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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 382

Cymothoidae from Fishes of Kuwait (Arabian Gulf) (Crustacea: Isopoda)

Thomas E. Bowman and Inam U. Tareen

SMITHSONIAN INSTITUTION PRESS

City of Washington

1983

ABSTRACT

Bowman, Thomas E., and Inam U. Tareen. Cymothoidae from Fishes of Kuwait (Arabian Gulf) (Crustacea: Isopoda), Smithsonian Contributions to Zoology, number 382, 30 pages, 20 figures, 2 tables, 1983.—Cymothoid isopods, parasites of fishes, are reported from Kuwait and the Arabian Gulf. Six genera (1 new) and 9 species (6 new) are reported, including the new genus Joryma and the new species Joryma sawayah, Anilocra monoma, Nerocila kisra, N. sigani, N. arres, and Catoessa gruneri. Nerocila phaiopleura Bleeker is illustrated and redescribed. Notes and illustrations are given for Cymothoa eremita (Brünnich) and a species of Mothocya that is not identified to species because the single specimen is immature. A key is given to the species of Nerocila having serrate margins on the uropodal endopods. Three species of Agana, A. engraulidis Barnard, A. brachysoma Pillai, and A. tartoor Pillai, are transferred to the new genus Joryma. Agarna malayi Tiwari, which had been transferred to Idusa, is returned to its original genus.

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

Bowman, Thomas E.

Cymothoidae from fishes of Kuwait (Arabian Gulf) (Crustacea, Isopoda)

(Smithsonian contributions to zoology ; no. 382)

Bibliography: p.

Supt. of Docs. no. : SI 1.27:382

1. Cymothoidae—Classification. 2. Fishes—Persian Gulf—Parasites. 3. Crustacea—Classification. 4. Crustacea—Persian Gulf—Classification. I. Tareen, Inam U. II. Title. III. Series.

QL1.S54 no. 382 591s [595.3′72] 83-600096 [QL444.M34]

Contents

Introduction

Acknowledgments

Anilocra Leach, 1818

Anilocra monoma, new species

Nerocila Leach, 1818

Nerocila (Nerocila) phaiopleura Bleeker, 1857

Key to Those Species of Nerocila with Serrate Uropodal Endopods

Nerocila (Emphylia) kisra, new species

Nerocila (Nerocila) sigani, new species

Nerocila (Nerocila) arres, new species

Catoessa Schioedte and Meinert, 1884

Catoessa gruneri, new species

Joryma, new genus

Joryma sawayah, new species

Mothocya Costa, 1851

Mothocya species

Cymothoa Fabricius, 1793

Cymothoa eremita? (Brünnich, 1783)

Discussion

Literature Cited

Requirements for Smithsonian Series Publication

Cymothoidae from Fishes of Kuwait (Arabian Gulf) (Crustacea: Isopoda)

Thomas E. Bowman and Inam U. Tareen

____________

Thomas E. Bowman, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Inam U. Tareen, Kuwait Institute for Scientific Research, Box 36825, Raas, Kuwait.

Introduction

It is not surprising that references to cymothoid isopods are found in some of the earliest works on natural history (e.g., Belon, 1553) for these crustaceans are conspicuous parasites of many fishes, including species that have provided us with food for centuries. Despite the antiquity of these published references, there are many parts of the world where the indigenous cymothoids are poorly known or completely unknown. One of the former regions is the Arabian Gulf, including Kuwait Bay, from which only 2 species of cymothoids have been reported, Ichthyoxenus asymmetrica Ahmed, 1970a, and Nerocila heterozota Ahmed, 1970b. To partly correct this deficiency we report herein on a collection of cymothoids from Kuwait made by the second author, supplemented by collections from Kuwait Bay made by Stephen A. Grabe of the consulting firm, Dames and Moore (marked “SG” in “Material” sections), and specimens from Kuwait submitted by Welton Lee through Ernest W. Iverson (marked “WL” in “Material” sections). Sample not labled “SG” or “WL” were collected by I.U. Tareen. Figures 1 and 2 show the localities where collections were made.

