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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 296

The Parasitic Copepods of Indo-West Pacific Lizardfishes (Synodontidae)

Roger Cressey and Hillary Boyle Cressey

SMITHSONIAN INSTITUTION PRESS

City of Washington

1979

ABSTRACT

Cressey, Roger, and Hillary Boyle Cressey. The Parasitic Copepods of Indo-West Pacific Lizardfishes (Synodontidae). Smithsonian Contributions to Zoology, number 296, 71 pages, 53 figures, 2 tables, 1979.—Eighteen species of parasitic copepods (12 new species) are described from Indo-West Pacific Lizardfishes. Twelve species are taeniacanthids, 10 of the genus Metataeniacanthus Pillai. Species of Metataeniacanthus are very host specific and aided in the identifications of the hosts. The caligid genus Abasia Wilson, formerly with 2 species is expanded to now include 6 species (3 new). The genus Alicaligus Shiino is placed in synonymy with Abasia. The species included are Metataeniacanthus aquilonius new species; M. conepigri new species; M. epigri new species; M. gibbsi new species; M. indiscretus new species; M. nudus new species; M. pacificus new species; M. solidus new species; M. synodi Pillai; M. vulgaris new species; Parataeniacanthus longicervis Pillai; Taeniacanthus sauridae Yamaguti and Yamasu; Abasia inflata new species; A. pillaii new species; A. platyrostris Pillai; A. pusilla new species; A. tripartita (Shiino); Lernanthropus temminckii Nordmann.

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–

The parasitic copepods of Indo-West Pacific lizardfishes (Synodontidae)

(Smithsonian contributions to zoology; no. 296)

Bibliography: p.

1. Copepoda—Pacific Ocean—Classification. 2. Copepoda—Indian Ocean—Classification. 3. Parasites—Fishes. 4. Synodontidae—Diseases. 5. Crustacea—Pacific Ocean—Classification. 6. Crustacea—Indian Ocean—Classification. I. Cressey, Hillarv Boyle. 1949– joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology: no. 296.

QL1.S54 no. 296 [QL444.C7] 591′.08s [595′.34] 79–11097

The Parasitic Copepods of Indo-West Pacific Lizardfishes (Synodontidae)

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

In 1973 Dr. I. Paperna of The Hebrew University sent the first author a collection of parasitic copepods from the Red Sea and East Africa containing a single specimen of a new species of Metataeniacanthus from a Red Sea Synodus. We subsequently examined several additional Red Sea and Indian Ocean Synodus housed in the Smithsonian collections to obtain additional material. These early collections contained not only more material of the Red Sea species but other new species as well. This led us to undertake a comprehensive examination of Indo-West Pacific lizardfishes for parasitic copepods. We examined 1954 lizardfishes and collected 18 species of parasitic copepods (12 new) described below. All of the species reported in this paper are considered by us to be specific parasites of synodontid fishes.

Representative lizardfishes from the Atlantic and Eastern Pacific Ocean were also examined. The Eastern Pacific hosts were generally devoid of parasitic copepods and those found were so few as not to indicate any host preferences. Of the Atlantic hosts examined, only Synodus foetens (L.) was commonly parasitized by a copepod, Abasia pseudorostris Wilson (1908), described from the same host and redescribed by Pillai (1963b).

The genus Metataeniacanthus Pillai is expanded from the original description of M. synodi to now include 10 species. The genus Abasia Wilson, formerly consisting of one Atlantic and one Indian Ocean species, now includes 6 species, 3 new and one synonymizing Shiino's Alicaligus tripartitus.

Based initially on the specificity of the species of Metataeniacanthus, the first author is publishing separately a revision of the Indo-West Pacific Synodus.

All material has been preserved in 70 percent alcohol and deposited in the collections of the Smithsonian Institution and British Museum (Natural History).

All figures were drawn with the aid of a Wild drawing tube. Specimens were also examined with a Cambridge stereoscan microscope. The descriptions and illustrations of Metataeniacanthus and Abasia incorporate observations made with the SEM; several SEM photographs of Abasia species are contained herein.

ACKNOWLEDGMENTS.—We thank the many curators of the following collections who either made lizardfish and copepod collections available or sent host specimens to us for study: The Australian Museum, Sydney; The Academy of Natural Sciences of Philadelphia, Philadelphia; The British Museum (Natural History), London: The Bernice P. Bishop Museum, Honolulu; The California Academy of Sciences, San Francisco; The Field Museum of Natural History, Chicago; Kyoto University, Kyoto; The Los Angeles County Museum, Los Angeles; The Museum of Comparative Zoology, Cambridge, Mass.; The National Museum of New Zealand, Wellington; The Queensland Museum, Brisbane; Rijksmuseum van Natuurlijke Historie, Lieden; Smithsonian Institution, Washington; The Western Australian Museum, Perth; National Taiwan University, Taiwan; Zoologisches Institut und Zoologisches Museum, Hamburg. We also thank Dr. Krishna Pillai and Mr. A. Chandran, Trivandrum, India for sending us parasitic copepods from Indian lizardfish.

The illustrations of Lernanthropus temminckii Nordmann were done by Nancy A. Zacks.

Metataeniacanthus Pillai, 1963

DIAGNOSIS.—Female: Lateral margins of carapace directed ventrad flanking oral area. Abdominal segments usually indistinctly divided. Abdomen about half total body length. Cephalic appendages as in Taeniacanthus. Maxilliped with a short claw, closely appressed to preceding segment. Leg 1 with broadly flattened rami. Second endopod segment of legs 2 and 3 with 1 inner seta. Legs 2–4 with 3–segmented rami. Caudal rami each with 1 lateral outer seta, one subterminal and 4 terminal setae (middle two much longer than outer two).

Male: Body form typically cyclopoid. Lateral margins of cephalon not directed ventrad. Abdomen 3–segmented. Appendages in general as in female except maxilliped with heavily toothed claw, expanded at tip.

TYPE-SPECIES.—Metataeniacanthus synodi Pillai, 1963a.

DISCUSSION.—When Pillai, 1963a, proposed the genus Metataeniacanthus based on the single species M. synodi he stated that the genus was provisional and its status depended on the discovery of additional species. The additional species described herein leave no doubt as to the validity of this genus.

The high degree of host specificity will enable R. Cressey to redefine the Indo-West Pacific lizardfishes of the genus Synodus. Most museum specimens in this group were identified as Synodus variegatus (Lacepède). As we began to collect different species of Metataeniacanthus from this material the misidentification of many of the hosts became apparent. Since no specialist was working in this host group R. Cressey was advised by ichthyologists that he should consider working out the systematics of Indo-West Pacific Synodus as well. The results of this work with Synodus will be reported elsewhere. Without firm host identifications the host-parasite data is of little value. The discovery of these additional species of Metataeniacanthus will enable the first author to revise the systematics of a host group based initially on the host-specifity of their parasitic copepods.

The host-parasite relationships present some intriguing problems. Why does Synodus hoshinonis harbor two species of Metataeniacanthus separated by the Malay archipelago? Why are some common Indo-West Pacific species such as Synodus binotatus Schultz without Metataeniacanthus? This host species is often collected with Synodus variegatus and S. englemani, both of which commonly harbor Metataeniacanthus. Why do individuals of Metataeniacanthus synodi usually orient with the head toward the distal end of the gill filament, contrary to all the other species of Metataeniacanthus? This peculiar orientation is unlike that of parasitic copepods of gill filaments in general. Usually the posterior of the parasite is oriented toward the distal end of the gill filament with the eggs exposed beyond the filament tip.

The genus is apparently restricted to the Indo-West Pacific. The spine and seta formula for legs 2–4 differ only on the third segment of the exopods as shown in Table 1.

Key to Species of Metataeniacanthus Females

Metataeniacanthus synodi Pillai

FIGURES 1–3, 45

MATERIAL EXAMINED.—Three paratype art deposited in the Smithsonian Institution by Dr. Pillai; collected from the gills of Synodus indicus (Day) at Trivandrum, India on 15 March 1961. Remaining material all from gills of Trachinocephalus myops (Schneider) at the following localities: 1 art from Sydney, Australia; 3 art from 11 hosts from Japan; 4 art from 7 hosts from Formosa Strait; 9 art from China coast; 7 art from 15 hosts from the Philippines (USNM collection); 3 art from 1 host from off Viet Nam; 5 art from 1 host from Gulf of Thailand; 4 art from I host from Gopaldur, India; 33 art from 4 hosts from the Arabian Sea (17°25′N, 71°39′E) R/V Anton Bruun, Cruise 4B; 8 art from 24 hosts from the Mozambique Channel (19°50′S, 36°21′E) R/V Anton Bruun, Cruise 8.

Female: Body form as in Figures 1a and 1b. Total length 2.1 mm, greatest width 0.58 mm (measured at widest part of cephalon). Cephalon wider than long (0.58 × 0.39 mm), fused with first thoracic segment. Thoracic segments 2–4 clearly articulated (see Figure 1a) or not (see Figure 1b). Area of egg sac attachment (Figure 1c) fused with abdomen. Abdomen about half total body length, segments measure 424, 324, 177, and 141 μm respectively; no ornamentation at posterior tip. Caudal rami (Figure 1d) about 3 times as long as wide (88.5 × 29.5 μm), each bearing one lateral, one subterminal and 4 terminal setae (longest 306.8 μm).

First antenna (Figure 1e) 7–segmented (Pillai, 1963a, considered the first antenna to be 6–segmented but his 3rd segment is actually subdivided and in light of other species described herein, we consider 7 segments a more accurate description); an aesthete is present on each of last 2 segments in addition to setae. Second antenna (Figure 1f) 3–segmented, first and second segments bearing one seta each, last segment with row of fine spinules along posterior edge continuing along edge of blunt process on ventral side, tip armed with 3 hooklike spines and 3 setae. Postantennal hook (Pillai's maxillary hook) (Figure 1g) slightly recurved. Oral area as in Figure 2a. Labrum posterior edge finely spinulose. Mandible, first maxilla, and second maxilla as in figure. Paragnath (Figure 2b) posterior distal edge lobed with a single row of spines in pits near base of lobes. Maxilliped (Figure 2c) with short distal hook (Pillai shows no distal hook but hook is held close to basal part of maxilliped and difficult to see).

TABLE1.—Spine and seta formulae for the exopod last segments of legs 2–4 of Metataeniacanthus species (Arabic numbers = setae, Roman = spines)

| legs 2 & 3 | leg 4
synodi, vulgaris, nudus | III:5 | III:5
pacificus | III:5 | III:4
epigri | III:5 | IV:3
indiscretus, aquilonius | III:4 | III:4
conepigri, gibbsi, solidus | III:4 | IV: 3

Legs 1–4 biramous; rami 3–segmented; Leg 1 (Figure 2d) basipod with plumose stout spine on outer distal corner and long plumose seta near interpodal plate; exopod (Figure 2e) first 2 segments each with a plumose seta on outer edge, inner edge of second segment with stout seta, third segment with 7 setae; endopod first 2 segments each with an inner seta, third segment with 7 setae, outer 2 much shorter than others; interpodal plate with posterior spinose process as in figure. Leg 2 (Figure 2f) basipod with outer naked seta; exopod first 2 segments each with slender, naked spine on outer distal corner, third segment with 3 slender, naked outer spines and 5 finely plumose distal to inner setae, inner margin of outermost seta densely plumose near base; a pair of short processes near base of outer spine of second segment and first 2 outer spines of third segment; a pair of setules near base of distalmost spine on third segment; endopod first 2 segments each with a single inner seta, third segment bearing 3 spines and 3 setae, outer 2 spines modified as in Figure 3a. Leg 3 (Figure 3b) similar to leg 2 except exopod outer spines with hyaline fringe, setules on exopod last segment of leg 2 replaced by blunt processes on leg 3, dense plumosities on outermost seta of exopod last segment not present in leg 3. Leg 4 (Figure 3c) basipod and exopod as in leg 3 except for lack of blunt processes near bases of outer spines on leg 4, each exopod segment with a setule near outer edge; endopod armed as in leg 3 except only 2 spines and 1 seta on distal segment. Leg 5 (Figure 3d) basal segment with outer seta; last segment with 4 setae, terminalmost longest, other 3 of about equal lengeth. Leg 6 (see Figure 1c) represented by 3 short setae at area of egg sac attachment.

