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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 311

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

Roger Cressey and Hillary Boyle Cressey

SMITHSONIAN INSTITUTION PRESS

City of Washington

1980

ABSTRACT

Cressey, Roger, and Hillary Boyle Cressey. Parasitic Copepods of Mackerel- and Tuna-like Fishes (Scombridae] of the World. Smithsonian Contributions to Zoology, number 311, 186 pages, 139 figures, 1980.—Forty-six species of parasitic copepods (one new genus, 8 new species) are reported from all known 46 species of Scombridae subfamily Scombrinae from the world. Literature records are included only where host and/or copepod identifications could be verified. Copepod families include: Bomolochidae; Shiinoidae; Caligidae; Euryphoridae; Tuxophoridae; Pseudocycnidae; Lernanthropidae; Lerneopodidae. The new bomolochid genus is Unicolax. The new species are U. collateralis, 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

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

Roger Cressey and Hillary Boyle Cressey

___________

Roger Cressey and Hillary Boyle Cressey, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

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 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 type-specimens 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 Guinea being described by Collette and Russo. The following list is adapted from information in Collette and Chao (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): circumtropical and subtropical

Grammatorcynus 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 (Lacépède): Indian and Western Pacific oceans

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 Pacific oceans

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 (Lacépède): Northwestern Pacific Ocean

Scomberomorus tritor (Cuvier): Gulf of Guinea to Mediterranean

Scomberomorus species: Northern Australia to New Guinea

Tribe SARDINI Jordan and Evermann (Bonitos)

Allothunnus fallai Serventy: circumglobal in southern hemisphere

Cybiosarda elegans (Whitley): Northern Australia to New Guinea

Gymnosarda unicolor (Rüppell): 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 Lacépède: 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 oceans

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, III.

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)

Muséum 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 collateralis, 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, 5a–c, 96, 109–113

MATERIAL EXAMINED. — Holotype art (USNM 172244), allotype art (USNM 172245) paratypes 11 art (USNM 172246) from the nasal sinuses of Scomberomorus brasiliensis (USNM 188424) from Brazil. In addition, there are 13 collections containing 61 art 2 art from S. brasiliensis from Panama (Atlantic), Colombia (Atlantic), Brazil and Argentina; 25 collections containing 70 art 40 art from S. maculatus from Cape Cod, New Jersey, Maryland, South Carolina, Georgia, Florida, Mississippi, Alabama, Louisiana, Texas, Panama (Atlantic), Surinam; 11 collections containing 28 art 20 art 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 1a Total length 2.62 mm, greatest width (widest part of cephalon) 1.20 mm. Genital segment (Figure 1b) wider than long (348 × 483 μm) with 3 dorsal setae at area of egg sac attachment. Abdomen 3-segmented, segments measuring (1 × w) 224 × 318 μm, 153 × 265 μ, 147 × 230 μm respectively; ventral surface of last segment with 2 patches of prominent spinules (see figure 1c). Caudal rami (Figure 1c) longer than wide (129 × 94 μ) with six setae; longest seta 590 μ

First antenna (Figure 1d) 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 corner, 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 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 ventro-lateral 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 two-thirds, 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 1b) 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 × 182 μm). Abdomen (Figure 4a) 2-segmented, segments measure (length × width) 88 × 118 μm, 82 × 106 μm, respectively; ventral surface of second segment with 2–3 rows of prominent spinules forming a proximal transverse band and 2 distal patches of spinules. Caudal rami (Figure 4b) slightly longer than wide (41 × 35 μm) with 6 setae; longest seta 572 μm.

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 4e) 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 5b) 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, 7a–b, 96

MATERIAL EXAMINED.—Holotype art (USNM 172247), allotype art (USNM 172248), paratypes 33 art 1 art (USNM 172249) from the nasal sinuses of Scomberomorus sierra from Buenaventura, Colombia (USNM 218565). In addition 13 collections containing 30 art 1 art from the nasal sinuses of S. concolor from Baja California and Sonora, Mexico; 26 collections containing 105 art 7 art from the nasal sinuses of S. sierra from Baja California, Sonora, Panama (Pacific), Colombia (Pacific), Peru.

FEMALE.—Body form as in Figure 5d. Total length 1.76 mm, greatest width 0.72 mm, Genital segment (see Figure 5e) wider than long (230 × 271 μm) with 3 naked setae at area of egg sac attachment. Abdomen (Figure 5e) 3-segmented segments measuring 171 × 206 μm, 118 × 177 μm, and 112 × 147 μm respectively, ventral surface of last segment with 2 patches of minute spinules (see Figure 6a). Caudal rami (Figure 6a) slightly longer than wide (59 × 53 μm) with 6 setae (longest 652 μm).

First antenna, second antenna, mandible, paragnath, first maxilla, second maxilla, and maxilliped similar to those of Holobomolochus divaricatus.

First leg similar to that of H. divaricatus. Legs 2–4 biramous, rami 3-segmented. Leg 2 (Figure 6b) similar to that of H. divaricatus except endopod third segment with row of blunt spinules at base, 2 outer spinulose spines, gradually tapering, with no terminal flagellum. Leg 3 (Figure 6c) coxopod, basipod, exopod similar to those of H. divaricatus except exopod spines relatively longer with distal spine longest; endopod similar to leg 2 except third segment with 2 rather than 3 setae and inner spine slightly longer than outer. Leg 4 similar to that of H. divaricatus. Leg 5 (Figure 6d) generally similar to that of H. divaricatus except free segment not as elongate, lateral and terminal spines relatively longer. Leg 6 represented by 3 setae at area of egg sac attachment.

MALE.—Body form as in Figure 6e. Total length 0.98 mm, greatest width 0.38 mm. Genital segment (Figure 7a) longer than wide (194 × 159 μm). Abdomen (see Figure 7a) 2-segmented, segments measuring 53 × 94 μm and 64 × 88 μm respectively, ventral surface of last segment with 2–3 irregular transverse rows of spinules near anterior edge and a small patch of spinules near insertion of each caudal ramus (see Figure 7b). Caudal rami (Figure 7b) longer than wide (47 × 29 μm), each with small ventral patch of slender spinules and 6 setae (longest 531 μm).

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.

REMARKS.—Females of this species can be distinguished 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 7c–g, 8, 96

MATERIAL EXAMINED.—Holotype art (USNM 172242) and paratypes 2 art (USNM 172243) from the nasal sinus of Scomberomorus cavalla (MCZ 17182) from Cuba. In addition, 9 collections containing 10 art 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 × 265 μm). Abdomen 3-segmented, segments measure (1 × w) 177 × 206 μm, 129 × 177 μm, and 112 × 159 μm 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 × 53 μm), each ramus with six setae and a patch of hairlike spinules on its ventral surface; longest seta 768 μm.

First antenna (Figure 7f) 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 8b) 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 8c) similar to that of H. divaricatus except exopod spines comparatively longer and more slender; endopod third segment outer spines slender and without lateral spinules. Leg 4 (Figure 8d) coxopod with row of stout spinules on outer distal corner; basipod with dorsal seta; exopod 3-segmented, first segment with slender outer spine with small terminal flagellum, row of small spinules at base of spine, second segment with inner seta and outer spine similar to that of first segment, row of small spinules at base of spine, third segment with 6 inner to terminal plumose setae and 2 outer spines similar to but slightly longer than those of first and second segments, a few minute spinules at base of proximal spines; endopod 3-segmented, first segment with inner seta, spinulose along distal half, second segment with inner seta similar to that of first segment, outer distal border with short row of hairs, outer edges of first and second segments haired, third segment slender, elongate with 2 spinulose spines flanking terminal seta, spines of about equal length, seta spinulose along distal half, short row of elongate spinules near base of each spine. Leg 5 (Figure 8e) similar to that of H. divaricatus except free segment not as elongate, spinules shorter. Leg 6 (see Figure 7d) represented by row of 3 setae on genital segment, inner 2 setae short, outer seta about 3 times longer.

MALE.—Unknown.

ETYMOLOGY.—The Latin asperatus (“rough with points or short, stiff hairs”) refers to the patches of hairs on the abdomen and caudal rami.

REMARKS.—Females of this species can be separated from H. divaricatus and H. nudiusculus by the presence of a patch of hairs on the ventral surface of each caudal ramus of H. asperatus (no ornamentation 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 cavalla 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.

Unicolax, 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.

Unicolax collateralis, new species

FIGURES 9–13, 97, 114–116a

MATERIAL EXAMINED.—Holotype art (USNM 172256), allotype art (USNM 172257) paratypes 12 art 23 art (USNM 172258) from the nasal sinus of Euthynnus alletteratus (USNM 203838) from St. George Bay, Lebanon. The following collections from Auxis species: 2 art from Woods Hole, Mass.; 2 art from St. George Bay, Lebanon; 2 art 3 art from Ghardaqa, Egypt; 2 collections containing 7 art 3 art from Hong Kong; 2 collections containing 2 art 1 art from Japan; 7 collections containing 12 art 4 art from the Philippines; 2 art from Chusan, China; 1 art from Hawaii. One collection containing 5 art from Cybiosarda elegans from Brisbane, Australia. The following collections from Euthynnus affinis: 2 art 2 art from Elat, Israel; 2 art from Mozambique; 2 art from Madagascar; 1 art from the Seychelles; 4 art from the Arabian Sea 24°N, 67°E; 2 art from Java; 16 collections containing 29 art 9 art from the Gulf of Thailand; 2 art from Formosa; 2 art from Hong Kong; 4 collections containing 8 art 1 art from the Philippines; 1 art from Palau; 1 art from Okinawa; 1 art from Tokyo; 4 art 7 art from Brisbane, Australia. The following collections from Euthynnus alletteratus: 1 art from the Caribbean (9°11′N, 77°50′W); 2 art from Brazil; 4 collections containing 29 art 16 art from St. George Bay, Lebanon. The following collections from Euthynnus lineatus: 1 art from Galapagos; 1 art from Lower California; 1 art from Mexico (Pacific); 2 art from Costa Rica (Pacific). One collection containing 1 art from Orcynopsis unicolor from St. George Bay, Lebanon. Two collections containing 5 art 3 art from Sarda australis from New South Wales, Australia. Two collections from Sarda orientalis containing 3 art from Pearl Islands, Panama, and Durban, South Africa. All collections from nasal sinuses of hosts.

FEMALE.—Body form as in Figure 9a. Total length 1.55 mm, greatest width 0.91 mm; length of cephalon 0.30 mm. Genital segment (Figure 9b) wider than long (200 × 312 μm). Abdomen (see Figure 9b) 3-segmented, segments measuring (1 × w) 153 × 206 μm, 100 × 171 μm, 94 × 141 μm respectively. First 2 segments without ornamentation, last segment with 2 large ventral patches of fine hairs (see Figure 9c). Caudal rami (Figure 9c) longer than wide (88 × 53 μm), each ramus with ventral patch of fine hairs and 6 setae as indicated in Figure 9c; longest seta 413 μm.