The number listed after each specimen is the length in mm from the anterior margin of the head to the posterior margin of the telson.

All specimens are deposited in the Division of Crustacea, National Museum of Natural History, Smithsonian Institution. (USNM = collections of the former United States National Museum, deposited in the National Museum of Natural History, Smithsonian Institution.)

ACKNOWLEDGMENTS.—We thank Mr. Grabe for making available the Dames and Moore material. Anne C. Cohen assembled and mounted the drawings. The syntype of Nerocila serra was borrowed from the Naturhistoriska Riksmuseet, Stockholm, through the courtesy of Charlotte Holmquist.

Anilocra Leach, 1818

Anilocra monoma, new species

FIGURES 3a–u, 4

MATERIAL.—From Acanthopagrus latus (Houttuyn): Fish market, Jun 1977, USNM 190883, 23 (paratype). From Hilsa ilisha (Hamilton-Buchanan): Doha, 30 Aug 1976, USNM 190881, 32 (holotype). From Nematalosus nasus (Bloch): SG 608 (seine), 5 Oct 1981, USNM 191044, juvenile 19.2 (paratype). From Johnius aneus: Kita, Mar 1977, USNM 190882, 2, 23.5, 46 (paratypes).

FIGURE 1.—Arabian Gulf along the coast of Kuwait, showing localities where cymothoid isopods were collected by Tareen.

FIGURE 2.—Kuwait Bay, showing stations where cymothoid isopods were collected by Grabe; Offshore trawl stations are connected by lines to inshore stations having same station number where seined collections were made; exact position of latter depended upon tide level.

ETYMOLOGY.—From the Greek mono (one) plus oma (suffix signifying tumor), referring to the presence of a swelling on only one margin of the dactyl in pereopods 1–4.

DIAGNOSIS.—Head with nearly straight lateral margins, only slightly concave at level of antennae 1 and 2. Antenna 1 with segment 3 produced anteriorly into rounded lobe. Dactyls of pereopods 1–4 with swelling on anterior margin, without swelling or with only faint suggestion of swelling on posterior margin. Telson broadly rounded, that of smaller specimens longer than wide, with distinct caudomedial lobe; that of larger specimens wider than long, without caudomedial lobe. Uropods slightly longer than telson, rami linear, subequal or endopod slightly longer.

ADDITIONAL DESCRIPTION.—Antenna 1 8-merous; segments 1–3 and 8 each with a few distal setules. Antenna 2 longer and broader than antenna 1, reaching midlength of pereonite 1, 8-merous. Mandible with dense cluster of apical setae on 3rd segment of palp. Maxilla 1 with 4 apical spines. Maxilla 2 with 2 spines on inner and outer lobes. Maxilliped palp with 3 apical spines. Pereopods gradually increasing in size except pereopod 7 much longer than pereopod 6; pereopods 5–7 with a few setules on distal segments. Exopods of pleopods slightly longer than endopods, with partial suture on lateral margin. Endopods of pleopod 2 with simple proximomedial lobe; endopods of pleopods 3 and 4 with complexly folded proximomedial lobes and 3 or 4 pockets on proximal third of posterior surface; endopod of pleopod 5 with very complex proximomedial lobes and surface folds, more so than shown in figure.

FIGURE 3.—Anilocra monoma, new species (a–d, i–k, l–l = 32 mm holotype from Hilsa ilisha; e, g, h = 23 mm paratype from Acanthopagrus latus; f = 46 mm paratype, u = 23.5 mm paratype, both from Johnius aneus: a, habitus, dorsal; b, head and pereon, lateral; c, head and pereonite 1, dorsal; d, head, ventral; e, f, telson and uropods, dorsal; g, antenna 1; h, antenna 2; i, mandible; j, maxilla 2; k, maxilla 1; l, maxilliped; m, apex of palp of maxilliped; n–l, pereopods 1–7; u, pleopod 1. Anilocra leptosoma, from Thailand: v, dactyl of pereopod 1, showing swellings on both margins.