Egg sac cylindrical, extending to beyond middle of abdomen.

Male: Unknown.

REMARKS.—Pillai collected this copepod from the gills of Synodus indicus. We examined many specimens of this species but did not recover this parasite. We did, however, find this copepod a common parasite of Trachinocephalus myops. Unlike other species of Metataeniacanthus this species generally attaches to the gill filament of the host with the head oriented toward the distal part of the filament. Most parasitic copepods that attach to gill filaments do so with the eggs exposed distally. We can offer no explanation as to why this species would orient its body in the opposite direction. Ovigerous females were collected with two different body forms (Figures 1a and 1b). No correlation could be found with host, site of infestation, geographic area, or size of parasite to account for this variation. The variation might depend on the age of the parasite, but this could not be properly evaluated from preserved material. All specimens considered were ovigerous.

Metataeniacanthus vulgaris, new species

FIGURES 4–5, 46

MATERIAL EXAMINED.—Holotype art (USNM 171326) and 4 art paratypes (USNM 171327) from the gills of 2 Synodus variegatus (Lacepède) collected by L. P. Schultz from Tutuila Reef, Samoa, 4 June 1939. All of the following collections are from the gills of the same host species. One collection containing 1 art from off Zanzibar. Twelve collections containing 65 art from the Seychelles. Five collections containing 10 art from off Ceylon. Three collections containing 9 art from off western Sumatra. One collection containing 1 art from off China. Seven collections containing 33 art from the Philippines. Three collections containing 22 art from the Celebes. Two collections containing 5 art and 1 art from Noumea, New Caledonia. Four collections containing 35 art from off Palau Islands. Eleven collections containing 42 art from off Queensland, Australia (Great Barrier Reef). One collection containing 7 art from Trobriand Islands, New Guinea. One collection containing 2 art from the Solomon Islands. Six collections containing 49 art from the Marshall Islands. Six collections containing 29 art from off Fiji. One collection containing 1 art from off Tonga Islands. One collection containing 2 art from off Samoa.

Female: Body form as in Figure 4a. Total length 3.03 mm, greatest width 0.58 mm (measured at widest part of cephalon). Cephalon wider than long (0.52 × 0.58 mm), fused with first thoracic segment. Fourth and fifth thoracic segments incompletely divided. Genital segment and area of egg sac attachment tachment (Figure 4b) wider than long (129 × 295 μm), not distinctly separated from abdomen. Abdomen more than half total body length (1.84 mm), unsegmented, cylindrical, with very fine spinules at posterior tip (see Figure 4c). Caudal rami (Figure 4c) longer than wide (94.4 × 35.4 μm), each bearing one lateral, one subterminal and 4 terminal setae (longest 289μm).

First antenna (Figure 4d) 7–segmented, an aesthaete on each of last 2 segments: segments measure 91, 71, 24, 15, 24, 18, and 29 μm long respectively; most setae on first 4 segments finely plumose. Second antenna (Figure 4e) 3–segmented; first and second segments bearing 1 seta each; last segment with row of fine spinules along posterior edge continuing along edge of blunt process on ventral side, tip of segment armed with larger blunt process also edged with fine spinules, 3 hooked spines and 3 setae. Postantennal hook (Figure 4f) recurved apically. Oral area as in Figure 4g. Mandible with 2 subequal bladelike processes at tip, each finely toothed. Paragnath distally trilobed with medial row of 8–11 blunt spinules recessed in pits. Second maxilla with 3 distal processes, shortest unarmed, longest with row of fine spinules, third with hooked spinules on each side. Maxilliped (Figure 4h) base with 2 inner, naked setae, short seta at base of terminal hook; hook with slight outward curve at tip and reaching to base of lateral setae on basal segment.

Legs 1–4 biramous; rami 3–segmented except 2–segmented exopod of first leg. Leg 1 (Figure 5a) basipod with stout plumose seta on outer corner and longer seta on inner edge near interpodal plate; exopod first segment with 1 outer seta, last segment with 3 short outer setae, 6 longer distal to inner setae; endopod first segment with short inner seta, second segment with longer inner seta, last segment with 7 setae (outer 2 short). Leg 2 (Figure 5b) basipod with naked seta on outer distal corner; exopod first segment with stout finely spinulose spine on outer distal corner, inner edge of segment with short, thick hairs, second segment with outer spinulose spine and inner seta, last segment bearing small papilla with minute sensilla near base of each 3 outer spinulose spines, terminal to inner edge with 5 setae; endopod first segment outer edge fringed with hairs, 1 inner distal seta, second segment similar to first, last segment with 3 outer spinulose spines, 3 terminal to inner setae (outermost short). Leg 3 (Figure 5c) similar to leg 2 except endopod last segment with 3 outer spines and only 2 terminal setae; outer distal spine of exopod first segment with rows of spinules near base not seen with light microscope but seen with SEM. Leg 4 (Figure 5d) similar to leg 3 except exopod last segment lacking papilla near base of 3 spines; endopod last segment with only 2 outer spines and 1 terminal seta. Leg 5 (Figure 5e) basal segment with dorsal outer seta; last segment with 4 setae of nearly equal length (2 outer, 2 terminal). Leg 6 (see Figure 4b) represented by 3 setae at area of egg sac attachment.

Egg sac cylindrical, nearly as long as abdomen, curving over abdomen.

Male: Body form similar to other males of genus. Total length 0.79 mm, greatest width 0.23 mm. Cephalon slightly longer than wide (0.26 × 0.23 mm). Genital segment longer than wide (177 × 132 μm). Abdomen 3–segmented, segments wider than long, measuring 29 × 82 μm, 29 × 75 μm, 44 × 67 μm respectively. Caudal rami longer than wide (59 × 29 μm), each ramus bearing one lateral, 2 subterminal and 3 terminal setae (longest 218 μm).

Cephalic appendages, except maxilliped, similar to female. Maxilliped (Figure 5f) basal segment with single seta, second segment with single seta and 2 raised, semicircular rows of blunt spinules, terminal claw with single seta, inner edge toothed, terminal teeth largest.

Leg 1 similar to female. Spine and seta formula for legs 2–4 follows. Roman numerals refer to spines, Arabic to setae.

Legs 2 and 3 lack sensillae at bases of exopod spines. Leg 5 similar to female except inner terminal seta about twice as long as other setae on leg. Leg 6 absent.

ETYMOLOGY.—The Latin vulgaris (“common”) alludes to the wide range of the host and its parasite.

REMARKS.—The female of this species can be separated from all known species of Metateaniacanthus by the single sensilla near the base of the exopod spines of legs 2 and 3. This species is a common parasite of the gill filaments of Synodus variegntus, occurring throughout most of its range except the southernmost Indian Ocean and mid-Pacific.

Metataeniacanthus solidus, new species

FIGURES 6, 7

MATERIAL EXAMINED.—Holotype art (USNM 171324) and 1 art paratype (USNM 171325) from the area behind the filaments of the pseudobranch and on the operculum inner surface of Synodus variegatus collected off Wailangilala Island, Fiji, 26 May 1965. One collection containing 2 art from the pseudobranch and inner surface of the operculum of the same host collected in the Caroline Islands. One collection containing 2 art from the pseudobranch of the same host collected at Bikini, Marshall Islands. One collection containing 1 art from the inner operculum of the same host species collected at Tonga Islands. One collection containing 1 art from the inner operculum on the same host species collected by B. Russell at Lizard Island, Queensland, 10 November 1975.

Female: Body form as in Figure 6a. Total length 3.64 mm, greatest width 0.53 mm (measured at widest part of cephalon). Cephalon as long as wide, fused with first thoracic segment. Second, third, and fourth thoracic segments distinctly divided. Genital segment and area of egg sac attachment wider than long (401 × 177 μm). Abdomen more than half body length (2.06 mm) unsegmented, without spinules at posterior tip, about 4 times as long as wide (2.06 × 0.51 mm). Caudal rami (Figure 6b) longer than wide (71 × 41 μm) armed with setae as in other species of genus, longest seta 313 μm.

First antenna (Figure 6c) 7–segmented, armed as in M. vulgaris. Second antenna (Figure 6d) armed as in M. vulgaris except terminal hooks less recurved. Oral area as in Figure 6e. Mandible as in other members of genus. Paragnath bilobed at tip with short row of spinules near tip. First maxilla with 2 short anteriorly directed setae and 2 long posteriorly directed setae. Second maxilla with 3 processes at tip, shortest unarmed, other 2 with hooklike spinules on two sides. Maxilliped (Figure 6f) base with 3 inner, naked setae, seta at base of terminal hook, hook slightly curved and not reaching to base of lateral setae on basal segment.

Legs 1–4 biramous, rami 3–segmented. Leg 1 (Figure 6g) basipod with stout plumose seta on outer distal corner and longer seta on inner edge near interpodal plate; exopod 3–segmented, first segment with plumose seta on outer distal corner, second segment with shorter plumose seta on outer margin and stout plumose seta on inner margin, third segment with 2 outer setae and 5 longer terminal to inner setae; endopod armed as in M. epigri. Leg 2 (Figure 6h) armed as in M. conepigri. Leg 3 (Figure 7a) as in M. conepigri except terminal spine on last endopod segment of M. solidus much longer relative to segment length. Leg 4 (Figure 7b) armed with spines and setae as in M. conepigri and exopod segments heavily sclerotized but outer edge not contorted inwardly as in M. conepigri. Leg 5 (Figure 7c) armed as in M. conepigri except terminal 2 setae equal in length (outermost shorter in M. conepigri). Leg 6 (Figure 7d) represented by 3 setae at area of egg sac attachment. Egg sac as in M. conepigri.

Male: Unknown.

ETYMOLOGY.—The Latin solidus (“dense,” “thick,” “hard”) alludes to the heavy sclerotization on the exopods of legs 2–4.

REMARKS.—This species can be separated from all other Metataeniacanthus by the nature of the sclerotized exopod of leg 4.

Although this copepod is found on the same host, Synodus variegatus, as M. vulgaris and in 3 of the 4 collections with M. vulgaris it occupies a different infestation site. It has never been collected from the primary gill filaments but, so far, has always been found either on the inner operculum wall or on the pseudobranch.

Metataeniacanthus epigri, new species

FIGURES 8–10, 47

MATERIAL EXAMINED.—Holotype (USNM 171313) and 28 art paratypes (USNM 171314) from the gills of Synodus englemani Schultz (paratypes USNM 152981) collected at Bikini, 8 July 1946. Seven collections containing 27 art from the Red Sea near Aquaba. A single collection containing 1 art from off Zanzibar. A single collection containing 3 art from the Comores Islands. Two collections containing 9 art from Nossi Bè, Malagasy Republic. Four collections containing 12 art and 1 art from Caragados Islands. Twenty three collections containing 236 art and 3 art from the Seychelles. Two collections containing 11 art from off Male, Maldive Islands. One collection containing 2 art from the South China Sea off Viet Nam. Seven collections containing 17 art from Big Hope Island and Endeavour Reef off Australia. Two collections containing 55 art from off the Philippines. Two collections containing 4 art from Indonesia. Three collections containing 24 art from near Trobriand Islands, New Guinea. Three collections containing 36 art from off New Britain. Three collections containing 6 art from off Ambon Island. One collection containing 75 art and 1 art from Celebes. Two collections containing 5 art from the Solomon Islands. Two collections containing 3 art and 1 art from the Marianas Islands. Four collections (exclusive of types) containing 68 art from the Marshall Islands. Three collections containing 14 art from Palau Islands. One collection containing 7 art from Tonga Islands. One collection containing 1 art from Fiji. One collection containing 1 art from Samoa. One collection containing 4 art from Tahiti. One collection containing 2 art from Gambier Islands. All above collections from the gill filaments of Synodus englemani.

