Parasitic Copepods of Mackerel - and Tuna-like Fishes (Scombridae ...

m 

Parasitic Copepods of Mackerel - 

and Tuna-like Fishes 

(Scombridae) 

of the World 

ROGER CRESSEY 

and 

HILLARY BOYLE CRESSEY 

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 311

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 311 

Parasitic Copepods of Mackerel 

and Tuna-like Fishes 



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. Cressey, 

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 

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 

Page

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 Mackereland 

Tuna-like Fishes 



Introduction 

There are 47 currently recognized species of the 

family Scombridae; plus an additional new species of 

Scomberomorus being described by Collette and Russo. 

One species, Gasterochisma melampus Richardson, is 

placed in a separate subfamily and has not been considered 

in this study. For approximately the last 20 

years the first author has been collecting and receiving 

collections of the copepods parasitic on these fishes. 

Of the 46 species of parasitic copepods reported here, 

40 species are represented in our collections. These 

40 species have been collected from 2422 individual 

fish from all 46 species of Scombrinae. As a result of 

this exhaustive collection we have presented a comprehensive 

account of the copepods parasitic on tunas 

and tuna-related fishes. 

Our primary objective has been to produce a baseline 

study of the copepods parasitic on scombrid fishes. 

Because of the unreliability of most past literature it 

has been impossible to assess the host-parasite relationship, 

especially the degree of host specificity. By compiling 

a comprehensive collection such as we report 

Roger Cressey and Hillary Boyle Cressey, Department of 

Invertebrate Zoology, National Museum of Natural History, 

Smithsonian Institution, Washington, D.C. 20560. 

Roger Cressey 

and Hillary Boyle Cressey 

here, resolving problems of synonymy with several 

copepod species, and working closely with scombrid 

taxonomists (R. H. Gibbs and B. B. Collette) to assure 

proper host identifications, we feel we have reached 

that objective. We do not feel that we have produced 

a definitive work, but hopefully, a solid foundation to 

which information in future collections can be added. 

Previous works by Lewis, Kabata, Shiino, and Pillai 

concerned with scombrid copepods are excellent and 

we have included some of their information where it 

fills gaps in our own collection data. For reasons stated 

above, however, we have not considered most of the 

earlier works in our host-parasite data. 

Because most species of scombrid fishes are commercially 

important, various aspects of their biology have 

been investigated for several years. One result of this 

interest has been the many reports of copepods parasitic 

on them since the last century. The major failing 

in this, however, has been the general lack of cooperative 

effort between parasitologists and ichthyologists. 

Too often, the copepod worker has been content to 

repeat unverified host identifications or unreliable 

common names. This casual approach to that aspect 

of the record has resulted in an account too confusing 

to be of much value in determining true relationships 

between the copepod species and their hosts. Except 

for a literature compilation by Silas and Ummerkutty

in 1967, no attempt has been made to critically 

examine the scombrid parasites in a comprehensive 

way to determine the taxonomic status of most of the 

described species. For example, Silas and Ummerkutty 

list 29 species of Caligus as reported parasites of the 

Scombrinae. As a result of our examinations of typespecimens 

of a number of these Caligus species, and 

the data based on our own collections, we have been 

able to place almost half of these in synonymy and 

reduce the number of Caligus species from these hosts 

to 15. 

We have included literature records in our data 

only in those cases where we could verify the host 

and/or the copepod identification. To include unverified 

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 

complete as reasonably possible to publish our results. 

We have, with each species, given a skeleton synonymy 

to provide a brief historical account of each. A complete 

synopsis of the species of Caligus was recently 

published by Margolis, Kabata, and Parker (1975) 

but a complete revision of the genus is badly needed. 

There are slightly over 200 currently recognized species 

of Caligus. Reducing the 29 species cited by Silas 

and Ummerkutty to 15 illustrates the extent of the 

problem. Extrapolating from that reduction of 29 

species to 15, it would not be unreasonable to expect 

that a generic revision might result in only 100 or so 

Caligus species as valid. 

The first author and B. B. Collette are planning a 

joint paper summarizing the host-parasite relationship 

from both ecological and evolutionary standpoints. 

We feel that this aspect can best be done separately 

and in conjunction with a scombrid specialist 

(Collette). In addition to collections made by us, 

most of the other collections (about one-third of the 

total) were made by Collette in the course of his 

studies on scombrids. 

KNOWN HOST SPECIES WITH DISTRIBUTION.—Below 

is a list of the known species of the scombrid subfamily 

Scombrinae with their general geographic distributions. 

Although there are 2 recognized species of 

Auxis, we were advised (B. B. Collette, pers. com.) 

that, for purposes of this study, we should refer to 

them as "Auxis species" until the genus has undergone 

revision. References to "Scomberomorus species" pertain 

to a new species from Northern Australia to New 

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

Guinea being described by Collette and Russo. The 

following list is adapted from information in Collette 

andChao (1975) and Klawe (1977). 

Tribe SCOMBRINI Bonaparte (Mackerels) 

Rastrelliger brachysoma (Bleeker) : South China Sea east 

to Fiji 

Rastrelliger faughni Matsui: South China Sea to Indonesia 

Rastrelliger kanagurta (Cuvier): Indian and South Central 

Pacific oceans 

Scomber australasicus (Cuvier) : Western & Central Pacific 

and Socorro Island (off Mexico) 

Scomber japonicus Houttuyn: circumglobal antitropical 

Scomber scombrus Linnaeus: North Atlantic Ocean and 

Mediterranean 

Tribe SCOMBEROMORINI Starks (Seerfishes) 

Acanthocybium solandri (Cuvier): circutntropical and subtropical 

Granimatorcynus bicarinatus (Quoy and Gaimard): Red Sea 

and Western Pacific Ocean 

Scomberomorus brasiliensis Collette, Russo, and Zavala- 

Camin: Western Atlantic (Yucatan to south Brazil) 

Scomberomorus cavalla (Cuvier): Tropical western Atlantic 

Ocean 

Scomberomorus commerson (Lacepede): Indian and Western 


Scomberomorus concolor (Lockington): Gulf of California 

and Monterey Bay 

Scomberomorus guttatus (Bloch and Schneider): Indian and 

western Pacific oceans 

Scomberomorus koreanus (Kishinouye): Indian and western 


Scomberomorus lineolatus (Cuvier): India to Indonesia 

Scomberomorus maculatus (Mitchill): Western Atlantic 

(Yucatan to Massachusetts) 

Scomberomorus multiradiatus Munro: Gulf of Papua 

Scomberomorus niphonius (Cuvier): Northwestern Pacific 

Ocean (China to Japan) 

Scomberomorus plurilineatus Fourmanoir: Mozambique 

Channel (coastal) and South Africa 

Scomberomorus queenslandicus Munro: Northern Australia 

Scomberomorus regalis (Bloch): Western Atlantic (especially 

West Indies) 

Scomberomorus semifasciatus (Macleay): Northern Australia 

to New Guinea 

Scomberomorus sierra Jordan and Starks: Eastern Pacific 

Ocean 

Scomberomorus sinensis (Lacepede): Northwestern Pacific 

Ocean 

Scomberomorus tritor (Cuvier): Gulf of Guinea to Mediterranean 

Scomberomorus species: Northern Australia to New Guinea

NUMBER 3 1 1 

Tribe SARDINI Jordan and Evermann (Bonitos) 

Allothunnus fallai Serventy: circumglobal in southern hemisphere 

Cybiosarda elegans (Whitley): Northern Australia to New 

Guinea 

Gymnosarda unicolor (Riippell) : Indian to central Pacific 

oceans 

Orcynopsis unicolor (Geoffroy St. Hilaire): Gulf of Guinea 

to Mediterraean 

Sarda australis (Macleay): southeast coast of Australia 

Sarda chiliensis (Cuvier), 2 subspecies: Northeastern (S. c. 

lineolata) and southeastern Pacific Ocean (S. c. chiliensis) 

Sarda orientalis (Temminck and Schlegel): Indian and 

Pacific oceans (coastal) 

Sarda sarda (Bloch): Atlantic Ocean and Mediterranean 

Tribe THUNNINI Starks (Tunas) 

Auxis rochei (Risso) : circumtropical 

Auxis thazard Lacepede: circumtropical 

Euthynnus affinis (Cantor): Indian and Pacific oceans 

Euthynnus alletteratus (Rafinesque): Atlantic Ocean 

Euthynnus lineatus Kishinouye: Eastern Pacific Ocean 

Katsuwonus pelamis (Linnaeus): circumtropical 

Thunnus alalunga (Bonnaterre): circumtemperate and 

tropical 

Thunnus albacares (Bonnaterre): circumsubtropical and 

tropical 

Thunnus atlanticus (Lesson): Western Atlantic Ocean 

Thunnus maccoyii (Castelnau): circumglobal in southern 

hemisphere 

Thunnus obesus (Lowe): circumsubtropical and tropical 

Thunnus thynnus (Linnaeus): North Pacific, North and 

South Atlantic (non-tropical) 

Thunnus tonggol (Bleeker): Indian and Western Pacific 

SPECIMENS EXAMINED.—Scombrids housed in or 

borrowed from the following places were examined 

for copepods by us or Collette. 

Australian Museum, Sydney (AMS) 

Academy of Natural Sciences, Philadelphia, Pa. 

British Museum (Natural History), London 

Bernice P. Bishop Museum, Honolulu, Hawaii 

California Academy of Sciences, San Francisco 

Chesapeake Biological Laboratory, Solomons, Md. 

CSIRO Marine Biological Laboratory, Cronulla, N.S.W., 

Australia 

Dominion Museum, Wellington, New Zealand 

Field Museum of Natural History, Chicago, 111. 

Hebrew University, Jerusalem 

Los Angeles County Museum of Natural History, Los 

Angeles, Calif. 

Museo Argentina de Ciencias Naturales, Buenos Aires 

Museum of Comparative Zoology, Harvard (MCZ) 

Muslum National d'Histoire Naturelle, Paris 

Museo di Storia Naturale, Genoa 

Museo de La Specola, Universita di Firenze, Florence 

Naturhistorisches Museum, Vienna 

National Museum of Natural Sciences, Ottawa 

Northwest Fisheries Center Auke Bay Laboratory, National 

Marine Fisheries Service, NOAA, Auke Bay, Alaska 

Queensland Museum, Brisbane 

Rijksmuseum van Naturlijke Histoire, Leiden 

J.L.B. Smith Institute of Ichthyology, Rhodes University, 

Grahamstown 

South African Museum, Cape Town 

Sea Fisheries Research Station, Haifa, Israel 

Scripps Institution of Oceanography, La Jolla, Calif. 

Senckenberg Museum, Frankfurt-am-Main 

Southeast Fisheries Center, National Marine Fisheries Service, 

NOAA (formerly Tropical Atlantic Biological Laboratory), 

Miami, Fla. 

Institute of Fisheries, University of British Columbia, Vancouver 

University of California, Los Angeles 

Rosenstiel School of Marine and Atmospheric Science, 

Miami, Fla. 

University of Michigan Museum of Zoology, Ann Arbor 

Smithsonian Institution, Washington, D.C. (USNM, former 

United States National Museum, collections in the National 

Museum of Natural History, Smithsonian Institution) 

Western Australia Museum, Perth 

Woods Hole Oceanographic Institution, Woods Hole, Mass. 

Zoological Museum, Copenhagen 

Zoological Museum, Oslo 

Representative collections of the following species 

will be deposited in the British Museum (Natural History, 

London), Museum National d'Histoire Naturelle 

(Paris), Australian Museum (Sydney), and California 

Academy of Sciences (San Francisco): Holobomolochus 

divaricatus, H. nudiusculus, Unicolax collateral!*, 

U. reductus, U. ciliatus, Ceratacolax euthynni, Shiinoa 

inauris, S. occlusa, Caligus coryphaenae, C. omissus, 

C. bonito, C. productus, C. cybii, Elytrophora brachyptera, 

E. indica, Pseudocycnus appendiculatus, 

Pseudocycnoides armata, and P. buccata. All type 

material and remaining collections are deposited in 

the Smithsonian Institution. 

ACKNOWLEDGMENTS.—This work began with a 

small collection of copepods parasitic on western 

Atlantic pelagic fishes given to one of us (R. Cressey) 

by Arthur Humes as a basis for a graduate research 

project in 1960. The initial collection was made by 

Robert Gibbs and Bruce Collette during various 

cruises of the R. V. Delaware, consisting primarily of 

copepods from sharks and scombrids. The shark parasites 

formed the basis for R. Cressey's doctoral dissertation 

revising the family Pandaridae (under the

direction of A. Humes). The remaining collection 

stimulated a continuing interest in scombrid copepods 

by the first author and later by the second. 

We thank the scientists and staffs of the many 

institutions cited above for making fish specimens 

available for examination for parasitic copepods. 

Through the years many people have sent us material, 

for which we are grateful. Outstanding among 

these are: Bruce B. Collette (National Marine Fisheries 

Service, Washington), Robert H. Gibbs (Smithsonian 

Institution, Washington), Arthur G. Humes 

(Boston University), Richard Shomora (National 

Marine Fisheries Service, Honolulu), and Vladimir 

Walters (National Marine Fisheries Service, Miami). 

Robert Gibbs was our principle consultant concerning 

problems in scombrid taxonomy during the early 

years of the work. Later, Bruce Collette began an 

intensive study of scombrid taxonomy and during the 

course of his work spent many hours gathering copepods 

for us. We extend a special thanks to him for 

his enthusiastic and continued support 

We were assisted in the illustrations by Nancy Zacks 

who rendered Figures 75-79 and 95 and by Michelle 

Wilcox who did the illustrations for Figures 82-84. 

Cynthia Hemmings assisted in the final preparation 

of the manuscript. The staff of the SEM laboratory 

all participated at various times in the preparation of 

the SEM photographs (Walter Brown, Mary Jacque 

Mann, and Susann Braden). We thank all of them 

for their assistance. 

The manuscript was reviewed by Bruce Collette and 

Brian Kensley and we are grateful for their many 

helpful suggestions. 

Holobomolochus Vervoort, 1969 

Holobomolochus divaricatus, new species 

FIGURES 1-4, ba-e, 96, 109-113 

MATERIAL EXAMINED. — Holotype 9 (USNM 

172244), allotype $ (USNM 172245) paratypes 11 9 

(USNM 172246) from the nasal sinuses of Scornberomorus 

brasiliensis (USNM 188424) from Brazil. 

In addition, there are 13 collections containing 61 $ 

2$ from S. brasiliensis from Panama (Atlantic), 

Colombia (Atlantic), Brazil and Argentina; 25 collections 

containing 70 $ 40 $ from S. maculatus from 

Cape Cod, New Jersey, Maryland, South Carolina, 


Georgia, Florida, Mississippi, Alabama, Louisiana, 

Texas, Panama (Atlantic), Surinam; 11 collections 

containing 28 9 20 $ and 3 immatures from S. regalis 

from Florida, Cuba, Haiti, Puerto Rico, Colombia 

(Atlantic), Venezuela and Surinam. All copepods 

were collected from the nasal sinuses of the host fish. 

FEMALE.—Body form as in Figure la. Total length 

2.62 mm, greatest width (widest part of cephalon) 

1.20 mm. Genital segment (Figure \b) wider than 

long (348 X 483 jxm) with 3 dorsal setae at area of 

egg sac attachment. Abdomen 3-segmented, segments 

measuring (1 X w) 224 X 318 /*m, 153 X 265 ^m, 

147 X 230 /*m respectively; ventral surface of last 

segment with 2 patches of prominent spinules (see 

figure \c). Caudal rami (Figure \c) longer than wide 

(129 X 94 /*m) with six setae; longest seta 590 /*m. 

First antenna (Figure Id) 5-segmented, second segment 

incompletely divided; first two segments with 15 

prominent plumose setae; segments 4 and 5 with one 

aesthete each. Second antenna (Figure 2a) of usual 

bomolochid type; terminal segment with numerous 

rows of spinules, 2 processes with spinules, 3 terminal 

setae and 4 elongate spines. Mandible, paragnath, 

first maxilla and second maxilla as in Figure 2b. 

Maxilliped (Figure 2c) terminal segment with prominent 

curved claw with slight projection on outer curve, 

and 3 plumose setae; basal segment with 1 plumose 

seta. 

Leg 1 (Figure 2d) exopod with stout, heavily plumose 

seta on outer corner; basipod with 2 patches of 

hairs; exopod 2-segmented, first segment with prominent 

rugose spine on outer distal corner, second segment 

with small spine at outer proximal comer, 

minute spine at outer distal corner and 2 stout flagellated 

spines best viewed dorsally, terminal to inner 

edge of segment with 6 plumose setae; endopod 

3-segmented, first segment with large patch of hairs 

and inner seta, second segment with smaller patch of 

hairs and inner seta, third segment with 5 terminal 

setae, all setae heavily plumose, outer edge of endopod 

segments haired. Legs 2-4 biramous, rami 3-segmented. 

Leg 2 (Figure 2e) coxopod with short row of spinules 

at outer distal corner; basipod with dorsal seta; exopod 

first segment with hairs on outer edge and spine on 

outer distal corner, second segment with inner seta 

and outer spine, slender spinules at base of spine, third 

segment with 6 inner to terminal setae and 3 outer 

spines; exopod spines broad at base, tapering distally 

with small terminal flagellum, fine hairs on both

NUMBER 3 1 1 

edges of spines; endopod first segment with inner 

seta and row of fine hairs along outer half of distal 

border, second segment similar except with 2 inner 

setae, third segment with 3 inner to terminal seta and 

2 outer spines similar to, but smaller than, exopod 

spines, outer edges of endopod segments haired; interpodal 

plate with long slender spinules. Leg 3 (Figure 

3a) coxopod with row of small spinules at outer distal 

corner; basipod with dorsal seta; exopod segments 

heavily sclerotized; first segment with outer spine, 

second segment with outer spine and inner seta, third 

segment with 2 outer spines and 6 terminal to inner 

setae, exopod spines heavily sclerotized, spines on 

second and third segments of about equal length, 

spine on first segment slightly shorter than others; 

endopod similar to leg 2 except third segment with 

only 2 setae; interpodal plate with short, stout spinules. 

Leg 4 (Figure 3b) coxopod and basipod similar to 

leg 3; exopod first segment with spinules along ventrolateral 

edge, hairs on inner edge and spine on outer 

distal corner, second segment with inner seta and 

outer spine, slender spinules at base of seta, shorter, 

stouter spinules at base of spine, third segment with 

6 inner to terminal setae, 2 outer spines, slender 

spinules at bases of 2 innermost setae, stouter spinules 

at bases of spines, spines slender, elongate with fine 

hair on edges and fine terminal flagellum; endopod 

first segment with patch of slender spinules near outer 

distal corner and inner seta spinulose along distal twothirds, 

second segment similar to first, third segment 

elongate, broadened distally with 2 terminal patches 

of stout spinules and 2 spines flanking terminal seta, 

spines elongate, of about equal length and finely haired 

on edges, seta spinulose along distal half; interpodal 

plate with short, stout spinules. Leg 5 (Figure 3c) 

basal segment with dorsal seta and small patch of 

slender spinules; free segment with 3 patches of 

slender spinules, one spine at about mid-point of outer 

margin and 2 spines flanking terminal, naked seta, 

spines elongate, simple with finely haired edges. Leg 

6 (see Figure 16) represented by 3 naked setae at 

area of egg sac attachment. 

MALE.—Body form as in Figure 3d. Total length 

1.18mm, greatest width 0.50mm (measured at widest 

part of cephalon). Genital segment (Figure 4a) longer 

than wide (236 X 182 pm). Abdomen (Figure 4a) 

2-segmented, segments measure (length X width) 

88 X 118 /un, 82 X 106 /un, respectively; ventral 

surface of second segment with 2-3 rows of promi- 

nent spinules forming a proximal transverse band and 

2 distal patches of spinules. Caudal rami (Figure 46) 

slightly longer than wide (41 X 35 /an) with 6 setae; 

longest seta 572 /tm. 

First antenna (Figure 4c) 5-segmented, second segment 

incompletely divided, antenna similar to female, 

but segments not as elongate. Second antenna, mandible, 

paragnath, first maxilla, second maxilla similar to, 

but proportionately smaller than, female. Maxilliped 

(Figure 4d*) second segment with 2 short setae near 

inner mid-margin, inner surface of segment with patch 

of elongate, blunt-tipped spinules and patch of smaller, 

rounder spinules; terminal claw with 2 setae and 

numerous closely spaced "teeth" along entire inner 

edge. Legs 1-4 biramous, rami 3-segmented except 

leg 4 endopod. 

Leg 1 (Figure Ae) coxopod heavily haired along 

outer distal edge; basipod with dorsal seta and ventral 

ornamentation as indicated in figure; exopod first segment 

with row of elongate spinules near base of spine 

on outer distal corner, second segment similar to first 

with the addition of an inner seta, third segment with 

4 inner to terminal setae and 3 outer spines, proximal 

2 spines similar to those of first and second segments, 

finely haired on edges, and with terminal flagellum 

and with row of spinules near base, distal spine elongate 

with row of very fine hairs along outer edge; 

endopod first segment enlarged, covering inner portion 

of exopod, stout hairs along distal edge and an inner 

seta, second segment only about half as wide as first 

segment with inner seta, small spinule on outer distal 

corner and row of fine hairs along outer half of distal 

edge, third segment with 4 terminal setae and small 

outer spinule, outer half of segment with 2 converging 

rows of small, rectangular, platelike spinules, outer 

edges of endopod segments heavily haired. Leg 2 

coxopod with short row of small spinules on outer 

distal corner; basipod with dorsal seta; exopod first 

segment with irregular rows of small bumplike spinules 

along outer edge, short row of slender spinules at base 

of spine on outer distal corner, second segment with 

inner seta and row of spinules at base of outer spine, 

third segment with 5 inner to terminal setae and 3 

outer to terminal spines, proximal 2 spines similar to 

spines of first and second segments, finely haired on 

edges with terminal flagellum and short row of spinules 

near base, terminal spine elongate with row of 

very fine hairs along outer two-thirds; endopod first 

segment with outer seta and row of hairs along outer

distal edge, second segment similar to first but with 

two inner setae, third segment with 3 inner to terminal 

setae, short row of spinules at base of 2 outer spines, 

inner spine about twice as long as outer, outer edges 

of endopod spines heavily haired. Leg 3 (Figure 5a) 

similar to leg 2 except endopod third segment with 

2 rather than 3 setae. Leg 4 (Figure 56) coxopod 

and basipod similar to leg 3; exopod similar to leg 3 

with the following exceptions: spines more slender; 

second segment with no spinules at base of outer spine; 

third segment with 4 rather than 5 setae; endopod 

2-segmented, first segment similar to leg 3, second 

segment elongate with stout spinules distally and near 

outer distal edge, 2 spines flanking naked, terminal 

seta, spines spinulose along edges, lacking terminal 

flagella, inner spine slightly longer than outer. Leg 5 

(Figure 5c) basal segment with dorsal seta and small 

patch of spinules at outer distal corner; free segment 

with patch of elongate spinules covering inner half of 

lower two-thirds of segment, terminally with inner 

spinulose spine and outer seta. 

ETYMOLOGY.—The Latin divaricatus ("spreading 

asunder at wide angle") refers to the characteristic 

appearance of the caudal rami of the female. 

REMARKS.—Females of this species can be distinguished 

from all previously described members of the 

genus by the nature of the exopod spines of leg 3 (see 

Discussion) ; they can be separated from the females of 

the two following species by the ornamentation of the 

ventral surface of the caudal rami and last abdominal 

segment. Holobomolochus divaricatus females have 

two large patches of stout spinules on the last abdominal 

segment, their caudal rami have no surface ornamentation; 

H. nudiusculus females have relatively 

small patches of minute hairs on the last abdominal 

segment; H. asperatus females have 2 large patches 

of hairs on the last abdominal segment and a patch 

of hairs on the ventral surface of each caudal ramus. 