FIGURE 4.—Anilocra monoma, new species (a–d = 23.5 mm paratype from Johnius aneus; e = 23 mm paratype from Acanthopagrus latus): a–d, right pleopods 2–5, posterior; e, uropod.

RELATIONSHIPS.—Only 2 known species of Anilocra have both a produced 3rd segment in antenna 1 and swellings in the dactyls of the anterior pereopods. Anilocra leptosoma Bleeker, known from Indonesia to the Gulf of Suez and South Africa, has a more slender body than A. monoma, the head is more abruptly narrowed at the level of antennae 1 and 2, the anterior pereopods have swellings on both sides, and the telson differs in shape, the sides being parallel for the proximal 2/3, then turning sharply inward and meeting in an angular apex. Anilocra dimediata Bleeker, known from Papua New Guinea, Indonesia, the Philippines, Vietnam, Sri Lanka, and Kerala, India, has a head shape similar to that of A. leptosoma, the posterolateral corners of pleonites 4 and 5 are acutely produced, and the telson is a third longer than wide.

Nerocila Leach, 1818

The Kuwait material of Nerocila is of considerable interest in that 3 of the 4 species are new.One of them is the second species in the heretofore monotypic subgenus Emphylia Koelbel, 1879, recently resurrected (Bowman, 1978). In all 3 of the new species the endopod of the uropod has a serrate lateral margin, a character known only in Nerocila trivitatta Bleeker, 1857, N. serra Schioedte and Meinert, 1881, and N. monodi Hale, 1940.

Nerocila (Nerocila) phaiopleura Bleeker, 1857

FIGURE 5

Nerocila phaiopleura Bleeker, 1857:25–26, pl. 1: fig. 3.

Nerocila phaeopleura Bleeker.—Miers, 1880:467.—Schioedte and Meinert, 1881:13–15, pl. 1: figs. 6, 7.—Gerstaecker, 1882:260.—Nierstrasz, 1915:75, 76, figs. 1, 2; 1918:113, 114, figs. 6, 7; 1931:124.—Barnard, 1925:392; 1936:164, 165, fig. 6a–c.—Chilton, 1926:180, 181, fig. 3a,b.—Monod, 1934:12.—Serène, 1937:69.—Morton, 1974:143 148, pl. 1.—Kensley, 1978:82, fig. 33D.—Bruce, 1982:316318, figs. 1, 4a–c.

MATERIAL.—From Chirocentrus dorab (Forskål): SG, 22 Sep 1981, USNM 191051, 18.4. From Dussumieria acuta (Cuvier and Valenciennes): SG 606 (trawl), 22 Apr 1982, USNM 191056, ovigerous 21.5. From unknown host, SG 610 (seine), 12 Apr 1982, USNM 191053, ovigerous 23.3.

DIAGNOSIS.—Head broadly rounded anteriorly. Antennae 1 widely separated. Pereopods 1–5 with slight swelling on proximal part of extensor margin of dactyl; pereopod 7 with marginal spines. Posteroventral corners of pereonites 1–4 rounded or quadrate, of pereonite 5 quadrate or slightly pointed, of pereonites 6 and 7 pointed but only slightly produced. Coxae broad, produced posteriorly slightly beyond their pereonites. Lateral processes of pleonites 1 and 2 broad, horizontal. Telson round-triangular, slightly wider than long. Uropods with long slender rami; exopod about 1.7 × longer than endopod. Uropod exopods each with dark stripe continued on lateral parts of pleon and posterior pereonites.

FIGURE 5.—Nerocila phaiopleura (a–d, k–o = 21.5 mm from Dussumieria acuta; e–j = 18.4 mm from Chirocentrus dorab): a, habitus, dorsal; b, head and pereon, lateral; c, head, dorsal; d, pleon and telson, dorsal; e, head, ventral; f–h, pereopods 1–3; i, pereopod 6: j, pereopod 7; k–o, pleopods 1–5.