Female: Body form as in Figure 8a. Total length 1.94 mm, greatest width 0.42 mm (measured at widest part of cephalon). Cephalon wider than long (0.34 × 0.42 mm) with low rounded rostrum. Genital segment and area of egg sac attachment (Figure 8b) wider than long (118 × 230 μm), not distinctly separated from abdomen and seen as a prominent ventral projection when viewed laterally. Abdomen almost half body length (0.94 mm), unsegmented, cylindrical, tapering gradually, with fine spinules near base of caudal rami (Figure 8c). Caudal rami (Figure 8c) almost 3 times as long as wide (94 × 32 μm), each ramus with 6 setae, innermost terminal finely plumose, longest 324 μm.

First antenna (Figure 8d) 7–segmented with aesthete on last 2 segments; segments measure 83, 68, 24, 24, 32, 24, and 35 μm long respectively, basal segments with 25 plumose setae. Second antenna (Figure 8e) 3–segmented, armed essentially as in other species. Oral area as in Figure 8f; labrum with fine spinules along posterior edge, mandible similar to other species, paragnath conspicuously trilobed and bearing row of spinules, remaining mouth parts as in other species. Maxilliped (Figure 8g) claw not reaching to base of setae on basal segment.

Legs 1–4 biramous, rami 3–segmented. Leg 1 (Figure 9a) basipod with stout plumose seta on outer corner and longer plumose seta on inner edge near interpodal plate; exopod 3–segmented, first segment with a seta on outer distal corner, second with a seta on outer and one on inner edge, last segment with 7 setae; endopod first segment with inner seta (segment indistinctly separated from basipod), second segment with an inner seta, last segment with 7 setae (outer 2 much shorter than other 5). Leg 2 (Figure 9b) basipod with naked seta on outer distal corner; exopod first segment with long, slender, finely spinulose spine on outer distal corner, inner edge with thick hairs, second segment with 2 blunt sensillae near base of outer spine and 1 inner seta, last segment with a pair of blunt sensillae near base of each of the 3 outer spines (Figure 9c) and terminal to inner edge with 5 finely plumose setae; endopod first segment outer edge fringed with hairs and one seta on inner edge, second segment outer edge with hairs and inner edge with hairs and one seta, last segment with 3 outer spines (proximal 2 about 1/2 length of distalmost) each with hyaline fringe on distal half of outer margin, terminal to inner margin of segment with 3 finely plumose setae plus hairs on inner margin. Leg 3 (Figure 9d) similar to leg 2 except exopod second segment more elongate and endopod last segment with one less seta. Leg 4 (Figure 10a) basipod with naked seta on outer distal corner; exopod first segment with long outer recurved spinulose spine and inner edge with hairs, second segment with short slender outer spine and one inner seta, last segment with 4 outer to terminal slender, spinulose spines and 3 inner finely plumose setae; endopod first and second segments outer edges with hairs and one long finely plumose seta on inner edge of each, last segment outer edge with few hairs, 2 short outer finely spinulose spines and one long terminal seta (distal half finely spinulose). Leg 5 (Figure 10b) basal segment with naked dorsal seta, last segment with 4 finely spinulose seta (innermost considerably longer than others). Leg 6 (see Figure 8b) represented by 3 naked setae at area of egg sac attachment.

Eggs sacs cylindrical, extending entire length of abdomen.

Male: Body form as in Figure 10c. Total length 0.96 mm, greatest width 0.37 mm. Cephalon somewhat wider than long (0.37 × 0.30 mm). Genital segment (Figure 10d) slightly longer than wide (153 × 142 μm). Abdomen (see Figure 10c) 3–segmented; segments wider than long, measuring 59 × 89 μm, 47 × 74 μm, and 38 × 59 μm, respectively. First abdominal segment with lateral lobate processes anteriorly. Caudal rami (Figure 10e) longer than wide (56 × 24 μm) each ramus bearing one lateral, 2 subterminal and 3 terminal setae (longest 236 μm).

Rostrum (Figure 10f) with ventral sclerotized plate bearing anteriorly directed spinelike process. Cephalic appendages as in female except maxilliped. Maxilliped (Figure 10g) second segment with saddle-shaped process on inner margin, edges of process ornamented with rows of spinules, 2 setae on inner margin of second segment; claw with heavily serrate inner margin, tip somewhat expanded and serrate as indicated in the figure.

Legs 1–4 similar to female except exopods lack sensillae at bases of outer spines and exopod segments relatively shorter. Leg 2 illustrated by Figure 10h. Leg 5 basically as in female, innermost terminal seta broken off on both specimens. Leg 6 absent.

ETYMOLOGY.—The Latin epigri (“wooden peg”) alludes to the sensillae at the bases of the outer exopod spines of female legs 2 and 3.

REMARKS.—This is a common parasite of Synodus englemani and found throughout most of its range, except, as in M. vulgaris, it has not been collected in the southernmost host range in the Indian Ocean or easternmost Pacific. The female can be easily separated from M. vulgaris females by the 2 sensillae at the bases of the outer exopod spines of legs 2 and 3 of M. epigri. The female can be separated from M. synodi females by the much longer setae on the fifth legs of M. epigri.

Metataeniacanthus conepigri, new species

FIGURES 11–12, 48

MATERIAL EXAMINED.—Holotype art (USNM 171311) and 6 art paratypes (USNM 171312) from the gills of Synodus jaculum Russell and Cressey collected at Palau Island, 26 August 1956. One collection containing 2 art from the pseudobranch of S. jaculum collected at the Comores Islands. One collection containing 21 art from the gills and pseudobranch of S. jaculum collected at Mahé, Seychelles. Two collections each containing 1 art from the gills of S. jaculum collected in the South China Sea. One collection containing 4 art from the gills of S. jaculum collected at Blanche Bay, New Britain. One collection containing 1 art from the pseudobranch of S. jaculum collected near Negros Island, Philippines. One collection containing 3 art from the pseudobranch of S. jaculum collected at Palau Islands, 6 April 1974. One collection containing 2 art from Queensland, Australia.

Female: Body form as in Figure 11a. Total length 1.79 mm, greatest width 0.42 mm (measured at widest part of cephalon). Genital segment and area of egg sac attachment (Figure 11b) wider than long (348 × 124 μm). Abdomen about half body length, unsegmented, cylindrical, without spinules posteriorly. Caudal ramus (Figure 11c) more than twice as long as wide (89 × 35 μm) and armed as in M. epigri except no setae are plumose, longest seta 283 μm.

First antenna, second antenna, oral appendages as in M. epigri. Maxilliped (Figure 11d) hook nearly reaching base of 2 mid-lateral setae.

Legs 1–4 biramous, rami 3–segmented. Leg 1 as in M. epigri. Leg 2 (Figure 11e) similar to M. epigri except exopod spines relatively much shorter and with one less exopod seta. Leg 3 (Figure 12a) as in M. epigri except for one less exopod seta. Leg 4 (Figure 12b) exopod with outer edge of last segment dorsally recurved inward with outer setae directed inward giving segment a twisted appearance; endopod as in M. epigri. Leg 5 (Figure 12c) stouter than M. epigri, setae on terminal segment non-plumose, innermost seta only slightly longer than adjacent seta (in M. epigri this seta is nearly twice the length of the adjacent one). Leg 6 (see Figure 11b) represented by 3 naked setae at area of egg sac attachment.

Egg sacs cylindrical, extending slightly beyond caudal rami.

Male: Unknown.

ETYMOLOGY.—The Latin con (“with”) plus epigri, alludes to the close systematic relationship of the hosts of these 2 parasites.

REMARKS.—This species can be separated from M. vulgaris by the 2 sensillae at the bases of the outer spines on the exopods of legs 2 and 3 of M. conepigri. It can be separated from M. synodi, M. epigri, as well as M. vulgaris by the presence of only 4 setae and 3 spines on the last exopod segment of M. conepigri (other 3 species have 4 setae). It can be further separated from M. epigri by the much longer terminal setae on the fifth leg of M. epigri.

The new species occurs throughout most of its host's range and is most often collected from on or under the pseudobranch.

Metataeniacanthus gibbsi, new species

FIGURES 13–15, 49

MATERIAL EXAMINED.—Holotype art (USNM 171315) and 6 art paratypes (USNM 171316) from the gills of Synodus indicus Day collected off East Africa (10°3′N, 51°15′E) during Cruise 9, Station 449, of the R/V Anton Bruun. Two additional collections containing 12 art collected as above off East Africa (Stations 442 and 465). One collection containing 113 art collected at 3°25′N, 47°14′E, “Meteor” Station 140. One collection containing 4 art collected during Cruise 1, Station 22. (Andaman Sea) as above. One collection containing 2 art from the Gulf of Mannar, Sri Lanka. One collection containing 1 art, 1 art from Trivandrum, India. One collection containing 1 art from Madras, India. One collection containing 2 art collected from Rawley Shoals, West Australia. All above material collected from the gills of Synodus indicus.

Female: Body form as in Figure 13a. Total length 1.34 mm, greatest width 0.28 mm (measured at widest part of cephalon). Cephalon slightly wider than long (0.27 × 0.28 mm). Cephalon folded laterally as in Figure 13b. Free thoraic segments incompletely divided dorsally. Genital segment and area of egg sac attachment (Figure 13c) wider than long (118 × 194 μm), not distinctly separated from abdomen. Abdomen more than half total body length (0.83 mm), unsegmented, cylindrical, and bearing prominent spinules on ventral surface between insertion of caudal rami (Figure 13d). Caudal rami (see Figure 13d) longer than wide (44 × 18 μm), each bearing 6 setae (longest 165 μm), 2 plumose as in figure.

First antenna (Figure 13e) 7–segmented, an aesthete on each of last 2 segments, segments measure 68, 53, 24, 18, 21, 18, and 24 μm long respectively. Second antenna (Figure 13f) armed with spines and setae as in other species of the genus; spinules along posterior edge and terminal process fine and in a single row. Oral area as in Figure 14a. Maxilliped (Figure 14b) with conspicuous hook, recurved at tip; hook nearly reaching base of 2 long setae of basal segment.

Legs 1–4 biramous, rami 3–segmented. Leg 1 (Figure 14c) armed as in other species of genus, interpodal plate with patch of scales along posterior edge. Leg 2 (Figure 14d) and Leg 3 (Figure 14e) similar to M. epigri except present species bears only 4 plumose setae on exopod last segment instead of 5. Leg 4 (Figure 15a) also as in M. epigri except outer exopod spines of present species naked. Leg 5 (Figure 15b) with 3 subequal terminal setae. Leg 6 (see Figure 13c) represented by 3 setae at area of egg sac attachment.

Egg sac extending slightly more than half the length of the abdomen.

Male: Body form as in Figure 15c. Total length 0.97 mm, greatest width 0.30 mm (measured at widest part of cephalon). Cephalon slightly wider than long (301 × 271 μm). Free thoracic segments indistinctly divided dorsally. Genital segment (Figure 15d) longer than wide (254 × 207 μm), posterior corners lobate, conforming to shape of spermatophore within. Abdomen 3–segmented, segments measure length and width respectively 71 × 83 μm, 59 × 77 μm, and 47 × 71 μm; last segment with spinules posteriorly between caudal rami. Caudal rami (Figure 15e) longer than wide (59 × 30 μm), longest seta 124 μm.

First antenna, second antenna, and mouth parts as in female. Maxilliped (Figure 15f) hook with serrate inner edge and macelike tip opposed by heavily ornamented area on basal segment as indicated in figure.

Leg 1 basically as in female except segmentation of exopod (Figure 15g) more distinct. Leg 2 (Figure 15h) armed with spines and setae as in female but lacking sensillae on exopod, exopod spines with heavier spinules than female, and exopod segments relatively shorter than in female. Leg 3 as in leg 2 except one less seta on endopod last segment. Leg 4 exopod similar to legs 2 and 3; endopod (Figure 15i) outer spines with heavier spinules and more spatulate than female. Leg 6 absent.

ETYMOLOGY.—Species is named for Dr. Robert Gibbs, who facilitated the loans of much of the host material examined in this study.

REMARKS.—The female of this copepod can be distinguished from all other female members of the genus by the patch of stout spinules on the abdomen between the bases of the caudal rami.