First antenna (Figure 9d) 7-segmented. Fourth seta on first segment modified to form a heavily sclerotized spine. Ornamentation of segments as follows. Segment 1: 3 plumose setae; 1 spine; 1 plumose seta. Segment 2: 4 antero-dorsal naked setae; 7 antero-ventral plumose setae; 3 ventral plumose setae; 2 ventral naked setae. Segment 3: 1 dorsal naked seta; 3 antero-ventral plumose setae; 1 ventral naked seta. Segment 4: no ornamentation. Segment 5: 1 dorsal naked seta; 1 antero-ventral naked seta; 2 ventral naked setae. Segment 6: 2 terminal ventral naked setae; 1 aesthete. Segment 7: 6 terminal naked setae; 1 subterminal naked seta; 1 aesthete. Second antenna (Figure 9e) 3-segmented; second segment with one plumose seta and one short, naked seta; third segment armed with several rows of short spinules, and with 4 hooklike terminal spines and 3 setae. Mouthparts as in Figure 10a. Labrum with 2 large patches of hairs, hairs longer along anterior edge. Mandible blade slender, accessory blade smaller. Paragnath with distal and lateral hairs; also distal edge with fine teeth. First maxilla with 3 long, plumose setae and one short, naked seta. Second maxilla with 2 blades, each with uniform spinules along edges. Maxilliped (Figure 10b) with 4 plumose setae; hook slightly curved, without accessory process.

Legs 1–4 biramous, each ramus 3-segmented except leg 1 exopod. Leg 1 (Figure 10c) coxopod with patch of stout hairs on outer corner and broad inner seta; basipod with outer seta, row of hairs above insertion of endopod, and inner blunt process, exopod first segment with stout outer spine terminating in short flagellum and second segment incompletely divided, with 2 short, outer flagellate spines, and 6 terminal to inner setae, endopod first segment with inner seta and short row of spinules (hairs?) along outer, lower edge, second segment similar to first, third segment with 5 terminal setae, outer edges of endopod segments heavily haired, all setae heavily plumose. Leg 2 (Figure 10d) coxopod with patch of slender spinules on outer corner, row of short hairs along mid lower edge, basipod with outer dorsal seta and triangular patch of minute spinules on midventral surface; exopod first segment with long, slender spinules on outer edge, serrate spine with terminal flagellum on outer distal corner, inner edge with small patch of hairs, second segment with inner seta and outer spine similar to that on first segment, third segment with 5 inner to terminal setae and 4 terminal to outer spines, proximal spine with short hairs on both edges and terminal flagellum, next 2 spines similar to those on first and second segments, distal spine elongate with broad outer serrations and short inner hairs; all exopod setae long, slender and plumose; endopod first segment with inner seta and row of fine hairs along distal edge, second segment enlarged with 2 inner setae and row of fine hairs on outer half of distal edge, third segment narrower than second with 3 inner to terminal setae and 2 short terminal spines with short lateral hairs and terminal flagella; endopod setae stout basally and heavily plumose; outer edges of endopod segments haired. Leg 3 (Figure 11a) coxopod with row of short spinules on outer distal corner, small central patch of blunt spinules and row of short hairs on mid-lower edge; basipod with outer dorsal seta; exopod first segment with outer distal spine, dorsolateral surface of segment with several 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, 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 11b) 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 11c) 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 11d. Total length 1.09 mm, greatest width 0.44 mm; length of cephalon 0.20 mm. Genital segment (Figure 11e) longer than wide (236 × 218 μm). Abdomen (see Figure 11e) 2-segmented, segments measuring 94 × 112 μm and 70 × 88 μm 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 × 35 μm); each ramus with ventral patch of fine hairs and 6 setae; longest seta 295 μm.

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 12d) coxopod with inner seta and spinules as indicated in Figure 12d; 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 12e) 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 13b) 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 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 collateralis (“standing side by side”) alludes to its occurrence in collections with U. mycterobius.

REMARKS.—Females of this species can be distinguished 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, 116b–f, 117

Parabomolochus anonymous Vervoort, 1965:3.

MATERIAL EXAMINED.—Two collections containing 31 art 2 art 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 × 247 μm). Abdomen 3-segmented, segments measure 82 × 153 μm, 53 × 141 μm, 70 × 123 μm length × width respectively. First 2 segments without ornamentation, last segment with 2 patches of fine hairs ventrally. Caudal rami each with patch of fine hairs and 6 setae; longest 171 μm.

First antenna (Figure 14b) 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 14d. 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 15b) 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 16b) 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.

MALE.—Body form as in Figure 16d. Total length 0.81 mm, greatest width 0.36 mm; length of cephalon 0.20 mm. Genital segment as long as wide (177 × 177 μm). Abdomen 2-segmented, segments measuring (length × width) 35 × 94 μm and 59 × 70 μm. Caudal rami slightly longer than wide (35 × 29 μm); longest seta 177 μm.

All appendages and ornamentation similar to (although proportionately smaller than) U. collateralis except leg 5.

Leg 5 (Figure 16e) similar to U. collateralis except patch of spinules on free segment covers lower two-thirds of inner half of segment and does not extend around the entire distal edge; also, spinules on both basal and free segments are 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 distinguished 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–c

Parabomolochus mycterobius Vervoort, 1965:11.

MATERIAL EXAMINED.—The following collections from Auxis species: 1 art 2 art from Massachusetts; 1 art from Ghardaqa, Egypt; 2 art from Formosa; 3 collections containing 4 art 1 art from Tokyo; 5 art from Luzon, Phillipines; 1 art 3 art from New South Wales; 1 art from Hawaii. The following collections from Euthynnus alletteratus: 1 art from Pensacola, Florida; 4 collections containing 5 art 5 art from St. George Bay, Lebanon. The following collections from E. affinis: 2 collections containing 2 art 2 art from Tokyo, Japan; 1 art from Kagoshima, Japan. All copepods collected from the nasal sinus of host.

Female.—Body form as in Figure 17a. Total length 2.55 mm greatest width 1.45 mm; length of cephalon 0.90 mm. Genital segment wider than long 241 × 454 μm. Abdomen 3-segmented, segments measuring 177 × 318 μm, 135 × 265 μm, 171 × 212 μm respectively. Caudal rami longer than wide 147 × 64 μm, longest seta 501 μm. Last abdominal segment and caudal rami with ornamentation similar to Unicolax collateralis.

Cephalic appendages and leg 1 similar to Unicolax collateralis.

Leg 2 (Figure 17b) coxopod with inner seta and row of fine spinules on outer distal corner; basipod with outer seta and 2 small patches of fine hairs; exopod first segment with large outer patch of fine hairs and spine on outer distal corner, second segment with inner seta and outer spine, third segment with 5 inner to terminal setae and 4 terminal to outer spines, exopod spines squat with short hairs along margins and terminal flagellum except distal spine on third segment elongate and tapered with fine hairs on outer edge, widely spaced hairs on inner edge; endopod first segment with stout inner seta, outer distal portion of segment with patch of short hairs, second segment enlarged with 2 stout inner setae and patch of short hairs on outer distal edge, third segment narrower than second, with 3 stout inner to terminal setae and 2 terminal spines similar to those on exopod, outer edges of endopod segments heavily haired. Leg 3 (Figure 17c) coxopod and basipod similar to leg 2; exopod first segment with patch of short hairs on mid-outer margin and spine on outer distal corner, second segment with inner seta and outer spine, third segment with 5 inner to terminal setae and 3 terminal to outer spines, distal exopod spine elongate and tapered, similar to corresponding spine of leg 2, other exopod spines slightly more elongate than corresponding spines of leg 2; endopod first segment with inner seta and patch of short hairs on outer distal corner, second segment similar to first, third segment with 2 inner to terminal setae, (inner seta plumose, terminal seta with short hairs of uniform length along outer margin) and 2 outer spines, outer edges of endopod segments heavily haired. Leg 4 (Figure 18a) coxopod with minute spinules along mid-distal margin; basipod with outer seta; exopod first segment with minute spinules covering most of dorsal surface, extending to small mid-ventral patch, spine on outer distal corner, 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 exopod spines of leg 3; endopod first segment with outer distal patch of short hairs and inner seta with short bristles along distal two-thirds, second segment with distal patch of short hairs and inner seta similar to that of first segment, third segment with distal patch of short hairs, and terminal seta flanked by 2 spines, seta with bristles along distal half, spines with bristled margins and terminal flagellum, outer edges of endopod segments haired. Leg 5 (Figure 18b) 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 1.36 mm, greatest width 0.52 mm; length of cephalon 0.35 mm. Genital segment longer than wide (336 × 289 μm). Abdomen 2-segmented, segments measuring (length × width) 100 × 129 μm and 70 × 106 μm respectively; second segment (see Figure 18d) ornamented ventrally with anterior, irregular row of minute hairs and distally with 2 patches of minute hairs. Caudal rami (Figure 18d) longer than wide (70 × 41 μm); each ramus with ventral patch of hairs and 6 setae; longest seta 383 μm.

Cephalic appendages similar to those of U. collateralis male.

Legs 1–4 biramous. Leg 1 (Figure 18e) 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 18f); 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 18f; 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 2–4. 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 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

MATERIAL EXAMINED.—Holotype art (USNM 172253) allotype art (USNM 172254) and 1 art 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 art from N.W. Coast of Madagascar; 2 art , 1 art from Travancore, India; 6 art from Cochin, India; 4 art from Batavia, Java; 2 art 1 art from Phuket, Thailand; 1 art from Hong Kong; 5 collections containing 23 art 1 art from the Philippines. The following collections from S. guttatus: 1 art from Calicut, India; 1 art from Padang, Sumatra; 2 art from Batavia, Java; 1 art from Singapore; 1 art from Phuket, Thailand; 2 collections containing 7 art from Hong Kong; 3 collections containing 13 art from China. The following collections from S. niphonius: 1 art from Korea; 1 art from China. The following collection from S. semifasciatus: 5 art 1 art from New Guinea. The following collections from S. tritor: 2 collections containing 4 art from Liberia; 8 art from Ghana; 2 art from Lagos, Nigeria.

FEMALE.—Body form as in Figure 20a. Total length 2.30 mm, greatest width 1.01 mm, length of cephalon 0.60 mm. Genital segment wider than long (265 × 330 μm). Abdomen 3-segmented, segments measuring 165 × 212 μm, 118 × 171 μm, 147 × 159 μm respectively. Caudal rami longer than wide (141 × 59 μm), longest seta 590 μm. Ventral side of last abdominal segment and caudal rami with patches of hairs similar to Unicolax collateralis.

First antenna (Figure 20b) 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 mid-lower edge; basipod with outer dorsal seta and row of short spinules along outer distal edges; exopod (Figure 20d) 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 21b) 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 21d. Total length 1.23 mm, greatest width 0.46 mm; length of cephalon 0.29 mm. Genital segment longer than wide (265 × 177 μm). Abdomen 2-segmented, segments measuring (length × width) 88 × 88 μm, and 88 × 70 μm, ventral surface of second segment (see Figure 21e) 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 21e) longer than wide (59 × 29 μm) each ramus with 6 setae and ventral patch of hairs; longest seta 413 μm.

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 21f). 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.