In addition the distal spine of the third exopod segment 

of leg 3 is only as long as, or shorter than, the 

adjacent spine; in H. nudiusculus and H. asperatus 

this spine is longer than the adjacent spine and more 

outwardly curved than other spines of the ramus (see 

Figures 3a, 6c, 8c). 

This copepod is a common parasite on Scomberomorus 

brasiliensis, S. maculatus, and S. regalis in the 

Western Atlantic from Cape Cod to southern Brazil. 


Holobomolochus nudiusculus, new species 

FIGURES 5d-e, 6, la-b, 96 


172247), allotype $ (USNM 172248), paratypes 

33 5 IS (USNM 172249) from the nasal sinuses of 

Scomberomorus sierra from Buenaventura, Colombia 

(USNM 218565). In addition 13 collections containing 

30 9 1

NUMBER 3 1 1 

ramus (see Figure 76). Caudal rami (Figure 7 b) 

longer than wide (47 X 29 /un), each with small 

ventral patch of slender spinules and 6 setae (longest 

531 /an). 

Cephalic appendages and legs 1-5 as in H. divaricatus 

male. 

ETYMOLOGY.—The Latin nudiusculus ("somewhat 

naked") refers to the small patches of minute hairs 

on the ventral surface of the last abdominal segment 

and the naked surface of the female caudal rami. 


from H. divaricatus and H. asperatus by the 

ornamentation of the ventral surface of the last abdominal 

segment. Holobomolochus nudiusculus has 2 

small patches of minute hairs on the last abdominal 

segment; H. divaricatus has 2 large patches of stout 

spinules; H. asperatus has 2 large patches of hairs on 

the last abdominal segment as well as a patch of hairs 

on each caudal ramus, unlike the caudal rami of H. 

nudiusculus. In addition the third segment endopods 

of legs 2 and 3 have small spinules near the bases of 

the outer spines; this ornamentation is lacking in H. 

asperatus and H. divaricatus. Females of this species 

can be distinguished from all other species of 

Holobomolochus by the nature of the exopod spines 

of leg 3 (see "Discussion"). 

Holobomolochus nudiusculus seems to be restricted 

to the eastern Pacific but common on its 2 hosts 

Scomberomorus concolor and S. sierra from California 

to Peru. 

Holobomolochus asperatus, new species 

FIGURES Ic-g, 8, 96 


172242) and paratypes 2$ (USNM 172243) from 

the nasal sinus of Scomberomorus cavalla (MCZ 

17182) from Cuba. In addition, 9 collections containing 

10 9 are from the nasal sinus of S. cavalla from 

Georgia, Florida, Texas, Cuba, Trinidad, and Brazil. 

FEMALE.—Body form as in Figure 7c. Total length 

1.79 mm, greatest width 0.75 mm; length of cephalon 

0.57 mm. Genital segment (Figure 7d) wider than 

long (212 X 265 /un). Abdomen 3-segmented, segments 

measure (1 X w) 177 X 206 ^n, 129 X 177 

fim, and 112 X 159 /*m respectively; ventral surface of 

last abdominal segment (see Figure 7e) with 2 prominent 

patches of hairlike spinules. Caudal rami (Figure 

7e) longer than wide (82 X 53 /un), each ramus with 

six setae and a patch of hairlike spinules on its ventral 

surface; longest seta 768 /*m. 

First antenna (Figure 7/) 7-segmented with an 

aesthete on segments 6 and 7. Second antenna, mandible, 

paragnath, first maxilla, second maxilla, and 

maxilliped similar to corresponding appendages in H. 

divaricatus. 

Leg 1 similar to that of H. divaricatus except endopod 

and basipod lack hairs found in H. divaricatus. 

Leg 2 (Figure 86) similar to that of H. divaricatus 

except exopod second segment lacks spinules at base 

of outer spine; endopod first and second segments each 

have a minute patch of small hairs on outer distal 

corner, outer spines on third segment are small. Leg 3 

(Figure 8

8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY 

on the rami of H. divaricatus or H. nudiusculus) and 

the lack of prominent patches of spinules on leg 4 

(present in H. divaricatus and H. nudiusculus). 

Holobomolochus asperatus can be distinguished from 

all other members of the genus by the nature of the 

leg 3 exopod spines (see Discussion). 

This species has been collected only from Scomberomorus 

cava.Ua from Georgia to northern Brazil. 

Although its range overlaps that of H. divaricatus, the 

2 parasites have never been collected from a common 

host species. 

Discussion 

The 3 species of Holobomolochus described here all 

have highly developed spines on the exopod of leg 3 

and the exopod segments themselves are heavily 

sclerotized. This character separates these 3 species 

from all previously described Holobomolochus. 

Although we do not feel that this difference alone 

warrants assigning these new species to a new genus, 

we feel that an improved understanding of the genus 

might result in a better basis for doing so. No other 

species of Holobomolochus has been collected from 

the nasal sinuses of the hosts. This unusual habitat 

may account for the unique nature of these 3 species. 

Vnicolax, new genus 

DIAGNOSIS.—Bomolochidae. Body form typical of 

family. Thoracic segments bearing legs 3-5 free in 

female, thoracic segments bearing legs 2-5 free in 

male. Abdomen 3-segmented in female, 2-segmented 

in male. Caudal rami in both sexes with 6 setae, 2 

much longer than other 4. First antenna 6- or 7-segmented 

in female, fourth seta on basal segment modified 

to form heavily sclerotized straight or nearly 

straight spine; first antenna in male 6-segmented with 

no modified setae. Second antenna and oral appendages 

typical of family. Maxilliped hook in female 

without accessory process; maxilliped basal segment 

in male with numerous small spinules. Legs 1-4 

biramous in both sexes; rami of legs 1-3 and leg 4 

exopod 3-segmented, leg 4 endopod 2- or 3-segmented. 

Female leg 2 endopod second segment with 2 inner 

setae, leg 3 endopod second segment with 1 inner seta. 

Rami of leg 1 in both sexes broad and flattened. 

TYPE SPECIES.—Unicolax collateralis new genus, 

new species. 

ETYMOLOGY.—The Latin, uni-colax (with "colax"), 

the suffix "colax" is common in other genera of the 

family and the name alludes to the new genus "with" 

others of the family. 

DISCUSSION.—Unicolax can be separated from all 

the other bomolochid genera except Bomolochus by 

the modified fourth seta on the basal segment of the 

first antenna. Holobomolochus, Acanthocolax, Pseudoeucanthus, 

Orbitacolax, Pseudorbitacolax, Pumiliopes, 

and Pumiliopsis have no modified setae or cuticular 

process on the first antenna; Boylea has the fifth seta 

modified; in Nothobomolochus the third, fourth, and 

fifth setae are modified; in Dicrobomolochus the second 

and third setae are modified; Ceratacolax and 

Tegobomolochus have a cuticular process in addition 

to the usual 15 plumose setae on the first antenna. 

Bomolochus is characterized by having the fourth 

seta of the first antenna modified, but in Bomolochus 

this seta is sharply curved and lightly sclerotized, 

whereas in Unicolax it is straight or nearly so, and 

heavily sclerotized. Unicolax can further be separated 

from Bomolochus as the rami of the first leg of Unicolax 

males are flattened and broad as in the females; 

the first leg of Bomolochus males does not resemble 

that of Bomolochus females. 

Unicolax has, so far, only been collected from the 

nasal sinuses of its hosts. 

Vnicolax collateralis, new species 

FIGURES 9-13, 97, 114-116* 


172256), allotype $ (USNM 172257) paratypes 12 9 

23 d (USNM 172258) from the nasal sinus of 

Euthynnus alletteratus (USNM 203838) from St. 

George Bay, Lebanon. The following collections from 

Auxis species: 2 9 from Woods Hole, Mass.; 2 S 

from St. George Bay, Lebanon; 2 9 3$ from 

Ghardaqa, Egypt; 2 collections containing 7 9 3$ 

from Hong Kong; 2 collections containing 2 9 1 $ 

from Japan; 7 collections containing 12 9 4$ from 

the Philippines; 2 9 from Chusan, China; 1 9 from 

Hawaii. One collection containing 5 9 from Cybiosarda 

elegans from Brisbane, Australia. The following 

collections from Euthynnus affinis: 2 9 2 3 from Elat, 

Israel; 2 9 from Mozambique; 2 9 from Madagascar; 

1 S from the Seychelles; 4 9 from the Arabian Sea 

24°N, 67°E; 2 9 from Java; 16 collections containing

NUMBER 311 

29 9 9 $ from the Gulf of Thailand; 2 9 from Formosa; 

2 2 from Hong Kong; 4 collections containing 

8 9 1 $ from the Philippines; 1 2 from Palau; 1 9 

from Okinawa; 1 $ from Tokyo; 4 2 7 $ from Brisbane, 

Australia. The following collections from 

Euthynnus alletteratus: 1 9 from the Caribbean 

(9°11'N, 77°50'W); 2 9 from Brazil; 4 collections 

containing 29 9 16 $ from St. George Bay, Lebanon. 

The following collections from Euthynnus lineatus: 1 

9 from Galapagos; 1 9 from Lower California; 1 9 

from Mexico (Pacific); 2 9 from Costa Rica (Pacific). 

One collection containing 1 9 from Orcynopsis unicolor 

from St. George Bay, Lebanon. Two collections 

containing 5 9 3

10 

with inner seta and outer spine, third segment with 5 

inner to terminal setae and 3 terminal to outer spines, 

exopod spines similar to corresponding spines of leg 2, 

except leg 3 lacks haired spine present on third segment 

of leg 2, all setae plumose; endopod first segment 

with inner seta and row of short hairs along 

distal edge, second segment similar to first, third segment 

with 2 inner to terminal setae and 2 terminal to 

outer spines, each with serrate edges and terminal 

flagellum, all setae plumose, outer edges of endopod 

segments haired. Leg 4 (Figure life) coxopod with 

central patch of small spinules and row of short hairs 

along mid-lower edge; basipod lacking ornamentation; 

exopod similar to leg 3 except first segment lacks dorsolateral 

spinules; endopod first segment with inner seta 

and row of fine hairs along distal edge, seta shorter 

than corresponding seta of leg 2, constricted slightly 

at midpoint, plumose distally finely serrate proximally, 

hairs on distal edge of segment slightly longer than 

corresponding hairs of leg 2, second segment similar 

to first, third segment with row of fine hairs distally 

and 2 spines flanking terminal seta, spines broadly 

serrate and with terminal flagellum, seta finely serrate 

on distal half, outer edges of endopod segments haired. 

Leg 5 (Figure lie) basal segment with outer dorsal 

seta and dense patch of short spinules on outer distal 

corner; free segment with 3 dense patches of short 

spinules, one spine on mid-outer margin and 2 spines 

flanking one terminal naked seta, spines with serrate 

edges and terminal flagellum. Leg 6 (see Figure 9b) 

represented by 2 long and one shorter seta on genital 

segment. 

MALE.—Body form as in Figure lid. Total length 


0.20 mm. Genital segment (Figure 11*) longer than 

wide (236X218 fan). Abdomen (see Figure lie) 

2-segmented, segments measuring 94 X 112 /*m and 

70 X 88 /*m respectively. Last abdominal segment 

ornamented ventrally with anterior row of fine spinules 

and 2 large patches of fine hairs (see Figure 12a). 

Caudal rami (Figure 12a) longer than wide (53 X 

35 fan) ; each ramus with ventral patch of fine hairs 

and 6 setae; longest seta 295 /un. 

First antenna (Figure 12b) 6-segmented. First segment 

with 5 plumose setae and no indication of 

modified spine present on female; fifth and sixth segments 

each with one aesthete. Second antenna, mandible, 

paragnath, first and second maxillae similar to 

female. Maxilliped (Figure 12c) with one seta and 


large patch of small, rounded spinules on second segment; 

last segment with 2 setae near base and row 

of teeth along inner edge. 

Legs 1-4 biramous. Leg 1 (Figure 12*/) coxopod 

with inner seta and spinules as indicated in Figure 

I2d; basipod with outer seta, blunt process between 

insertion of rami, and inner process sclerotized at 

base with terminal, fringed membrane, spinules near 

base of this process; exopod 3-segmented, first segment 

with outer spine, second segment with outer spine and 

inner seta, third segment with 2 outer spines and 5 

setae; endopod first segment with inner seta, distal 

outer edge of segment with row of hairs, second segment 

similar to first, third segment with 5 setae 

terminally and one outer spine, minute spinules near 

bases of setae; all setae heavily plumose. Leg 2 (Figure 

12tf) coxopod with short row of spinules on outer 

distal corner and row of hairs on mid-distal edge; 

basipod with naked seta dorsally near outer edge; 

exopod first segment with outer distal serrate spine 

with terminal flagellum, row of fine spinules near 

base of spine, dorso-lateral surface of segment with 

several rows of short, rounded spinules, second segment 

with inner seta and outer spine similar to that 

on first segment, third segment with 5 inner to terminal 

setae and 3 terminal to outer spines, proximal 

2 spines similar to those on first and second segments, 

distal spine elongate with broad outer serrations and 

short inner hairs; endopod first segment with inner 

seta and short row of hairs along distal edge, second 

segment similar, third segment with 3 inner to terminal 

setae and 2 outer spines, each finely haired on 

edges and with terminal flagellum; outer edge of 

endopod segments heavily haired. Leg 3 (Figure 13a) 

similar to leg 2, with following exceptions; exopod 

second segment lacks outer spine and fine spinules; 

endopod third segment with only 2 setae, inner spine 

about twice as long as outer, ornamentation on edges 

of each spine serrate rather than haired as in leg 2. 

Leg 4 (Figure 13ft) coxopod, basipod, and exopod 

similar to leg 3 except exopod third segment with 

only 4 setae; endopod 2-segmented, first segment with 

inner seta, second segment elongate with row of fine 

hairs distally and 2 spines flanking terminal seta, 

spines finely serrate with terminal flagella, inner spine 

about one-third longer than outer, seta finely serrate 

along distal half. Leg 5 (Figure 13c) basal segment 

with outer dorsal seta and dense patch of spinules on 

outer distal corner; free segment with terminal outer

NUMBER 3 1 1 11 

seta extending almost to end of genital segment, and 

inner spine broadly serrate on edges and with terminal 

flagellum; segment with dense patch of spinules on 

distal half. 

ETYMOLOGY.—The Latin collateredis ("standing 

side by side") alludes to its occurrence in collections 

with U. mycterobius. 


from U. mycterobius, U. ciliatus, and 

U. reductus principally by the nature of the spines 

on the exopod third segment of leg 2 as the proximal 

spine of U. collateralis is finely haired on both edges 

and the next two spines have broad serrations on both 

side. This is unlike the ornamentation of corresponding 

spines on any of the other above species. Unicolax 

collateralis can be distinguished from U. anonymous, 

which it most closely resembles, by having 3 spines, 

5 setae on the exopod third segment of leg 4; U. 

anonymous has 3 spines, 4 setae on the corresponding 

segment. The ventral surface of the endopod segments 

of legs 2-4 are densely haired in U. anonymous, 

whereas U. collateralis has only a single row of hairs 

along the distal edge of some of its endopod segments. 

Males of this species can be distinguished from those 

of U. anonymous by the ornamentation of leg 5. Unicolax 

collateralis has a patch of stout spinules that 

extends from about the middle of the free segment 

distally and around the entire distal edge; in U. 

anonymous the spinules are distinctly elongate and 

the patch extends distally from above the middle of 

the segment and does not continue around to the 

outer distal edge. Males can be distinguished from 

U. mycterobius, U. ciliatus, and U. reductus by the 

ornamentation of the ventral surface of the last abdominal 

segment. In U. collateralis this segment has 

a single, even row of spinules near the anterior 

border, and 2 large patches of hairs. Unicolax mycterobius 

has similar ornamentation, but the anterior 

row of spinules is irregular and the spinules are 

minute. 

This copepod is circumglobal and occurs in the 

nasal sinuses of a variety of scombrid hosts (species 

of Auxis, Cybiosarda, Euthynnus, Orcynopsis, and 

Sarda). It is often collected with U. mycterobius, 

which parasitizes some of the same hosts (species of 

Auxis and Euthynnus). 

Unicolax anonymous (Vervoort, 1965), new 

combination 

FIGURES 14-16, 98, 116fr—/, 117 

Parabomolochus anonymous Vervoort, 1965:3. 

MATERIAL EXAMINED.—Two collections containing 

319 2 S from the nasal rosettes of Euthynnus alletteratus 

from Ghana and the Gulf of Mexico. 

FEMALE.—Body form as in Figure 14a. Total length 

of cephalon 0.37 mm. Genital segment wider than 

long (177 X 247 /*m). Abdomen 3-segmented, segments 

measure 82 X 153 /*m, 53 X 141 /*m, 70 X 

123 /urn length X width respectively. First 2 segments 

without ornamentation, last segment with 2 patches 

of fine hairs ventrally. Gaudal rami each with patch of 

fine hairs and 6 setae; longest 171 ^m. 

First antenna (Figure 146) 7-segmented. Fourth 

seta on first segment modified to form heavily sclerotized 

spine; one aesthete on each of last 2 segments. 

Second antenna (Figure 14c) 3-segmented, last segment 

with several rows of hooklike spinules, 4 setae, 

and 4 hooklike terminal spines. Mandible, paragnath, 

first maxilla, second maxilla as in Figure 14a\ Paragnath 

with outer row of teethlike spinules and long 

hairs; first maxilla with 3 long plumose setae and 

one short, naked seta. Maxilliped (Figure 14e) with 

one plumose seta on basal segment, 3 plumose setae 

on last segment; hook slightly curved, without accessory 

process. 

Legs 1-4 biramous, each ramus 3-segmented except 

leg 1 exopod. Leg 1 (Figure 15a) coxopod with 

broad inner seta; basipod with outer seta, one small 

blunt spinule near lower inner edge and two large 

patches of hairs; exopod first segment with outer 

spine, second segment with 2 short outer spines and 

6 terminal to inner setae; endopod first and second 

segments each with inner seta and patch of hairs on 

lower to outer edge, third segment with 5 setae, outer 

edges of endopod segments heavily haired. Leg 2 

(Figure 156) coxopod with inner seta, outer distal 

corner with short curved spinules; basipod with outer 

seta and patch of small spinules between insertion of 

rami; exopod first segment with long, slender spinules 

on outer edge and stout broadly serrate spine with 

terminal flagellum on outer distal corner, second segment 

with inner seta and outer spine similar to that 

of first segment, third segment with 5 inner to terminal 

setae and 4 terminal to outer spines, proximal 

spine with short hairs along both sides and terminal


flagellum, next 2 spines similar to those of first and 

second segments, distal spine elongate with broad 

outer serrations and short inner hairs; endopod first 

segment with inner seta and large patch of short 

hairs distally, second segment enlarged, with 2 inner 

setae and patch of short hairs on outer distal corner, 

third segment narrower than second with 3 inner to 

terminal setae and 2 short terminal spines each with 

hairs on edges and terminal flagellum, outer edges 

of endopod segments haired. Leg 3 (Figure 16a) 

coxopod and basipod similar to leg 2; exopod first 

segment with outer distal spine, dorso-lateral surface 

of segment with several uneven rows of short spinules 

giving bumpy appearance, second segment with inner 

seta and outer spine, third segment with 5 inner to 

terminal setae and 3 terminal to outer spines, exopod 

spines similar to corresponding spines of leg 2, except 

leg 3 lacks haired spine present on third segment of 

leg 2; endopod first segment with inner seta and 

patch of hairs covering distal half of segment, second 

segment with inner seta and patch of hairs on outer 

distal corner, third segment with 2 inner to terminal 

setae and 2 terminal to outer spines, inner spine 

longer, each with serrate edges and terminal flagellum, 

outer edges of endopod segments haired. Leg 4 

(Figure 166) coxopod with row of short hairs on 

mid-distal edge; basipod with outer seta; exopod 

first segment with spine on outer distal corner, second 

segment with inner seta and outer spine, third segment 

with 4 inner to terminal setae and 3 terminal 

to outer spines, exopod spines similar to corresponding 

spines of leg 3; endopod first segment with inner 

seta, proximal half plumose, distal portion edged with 

short bristles, segment with large patch of short hairs 

distally, second segment with inner seta similar to 

that of first segment, and patch of short hairs on outer 

distal corner, third segment with 2 spines flanking 

terminal seta, spines broadly serrate distally, seta with 

short bristles on lower half, distal edge of segment 

with short spinules, outer edges of endopod segments 

haired. Leg 5 (Figure 16c) basal segment with outer 

seta and dense patch of short spinules on outer distal 

corner; free segment with 3 dense patches of short 

spinules, one spine on mid-outer margin, and 2 spines 

flanking one naked seta distally, spines each with fine 

serrations along distal half and terminal flagellum. 

Leg 6 (see Figure 14a) represented by 3 setae on 

genital segment. 



0.20 mm. Genital segment as long as wide (177 X 

177 /an). Abdomen 2-segmented, segments measuring 

(length X width) 35 X 94 /un and 59 X 70 /un. 

Caudal rami slightly longer than wide (35 X 29 /*m) ; 

longest seta 177 pin. 

All appendages and ornamentation similar to (although 

proportionately smaller than) U. collaterals 

except leg 5. 

Leg 5 (Figure 16«) similar to U. collaterals except 

patch of spinules on free segment covers lower twothirds 

of inner half of segment and does not extend 

around the entire distal edge; also, spinules on both 

basal and free segments ar*» longer and more slender 

than those of U. collateralis. 

REMARKS.—Unicolax anonymous was described by 

Vervoort (1965) as Parabomolochus anonymous from 

the nasal sinus of Euthynnus alletteratus from the 

Gulf of Guinea. 

We believe the copepods above to be the same 

species as Vervoort's even though our descriptions 

differ on some points. Vervoort (1965:6) reported 

"no spinules on the abdominal somites" of the female. 

We, however, found 2 patches of hairs on the ventral 

surface of the last abdominal segment as well as a 

ventral patch of hairs on each caudal ramus. Vervoort 

(1965:11) further reported that "the external 

margin of the first exopod segment [of legs 2-4] is 

strongly haired." We found this true only of leg 2; 

on legs 3 and 4 the first exopod segment was ornamented 

by several irregular rows of bumplike spinules. 

In spite of these differences we feel we are dealing 

with the same species for the following reasons: the 

copepods reported here are from the same host species 

and the same locality (and the Gulf of Mexico as 

well) as those described by Vervoort; Vervoort reported 

collecting Ceratocolax euthynni from the same 

individual hosts as U. anonymous, we also found 

C. euthynni in both our collections with U. anonymous; 

apart from the differences mentioned above, 

our descriptions are in agreement. 

Females of U. anonymous can be distinguished 

from U. mycterobius, U. ciliatus, and U. reductus 

by the nature of the spines on the exopod third segment 

of leg 2. The proximal spine in U. anonymous 

is finely haired on both sides while the remaining 

spines are broadly serrate; this ornamentation is unlike 

that of the corresponding spines of any other of 

the above species. Unicolax anonymous can be dis-

NUMBER 3 1 1 13 

tinguished from U. collateralis, which it most closely 

resembles, by the ornamentation of the exopod third 

segment of leg 4; U. anonymous has 3 spines, 4 setae; 

U. collateralis has 3 spines, 5 setae. Further, the 

endopod segments of legs 2-4 in U. anonymous each 

have a dense patch of hairs; U. collateralis has only 

a single row of hairs near the distal border of some 

of its endopod segments. 