This parasite was collected from nearly all collections of the host species and, like the host, it has not been collected outside of the Indian Ocean.

Metataeniacanthus pacificus, new species

FIGURES 16–17, 50

MATERIAL EXAMINED.—Holotype art (USNM 171322), and 5 art paratypes (USNM 171323) from the gills of 1 Synodus hoshinonis Tanaka from MacClesfield Bank, South China Sea, 13 June 1964. One collection containing 10 art from the gills of the same host collected from the same area. One collection containing 10 art from the gills of the same host collected in the Arafura Sea (Challenger Expedition).

Female: Body form as in Figure 16a. Total length 1.25 mm, greatest width 0.24 mm (measured at widest part of cephalon). Cephalon slightly longer than wide (0.29 × 0.24 mm) fused with first thoracic segment (bearing leg 1). Thoracic segments 2, 3, and 4 separate (see Figures 16a and 16b). Genital segment and area of egg sac attachment (Figure 16c) wider than long (165 × 71 μm), separated from fifth thoracic segment and abdomen by slight lateral constriction. Abdomen about one-third total body length (0.45 long × 0.18 mm wide), widest anteriorly and tapering posteriorly with patch of spinules between caudal rami (see Figure 16d). Caudal rami (Figure 16d) longer than wide (45 × 18 μm), each ramus with 3 plumose and 3 naked setae, longest seta 236 μm.

First antenna (Figure 16e) 7–segmented (third and fourth segments incompletely divided), one aesthete on each of last 2 segments; segments measure 59, 47, 18, 12, 26, 18, and 29 μm long respectively. Second antenna (Figure 16f) 3–segmented, armed as in other species of genus with spinules along posterior edge of last segment somewhat more widely spaced than other species, except the species to be described from Synodus fuscus. Postantennal hook (Figure 16g) long, slender, recurved apically. Oral area (Figure 17a) as in other species of genus except paragnath distinctly trilobed apically and without medial row of spinules; first maxilla with 2 long and 2 short setae. Maxilliped (Figure 17b) hook short and slightly curved, not reaching to base of lateral setae.

Legs 1–4 biramous, rami 3–segmented. Leg 1 (Figure 17c) armed as in other species of genus; posterior edge of interpodal plate with bilobed spinulose process. Leg 2 (Figure 17d) basipod with outer distal seta; exopod first segment with naked spine on outer distal corner, second segment with 2 long sensillae near base of outer spine one inner seta, last segment with 2 sensillae near base of each of 3 outer spines (Figure 17e), terminal to inner edge with 5 plumose setae; endopod first segment with inner seta, second segment with inner seta, last segment with 3 short outer spines and 3 terminal to inner setae (1 short, 2 long). Leg 3 as in leg 2 except endopod last segment with only 2 terminal setae. Leg 4 (Figure 17f), basipod with outer seta; exopod first segment with slender outer spine, second segment with outer spine and inner seta, last segment with 3 outer spines and 4 terminal to inner setae; endopod first and second segments similar to legs 2 and 3, last segment with 2 short outer spines and one long naked terminal seta. Leg 5 (Figure 17g) basal segment with outer seta, last segment with 4 naked setae (2 outer and 2 terminal, innermost twice length of others). Leg 6 (see Figure 16c) represented by 3 short setae on genital segment.

Egg sac cylindrical, extending beyond tip of abdomen.

Male: Unknown.

ETYMOLOGY.—The Latin pacificus alludes to the parasite collected from S. hoshinonis in the Pacific part of its range only.

REMARKS.—Metataeniacanthus pacificus can be separated from all other species of the genus by the long setiform sensillae at the bases of the outer exopod spines of legs 2 and 3. It is also the only species with the combination of 5 setae and 3 spines on the last exopod segments of legs 2 and 3, and 4 setae and 3 spines on the corresponding segment of leg 4.

This parasite is so far recorded only from its host in the South China Sea and the Arafura Sea and appears to be restricted to Synodus hoshinonis occurring in the western Pacific Ocean. Metataeniacanthus nudus, described below, parasitizes Indian Ocean members of the same host species.

Metataeniacanthus nudus, new species

FIGURES 18–19, 50

MATERIAL EXAMINED.—Holotype art (USNM 171320), and 8 art paratypes (USNM 171321) from the area of the pseudobranch of 6 Synodus hoshinonis Tanaka (USNM 217772) collected from the Andaman Sea during Cruise I of the R/V Anton Bruun, Station 21 (9°54′N, 97°42′E), 24 March 1973. Two additional collections containing 3 art from the same site on the same species of host from the Andaman Sea (10°37′N, 97°34′E and 13°28′N, 97°19′E), R/V Anton Bruun, Cruise I, stations 22 and 37.

Female: Body form as in Figure 18a. Total length 2.94 mm, greatest width 0.65 mm (measured at widest part of cephalon). Cephalon wider than long (0.65 × 0.48 mm), fused with first thoracic segment. Second, third, and fourth thoracic segments free, each narrower than the segment anterior to it. Fifth segment incompletely separated from genital segment. Genital segment and area of egg sac attachment (Figure 18b) wider than long (289 × 89 μm), incompletely separated from abdomen. Abdomen less than half total length, 4–segmented, segments measuring 342, 242, 89, and 94 μm long respectively, last segment with short ventral row of spinules near insertion of caudal rami. Caudal rami (Figure 18c) longer than wide (48 × 32 μm), armed with one lateral, one subterminal, and 4 terminal setae, middle 2 with short plumosities.

First antenna (Figure 18d) 7–segmented, armed as in other members of genus; segments measure 53, 74, 30, 41, 65, 56, and 63 μm long; respectively. Second antenna (Figure 18e) 3–segmented, armed with spines and setae as in other members of genus. Oral area as in Figure 18f. Mandible as in other members of genus. Paragnath with weakly lobate tip. First maxilla bearing 2 long and 2 short setae. Second maxilla as in other members of genus. Maxilliped (Figure 18g) base with 2 inner, short setae, terminal hook long, reaching beyond base of 2 inner setae, tip strongly recurved.

Legs 1–4 biramous; rami 3–segmented. Leg 1 (Figure 19a) armed with spines and setae as in all other species of genus. Leg 2 (Figure 19b) armed with spines and setae as in M. vulgaris, spines on last endopod segment more prominent than in other species. Leg 3 as in leg 2 except one less seta on endopod last segment. Leg 4 (Figure 19c) exopod spines and setae as in M. vulgaris; endopod last segment with 2 prominent spines and 1 seta, seta with short spinules on edge rather than plumosities. Leg 5 (Figure 19d) with 1 lateral, 1 subterminal, and 2 terminal setae, all spinulose, innermost with stout base. Leg 6 (Figure 18b) represented by 3 setae at area of egg sac attachment.

Egg sac short, extending only to end of second abdominal segment.

Male: Unknown.

ETYMOLOGY.—The Latin nudus (“naked”) alludes to the lack of sensilla at the bases of the outer exopod spines of legs 2–4.

REMARKS.—This species can be separated from all other Metataeniacanthus by the lack of sensillae at the bases of the outer exopod spines of legs 2–4. Furthermore, it is unique within the genus in that the maxilliped claw extends well beyond the bases of the setae on the first maxilliped segment.

This species, collected from Synodus hoshinonis in the Andaman Sea, has same host species as M. pacificus. The two parasites, however, are geographically separated by the Malay archipelago. Examination of specimens of S. hoshinonis from both geographic areas, each with its own Metataeniacanthus species as a parasite, are inconclusive, and they are considered here as one species. The Indian Ocean population of the host extends to East Africa and the Red Sea. The parasite (M. nudus), however, has so far only been collected in the Andaman Sea.

Metataeniacanthus indiscretus, new species

FIGURES 20–22, 51

MATERIAL EXAMINED.—Holotype art (USNM 171317), allotype art (USNM 171318) and 21 art 1 art paratypes (USNM 171319) collected during Cruise 1 of the R/V Anton Bruun, Station 22 (10°37′N, 97°34′E) from the gills and pseudobranch of Synodus species (undescribed). Three collections containing 22 art from the gills and pseudobranchs of the same host in the South China Sea (16°N, 114°E). One collection containing 3 art from the gills of the same host from Netherlands Indies (Indonesia) by Longley. One collection containing 4 art from the gills of the same host collected in the Andaman Sea during Cruise 1, Station 28 of the R/V Anton Bruun.

Female: Body form as in Figure 20a. Total length 1.28 mm, greatest width 0.29 mm (measured at widest part of cephalon). Cephalon somewhat wider than long (0.29 × 0.25 mm), fused with first thoracic segment. Remaining thoracic segments separate but incompletely divided dorsally. Genital segment about as long as wide and narrower than abdomen. Abdomen more than half total body length and inflated in ovigerous females, unsegmented, cylindrical, no spinules could be seen at posterior tip. Caudal rami small, longer than wide (44 × 24 μm), longest seta 218 μm, armed as in other members of the genus.

First antenna (Figure 20b) 6–segmented, an aesthete on each of last 2 segments, segments measure 33, 27, 25, 21, 18, and 24 μm respectively. Second antenna (Figure 20c) armed as in other members of the genus, last segment with a row of heavy spinules along posterior edge more widely spaced than in most species of the genus; terminal spinulose process with finer spinules. Postantennal hook (Figure 20d) recurved apically, large and generally visible from dorsal aspect. Oral area as in Figure 20e. Spinules along edge of both terminal processes heavy and more widely spaced than in other species of the genus. Maxilliped (Figure 20f) basal segment partly divided near mid-area, proximal part narrower than distal, terminal hook short, not reaching base of lateral setae on basal segment.

Legs 1–4 biramous, rami 3–segmented. Leg 1 (Figure 20g) basipod with stout plumose seta on outer distal corner and another on inner margin; exopod first segment with recurved seta on outer distal corner, second segment with one inner and one outer seta, third segment with 7 inner to outer setae; endopod first and second segments each with one inner seta, third segment with 7 inner to outer setae. Leg 2 (Figure 20h) basipod with naked seta on outer distal corner; exopod first segment with finely spinulose seta on outer distal corner, second segment with an inner plumose seta and on outer finely spinulose seta, third segment with 3 outer to terminal finely spinulose setae and 4 terminal to inner plumose setae, a pair of short sensillae near bases of spinulose setae of second and third segments; endopod first and second segments each with an inner plumose seta, third segment with 3 short, outer spines and 3 terminal setae. Leg 3 (Figure 21a) armed as in leg 2 except one less plumose seta on third segment of exopod and endopod. Leg 4 (Figure 21b) armed as in leg 3 except no sensillae at base of outer exopod setae and only one spine and one seta on terminal endopod segment. Leg 5 (Figure 21c) unsegmented, bearing one outer and 3 terminal setae, ramus weakly sclerotized and quite unlike other known members of the genus. Leg 6 (Figure 21d) represented by 3 short setae at area of egg sac attachment.

Male: Body form as in Figure 21e. Total length 0.65 mm, greatest width 0.23 mm (measured at widest part of cephalon). Cephalon about as long as wide. Genital segment (Figure 22a) longer than wide (135 × 100 μm). Abdomen (see Figure 21e) 3–segmented, segments incompletely divided, measuring 35 × 56 μm, 30 × 53 μm, and 35 × 47 μm length and width respectively. Caudal ramus (Figure 22b) about twice as long as wide (35 × 18 μm), longest seta 130 μm.

First antenna (Figure 22c) armed as in female except aesthete on penultimate segment nearly as long as entire antenna. Second antenna as in female except terminal segment relatively longer. Postantennal hook (Figure 22d) small, pointed. Oral area as in female. Maxilliped (Figure 22e) hook with serrate inner edge and macelike tip opposed by ornamented area on second segment.

Leg 1 as in female. Leg 2 (Figure 22f) armed with spines and setae as in female but lacking sensillae on exopod, segments of exopod relatively shorter than in female. Leg 3 as in leg 2 except endopod last segment with one less seta. Leg 4 (Figure 22g) armed with spines and setae as in female but segments in both rami relatively shorter. Leg 5 (Figure 22h) unsegmented, bearing one lateral and 3 terminal naked setae. Leg 6 absent.