REMARKS.—Females of this species can be distinguished 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 21e).

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.

Unicolax reductus, new species

FIGURES 23–26, 99, 119d–f, 120–122a,b

MATERIAL EXAMINED. — Holotype art (USNM 172259), allotype art (USNM 172260), paratypes 15 art (USNM 172261) from Katsuwonus pelamis (USNM 176974) nasal sinus from New South Wales. Additional material from the same host: 24 art 2 art from Tahiti and 25 art art from Japan. All copepods collected from the nasal sinuses of the hosts.

FEMALE.—Body form as in Figure 23a. Total length 2.94 mm, greatest width 1.50 mm; length of cephalon 0.94 mm. Genital segment (Figure 23b) wider than long (289 × 501 μm). Abdomen (see Figure 23b) 3-segmented, segments measuring 206 × 247 μm, 188 × 230 μm, 230 × 200 μm respectively; segments without ornamentation. Caudal rami (Figure 23c) longer than wide (206 × 88 μm) each ramus bearing 6 setae and no other ornamentation, longest seta 401 μm.

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 subterminal naked seta; one terminal aesthete. Second antenna (Figure 23e) similar to U. collateralis and ciliatus. Labrum similar to U. collateralis. Remaining mouthparts as in Figure 23f. 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 24b) 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 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 flagellum. Leg 6 (see Figure 23b) represented by 3 short setae, of about equal length, on genital segment.

MALE.—Body form as in Figure 25c. Total length 1.78 mm, greatest width 0.78 mm; length of cephalon 0.39 mm. Genital segment (Figure 25d) slightly wider than long (277 × 300 μm), abdomen 2-segmented, segments measuring (length × width) 159 × 177 μm, and 194 × 147 μm; second segment with two ventral rows of minute spinules (see Figure 25e). Caudal rami (Figure 25e) longer than wide (141 × 73 μm) with no ornamentation other than 6 setae (longest 371 μm).

First antenna similar to that of U. collateralis male. Remaining cephalic appendages (except maxilliped) 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 26b) 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 evthynni Vervoort, 1965

FIGURES 27–31, 100, 122c–f, 123–125

Ceratacolax euthynni Vervoort, 1965:26.

MATERIAL EXAMINED.—34 collections containing 94 art and 63 art from the following hosts and localities: 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 28b) 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, 30d, 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), new combination

FIGURES 32, 33, 100

Bomolochus kanagurta Pillai, 1965:51.

MATERIAL EXAMINED.—5 collections containing 9 art from the gills of Rastrelliger kanagurta from China, India (Madras), Red Sea; R. faughni from the Philippines.

FEMALE.—Body form as in Figure 32a. Total length 1.50 mm. Abdomen 3-segmented, segments measure 118 × 206 μm, 95 × 195 μm, 95 × 177 μm, length by width respectively. Last abdominal segment (Figure 32b) with 2 oblique patches of spinules. Caudal rami (see Figure 32b) about twice as long as wide (94 × 55 μm); 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 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 33b) 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), new combination

FIGURES 34, 35a–b, 100

Bomolochus aculeatus Pillai, 1962a:610.

MATERIAL EXAMINED.—2 collections containing 15 art from the nasal sinuses of 2 Rastrelliger faughni from Manila and Lingayan Gulf, Philippines.

FEMALE.—Body form as in Figure 34a. Total length 1.71 mm. Greatest width 0.73 mm. Rostrum protruding from between bases of first antennae and bearing a pair of ventral hooks (see Figure 34b).

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 34b) 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 34e) 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 34f). 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,b) 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–e

Pumiliopes capitulatus Cressey and Boyle, 1973:1.

MATERIAL EXAMINED.—23 collections containing 35 art from the orbits of the following hosts and localities: Rastrelliger kanagurta the Red Sea, Sri Lanka, India (Madras), western Indian Ocean, Philippines, Java; R. faughni from Philippines; Scomber japonicus from Gulf of Guinea, Mauritania, Zanzibar; S. australasicus from Philippines.

FEMALE.—Body form as in Figure 35c. Total length 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 35d) 3-segmented; segments measure 71, 38, and 65 pm long respectively; each of last 2 segments narrower than preceeding one; first 2 segments each with transverse posteroventral band of fine spinules, last segment with 2 ventral patches of scales and row of fine spinules on each outer posterior corner. Caudal rami (Figure 35e) nearly 3 times as long as wide. Each ramus with patch of 25–30 scales on ventral surface and 6 setae, longest with stout base.

First antenna (Figure 36a) 5-segmented; basal part (first 2 segments) 289 μm long, remaining segments 35, 18, and 24μm long respectively, no aesthetes on either of last 2 segments. Rostrum without hooks. Second antenna (Figure 36b) 3-segmented; basal segment with seta on outer distal corner; second segment with short, blunt medial seta; last segment subdivided, outer part with row of 14–16 prominent recurved hooklets and short terminal seta, inner part armed with stout claw arising near midlength, 3 stout setae constricted in their posterior third, and 2 additional setae (one very short). Labrum without ornamentation. Mouthparts as in Figure 36e. Labium represented by row of spinules posterior to mouth. Maxilliped (Figure 36d) with short, heavily sclerotized, nearly straight hook, without accessory process.

Legs 1–4 biramous, rami 2-segmented except for leg 4 endopod (3-segmented). We will omit a detailed description of legs 1–4 except to provide illustrations of each and to note that in the present material the exopods bear only 1 setule on outer edge rather than 2 or 3 as reported in the original description. Leg 5 (Figure 37d) with patches of scales, a short seta near midlength of outer edge and 3 terminal setae (outermost shortest and middle longest). Leg 6 represented by 3 prominent setae on midlateral border of genital segment.

Egg sacs flattened (probably due to their position on the host between the orbit and adipose eyelid).

MALE.—Unknown.

REMARKS.—We originally described this species based on 2 specimens collected from the orbit of Clupanodon punctatus from Hong Kong. The specimens reported here vary somewhat from the original description. The most obvious difference is the greater number of scalelike spinules in the patches on the caudal rami and abdomen of the scombrid parasites. Also, the scombrid copepods have only 1 setule (inappropriately referred to as setae in our original description) on the last exopod segments of legs 2 and 3.

Considering the relative numbers of clupeid and scombrid hosts examined during the 2 studies, we found this species to be more common on the scombrid hosts; for that reason we consider them to be the preferred hosts. Since clupeids are a primary food of these scombrids, it is understandable that they would occassionally be found on clupeids because of the ecological relationship.

Shiinoa Kabata, 1968

Shiinoa inauris Cressey, 1975

FIGURES 38, 39a–d, 101

Shiinoa inauris Cressey, 1975:211.

MATERIAL EXAMINED.—14 collections containing 22 art 15 art from the nasal lamellae of Scomberomorus regalis from Cuba, Puerto Rico, Surinam, and Venezuela; Scomberomorus maculatus from Massachusetts, Florida, Texas; Scomberomorus brasiliensis from Surinam, Brazil.

FEMALE.—Body form cylindrical (Figure 38a). Total length 3.56 mm (3.45–3.67 mm) based on an average of 3 specimens. Greatest width 0.71 mm. Rostrum produced anteriorly, curved ventrally with distal half covering terminal portion of second antennae (Figure 38b). These 2 elements form a loop by which the female attaches to the nasal lamella through a hole in the lamella. Body segmentation indistinct. Posterior outer corners of genital segment produced to form short, dorsally directed processes; tips of processes covered with knobs or bumps. Abdomen 4-segmented. Caudal rami about 3 times longer than wide, bearing 6 short setae.

First and second antennae separated from mouth area. First antenna (Figure 38b) with 5–6 segments, each bearing short naked setae. Second antenna well-developed. Each antenna forming a ring and parallel to each other; distal third within the curvature of the rostrum. Mandible, first maxilla, and second maxilla as in Figure 38c. Maxilliped absent.

Leg 1 (Figure 38d) 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 corner 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 Kabata, 1968

FIGURES 39e-h, 101, 126d-f, 127–128a,b

Shiinoa occlusa Kabata, 1968a:497.

MATERIAL EXAMINED.—21 collections containing 41 art 11 art 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 Müller, 1785

Caligus coryphaenae Steenstrup and Lütken, 1861:352.

FIGURES 40, 41 a–b, 102; 128c–f, 129, 130

Caligus coryphaenae Steenstrup and Lütken, 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.

MATERIAL EXAMINED.—138 collections containing 316art and 281 art from the body surface and gills of the following hosts and localities: Acanthocybium 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 × 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 40b.

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 40d) 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 40f) 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 μ distalmost to proximalmost respectively.

MALE.—Body form as in Figure 41b. 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 genital segment.

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. regalis.

Caligus regalis Leigh-Sharpe, 1930

FIGURES 41c–g, 102

Caligus regalis Leigh-Sharpe, 1930:5.

Caligus euthynus Kurian, 1961:63.

Caligus alveolares Heegaard, 1962:156.

MATERIAL EXAMINED.—5 collections containing 149 art 113 art from the body surface of Euthynnus affinis collected off Nosy Bé, Malagasy Republic. Types of C. alveolaris (AMS P. 16408) from the same host species, reported as E. allitteratus [sic] from north Queensland, Australia.

FEMALE.—Body form as in Figure 41c. Total length 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 × 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 μm wide, rostral space between each lunule about 950 μm. Cephalic appendages as in C. coryphaenae except sternal furca not as divergent in C. regalis (Figure 41d).

Legs 1–4 as in C. coryphaenae; the stout spine on the first exopod segment of leg 3 (Figure 41e) is curved inwards rather than straight as in C. coryphaenae. The apical seta on the exopod of leg 4 (Figure 41f) is only slightly longer than the adjacent seta, setae measure 189, 177, 177, 177, and 183 μm distal to proximal respectively.

MALE.—Body form as C. coryphaenae. Total length 3.51 mm (3.37–3.53 mm) based on an average of 5 specimens. Greatest width 1.73 mm (measured at widest part of cephalon). Genital segment (Figure 41g) wider than long (914 × 711 μm). 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 2-segmented, segments measure 247 and 479 μm 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).

MATERIAL EXAMINED.—207 collections containing 2286 art , 657 art from the mouth, gills, and body surface of the following hosts and localities: Acanthyocybium solandri from the northeast Malagasy Republic, Seychelles, Line Islands, Campeche Bay; Scomberomorus cavalla from east coast United States (several localities); Scomberomorus regalis from Dominican Republic, Puerto Rico, Virgin Islands; Scomberomorus species from Australia (east coast); Scomberomorus tritor from Nigeria; Sarda sarda from Massachusetts; Sarda orientalis from Panama (Pacific); Sarda chiliensis lineolatus from Baja California; Auxis species from Panama (Pacific), Mexico (Pacific); Euthynnus affinis from Hawaii; Euthynnus alletteratus from Bermuda, Anagada Passage (W. Indies); Katsuwonus pelamis from northwest Malagasy Republic, Tahiti, Line Islands, Suitcases Bank (E. Pacific), Peru, Ecuador, Gulf of Mexico (Alabama), New Jersey, Dominican Republic, Venezuela, Brazil, Gulf of Guinea; Thunnus alalunga from North Carolina; Thunnus albacares from northwest Malagasy 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 42b); 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 44e) with short claw and 2 prominent bossed areas on basal segment. Genital segment and abdomen as in Figure 44d.