Unicolax anonymous males have, on the free segment 

of leg 5, a patch of elongate spinules that extends 

from above the middle of the segment to the 

inner distal border; the spinules do not continue 

around the entire distal border of the segment. This 

ornamentation is unlike that of other species of Unicolax 

males. 

This copepod has, so far, only been reported from 

the nasal sinuses of Euthynnus alletteratus from the 

Gulf of Guinea and the Gulf of Mexico. It is a relatively 

small copepod and may, as Vervoort noted, 

frequently be overlooked. 

Unicolax mycterobius (Vervoort, 1965), 

new combination 

FIGURES 17-19, 99, 118, 119a-

14 

along distal half, spines with bristled margins and 

terminal flagellum, outer edges of endopod segments 

haired. Leg 5 (Figure 186) basal segment with small 

outer patch of short hairs and dorsal seta; free segment 

with dense patches of hairs, one naked spine 

with terminal flagellum on mid-outer margin, one 

naked terminal seta flanked by 2 spines, outer spine 

naked with terminal flagellum, inner spine with short 

bristles along margins and terminal flagellum. Leg 6 

(see Figure 17a) represented by 3 setae on genital 

segment. 

MALE.—Body form as in Figure 18c. Total length 


0.35 mm. Genital segment longer than wide (336 X 

289 /*m). Abdomen 2-segmented, segments measuring 

(length X width) 100 X 129 /an and 70 X 106 (an 

respectively; second segment (see Figure \8d) ornamented 

ventrally with anterior, irregular row of minute 

hairs and distally with 2 patches of minute hairs. 

Caudal rami (Figure I8d) longer than wide (70 X 

41 /an); each ramus with ventral patch of hairs and 

6 setae; longest seta 383 /xm. 

Cephalic appendages similar to those of U. collateralis 

male. 

Legs 1-4 biramous. Leg 1 (Figure \8e) coxopod 

with broad inner seta and 2 short rows of fine spinules, 

one on outer distal corner, one on mid-distal 

edge; basipod with outer seta, blunt process near 

insertion of rami, and inner spine sclerotized at base 

and with fringed membrane distally, a large, prominent 

patch of rounded spinules surrounding base of 

spine and extending to mid portion of segment (see 

Figure 18/); exopod first segment with stout spine on 

outer distal corner, row of fine spinules surrounding 

base of spine, second segment incompletely divided 

with 6 inner to terminal setae, one outer setiform 

spine and 2 stout outer spines (one dorsal) as indicated 

in Figure 18/; endopod 3-segmented, similar to 

U. collateralis except first 2 segments each with large 

patch of hairs on distal half, and third segment with 

spinules more prominent than those of U. collateralis 

near bases of setae. Leg 2 (Figure 19a) coxopod with 

inner plumose seta and row of hairs on mid-distal 

margin; basipod with naked seta on outer dorsal 

corner; exopod similar to that of U. collateralis except 

first segment with large patch of fine hairs 

around outer ventro-lateral portion of segment, also, 

spinules at base of proximal 4 spines of ramus stout, 

some almost half as long as accompanying spines; 


endopod similar to that of U. collateralis except first 

and second segments each with large patch of fine hairs 

along outer and distal borders, third segment with 

comparatively small patch of fine hairs along outer 

edge, and several short, irregular rows of minute spinules 

as well as 2 or 3 small patches of bumps along 

distal margin at bases of terminal 2 spines and 3 setae. 

Leg 3 (Figure 19b) similar to leg 2 with following 

exceptions: exopod first segment with 2 patches of 

ventro-lateral spinules, proximal patch hairlike, distal 

patch small, fine spinules; second segment lacks outer 

spine and accompanying basal spinules; endopod third 

segment with only 2 inner to terminal setae, outer 

spines slightly larger than those of leg 2, with inner 

spine about one-third longer than outer. Leg 4 (Figure 

19c) coxopod, basipod, exopod similar to leg 3 except 

ventro-lateral spinules of first segment all very fine 

and third segment with only 4 inner to terminal setae; 

endopod completely or incompletely divided into 3 

segments (see "Remarks"), first and second segments 

similar to those of legs 2 and 3 except second segment 

lacks inner seta, third segment with 2 or 3 short, 

irregular rows of fine spinules distally and 2 spines 

flanking terminal seta, spines of about equal length, 

each finely serrate and with terminal flagellum, seta 

finely spinulose along distal two-thirds. Leg 5 similar 

to that of U. collateralis, outer seta extending to about 

half the length of genital segment. 

REMARKS.—Vervoort (1965) first described this 

species as Parabomolochus mycterobius. His description 

was based on a single collection of 6 females and 

2 males from the nasal sinus of Auxis thazard from 

the Gulf of Guinea. Our collections have enabled us 

to add details to Vervoort's original description. 

Unicolax mycterobius females are the only known 

members of the genus in which all the exopod spines 

of legs 2-4 are edged with fine hairs; the distalmost 

spine has fine hairs on the outer edge only, all other 

exopod spines have hairs on both edges. Females of 

other species of the genus have stout serrations on 

some or all of the exopod spines of legs 2-A. The free 

segment of leg 5 of U. mycterobius females has 3 

patches of fine hairs; other species have patches of 

distinctly stout spinules on the corresponding segment. 

Males of the species can be separated principally 

by the nature of legs 1 and 4. Leg 1 basipod has a 

very large, prominent patch of rounded spinules near 

the inner distal corner. Although males of other species 

have ornamentation at the corresponding site, in

NUMBER 311 15 

no other species is the patch as large or the spinules as 

prominent as in U. mycterobius. 

Leg 4 endopod is 2 or 3 segmented, the variation 

seems to be geographical. In the 7 collections containing 

U. mycterobius males, 3 (containing 6 males) 

were from the Pacific (Tokyo and New South Wales), 

4 (containing 8 males) were from Massachusetts and 

Lebanon. Males from the Pacific have 3-segmented leg 

4 endopods with clear articulation between the second 

and third segments. Males from Massachusetts and 

Lebanon have 2-segmented endopods, yet retain evidence 

of the third segment in that they have a patch 

of fine hairs mid-way along the outer edge of the last 

segment where the articulation would be. No other 

differences were found between males of the 2 populations. 

Copepods from both populations were found 

on species of Auxis and Euthynnus; females collected 

with males from both populations did not appear to be 

different from each other. 

Males of this species also have more prominent 

spinules at the base of exopod spines of legs 2—4 and 

denser patches of hairs on endopod segments of legs 

2-4 than males of other species. 

Unicolax mycterobius occurs from the northwestern 

Atlantic to the Western Pacific and is found in the 

nasal sinuses of species of Auxis and Euthynnus. 

Unicolax ciliatus, new species 

FIGURES 20-22, 98 


172253) allotype $ (USNM 172254) and 1 9 paratype 

(USNM 172255) from the nasal sinus of Scomberomorus 

plurilineatus (USNM 273221) E. African 

Marine Fisheries, Zanzibar Channel, 6 October 1965. 

The following collections from S. commerson: 2 9 

from N.W. Coast of Madagascar; 2 9, 1 £ from 

Travancore, India; 6 9 from Cochin, India; 4 9 from 

Batavia, Java; 2 9 1 $ from Phuket, Thailand; 1 9 

from Hong Kong; 5 collections containing 23 9 1 i 

from the Philippines. The following collections from 

S. guttatus: 1 9 from Calicut, India; 1 9 from 

Padang, Sumatra; 2 9 from Batavia, Java; 1 9 from 

Singapore; 1 9 from Phuket, Thailand; 2 collections 

containing 7 9 from Hong Kong; 3 collections containing 

13 9 from China. The following collections 

from S. niphonius: 1 9 from Korea; 1 9 from China. 

The following collection from S. semifasciatus: 5 9 

1 $ from New Guinea. The following collections from 

S. tritor: 2 collections containing 4 9 from Liberia; 

8 9 from Ghana; 2 9 from Lagos, Nigeria. 


2.30 mm, greatest width 1.01 mm, length of cephalon 

0.60 mm. Genital segment wider than long (265 X 

330 juin). Abdomen 3-segmented, segments measuring 

165 X 212 ,un, 118 X 171 tan, 147 X 159 /un respectively. 

Caudal rami longer than wide (141 X 59 fun), 

longest seta 590 /an. Ventral side of last abdominal 

segment and caudal rami with patches of hairs similar 

to Unicolax collateralis. 

First antenna (Figure 206) 7-segmented; segments 

armed similarly to U. collateralis; spine on first segment 

about twice as long as adjacent plumose setae. 

Second antenna, mouthparts, and maxilliped similar 

to U. collateralis. 

Legs 1-4 biramous, each ramus 3-segmented. Leg 1 

coxopod and endopod similar to U. collateralis; basipod 

similar to U. collateralis except outer plumose 

seta more robust (see Figure 20c) ; exopod (Figure 

20c) distinctly 3-segmented, first segment with outer 

crenulated spine with terminal flagellum, second segment 

with short outer spine with terminal flagellum 

about twice as long as spine and 3 terminal to inner 

plumose setae, third segment with outer spine similar 

to that on second segment and 3 terminal plumose 

setae. Leg 2 endopod similar to U. collateralis, coxopod 

with patch of long slender spinules near outer distal 

corner and short row of minute hairs along midlower 

edge; basipod with outer dorsal seta and row of 

short spinules along outer distal edges; exopod (Figure 

20a") first segment with large patch of long, slender 

spinules on outer edge, patch of hairs on inner edge 

and outer distal spine with short hairs on edges and 

short, terminal flagellum, second segment with long 

inner plumose seta and outer spine similar to that on 

first segment, third segment with 5 inner to terminal 

plumose setae and 4 outer to terminal spines, proximal 

3 spines similar to spines of first and second segments, 

terminal spine elongate with outer serrations and short, 

inner hairs. Leg 3 (Figure 21a) coxopod with row of 

short spinules on outer distal corner; basipod with 

outer dorsal seta and short row of short spinules on 

outer distal corner; exopod first segment with outer 

distal spine with serrate rather than haired edges as 

in leg 2, and terminal flagellum, inner edge of segment 

haired, dorsolateral surface of segment with several 

rows of short spinules giving bumpy appearance,


second segment with inner plumose seta and outer 

spine similar to that on first segment, third segment 

with 5 inner to terminal plumose setae and 3 outer to 

terminal spines, proximal 2 spines similar to those of 

first and second segments, distal spine elongate, similar 

to corresponding spine of leg 2; endopod first segment 

with inner plumose seta and short row of fine 

hairs near outer distal corner, second segment armed 

similarly to first but segment slightly more elongate, 

third segment with 2 inner to terminal plumose setae 

and 2 outer to terminal spines, each with serrate edges 

and terminal flagellae, inner spine about twice as long 

as outer spine, outer edges of endopod segments haired. 

Leg 4 (Figure 2\b) coxopod with short row of small 

spinules on outer distal corner; basipod with outer 

dorsal seta; exopod first segment with spine on outer 

distal corner and hairs on inner edge, second segment 

with inner plumose seta and outer spine, third segment 

with 5 inner to terminal plumose setae and 3 outer to 

terminal spines, exopod spines slightly more elongate 

than corresponding spines of leg 3, but similarly armed; 

endopod first segment with short row of short hairs 

on outer half of distal edge and inner seta constricted 

near mid-point, plumose above constriction, finely 

serrate below it, second segment similar to first, third 

segment with fine hairs along outer half of distal edge 

and 2 spines with serrate edges and terminal flagella 

flanking a terminal seta finely serrate on lower half, 

outer edges of endopod segments haired. Leg 5 (Figure 

21c) basal segment with dorsal seta and row of minute 

spinules on outer half of distal edge; free segment 

with 2 outer patches of spinules, one near mid-segment, 

one distal, both patches with small, widely-spaced 

spinules, large dense patch of more elongate spinules 

from mid-inner edge to distal edge of segment, one 

spine on mid-outer margin and 2 spines flanking 

terminal, naked seta, spines with finely serrate edges 

and terminal flagellum. Leg 6 represented by 2 long 

and one shorter seta on genital segment. 

MALE.—Body form as in Figure 21 d. Total length 


0.29 mm. Genital segment longer than wide (265 X 

177 /an). Abdomen 2-segmented, segments measuring 

(length X width) 88 X 88 /*m, and 88 X 70 /*m, 

ventral surface of second segment (see Figure 21«) 

ornamented with 4 rows of slender, elongate spinules, 

2 proximal rows that meet at mid-point of segment 

and 2 separate distal rows. Caudal rami (Figure 21«?) 

longer than wide (59 X 29 jan) each ram us with 6 

setae and ventral patch of hairs; longest seta 413 /*m. 

First antenna similar to that of U. collateralis male 

except distal seta on third segment heavily plumose 

and extends beyond aesthete of terminal segment. 

Remaining cephalic appendages similar to those of 

U. collateralis male. 

Legs 1-4 biramous. Leg 1 similar to that of U. collateralis 

male except endopod third segment with 

outer spine haired on outer edge and with terminal 

flagellum (see Figure 21/). Legs 2-5 (Figures 22a—d 

respectively) similar to those of U. collateralis male 

with following exceptions: leg 3 endopod third segment 

outer 2 spines each with lateral serrations as 

broad as those of exopod spines; leg 4 endopod first 

segment inner seta extends beyond spines of second 

segment almost as far as terminal seta, spines of second 

segment of about equal length, serrations of outer 

spine broader than those of inner spine; leg 5 basal 

segment with single row of small spinules on outer 

distal edge, free segment with single row of distal 

spinules near bases of terminal spine and seta, spinules 

on inner lateral margin of segment limited to a few 

irregular rows. 

ETYMOLOGY.—The Latin ciliatus ("hairy") alludes 

to the hairs rather than serrations on the spines of the 

exopod of leg 2. 


from all others of the genus by the nature of 

the spines on the exopod third segment of leg 2. The 

first 3 spines are finely haired on both sides, the distal, 

elongate spine has conspicuous, broad serrations on 

the outer edge. Unicolax anonymous, U. collateralis, 

and U. reductus have broad, lateral serrations on the 

last 3 spines of the corresponding segment. Unicolax 

mycterobius has fine hairs rather than serrations on 

the distal elongate spine. 

Unicolax ciliatus is the only species in which the 

ornamentation of the exopod spines of leg 2 differs 

from that of legs 3 and 4. The modified spine of the 

first antenna is comparatively longer and more slender 

in U. ciliatus than in other species. 

Males of this species are characterized by having, 

on the ventral surface of the last abdominal segment, 

a row of spinules near the insertion of each caudal 

ramus and a proximal row of spinules in each right 

and left half; the 2 proximal rows meet along the 

mid line of the segment, the 2 distal rows are interrupted 

medially (see Figure 21

NUMBER 3 1 1 17 

This species is found from the western Pacific to 

the Gulf of Guinea and is, so far, the only species of 

Unicolax found on species of Scomberomorus. 

Vnicolax reductus, new species 

FIGURES 23-26, 99, \\9d-f, 12O-122a,6 


172259), allotype $ (USNM 172260), paratypes 

15 5 (USNM 172261) from Katsuwonus pelamis 

(USNM 176974) nasal sinus from New South Wales. 

Additional material from the same host: 24 9 2$ 

from Tahiti and 25 9 1 & from Japan. All copepods 

collected from the nasal sinuses of the hosts. 



0.94 mm. Genital segment (Figure 236) wider than 

long (289 X 501 /*m). Abdomen (see Figure 236) 

3-segmented, segments measuring 206 X 247 /*m, 

188 X 230 Aim, 230 X 200 /un respectively; segments 

without ornamentation. Caudal rami (Figure 23c) 

longer than wide (206 X 88 /an) each ramus bearing 

6 setae and no other ornamentation, longest seta 

401 fim. 

First antenna (Figure 23d) 6-segmented. Ornamentation 

of segments as follows. Segment 1: 3 plumose 

setae; 1 seta modified to form a sclerotized spine; 

1 plumose seta. Segment 2: 5 antero-dorsal naked 

setae; 10 long plumose setae in a diagonal line across 

ventral surface of segment; 4 short, ventral plumose 

setae; 1 ventral naked seta. Segment 3: one anteroventral 

naked seta; one postero-ventral sparcely plumose 

seta. Segment 4: one ventral naked seta; one 

dorsal naked seta. Segment 5: 2 terminal ventral naked 

setae; one ventral aesthete. Segment 6: 6 terminal 

naked setae; one sub terminal naked seta; one terminal 

aesthete. Second antenna (Figure 23*) similar to 

U. collaterals and ciliatus. Labrum similar to U. collateralis. 

Remaining mouthparts as in Figure 23/. 

Maxilliped (Figure 24a) basal segment with one plumose 

seta; second segment with 3 plumose setae; hook 

bent, with elbowlike shape, and without accessory 

process. 

Legs 1-4 biramous, each ramus 3-segmented except 

leg 1 exopod and leg 4 endopod. Leg 1 (Figure 246) 

coxopod with broad inner seta, patch of small hairs 

and a blunt process near inner distal edge; basipod with 

outer patch of small hairs; exopod first segment with 

outer spine with fine serrations and terminal flagellum, 

second segment incompletely divided, with one outer 

flagellated spine and 6 terminal to inner plumose 

setae; endopod first segment with inner plumose seta 

and small hairs on distal and outer portion of segment, 

second segment with inner plumose seta, third segment 

with 5 terminal plumose setae. Leg 2 (Figure 

24c) coxopod with few, widely spaced hairs along 

distal edge; basipod with outer dorsal seta and patch 

of fine spinules on distal edge near insertion of endopod; 

exopod first segment with patch of fine spinules 

on lower, outer portion of segment, row of several, 

stout spinules near base of spine on outer distal corner, 

spine with few, broad serrations and fine terminal 

flagellum, second segment with short, inner, naked 

seta, row of stout spinules near base of outer spine 

similar to that on first segment, third segment with 

4 inner to terminal, sparcely plumose setae, and 3 

outer to terminal spines, proximal 2 spines each with 

row of stout spinules near base and similar to those of 

first and second segments, terminal spine elongate 

with few, broad outer serrations; endopod first segment 

with stout, inner, plumose seta and patch of 

fine hairs on lower, outer edge of segment, second 

segment enlarged, with 2 inner, plumose setae and 

patch of fine hairs on outer, distal corner, third segment 

with 3 inner to terminal plumose setae and 2 

small, outer to terminal spines, each with finely serrate 

edges, outer edges of endopod segments haired. 

Leg 3 (Figure 24d) coxopod without ornamentation; 

basipod with outer, dorsal seta and small patch of 

fine spinules near insertion of endopod; exopod similar 

to leg 2 except spinules on lower, outer portion of first 

segment slightly larger and more sparcely placed than 

corresponding spinules of leg 2; endopod first segment 

with inner, sparcely plumose seta and patch of fine 

spinules on outer half of distal edge, second segment 

similar to first, third segment with 2 inner to terminal 

sparcely plumose setae, 2 outer to terminal spines each 

with broadly serrate edges and terminal flagellum 

(terminal spine longer than outer spine) and small 

patch of spinules near base of spines, outer edges of 

endopod segments with short hairs. Leg 4 (Figure 

25a) coxopod and basipod similar to leg 3; exopod 

similar to leg 3 with following exceptions: first segment 

with very few spinules on outer portion of segment; 

third segment with only 3 rather than 4 

sparcely plumose setae; also spines slightly longer than 

corresponding spines of leg 3; endopod first segment

18 

with inner seta finely serrate distally and patch of 

spinules along outer distal edge, few spinules on outer 

edge, second segment incompletely divided with triangular 

patch of spinules proximal to division indicating 

probable fusion of second and third segments, 

uneven row of stout spinules distally near bases of 

terminal seta flanked by 2 spines, seta finely serrate 

distally, outer spine similar to those on exopod, inner 

spine with fine serrations, outer edge of segment with 

two patches of short hairs. Leg 5 (Figure 25b) basal 

segment with naked, dorsal seta, patch of spinules on 

outer, distal corner; free segment with patch of spinules 

on inner edge and a row of stout spinules near 

base of each of the following: one, terminal, naked 

seta; 3 spines, one on mid-outer edge, one on each 

side of terminal seta, all spines stout with broad serrations 

distally and fine, terminal flagellurn. Leg 6 (see 

Figure 23b) represented by 3 short setae, of about 

equal length, on genital segment. 



0.39 mm. Genital segment (Figure 25d) slightly 

wider than long (277 X 300 /mi), abdomen 2-segmented, 

segments measuring (length X width) 159 

X 177 /an, and 194 X 147 /on; second segment with 

two ventral rows of minute spinules (see Figure 25^). 

Caudal rami (Figure 25«) longer than wide (141 X 

73 fixn) with no ornamentation other than 6 setae 

(longest 371 pin). 

First antenna similar to that of U. collateralis male. 

Remaining cephalic appendages (except rnaxilliped) 

similar to those of female. Maxilliped (Figure 26a) 

second segment inflated with 2 naked setae and numerous 

irregular rows of low, rounded spinules; terminal 

segment claw-like with 2 naked setae and row of 

teethlike spinules along inner edge to apex. 

Legs 1-4 biramous. Leg 1 (Figure 266) basipod 

with stout outer seta, patch of hairs near insertion of 

rami, inner spine sclerotized at base with terminal 

fringed membrane, patch of spinules near base of 

spine extending midway up segment; exopod 3-segmented, 

first segment with outer spine with stout 

terminal flagellum, second segment with inner seta 

and outer spine similar to that on first segment, third 

segment with 5 setae and 2 outer spines, proximal 

spine similar to previous 2, distal spine simple, not 

heavily sclerotized; endopod 3-segmented, similar to 

U. mycterobius male except patches of hairs on first 

and second segments smaller. Leg 2 (Figure 26c) 


similar to female except exopod setae comparatively 

longer; endopod setae longer than and not as robust 

as in female and second segment not as inflated. Legs 

3 and 4 similar to female. Leg 5 (Figure 26d) basal 

segment with dorsal naked seta; free segment ornamented 

similarly to female, but segment itself not as 

robust. 

ETYMOLOGY.—The Latin reductus ("reduced") 

alludes to the reduced number of setae on exopods 

of legs 2-4 and the reduced number of segments of 

the endopod of leg 4. 

REMARKS.—Females of U. reductus can be distinguished 

from other Unicolax by the following characters: 

the ventral surface of the last abdominal segment 

has' no ornamentation; the caudal rami have 

only six setae each, with no ventral hairs or spinules 

(other species have a ventral patch of hairs on each 

ramus); leg 2 exopod third segment has 3 spines, 

4 setae (other species have 4 spines, 5 setae); leg 3 

exopod third segment has 4 setae (other species have 

5) ; leg 4 exopod third segment has 3 setae (U. collateralis, 

U. mycterobius, and U. ciliatus have 5, U. 

anonymous has 4) ; leg 4 endopod is 2-segmented (in 

all other species it is 3-segmented). 

Males of U. reductus differ from others of the genus 

by the following characters: the ventral surface of 

the last abdominal segment is ornamented by only 2 

distal rows of minute spinules; the caudal rami have 

no ornamentation on the ventral surface (other males 

have a patch of hairs on the ventral surface of each 

ramus). Legs 3, 4, and 5 are strikingly similar to those 

of the female, while the legs of males of others in 

the genus differ in several ways from their female 

conspecifics. 

Unicolax reductus is the only member of the genus 

found on Katsuwonus pelamis and is probably restricted 

to that host species. 

Because of the number and nature of its unique 

characteristics, it is the most easily identified member 

of the genus. 

Ceratacolax Vervoort, 1965 

Ceratacolax enthynni Vervoort, 1965 

FIGURES 27-31, 100, 122c-/, 123-125 

Ceratacolax euthynni Vervoort, 1965:26. 