ETYMOLOGY.—The Latin indiscretus (“undivided”) alludes to the fusion of segments of the fifth leg.

REMARKS.—The female of this species can be separated from all known Metataeniacanthus females by the nature of leg 5. In all other species leg 5 is clearly 2–segmented. In M. indiscretus leg 5 is unsegmented.

The host of this parasite is a new species being described separately by the first author.

Metataeniacanthus aquilonius, new species

FIGURES 23–24, 52

MATERIAL EXAMINED.—Holotype art (USNM 171309) 64 paratype art (USNM 171310) collected from the gills of Synodus macrops Tanaka collected off Kochi, Japan, 22 October 1976. One collection containing 1 art from the Gulf of Aden. Two collections containing 5 art from off Somalia, R/V Anton Bruun, Cruise 9, Stations 463, 468. One collection from the Arabian Sea (17°25′N, 70°39′E), Anton Bruun, Cruise 4B, Station 202A. One collection containing 1 art from the Andaman Sea (92°53′E, 11°49′N). One collection containing 1 art from off Hong Kong. One collection containing 1 art from the Formosa Strait (26°N, 121°E). One collection containing 10 art from off Owase, Japan. One collection containing 1 art from the Japan Sea off the Shimane Prefecture. All collections above were from the gills and pseudobranch (Hong Kong coll.) of Synodus macrops.

Female: Body form as in Figure 23a. Total length 2.95 mm, greatest width 0.40 mm (measured at widest part of cephalon). Cephalon slightly wider than long (0.43 × 0.41 mm). Thoracic segments bearing legs 2–5 distinct but not divided dorsally. Genital segment and area of egg sac attachment (Figure 23b) wider than long (0.29 × 0.13 mm), Abdomen more than half total body length (1.67 mm), unsegmented, cylindrical, and bearing fine spinules between insertion of caudal rami (Figure 23c). Caudal rami (see Figure 23c) longer than wide (100 × 35 μm), each bearing 6 setae, longest 2 with short plumosities (longest 277 μm).

First antenna (Figure 23d) 7–segmented, an aesthete on each side of last 2 segments, segments measure 65, 27, 18, 24, 24, 21, and 32 μm long respectively, Second antenna (Figure 23e) tip armed with 3 recurved spines and 3 setae, spinules along edge of last segment somewhat stouter and not as fine as in most other species described here. Postantennal hook (Figure 23f) strongly recurved. Oral area (Figure 23g) basically as in other species; paragnath tip weakly lobed. Maxilliped (Figure 23h) hook slightly recurved, not reaching base of 2 lateral setae.

Legs 1–4 biramous, rami 3–segmented. Leg 1 (Figure 24a) armed as in other species of genus. Leg 2 (Figure 24b) armed with spines and setae as in M. gibbsi; 2 sensillae at bases of exopod spines of last 2 segments, distalmost outer spine of last endopod segment about twice length of proximal 2. Leg 3 (Figure 24c) armed as in leg 2 except one less seta on endopod last segment. Leg 4 (Figure 24d) armed as in leg 3 except sensillae lacking at bases of exopod spines and one less spine and seta on endopod last segment. Leg 5 (Figure 24e) with setae as in other members of genus, innermost only slightly longer than others on free segment, single row of spinules near base of innermost seta. Leg 6 (see Figure 23b) represented by 3 short setae at area of egg sac attachment.

Egg sac not extending to caudal rami.

Male: Unknown.

ETYMOLOGY.—The Latin aquilonius (“north”) alludes to its range in the northern hemisphere of the Indo-West Pacific.

REMARKS.—This species can be separated from all other species of Metataeniacanthus except M. indiscretus by the spine and seta formula (see Table 1). It can be separated from M. indiscretus by the nature of leg 5. The fifth leg of the new species is 2–segmented whereas in M. indiscretus it is one-segmented.

Metataeniacanthus aquilonius has so far been found throughout the range of its host, extending from the Gulf of Aden across the northern Indian Ocean into the West Pacific to Japan. The copepod has been collected from the gills and the pseudobranch of Synodus macrops.

Parataeniacanthus Pillai, 1963

Parataeniacanthus longicervis Pillai, 1963

FIGURES 25, 26

MATERIAL EXAMINED.—13 art 1 art from the gills of Saurida undosquamis (Richardson) from the following localities: R/V Anton Bruun, Cruise 4B, Station 209A (20°49′N, 60°41′E) and Station 216A (21°49′N, 68°55′E), Cruise 1, Station 28A (11°52′N, 92°49′E), R/V Cape St Mary, Cruise 20, South China Sea (16°6′N, 114°41′E); 2 art from the gills of Saurida elongata (Temminck and Schlegel) from Hong Kong.

Female: The female of this species was described by Pillai (1963) and only a few points will be added here to that description. Illustrations of diagnostic features are included so that the reader can identify material of this species without reference to the original description. Body form as in Figure 25a. Total length 4.07 mm. Greatest width 1.16 mm (measured at widest part of cephalon). Caudal rami (Figure 25c) twice as long as wide (248 × 118 μm). Oral area typically taeniacanthid. Maxilliped (Figure 25d) with variable number of teeth (6–9) on prominent claw.

Legs 1–4 biramous, rami 3–segmented. Leg 2 (Figure 25e) as in Pillai except 3 outer spines of endopod fringed on outer edge. Leg 3 (Figure 25f) as in Pillai except very small spinules near base of exopod distalmost outer spine (Pillai notes these on leg 4 only). Leg 4 (Figure 25g) as in Pillai with pair of small spinules near base of each outer spine on segments 2 and 3. Leg 5 (Figures 26a) lamelliform, innermost seta (short claw of Pillai) heavily sclerotized, truncate, spinelike. Leg 6 (see Figure 25b) represented by 3 short setae at area of egg sac attachment.

Male: Body form as in Figure 26b. Total length 3.18 mm. Greatest width 0.83 mm (measured at widest part of cephalon). Cephalon 0.64 mm long. Genital segment slightly wider than long (354 × 324 μm). Abdomen 4–segmented, each segment wider than long, measuring 354 × 277 μm, 313 × 248 μm, 248 × 218 μm, and 195 × 177 μm respectively. Caudal ramus nearly 3 times as long as wide (153 × 57μm), longest seta 678 μm. Appendages as in female except maxilliped (Figure 26d). Maxilliped basal segment with row of rounded teeth, claw with distal double row of teeth.

REMARKS.—This species is apparently an occasional parasite on species of Saurida. Pillai (1963a: 120) reported it from Saurida tumbil (Bloch) but we did not recover it from this host. Unlike Metataeniacanthus, legs 1–4 of both sexes are identical. In Metataeniacanthus the legs of males are more delicate and less modified.

Taeniacanthus Sumpf, 1871

Taeniacanthus sauridae Yamaguti and Yamasu, 1959

FIGURES 27, 28

MATERIAL EXAMINED.—13 art 1 art from gills of 12 Saurida elongata (Temminck and Schlegel) (USNM 136243) collected in the Philippines (1909 Albatross collection), 1 art from the gills of Saurida gracilis (Quoy and Gaimard), 2 art from the gills of Saurida longimanus Norman (USNM 136050 and 136213) also from same collection area as above, 1 art from the gills of Saurida undosquamis (Richardson) collected during Cruise 1 of the R/V Anton Bruun in the Andaman Sea (9°13′N, 97°5′E), 3 March 1963.

Female: The female of this species was adequately described by the authors and no further description will be included here except where differences were noted or to enable the reader to identify this parasite without reference to the original description. Body form as in Figure 27a. First antenna 7–segmented. Maxilliped (Figure 27b) basal segment with 2 setae on inner margin (Y and Y note only 1), 2 short setae near base of claw, claw bifid at tip. Leg 2 (Figure 27c) as in Y and Y but comblike folds on outer margins of exopod segments and last 2 endopod segments seen as sclerotized areas on specimens recorded above. Legs 3 and 4 as in Figures 27d and 27e.

Male: Body form as in Figure 27f. Total length 0.84 mm, greatest width 0.34 mm (measured at widest part of cephalon). Cephalon somewhat wider than long (336 × 271 μm). Thoracic segment bearing leg 1 fused with cephalon. Thoracic segments bearing legs 2–5 free. Genital segment (Figure 28a) wider than long (153 × 106 μm). Abdomen (see Figure 27f) 3–segmented, each segment wider than long and measuring 83 × 41 μm, 77 × 35 μm, and 71 × 41 μm respectively. Caudal rami about twice as long as wide (47 × 20 μm), longest seta 212 μm.

Oral area and mouthparts as in female. Maxilliped (Figure 28b) basal segment with prominent seta near inner mid-margin, 2 rows of knoblike spinules on distal two-thirds with a short row of pointed spinules proximal to knobs; claw with a basal seta and toothed as in the figure.

Leg 1 as in female. Legs 2–4 biramous, rami 3–segmented. Leg 2 (Figure 28c) exopod first segment with slender spine on outer distal corner, second segment with inner plumose seta and outer distal spine, last segment with 3 outer spines and 5 terminal to inner setae; endopod first segment with inner plumose seta, second segment with 2 inner setae, last segment with 3 stout, toothed, outer spines and 3 inner setae. Leg 3 (Figure 28d) exopod as in leg 2 except one less spine and one additional seta on last segment, endopod as in leg 2 except terminalmost toothed spine of last segment much longer than other 2, and 1 less inner seta. Leg 4 (Figure 28e) exopod as in leg 3; endopod first and second segments each with an inner seta, last segment with 2 toothed spines and 1 terminal seta. Leg 5 (Figure 28f) basipod with an outer seta; free segment with one outer lateral and 3 terminal setae, all setae of about equal length (inner 2 slightly longer than outer 2). Leg 6 absent.

REMARKS.—The original description of this copepod was based on the collection of 4 art from the gills of Saurida argyrophanes (Richardson), now considered (Norman) a synonym of Saurida undosquamis, collected from the Inland Sea of Japan. This copepod appears to be an occasional parasite on Indo-Pacific Saurida based on the collections reported here. The females can be easily separated from other species of the genus by the peculiar bifurcate maxilliped claw.

Taeniastrotus Cressey, 1969

Taeniastrotus species

As this paper was going to press we received a collection of what appears to be new species of Taeniastrotus Cressey. The parasites (8 art) were collected from the body surface “between the scales” of Trachinocephalus myops at Trivandrum, India by Mr. A. Chandran of the University of Kerala.

We expect to publish a detailed description of this new species separately.

Abasia Wilson, 1908

DIAGNOSIS.—Female: Frontal lunule absent or present. Rostrum prominent, generally produced beyond anterior margin of first antenna. Lateral margins of carapace folded ventrally. Thoracic area reduced. Genital segment large, comprising about one–third total body length. Abdomen 2–segmented. Caudal rami small, bearing 6 setae, terminal 3 setae about equal in length. First antenna typically caligoid. Second antenna uncinate. First maxilla present (second maxilla of Pillai, 1963b). Second maxilla with stout terminal flagellum (first maxilliped of Pillai, 1963b). Maxilliped robust with well-developed claw (second maxilliped of Pillai, 1963b). Sternal furca absent. Legs 1–4 as in other caligoids but showing varying degrees of reduction.

Male: Frontal lunules present. Lateral margins of carapace not folded ventrally. Second antenna dactylate. Other appendages generally as in female.

TYPE-SPECIES.—Abasia platyrostris Wilson, 1908; gender, f.

DISCUSSION.—The new species of Abasia described herein have characters differing from one another to the extent that some authors might consider them to represent more than one genus. In view of the morphological characters that they have in common, their preference for synodontid fishes, we prefer to consider the group as members of one plastic genus. The reduction of the female third leg is represented in several stages as indicated by Table 2.