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. Certainly 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 pelamis 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 71 art and 25 art 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 from northwest Malagasy Republic; Thunnus albacares from Queensland.

FEMALE.—Body form as in Figure 45a. Range of 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.

MATERIAL EXAMINED.—76 collections containing 155 art and 58 art 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 48a. 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.

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 50e) 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.

MATERIAL EXAMINED.—8 collections containing 8 art 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.

FEMALE.—Body form as in Figure 51a. Total length 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 μm). Genital segment (Figure 51b) longer than wide (986 × 725 μm), widest posteriorly, posterior corners rounded and only slightly produced. Abdomen (see Figure 51b) incompletely divided into 2 segments, about twice as long as wide (681 × 362 μm), posterior ventral surface with patches of spinules as in Figure 51c. Caudal rami (Figure 51c) longer than wide (159 × 106 μm) bearing 6 plumose setae as indicated in figure.

Oral area as in Figure 52c. First antenna (Figure 51d) first segment with 27 setae (all but 2 are plumose), last segment with 13 naked setae. Second antenna (Figure 51f) 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 52e) 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 53b 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 μm 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 μm) whereas in our material this seta is much longer than the adjacent seta (177 in 112 μm). 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

MATERIAL EXAMINED.—58 collections containing 147 art 41 art 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 art 3 art paratypes (USNM 172241) from the gills of S. sierra from San Juan Lagoon, Mexico (USNM 219623).

FEMALE.—Body form as in Figure 53c. Total length 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 μm) slightly more than least distance (218 μm) between lunules (interlunular space). Genital segment (Figure 53d) slightly longer than wide (1.16 × 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 × 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 53e. Caudal rami (see Figure 53e) longer than wide (142 × 94 μm) armed with 6 plumose setae as indicated in figure, longest seta 307 μm.

Oral area as in Figure 54a. First antenna (Figure 54b) first segment with 27 setae, all plumose except 2 in middle anterior group; last segment with 13 naked setae. Second antenna (Figure 54d) 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 54f) 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 55b) 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 conspicuous 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 μm 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 56b) longer than wide (884 × 580 μm). Abdomen (Figure 56b) 2-segmented, first segment nearly square (348 × 348 μm), second segment longer than wide (449 × 334 μm) 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 56f) 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 56b.

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 (= S. 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 non-scombrid 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 biseriodentattis 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 art 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 art and art 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 localities), Hong Kong; Scomberomorus queenslandicus from Onslow (W. Australia), Ambon (Moluccas); Scomberomorus plurilineatus from Zanzibar.

FEMALE.—Body form as in Figure 57a. Total length 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 × 0.75 mm). Abdomen about twice as long as wide (0.68 × 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 μm) greater than diameter of lunule (207 μm). First antenna of usual type of genus except last segment about 4 times longer than wide (177 × 44 μm). 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 57e) 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 57e). Leg 3 similar to other species of genus, heavily sclerotized spine on exopod first segment (Figure 58a) recurved inwardly. Leg 4 (Figure 58b) 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 μm 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 μm) less than least space (interlunular space) between them (265 μ). 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 58e) base with 2 sclerotized processes on inner margin (not shown on Shen's figure), claw with a short sclerotized process near base of seta near inner midmargin.

Leg 1 basipod covered with conspicuous spinules; coxopod with row of spinules on posterior margin; exopod (Figure 58g) with 3 fringed spines and a short plumose seta at tip, inner lateral margin with 3 short setae; endopod lacking. Leg 2 (Figure 59a) endopod first segment with a group 4–5 spinules on outer distal corner, second segment with a double row of 8–9 stout spinules on outer edge, last segment without spinules on proximal outer corner. Leg 3 exopod first segment spine only slightly recurved inwards. Leg 4 (Figure 59b) exopod 2-segmented, distalmost spine slightly longer than adjacent spine, all spines relatively short.

IMMATURE MALE.—Body form as in Figure 59c. Total length 3.60 mm (3.50–3.80 mm) based on an average of 5 specimens. As in female except as follows. Second antenna with short claw, basal segment with 2 rugose areas, no posterior process. Postantennal process prominent, recurved with inner angle less than 45°. Maxilliped (Figure 59f) very large, basal segment with heavily trilobed process on inner margin opposite tip of claw. Legs 1–4 as in female. Caudal ramus with ventral patch of spinules; spinules also present on abdomen at outer distal corner (see Figure 59d).

REMARKS.—The description of this species was based on immature specimens. When we examined the adult art specimen of Caligus auxisi sent to us by Dr. Pillai we were struck by a number of similarities 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 art 1 art), 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 (1 art) 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.

MATERIAL EXAMINED.—19 collections containing 158 art and 59 art 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.

FEMALE.—Body form as in Figure 60a. Total length 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 60b) trapizoidal (widest posteriorly), somewhat wider than long (1.6 × 1.8 mm), comprising about 20 percent total length, posterior corners not produced. Abdomen (see Figure 60b) incompletely divided into 2 segments, widest anteriorly and tapering to caudal rami, about 2½ times longer than wide (2.03 × 0.79 mm) comprising about 25 percent of total length. Caudal rami (Figure 60c) small, somewhat longer than wide (236 × 200 μm) armed with plumose setae as in the figure.

Oral area as in Figure 60d. First antenna (Figure 60e) 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 61d) of usual caligoid type with 2 terminal, fringed, flagella. Maxilliped (Figure 61e) a stout claw. Sternal furca (Figure 61f) 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 62b) 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 μm 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 62d. Total length 3.24 mm (3.00–3.45 mm) based on an average of 5 specimens. Genital segment (Figure 63a) somewhat longer than wide (841 × 739 μm). Abdomen 2-segmented, somewhat longer than wide (580 × 450 μm), second segment about 2½ times length of first. Caudal rami as in female.

Cephalic appendages as in female except second antenna (Figure 63b) 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.

MATERIAL EXAMINED.—30 collections containing 97 art and 4 art 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 alletteratus (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.

FEMALE.—Body form as in Figure 64a. Range of total length 3.00–4.65 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 65e) 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 66b) 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 66e) with rugose patch on basal half; maxilliped of male (Figure 66f) with sclerotized process on basal segment opposing tip of claw; fifth and sixth legs represented by 3 setae at mid-margin and 2 setae at posterior corner of genital segment respectively; abdomen 2-segmented with first segment about half as long as second (Figure 66d).

REMARKS.—We have recorded this species here from 9 species of scombrids from all 4 tribes. Although it is found on several species of hosts it seems to prefer species of Sarda (approximately 13 percent infestation of specimens examined). Although this species has wide distribution it has not been recorded from the Indian Ocean. We consider Parapetalus species of Silas and Ummerkutty to be C. pelamydis based on their illustrations.

Caligus infestans Heller, 1868

FIGURES 67–69, 104

Caligus infestans Heller, 1868:167.

Caligus sphyraenae Nunes-Ruivo and Fourmanoir, 1956:71.

Caligus maculatus Heegaard, 1962:157.

MATERIAL EXAMINED.—5 collections containing 7 art and 1 art from the gills and body surface of 5 Scomberomorus commerson from Indonesia, Thailand, and Mozambique.

FEMALE.—Body form as in Figure 67a. Total length 5.14 mm (based on an average of 4 specimens ranging 4.80–5.55 mm). Greatest width 2.18 mm (1.95–2.40 mm). Cephalon comprises nearly half total length. Genital segment somewhat longer than wide (1.8 × 1.4 mm) with posterior distal corners lobed and extending to about middle of abdomen. Abdomen 2-segmented, longer than wide (0.77 × 0.48 mm). Caudal rami (Figure 67c) about as long as wide (271 × 248 μm) with outer edge about twice length of inner edge, one small, subterminal seta at inner distal corner and 4 terminal plumose setae (outermost much shorter than others). Pillai (1971:160) states that there is a ventral patch of fine denticles on the caudal ramus (anal laminae). These are actually on the dorsal surface rather than ventral and appear as sclerotized bumps in the outer proximal quarter.

Oral area as in Figure 67d. First antenna (Figure 67e) typically caligoid but posterior 6 setae of last segment with tips feathered giving them a frayed appearance. Second antenna (Figure 68a) with usual hookshaped last segment, basal segment with posterior, pointed, sclerotized process. Tip of second maxilla (Figure 68b) with usual 2 fringed processes but tips of processes somewhat expanded. Maxilliped (Figure 68c) with stout terminal hook and sclerotized process on basal segment opposite tip of hook. Sternal furca (Figure 68d) with widely divergent tines, tips of tines pointed and somewhat recurved inwards.

Leg 1 (Figure 68e) basipod with large patch of spinules; exopod first segment with short spine on outer distal corner and row of long spinules along inner edge, last segment armed as in other species of the genus but 3 inner setae with plumosities stouter than usual from bases to tips; endopod weakly developed. Leg 2 (Figure 68f) exopod, first segment 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 corner, 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 69b) exopod first segment outer edge produced as approximate right angle at distal two-thirds; last segment with distalmost spine 2½ times length of others (442 × 177 μm). Leg 5 represented by 3 short plumose setae near edge of genital segment at distal three-fourths.

MALE.—Body form as in Figure 69c. Total length 4.95 mm. Greatest width 2.10 mm. Genital segment about as long as wide. Abdomen (Figure 69d) 2-segmented, first segment shorter than second (second about 3 times length of first). Appendages as in female except second antenna (Figure 69e) with 2 rugose areas on basal segment, a short, recurved, terminal claw, a seta near base of terminal claw.

REMARKS.—This species is easily separated from the other species of Caligus reported from scombrids by the peculiar shape of the first segment of the exopod of leg 4.

Heller described this species from Scomber species from Java. According to B. Collette (pers. comm.) the specimens reported by Heller as Scomber species from Java must have been Rastrelliger species since no true Scomber were taken from Java during the Novara cruise that was the basis of his report. Since then it has been reported by Bassett-Smith 1898b: 360; Nunes-Ruivo and Fourmanoir 1956:69; Heegaard 1962:157; Kirtisinghe 1964:52; Kabata 1965:119; and herein from Scomberomorus commerson and S. queenslandicus from the Indo-West Pacific. Kabata 1965:119 also reported a single male from Euthynnus affinis (as E. alletteratus). It seems apparent that this species is primarily a parasite on the body surface of Scomberomorus commerson. It is probably more common than our records indicate as most of our collections from this host were from preserved specimens and body surface parasites were no longer present.

Caligus diaphanus Nordmann, 1832

FIGURES 70–72

Caligus diaphanus Nordmann, 1832:26.

Caligus caudatus Kroyer, 1838:16.

Caligus isonyx Steenstrup and Lütken, 1861:354.

MATERIAL EXAMINED.—4 art 1 art from the gills of a specimen of Scomberomorus tritor collected from the mouth of the Belas River, Ghana.