MATERIAL EXAMINED.—34 collections containing 

94 9 and 63 S from the following hosts and localities:

NUMBER 3 1 1 19 

Euthynnus alletteratus from Massachusetts, New 

Jersey, Florida (west coast), and Ghana; Sarda sarda 

from Massachusetts, Rhode Island, Chesapeake Bay, 

Florida (east and west coasts), Mississippi, Texas, 

Venezuela, Brazil, Spain (Cadiz), Gulf of Guinea, 

and South Africa (Port Elizabeth). All collections 

from the nasal sinuses of the hosts. 

Vervoort provided a good discription of both the 

male and female of this species. We have figured both 

sexes completely and will comment only on those 

characteristics not included in, or which differ from, 

Vervoort's description. 

FEMALE.—Ventral surface of last abdominal segment 

and caudal rami (Figure 27d) with patches of 

prominent spinules (omitted by Vervoort). Leg 1 

(Figure 286) exopod 2-segmented (Vervoort indicated 

1), second segment incompletely divided; first segment 

with one crenate spine on outer distal corner, 

second segment with 3 outer spines (omitted by Vervoort) 

and 6 inner to terminal setae. 

MALE.—Ventral surface of last abdominal segment 

and caudal rami with patches of spinules as indicated 

in Figures 27c and d (Vervoort omitted spinules on 

caudal rami). Legs 1-3 (Figures 30c, 30rf, 31a) endopod 

third segment, second from innermost seta with 

row of 4-5 long, slender spinules along proximal outer 

edge (spinules omitted by Vervoort). 

REMARKS.—This bomolochid can be easily separated 

from other known members of the family on the 

basis of the following characteristics. The structure of 

the first antenna is unique by the presence of a long, 

curved, heavily sclerotized dorsal hook situated at the 

junction of the first and second segments; this hook 

is in addition to the usual row of 15 plumose setae on 

the basal segments and apparently does not represent 

a modified seta or setae as in some other bomolochid 

genera. The genital segment includes 3 flaplike structures 

at the area of egg sac attachment; within these 

structures are 3 spinulose setae (not easily visible without 

dissection) representing leg 6. Leg 2 exopod and 

both rami of legs 3 and 4 are characterized by patches 

of stout spinules along outer edges, and by setae that 

are spinulose rather than plumose. Leg 4 endopod is 

conspicuously elongate, especially the terminal segment. 

Males are characterized by a first leg that is neither 

flattened nor modified as in the female. Legs 2 and 3 

endopod second segment both have 2 inner setae. 

The setae of the male legs are all plumose in contrast 

to the spinulose setae of the female. Perhaps the most 

unique feature of the male is the presence of long 

slender spinules on the base of one seta on each 

endopod last segment of legs 1-3. 

In both sexes there is some individual variation in 

the pattern of spinules of patches on the last abdominal 

segment and caudal rami. In the female, particularly, 

spinules on the caudal rami may appear as a 

single longitudinal row or as a patch of 2-3 rows. 

We could not correlate this variation to host or geographic 

distribution. The variation seems to occur 

randomly with occasional variation on the same individual 

from right to left ramus. 

Nothobomolochus Vervoort, 1962 

Nothobomolochus kanagurta (Pillai, 1965), 


FIGURES 32, 33, 100 

Bomolochus kanagurta Pillai, 1965:51. 

MATERIAL EXAMINED.—5 collections containing 9 9 

from the gills of Rastrelliger kanagurta from China, 

India (Madras), Red Sea; R. faughni from the 

Philippines. 


1.50 mm. Abdomen 3-segmented, segments measure 

118 X 206 /an, 95 X 195 /an, 95 X 177 pin, length 

by width respectively. Last abdominal segment (Figure 

32b) with 2 oblique patches of spinules. Caudal 

rami (see Figure 326) about twice as long as wide 

(94 X 55 jan); each ramus with a lateral seta, 2 subterminal 

and 2 terminal setae, one of which is much 

larger, its base nearly as wide as distal end of ramus. 

First antenna (Figure 32c) with 3 modified setae on 

basal segment, outer 2 longer than more heavily 

sclerotized middle seta; remaining setae of usual 

bomolochid type; an aesthete on each of last 2 segments. 

Second antenna with rows of hooklets on second 

segment and armed with terminal setae typical 

of the genus. Oral appendages typical bomolochid. 

Maxilliped (Figure 32d) with recurved hook bearing 

a short, blunt-tipped accessory process, base armed 

with 3 prominent plumose setae. 

Legs 1-4 biramous, each ramus 3-segmented. Leg 1 

endopod with broad segments as in other bomolochids. 

Leg 2 (Figure 33a) coxopod with a group of long 

spinules on outer distal corner and a row of short

20 

spinules at articulation with basipod; basipod with a 

cluster of long spinules arranged in a circle near inner 

margin and a long, stout seta at outer distal corner; 

exopod first segment with a patch of long spinules 

along outer margin and a heavy serrated spine at 

outer distal corner, second segment with serrated spine 

at outer distal corner and an inner seta, third segment 

with 3 serrated spines, a fringed terminal spine, 

and 5 terminal to inner setae (last 5 spines armed on 

1 margin only) ; endopod first 2 segments each with 

a row of spinules on distal margin in addition to usual 

inner setae, last segment with 2 short, outer, lightly 

fringed spines and 3 terminal setae. Leg 3 (Figure 

336) coxopod with a row of stout spinules at outer 

distal corner; basipod with 3 patches of spinules (outer 

2 fine, inner patch heavier) and a seta on outer distal 

corner; exopod similar to leg 2 except no spinules on 

first segment and 1 less spine on last segment; endopod 

as in leg 2 except 1 less seta on last segment. Leg 4 

(Figure 33c) as in leg 3 except 1 less patch of fine 

spinules on basipod; 1 less spine and seta on exopod 

last segment; endopod patches of spinules larger, 

setae with short spinules rather than usual plumosities, 

last segment with only 1 long seta. Leg 5 (Figure 33d) 

with 2 prominent patches of spinules in distal third, 

innermost with longer spinules; 1 lateral, naked seta 

and 3 terminal setae, outer 2 finely spinulose (all setae 

of about equal length). 

MALE.—We did not collect males of this species 

but Pillai noted (1965:53) that the second segment 

of the maxilliped bears 2 rows of "tubercles" on inner 

margin. 

REMARKS.—This species has been collected only 

from species of Rastrelliger from the Indo-West 

Pacific. 

Orbitacolax Shen, 1957 

Orbitacolax aculeatus (Pillai, 1962), 


FIGURES 34, 35a-b, 100 

Bomolochus aculeatus Pillai, 1962a: 610. 


15 9 from the nasal sinuses of 2 Rastrelliger faughni 

from Manila and Lingayan Gulf, Philippines. 


1.71 mm. Greatest width 0.73 mm. Rostrum protrud- 


ing from between bases of first antennae and bearing 

a pair of ventral hooks (see Figure 346). 

Abdomen 3-segmented, ventral surface covered with 

spinules. Caudal rami about twice as long as wide, 

each with one lateral, 2 subterminal, and 3 terminal 

setae (innermost much stouter and longer than 

others). First antenna (Figure 346) bearing 15 

plumose and 6 naked setae along outer edge and 1 

plumose and 3 naked setae directed posteriorly on 

first 3 segments, fourth and fifth segments with 3 setae 

each, last segment with 7 short and 1 long terminal 

setae. Second antenna (Figure 34c) with 6 rows of 

small hooks along outer edge of second segment and 

6 setae at tip (Pillai indicates 5). Labrum with 2 

dense patches of spatulate spinules, posterior corners 

produced as spinulose knobs. Other oral appendages 

as in Figure 34d. Maxilliped (Figure 34e) hook with 

accessory process, posterior edge of basal segment with 

spinules (not indicated by Pillai). 

We will not repeat a full description of legs 1-5 but 

rather restrict our discussion to those points where our 

material differs with the description of Pillai's. Pillai 

illustrated the basipod of leg 1 with 1 large patch of 

spinules; in our specimens there are 2 (see Figure 34/). 

Leg 2 as described by Pillai except endopod second 

segment with 2 setae rather than one. Leg 3 exopod 

similar to that of leg 2; endopod first and second segments 

each with one inner seta, third segment with 2 

naked subequal outer spines and 2 inner setae. Pillai 

shows no setae on first and second segments and only 

one outer spine on third segment. Legs 4 and 5 (Figure 

35a,6) as described by Pillai. 

REMARKS.—In spite of the differences between our 

material and the description given by Pillai we feel 

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 Rastrelliger 

from India and the Philippines. 

Pumiliopes Shen, 1957 

Pumiliopes capitulatus Cressey and Boyle, 1973 

FIGURES 35c-e, 36, 37, 100, 126a-c 

Pumiliopes capitulatus Cressey and Boyle, 1973:1. 


35 5 from the orbits of the following hosts and localities: 

Rastrelliger kanagurta the Red Sea, Sri Lanka,

NUMBER 311 21 

India (Madras), western Indian Ocean, Philippines, 

Java; R. faughni from Philippines; Scomber japonicus 

from Gulf of Guinea, Mauritania, Zanzibar; S. australasicus 

from Philippines. 


of holotype 1.53 mm, greatest width 0.75 mm. Cephalon 

about as long as wide. Genital segment (Figure 

35d) wider than long. Abdomen (see Figure 35


as in Figure 38c. Maxilliped absent. 

Leg 1 (Figure 38rf) biramose; exopod 2-segmented, 

first segment with a short toothed spine on outer distal 

corner, second segment with 4 toothed spines and 3 

weak setae along inner margin; endopod 3-segmented, 

first segment unarmed, a non-articulated spine on 

outer distal corner of second segment, last segment 

with 2 non-articulated terminal spines and a small 

seta on inner margin. Leg 2 (Figure 39a) as in leg 1 

except one less spine and seta on exopod last segment 

and an additional seta on endopod last segment. Leg 3 

(Figure 39b) 1-segmented, bearing a spine on inner 

distal corner and 2 short terminal setae. Legs 4, 5, and 

6 absent. 

Egg sacs typically cyclopoid. 

MALE.—Body form as in Figure 38a. All males 

collected were attached to females. Body segmentation 

well defined. Abdomen 4-segmented. Caudal rami as 

in female. 

First antenna 8-segmented each with short setae. 

Second antenna (Figure 39c) 3-segmented; first segment 

with 2 short setae on inner margin, second segment 

with a seta on outer distal comer and 3 setae on 

a sclerotized ridge at inner distal corner, third segment 

in form of a bifurcate claw with an accessory spine 

near base. Legs 1-3 similar to female except last 

exopod segment of leg 2 with a long, heavily sclerotized 

spine (Figure 39d). 

REMARKS.—This species so far has been collected 

only from species of Scomberomorus from the western 

Atlantic. For a more complete description the reader 

should consult the original description (Cressey, 1975). 

Shiinoa occlusa Rabat a, 1968 

FIGURES 39e-h, 101, \2U-f, 127-128a,fc 

Shiinoa occlusa Kabata, 1968a:497. 


41 9 11 $ from the nasal lamellae of the following 

hosts and localities: Grammatorcynus bicarinatus from 

North Celebes, Solomon Islands, Palau, Caroline Islands; 

Gymnosarda unicolor from Solomon Islands; 

Scomberomorus commerson from Mozambique, Pakistan, 

Gulf of Thailand, Solomon Islands, Philippines, 

Palau; S. guttatus from China; S. niphonius from 

Japan; S. queenslandicus from Papua; S. tritor from 

Canary Islands; Acanthocybium solandri from Kapingarmarangi 

Atoll. 

When Kabata described this species he stated that 

his description was based on an immature female. The 

first author examined that specimen during the course 

of another study and compared some of the present 

material with it. It was concluded that Kabata's specimen 

was a nonovigerous female adult. We have found 

no differences between the specimens reported here 

and Kabata's description except for the presence of 

egg sacs on some of our specimens. We will point out 

the differences between this species and S. inauris 

rather than repeat a full description here. We have, 

however, provided several SEM photographs of a 

female S. occlusa (from Gymnosarda unicolor). Females 

of S. occlusa can be distinguished from S. inauris 

by the following: the abdomen of S. occlusa is 

about one-sixth of the total body length (one-third in 

S. inauris) ; the exopods of legs 1 and 2 of S. occlusa 

are 3-segmented (2-segmented in S. inauris). Males 

of the 2 species can be separated by the following 

characters: distal segment of the second antenna of 

S. occlusa with 3 clawlike terminal spines (a bifid 

claw in S. inauris); 3-segmented exopods of legs 1 

and 2 of S. occlusa (2-segmented in S. inauris); 2 

stout, but unequal, spines at tip of leg 2 exopod of 

S. occlusa (1 stout spine in S. inauris). 

REMARKS.—This species is very similar to S. inauris 

but easily separated by the characters cited above. The 

species so far has been collected from the Western 

Pacific, Indian Ocean, and Eastern Atlantic. It should 

be pointed out that a third species (S. elagata Cressey, 

1976) has been described from the carangid Elagatus 

bipinnulata (Quoy and Gaimard) from the Western 

Pacific. 

Caligus Muller, 1785 

Caligus coryphaenae Steenstrup and Liitken, 1861 

FIGURES 40, 41a-6, 102; 128c-/, 129, 130 

Caligus coryphaenae Steenstrup and Liitken, 1861:352. 

Caligus scutatus Milne-Edwards, 1840:453. 

Caligus thymni Dana 1852:56. 

Caligus bengoensis Scott, 1894:130. 

Caligus aliuncus Wilson, 1905:576. 

Caligus elongatus Heegaard, 1943:11. 

Caligus tesserifer Shiino, 1952:89. 


316 9 and 281 $ from the body surface and gills of 

the following hosts and localities: Acanthocybium

NUMBER 3 1 1 23 

solandri from northeast coast of Malagasy Republic, 

Auxis species from Gulf of Guinea; Euthynnus alletteratus 

from Puerto Rico (Atlantic) ; Katsuwonus 

pelamis from northwest Malagasy Republic, Christmas 

Island (Pacific), Peru, Ecuador, east coast of United 

States (several localities), Puerto Rico (Atlantic), 

Venezuela, Brazil (north coast), Cape Verde Islands, 

Gulf of Guinea; Thunnus alalunga from New Jersey; 

Thunnus albacares from Christmas Island (Pacific), 

east coast of United States (several localities), Brazil, 

Gulf of Guinea; Thunnus atlanticus from Puerto Rico 

(Atlantic); Thunnus obesus from Christmas Island 

(Pacific), east coast of United States (several localities), 

Surinam, Brazil (north coast), Cape Verde Islands; 

Thunnus thynnus from east coast of United 

States (several localities). 

Shiino (1959a), Pillai (1962b), and Lewis (1967) 

have provided good descriptions and figures for this 

well known species. We will restrict our consideration 

of this species to those characters that serve to distinguish 

it from the other Caligus species found on 

scombrid hosts. 

FEMALE.—Body form as in Figure 40a. Lewis, et al. 

(1969:423) provided morphometric data for 36 

specimens from 4 host species from the Indian Ocean. 

The average total length for his material was 5.57 

mm. Our material from the Indian Ocean and Pacific 

areas did not differ significantly from this mean. The 

Atlantic specimens, however, tended to be larger (5.8- 

6.5 mm based on 10 specimens from various localities). 

The cephalon comprises about one-half of the total 

length with the genital segment and abdomen each 

comprising about one-fourth. 

Frontal lunules widely spaced; space between lunules 

(1.31 mm) more than twice greatest diameter of 

either lunule (0.58 mm). 

Genital segment slightly longer than wide (1.5 X 

1.4 mm) with posterior outer corners produced somewhat 

beyond origin of abdomen. Abdomen 3-segmented; 

1st segment constricted in posterior two-thirds 

giving the appearance of 2 segments, second segment 

shortest, third segment with 2 distal lobes separating 

caudal rami; segments measure 0.9, 0.3, and 0.37 mm 

respectively. Caudal ramus as in Figure 406. 

Postantennal spine lacking. Postoral spine wide at 

base, triangular. Sternal furca (Figure 40c) with 

short, widely divergent tines; furca with an accessory 

sclerotized cuticular process on each side. 

Leg 1 (Figure 40rf) basipod with a patch of short 

spinules, patches of longer hairs, a short plumose seta 

on inner margin, a long, very plumose, seta at outer 

distal corner; exopod 3 distal spines each with prominent 

fringes, inner 2 with accessory process; endopod 

reduced to a short, sclerotized process. Leg 2 endopod 

(Figure 40e) with patches of long spinules on outer 

margin of each segment. Leg 3 exopod (Figure 40/) 

with prominent thumblike spine on outer distal corner 

of first segment, each of last 2 segments with long 

setules on outer margin in addition to usual spines and 

setae. Leg 4 exopod (Figure 41a) 3-segmented, first 

and second segment each with a prominent fringed 

spine on outer distal corner; last segment with 3 

fringed spines, distalmost longest, all spines with a 

fringe at base; spines measure 366, 236, 153, 206, and 

247 /un distalmost to proximalmost respectively. 

MALE.—Body form as in Figure 416. Total length 

5.3 mm. Cephalon comprises more than half total 

length. Appendages as in female except second antenna 

with accessory process on claw. Maxilliped with 

small sclerotized area opposite tip of claw. Sternal 

furca similar to female except tines not quite as divergent. 

Thoracic appendages as in female. Legs 5 and 6 

represented by setae at the posterior corners of the 


REMARKS.—This species has been reported many 

times since its original description in 1861. It is common 

on the body surface of scombrids of the tribe 

Thunnini. The single collection from Acanthocybium 

solandri (Scomberomorini) is the only exception reported 

here of all scombrids examined. It is also common 

on species of Coryphaena and is found in all except 

polar oceans (see Figure 102). Margolis, Kabata, 

and Parker, 1975:25 provide a complete synonymy to 

this species. 

Further discussion of this species follows the description 

of C. regedis. 

Caligus regalis Leigh-Sharpe, 1930 

FIGURES *\c-g, 102 

Caligus regalis Leigh-Sharpe, 1930:5. 

Caligus euthynus Kurian, 1961:63. 

Caligus alveolaris Heegaard, 1962:156. 

MATERIAL EXAMINED.—5 collections containing 149 

5, 113 $ from the body surface of Euthynnus 

affinis collected off Nosy Be, Malagasy Republic. 

Types of C. alveolaris (AMS P. 16408) from the


same host species, reported as E. allitteratus [sic] 

from north Queensland, Australia. 


4.62 mm (4.27-4.73 mm) based on an average of 5 

specimens. Greatest width 2.1 mm (measured at 

widest part of cephalon). Cephalon comprises about 

one-half total body length. Genital segment somewhat 

longer than wide (1.77 X 1.55 mm), posterior corners 

produced to form rounded lobes. Abdomen 2-segmented, 

segments measure 1.02 and 0.43 mm long 

respectively, comprising about one-third total body 

length. Caudal rami as in Caligus coryphaenae except 

outer terminal seta much stouter in C. regalis. 

Frontal lunules widely spaced, each about 400 /*m 

wide, rostral space between each lunule about 950 /xrn. 

Cephalic appendages as in C. coryphaenae except 

sternal furca not as divergent in C. regalis (Figure 

Legs 1-4 as in C. coryphaenae; the stout spine on 

the first exopod segment of leg 3 (Figure 41«) is 

curved inwards rather than straight as in C. coryphaenae. 

The apical seta on the exopod of leg 4 

(Figure 41/) is only slightly longer than the adjacent 

seta, setae measure 189, 177, 177, 177, and 183 /*m 

distal to proximal respectively. 

MALE.—Body form as C. coryphaenae. Total length 


specimens. Greatest width 1.73 mm (measured at 

widest part of cephalon). Genital segment (Figure 

41g) wider than long (914 X 711 /un). Caudal rami 

with outermost terminal seta stout and about as wide 

at base as other 3 (much smaller in C. coryphaenae). 

Legs 5 and 6 at posterior outer corners. Abdomen 2segmented, 

segments measure 247 and 479 /tin respectively. 

REMARKS.—Pillai, 1962b, discussed the relationship 

between this species and C. coryphaenae and provided 

a comprehensive description of C. regalis (as C. 

euthynus). All collection records to date indicate that 

C. regalis is specific to Euthynnus affinis and C. 

coryphaenae is primarily found on other species of 

scombrids of the tribe Thunnini. 

These 2 species of Caligus differ from other members 

of the genus in lacking a postantennal process 

(Lewis 1967:105, considered the group of sensillae 

at the usual site of the postantennal process as representing 

the process). When the genus Caligus has 

undergone a much needed revision it may be that 

these 2 species should be removed from the genus. 

We have placed Heegaard's C. alveolaris and 

Kurian's C. euthynus in synonymy with C. regalis of 

Leigh-Sharpe. Although Leigh-Sharpe's figures are 

poor there is little doubt that they illustrate the copepods 

discussed here, especially considering that he 

collected his material from the same host-species (as 

E. yaito). 

Caligus productus Dana, 1852 

FIGURES 42-44, 103 

Caligus productus Dana, 1852:56. 

Caligus alalongae Kroyer, 1863:129. 

Caligus monacanthi Kroyer, 1863:133. 

Caligus lobatus Wilson, 1935:1. 

Caligus katuwo Yamaguti, 1936:6. 

Caligus dentatus Heegaard, 1962:160. 

Caligus microdontus Heegaard, 1964:139. 

This species has been recorded many times from 

scombrids and occasionally non-scombrid hosts (see 

Lewis, 1967:116). Shiino, 1959b and Lewis, 1967 

have provided good descriptions and we do not feel 

a need to repeat another here. We have, however, 

provided pertinent illustrations and a consideration 

of certain characters to enable the reader to identify 

this species without resorting to other literature. A 

complete synopsis of the literature has been provided 

by Margolis, Kabata, and Parker (1975:64). 


2286 9 , 657

NUMBER 3 1 1 25 

Republic, Mozambique Channel, Chagos Islands, 

Somalia, Palau, Line Islands, Brazil (north coast), 

Dominican Republic (Atlantic), Bermuda, east coast 

United States (several localities) ; Thunnus atlanticus 

from West Indies (several localities), Nicaragua; 

Thunnus thynnus from east coast United States 

(several localities), Gulf of Mexico, Japan. 

FEMALE.—Body form as in Figure 42a. Range of 

total length (including data of Lewis 1967, 1968) 

3.85-5.45 mm. We did not find any significant differences 

in total length between our measurements of 

Atlantic specimens and those of ours and Lewis' from 

the Indian and Pacific oceans. Examination of specimens 

from several different hosts from widely separated 

geographic areas revealed no significant variations 

in form. It appears that this is a very stable 

species, well established as a parasite of scombrids 

except members of the tribe Scombrini {Scomber and 

Rastrelliger). 

Caligus productus females can be easily separated 

from other species of the genus by a combination of 

the following characters: the outer distal corners of 

the genital segment are rounded and extend well 

beyond the insertion of the abdomen; the ventral 

surface of the abdomen bears a medial subterminal 

patch of minute spinules and a terminal transverse 

patch of fine hairs (see Figure 426); the last exopod 

segment of leg 1 lacks the usual 3 lateral setae; the 

outer distal corner of the first endopod segment of 

leg 2 bears a row of long spinules, the second segment 

bears a double row of stout spines (7-9 per row) 

along its outer edge; leg 4 exopod is 2-segmented, the 

last segment bearing one lateral and 3 terminal setae 

(outermost seta slightly longer than others). 

MALE.—Cephalothoracic appendages as in female 

except second antenna (Figure 44«) with short claw 

and 2 prominent bossed areas on basal segment. Genital 

segment and abdomen as in Figure 440*. 