DISTRIBUTION.—A widespread species, recorded from Hong Kong (Morton, 1974; Bruce, 1982); Thale Luang (as Talé Sap), a lagoon on the east coast of the Malay Peninsula, Thailand (Chilton, 1926); Gulf of Siam (Monod, 1934, Serène, 1937); from the East Indies at Jakarta, Java (Bleeker, 1857), and the west coast of Sumatra (Nierstrasz, 1918); the Bay of Bengal (Barnard, 1936); and the coast of Natal, South Africa (Barnard, 1925). The Kuwait specimens extend the known distribution greatly.

HOSTS.—“Spratelle kowale” and “Spratelle tembang” (Bleeker, 1857) (probably species of Spratelloides); Chirocentrus dorab (Forskål) (Barnard, 1925); Dusssumieria acuta Cuvier and Valenciennes (Monod, 1934); Xiphias gladius Linnaeus (Barnard, 1936); Sardinella gibbosa (Bleeker), S. perforata (Cantor), Dussumieria hasselti Bleeker, and Decapterus maruadsi (Temminck and Schlegel), (Morton, 1974); Sardinella zunasi (Bleeker) (Bruce, 1982).

Note that both of the hosts in Kuwait have been reported previously as hosts of N.phaiopleura.

Morton (1974) found a constant position for N. phaiopleura on 4 host species of fishes at Hong Kong. The isopod was always found on the posterior third of the body, overlying the lateral line and facing the head of the fish. This position makes accessible the streak of red muscle underlying the lateral line, which contains much fat and myoglobin. It also allows the isopod's abdomen to be raised for pleopod ventilation without risking dislodgement when the fish's tail beats sideways. In contrast, when infesting Dussumieria acuta at Kuwait, N. phaiopleura was attached behind the operculum and above the pectoral fin.

ETYMOLOGY.—The specific name is derived from the Greek phaios (dusky) plus pleura (side) and refers to the 2 lateral stripes. Bleeker (1857) headed his description with the vernacular “Nérocile à flancs bruns.” The correct latinization would have been phaeopleura rather than Bleeker's phaiopleura, and the former has been used by all authors subsequent to Bleeker. However, ICZN Article 32(a) states that improper latinization is not an inadvertent error, hence phaiopleura is the correct original spelling and must be used for this species.

The remaining species of Nerocila considered in this report are characterized by the serrate margin of the uropodal endopod. A key to these species is given below.

Key to Those Species of Nerocila with Serrate Uropodal Endopods

(* = Kuwait species)

Nerocila (Emphylia) kisra, new species

FIGURES 6–8

MATERIAL (all specimens except the holotype are paratypes).—From Therapon puta (Cuvier and Valenciennes): Nov 1976, USNM 183794, with empty marsupium 24.4, 2 non-ovigerous 25.7, 26.6, ovigerous 28.5; Ras-al-Ard, Jan 1977, USNM 183790, 3 ovigerous 16.5, 17.3, 18.6; SG 615 (trawl), 28 Oct 1981, USNM 191054, 24; SG 602 (seine), 25 Sep 1981, USNM 191052, 18.0. From Scomberomorus species: Doha, 16 Feb 1977, USNM 183795, ovigerous 30.2. From Lutjanus species: Fish market, Apr 1980, USNM 183793, with empty marsupium 22.8; 2 ovigerous 25.0, 30.0. From Polydactylus species, caudal peduncle: Jan 1980, USNM 183792, with empty marsupium 25.0. From Ilisha indica (Swainson), on tail: Mar 1978, USNM 183791, with empty marsupium, anterior body not molted, 21.5. From Helotes sexlineatus (Quoy and Gaimard): Ras-al-Ard, Dec 1976, USNM 183788, ovigerous 28.4 (holotype); USNM 183789, 2 ovigerous 28.2, 28.2. From Acanthopagrus cuvieri (Day): Fish market, Mar 1977, USNM 189266, 2 ovigerous 19.6, 24.0. From Pelates quadrilineatus (Bloch), on lower jaw: SG 605 (seine), 24 Sep 1981, USNM 191047, 18.4; SG 604 (seine), 25 Sep 1981, USNM 191050, ovigerous 19.8. From Kathala axillaris (Cuvier): SG, 31 Oct 1981, USNM 191048, 16. From unknown host: Sep 1981, SG 605 (seine), USNM 191049, 18.8; SG 615 (trawl), USNM 191046, with embryos 23.