TABLE 2.—Segmentation of the third leg exopods and endopods of the species of Abasia

| Exopod | endopo
pseudorostris | 3 | 2
platyrostris | 2 | 2
tripartite | 2 | 1
pusilia | 2 | 0
pillaii | 2 | 0
inflata | 1 | 0

If the reduction were in discrete groups, the case for more than one genus might be supported. Abasia pusilla and A. pillaii are the only 2 with similar segmentation. The close relationship of these 2 species has been pointed out above. Pillai (1963b) points out that although Wilson (1908) states that A. pseudorostris attaches to the roof of the mouth, with the anterior of the cephalon buried in the flesh, the structure of the anterior of the animal does not support this claim. The senior author's experience collecting this Atlantic species is that the copepod is found in the gill area, not buried, and was never collected from the roof of the mouth.

In 1974, Kabata published SEM photos of the mouth tubes of 3 species of caligid copepods. These photos and the diagramatic illustrations demonstrate the presence of a tooth structure, in addition to the mandible, within the mouth tube, near the tip of the labium. He designated this structure as the strigil (p. 1587) and described it in some detail. We found the same structure in SEM photos of the mouth tubes of Abasia species.

Key to Species of Abasia Females

Abasia platyrostris Pillai, 1963

FIGURES 29, 53

MATERIAL EXAMINED.—9 art 3 art from the buccal cavity of 2 Synodus ulae Schultz from Honolulu, Hawaii. One art from the buccal cavity of Saurida undosquamis (Richardson) from the Arabian Sea (20°49′N, 69°41′E) R/V Anton Bruun, Cruise 4B, Station 209A. One art paratype donated to the Smithsonian collections by Dr. Pillai from the buccal cavity of Synodus indicus collected at Trivandrum, India. One art from all Western Australia.

Female: Body form as in Figure 29a. This species was well described by Pillai and only a few additional details are added here. Caudal rami (Figure 29b) longer than wide (100 × 65 μm) armed with plumose setae as in the figure, longest 147 μm. First antenna (Figure 29c) 2–segmented; first segment armed with 23 setae, distalmost setae armed with stout spinules (“conspicuously pectinate” of Pillai, p. 7); second segment armed with 12 setae and 1 aesthete.

Leg 2 (Figure 29d) rami 2–segmented; exopod first 2 segments each with a stout, inwardly directed spine on the outer distal corners; endopod first segment outer distal corner with a row of stout spinules; interpodal plate with a prominent distal fringe. Leg 3 (Figure 29e) rami 2-segmented; exopod last segment with 3 spines and 3 short setae (Pillai shows only 2 setae); endopod segments with a row of stout spinules on outer edges. Leg 4 (Figure 29f) ramus 2-segmented with a spine on the outer distal corner of the first and 3 short spines on the distal end of the second segment, all spines with a short row of spinules near their bases. Legs 5 and 6 each reduced to a single seta on genital segment.

REMARKS.—Pillai (1963b:5), reported this copepod from the buccal cavity of Saurida tumbil (Bloch) and Synodus indicus (Day) all from Trivandrum, India. Kazachenko (1975:212) reports this copepod from the gills of S. tumbil and S. undosquamis (Richardson) from off the coast of northwestern Australia. Our examinations of synodontid fishes resulted in the additional record of the parasite from Synodus ulae Schultz. It seems that this copepod, although probably restricted to synodontid fishes, is an occasional parasite on a number of species within that family. In view of the past unsettled status of Indo-West Pacific Synodus systematics it seems reasonable to treat the record of this parasite from S. indicus as tentative.

Abasia pusilla, new species

FIGURES 30, 31, 39a–e, 53

MATERIAL EXAMINED.—Holotype art (USNM 171333), allotype (USNM 171334) and 5 paratypes (USNM 171335), 2 art, (1 art prepared for SEM study) from the gills of 5 Synodus ulae from Wakanoura, Japan. Three art from 1 Synodus ulae from Misaki, Japan; 1 art from the gills of Synodus ulae from Kochi, Japan; 1 art from the gills of Synodus ulae from Japan: and 1 art from Okinawa.

Female: Body form as in Figures 30a and 30b. Total length 4.96 mm, greatest width 2.23 mm. Cephalon slightly longer than wide (1.77 × 1.58 mm) with lateral margins folded ventrally to encompass host gill filament. Thoracic segments indistinctly separated from cephalon. Genital segment wider than long (2.23 × 1.57 mm), globose. Abdomen unsegmented, somewhat longer than wide (0.62 × 0.51 mm), constricted in posterior third, probably indicating fused segmentation. Caudal rami (Figure 30c) longer than wide (118 × 71 μm) each bearing 6 plumose setae, longest 165 μm.

Rostrum projecting forward with indistinct frontal lunules (SEM photo. Figure 39a, indicates lunule is well developed but difficult to see with light microscope) and bearing a well-defined rugose area ventrally at mid point of anterior margin (Figure 39b). First antenna (Figure 30d) of usual caligid type, first segment with 24 sparsely plumose setae, second segment with 12 setae and 1 aesthete. Second antenna (Figure 30e) with robust basal segment and a short claw bearing 2 short setae. Tip of mandible (Figure 30f) bearing 12 teeth. First maxilla (Figure 30g) as in the figure. Second maxilla (Figure 30h) second segment with a prominent spine on inner distal corner (Figure 39c), terminal claw with rows of spinules as indicated in the figure and and Figure 39d. Maxilliped (Figure 30i) with stout base and short, recurved, terminal claw.

Leg 1 (Figure 30j) biramose; basipod with an inner and an outer seta; exopod 2–segmented, first segment with a short spine on outer distal corner, second segment with 4 terminal setae, outer 2 with lateral spinules, innermost with short plumosities, remaining seta naked; endopod reduced, bearing one, short, terminal seta. Leg 2 (Figure 31a) coxopod with broad hyaline membrane along outer edge; basipod with short seta on outer edge and broad hyaline membrane along inner margin; exopod 3–segmented, first segment with broad hyaline membrane on outer edge, a long finely spinose spine on outer distal corner, and an inner seta, second segment with a long finely spinose spine on outer distal corner (Figure 39e) and an inner seta, last segment with 2 outer spines and 6 setae; endopod 3–segmented, first segment with a row of spinules along outer distal third and an inner seta, second segment with a row of spinules along outer margin and 2 inner setae, last segment with a short row of spinules on outer margin and 6 setae; interpodal plate with a broad posterior hyaline membrane. Leg 3 (Figure 31b) coxopod and basipod forming a lamella, outer distal corner bearing a seta, inner margin bearing a plumose seta near exopod and a broad hyaline membrane to interpodal plate; exopod 2–segmented, first segment heavily sclerotized with stout spine on outer distal corner, second segment with 2 terminal setae; endopod represented by single seta. Leg 4 (Figure 31c) coxopod and basipod indistinctly divided with seta on outer distal corner; exopod 2–segmented, first segment with short, stout spine on outer distal corner, second segment with 2 terminal, short, stout spines; entire leg weakly developed. Legs 5 and 6 absent.

Male: Body form as in Figure 31d. Total length 2.10 mm, greatest width 0.88 mm. Cephalon slightly longer than wide (0.99 × 0.88 mm), lateral margins not folded ventrally; frontal lunules more developed than female. Genital segment (Figure 31e) about as long as wide (0.61 × 0.59 mm), rounded, widest near middle. Abdomen 2–segmented (see Figure 31e), first segment much wider than long (206 × 53 μm), second segment only slightly wider than long (248 × 230 μm). Caudal rami 118 μm long and 71 μm wide, each with 6 plumose setae, 2 short lateral, 3 terminal, 1 subterminal; longest seta 153 μm.

Appendages as in female except as follows. Second antenna with dactylate tip (Figure 31f). Maxilliped with claw longer than in female relative to basal segment. Legs 1–4 as in female. Legs 5 and 6 represented by 2 setae and 1 seta on lateral margins of genital segment as indicated in Figure 31e.

ETYMOLOGY.—The Latin pusilla (“small”) refers to the size of the abdomen when compared to the other species of the genus.

REMARKS.—This species can be separated from the previously described species (A. pseudorostris Wilson and A. platyrostris Pillai) by the reduction of the leg 3 endopod. It can be further separated from these 2 species plus the following 2 new species by the reduced development of the abdomen in A. pusilla.

This species so far is restricted to Synodus ulae from Japan. In 2 collections this species was found with another species of Abasia, described below.

Abasia inflata, new species

FIGURES 32, 33, 39f, 40, 53

MATERIAL EXAMINED.—Holotype art (USNM 171328) and 3 paratype art (USNM 171329) from the gills of Saurida gracilis from the Philippines. One art from the Philippines. One art from Trobriand Islands, New Guinea. One art from New Georgia, Solomons Islands. Four art from Ambon and Kabaena Islands, Molluccas. All other material from the gills of the same host species as the types.

Female: Body form as in Figure 32a–c. Total length 2.25 mm, greatest width 0.96 mm. Cephalon slightly longer than wide (0.90 × 0.73 mm), dorsal surface papillose (Figures 39f, 40a). Genital segment about as long as wide (0.88 × 0.90 mm), wider than cephalon. Abdomen 2–segmented; first segment inflated with other posterior corners produced, extending to or slightly beyond caudal rami, about as wide as genital segment; second segment (Figure 32d) much smaller. Caudal rami (see Figure 32d), wider than long (47 × 35 μm), with 6 plumose setae (longest 94 μm).

Rostrum (Figure 32e) with 2 fleshy lobes projecting beyond first antennae. Frontal lunules absent (not seen with light or scanning electron microscope). Mid-ventral rugose area (Figures 40b,c) present as in previous species. First antenna (Figure 32f) of usual caligoid type; basal segment relatively smaller than in preceeding species, no asthetes seen. Second antenna (Figure 32g) uncinate, claw more curved than in proceeding species. Mouth tube with mandible and strigil (see Kabata, 1974:1587) within (Figure 40d). Mandible (Figure 32h) as in other species of the genus. First maxilla (Figure 32i) simple, with tip bearing 3 setae, one very broad, with spinules on distal half. Second maxilla (Figure 32j) with short distal claw (Figure 40e) bearing rows of spinules, an inner distal spinulose spine, and a palmate process (Figure 40f) on outer edge near base of distal claw. Maxilliped (Figure 32k) as in preceeding species.

Leg 1 (Figure 33a) weakly developed; exopod 1–segmented, bearing a lateral outer spine where segmentation occurs in other species, 4 short terminal setae, armed as in the figure; endopod small, bearing a short distal seta. Leg 2 (Figure 33b) with broad hyaline membranes as in previous species; exopod 2–segmented, first segment with well-developed outer distal spine and an inner seta, last segment with an outer well-developed spine, 2 weak outer spines and 4 terminal to inner plumose setae; endopod 2–segmented (segmentation weak), first segment with an inner plumose seta, last segment with 5 outer to inner plumose setae. Leg 3 (Figure 33c) reduced to a single segment bearing 1 subterminal spine and 2 short terminal spines, longest armed with rows of spinules. Legs 4–6 absent.

Male: Unknown.

ETYMOLOGY.—The Latin inflata (“swollen”) alludes to the 2 lobes on the rostrum.

REMARKS.—This species is characterized by the fleshy lobes of the rostrum and the absence of frontal lunules in the female (males were not found but we suspect that since the rostral area of males is usually less modified a frontal lunule would probably be seen).

To date this species has only been found on the gills of Saurida gracilis and restricted to the western Pacific Ocean.

Abasia tripartita (Shiino, 1955)

FIGURES 34, 35, 41, 42, 43, 44a, b, 53

Alicaligus tripartitius Shiino 1955:56–61.

MATERIAL EXAMINED.—One art and 1 art from the gills of Synodus ulae Schultz from Wakanoura, Japan. One art from the same host from Misaki, Japan. Three collections containing 15 art and 2 art from the gills of Synodus hoshinonis from Japan.

Female: Body form as in Figure 34a,b. Total length 3.55 mm, greatest width 2.04 mm (measured at widest part of genital segment). Cephalon slightly wider than long (1.42 × 1.27 mm), folded laterally. Thoracic segments fused with cephalon dorsally. Genital segment wider than long (2.04 × 1.37 mm), globular in dorsal view but flattened dorsoventrally, dorsal surface papillose (Figure 41a). Abdomen 2–segmented, nearly as long as wide (1.09 × 1.26 mm); first segment encompassing second laterally, distal corners extend beyond caudal rami. Caudal rami (Figure 34c) longer than wide (106 × 54 μm) armed with 2 outer lateral, 3 terminal, and 1 subterminal plumose setae, longest 189 μm.