REMARKS.—This species is common on scombrid 

fishes throughout the circumtropical and subtropical 

area. We have recorded it here from 15 species of 

hosts, infesting the buccal area, gills and body surface. 

Data presented by Lewis, et al, 1969, and our own 

collection data indicate the parasite is most commonly 

found in the buccal area, next in the gill area, and 

occasionally on the body surface. The fewer records 

from the body surface could result from the more 

exposed site and consequently the copepods more 

easily drop off prior to examination of the host. Cer- 

tainly our own records, based in large part on the 

examination of preserved hosts, would be biased in 

that regard. 

Although we collected Caligus productus from 14 

species of scombrid fishes it is common (found on 

more than 25% of the specimens examined) on only 

4 species; Katsuwonus petamis 52%, Thunnus albacares 

45%, T. atlanticus 92%, and T. thynnus 28%. 

It is curious that, although this parasite was found 

on 8 of the 13 species of Thunnini, we did not 

collect it from any of the 40 specimens of T. obesus 

examined. 

This species was not found on any species of the 

tribe Scombrini (Rastrelliger and Scomber). Leigh- 

Sharpe (1926:384) reported this species from Scomber 

scombrus and placed the previously described Caligus 

scomberi Bassett-Smith in synonymy with it. Later 

writers (see Margolis, Kabata, and Parker 1975:74) 

and our examination of the type of Caligus scomberi 

agree that it is a synonym of Caligus pelamydis 

Kroyer, 1863 and the records of C. productus from 

Scomber scombrus discounted. 

Caligus asymmetricus Kabata, 1965 

FIGURES 45-47, 104 

Caligus asymmetricus Kabata, 1965:109. 

Caligus thynni Pillai, 1963:89. 

In addition to Pillai's original description, Lewis 

(1967:131) redescribed this species and presented an 

account of its rather confusing history. In view of 

these recent works we will not present another full 

description of C. asymmetricus, but, as with some 

other Caligus species from scombrids, discuss only the 

salient features and provide full illustrations to enable 

the reader to identify this species without consulting 

the earlier works. 

MATERIALS EXAMINED.—22 collections containing 

719 and 25 $ from the gills, gill arches, and roof of 

the mouth (1 collection) of the following hosts and 

localities: Grammatorcynus bicarinatus from Palau, 

Marshall Islands, Kapingamarangi Atoll; Scomberomorus 

commerson from New South Wales, East 

Indies; Sarda australis from New South Wales; Sarda 

orientalis from Seychelles, Hawaii; Cybiosarda elegans 

from western Australia; Auxis sp. from New South 

Wales; Euthynnus affinis from New South Wales, 

northwest Malagasy Republic; Katsuwonus pelamis

26 

from northwest Malagasy Republic; Thunnus albacares 

from Queensland. 


total length 3.00-4.20 mm (3.49 mm average of 22 

specimens). Those from Cybiosarda elegans (western 

Australia) averaged somewhat longer (3.78 mm of 9 

specimens) than those from other hosts from other 

areas of the Pacific and Indian oceans. Those measured 

from other areas and other hosts did not vary 

significantly. 

Caligus asymmetricus can be separated from other 

Caligus species found on scombrids by the combination 

of the following characters: abdomen short 

(about 10 percent of total length); sternal furca 

narrow, tines often asymmetrical or otherwise distorted; 

leg 2 endopod first segment with long spinules 

at outer distal corner, second segment with a row of 

6-8 stout spines; leg 4 exopod 2-segmented, outermost 

seta longest, other 4 exopod setae all about equal in 

length to each other. 

MALE.—Characters described above apply to males 

except abdomen comprises about 15 percent of total 

body length. 

REMARKS.—Caligus asymmetricus was reported by 

Pillai (1963:89) from Euthynnus affinis from India, 

by Kabata (1965:110) from the same host (reported 

as E. alletteratus) from Queensland, Australia, and 

by Lewis (1967:131) from the same host (reported 

as E. yaito) from Hawaii. We record this species from 

9 scombrid species within the Indo-West Pacific. Although 

the records are scattered throughout 3 of the 

4 tribes of Scombrinae, our collections indicate that 

Indo-West Pacific species of the tribe Sardini may be 

preferred hosts (17 percent infestation rate on 3 host 

species as opposed to 4 percent infestation rate on 2 

species of Scomberomorini, 2 percent infestation rate 

on 4 species of Thunnini and no records from the 

5 Indo-West Pacific species of Scombrini). 

Kabata (1965:110) notes that both his and Pillai's 

specimens were collected with specimens of Caligus 

bonito (reported as C. kuroshio). Of the 22 collections 

reported here only 2 were accompanied by C. bonito, 

nor did we find any other species common in collections 

of C. asymmetricus. 

We found that the peculiar asymmetry or distortion 

of the tines of the sternal furca reported previously in 

the collections from India, Australia, and Hawaii did 

not occur in all collections but rather seems to occur 

randomly within populations. The sternal furcae of 


our specimens from Sarda orientalis from Hawaii are 

usually distorted, including one specimen with 1 central 

tine rather than the usual 2. In certain other 

collections the furcae were normally developed. 

Caligus bonito Wilson, 1905 

FIGURES 48-50, 105 

Caligus bonito Wilson, 1905:589. 

Caligus sarda Pearse, 1952:17. 

Caligus kuroshio Shiino, 1959b: 51. 

This species has been reported many times (see 

Margolis, Kabata, and Parker, 1975:18 for complete 

literature history) and has been recently redescribed 

by Pillai, 1971:161 and Lewis, 1967:124. Because 

of these recent good descriptions it is unnecessary to 

completely redescribe this species again. We discuss 

only certain taxonomic features that we consider important 

in distinguishing this species from other 

Caligus found on scombrid fishes. We have, however, 

provided illustrations so the reader can identify specimens 

without consulting other work. 


155 9 and 58 $ from the mouth and gills of the following 

hosts and localities: Euthynnus affinis from 

New South Wales; E. alletteratus from Florida (Gulf 

coast), Dominican Republic (Atlantic); E. lineatus 

from Galapagos Islands, Panama (Pacific), Baja California 

(Pacific) ; Thunnus thynnus from Florida (Atlantic) 

; Scomberomorus regalis from Antigua; Sarda 

sarda from western North Atlantic (several localities 

Massachusetts to Brazil); Florida (north Gulf coast), 

Rio de Janeiro, Norway, Tunisia, Gulf of Guinea, 

Angola, South Africa (Port Elizabeth) ; Sarda australis 

from New South Wales; Sarda chiliensis chiliensis 

from Peru; Sarda chiliensis lineolatus from Cedros 

Islands, California; Sarda orientalis from Red Sea 

(Elat), South Africa (Port Elizabeth), India 

(Cochin), China, Galapagos; Gymnosarda unicolor 

from Bikini. 

FEMALE.—Body form as in Figure 48fl. Total length 

6.45 mm. (average of 37 specimens 5.10-8.10 mm 

range). Measurements taken from collections from 

Sarda sarda, S. australis, and S. chiliensis. No significant 

differences could be seen between C. bonito from 

different host species or geographic areas, except that 

specimens from warmer water tend to be smaller than 

those from colder.

NUMBER 3 1 1 27 

Caligus bonito females can be easily separated from 

other species of the genus except Caligus omissus new 

species by a combination of the following characters: 

cephalon, genital segment, and abdomen each comprising 

about one-third of the body length (abdomen 

widest anteriorly), endopod first and second segments 

of leg 2 with a group of prominent spinules at outer 

distal corner and a double row of stouter spinules 

along outer edge respectively (see Figures 49e,f), tines 

of sternal furca (Figure 49c) nearly parallel and blunt 

tipped, lateral setae of leg 1 exopod last segment with 

stout spinules on basal fourth (see Figure 49d), leg 

4 exopod 2-segmented with terminalmost seta about 

twice as long as others. See discussion of characters 

separating C. bonito from C. omissus new species in 

the description of the new species. 

MALE.—Body form as in Figure 50c. Total lengths 

of males are approximately 80 percent that of the 

females in any collection. The ranges would be proportionately 

the same as given for the females. Male 

as in female except in the following characters: 

abdomen 2-segmented; second antenna (Figure 50d) 

second segment with 2 rugose areas, claw with 2 short 

processes at tip and a seta on inner margin; maxilliped 

(Figure 50«) basal segment with sclerotized bifid knob 

on margin opposite tip of claw. 

DISCUSSION.—We have recorded this species here 

from 11 scombrid species. From our data Caligus 

bonito seems to be primarily a parasite of scombrids 

of the tribe Sardini (collected from 7 species) ; to a 

lesser extent a parasite of species of Thunnini (from 

4 species) ; rarely found on members of the Scomboromorini 

(1 species) ; not found on the tribe 

Scombrini (Rastrelliger and Scomber). It is recorded 

from all except polar oceans and previous 

records are cited by Lewis (1967:125) (including 

nonscombrid hosts). 

Lewis (1967:131) discussed the difficulties distinguishing 

this species from Caligus productus and 

C. quadratus Shiino. Although these 3 species superficially 

resemble each other, the nature of the armature 

on the first, second, and fourth legs should easily 

separate them. Caligus quadratus is easily separated 

from C. productus by the normally developed setae 

on the leg 1 exopod of C. quadratus (absent in C. 

productus) and from C. bonito by the stout spinules 

present on these same setae of C. bonito (normal 

plumosities on C. quadratus). 

After examining the type-specimens of Caligus sarda 

Pearse (USNM 92667) we have placed it in synonymy 

with Caligus bonito Wilson. 

Caligus mutabilis Wilson, 1905 

FIGURES 51, 52, 53a-b, 105 

Caligus mutabilis Wilson, 1905:573. 


from the gills or gill chambers of Scomberomorus 

brasiliensis from Brazil and Costa Rica; S. cavalla 

from Surinam; S. maculatus from west coast of 

Florida; Scomber japonicus from Campeche. 


2.78 mm. Greatest width 1.12 mm. Frontal lunules 

large, greatest width of lunule more than least distance 

between each lunule (307 vs 254 fan). Genital segment 

(Figure 516) longer than wide (986 X 725 /*m), 

widest posteriorly, posterior corners rounded and only 

slightly produced. Abdomen (see Figure 516) incompletely 

divided into 2 segments, about twice as long 

as wide (681 X 362 fin), posterior ventral surface 

with patches of spinules as in Figure 51c. Caudal 

rami (Figure 51c) longer than wide (159 X 106 /an) 

bearing 6 plumose setae as indicated in figure. 

Oral area as in Figure 52c. First antenna (Figure 

5ld) first segment with 27 setae (all but 2 are plumose), 

last segment with 13 naked setae. Second 

antenna (Figure 51/) with spatulate posterior process, 

otherwise of usual form of genus. Maxillae and maxilliped 

as in Caligus bonito. Sternal furca (Figure 52b) 

tines slightly divergent, blunt tipped. 

Leg 1 (Figure 52d) basipod with large patch of 

fine spinules and 2 plumose setae; exopod first segment 

with row of spinules on inner edge and short 

spine at outer distal corner, second segment with 3 

terminal spines (2 with accessory processes), a terminal 

seta plumose on outer edge and 3 lateral setae, 

each with outer edge armed with stout spinules at 

base becoming finer toward tip and short plumosities 

along inner edge. Leg 2 (Figure 52«) exopod similar 

to other species of genus, endopod first segment with 

a patch of long spinules at outer distal corner, second 

segment with a dense patch of long spinules along 

outer edge, last segment with 3-4 shorter spinules at 

proximal outer edge, all segments with plumose setae 

as in figure. Leg 3 (Figure 53a) similar to other 

species of genus, thumblike spine at outer distal corner 

of exopod first segment only slightly recurved. Leg 4


(Figure 536) exopod 2-segmented, first segment with 

prominent spine at outer distal corner, second segment 

with 3 terminal and 1 lateral spine, distalmost longest 

(setae measure 177, 112, 83, 83, 124 fim distal to 

proximal respectively). Leg 5 represented by 3 short 

setae near posterior corner of genital segment. 

MALE.—None collected. 

REMARKS.—This species is apparently common on 

a number of host species in the western Atlantic and 

Gulf of Mexico (scombrid and non-scombrid). It 

has also been reported from the Gulf of California 

and Pacific coast of Mexico from a number of host 

species by Causey (1960:329) and Wilson (1937:27). 

We describe here a new, closely related, species of 

Caligus from eastern Pacific Scomberomorus and it 

seems likely that at least some of the specimens reported 

by these 2 authors may have been the new 

species. None of that material has been deposited in 

the Smithsonian collections so we have not been able 

to verify the identifications. 

Examination of the type-specimens (USNM 6155) 

from Woods Hole, Mass, indicate variation from our 

specimens on at least 2 points. The outermost of the 

4 terminal setae on the caudal rami of the type specimens 

is less than half the length of the outer edge of 

the ramus, whereas in our specimens that seta is 

almost as long as the outer edge of the ramus. The 

longest seta on the exopod of leg 4 of the type specimens 

is only slightly longer than the adjacent seta 

(218 and 177 /*m) whereas in our material this seta 

is much longer than the adjacent seta (177 in 112 

/*m). We do not consider these differences important 

enough to warrant describing a new species but rather 

may reflect ecological differences between the 2 groups 

of material. 

Caligus omissus, new species 

FIGURES 53c-e, 54-56, 105 


147 9 41 S from the gills, gill arches, gill rakers, inner 

operculum, and roof of the mouth from 41 specimens 

(114 examined) of Scomberomorus sierra from Peru, 

Colombia, Panama, Costa Rica, Pacific coast of Mexico, 

and Baja California; from 7 specimens (47 examined) 

of Scomberomorus concolor from off Sonora, 

Mexico and Gulf of California. Holotype (USNM 

172239), allotype (USNM 172240) and 40$ 3* 

paratypes (USNM 172241) from the gills of S. sierra 

from San Juan Lagoon, Mexico (USNM 219623). 


2.96 mm (2.78-308 mm) based on an average of 6 

specimens from the type series. Greatest width 1.15 

mm (1.12-1.19). Cephalon comprises about 40 percent 

of total length. Frontal lunules prominent, 

diameter of lunule (266 /*m) slightly more than 

least distance (218 /*m) between lunules (interlunular 

space). Genital segment (Figure 53d) slightly longer 

than wide (1.16 X 1-09 mm), posterior corners 

rounded, only slightly produced, comprising about 30 

percent of total length. Abdomen (see Figure 53d) 

1-segmented, nearly 3 times as long as wide (0.96 X 

0.36 mm), usually slightly wider in posterior third 

but never noticeably wider in anterior half; distal 

ventral surface with patches of spinules as in Figure 

53*. Caudal rami (see Figure 53e) longer than wide 

(142 X 94 fixn) armed with 6 plumose setae as indicated 

in figure, longest seta 307 /*m. 

Oral area as in Figure 54a. First antenna (Figure 

546) first segment with 27 setae, all plumose except 

2 in middle anterior group; last segment with 13 

naked setae. Second antenna (Figure 54rf) with 

prominent hook, basal segment with conspicuous 

pointed posterior process. Post antennal process (Figure 

54c) a prominent recurved hook. Postoral process 

(Figure 54e) with slight outward curve, extending 

well beyond tip of mouth tube. Maxilliped (Figure 

54/) with a strongly sclerotized, recurved claw. 

Sternal furca (Figure 54g) with widely divergent 

tines, each with a blunt tip. 

Leg 1 (Figure 55a) basipod with a patch of 

prominent spinules covering two-thirds of ventral 

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 

accessory spines), 1 terminal seta (plumose on outer 

edge), and 3 lateral setae with stout spinules on base 

of outer edge of each followed by short plumosities; 

endopod reduced to a short process with 2 small distal 

spines. Leg 2 (Figure 556) as in other species of the 

genus except exopod spine on outer distal corner of 

last segment very small; endopod first segment with 

3-4 stout spinules on outer distal corner, second segment 

with a double row of stout spinules along outer 

edge, a patch of 4-5 short spinules on proximal outer 

edge of last segment, all segments with plumose setae 

as in figure. Leg 3 (Figure 55c) basipod with con-

NUMBER 3 1 1 29 

spicuous spinules on outer quarter and a patch of 

finer spinules at inner, proximal corner; endopod first 

segment with stout, recurved claw on outer distal 

corner, a short spine on outer distal corner of second 

segment, 3 short spines on outer edge of last segment 

and each segment with plumose setae as in figure; 

endopod with plumose setae as indicated in figure. 

Leg 4 (Figure 55d) exopod 2-segmented, first segment 

with a prominent spine on outer distal corner, second 

segment with one lateral spine and 3 terminal spines, 

distalmost spine longest, spines measure 201, 124, 106, 

100, and 130 /tun distal to proximal spine respectively. 

Leg 5 represented by 3 short plumose setae near 

distal corner of genital segment. 

MALE.—Body form as in Figure 56a. Total length 

3.15 mm (3.08-3.23 mm), based on an average of 

2 specimens. Greatest width 1.27 mm. Cephalon comprising 

about one-half total length. Genital segment 

(Figure 566) longer-than wide (884 X 580 /an). 

Abdomen (Figure 566) 2-segmented, first segment 

nearly square (348 X 348 /*m), second segment longer 

than wide (449 X 334 /*m) with ventral patches of 

spinules as in female. Caudal ramus as in female. 

Appendages as in female except as follows. Postantennal 

process (Figure 56d) a prominent, sharply 

recurved, hook. Second antenna (Figure 56c) with 

2 rugose areas on basal segment and a short bifid claw. 

Maxilliped (Figure 56/) basal segment with a pair of 

sclerotized processes on inner margin opposite tip of 

claw. Sternal furca (Figure 56g) with tines less 

divergent than female. 

Legs 1-4 as in female. Legs 5 and 6 each represented 

by 2 short plumose setae on the genital segment 

as in Figure 566. 

ETYMOLOGY.—The Latin omissus ("neglected") 

alludes to its common association with eastern Pacific 

Scomberomorus species but not collected or, if so, 

not recognized as new. 

REMARKS.—This new species seems to be closely 

related to Caligus mutabilis and C. bonito to a lesser 

extent. These 3 Caligus share a number of characters: 

the similar pattern of fine spinule patches on the 

distal-ventral surface of the abdomen, a prominent 

posterior process arising from the base of the second 

antennae, inner lateral setae of leg 1 with spinules 

along the basal part of the outer edge, prominent, 

stout spinules along the outer edges of the leg 2 

endopod segments, and a 2-segmented exopod of 

leg 4. The new species can be separated from the 

other 2 species as well as all other species of Caligus 

found on scombrids by a combination of the following 

characters: abdomen narrowest anteriorly, widely 

divergent tines on the sternal furca, distal process 

on the base of the second antenna pointed, basipod 

of leg 1 with a large patch of prominent spinules 

(fine spinules in C. mutabilis and C. bonito), the 

spinules on the outer basal margin of the lateral setae 

of leg 1 are fingerlike (short and stout in C. mutabilis 

and C. bonito). 

Wilson (1937:27) reported C. mutabilis from 

Scomberomorus maculatus (=5. sierra) from Mexico 

(Pacific). We suspect that this material was actually 

the new species reported here. Causey (1960:329) 

reported C. mutabilis from the Gulf of California and 

Pacific coast of Mexico from several species of fishes 

including Scomberomorus sierra. It seems likely to us 

that at least the specimens from S. sierra were actually 

C. omissus and some or all of those from the nonscombrid 

hosts may have been as well. 

Caligus omissus was collected from 36 percent (41 

of 114) of the specimens of S. sierra and 15 percent 

(7 of 47) of the S. concolor examined. This would indicate 

that these species may well be the preferred 

hosts for this species. Examination of collections from 

non-scombrid hosts from the Eastern Pacific may well 

turn up additional material of this new species. 

Caligus biseriodentatus Shen, 1957 

FIGURES 57-59, 105 

Caligus biseriodentatus Shen, 1957:352. 

Caligus proboscidatus Heegaard, 1962:161. 

Caligus obovatus Heegaard, 1962:166. 

Caligus auxisi Pillai, 1963:85. 

MATERIAL EXAMINED.—1 $ from the inner surface 

of the operculum of Auxis thazard from Trivandrum, 

India (a specimen of Caligus auxisi donated to the 

Smithsonian by Dr. Pillai) ; holotype (AMS P16418) 

of C. obovatus Heegaard from Scomberomorus 

queenslandicus from Queensland. Holotype (AMS 

PI6420) of C. proboscidatus Heegaard from Scomberomorus 

queenslandicus; from Queensland; 237 

immature 9 and $ from mouth, gill arches, and 

inner operculum of the following hosts and localities: 

Scomberomorous commerson from northwest Malagasy 

Republic, Somalia, Ceylon, East Indies, Philippines, 

China; Scomberomorus guttatus from Arabian 

Sea, Ceylon, Thailand, East Indies (various locali-

30 

ties), Hong Kong; Scomberomorus queenslandicus 

from Onslow (W. Australia), Ambon (Moluccas); 

Scomberomorus plurilineatus from Zanzibar. 


3.15 mm. Greatest width 1.50 mm (measured at 

widest part of cephalon) ; cephalon comprising about 

40 percent of total length. Genital segment somewhat 

longer than wide (0.90 X 0.75 mm). Abdomen about 

twice as long as wide (0.68 X 0.30 mm), 1-segmented; 

dorsal surface distal half with a large patch 

of spinules, ventral surface distal half with groups of 

spinules as in Figure 57b. Caudal rami slightly longer 

than wide, each with a seta on middorsal surface and 

6 subterminal to terminal setae. 

Space between frontal lunules (254 fun) greater 

than diameter of lunule (207 /un). First antenna of 

usual type of genus except last segment about 4 

times longer than wide (177 X 44 /*m). Postantennal 

process fingerlike. Second antenna with usual claw, 

base with posterior spatulate process. Postoral process 

slender. First maxilla represented by 2 short and 1 

long setae (nearly as long as postoral process). Second 

maxilla typical of the genus. Maxilliped with stout 

recurved claw at tip. Sternal furca (see Figure 57c) 

tines nearly parallel and, together, wider than base. 

Leg 1 (Figure 57d) basipod covered with conspicuous 

spinules, a short plumose seta on distal 

margin and a longer plumose seta at outer corner; 

exopod first segment with row of toothlike spinules 

on inner edge, second segment with usual 3 distal 

spines, each with inconspicuous fringe and no accessory 

process, one terminal plumose seta and 3 short 

lateral setae, each with 2-3 stout spinules on outer 

basal border; endopod reduced. Leg 2 of usual form 

of the genus except setae of both rami with a row of 

15-20 conspicuous spinules on proximal outer edge 

(Figure 57«) followed by usual plumosites; endopod 

first segment with group of 3—4 stout spinules on outer 

distal corner, second segment with a double row of 

stout spinules (7-8 spinules in each) along outer edge 

(Figure 57*). Leg 3 similar to other species of genus, 

heavily sclerotized spine on exopod first segment (Figure 

58a) recurved inwardly. Leg 4 (Figure 58&) 

basipod with scattered stout spinules on dorsal surface; 

exopod first segment with a row of spinules on 

outer edge and outer distal spine, second segment 

with 4 spines, distalmost longest and each with a 

fringe at base, spines measure 159, 118, 106, 94, and 

106 ftm distal to proximal respectively. 


MALE.—According to Pillai (1963:87) the second 

antenna basal segment with a bossed pad opposite tip 

of claw. The postantennal process (first maxilla of 

Pillai) a long "sickle-shaped" claw. Maxilliped base 

with 4 low projections on inner border. Total length 

2.70 mm. 

IMMATURE FEMALE.—Body form as in Figure 58d. 