FEMALE.—Body form as in Figure 70a. Total length 3.48 mm, greatest width (measured at widest part of cephalon) 1.59 mm. Genital segment (Figure 70b) somewhat wider than long (1.09 × 1.31 mm), globose, comprising about one-third of the total body length. Abdomen (see Figure 70b) 2-segmented, segments measure 0.55 × 0.48 mm and 0.29 × 0.35 mm (length × width) respectively. A patch of fine spinules at the outer distal corners of the last abdominal segment. Caudal rami (Figure 70c) longer than wide (153 × 106 μm) with 6 plumose setae, terminal 3 nearly equal in length (longest 224 μm).

First antenna (Figure 70d) first segment with 27 setae, most with fine plumosities, last segment with 14 naked setae, segments measure 118 μm and 100 μm respectively along posterior margins. Second antenna (Figure 70f) and stout recurved claw. Mandible within mouth tube, tip with 12 teeth. First maxilla (Figure 70g) reduced to 3 setae near base of postoral process. Second maxilla (Figure 71a) with 2 fringed subequal processes at tip, a hyaline process near tip of last segment. Maxilliped (Figure 71b) in form of a stout uncinate claw. Sternal furca (Figure 71c) with nearly parallel tines, tines rounded at tip and somewhat recurved. Ventral cephalic area as in Figure 71d.

First leg (Figure 71e) basipod with plumose seta at outer distal corner and a shorter plumose seta near inner mid-margin; exopod first segment with a short spine on outer distal corner and a row of spinules along inner margin, second segment with 4 outer to terminal spines and 3 inner plumose setae. Second leg (Figure 71f) basipod with a short plumose seta on outer distal corner; exopod first and second segments each with a stout fringed spine on outer distal 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 μm) on outer distal corner, second segment with fringed seta (136 μm) on outer distal corner, last segment with 3 fringed setae (153, 165, and 206 μm long respectively), a fringe at base of each seta. Fifth leg represented by 4 setae near the posterior distal corner of the genital segment.

MALE.—Body form as in Figure 72c. Total length 2.39 mm, greatest width 1.30 mm. Genital segment about as long as wide (0.43 × 0.45 mm). Abdomen 2-segmented segments measure 88 × 250 μm and 292 × 280 μm (length × 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 72e) with accessory process at distal two-thirds. Maxilliped (Figure 72f) 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 material collected by us from Trigla lineata from off Naples, Italy.

Caligus savala Gnanamuthu, 1948

FIGURE 73a–f

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 art from the body surface of Euthynnus 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 73b) with pointed posterior process on base. Postantennal process (Figure 73c) not strongly curved. Sternal furca (Figure 73d) 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 73f) 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, 74a–b

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 73i) 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 74b) 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).

Caligus amblygenitalis Pillai, 1961

FIGURE 74c–g

Caligus 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 2-segmented, bearing 4 spines (no lateral spine on last segment which is usually present in other species when exopod is 2-segmented), distalmost spine about one-third longer than adjacent spine.

Caligus pseudokalumai Lewis, 1968

Caligus 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.

MATERIAL EXAMINED.—78 collections containing 771 art 269 art from the gill area of the following hosts and localities: Thunnus alalunga from Ile 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 Ile Amsterdam, Seychelles, Christmas Island (Pacific), 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.

FEMALE.—Body form as in Figure 75a. Total length 8.26 mm, greatest width 4.71 mm (measured at widest part of cephalon). Genital segment longer than wide (2.61 × 1.89 mm). Abdomen 2-segmented, segments measure (l × w) 0.72 × 0.87 mm, 0.68 × 0.85 mm respectively; first segment partially hidden in dorsal view by flaps of genital segment. Caudal rami (Figure 75b) longer than wide (680 × 400 μ), 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 75f) 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 one-segmented 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 77e) longer than wide (1.74 × 1.53 mm). Abdomen 2-segmented, segments measure (l × w) 0.56 × 0.72 mm, 0.52 × 0.69 mm respectively. Caudal rami similar to female. First antenna 2-segmented, similar to female. Second antenna (Figure 77f) 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 Ile 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 77b 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 Ile 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.

MATERIAL EXAMINED.—9 collections containing 128 art 27art from the gill area of Thunnus obesus from Ile 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 78b) longer than wide (2.97 × 2.20 mm). Abdomen (see Figure 78b) first segment crescent-shaped, produced at posterior corners to envelope anterior half of second segment; segments measure (l × w) 1.01 × 1.50 mm, 1.02 × 1.10 mm respectively. Caudal rami longer than wide (0.65 × 0.43 mm) similar to E. brachyptera. First antenna (Figure 78c) 2-segmented, first segment with 25 setae, one modified into sclerotized spine; second segment with one subterminal seta, 13 terminal setae. Second antenna as in Figure 78d. Mouth tube, mandible, post oral process, and first maxilla as in Figure 78e; post oral process longer than mouthtube. Second maxilla, maxilliped, sternal furca, and legs 1–3 similar to E. brachyptera. Leg 4 (Figure 78f) similar to E. brachyptera except exopod spines relatively longer in E. indica. Legs 5 and 6 (Figure 78g) represented by 1 and 3 setae respectively on lateral genital segment.

MALE.—Body form as in Figure 79a. Total length 8.33 mm, greatest width 4.16 mm (measured at widest part of cephalon). Genital segment longer than wide (1.97 × 1.68 mm). Abdomen 2-segmented, segments measure 0.65 × 0.94 mm, 0.87 × 0.85 mm respectively. Caudal rami longer than wide (0.71 × 0.40), similar to female. First antenna similar to female. Second antenna (Figure 79b) similar to E. brachyptera male except basal segment with only small process. Mouth tube, mandible, post oral process, first maxilla, second maxilla similar to female. Maxilliped and sternal furca similar to E. brachyptera male. Legs 1–6 similar to female.

REMARKS.—So far this species has been collected only from Thunnus obesus and may well be restricted to that host. It occurs in collections with E. brachyptera but we have no data to indicate if these 2 species occupy the same microhabitat. We have included Shiino's records of this copepod on the map (Figure 105).

Gloiopotes Steenstrup and Lütken, 1861

Gloiopotes hygomianus Steenstrup and Lütken, 1861

FIGURE 79c–i

Gloiopotes hygomianus Steenstrup and Lütken, 1861:363.

MATERIAL EXAMINED.—27 collections containing 77 art 56 art from the body surface of Acanthocybium solandri from the following localities: Seychelles Islands, Malagasy Republic, Ile Amsterdam, Christmas Island (Pacific), Washington Island, Hawaii, Socorro Island, Revillagigedo, Puerto Rico, Nicaragua, North Atlantic (39°N, 41°W), North Carolina, Brazil, Canary Islands, Cape Verde Islands, Azores Islands.

Shiino (1960:533) provided a good description of this species and we will provide description and illustrations to enable the reader to recognize this species without resorting to earlier works. For a comprehensive account consult the work of Shiino (1960) and Cressey (1967).

FEMALE.—Body form as in Figure 79c. Total length 16.7 mm (15.0–17.7 mm.) greatest width 6.2 mm (6.0–6.5 mm). Measurements based on an average of 8 specimens. Caudal rami much longer than wide. Frontal lunules absent. First antenna 2-segmented (distal half of last segment sclerotized giving the impression of 2 segments). Second antenna clawlike, heavily sclerotized at tip. Sternal furca (Figure 79d) with accessory process lateral to the base of each tine. Leg 1 exopod last segment (Figure 79f) with 2 bifid spines plus an accessory spine giving trifid appearance; and unmodified spine on outer distal corner and 3 lateral setae. Leg 4 (Figure 79h) exopod 3-segmented, last segment with 3 short terminal spines.

MALE.—Body form as in Figure 79i. Total length 12.1 mm (11.7–12.5 mm). Greatest width 4.8 mm (4.2–5.7 mm). Measurements based on 6 specimens. Appendages similar to female except second antenna with an accessory process on claw.

REMARKS.—This species is restricted to Acanthocybium solandri and probably found throughout the range of the host. The 4 other species of the genus are parasites of billfishes (Istiophoridae). The genus Gloiopotes was revised by the first author (Cressey, 1967).

Caligulus Heegaard, 1962

Caligulus longispinosus Heegaard, 1962

FIGURE 80

Caligulus longispinosus Heegaard, 1962:171.

We did not collect this species but we have examined the type male deposited in the Australian Museum (AMS P. 16429). Heegaard stated that a female holotype and male allotype were deposited but only 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 80d), leg 1 (Figure 80e), 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 Fourmanoir, 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 81b–g

Tuxophorus cervicornis Heegaard, 1962:172.

MATERIAL EXAMINED.—2 collections containing 2 art from the body surface of Scomberomorus commerson from Coffs Harbor, New South Wales, Australia, and Karachi, Pakistan.

FEMALE.—Body form as in Figure 81b. Total length 6.80 mm. Greatest width 3.15 mm. Cephalon slightly longer than wide (3.10 × 3.15 mm). Thoracic segment bearing leg 4 with dorso-lateral winglike plates. Genital segment slightly wider than long (2.10 × 2.25 mm, including posterior processes); outer posterior corners produced. Abdomen 3-segmented. Caudal ramus about 6.5 times longer than wide (1.95 × 0.30 mm). Oral appendages as in T. collettei except sternal furca with pointed tines (Figure 81e) 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 EXAMINED.—Holotype art (USNM 172250), allotype (USNM 172251), and 13 art 1 art 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.

FEMALE.—Body form as in Figure 82a. Total length (including caudal rami) 9.0 mm, greatest width (measured at widest part of cephalon) 2.33 mm. Cephalon somewhat longer than wide (2.55 × 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 × 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 × 0.58 mm) narrowing near mid-margin and near distal end. Caudal rami about 3.8 times longer than wide (490 × 129 μm), widest proximally; 2 short setae on each lateral margin at distal two-thirds, 1 subterminal inner seta, 3 short, plumose, terminal setae.

First antenna (Figure 82d) 2-segmented; first segment with 13 short, stout, plumose setae, second segment with 11 naked setae. Second antenna (Figure 82e) 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 3-segmented, 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 μm 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.

MALE.—Body form as in Figure 84c. Total length (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 × 2.33 mm). General morphology of cephalon as in female. Genital segment (Figure 84d) longer than wide (1.3 × 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 × 0.73 mm), small spinules on ventral surface. Caudal rami as in female (590 × 141 μm). Cephalic appendages as in female except second antenna (Figure 84e) with large rugose areas on basal segments and maxilliped with 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.

MATERIAL EXAMINED.—43 collections containing 85 art 4 art 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 85e) with 2 stout and 1 weak setae at tip. Second maxilla (Figure 85f) 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 × 370 μm length by width). Abdomen (Figure 87b) longer than wide (300 × 220 μm). Caudal rami (see Figure 87b) somewhat longer than abdomen and about 4 times as long as wide (450 × 110 μm) 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 87e) 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 87f) 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.

Pseudocycnoides Yamaguti, 1963

Pseudocycnoides armatus (Bassett-Smith, 1898)

FIGURES 88–91a–b, 107, 136b–f, 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.