Rostrum with distinct frontal lunules (Figures 41b–f, 42a) and ventral rugose area (Figure 42b,c). First antenna (Figure 34d) with 2 segments, about equal in length, armed as in figure, an aesthete on last segment. Second antenna (Figure 34e) not as robust as other species of genus; claw recurved, bearing 2 setae. Mouth tube truncate at tip (Figure 42d). Tip of mandible (Figure 34f) bearing 12 teeth. Strigil within mouth tube (Figure 42e,f). First maxilla (Figure 34g) basically as in other species of genus, bearing one long, stout seta and 2 shorter ones. Second maxilla (Figure 34h; Figure 43a,b) armed as in other species, palmate process (Figure 43c) near base of terminal processes small, longest terminal process with rows of spinules. Maxilliped (Figure 34i) stout, armed as in other species of genus.

Leg 1 (Figure 35a) biramous, basipod with an inner and outer seta; exopod 2–segmented, first segment with a short spine on outer distal corner, second segment with 3 spines and a sparsely plumose seta distally (Figure 43d,e): endopod reduced to a short process bearing a small distal knob. Leg 2 (Figure 35b) basipod with an outer naked seta; exopod 3–segmented, first segment with a prominent outer spinulose spine (Figure 43f) and an inner seta, second segment armed as in first, third segment with 3 outer spines (distalmost longest and fringed with a hyaline membrane along outer edge and short plumosities on inner) and 5 setae; endopod 3–segmented, first segment with patch of prominent spinules on outer distal corner (Figure 44a) and an inner seta, second segment with prominent spinules along outer edge and 2 inner setae, last segment with 6 setae. Leg 3 (Figure 35c) basipod lamelliform, bearing a naked seta and patch of setules on outer edge, a short hyaline membrane and plumose seta on inner edge, and a patch of spinules near insertion of interpodal plate; exopod 2–segmented, first segment heavily sclerotized with a stout spine distally, second segment with 4 setae, outer 3 naked and broad, innermost finely plumose; endopod 1–segmented bearing a patch of setules distally and an inner seta. Leg 4 (Figure 35d; Figure 44b) as in A. platyrostris. Legs 5 and 6 absent. Egg strings of usual caligoid type.

Male: Body form as in Figure 35e. Total length 2.30 mm, greatest width 1.20 mm (measured at widest part of cephalon). Cephalon about as long as wide. Genital segment (Figure 35f) somewhat wider than long (543 × 502 μm), widest anteriorly. Abdomen 2–segmented, first segment wider than long (35 × 183 μm), second segment longer than wide (271 × 230 μm). Caudal rami armed as in female, about twice as long as wide (136 × 71 μm), longest seta 295 μm. Frontal lunule and rostral area (Figure 35g) well developed. Appendages as in female except second antenna more dactylate with second segment bearing rugose areas on inner surface (Figure 35h).

REMARKS.—The specimens described here agree with the description by Shiino in 1955 of the new genus and species, Alicaligus tripartitus, from Sarda orientalis in Japan. Considering the diversity within the genus Abasia demonstrated by the species described in this paper, it seems obvious that Alicaligus is also a member of Abasia and we have regarded it here as such. Shiino's reasons for regarding his material as representing a new genus seemed valid at the time, but the additional material described here clarifies the concept of Abasia. The host for Shiino's material must be regarded as accidental. As part of a comprehensive survey of copepods parasitic on scombrid fish, we have examined several specimens of Sarda orientalis and have not recovered A. tripartita. In view of the host preferences of the other species of Abasia, it seems reasonable to regard synodontid fishes as the preferred hosts for A. tripartita as well.

We have changed the specific name, tripartitus, to the feminine form, tripartita, to agree with the feminine Abasia.

Abasia pillaii, new species

FIGURES 36, 37, 44c,d, 53

MATERIAL EXAMINED.—Holotype art (USNM 171330), allotype art (USNM 171331), and 4 art 1 art paratypes (USNM 171332) collected from the gills of 4 Synodus englemani caught during the Albatross expedition in the Philippines. Three art 2 art from the gills of the same host from off Negros Island, Philippines. Three art from the gills of the same host caught off Ambon Island, Moluccas (Amboyna, British Museum 58.4.21.473),

Female: Body form as in Figure 36a,b. Total length 2.47 mm, greatest width 1.23 mm (measured at widest part of genital segment). Cephalon longer than wide, 1.45 × 0.90 mm respectively. Genital segment wider than long (1.23 × 0.90 mm), wider than cephalon. Abdomen 2–segmented; first segment somewhat inflated with distal end extending slightly beyond caudal rami, second segment small and enveloped by first segment anteriorly and laterally. Caudal rami (Figure 36c) small (70 × 53 μm), armed with 6 setae as in other members of the genus, longest seta 153 μm.

Rostrum somewhat produced with frontal lunules well-developed, rugose area on ventral surface as in other species of the genus. First antenna (Figure 36d) first segment with 24 sparsely plumose setae, second segment with 11 setae and 2 aesthetes. Second antenna (Figure 36e) with heavily sclerotized terminal claw bearing 2 short setae near base. Mandible of usual caligoid type with 12 teeth at tip (Figure 36f). Strigil (Figure 44c,d) comblike blade posterior to mandible within mouth tube. First maxilla (Figure 36g) simple, with 3 setae as in other species of the genus. Second maxilla (Figure 36h) with stout terminal claw bearing rows of spinules, a short, spinulose, subterminal spine and a spinose spatulate process near base of terminal spines (Figure 36i). Maxilliped (Figure 36j) basal segment stout with a short recurved claw.

Leg 1 (Figure 37a) basipod with a seta on the outer distal corner and another on posterior margin medial to endopod; exopod 2–segmented, first segment with a weak spine on outer distal corner, second segment with a short outer spine, 2 partially spinulose terminal spines, 1 longer terminal seta and 1 very short inner seta; endopod rudimentary, bearing a short terminal seta. Leg 2 (Figure 37b) armed as in A. pusilla except endopod first segment of A. pillaii with spinules along entire edge (rather than distal third) and no spinules on outer edge of last segment. Leg 3 (Figure 37c) as in A. pusilla except spine on outer distal corner of A. pillaii much heavier and longest terminal seta of last segment spinulose in A. pillaii. Leg 4 similar to that of A. pusilla. Legs 5 and 6 absent. Egg strings of usual caligoid type.

Male: Body form as in Figure 37d. Total length 1.49 mm. Greatest width 0.74 mm. Genital segment (Figure 37e) ovate, as wide as long (406 × 406 μm). Abdomen 2–segmented; first segment small and weakly sclerotized; second segment slightly longer than wide (207 × 171 μm) with sclerotized integument around most of segment. Caudal rami longer than wide (82 × 57 μm) armed as in female.

Appendages as in female except second antenna (Figure 37f) with shorter claw and rugose area on basal segment as indicated in figure. Leg 4 (Figure 37g) weakly developed with a spine on outer distal corner of first segment and 3 spines on the last segment as indicated in the figure. Legs 5 and 6 represented by setae on the lateral margins of the genital segment (see Figure 37e).

ETYMOLOGY.—This species is named for Dr. N. Krishna Pillai as a tribute to his outstanding contributions to the study of Indian Ocean parasitic copepods.

REMARKS.—This species seems to be very close to A. pusilla. The 2 species can be easily separated by the greater development of the genital segment of the female of A. pillaii (compare Figures 30a and 36a). The appendages of these 2 species are very similar and suggest that they are more closely related to each other than to any other species. The new species can be separated from the other known species of Abasia by the nature of the female third leg. In A. pusilla and pillaii the endopod of leg 3 is reduced to a single seta with no discernable ramus. The female leg 3 exopod of A. inflata is further reduced to a single weakly developed ramus representing the entire third leg.

This new species so far has only been found on Synodus englemani caught in the Moluccas and Philippines. It is interesting to note that its closest relative, A. pusilla, has only been reported from S. ulae in Japan.

Lernanthropus Blainville, 1822

Lernanthropus temminckii Nordmann, 1864

FIGURE 38

MATERIAL EXAMINED.—Eight art from the gills of Saurida tumbil (Bloch) collected at Trivandrum, India, October 1977 by Mr. A. Chandran. One art from the gills of Saurida tumbil collected in the Bay of Bengal during Cruise 1 of the R/V Anton Bruun, Station 47B. One art from the gills of Saurida tumbil collected in the Gulf of Thailand, 17 June 1961. One art from the gills of Saurida undosquamis (Richardson) collected off Somalia during Cruise 9 of the R/V Anton Bruun, December 1964.

Female: Body form as in Figure 38. The average total length for 6 of the specimens from Trivandrum is 5.52 mm (4.80–5.85 mm). The total length of the specimen collected off Somalia is 7.95 mm. Caudal rami about 4 times as long as wide (236 × 59 μm), widest proximally, gradually narrowing to tip; 2 well-developed setae in proximal third and 2 small terminal setae (spines?).

Leg 1 exopod with 5 short stout spines; endopod with a single seta about as long as ramus. Leg 2 exopod with 3 terminal spines, inner spine longest; endopod reduced to an unarmed, knoblike process. Legs 3 and 4 produced as long slender processes.

REMARKS.—Nordmann described this copepod (1864:501) from “Saurus lacerta” from the East Indies. It had not been recorded since. It appears to be specific, although not common, to Saurida species and so far only known from the Indian Ocean and South Pacific.

Appendix

Indo-West Pacific Lizardfish Examined and Their Copepod Parasites

(number of fish examined in parentheses)

Synodus englemani Schultz (227)
Metataeniacanthus epigri, new species
Abasia pillaii, new species

Synodus species (undescribed) (8)
Metataeniacanthus indiscretus, new species

Synodus hoshinonis Tanaka (43)
Metataeniacanthus nudus, new species
Metataeniacanthus pacificus, new species
Abasia tripartita (Shiino)

Synodus indicus (Day) (112)
Metataeniacanthus gibbsi, new species

Synodus jaculum Russell and Cressey (62)
Metataeniacanthus conepigri, new species

Synodus macrops Tanaka (70)
Metataeniacanthus aquilonius, new species

Synodus ulae Schultz (60)
Abasia platyrostris Pillai
Abasia pusilla, new species
Abasia tripartita (Shiino)

Synodus variegatus (Lacepède) (654)
Metataeniacanthus vulgaris, new species

Trachinocephalus myops (Schneider) (165)
Metataeniacanthus synodi Pillai
Taeniastrotus species

Saurida elongata (Temminck and Schlegel) (3)
Parataeniacanthus longicervis Pillai
Taeniacanthus sauridae Yamaguti and Yamasu

Saurida gracilis (Quoy and Gaimard) (161)
Taeniacanthus sauridae Yamaguti and Yamasu
Abasia inflata, new species

Saurida longimanus Norman (20)
Taeniacanthus sauridae Yamaguti and Yamasu

Saurida tumbil (Bloch) (55)
Lernanthropus temminckii Nordmann

Saurida undosquamus (Richardson) (103)
Parataeniacanthus longicervis Pillai
Taeniacanthus sauridae Yamaguti and Yamasu
Abasia platyrostris Pillai
Lernanthropus temminckii Nordmann

Synodus binotatus Schultz (131)

Synodus capricornis Cressey and Randall (5)

Synodus doaki Russell and Cressey (10)

Synodus fuscus Tanaka (16)

Synodus kaianus (Gunther) (23)

Synodus rubromarmoratus Russell and Cressey (13)

Synodus sageneus (Waite) (6)

Synodus similis McCullough (5)
No parasitic copepods

Literature Cited

Kabata, Z.

1974.   Mouth and Mode of Feeding of Caligidae (Copepoda) Parasites of Fishes, as Determined by Light and Scanning Electron Microscopy. Journal of the Fisheries Research Board of Canada, 31(10):1583–1588.

Kazachenko, V. N.

1975.   Parasitic Crustaceans (Copepoda) from Fish of Tropical Pacific and Indian Oceans. Bulletin of the Pacific Scientific Research Institute of Fisheries and Oceanography (TINRO), 98:211–217. [In Russian.]

