseudocycnus appendiculatus Heller, female: a, leg 2 (125X ) ; b, same (500X) ; c, leg 3 (400X); rf, tip of leg 3 (1200X); male: e, genital segment, ventral (150X); /, spinules on genital segment, ventral (5000X). NUMBER 311 181

FIGURE 134.—Pseudocycnus appendiculatus Heller, male: a, caudal ramus, ventral (200X) b, spinules on caudal ramus (2000X ) ; c, mouth tube (400X ) ; d, tip of mouth tube (2000X ) e, first maxilla (400 X ); /, base of maxilliped (500X ). 182 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 135.—Pseudocycnus appendiculatus Heller, male: a, leg 1 (400x); b, leg 1 exopod (2000X); c, leg 1 endopod (4000X); d, leg 2 (400X); e, spinules on 'leg 2 basipod (5500X); /, leg 3 (625X). NUMBER 311 183

FIGURE 136.—Pseudocycnus appendiculatus Heller, male: a, base of leg 5 (5000X)- Pseudo- cycnoides armatus (Bassett-Smith), male: b, spinules on genital segment (6550X ) ; c, abdomen and caudal rami, ventral (250X); d, caudal ramus, ventral (750x); e, tip of caudal ramus, ventral (1600x ); /, leg 1 (500X ). 184 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 137.—Pseudocycnoides armatus (Bassett-Smith), male: a, legs 2 and 3 (500x); b, leg 2 tip of exopod (3100 X ); c, leg 2 tip of endopod (2000X ) ; d, leg 2 endopod spinules (7000X ) ; e, leg 2 exopod tip and leg 3 (1050X ) ; /, tip of leg 3 (5250X )• NUMBER 311 185

FIGURE 138.—Pseudocycnoides buccata (Wilson), female: a, cephalon, ventral (75X); b, second antenna (37OX); c, mouth tube (600x); d, tip of mouth tube (2500X); e, first maxilla (1250X); /, tip of second maxilla (1450X). 186 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 139.—Pseudocycnoid.es buccata (Wilson), female: a, tip of second maxilla (1225X); b, same (1450x); c, same (4400X).

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