MATERIAL EXAMINED.—31 collections containing 149 art 11 art from the following hosts and localities: Scomberomorus commerson from Red Sea (Suez), Malagasy Republic, India (Cochin), Pakistan, Sri Lanka, Gulf of Thailand, Australia (NSW and Queensland), Philippines, Hong Kong; S. guttatus from Borneo (Padang), Hong Kong; S. semifasciatus from New Guinea; S. lineolatus from India (Palk Strait); S. koreanus from India (Bombay), Sumatra, China; S. queenslandicus from Papua New Guinea; S. plurilineatus from South Africa (Natal).

FEMALE.—Body form (Figure 88a) elongate with winglike process on each side of second free thoracic segment and anteriormost genital segment (fused with third thoracic segment). Total length 8.4 mm (7.8–8.7 mm based on 5 specimens). Greatest width 1.2 mm (1.05–1.35 mm based on 5 specimens). Genital segment comprising well over half total body length. Abdomen and caudal rami fused; each ramus tapering to a point. First antenna (Figure 88b) 7-segmented, first segment with no ornamentation, second segment with prominent sclerotized spine at anterodistal corner and other setae as in the figure; all segments with membranelike covering. Second antenna (Figure 88c) terminal segment clawlike with prominent medial spine and small proximal seta. Mouthtube (Figure 88d) tapering posteriorly; labrum with 4 patches of bumplike spinules; mandible with 7 teeth. First maxilla as in P. buccata Wilson. Second maxilla (Figure 88e) basal segment with large patch of spinules; second segment elongate, curved distally, together with short third segment forming a claw with rows of numerous stout spines along outer distal edge. Maxilliped (Figure 89a) basal segment enlarged, visible from dorsal aspect; terminal segment forming stout, heavily sclerotized claw, darkened and rugose at tip with small accessory spine on inner curve.

Leg 1 (Figure 89b) biramous, rami not distinctly separate from basipod; coxopod with small patch of slender spinules and triangular process on outer distal corner; exopod 1-segmented with 2 terminal spines; endopod 1-segmented with medial inner spine and 4 terminal spines, innermost shortest. Leg 2 (Figure 89c) basipod with patches of slender spinules and stout outer spine; rami 1-segmented, exopod with 4 terminal spines, outermost stout, toothed along outer edge; endopod with patches of slender spinules and 3 terminal spines. Leg 3 (Figure 89d) uniramous, ramus with slender spinules and 3 terminal spines, outermost spine stout, toothed along outer edge. Leg 4 represented by a simple seta on anterolateral genital segment. Egg strings usually as long as body.

MALE.—Body form as in Figure 8e. Total length 2.39 mm. Greatest width 0.59 mm. Genital segment more than half total body length. Abdomen (Figure 90a) without ornamentation. Caudal rami (see Figure 90a) with numerous spinules near tips and 5 setae. First antenna (Figure 90b), second antenna, mouthtube, mandible, and first maxilla as in female. Second maxilla (Figure 90c) similar to female, but spines on last 2 segments more numerous and more closely spaced. Maxilliped (Figure 90d) basal segment with 3 patches of small spinules and process on inner edge; distal 2 segments clawlike, each segment with 1 accessory spine.

Leg 1 (Figure 90e) 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 91b) 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 scomberomori (Yamaguti, 1939)

FIGURES 91c–g

Pseudocycnus scomberomori Yamaguti, 1939:457.

Pseudocycnoides scomberomori.—Yamaguti, 1963:172.

MATERIAL EXAMINED.—7 collections containing 22 art from the gills of Scomberomorus niphonius from Korea, North China, Japan.

FEMALE.—Body form as in Figure 91c. 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 91d) claw longer and tip more recurved. Leg 1 (Figure 91e) basipod with large patch of spinules (spinules on outer edge only in P. armatus). Leg 2 (Figure 91f) 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 91g) 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), new combination

FIGURES 92–94a–c, 107, 138, 139

Pseudocycnus buccata Wilson, 1922:79.

Cybicola elongata Pearse, 1951:365.

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

MATERIAL EXAMINED.—125 collections containing 667 art 10 art from the gill filaments of following hosts and localities: Scomberomorus sierra from Peru, Colombia (Pacific), Panama (Pacific), Costa Rica (Pacific), Gulf of California, Baja California; S. concolor from Gulf of California, California; S. regalis from Brazil (north coast), Surinam, Gulf of Venezuela, Virgin Islands, Haiti, Cuba, Florida (Atlantic); S. maculatus from Mexico (Vera Cruz), Florida (west coast), Alabama, United States east coast (several localities from Florida to Cape Cod); S. brasiliensis from Brazil (Rio de Janeiro to north coast), Trinidad, Panama (Atlantic), Costa Rica, British Honduras, Mexico (Mazatlán); S. cavalla from Brazil, Trinidad, Cuba, Gulf of Mexico, Cape Cod.

FEMALE.—Body from as in Figure 92a. Total length 5.14 mm. Greatest width 1.20 mm. Thoracic segments bearing legs 2 and 3 free. Genital segment comprising about 60 percent total body length; ventral surface with a patch of spinules between bases of leg 3, 3 patches of spinules between leg 4 setae, and another patch distal to leg 4 setae. Considerable variation of total length occurs in this species and this will be considered in a separate study by the first author and B. B. Collette. Abdomen (Figure 92a) somewhat wider than long (652 × 493 μm). Caudal rami about twice as long as wide (362 × 188 μ), 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 92f) 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 well-developed 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 genital segment.

Egg string long.

MALE.—Body form as in Figure 93d. Total length 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 × 0.53 mm); abdomen (Figure 93c) nearly twice as wide as long (259 × 147 μm) with a ventral patch of spinules near each outer distal corner. Caudal rami (see Figure 93e) about twice as long as wide (123 × 64 μm), 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 93f) similar to female except claw longer and more pointed. Mouth tube (Figure 94a) with 3 patches of spinules on labrum. 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 94b) 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 94e–g

Lernanthropus kanagurta Tripathi, 1962:194.

MATERIAL EXAMINED.—1 collection containing 1 art from the gills of Rastrelliger brachysoma from Borneo.

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 94f 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.

MATERIAL EXAMINED.—58 collections containing 114 art from the axil of the pectoral fin of the following hosts and localities: Acanthocybium solandri from Malagasy Republic, Ile 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

MATERIAL EXAMINED.—3 collections containing 3 art 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 95 c, 108

Anchorella scombri Kurz, 1877:403.

Clavella scombri.—Brian, 1906:116.

Clavellisa scombri.—Wilson, 1915:694.

MATERIAL EXAMINED.—10 collections containing 17 art 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 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-to-rare 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 capitulatus (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)

Pumiliopes capitulatus (8)

Clavellisa scombri (9)

Caligus pelamydis (4)

Caligus mutabilis (1)

Clavelopsis saba

Scomber australasicus (17)

Clavellisa scombri (1)

Pumiliopes capitulatus (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)

Shiinoa occlusa (5)

Caligus asymmetricus (4)

Scomberomorus brasiliensis (62)

Pseudocycnoides buccata (39)

Holobomolochus divaricatus (14)

Caligus mutabilis (4)

Shiinoa inauris (3)

Scomberomorus cavalla (36)

Pseudocycnoides buccata (18)

Holobomolochus asperatus (10)

Caligus mutabilis (2)

Caligus productus (1)

Scomberomorus commerson (113)

Pseudocycnoides armatus (20)

Unicolax ciliatus (20)

Shiinoa occlusa (11)

Caligus biseriodentatus (11)

Caligus cybii (10)

Caligus infestans (5)

Caligus asymmetricus (2)

Brachiella magna (2)

Tuxophorus cervicornis (2)

Scomberomorus concolor (47)

Pseudocycnoides buccata (14)

Holobomolochus nudiusculus (13)

Caligus omissus (7)

Scomberomorus guttatus (55)

Caligus biseriodentatus (16)

Unicolax ciliatus (13)

Pseudocycnoides armatus (2)

Shiinoa occlusa (1)

Scomberomorus koreanus (17)

Caligus cybii (11)

Pseudocycnoides armatus (3)

Scomberomorus lineolatus (10)

Pseudocycnoides armatus (1)

Unicolax ciliatus (1)

Caligus biseriodentatus (1)

Scomberomorus maculatus (76)

Pseudocycnoides buccata (27)

Holobomolochus divaricatus (24)

Shiinoa inauris (7)

Caligus mutabilis (1)

Scomberomorus multiradiatus (4)

Pseudocycnoides armatus (2)

Scomberomorus niphonius (18)

Pseudocycnoides scomberomori (6)

Unicolax ciliatus (2)

Caligus pelamydis (2)

Shiinoa occlusa (1)

Scomberomorous plurilineatus (3)

Pseudocycnoides armatus (3)

Scomberomorus regalis (38)

Pseudocycnoides buccata (12)

Holobomolochus divaricatus (11)

Shiinoa inauris (5)

Caligus productus (3)

Caligus bonito (1)

Brachiella thynni (1)

Tuxophorus collettei (1)

Scomberomorus semifasciatus (17)

Pseudocycnoides armatus (2)

Unicolax ciliatus (2)

Caligus cybii (1)

Scomberomorus sierra (116)

Pseudocycnoides buccata (48)

Caligus omissus (39)

Holobomolochus nudiusculus (28)

Scomberomorus sinensis (8)

Caligus cybii (2)

Brachiella magna (1)

Scomberomorus tritor (21)

Unicolax ciliatus (4)

Shiinoa occlusa (1)

Caligus productus (1)

Caligus diaphanus (1)

Scomberomorus queenslandicus (19)

Caligus biseriodentatus (11)

Pseudocycnoides armatus (2)

Unicolax ciliatus (1)

Shiinoa occlusa (1)

Scomberomorus species (3)

Unicolax ciliatus (1)

Caligus productus (1)

Caligus cybii (1)

SARDINI

Sarda sarda (104)

Caligus bonito (23)

Ceratacolax euthynni (21)

Caligus pelamydis (7)

Caligus productus (1)

Sarda australis (20)

Caligus bonito (13)

Caligus pelamydis (11)

Caligus asymmetricus (2)

Unicolax collateralis (2)

Sarda chiliensis (44)

Caligus bonito (24)

Caligus pelamydis (2)

Caligus productus (1)

Sarda orientalis (27)

Caligus bonito (6)

Caligus asymmetricus (4)

Unicolax collateralis (4)

Caligus productus (1)

Gymnosarda unicolor (7)

Caligus bonito (1)

Caligus productus (1)

Shiinoa occlusa (1)

Cybiosarda elegans (8)

Caligus asymmetricus (2)

Unicolax collateralis (1)

Orcynopsis unicolor (6)

Unicolax collateralis (1)

Allothunnus fallai (3)

Elytrophora brachyptera (3)

THUNNINI

Auxis species (65)

Unicolax collateralis (19)

Unicolax mycterobius (9)

Caligus productus (2)

Caligus asymmetricus (1)

Caligus coryphaenae (1)

Caligus pelamydis (1)

Euthynnus affinis (71)

Unicolax collateralis (29)