Nordmann, Alexander V.

1864.   Neue Beiträge zur Kenntniss Parasitischer Copepoden. Bulletin de la Société Impériale des Naturalistes de Moscou, 27(4):461–520.

Pillai, N. Krishna

1963a. Copepods of the Family Taeniacanthidae Parasitic on South Indian Fishes. Crustaceana, 6(2):110–128.

1963b. Observations on the Genus Abasia (Copepoda) with the Description of a New Species. Crustaceana, 5(1): 1–9.

Shiino, Sueo M.

1955.   Alicaligus tripartitus gen. et sp. nov. A Caligid Copepod Found on the Gills of Sarda orientalis (T&S). Pacific Science, 9:56–61.

Wilson, Charles Branch

1908.   North American Parasitic Copepods: New Genera and Species of Caliginae. Proceedings of the United States National Museum, 33:596–627.

Yamaguti, Satyu, and T. Yamasu

1959.   Parasitic Copepods from the Fishes of Japan with Descriptions of 26 New Species and Remarks on Two Known Species. Biological Journal of Okayama University, 5(3–4):89–165.

Figures

FIGURE 1.—Metataeniacanthus synodi, female: a, b, dorsal; c, area of egg sac attachment; d, caudal rami; e, first antenna; f, second antenna; g, postantennal hook.

FIGURE 2.—Metataeniacanthus synodi, female: a, oral area; b, paragnath; c, maxilliped; d, first leg and interpodal plate; e, exopod of first leg; f, second leg.

FIGURE 3.—Metataeniacanthus synodi, female: a, last two segments of leg 2 endopod; b, third leg; c, fourth leg; d, fifth leg.

FIGURE 4.—Metataeniacanthus vulgaris, new species, female: a, dorsal; b, area of egg sac attachment; c, caudal rami; d, first antenna; e, second antenna; f, postantennal hook; g, oral area; h, maxilliped.

FIGURE 5.—Metataeniacanthus vulgaris, new species, female: a, first leg; b, second leg; c, third leg; d, fourth leg; e, fifth leg; male: f, maxilliped.

FIGURE 6.—Metataeniacanthus solidus, new species, female: a, dorsal; b, caudal rami; c, first antenna; d, second antenna; e, oral area; f, maxilliped; g, first leg; h, second leg.

FIGURE 7.—Metataeniacanthus solidus, new species, female: a, third leg; b, fourth leg; c, fifth leg; d, sixth leg at area of egg sac attachment.

FIGURE 8.—Metataeniacanthus epigri, new species, female: a, dorsal; b, area of egg sac attachment; c, caudal rami; d, first antenna; e, second antenna; f, oral area; g, maxilliped.

FIGURE 9.—Metataeniacanthus epigri, new species, female: a, first leg; b, second leg; c, second leg, exopod last segment; d, third leg.

FIGURE 10.—Metataeniacanthus epigri, new species, female: a, fourth leg; b, fifth leg; male: c, dorsal; d, genital segment and abdomen; e, caudal rami; f, rostral area, ventral; g, maxilliped; h, second leg.

FIGURE 11.—Metataeniacanthus conepigri, new species, female: a, dorsal; b, area of egg sac attachment; c, caudal rami; d, maxilliped; e, second leg.

FIGURE 12.—Metataeniacanthus conepigri, new species, female: a, third leg; b, fourth leg; c, fifth leg.

FIGURE 13.—Metataeniacanthus gibbsi, new species, female: a, dorsal; b, anterior half, lateral; c, area of egg sac attachment; d, caudal rami; e, first antenna; f, second antenna.

FIGURE 14.—Metataeniacanthus gibbsi, new species; female: a, oral area; b, maxilliped; c, first leg and interpodal plate; d, second leg; e, third leg.

FIGURE 15.—Metataeniacanthus gibbsi, new species, female: a, fourth leg; b, fifth leg; male: c, dorsal; d, genital segment and abdomen; e, caudal rami; f, maxilliped; g, first leg exopod; h, second leg; i, fourth leg endopod.

FIGURE 16.—Metataeniacanthus pacificus, new species, female: a, dorsal; b, anterior half, lateral; c, area of egg sac attachment; d, caudal rami; e, first antenna; f, second antenna; g, postantennal hook.

FIGURE 17.—Metataeniacanthus pacificus, new species, female: a, oral area; b, maxilliped; c, first leg; d, second leg; e, last segment of second leg exopod; f, fourth leg; g, fifth leg.

FIGURE 18.—Metataeniacanthus nudus, new species, female: a, dorsal; b, area of egg sac attachment; c, caudal rami; d, first antenna; e, second antenna; f, oral area; g, maxilliped.

FIGURE 19.—Metataeniacanthus nudus, new species, female: a, first leg; b, second leg; c, fourth leg; d, fifth leg.

FIGURE 20.—Metataeniacanthus indiscretus, new species, female: a, dorsal; b, first antenna; c, second antenna; d, postantennal hook; e, oral area; f, maxilliped; g, first leg; h, second leg.

FIGURE 21.—Metataeniacanthus indiscretus, new species, female: a, third leg; b, fourth leg; c, fifth leg; d, sixth leg at area of egg sac attachment; male: e, dorsal.

FIGURE 22.—Metataeniacanthus indiscretus, new species, male: a, genital segment and abdomen; b, caudal rami; c, first antenna; d, postantennal hook; e, maxilliped; f, second leg; g, fourth leg; h, fifth leg.

FIGURE 23.—Metataeniacanthus aquilonius, new species, female: a, dorsal; b, area of egg sac attachment; c, caudal rami; d, first antenna; e, second antenna; f, postantennal hook; g, oral area; h, maxilliped.

FIGURE 24.—Metataeniacanthus aquilonius, new species, female: a, first leg; b, second leg; c, third leg; d, fourth leg; e, fifth leg.

FIGURE 25.—Parataeniacanthus lorigicervis Pillai, female: a, dorsal; b, area of egg sac attachment; c, caudal rami; d, maxilliped; e, second leg; f, third leg; g, fourth leg.

FIGURE 26.—Parataeniacanthus longicervis Pillai, female: a, fifth leg; male: b, dorsal; c, oral area including maxilliped.

FIGURE 27.—Taeniacanthus sauridae Yamaguti & Yamasu, female: a, dorsal; b, maxilliped; c, second leg; d, third leg; e, fourth leg; male: f, dorsal.

FIGURE 28.—Taeniacanthus sauridae Yamaguti & Yamasu, male: a, genital segment and abdomen; b, maxilliped; c, second leg; d, third leg; e, fourth leg; f, fifth leg.

FIGURE 29.—Abasia platyrostris Pillai, female: a, dorsal; b, caudal rami; c, first antenna; d, second leg; e, third leg; f, fourth leg.

FIGURE 30.—Abasia pusilla, new species, female: a, dorsal; b, lateral; c, caudal rami; d, first antenna; e, second antenna; f, tip of mandible; g, first maxilla; h, second maxilla; i, maxilliped; i, first leg.

FIGURE 31.—Abasia pusilla, new species, female: a, second leg; b, third leg; c, fourth leg; male: d, dorsal; e, genital segment and abdomen, ventral; f, second antenna.

FIGURE 32.—Abasia inflata, new species, female: a, dorsal; b, lateral; c, lateral; d, second abdominal segment and caudal rami; e, rostrum, ventral; f, first antenna; g, second antenna; h, tip of mandible; i, first maxilla; j, second maxilla; k, maxilliped.

FIGURE 33.—Abasia inflata, new species, female: a, first leg; b, second leg; c, third leg.

FIGURE 34.—Abasia tripartita (Shiino), female: a, dorsal; b, lateral; c, caudal rami; d, first antenna; e, second antenna; f, tip of mandible; g, first maxilla; h, second maxilla; i, maxilliped.

FIGURE 35.—Abasia tripartita (Shiino), female: a, first leg; b, second leg; c, third leg; d, fourth leg; male: e, dorsal; f, genital segment and abdomen, ventral; g, frontal lunule and adjacent rostral area, ventral; h, second antenna.

FIGURE 36.—Abasia pillaii, new species, female: a, dorsal; b, lateral; c, caudal rami; d, first antenna; e, second antenna; f, tip of mandible; g, first maxilla; h, second maxilla; i, tip of second maxilla; j, maxilliped.

FIGURE 37.—Abasia pillaii, new species, female: a, first leg; b, second leg; c, third leg; male: d, dorsal; e, genital segment and abdomen, ventral; f, second antenna; g, fourth leg.

FIGURE 38.—Lernanthropus temminckii Nordmann, female: a, dorsal; b, ventral; c, lateral.

FIGURE 39.—Abasia pusilla, new species, female: a, frontal lunule, × 2475; b, rostrum, ventral, × 1765; c, basal spine of second maxilla, × 5600; d, basal spine and base of distal process on second maxilla, × 1375; e, exopod outer spine of leg 2, × 1680. Abasia inflata, new species, female: f, dorsal surface of carapace, × 2430.

FIGURE 40.—Abasia inflata, new species, female: a, dorsal surface of carapace, × 6075; b, rostrum, ventral, × 3000; c, rostrum, ventral, × 15,000; d, tip of mouth tube, labrum removed, × 3600; e, tip of second maxilla, × 1440; f, process near tip of second maxilla, × 5000.

FIGURE 41.—Abasia tripartita (Shiino), female: a, dorsal surface of genital segment, × 1500; b, cephalon, anterior view, × 165; c, rostral area, ventral, × 300; d, cephalon, ventral view of anterior half, × 270; e, frontal lunule, × 1265; f, frontal lunule, × 1470.

FIGURE 42.—Abasia tripartita (Shiino), female: a, edge of frontal lunule, × 3000; b, rostrum, × 2000; c, detail of rostrum, × 15,000; d, tip of mouth tube, labrum uppermost, × 560; e, tip of mouth tube, labrum removed, × 1430; f, strigil within mouth tube, × 2600.

FIGURE 43.—Abasia tripartita (Shiino), female: a, tip of second maxilla, × 1000; b, spine on distal corner of second maxilla, × 2500; c, process at base of flagellum of second maxilla, × 4700; d, tip of leg 1 exopod, × 1400; e, spines at tip of leg 1 exopod, × 3000; f, exopod spines of leg 2, × 1200.

FIGURE 44.—Abasia tripartita (Shiino), female: a, spines on outer distal corner of endopod first segment of leg 2, × 4000; b, tip of leg 4, × 1200. Abasia pillaii, new species, female: c, mandible and strigil within mouth tube, × 1800; d, mandible and strigil within mouth tube, × 2000.

FIGURE 45.—Distribution of Metataeniacanthus synodi Pillai, 1963, and its host Trachinocephalus myops (Schneider, 1801). (Dots within circles = unparasitized.)

FIGURE 46.—Distribution of Metataeniacanthus vulgaris, new species, and M. solidus, new species, and their host Synodus variegatus (Lacepède, 1803). (Dots within circles = unparasitized.)

FIGURE 47.—Distribution of Metataeniacanthus epigri, new species and its host Synodus englemani Schultz, 1953. (Dots within circles = unparasitized.)

FIGURE 48.—Distribution of Metataeniacanthus conepigri, new species and its host Synodus jaculum Russell and Cressey, 1979. (Dots within circles = unparasitized.)

FIGURE 49.—Distribution of Metataeniacanthus gibbsi, new species and its host Synodus indicus (Day, 1973). (Stars within circles = unparasitized.)

FIGURE 50.—Distribution of Metataeniacanthus pacificus, new species and M. nudus, new species and their host Synodus hoshinonis Tanaka, 1917. (Dots within circles = unparasitized.)

FIGURE 51.—Distribution of Metataeniacanthus indiscretus, new species and its host Synodus species (undescribed). (Dots within circles = unparasitized.)

FIGURE 52.—Distribution of Metataeniacanthus aquilonius, new species and its host Synodus macrops Tanaka, 1917. (Dots within circles = unparasitized.)

FIGURE 53.—Distribution of the 5 known species of Abasia in the Indo-West Pacific.

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