Total length 2.90 mm (3.00-2.70 mm) based on an 

average of 5 specimens. Frontal lunules greatest 

width (236 fim) less than least space (interlunular 

space) between them (265 /*m). First antenna typical 

of the genus. Second antenna with strong claw and 

pointed posterior process on base. Postantennal process 

a short recurved hook. Postoral process a sharply 

pointed process, tip not extending beyond tip of 

mouth tube. Second maxilla typical of the genus. 

Maxilliped (Figure 58

NUMBER 311 31 

between it and C. biseriodentatus. They are as follows: 

space between lunules slightly greater than 

diameter of either lunule; second antenna with a 

posterior process; one of the setae of the first maxilla 

is exceptionally long; base of furca is narrower than 

spread of tines; basipod of leg 1 covered with stout 

spinules; lateral setae on leg 1 exopod very short; 

armature of leg 2 endopod the same; leg 4 exopod 

2-segmented and terminal spines relatively short and 

distalmost spine only slightly longer than adjacent 

spine; armature of caudal rami simliar. Based on these 

common characters we concluded that Caligus auxisi 

represents the adult form of Caligus biseriodentatus. 

We feel that the adult form of this species probably 

prefers a non-scombrid host—possibly a prey species. 

The first author has experienced a similar circumstance 

in a species of Caligus from the Gulf of Mexico 

with the immature stages common on one species of 

fish and the adults found only on others. 

Discussion 

The preceding 4 species (C. bonito, C. mutabilis, 

C. omissus, and C. biseriodentatus) are morphologically 

closely related as pointed out below. They show 

an interesting distribution and host selection. Caligus 

bonito, possibly the ancestral form, is circumglobal 

in distribution and seems to prefer hosts of the tribes 

Thunnini and Sardini, with an infestation rate of 17 

percent in the 11 species infested. A single collection 

was made from Scomberomorus regalis (1 5 1 $ ), 

which was probably an adventitious association. 

Caligus mutabilis apparently infests several species of 

hosts in the Western Atlantic but, of the scombrids 

of that area, it is found on species of Scomberomorus 

with a 3 percent infestation rate on the 3 species infested. 

A single collection (19) was made from 

Scomber japonicus. Caligus omissus apparently is 

restricted to the eastern Pacific and is a common 

parasite on the 2 species of Scomberomorus endemic 

to that area with a 29 percent infestation rate. We 

have not examined non-scombrid hosts of that area 

but we suspect that, as in C. mutabilis in the western 

Atlantic, it occurs on non-scombrid hosts as well. The 

scombrid hosts of Caligus biseriodentatus are Auxis 

species and Scomberomorus. The parasite occurs on 

Scomberomorus only in its immature stages. The 

adults have been collected from Auxis. We believe 

that some other host species is preferred for the 

adult as the only collection of this species in the adult 

form was a single collection from Auxis. 

These 4 species have the following characters in 

common: an elongate, 1-segmented abdomen; second 

antenna with a posterior process on the basal segment; 

caudal ramus longer than wide; leg 1 basipod with 

a prominent patch of spinules; leg 1 lateral setae with 

stout spinules on the outer basal part; leg 2 endopod 

second segment with conspicuous stout spinules on 

the outer edge; leg 4 exopod 2-segmented. Other 

Caligus species possess some of these characters but 

none of those found on scombrids share as many. 

It is interesting to note that 2 of these species 

(C. mutabilis and C. biseriodentatus) apparently 

utilize one host species for the immature forms and 

another for the adult. The relationship between 

Caligus mutabilis and its various hosts is being studied 

by the first author as part of a study of the parasitic 

copepods of the teleost fishes of Charlotte Harbor, 

Florida. 

No counterpart Caligus has been collected from the 

21 specimens of the eastern Atlantic endemic, Scomberomorus 

tritor (the record of Caligus diaphanus 

probably represents an accidental infestation because 

species of Trigla are its usual hosts). 

Caligus cybii Bassett-Smith, 1898 

FIGURES 60-63, 104 

Caligus cybii Bassett-Smith, 1898a: 6. 

Caligus brevisoris Shen, 1957:357. 

Caligus quinqueabdominalis Heegaard 1962:162. 


158 9 and 59 $ from the gills and gill area of the 

following hosts and localities: Scomberomorus commerson 

from Red Sea (Suez), northwest Malagasy 

Republic, Andaman Sea, Gulf of Thailand, Philippines, 

Hong Kong: Scomberomorus koreanus from 

Bombay, Sumatra, Japan; Scomberomorus species 

from New Guinea; Scomberomorus semifasciatus 

from the Gulf of Carpenteria; Scomberomorus sinensis 

from Hong Kong. 


4.84 mm (4.65-5.25 mm), based on an average of 

7 specimens. Greatest width 1.65 mm. Cephalon 

comprises about 40 percent of total length. Genital 

segment (Figure 606) trapizoidal (widest posteriorly), 

somewhat wider than long (1.6 X 1-8 mm), compris-

32 

ing about 20 percent total length, posterior corners 

not produced. Abdomen (see Figure 60fe) incompletely 

divided into 2 segments, widest anteriorly and 

tapering to caudal rami, about 1 x /% times longer than 

wide (2.03 X 0.79 mm) comprising about 25 percent 

of total length. Caudal rami (Figure 60c) small, 

somewhat longer than wide (236 X 200 /nm) armed 

with plumose setae as in the figure. 

Oral area as in Figure 60a". First antenna (Figure 

60«) with 27 setae (most plumose) on first segment 

and 13 naked setae on last segment. Postantennal 

spine (Figure 61a) very small. Postoral spine as in 

Figure 61c. Second maxilla (Figure 61a 1 ) of usual 

caligoid type with 2 terminal, fringed, flagella. Maxilliped 

(Figure 61 e) a stout claw. Sternal furca (Figure 

61/) with divergent tines, not sharply pointed. 

Leg 1 (Figure 61g) basipod with a patch of fine 

spinules, a short plumose seta on posterior border, and 

a long plumose seta on outer distal corner; exopod 

first segment with a row of long spinules along most 

of inner edge and a short, finely serrate spine on 

outer distal corner, last segment with 4, finely serrate, 

spines without accessory spines and 3 setae on inner 

margin (plumosities of setae thickened) ; endopod 

reduced to a short process with 2 small spines at tip. 

Leg 2 (Figure 62a) coxopod with a patch of fine 

spinules on outer surface; basipod with very short 

spine at outer distal corner; exopod first 2 segments 

with heavily sclerotized, finely serrate, inwardly 

directed spines on outer distal corner, last segment 

with 2 short spines and a longer terminal spine with 

prominent plumosities along inner margin, all segments 

with plumose setae in the figure; endopod 

second segment with a prominent patch of long 

spinules on outer edge, last segment with a small patch 

of similar spinules at outer proximal half, all segments 

with plumose setae as in the figure. Leg 3 (Figure 

626) typical of the genus, outer margin of basipod 

with heavy spinules; exopod first segment with stout, 

inwardly directed spine on outer distal corner, other 

2 segments with plumose setae as in figure; endopod 

reduced with 3 incompletely separated segments bearing 

plumose setae as in figure. Leg 4 (Figure 62c) 

exopod 3-segmented; the first 2 each bearing a prominent 

spine on outer distal corner, last segment with 

3 terminal spines, terminalmost longest, the 5 spines 

of the exopod measure 171, 124, 142, 142, and 200 

f*m proximal to distal respectively, each with a conspicuous 

fringe at base and on edges as figured. Leg 


5 represented by a pair of small plumose setae at 

outer posterior corners of genital segment. 

MALE.—Body form as in Figure 62rf. Total length 


specimens. Genital segment (Figure 63a) somewhat 

longer than wide (841 X 739 /xm). Abdomen 2segmented, 

somewhat longer than wide (580 X 

450 /Am), second segment about 2/2 times length of 

first. Caudal rami as in female. 

Cephalic appendages as in female except second 

antenna (Figure 636) with rugose areas on basal 

segment and a bifid claw; postoral process (Figure 

63c) with an accessory process and spine near distal 

two-thirds; maxilliped (Figure 63d) basal segment 

with sclerotized processes opposite tip of claw, claw 

more robust than in female. 

Legs as in female except leg 5 represented by 3 

short, plumose, setae at each lateral mid-margin and 

leg 6 represented by 3 similar setae at posterior distal 

corners of genital segment (see Figure 63a). 

REMARKS.—So far this species is reported only from 

Indo-West Pacific species of Scomberomorus. Collection 

data indicates this parasite is usually found on the 

gills, gill arches, operculum, and pseudobranch of 

the host. 

Caligus pelamydis Kroyer, 1863 

FIGURES 64-66, 104 

Caligus pelamydis Kroyer, 1863:124. 

Caligus scomberi Bassett-Smith, 1896:11. 

Caligus scombri Scott, T., 1901:148. 

Parapetalus sp. Silas and Ummerkutty, 1967:908. 

This species has been recorded many times from 

scombrid and non-scombrid hosts (see Lewis, 1967: 

139). Shiino, 1965 and Lewis, 1967 have provided 

good descriptions of this species. We will discuss only 

those features of special taxonomic importance in 

addition to including figures. A complete synopsis 

of the literature has been provided by Margolis, 

Kabata, and Parker, 1975:59. 


97 9 and 4 $ from the gill arches, gills, and inner 

wall of the operculum of the following hosts and 

localities: Scomber scombrus from France; Scomber 

japonicus from Gulf of Mexico, Florida (Atlantic), 

Gulf of Guinea; Scomberomorus niphonius from 

Japan; Auxis species from New South Wales; Euthynnus 

affinis from New South Wales; Euthynnus allet-

NUMBER 311 33 

teratus (locality unknown); Sarda sarda from Argentina, 

Gulf of Guinea, Tunis, South Africa (Port 

Elizabeth); Sarda australis from New South Wales; 

Sarda chiliensis lineolatus from California. 


total length 3.00-^.65 mm. As in other species of 

Caligus with widespread distribution, the size differences 

seem to be correlated with water temperature 

rather than host species. 

Caligus pelamydis females can be separated from 

other species of Caligus found on scombrids by the 

following combination of characters: cephalon 40-45 

percent of total length; abdomen 23-25 percent of 

total length; second and third segments of leg 2 

endopod (see Figure 65«) each with a large patch of 

fine spinules along outer edge; leg 4, 3-segmented 

with last segment produced distally to give segment a 

triangular shape (see Figure 666) and prominent 

fringes at bases of all setae, tines of sternal furca 

spatulate and as wide or wide than base (Figure 

65c). 

MALE.—Body form as in Figure 66c. Range of total 

length 2.48-2.55 mm (based on 3 specimens from 

Sarda sarda, Gulf of Guinea). Males are generally 

rare in collections (comprising only about 4 percent 

of the total sample studied). Shiino (1965:411), 

described the male of this species comparing it to an 

immature female in the same collection. Certain appendages 

differ between the sexes but some of those 

noted by Shiino were probably due to the immaturity 

of the female. In our sample we found the following 

differences based on adults of both sexes: second 

antenna of male with rugose patch along posterior 

edge of basal segment; postoral process of male (Figure 

66

34 

with long inwardly directed spine, on outer distal 

corner, second segment with somewhat shorter similar 

spine, last segment with similar spine about half 

length of that of the first segment, exopod segments 

otherwise armed with plumose setae as in the figure; 

endopod first segment with a row of short spinules 

along outer edge and a patch of longer spinules at 

outer distal comer, second segment with a dense patch 

of long spinules along outer edge, third segment with 

a few long spinules at outer proximal edge, endopod 

with plumose setae as in the figure. Leg 3 (Figure 

69a) exopod first segment with stout, slightly recurved 

spine, on outer distal corner, second segment 

with short, weak, spine on outer distal corner, 

last segment with 3 weak spines and 4 plumose setae; 

endopod with 2 segments bearing plumose setae as 

in the figure. Leg 4 (Figure 696) exopod first segment 

outer edge produced as approximate right angle at 

distal two-thirds; last segment with distalmost spine 

2/2 times length of others (442 X 177 /*m). Leg 5 

represented by 3 short plumose setae near edge of 

genital segment at distal three-fourths. 

MALE.—Body form as in Figure 69

NUMBER 3 1 1 35 

corner and an inner seta, last segment with 3 outer 

spines and 5 terminal to inner setae; endopod first 

segment with an inner seta, second segment with a 

patch of long spinules along outer edge and 2 inner 

setae, last segment with a patch of long spinules on 

outer edge and 6 setae. Third leg (Figure 72a) 

basipod with scattered short spinules near outer margin; 

exopod first segment with a prominent spine on 

outer distal corner, second segment with a short spine 

on outer distal corner and inner seta, last segment 

with 3 short outer spines and 4 terminal to inner 

setae; endopod first segment with an inner seta, last 

segment with 6 terminal setae. Fourth leg (Figure 

72b) basipod with a small seta on outer distal corner; 

exopod first segment with a fringed seta (112 /Am) 

on outer distal corner, second segment with fringed 

seta (136 /*m) on outer distal corner, last segment 

with 3 fringed setae (153, 165, and 206 /*m long 

respectively), a fringe at base of each seta. Fifth leg 

represented by 4 setae near the posterior distal corner 

of the genital segment. 


2.39 mm, greatest width 1.30 mm. Genital segment 

about as long as wide (0.43 X 0.45 mm). Abdomen 

2-segmented segments measure 88 X 250 fixn and 

292 X 280 jan (length X width) respectively. Caudal 

rami as in female. Appendages as in female except 

as follows. Second antenna (Figure 72d) with accessory 

process on claw and bossed areas on segments as 

illustrated. Postoral process (Figure 72«) with accessory 

process at distal two-thirds. Maxilliped (Figure 

72/) base with 2 well-developed processes opposing 

tip of claw, portion of claw distal to seta proportionately 

shorter than in female. Fifth leg represented by 

4 setae at mid-lateral margin of genital segment. Sixth 

leg represented by 3 setae at posterior distal corners 

of genital segment. 

REMARKS.—This species has been reported by 

numerous authors from the west coasts of Europe and 

Africa (see Margolis, et al, 1975). It is apparently 

commonly found on species of Trigla but has been 

reported from several other hosts as well. Our report 

here is the first from a scombrid host and in light 

of the many previous literature citations it should not 

be considered a primary parasite of scombrids. We 

have included its description here as no complete 

description exists in modern literature. The specimens 

studied in this description were compared with ma- 

terial collected by us from Trigla lineata from off 

Naples, Italy. • 

Caligus savala Gnanamuthu, 1948 

FIGURE 73a-/ 

Caligus savala Gnanamuthu, 1948:591. 

Caligus affinis Kurian, 1961:71. 

Caligus acutus Kirtisinghe, 1964:66. 

Gnanamuthu described this species from specimens 

collected from the plankton and a single male from 

the body surface of Trichiurus savala Cuvier. Kurian 

collected 2 9 from the body surface of Euthynntu 

affinis (described as C. affinis). Kirtisinghe, 1964, 

noted that Caligus affinis was preoccupied and renamed 

the species Caligus acutus Kirtisinghe. Pillai, 

1971:164 redescribed the species from type material 

of C. savala deposited in the British Museum and 

placed C. affinis and C. acutus in synonymy with C. 

savala. 

The single record of this species from a scombrid 

indicates that its occurrence on such a host was 

probably accidental. 

The female may be characterized as follows. Body 

form as in Figure 73a. Second antenna (Figure 736) 

with pointed posterior process on base. Postantennal 

process (Figure 73c) not strongly curved. Sternal 

furca (Figure 73a") with tines only slightly divergent 

and spreading only slightly wider than base. Leg 2 

endopod with patches of slender spinules along outer 

margin of each (entire margin of segment 1 with 

spinules—usually only at outer corner of other 

species). Stout spine at the outer distal corner of 

leg 3 exopod first segment not curved inwardly. Leg 

4 exopod (Figure 73/) 2-segmented, distalmost spine 

about one-third longer than adjacent spine. 

We did not collect this species and have based the 

brief description and figures on Pillai's redescription 

in 1971. 

Caligus macarovi Gussev, 1951 

FIGURES 73g-i, 74o-6 

Caligus macarovi Gussev, 1951:408. 

Caligus fulvipurpureus Shiino, 1954:150. 

Gussev described this species from specimens collected 

from 4 species of fish including Auxis and 

Cololabis saira (Brevoort) from the Sea of Japan.


Shiino later (1954:150) reported it (as C. fulvipurpureus) 

from the body surface of several Cololabis 

saira from Japan and again (Shiino, 1959b:277) 

from off Mexico (Pacific) as C. macarovi. This parasite 

apparently is common on Pacific Cololabis (also 

reported from this host by Kazachenko, et al, 1972: 

224). It is not surprising that a copepod common on 

a prey species of fish (Cololabis) would occasionally 

be found on its predator species. The female of this 

species is characterized by the following. Cephalon 

comprises about 40 percent of total length (see Figure 

73g). Genital segment about as wide as cephalon, 

sides nearly parallel. Abdomen one-segmented, about 

twice as long as wide. Second antenna with usual 

claw, posterior process on basal segment. Postantennal 

spine short, angle of inner curve greater than 90° 

(see Figure 73h). Maxilliped with 2-segmented claw. 

Sternal furca (Figure 73*) tines not widely divergent, 

not sharply pointed. Leg 1 basipod with patch of 

spinules, 3 inner lateral setae of last segment with 

spinules on outer basal part followed by fringe. Leg 

2 endopod segments (Figure 74a) each with a dense 

patch of spinules on outer edge. Leg 4 (Figure 74fc) 

exopod 2-segmented, last segment without midlateral 

spine, distalmost spine longer than adjacent spine. 

Male as in female except in body form. Second antenna 

with short claw and rugose areas on basal 

segment. Maxilliped with rugose area on basal segment 

opposite tip of claw. 

We did not collect this parasite during the course 

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). 

Caligns amblygenitalis Pill a i, 1961 

FIGURE l\c-g 

Caligns amblygenitalis Pillai, 1961:98. 

Pillai describted this species from a single female 

collected from the body surface of Euthynnus affinis 

from Vizhingom, India. It has not been recorded 

since. 

The female is distinguished by the following characters. 

Frontal lunules widely separated, interlunular 

space about 3 times diameter of lunule (according to 

Pillai's figure). Genital segment about as long as 

cephalon and with lobed posterior corners. Abdomen 

1-segmented, about twice as long as wide. Second 

antenna with posterior process on basal segment. 

Postantennal process (first maxilla of Pillai) long 

and blunt. Sternal furca with widely divergent tines, 

spread of tines wider than base. Leg 1 lateral setae 

with spinules on basal part of outer edge. Leg 2 

endopod segments with a patch of narrow spinules 

on the outer edge of each, second spine on exopod 

with strong teeth on outer edge. Leg 4 exopod 2segmented, 

bearing 4 spines (no lateral spine on last 

segment which is usually present in other species when 

exopod is 2-segmented), distalmost spine about onethird 

longer than adjacent spine. 

Caligus pseudokalumai Lewis, 1968 

Caligns pseudokalumai Lewis, 1968:59. 

This species was described from a single female 

collected from the body surface of a Gymnosarda 

unicolor from Eniwetok Atoll. Lewis compared this 

species with a previously described species, Caligus 

kalumai Lewis, 1964, collected from Acanthurus 

guttatus from Hawaii. The differences between the 

2 species are slight and Lewis suggested that their 

close relationship should be more closely examined 

when additional material is available. We feel that 

this species represents an accidental infestation and 

Gymnosarda unicolor is not its usual host. Lewis has 

provided a good description with illustrations and 

since this is undoubtedly not a scombrid parasite we 

will not repeat a description here. 

Elytrophora Gerstaecker, 1853 

Elytrophora brachyptera Gerstaecker, 1853 

FIGURES 75-77, 106 

Elytrophora brachyptera Gerstaecker, 1853:58. 

Elytrophora hemiptera Wilson, 1921:4. 


771 9 269 $ from the gill area of the following hosts 

and localities: Thunnus alalunga from He Amsterdam, 

Chile, New Jersey, Brazil (north coast), North 

Atlantic (25°N, 35°W), Azores Islands; T. albacares 

from Chagos, Australia (NSW), Christmas Island 

(Pacific), Hawaii, New Jersey, Brazil (north coast) ; 

T. atlanticus from St. Thomas Island; T. maccoyii 

from Western Australia (Albany) ; T. obesus from 

He Amsterdam, Seychelles, Christmas Island (Pacific),

NUMBER 3 1 1 37 

Hawaii, Juan Fernandez Islands, New Jersey, Brazil 

(north coast), Cape Verde Islands, Canary Islands, 

Azores Islands; T. thynnus from South Africa (west 

coast), Eastern Pacific, Western North Atlantic, New 

Jersey, Portugal (Lisbon); Allothunnus fallai from 

New Zealand, California. 


8.26 mm, greatest width 4.71 mm (measured at widest 

part of cephalon). Genital segment longer than 

wide (2.61 X 1-89 mm). Abdomen 2-segmented, segments 

measure (1 X w) 0.72 X 0.87 mm, 0.68 X 

0.85 mm respectively; first segment partially hidden 

in dorsal view by flaps of genital segment. Caudal 

rami (Figure 756) longer than wide (680 X 400 /*m), 

each with 6 setae (2 very small). First antenna (Figure 

75c) 2-segmented. Second antenna (Figure 75d) 

basal segment with prominent triangular process, 

terminal segment clawlike with 1 short spine and 1 

seta. Mouthtube, mandible, post oral process and first 

maxilla as in Figure 75e; mandible with 12 teeth; 

post oral process about as long as mouthtube; first 

maxilla with 3 small setae. Second maxilla (Figure 

75/) terminal segment with 2 prominent pectinated 

membranes at about mid-point, terminally with 2 

long curved, subequal fringed processes. Maxilliped 

(Figure 75g) claw distal half finely grooved. Sternal 

furca (Figure 76a) with stout, widely divergent tines. 

Leg 1 (Figure 76b) biramous, endopod first segment 

without ornamentation, second segment spinulose 

along outer edge, 3 inner setae; exopod onesegmented 

with 3 stout, fringed outer spines, 1 

terminal plumose seta, 3 inner plumose setae. Leg 2 

(Figure 76c) biramous, rami 3-segmented, endopod 

first and second segments each with spinulose outer 

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 

seta, second segment with 2 inner setae, third segment 

small, with 4 setae; exopod as in Figure 77a. Leg 4 

(Figure 77b) endopod 2-segmented, first segment 

with inner seta, second segment incompletely divided 

with one short inner seta and 3 short terminal setae; 

exopod first segment with outer spinulose spine, 

second segment similar, with inner seta, third segment 

with 3 outer to terminal spines and 4 short inner 

setae. Legs 5 and 6 represented by 1 and 3 setae 

respectively on lateral genital segment. 

MALE.—Body form as in Figure 77d\ Total length 

6.67 mm, greatest width 3.62 mm (measured at widest 

part of cephalon). Genital segment (Figure 77«) 

longer than wide (1.74 X 1.53 mm). Abdomen 2segmented, 

segments measure (1 X w) 0.56 X 0.72 

mm, 0.52 X 0.69 mm respectively. Caudal rami similar 

to female. First antenna 2-segmented, similar to 

female. Second antenna (Figure 77/) terminal segment 

long, recurved claw with stout spine on inner 

margin; triangular process on basal segment not as 

prominent as in female. Remaining cephalic appendages 

except maxilliped similar to female. Maxilliped 

modified as in Figure 77g. Sternal furca slender, tines 

less divergent than in female. Legs 1-6 similar to 

female, except leg 4 endopod setae longer. 

REMARKS.—Examination of Wilson's types of E. 

hemiptera indicate that they are actually E. brachyptera 

and we have placed his species in synonomy. 