Caligus asymmetricus (5)

Caligus regalis (5)

Pseudocycnus appendiculatus (4)

Unicolax mycterobius (3)

Caligus pelamydis (2)

Caligus productus (1)

Caligus bonito (1)

Euthynnus alletteratus (64)

Caligus coryphaenae (9)

Unicolax collateralis (8)

Ceratacolax euthynni (7)

Caligus productus (5)

Caligus bonito (4)

Pseudocycnus appendiculatus (3)

Unicolax mycterobius (3)

Unicolax anonymous (2)

Caligus pelamydis (1)

Euthynnus lineatus (15)

Unicolax collateralis (4)

Caligus bonito (3)

Ceratacolax euthynni (2)

Unicolax mycterobius (1)

Katsuwonus pelamis (132)

Caligus coryphaenae (51)

Caligus productus (51)

Pseudocycnus appendiculatus (7)

Unicolax reductus (3)

Caligus asymmetricus (1)

Thunnus alalunga (12)

Elytrophora brachyptera (7)

Caligus coryphaenae (1)

Caligus productus (1)

Pseudocycnus appendiculatus (1)

Thunnus albacares (110)

Caligus productus (49)

Elytrophora brachyptera (39)

Caligus coryphaenae (32)

Pseudocycnus appendiculatus (21)

Brachiella thynni (8)

Caligus asymmetricus (1)

Thunnus atlanticus (76)

Caligus productus (70)

Caligus coryphaenae (9)

Elytrophora brachyptera (1)

Thunnus maccoyii (2)

Elytrophora brachyptera (2)

Thunnus obesus (40)

Elytrophora brachyptera (20)

Caligus coryphaenae (18)

Brachiella thynni (10)

Elytrophora indica (9)

Pseudocycnus appendiculatus (3)

Thunnus thynnus (57)

Caligus coryphaenae (16)

Caligus productus (16)

Elytrophora brachyptera (11)

Pennella species (3)

Brachiella thynni (2)

Caligus bonito (1)

Pseudocycnus appendiculatus (1)

Thunnus tonggol (26)

Pseudocycnus appendiculatus (7)

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

Scomberomorus maculatus (32)

Scomberomorus regalis (29)

Scomberomorus brasiliensis (23)

Holobomolochus nudiusculus

Scomberomorus concolor (28)

Scomberomorus sierra (24)

Holobomolochus asperatus

Scomberomorus cavalla (28)

Unicolax collateralis

Euthynnus affinis (41)

Auxis species (29)

Euthynnus lineatus (27)

Sarda orientalis (21)

Orcynopsis unicolor (17)

Cybiosarda elegans (13)

Euthynnus alletteratus (13)

Unicolax anonymous

Euthynnus alletteratus (3)

Unicolax mycterobius

Auxis species (14)

Euthynnus alletteratus (5)

Euthynnus affinis (4)

Unicolax ciliatus

Scomberomorus guttatus (24)

Scomberomorus tritor (19)

Scomberomorus commerson (18)

Scomberomorus semifasciatus (12)

Scomberomorus niphonius (11)

Scomberomorus lineolatus (10)

Unicolax reductus

Katsuwonus pelamis (2)

Ceratacolax euthynni

Sarda sarda (20)

Euthynnus alletteratus (14)

Nothobomolochus kanagurta

Rastrelliger faughni (14)

Rastrelliger kanagurta (4)

Rastrelliger species

Orbitacolax aculeatus

Rastrelliger faughni

Pumiliopes capitulatus

Rastrelliger faughni (14)

Rastrelliger kanagurta (10)

Scomber japonicus (2)

Scomber scombrus (1)

SHIINOIDAE

Shiinoa inauris

Scomberomorus regalis (13)

Scomberomorus maculatus (9)

Scomberomorus brasiliensis (5)

Shiinoa occlusa

Grammatorcynus bicarinatus (14)

Gymnosarda unicolor (14)

Scomberomorus commerson (10)

Scomberomorus niphonius (6)

Scomberomorus queenslandicus (5)

Scomberomorus guttatus (1)

Scomberomorus tritor (1)

Acanthocybium solandri (1)

CALIGIDAE

Caligus coryphaenae

Thunnus obesus (45)

Katsuwonus pelamis (38)

Thunnus albacares (29)

Thunnus thynnus (28)

Euthynnus alletteratus (14)

Thunnus atlanticus (12)

Thunnus alalunga (8)

Auxis species (1)

Caligus regalis

Euthynnus affinis (70)

Caligus productus

Thunnus atlanticus (92)

Thunnus albacares (45)

Katsuwonus pelamis (39)

Thunnus thynnus (28)

Acanthocybium solandri (15)

Sarda orientalis (13)

Euthynnus alletteratus (8)

Thunnus alalunga (8)

Scomberomorus regalis (8)

Scomberomorus tritor (5)

Scomberomorus cavalla (3)

Auxis species (3)

Sarda chiliensis lineolatus (2)

Euthynnus affinis (1)

Sarda sarda (1)

Scomberomorus species

Caligus asymmetricus

Cybiosarda elegans (25)

Sarda orientalis (21)

Grammatorcynus bicarinatus (11)

Sarda australis (10)

Euthynnus affinis (7)

Auxis species (2)

Scomberomorus commerson (2)

Katsuwonus pelamis (1)

Thunnus albacares (1)

Caligus bonito

Sarda australis (65)

Sarda chiliensis chiliensis (54)

Sarda chiliensis lineolatus (35)

Sarda orientalis (39)

Euthynnus lineatus (20)

Gymnosarda unicolor (14)

Euthynnus alletteratus (6)

Scomberomorus regalis (3)

Thunnus thynnus (2)

Euthynnus affinis (1)

Caligus mutabilis

Scomberomorus brasiliensis (6)

Scomberomorus cavalla (6)

Scomberomorus maculatus (1)

Scomber japonicus (1)

Caligus omissus

Scomberomorus sierra (34)

Scomberomorus concolor (15)

Caligus biseriodentatus

Scomberomorus queenslandicus (58)

Scomberomorus guttatus (29)

Scomberomorus commerson (10)

Scomberomorus lineolatus (10)

Auxis species

Caligus cybii

Scomberomorus koreanus (65)

Scomberomorus sinensis (25)

Scomberomorus species (20)

Scomberomorus commerson (9)

Scomberomorus semifasciatus (6)

Caligus pelamydis

Sarda australis (55)

Scomberomorus niphonius (11)

Sarda chiliensis lineolatus (10)

Sarda sarda (7)

Euthynnus affinis (3)

Euthynnus alletteratus (2)

Auxis species (2)

Scomber japonicus (1)

Scomber scombrus (1)

Caligus infestans

Scomberomorus commerson (4)

Caligus diaphanus*

Scomberomorus tritor (5)

Caligus savala*

Euthynnus affinis

Caligus macarovi*

Auxis species

Caligus amblygenitalis*

Euthynnus affinis

Caligus pseudokalumai

Gymnosarda unicolor

EURYPHORIDAE

Elytrophora brachyptera

Allothunnus fallai (100)

Thunnus maccoyii (100)

Thunnus alalunga (58)

Thunnus obesus (50)

Thunnus albacares (35)

Thunnus thynnus (19)

Thunnus atlanticus (1)

Elytrophora indica

Thunnus obesus (23)

Gloiopotes hygomianus

Acanthocybium solandri (43)

Caligulus longispinosus

Euthynnus affinis

TUXOPHORIDAE

Tuxophorus cybii

Scomberomorus commerson

Acanthocybium solandri

Tuxophorus cervicornis

Scomberomorus commerson (2)

Tuxophorus collettei

Scomberomorus regalis (3)

PSEUDOCYCNIDAE

Pseudocycnus appendiculatus

Thunnus tonggol (27)

Thunnus albacares (19)

Thunnus obesus (8)

Euthynnus affinis (6)

Euthynnus alletteratus (5)

Katsuwonus pelamis (5)

Pseudocycnoides armatus

Scomberomorus plurilineatus (100)

Scomberomorus koreanus (18)

Scomberomorus commerson (18)

Scomberomorus semifasciatus (12)

Scomberomorus queenslandicus (11)

Scomberomorus lineolatus (10)

Scomberomorus guttatus (4)

Pseudocycnoides scomberomori

Scomberomorus niphonius (33)

Pseudocycnoides buccata

Scomberomorus brasiliensis (63)

Scomberomorus cavalla (50)

Scomberomorus sierra (41)

Scomberomorus maculatus (36)

Scomberomorus regalis (32)

Scomberomorus concolor (30)

LERNANTHROPIDAE

Lernanthropus kanagurta*

Rastrelliger brachysoma (3)

LERNEOPODIDAE

Brachiella thynni

Acanthocybium solandri (61)

Thunnus obesus (25)

Thunnus albacares (7)

Thunnus thynnus (4)

Scomberomorus regalis (3)

Brachiella magna

Scomberomorus sinensis (13)

Scomberomorus commerson (2)

Clavellisa scombri

Scomber australasicus (6)

Scomber japonicus (2)

Clavellopsis saba

Scomber japonicus

___________

* Probably not usually scombrid parasites.

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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.

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.

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.

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.

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.

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; f, first antenna; g, second antenna.

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.

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.

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

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.

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

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.

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

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

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

FIGURE 18.—Unicolax mycterobius (Vervoort), new genus, female: a, leg 4; b, leg 5; male: c, dorsal; d, last abdominal segment and caudal rami, ventral; e, leg 1; f, 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.

FIGURE 20.—Unicolax ciliatus, new genus, new species, female: a, dorsal; b, first antenna; c, leg 1 exopod; d, leg 2 exopod.

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; f, leg 1 endopod.

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

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; f, mandible, paragnath, first maxilla, second maxilla.

FIGURE 24.—Unicolax reductus, new genus, new species, female: a, maxilliped; b, leg 1; c, leg 2; d, leg 3.

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.

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; f, second antenna; g, mandible, paragnath, first and second maxilla.

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

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; f, first antenna.

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.

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

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

FIGURE 34.—Orbitacolax aculeatus (Pillai), female: a, dorsal; b, first antenna; c, second antenna; d, oral area; e, maxilliped; f, leg 1.

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 rami, ventral.

FIGURE 36.—Pumiliopes capitulatus Cressey and Boyle, female: a, first antenna; b, second antenna; c, oral area; d, maxilliped; e, leg 1.

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

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

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: f, genital segment and abdomen; g, caudal ramus; h, second antenna.

FIGURE 40.—Caligus coryphaenae Steenstrup and Lütken, female: a, dorsal; b, caudal ramus, ventral; c, sternal furca; d, leg 1; e, leg 2 endopod; f, leg 3 exopod.

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

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

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

FIGURE 44.—Caligus productus Müller, female: a, leg 3; b, leg 4; male: c, dorsal; d, genital segment and abdomen; e, second antenna.

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

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

FIGURE 47.—Caligus asymmetricus Kabata, female: a, leg 3; b, leg 4; male: c, dorsal; d, genital segment and abdomen, ventral; e, second antenna.

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.

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

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

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; e, postantennal process; f, second antenna.