ETYMOLOGY.—The name kisra is an acronym derived from the Kuwait Institute for Scientific Research.

DIAGNOSIS.—Head narrowed anteriorly. Antenna 1 with inflated 1st segments nearly in contact. Pereopods without swelling in dactyls; pereopods 3, 6, and 7 with marginal spines. Pereopod 4 with largest dactyl. Posteroventral corners of pereonites 1–4 not produced into points, that of pereonite 5 sometimes slightly produced, that of pereonite 6 usually slightly or moderately produced, that of pereonite 7 slightly to strongly produced. Coxae produced into successively longer pointed processes, always exceeding posteroventral corners of pereonites. Pleotelson 1.2–1.4 × as wide as long, ending in weak caudomedial lobe. Exopod of uropod sublinear, distinctly longer than endopod; endopod with notch on medial margin and moderately coarse to finely serrate lateral margin.

ADDITIONAL DESCRIPTION.—Antenna 1 8-merous; last 5 segments (= flagellum) each with dense posterodistal cluster of esthetes. Antenna 2 11-merous; segments 6–10 each with anterodistal row of fine setae; segment 11 with cluster of apical setae. Mandibular palp with a few scattered setae at apex of 3rd segment. Maxilla 1 with 4 apical spines, one much stouter than others. Maxilla 2 with 2 spines on inner lobe and 1 on outer lobe; lobes covered with fine comb-like scales. Maxilliped distal palp segment with 3 terminal spines and smaller spine at about midlength of medial margin. Pleopods 1–5 with lateral accessory gills on protopod, pleopods 1–4 with medial accessory gills on endopod; lateral accessory gills of pleopods 2–5 bilobed, proximal lobe shown extended in figures, but was folded distally in specimen examined; appendix masculina of pleopod 2 slightly more than half length of endopod; endopods of pleopods 3 and 4 with pocket on posterior surface with lateral opening; endopod of pleopod 5 with complex folds.

VARIATIONS.—Body length varied considerably, from 16.5–30.2 mm for ovigerous females. The degree to which the coxae and posteroventral corners of the pereonites are produced is somewhat variable (Figure 6a,h,g). Pereonite 7 may be distinctly pointed or produced into only a small point. The length and direction of the lateral processes of pleonites 1 and 2 vary (Figure 6i–k), as do the size and spacing of the teeth on the uropodal endopod (Figure 6f–h).

FIGURE 6.—Nerocila kisra, new species (a, c–f= 28.4 mm holotype from Helotes sexlineatus; b = paratype from H. sexlineatus; g, i–k = 22.8 mm paratype from Lutjanus sp.; h = 25 mm paratype from Polydactylus sp.): a, habitus, dorsal; b, head and pereon, lateral; c, head, dorsal; d, head, ventral; e, antenna 1; f, antenna 2; g, h, pereon, lateral; i, j, pleon, dorsal; k, pleon and telson, dorsal.

FIGURE 7.—Nerocila kisra, new species, 28.4 mm holotype from Helotes sexlineatus: a, left mandible; b, apex of maxilla 1; c, apex of maxilla 2; d, left maxilliped; e–k, pereopods 1–7.

FIGURE 8.—Nerocila kisra, new species (a–e = 28.4 mm holotype from Helotes sexlineatus; f = from H. sexlineatus; g, h = paratype from Therapon puta): a–e, pleopods 1–5; f–h, uropod.

Nerocila (Nerocila) sigani, new species

FIGURE 9

Material.—From caudal fin of rabbit fish, Siganus oramin (Block and Scheider): Ras-al-Ard, Dec 1977, USNM 190714, non-ovigerous 18.6 (holotype).

ETYMOLOGY.—From the generic name of the host.