Hewitt (1968) made a tentative revision of the genus 

Elytrophora in which he considered E. brachyptera 

to be the only species with 2 additional subspecies 

{atlantica and indica). We feel that E. indica is a 

valid species and have indicated it as such below. 

Elytrophora brachyptera is a circumglobal species 

found on several species of Thunnini. 

We noticed that material collected from colder 

waters (New Zealand and lie Amsterdam) show a 

variation in the armature of leg 4. One of the 4 inner 

setae is strongly developed whereas in specimens from 

more temperate waters all of these setae are reduced 

(compare Figures 77& and c). We did not feel this 

variation warranted designating the colder water 

material as a new species. These variant specimens 

were collected from the lie Amsterdam specimen of 

Thunnus obesus and the southernmost specimens of 

Allothunnus. 

Elytrophora indica Shiino, 1958 

FIGURES 78, 79a,b, 106 

Elytrophora indica Shiino, 1958:107. 

Elytrophora brachyptera indica Shiino.—Hewitt, 1968:124. 


128 $ 27 $ from the gill area of Thunnus obesus from 

He Amsterdam, Seychelles, Christmas Island (Pacific), 

Juan Fernandez Islands. 

FEMALE.—Body form as in Figure 78a. Total 

length 10.15 mm, greatest width 5.80 mm (measured 

at widest part of cephalon). Genital segment (Figure 

78fc) longer than wide (2.97 X 2.20 mm). Abdomen

38 

(see Figure 786) first segment crescent-shaped, produced 

at posterior corners to envelope anterior half 

of second segment; segments measure (1 X w) 1.01 X 

1.50 mm, 1.02 X 1.10 mm respectively. Caudal rami 

longer than wide (0.65 X 0-43 mm) similar to E. 

brachyptera. First antenna (Figure 78

NUMBER 311 39 

a partially dissected male was in the vial when we 

received it from the Australian Museum. Examination 

of this specimen and Heegaard's figures of the 

female lead us to believe that this species was erroneously 

placed in the Euryphoridae by the author. 

His basis for doing so was the presence of dorsal 

plates covering the fourth thoracic segment. His figures 

do not indicate these and the male specimen does 

not have them. Unfortunately, lacking the female, we 

could not resolve the problem of its true taxonomic 

position. We hope that additional material of this 

interesting species will be collected eventually and its 

taxonomic position made clear. 

We have refigured the male maxilliped (Figure 

80c), sternal furca (Figure SOd), leg 1 (Figure 80*), 

and leg 2 (Figure 80f). 

The original collection was made from the body 

surface of Euthynnus affinis (reported as E. alletteratus) 

from Howick Island, north Queensland, 

Australia. 

The total length of the male is 3.35 mm. 

Tuxophorus Wilson, 1908 

Tuxophorus cybii Nunes-Ruivo and Founnanoir, 

1956 

FIGURE 81a 

Tuxophorus cybii Nunes-Ruivo and Fourmanoir, 1956:76. 

Tuxophorus solandri Kurian, 1961:72. 

We did not collect this species but we have reproduced 

the figure of the female based on the illustration 

of Nunes-Ruivo and Fourmanoir. In 1961 Kurian 

described a new species, T. solandri, based on a single 

specimen from the body surface of Acanthocybium 

solandri from India. Kurian was apparently unaware 

of the description of T. cybii earlier as he made no 

mention of it. Comparisons of the 2 descriptions leave 

no doubt that they represent the same species and, 

consequently, we have placed Kurian's T. solandri 

in synonymy with T. cybii. 

Furthermore, we suspect that T. cervicornis of Heegaard 

(described below) may also be a synonym of 

T. cybii. The only difference we find between Heegaard's 

species and T. cybii and T. solandri is the 

highly developed processes on the outer distal corners 

of T. cervicornis (absent in the other 2). This could 

be accounted for if the descriptions of T. cybii and 

T. solandri are based on immature specimens. Each 

description is based on a single specimen and neither 

Nunes-Ruivo and Fourmanior or Kurian show egg 

strings in the illustrations (Heegaard does). 

One of our collections of T. cervicornis is from 

Pakistan, extending the range of that species to within 

the known range of T. cybii. 

Since we did not collect specimens of T. cybii or 

immature T. cervicornis we feel it would be premature 

to place T. cybii in synonymy. We trust that future 

collection will bear out our suspicions regarding the 

synonymy of these 2 species. 

Tuxophorus cervicornis Heegaard, 1962 

FIGURE 81 b-g 

Tuxophorus cervicornis Heegaard, 1962:172. 


from the body surface of Scomberomorus commerson 

from Coffs Harbor, New South Wales, Australia, and 

Karachi, Pakistan. 

FEMALE.—Body form as in Figure 81fe. Total length 

6.80 mm. Greatest width 3.15 mm. Cephalon slightly 

longer than wide (3.10 X 3.15 mm). Thoracic segment 

bearing leg 4 with dorso-lateral winglike plates. 

Genital segment slightly wider than long (2.10 X 2.25 

mm, including posterior processes) ; outer posterior 

corners produced. Abdomen 3-segmented. Caudal 

ramus about 6.5 times longer than wide (1.95 X 0.30 

mm). Oral appendages as in T. collettei except sternal 

furca with pointed tines (Figure 81tf) rather than 

spatulate. Leg 1 as in T. collettei except spinules on 

terminal spines thicker in T. cervicornis. Legs 2-3 

as in T. collettei. Leg 4 (Figure 81g) exopod 3-segmented, 

terminalmost spine shorter (relative to adjacent 

spine) than in T. collettei. 

MALE.—Unknown. 

REMARKS.—This species has been reported from 

Scomberomorus commerson from northern Australia 

and Pakistan. See the discussion following the description 

of T. cybii for comparisons with that species. 

Tuxophorus collettei, new species 

FIGURES 82-84 

MATERIAL EX MINED.—Holotype $ (USNM 

172250), allotype (USNM 172251), and 13 $ 1 $


paratypes (USNM 172252 collected from the body 

surface of Scomberomorus regalis caught off the Virgin 

Islands, 8 June 1967 during a cruise of the R.V. 

Oregon II. Collected by C. A. Child. 


(including caudal rami) 9.0 mm, greatest width 

(measured at widest part of cephalon) 2.33 mm. 

Cephalon somewhat longer than wide (2.55 X 2.33 

mm). Thoracic segments bearing legs 1-3 fused, incorporated 

within posterior lateral lobes of cephalon. 

Thoracic segment bearing leg 4 with lateral winglike 

plates, posterior margins of plates reaching to "shoulders" 

of genital segment. Genital segment longer than 

wide (1.58 X 1-35 mm) with lateral margins parallel 

giving segment a boxlike shape; distal corners rounded 

and slightly produced, ventral surface with 2 lateral 

patches of stout spinules (Figure 82b) ; a dorsal, 

heavily sclerotized process near each distal corner 

(Figure 82a). Abdomen about 6 times longer than 

wide (3.45 X 0-58 mm) narrowing near mid-margin 

and near distal end. Caudal rami about 3.8 times 

longer than wide (490 X 129 fim), widest proximally; 

2 short setae on each lateral margin at distal twothirds, 

1 subterminal inner seta, 3 short, plumose, 

terminal setae. 

First antenna (Figure 82*/) 2-segmented; first segment 

with 13 short, stout, plumose setae, second segment 

with 11 naked setae. Second antenna (Figure 

82tf) robust with stout recurved claw, claw with short 

seta near base of longer seta near middle of outer 

edge. Mandible of usual caligoid type with 12 teeth 

at tip. First maxilla represented by 3 short setae 

lateral to mouthtube near base of post-oral process. 

Second maxilla (Figure 83a) long and slender, a patch 

of long hairs (compound setae?) at junction of first 

and second segment, a palmate fringe near midmargin 

of second segment; 2 terminal fringed flagella, 

outermost about twice length of inner. Maxilliped 

(Figure 83b) basal segment with a short, stout seta 

near inner mid-margin; second segment in form of a 

recurved claw bearing a short seta near middle. Sternal 

furca (Figure 83c) with short, divergent, truncate 

tines, tines with lateral flanges on basal half; accessory 

processes lateral to furca. 

Legs 1-3 biramose. Leg 1 (Figure 83d) basipod 

with posterior seta and plumose seta on outer distal 

corner; exopod first segment with row of hairs along 

posterior border and short spine on outer distal corner; 

second segment with 3 terminal spines, outermost with 

inner row of spinules, others bifid and with broad 

serrations on inner and outer edges, each spine with 

a fringe near its base, a short terminal seta and 3 

lateral plumose setae: endopod a small process near 

base of exopod bearing a row of hairs near tip. Leg 

2 (Figure 83e) basipod unarmed except a prominent 

fringe on both inner and outer margins; exopod 3segmented, 

first segment with a stout serrate spine, 

inwardly directed, on outer distal corner and an inner 

seta, second segment armed as first except spine smaller 

and more finely serrate, third segment with 2 outer 

spines (terminalmost finely fringe) and 6 plumose 

setae; endopod first segment outer edge with a row 

of long spinules and an inner seta, second segment 

with 2 inner setae and spinules on outer margins 

(smaller than those on first segment), third segment 

with spinules on outer edge and 6 plumose setae. 

Third leg (Figure 84a) basipod with patch of spinules 

on outer half and smaller patch of finer spinules 

near inner margin; exopod first segment with stout 

spine on outer distal corner and fringe on outer 

margin, second segment with spine on outer distal 

corner, inner seta, and patch of long spinules on outer 

edge, last segment with 3 outer spines, 4 short terminal 

to inner setae, and long spinules on outer edge; 

endopod first segment with patch of spinules along 

outer edge and inner seta, second segment with patch 

of spinules on outer edge and 6 plumose setae. Leg 4 

(Figure 84b) basipod unarmed; exopod 3 segmented, 

first and second segments each with a fringed spine 

on outer distal corner, last segment with 3 fringed 

spines measuring 82, 112, and 236 fim respectively, 

each exopod spine with a fringe near its base. No 

evidence of legs 5 or 6 could be found. 

Egg strings of usual caligoid type, reaching well 

beyond tip of abdomen. 


(including caudal rami) 6.15 mm, greatest width 

(measured at widest part of cephalon) 2.68 mm. 

Cephalon about as wide as long (2.68 X 2.33 mm). 

General morphology of cephalon as in female. Genital 

segment (Figure 84a 1 ) longer than wide (1.3 X 0.9 

mm), ventral surface with spinules as indicated in 

figure. Abdomen similar to that of female except only 

3.7 times as long as wide (2.68 X 0.73 mm), small 

spinules on ventral surface. Caudal rami as in female 

(590 X 141 fim). Cephalic appendages as in female 

except second antenna (Figure 84e) with large rugose 

areas on basal segments and maxilliped with

NUMBER 3 1 1 41 

small rugose knob on basal segment opposing tip of 

claw. 

Legs 1-4 as in female. Legs 5 and 6 absent. 

REMARKS.—Of the 5 previously described species 

(not including T. tylosuri, which was replaced in the 

genus Caligus by Pillai, 1961) the new species can 

be separated from T. wilsoni Kirtisinghe and T. 

caligodes Wilson by the long, slender caudal ramus 

of the new species (nearly square in T. wilsoni and 

T. caligodes). It can be separated from T. cybii Nunes- 

Ruivo and Fourmanoir, T. solandri Kurian (synonymous 

with T. cybii), and T. cervicornis Heegaard by 

the much longer abdomen, the spatulate tines of the 

sternal furca, and the relatively longer inner seta on 

the end of leg 4 (twice as long as other leg 4 setae) 

of T. collettei. 

ETYMOLOGY.—This species is named for Dr. Bruce 

Collette whose enthusiastic support and collecting 

efforts made this paper considerably more comprehensive 

than it would have been otherwise. 

Pseudocycnus Heller, 1868 

Pseudocycnus appendiculatus Heller, 1868 

FIGURES 85-87, 131-136a 

Pseudocycnus appendiculatus Heller, 1868:218. 

Pseudocycnus spinosus Pearse, 1952:30. 

Pseudocycnus thynnus Brandes, 1955:190. 


85 9 4 $ from the gills of the following hosts and 

localities: Thunnus tonggol from Pakistan, Gulf of 

Thailand; T. albacares from Seychelles Island, Somalia, 

Australia (Queensland), Philippines, Caroline 

Islands, Kapingamarangi Atoll, Peru, Costa Rica 

(Pacific), Juan Fernandez Islands, Gulf of Guinea; 

T. obesus from Juan Fernandez Islands, North Atlantic 

(40°N, 49°W); Katsuwonus pelamis from 

Malagasy Republic, Surinam, Gulf of Guinea; Euthynnus 

alletteratus from Florida (West Coast), Gulf 

of Mexico; E. affinis from Malagasy Republic, Gulf 

of Thailand. 

FEMALE.—Body form as in Figure 85a. Thoracic 

segments bearing legs 2 and 3 free, without lateral 

processes. Abdomen short, incompletely separated from 

genital segment. Caudal rami long, about one-half 

length of genital segment, fused with abdomen. First 

antenna (Figure 85b) 7-segmented; first segment 

without setae; second segment with 4 setae, outer distalmost 

stout (not as prominent as in Pseudocycnoides), 

other segments with setae as in the figure. 

Second antenna (Figure 85c) with heavily sclerotized 

claw, 2 short, stout setae on inner margin. Mouthtube 

(Figure 85d) without ornamentation on labrum. Mandible 

(Figure 85d) with 7 teeth at tip. First maxilla 

(Figure 85«) with 2 stout and 1 weak setae at tip. 

Second maxilla (Figure 85/) with irregular rows of 

stout spinules along outer edge of distal 2 segments. 

Maxilliped (Figure 86a) with wide basal articulation, 

basal segment narrows proximally to articulate with 

claw; claw with bifid tip and stout accessory process 

near mid-margin. 

Leg 1 (Figure 86b) biramous; basipod with 2 patches 

of long spinules; exopod 1-segmented, bearing 3 

short, stout spines at tip; endopod mostly obscured 

by fleshy tissuelike processes from basipod, bearing 

2 spines and 2 setae at tip. Leg 2 (Figure 86c) basipod 

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; 

endopod with 3 stout, terminal spines, 2 groups of 

spinules as figured; both rami 1-segmented. Leg 3 

(Figure 86d) basipod with a large patch of spinules; 

exopod fused with basipod and bearing 3 spines at tip. 

Leg 4 represented by a single lateral seta at proximal 

part of genital segment. Egg strings long. 

MALE.—Body form as in Figure 87a. Total length 

2.70 mm. Greatest width 0.75 mm. Genital segment 

less than half total body length (520 X 370 /xm 

length by width). Abdomen (Figure 87b) longer than 

wide (300 X 220 fim). Caudal rami (see Figure 81b) 

somewhat longer than abdomen and about 4 times 

as long as wide (450 X HO [tin) ventral surface with 

5 patches of sclerotized knobs. 

First antenna (Figure 87c) 6-segmented (fourth 

segment of male is fused fourth and fifth segments 

of female), segment with bulbous spine on outer distal 

corner, remaining segments with setae as in the figure. 

Maxilliped (Figure 87d) base with patches of short 

spinules on inner margin and a heavily sclerotized 

claw, a short seta at outer distal corner of penultimate 

segment. Other cephalic appendages as in female except 

first maxilla with 1 less seta and second maxilla 

with more spinules at tip. 

Leg 1 (Figure 81 e) basipod with 2 patches of stout 

spinules, a seta at outer distal corner, a prominent, 

heavily sclerotized process arising from inner mid-


margin, process extends to tip of exopod; exopod with 

1 spine at tip and surface ornamentation as in the 

figure; endopod an elongate process modified at tip 

giving it a tubelike appearance (see SEM photo). Leg 

4 (Figure 87/) a prominent lateral process with a 

single seta at tip. Other legs as in the female. 

REMARKS.—This species has been recorded many 

times from scombrids and is apparently circumglobal 

in distribution. Kabata (1970:171) summarized the 

known hosts and distribution to that time. 

Our collection indicates that this parasite is most 

common on scombrids of the tribe Thunnini and to a 

lesser extent Sardini. The 6 species of Thunnini infested 

were parasitized at a rate of nearly 10 percent 

and the single species of Sardini about 2 percent. 

Females of this species show considerable variation 

in total length. Lewis, et al. (1969) have presented 

data based on hosts and locality. Total lengths range 

from 11-18 mm with the longer specimens from 

colder waters. 

Pseudocycnoid.es Yamaguti, 1963 

Pseudocycnoides armatus (Bassett-Smith, 1898) 

FIGURES 88-91a-fe, 107, 1366-/, 137 

Hellaria armata Bassett-Smith, 1898a: 10. 

Cybicola armata.—Bassett-Smith, 1898b:371 

Pseudocycnus armatus.—Kirtisinghe, 1935:339. 

Paracycnus lobosus Heegaard, 1962:182. 

Pseudocycnoides armatus.—Yamaguti, 1963:172. 

Pseudocycnoides rugosa Kensley and Grindley, 1973:104. 


149 5 11

NUMBER 3 1 1 43 

Leg 1 (Figure 90*) biramous, each ramus 1-segmented 

and not distinctly separate from basipod; 

basipod with patch of slender spinules, triangular 

process, and several lobate processes; exopod with 

dense outer patch of spinules, smaller spinules in semicircle 

and 2 medial lobes, 2 spines and 3 lobed 

processes distally; endopod produced as a slender, 

elongate, heavily sclerotized spine with several small 

distal processes, spine with membranelike covering. 

Leg 2 (Figure 91a) biramous, each ramus 1-segmented; 

basipod with dense inner patch of spinules, 

2 lighter patches of spinules and long, slender outer 

spine; exopod with irregular distal row of short 

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 

of spinules, 1-2 rows of hairs at bases of 3 terminal 

spines, innermost spine elongate, densely spinulose 

around entire distal three-fourths, tapering gradually. 

Leg 3 (Figure 916) similar to female. Legs 4 and 5 

represented by simple spines on lateral genital segment. 

REMARKS.—This species is common on species of 

Scomberomorus throughout the Indo-West Pacific 

(except S. niphonius). We examined the type material 

of P. rugosa Kensley and Grindley and concluded that 

their material was actually P. armatus. Their conclusion 

that P. rugosa represented a new species was 

based on comparisons with poor literature descriptions 

of P. armatus (B. Kensley, pers. comm.). We place 

P. rugosa in synonymy with P. armatus. 

Pseudocycnoides scomberotnori (Yamaguti, 1939) 

FIGURES 91 c-g 

Pseudocycnus scomberomori Yamaguti, 1939:457. 

Pseudocycnoides scomberomori.—Yamaguti, 1963:172. 


from the gills of Scomberomorus niphonius from 

Korea, North China, Japan. 

FEMALE.—Body form as in Figure 9It. Total length 

4.8 mm. Greatest width 0.9 mm. Abdomen and caudal 

rami of about equal length (0.6 mm) ; abdomen much 

shorter in P. armatus. Appendages as in P. armatus 

except as described below. Maxilliped (Figure 91 d) 

claw longer and tip more recurved. Leg 1 (Figure 9\e) 

basipod with large patch of spinules (spinules on outer 

edge only in P. armatus). Leg 2 (Figure 91/) endopod 

longest spine extending beyond tip of exopod spines 

and heavily spinulose (longest spine on endopod of 

P. armatus much shorter and with few spinules). Leg 

3 (Figure 91 g) outer spine not as robust as in P. 

armata. 

MALE.—Unknown. 

REMARKS.—This species is apparently restricted to 

Scomberomorus niphonius, which is endemic to waters 

of North China and Japan. Yamaguti described this 

parasite from S. chinensis (probably S. niphonius) 

from the Inland Sea of Japan. 

Pseudocycnoides scomberomori is closely related to 

P. armatus but easily separated from it by the characters 

cited above. 

Pseudocycnoides buccata (Wilson, 1922), 


FIGURES 92-94a-c, 107, 138, 139 

Pseudocycnus buccata Wilson, 1922:79. 

Cybicola elongate Pearse, 1951:365. 

Pseudocycnopsis buccata (Wilson).—Yamaguti, 1963:172. 


667 5 10


bearing 2 short terminal setae. Entire body covered 

with dense, fine hairs. First antenna (Figure 92c) 

7-segmented, no ornamentation on first segment, remaining 

segments with short, naked setae as indicated 

in the figure. Second antenna (Figure 92d) a simple 

claw with 2 short, stout setae on inner edge of claw. 

Mouth tube (Figure 92e) without surface ornamentation. 

Mandible with 6-7 teeth. First maxilla (Figure 

92/) bearing 3 setae at tip. Second maxilla (Figure 

92g) similar to that of P. armatus except no patch 

of spinules on basal segment. Maxilliped (Figure 

93a) with a short, heavily sclerotized claw (Figure 

93b), basal segment with a thumblike process at inner 

proximal corner as in P. armatus. 

Legs 1-3 rudimentary. Leg 1 (Figure 93c) rami 

1-segmented; exopod with 5 short terminal spines; 

endopod bearing a short terminal spine; both rami 

covered with dense hairs and difficult to study. Leg 2 

(Figure 93c) rami 1-segmented; basipod with a welldeveloped 

seta at outer distal corner; exopod bearing 

4 terminal spines; endopod with 1 easily visible spine, 

2 very short spines, and spinules as in the figure. Leg 3 

(Figure 93c) reduced to a short knob bearing a short 

spine. Leg 4 represented by a lateral seta on anterior 


Egg string long. 


1.88 mm. Greatest width 0.55 mm. Thoracic segments 

bearing legs 2 and 3 incompletely separated. Genital 

segment about twice as long as wide (1.04 X 0.53 

mm) ; abdomen (Figure 93e) nearly twice as wide as 

long (259 X 147 /xm) with a ventral patch of spinules 

near each outer distal corner. Caudal rami (see Figure 

93e) about twice as long as wide (123 X 64 fim), 

bearing 2 prominent lateral setae, 2 small lateral setae 

and a terminal spine, ventral surface covered with 

spinules. First antenna as in female. Second antenna 

(Figure 93/) similar to female except claw longer and 

more pointed. Mouth tube (Figure 94a) with 3 

patches of spinules on lab rum. First maxilla (Figure 

94a) with 2 stout setae at tip, setae relatively longer 

than in female. Second maxilla as in S. armatus except 

no spinules on basal segment. Maxilliped (Figure 94fc) 

with 2-segmented claw and spinules on proximal inner 

corner of base. 

Leg 1 (Figure 94c) biramous, each ramus 1-segmented 

and not distinctly articulated with basipod; 

basipod with a patch of spinules and a triangular 

process and a seta at outer distal corner; exopod with 

patches of spinules and spinulose lobes in distal third, 

2 short spines at tip; endopod modified as a curved, 

heavily sclerotized process, lobed at tip and most of 

its length enclosed in a membrane (compare with 

P. armatus). Leg 2 similar to P. buccata except 

exopod inner 3 spines smaller and distal half with 

widely spaced spinules; endopod with 2 distinct 

patches of spinules. Leg 3 (Figure 94d) reduced to a 

spinulose process with a short spine at tip. 

REMARKS.—Yamaguti (1963:172) removed this 

species from Pseudocycnus in which it had been placed 

by Wilson. He designated the new genus Pseudocynopsis 

for this species and another new genus Pseudocycnoides 

to include P. armatus and also P. 

scomberomori (Yamaguti), both originally described 

as species of Pseudocycnus. We agree with Yamaguti 

that these 3 species should be removed from Pseudocycnus 

but we do not feel that placing P. buccata in 

a genus separate from P. armatus and P. scomberomori 

is justified. Consequently we have placed this 

species in the genus Pseudocycnoides. The original 

description of P. buccata was superficial and inaccurate 

and was undoubtedly the basis for Yamaguti's 

conclusions. Our studies of these 3 species indicate 

them to be much more closely related than previously 

known. These conclusions are based on the comparative 

morphology, not only of the female, but first 

descriptions of males of P. buccata and P. armatus as 

well. It should be pointed out that the first pair of 

legs in all 3 of these species is very small, between 

the bases of the maxillipeds, and not easily seen. 