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

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

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

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

FIGURE 56.—Caligus omissus, new species, male: a, dorsal; b, genital segment and abdomen, ventral; c, second antenna; d, postantennal process; e, postoral process; f, 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.

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

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; f, maxilliped.

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.

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

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

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

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

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

FIGURE 66.—Caligus pelamydis Kroyer, female: a, leg 3; b, leg 4; male: c, dorsal; d, genital segment and abdomen, ventral; e, postoral process; f, maxilliped.

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.

FIGURE 68.—Caligus infestans Heller, female: a, second antenna; b, tip of second maxilla; c, maxilliped; d, sternal furca; e, leg 1; f, leg 2.

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

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; f, second antenna; g, postoral process.

FIGURE 71.—Caligus diaphanus Nordmann, female: a, second maxilla; b, maxilliped; c, sternal furca; d, oral area; e, leg 1; f, leg 2.

FIGURE 72.—Caligus diaphanus Nordmann, female: a, leg 3; b, leg 4; male: c, dorsal; d, second antenna; e, postoral process; f, maxilliped.

FIGURE 73.—Caligus savala Gnanamuthu, female: a, dorsal; b, second antenna; c, postantennal process; d, sternal furca; e, leg 2 endopod; f, 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.)

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; f, 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 Gerstaecker, female: a, dorsal; b, caudal ramus; c, first antenna; d, second antenna; e, mouth tube and postoral process; f, second maxilla; g, maxilliped.

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

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; f, second antenna; g, maxilliped.

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; f, leg 4; g, leg 6.

FIGURE 79.—Elytrophora indica Shiino, male: a, dorsal; b, second antenna. Gloiopotes hygomianus Steenstrup and Lütken, 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.

FIGURE 80.—Caligulus longispinosus Heegaard, female: a, dorsal; male: b, dorsal; c, maxilliped; d, sternal furca; e, leg 1; f, leg 2 (female after Heegaard, 1962).

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; f, leg 1; g, leg 4. (T. cybii after N.-R and Fourmanoir, 1956.)

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; f, postoral process.

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

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; f, maxilliped.

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

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

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

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

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

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

FIGURE 91.—Pseudocycnoides armatus (Bassett-Smith), male: a, leg 2; b, leg 3. Pseudocycnoides scomberomori (Yamaguti), female: c, dorsal; d, maxilliped; e, leg 1; f, leg 2; g, leg 3.

FIGURE 92.—Pseudocycnoides buccata (Wilson), female: a, dorsal; b, abdomen and caudal rami, young female, ventral; c, first antenna; d, second antenna; e, mouth tube; f, 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; f, second antenna.

FIGURE 94.—Pseudocycnoides buccata (Wilson), male: a, first maxilla, mandible and mouth tube; b, maxilliped; c, leg 1; d, leg 3. Lernanthropus kanagurta Tripathi, female: e, dorsal; f, second antenna; g, leg 1.

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. (B. thynni after Shiino, 1956; C. saba after Shiino, 1959c.)

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

FIGURE 97.—Distribution of Unicolax collateralis, new species.

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

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

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. occulusa Kabata.

FIGURE 102.—Distribution of Caligus coryphaenae Steenstrup and Lütken, and C. regalis 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.

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

FIGURE 105.—Distribution of Caligus bonito Wilson, C. omissus, new species, C. mutabilis Wilson, and C. biseriodentatus Shen.

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

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

FIGURE 108.—Distribution of Brachiella thynni Cuvier, Clavellisa scombri (Kurz), B. magna Kabata, and Clavellopsis saba Yamaguti. (Includes Kabata, 1968b B. magna.)

FIGURE 109.—Holobomolochus divaricatus, new species, female: a, rostral area (1775×); b, abdomen and caudal rami, ventral (540×); c, abdomen, outer distal corner (2950×); d, labrum (1000×); e, paragnath (3145×); f, leg 2 exopod (500×).

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

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

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

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

FIGURE 114.—Unicolax collateralis, new species, female: a, second antenna (750×); b, hooks on second antenna (2400×); c, same (2600×); d, labrum (600×); e, oral area (1500×); f, tip of mandible (10,000×).

FIGURE 115.—Unicolax collateralis, new species, female: a, tip of paragnath (10,000×); b, leg 2 exopod (1000×); c, spines on leg 2 exopod (2000×); d, hyaline fringe of leg 2 exopod outer edge (8000×); e, spine on leg 2 exopod (2500×); f, leg 4 endopod last segment (1000×).

FIGURE 116.—Unicolax collateralis, new species, female: a, tip of leg 4 endopod (2000×). Unicolax anonymous (Vervoort), female: b, cephalon, anterior view (600×); c, first and second antenna, anterior view (700×); d, modified seta of first antenna (2300×); e, hooks on second antenna (7500×); f, oral area (750×).

FIGURE 117.—Unicolax anonymous (Vervoort), female: a, leg 2 exopod (1650×); b, leg 3 exopod outer edge (1750×). Unicolax mycterobius (Vervoort), female: c, cephalon, anterior view (200×); d, surface of cephalon dorsal to insertion of first antennae (1400×); e, cephalon, anteroventral view (200×); f, modified seta of first antenna (400×).

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

FIGURE 119.—Unicolax mycterobius (Vervoort), female: a, leg 1 exopod first segment (700×); b, leg 1 basipod and exopod insertion (1050×); c, leg 4 endopod tip (750×). Unicolax reductus, new species, female: d, distal end of caudal ramus (1500×); e, modified seta of first antenna (500×); f, second antenna (1000×).

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

FIGURE 121.—Unicolax reductus, new species, female: a, leg 3 (260×); b, leg 4 endopod first segment spinules (2000×); c, leg 3 exopod first segment (1000×); d, leg 3 exopod first segment spinules (5000×); e, leg 4 (240×); f, leg 4 exopod (650×).

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

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

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

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

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

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

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

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

FIGURE 130.—Caligus coryphaenae Steenstrup and Lütken, female: a, leg 2 exopod (260×); b, spine on leg 2 first segment (650×); c, spinules on outer edge of leg 2 endopod (1000×); d, tip of leg 4 (290×); e, spine on leg 4 second segment (600×); f, fringe at base of terminal seta on leg 4 (1200×).

FIGIRE 131.—Pseudocycnus appendiculatus Heller, female: a, cephalon, anterior view (55×); b, same, between bases of first antennae (300×); c, anal opening (240×); d, same (373×); e, second antenna (105×); f, second maxilla (500×).

FIGURE 132.—Pseudocycnus appendiculatus Heller, female: a, maxilliped (200×); b, tip of maxilliped (500×); c, leg 1 (420×); d, leg 1 exopod base of terminal spine (1800×); e, leg 1 endopod (750×); f, same, lateral view (800×)

FIGURE 133.—Pseudocycnus appendiculatus Heller, female: a, leg 2 (125×); b, same (500×); c, leg 3 (400×); d, tip of leg 3 (1200×); male: e, genital segment, ventral (150×); f, spinules on genital segment, ventral (5000×).

FIGURE 134.—Pseudocycnus appendiculatus Heller, male: a, caudal ramus, ventral (200×); b, spinules on caudal ramus (2000×); c, mouth tube (400×); d, tip of mouth tube (2000×); e, first maxilla (400×); f, base of maxilliped (500×).

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

FIGURE 136.—Pseudocycnus appendiculatus Heller, male: a, base of leg 5 (5000×). Pseudocycnoides armatus (Bassett-Smith), male: b, spinules on genital segment (6550×); c, abdomen and caudal rami, ventral (250×); d, caudal ramus, ventral (750×); e, tip of caudal ramus, ventral (1600×); f, leg 1 (500×).

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

FIGURE 138.—Pseudocycnoides buccata (Wilson), female: a, cephalon, ventral (75×); b, second antenna (370×); c, mouth tube (600×); d, tip of mouth tube (2500×); e, first maxilla (1250×); f, tip of second maxilla (1450×).

FIGURE 139.—Pseudocycnoides buccata (Wilson), female: a, tip of second maxilla (1225×); b, same (1450×); c, same (4400×);

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Taxonomic keys in natural history papers should use the alined-couplet form in the zoology and paleobiology series and the multi-level indent form in the botany series. If cross-referencing is required between key and text, do not include page references within the key, but number the keyed-out taxa with their corresponding heads in the text.

Synonymy in the zoology and paleobiology series must use the short form (taxon, author, year:page), with a full reference at the end of the paper under “Literature Cited.” For the botany series, the long form (taxon, author, abbreviated journal or book title, volume, page, year, with no reference in the “Literature Cited”) is optional.

Footnotes, when few in number, whether annotative or bibliographic, should be typed at the bottom of the text page on which the reference occurs. Extensive notes must appear at the end of the text in a notes section. If bibliographic footnotes are required, use the short form (author/brief title/page) with the full reference in the bibliography.

Text-reference system (author/year/page within the text, with the full reference in a “Literature Cited” at the end of the text) must be used in place of bibliographic footnotes in all scientific series and is strongly recommended in the history and technology series: “(Jones, 1910:122)” or “…. Jones (1910:122).”

Bibliography, depending upon use, is termed “References,” “Selected References,” or “Literature Cited.” Spell out book, journal, and article titles, using initial caps in all major words. For capitalization of titles in foreign languages, follow the national practice of each language. Underline (for italics) book and journal titles. Use the colon-parentheses system for volume/number/page citations: “10(2):5–9.” For alinement and arrangement of elements, follow the format of the series for which the manuscript is intended.

Legends for illustrations must not be attached to the art nor included within the text but must be submitted at the end of the manuscript—with as many legends typed, double spaced, to a page as convenient.

Illustrations must not be included within the manuscript but must be submitted separately as original art (not copies). All illustrations (photographs, line drawings, maps, etc.) can be intermixed throughout the printed text. They should be termed Figures and should be numbered consecutively. If several “figures” are treated as components of a single larger figure, they should be designated by lowercase italic letters (underlined in copy) on the illustration, in the legend, and in text references: “Figure 9b.” If illustrations are intended to be printed separately on coated stock following the text, they should be termed Plates and any components should be lettered as in figures: “Plate 9b.” Keys to any symbols within an illustration should appear on the art and not in the legend.

A few points of style: (1) Do not use periods after such abbreviations as “mm, ft, yds, USNM, NNE, AM, BC.” (2) Use hyphens in spelled-out fractions: “two-thirds.” (3) Spell out numbers “one” through “nine” in expository text, but use numerals in all other cases if possible. (4) Use the metric system of measurement, where possible, instead of the English system. (5) Use the decimal system, where possible, in place of fractions. (6) Use day/month/year sequence for dates: “9 April 1976.” (7) For months in tabular listings or data sections, use three-letter abbreviations with no periods: “Jan, Mar, Jun,” etc.

Arrange and paginate sequentially EVERY sheet of manuscript—including ALL front matter and ALL legends, etc., at the back of the text—in the following order: (1) title page, (2) abstract, (3) table of contents, (4) foreword and/or preface, (5) text, (6) appendixes, (7) notes, (8) glossary, (9) bibliography, (10) index, (11) legends.