This species is found on all American species of 

Scomberomorus (western Atlantic and eastern Pacific) 

and is relatively common on all its hosts. 

Examination of the type-specimen of Cybicola 

elongata Pearse (USNM 88538) leaves no doubt that 

it is the same as Wilson's species. The host for Pearse's 

material was Scomberomorus maculatus. 

The genus Pseudocycnopsis Yamaguti should be 

considered a synonym of Pseudocycnoides Yamaguti. 

Lernanthropus Blainville, 1822 

Lernanthropus kanagurta Tripathi, 1962 

FIGURE 94«-g 

Lernanthropus kanagurta Tripathi, 1962:194. 

MATERIAL EXAMINED.—1 collection containing 1 9 

from the gills of Rastrelliger brachysoma from Borneo.

NUMBER 311 45 

FEMALE.—Body form as in Figure 94e. Total length 

3.23 mm. Greatest width 1.35 mm. Cephalon about 

one-fourth total body length (not including posterior 

processes) and only about one-half greatest width. The 

single specimen was somewhat mutilated so we were 

not able to examine all of the appendages. The form 

of the second antenna and leg 1 are as in Figures 94/ 

and 94g. 

REMARKS.—When Tripathi described this copepod 

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) 

or some other host is preferred. 

Brachiella Cuvier, 1830 

Brachiella thynni Cuvier, 1830 

FIGURES 95a, 108 

Brachiella thynni Cuvier, 1830:257. 

Thynnicola ziegleri Miculicich, 1904:47. 


114 9 from the axil of the pectoral fin of the following 

hosts and localities: Acanthocybium solandri from 

Malagasy Republic, He Amsterdam, Seychelles, Palau 

Island, Yap Island, Kapingamarangi Atoll; Raroia 

Island, Line Islands, California, Nicaragua (Atlantic), 

Campeche, Brazil (north coast), Surinam, Azores; 

Thunnus albacares from Christmas Island (Pacific), 

Juan Fernandez Islands, off New Jersey, Azores; 

Thunnus obesus from Seychelles, Christmas Island 

(Pacific), Hawaii, mid-North Atlantic (39°N, 44°W), 

Azores; Thunnus thynnus from east coast of United 

States, Portugal; Scomberomorus regalis from Leeward 

Islands. 

This species has been reported many times from 

scombrids and other hosts. It is usually attached to its 

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 

a description here. 

Brachiella magna Kabata, 1968 

FIGURES 95b, 108 

Brachiella magna Kabata, 1968b;508, fig. 2a-j 


from the gills and nasal lamellae of Scomberomorus 

commerson from Australia (New South Wales) and 

S. sinensis (gills) from Hong Kong. 

The original description was based on a collection 

from the gills of S. commerson from off Queensland, 

Australia. Considering the number of specimens of the 

2 hosts examined (121) it is apparently uncommon on 

and known only from west Pacific Scomberomorus. 

Kabata has provided a good description with illustrations. 

Clavellisa Wilson, 1915 

Clavellisa scombri (Kurz, 1877) 

FIGURES 95C, 108 

Anchorella scombri Kurz, 1877:403. 

Clavella scombri.—Brian, 1906:116. 

Clavellisa scombri.—Wilson, 1915:694. 


17 9 from the following hosts and localities: Scomber 

japonicus from Philippines, Peru, Gulf of Mexico 

(Campeche and Alabama), Sierra Leone, Liberia; 

Scomber australasicus from Taiwan, Australia. 

This species was originally described from material 

collected from the gills of Scomber scombrus from 

Trieste. Our material was collected from the other 2 

species of Scomber. Since we examined 97 specimens 

of the type host without recovering this species we 

wonder if perhaps the earlier hosts were misidentified. 

Whatever the case, it seems apparent that this species 

is restricted to species of Scomber and is found 

wherever its hosts are. Later records by Yamaguti 

(1939) and Shiino (1959c) cite Scomber japonicus as 

the host species. Yamaguti (1939) redescribed this 

species and provided good illustrations. He inadvertently 

omitted this species in his 1963 synoptic survey 

of fish copepods. 

Clavellopsis Wilson, 1915 

Clavellopsis saba Yamaguti, 1939 

FIGURES 95d, 108 

Clavellopsis saba Yamaguti, 1939:558. 

This species was described from Japan from 

Scomber japonicus and reported again by Shiino 

(1959c: 367) from the same host from the Sea of

46 

Japan (Tsunodayama). During our studies we examined 

11 specimens of S. japonicus from Japan but 

did not recover this copepod. 

Shiino (1959c) has provided a good description 

with illustrations. 

Pennella Oken, 1816 

Pennella species 

Members of this genus have been reported many 

times from large marine pelagic animals. It is easily 

recognized attached to the body of the host; the head 

buried in the muscle tissue and the long wormlike 

body trailing free. Specimens have been recorded 

several inches long. 

This genus is in need of revision and species names 

should be used with caution. At least 6 species have 

been reported from whales, 7 from flyingfishes, and 4 

from swordfish. A revision would undoubtedly reduce 

these numbers of Pennella species. 

We have collected Pennella from Acanthocybium 

solandri and Thunnus thynnus. Because of the confusion 

surrounding the taxonomy of this genus we 

have not assigned species names to our specimens. 

Summary 

Our data, relative to infestation rates, are subject 

to considerable bias because of variables in collecting 

methods throughout the course of the study. Much of 

the material was collecetd from preserved fish in 

museum collections. In these cases copepods usually 

found on the body surfaces of the hosts are lost. Consequently, 

rate of infestation data for each species, 

based on data from the entire collection, is biased. 

Nevertheless, we have included infestation rates in 

spite of this, as we feel that they do reflect, in a general 

way, host preferences and relative common-torare 

occurrences of the parasite species on their hosts. 

Selected collections will be considered in more detail 

in the joint work with Collette. 

The following is a list of scombrid fishes with the 

parasitic copepods collected from them during the 

course of this study. The number in parentheses after 

the host name indicates the number of individuals examined. 

The number in parentheses after the copepod 

name indicates the number of fish infested. The 


copepods are listed with each host in descending order 

of frequency. 

SCOMBRINI 

Rastrelliger kanagurta (108) 

Pumiliopes capitulates (11) 

Nothobomolochus kanagurta (4) 

Rastrelliger brachysoma (33) 

Lernanthropus kanagurta (1) 

Rastrelliger faughni (14) 

Pumiliopes capitulatus (2) 

Nothobomolochus kanagurta (2) 

Orbitacolax aculeatus (2) 

Scomber scombrus (97) 

Caligus pelamydis (1) 

Scomber japonicus (500) 


Clavellisa scombri (9) 


Caligus mutabilis (1) 

Clavelopsis saba 

Scomber australasicus (17) 

Clavellisa scombri (1) 


SCOMBEROMORINI 

Acanthocybium solandri (61) 

Brachiella thynni (37) 

Gloiopotes hygomianus (26) 

Caligus productus (9) 

Caligus coryphaenae (1) 

Shiinoa occlusa (1) 

Pennella species (1) 

Grammatorcynus bicarinatus (37) 


Caligus asymmetricus (4) 

Scomberomorus brasiliensis (62) 

Pseudocycnoides buccata (39) 

Holobomolochus divaricatus (14) 


Shiinoa inauris (3) 

Scomberomorus cavalla (36) 


Holobomolochus asperatus (10) 



Scomberomorus commerson (113) 

Pseudocycnoides armatus (20) 

Unicolax ciliatus (20) 

Shiinoa occlusa (11)

NUMBER 3 1 1 47 

Caligus biseriodentatus (11) 

Caligus cybii (10) 

Caligus infestans (5) 


Brachiella magna (2) 

Tuxophorus cervicornis (2) 

Scomberomorus concolor (47) 

Pseudocycnoid.es buccata (14) 

Holobomolochus nudiusculus (13) 

Caligus omissus (7) 

Scomberomorus guttatus (55) 





Scomberomorus koreanus (17) 



Scomberomorus lineolatus (10) 




Scomberomorus maculatus (76) 





Scomberomorus multiradiatus (4) 


Scomberomorus niphonius (18) 

Pseudocycnoides scomberomori (6) 




Scomberomorous plurilineatus (3) 


Scomberomorus regalis (38) 





Caligus bonito (1) 


Tuxophorus collettei (1) 

Scomberomorus semifasciatus (17) 




Scomberomorus sierra (116) 


Caligus omissus (39) 

Holobomolochus nudiusculus (28) 

Scomberomorus sinensis (8) 


Brachiella magna (1) 

Scomberomorus tritor (21) 




Caligus diaphanus (1) 

Scomberomorus queenslandicus (19) 





Scomberomorus species (3) 




Sardasarda (104) 


Ceratacolax euthynni (21) 



Sar


Auxis species (65) 

Unicolax collaterals (19) 

Unicolax mycterobius (9) 





THUNNINI 

Euthynnus affinis (71) 

Unicolax collateralis (29) 


Caligus re gaits (5) 

Pseudocycnus appendiculatus (4) 





Euthynnus alletteratus (64) 








Unicolax anonymous (2) 


Euthynnus lineatus (15) 





Katsuwonus pelamis (132) 




Unicolax reductus (3) 


Thunnus alalunga (12) 

Elytrophora brachyptera (7) 




Thunnus albacares (110) 







Thunnus atlanticus (76) 




Thunnus maccoyii (2) 


Thunnus obesus (40) 




Elytrophora indica (9) 


Thunnus thynnus (57) 




Pennella species (3) 




Thunnus tonggol (26) 


Below is a list of the copepods and the fishes on 

which they were found. The number in parentheses 

after the host name indicates the infestation rate 

(number of infested fish/number of fish examined). 

We remind the reader of the bias in the infestation 

rates discussed above. No number after a fish name 

indicates the record is from the literature. 

BOMOLOCHIDAE 

Holobomolochus divaricatus 




Holobomolochus nudiusculus 



Holobomolochus asperatus 


Unicolax collateralis 


^4uxu species (29) 


Sarda orientalis (21) 

Orcynopsis unicolor (17) 

Cybiosarda elegans (13) 


Unicolax anonymous 


Unicolax mycterobius 



Euthynnus affinis (4)

NUMBER 311 49 

Unicolax ciliatus 

Scomberomorus gut talus (24) 






Unicolax reductus 


Ceratacolax euthynni 

Sarda sarda (20) 


Nothobomolochus kanagurta 



Rastrelliger species 

Orbitacolax aculeatus 

Rastrelliger faughni 

Pumiliopes capitulatus 





SHIINOIDAE 

Shiinoa inauris 




Shiinoa occlusa 


Gymnosarda unicolor (14) 







CALIGIDAE 

Caligus coryphaenae 









Caligus regalis 


Caligus productus 






Sarda orient alls (13) 







Sarda chiliensis lineolatus (2) 



Scomberomorus species 

Caligus asymmetricus 

Cybiosarda elegans (25) 



Sarda australis (10) 


^4uxu species (2) 




Caligus bonito 


Sarda chiliensis chiliensis (54) 




Gymnosarda unicolor (14) 





Caligus mutabilis 





Caligus omissus 



Caligus biseriodentatus 





species 

Caligus cybii 



Scomberomorus species (20)




Caligus pelamydis 










Caligus infestans 


Caligus diaphanus* 


Caligus savala* 

Euthynnus affinis 

Caligus macarovi* 

Auxis species 

Caligus amblygenitalis* 


Caligus pseudokalumai 

Gymnosarda unicolor 

Elytrophora brachyptera 

Allothunnus fallal (100) 

Thunnus maccoyii (100) 






Elytrophora indica 


EURYPHORIDAE 

Gloiopotes hygomianus 


Caligulus longispinosus 


TUXOPHORIDAE 

Tuxophorus cybii 

Scomberomorus commerson 

Acanthocybium solandri 

Tuxophorus cervicornis 


Tuxophorus collettei 


PSEUDOCYCNIDAE 

Pseudocycnus appendiculatus 

Thunnus tonggol (27) 






Pseudocycnoides armatus 

Scomberomorus plurilineatus (100) 







Pseudocycnoides scomberomori 


Pseudocycnoides buccata 







LERNANTHROPIDAE 

Lernanthropus kanagurta* 

Rastrelliger brachysoma (3) 

LERNEOPODIDAE 

Brachiella thynni 






Brachiella magna 



Clavellisa scombri 

Scomber australasicus (6) 


Clavellopsis saba 

Scomber japonicus 

* Probably not usually scombrid parasites.

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1896. Notes on the Parasitic Copepoda of Fish Obtained 

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8-16. 

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Bombay. Annals and Magazine of Natural History, 

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1955. Uber eine neue Art der Parasitischen Copepoden: 

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Institute, University of Kerala, Trivandrum, 

8:87-130. 

1962a. On a New Species of Bomolochus (Copepoda) 

with Remarks on Orbitacolax Shen. The Journal 

of Parasitology, 48(4):610-612. 

1962b. Observations on the Synonymy of Caligus 

coryphaenae Stp-Lutk. Annals and Magazine of 

Natural History, series 5, volume 13:513-522. 

1963. Copepods Parasitic on South Indian Fishes— 

Family Caligidae. Journal of the Marine Biological 

Association of India, 5(1): 68—96. 

1965. Copepods Parasitic on South Indian Fishes: Family 

Bomolochidae, 3. Journal of the Bombay Natural 

History Society, 62 (1): 38-55. 

1971. Notes on Some Copepod Parasites in the Collection 

of the British Museum (N.H.), London. 

Journal of the Marine Biological Association of 

India, 11:149-174. 

Scott, T. 

1894. Report on Entomostraca from the Gulf of Guinea, 

Collected by John Rattray, B.Sc. Transactions of 

the Linnean Society, London, 6(2): 1-161. 

1901. Notes on Some Parasites of Fishes. Report of the 

Fishery Board for Scotland, 19(3) : 120-153. 

Shen, C. 

1957. Parasitic Copepods from Fishes of China, Part II: 

Caligoida, Caligidae (1). Ada Zoologica Sinica, 

9(4):351-377. 

Shiino, S. M. 

1952. Copepods Parasitic on Japanese Fishes, 1: On the 

Species of Caligus and Lepeophtheirus. Report 

of the Faculty of Fisheries, Prefectural University 

of Mie, 1:79-113. 

1954. A New Fish-Louse Found on the Mackerel-Pike. 

Annotationes Zoological Japonenses, 27(3):150- 

153. 

1956. Copepods Parasitic on Japanese Fishes, 12: Family 

Lernaeopodidae. Report of the Faculty of Fisheries, 

Prefectural University of Mie, 2(2):269-311.

NUMBER 3 1 1 53 

1958. Parasitic Copepods from Fishes Collected in the 

Indian Ocean. Annual Report of the Prefectural 

University of Mie, section 2 (Natural Science), 

2(3):98-113. 

1959a. Revision der auf Goldmakrele, Coryphaena hippurus 

L. schmarotzenden Caligidenarten. Annual 

Report of the Prefectural University of Mie, section 

2 (Natural Science), 3(1): 1-34. 

1959b. Ostpazifische Parasitierende Copepoden. Report 

of the Faculty of Fisheries, Prefectural University 

of Mie, 3(2):267-333. 

1959c. Sammlung der parasitischen Copepoden in der 

Prafturuniversitat von Mie. Report of the Faculty 

of Fisheries, Prefectural University of Mie, 3(2): 

334-374. 

1960. Parasitic Copepods of Fishes from the Eastern 

Pacific. Report of the Faculty of Fisheries, Prefectural 

University of Mie, 3(3) :527-541. 

1965. Parasitic Copepods of the Eastern Pacific Fishes, 

5: Caligus. Report of the Faculty of Fisheries, 

Prefectural University of Mie, 5:391-420. 

Silas, E. G., and A.N.P. Ummerkutty. 

1967. Parasites of Scombroid Fishes, Part II: Parasitic 

Copepoda. Symposium on Scombroid Fishes, 

3:876-993. 

Steenstrup, J., and C. Liitken. 

1861. Bidrag til Kundskab om det aabne Havs Synltekrebs 

og Lernaeer samt om nogle andre nye eller hiltil 

kun ufuldstaen digt kjendte parasitiske Copepoder, 

5:341-432. 

Tripathi, Y. R. 

1962. Parasitic Copepods from Indian Fishes, III: 

Family Anthosomatidae and Dichelesthiidae. Pro- 

ceedings of the First All-India Zoological Congress, 

2:191-217. 

Vervoort, W. 

1965. Three New Species of Bomolochidae (Copepoda, 

Cyclopoida) from Tropical Atlantic Tunnies. 

Zoologische Verhandelingen, 76:3—40. 

Wilson, C. B. 

1905. North American Parasitic Copepods Belonging to 

the Family Caligidae, 1: The Caliginae. Proceedings 

of the United States National Museum, 

28:479-672. 


the Lernaeopodidae, with a Revision of the Entire 

Family. Proceedings of the United States National 

Museum, 47(2063): 565-729. 

1921. New Species and a New Genus of Parasitic 

Copepods. Proceedings of the United States National 

Museum, 59(2354): 1-17. 


the Family Dichelesthiidae. Proceedings of the 

United States National Museum, 60(2400): 1-100. 

1935. New Parasitic Copepods. Smithsonian Miscellaneous 

Collections, 91 (19): 1-9. 

1937. Parasitic Copepods Taken During the Third Hancock 

Expedition to the Galapagos Islands. Allan 

Hancock Pacific Expedition, 2(4): 23-30. 

Yamaguti, S. 

1936. Parasitic Copepods from Fishes of Japan, part 2: 

Caligoida, I. 22 pages, [private publication.] 

1939. Parasitic Copepods from Fishes of Japan, part 5: 

Caligoida, III. Volumen Jubilare Pro Prof. Sadao 

Yoshida 3:443-487. 

1963. Parasitic Copepoda and Branchiura of Fishes. 

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.


FIGURE 3.—Holobomolochus divaricatus, new species, female: a, leg 3; b, leg 4; c, leg 5; male: 

d, dorsal.

NUMBER 3 1 1 57 

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. 

E

58 


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 


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.


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 


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 


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

NUMBER 3 1 1 67 

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. 

E


B 

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

NUMBER 3 1 1 69 

FIGURE 16.—Unicolax anonymous (Vervoort), new genus, female: a, leg 3; b, leg 4; c, leg 5; 

male: d, dorsal; e, leg 5. 

E


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

B

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.


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 


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 


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.


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.


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.


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. 

D

NUMBER 311 83 

B 

D 

FIGURE 30.—Ceratacolax euthynni Vervoort, male: a, second antenna; b, maxilliped; c, leg 1; 

d, leg 2.


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

s

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 


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

NUMBER 311 87 

FIGURE 34.—Orbitacolax aculeatus (Pillai), female: a, dorsal; b, first antenna; c, second antenna; 

d, oral area; e, maxilliped; /, leg 1. 

D

88 


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. 

D 

E

NUMBER 3 1 1 89 

B 

FIGURE 36.—Pumiliopes capitulatus Cressey and Boyle, female: a, first antenna; b, second antenna 

; c, oral area; d, maxilliped; e, leg 1. 

D


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

d, leg 5. 

B 

D

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 


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 

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. 

E 

B 

93

94 


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 

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. 

B

96 


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 

FIGURE 44.—Caligus productus Muller, female: a, leg 3; b, leg 4; male: c, dorsal; ,d, genital 

segment and abdomen; e, second antenna. 

E 

B

98 


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 

D 

FIGURE 46.—Caligus asymmetricus Kabata, female: a, second maxilla; b, maxilliped; c, sternal 

furca; d, leg 1; e, leg 2. 

E 

99

100 


FIGURE 47.—Caligus asymmetricus Kabata, female: a, leg 3; b, leg 4; male: c, dorsal; d, genital 

segment and abdomen, ventral; e, second antenna.

NUMBER 311 

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. 

101

102 


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 

FIGURE 50.—Caligus bonito Wilson, female: a, leg 3; b, leg 4; male: c, dorsal; d, second 

antenna; e, distal portion of maxilliped. 

D

104 


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. 

B 

D

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 

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 


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

B

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.


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 

E 

B 


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.


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 


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 

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

115

116 

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 

FIGURE 64.—Caligus pelamydis Kroyer, female: a, dorsal; b, genital segment and abdomen, 

ventral; e, oral area; d, first antenna; e, second antenna. 

B


FIGURE 65.—Caligns pelamydis Kroyer, female: a, second maxilla; b, maxilliped; c, sternal 

furca; d, leg 1; e, leg 2. 

E

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 


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 

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. 

F

122 


FIGURE 69.—Caligus infestans Heller, female: a, leg 3; b, leg 4; male: c, dorsal; d, genital 

segment and abdomen, ventral; e, second antenna. 

B

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 


FIGURE 71.—Caligns diaphanus Nordmann, female: a, second maxilla; b, maxilliped; c, sternal 

furca; d, oral area; e, leg 1; /, leg 2.


FIGURE 72.—Caligus diaphanus Nordmann, female: a, leg 3; b, leg 4; male: c, dorsal; d, second 

antenna; e, postoral process; /, maxilliped. 

B

126 


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 amblygenitalis 

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.)


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, maxilliped.

NUMBER 3 1 1 129 

FIGURE 76.—Elytrophora brachyptera Gerstaecker, female: a, sternal furca; b, leg 1; c, leg 2; 

d, leg 3; e, leg 3 endopod. 

D

130 


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 

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. 

E

132 


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, maxilliped; 

d, sternal furca; e, leg 1; /, leg 2 (female after Heegaard, 1962).


D 

B 

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 

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 


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 

FIGURE 86.—Pseudocycnus appendiculatus Heller, female: a, maxilliped; b, leg 1; c, leg 2; 

d, leg 3. 

D


D 

FIGURE 87.—Pseudocycnus appendiculatus Heller, male: a, dorsal; b, abdomen and caudal 

rami, ventral; c, first antenna; d, maxilliped; e, leg 1; /, leg 4. 

B

NUMBER 3 1 1 141 

FIGURE 88.—Pseudocycnoides armatus (Bassett-Smith), female: a, dorsal; b, first antenna; 

c, first and second antennae; d, mouth tube; e, second maxilla. 

D


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 


FIGURE 91.—Pseudocycnoides armatus (Bassett-Smith), male: a, leg 2; b, leg 3. Pseudocycnoides 

scomberomori (Yamaguti), female: c, dorsal; d, maxilliped; e, leg 1; /, leg 2; 

g. ^g 3.


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.


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 

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. 

E

148 


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.


• 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). 

FIGURE 101.—Distribution of Shiino inauris Cressey and S. occlusa Kabata. 

o inauns 

• occlusa


• 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 


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 


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 


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).


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 


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).


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)-


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 segment 

(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 


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 ) ; «, cephalon, 

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).


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).


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).


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 )•


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).


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).


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 


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 

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)- 

173


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).


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 ).


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 ).


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 ) •


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)-


*—— 

p^ - ' £ 

1 

• 

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). 

D

NUMBER 311 

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 ). 

181


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).


FIGURE 136.—Pseudocycnus appendiculatus Heller, male: a, base of leg 5 (5000X)- Pseudocycnoides 

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 ).


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 )•


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 

FIGURE 139.—Pseudocycnoid.es buccata (Wilson), female: 

a, tip of second maxilla (1225X); b, same (1450x); c, 

same (4400X). 

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