DIAGNOSIS.—Head broadly rounded anteriorly, produced into shelf anterior to antennae 1. Antennae 1 separated by nearly 0.8 length of proximal segment. Pereopods without swelling in dactyls; pereopods 6 and 7 with marginal spines; pereopods 1–5 with subequal dactyls. Posteroventral corners of all pereonites produced into points. Coxae produced into successively longer pointed processes, those of coxae 1–5 subequal to posteroventral processes of pereonites, those of coxae 6, 7 distinctly longer. Pleotelson about 1/3 wider than long, smoothly rounded, with no trace of caudomedial lobe. Exopod of uropod linear, slightly longer than endopod; endopod with deep notch on medial margin and serrate lateral margin.

ADDITIONAL DESCRIPTION.—Antennae 1 and 2 as in N. kisra, except esthete clusters in antenna 1 are more distal than posterodistal. Mouthparts not removed in order to avoid damaging buccal region of single specimen. Pleopods essentially as in N. kisra, with minor differences the taxonomic significance of which cannot be evaluated from the single specimen.

Nerocila (Nerocila) arres, new species

FIGURES 10–12

Material.—From Epinephalus tauvina (Forskål), caudal fin: Seif fish market, Kuwait City, 13 Feb 1977, USNM 189268, 26.2 (paratype). From Acanthopagrus latus (Houttuyn): north of platform, 29 Apr 1977, USNM 189265, ovigerous 29.0 (paratype). From Nemipterus japonicus (Bloch), caudal fin: Jan 1977, USNM 189264, ovigerous 30.5 (holotype). From Nemipterus tolu (Cuvier and Valenciennes), caudal fin: SG 611 (trawl), 12 Apr 1982, USNM 191055, ovigerous 28.4 (paratype).

ETYMOLOGY.—From serra spelled backwards, referring to its similarity to Nerocila serra.

DIAGNOSIS.—Head rounded anteriorly, produced into broad shelf anterior to antennae 1. Antennae 1 separated by 0.2–0.7 length of proximal segment. Pereopods 1–5 with swelling in dactyls (Acanthopagrus and Nemipterus specimens), or only pereopod 5 with swelling (Epinephalus specimen); dactyls of pereopods 1–5 subequal. Pereopod 3 with 3 marginal spines on propus in Nemipterus specimen, without spines in other specimens. Pereopods 6 and 7 with marginal spines. Posteroventral corners of all pereonites produced into points, those of pereonites 1–5 subequal, those of pereonites 6 and 7 progressively longer. Coxae produced into pointed processes subequal to posteroventral processes of pereonites. Pleotelson nearly 1/3 wider than long; posterior end damaged in Epinephalus and Acanthopagrus specimens, with low, broad, caudomedial lobe in Nemipterus specimen. Exopod of uropod sublinear, curving medially, slightly longer than endopod; endopod with deep notch on lateral margin and coarsely serrate medial margin.

ADDITIONAL DESCRIPTION.—Antennae 1 and 2 as in N. kisra, but segments 1 and 2 of antenna 1 may be partly fused and number of segments may differ by 1 or 2. Mouthparts not examined. Pleopods similar to those of N. kisra and N. sigani.

RELATIONSHIPS.—Uropods with serrate endopod margins have been reported in 3 species of Nerocila: N. trivittata Bleeker, 1857, N. serra Schioedte and Meinert, 1881, and N. monodi Hale, 1940. Monod considered N. serra to be a junior synonym of N. trivittata (see Niestrasz, 1931:124, footnote; Hale, 1940:301). Monod's opinion may very well be correct; the color patterns appear to be identical, and there is nothing in Bleeker's brief description and illustrations with few details that conflicts with the account of Schioedte and Meinert. The type-locality of N. trivittata is Amboina, that of N. serra Bangka, both in Indonesia. But Bleeker did not mention some important characters now known to be useful in distinguishing species of Nerocila. Hoping to better characterize N. trivittata and N. serra, we attempted to compare type material of the 2 species. From the Naturhistoriska Riksmuseet we obtained, through the courtesy of Dr. Charlotte Holmquist, one of the 2 syntypes of N. serra used by the authors in preparing the description. Unfortunately, we have not been able to locate Bleeker's single specimen of N. trivittata. It is not in the Rijksmuseum van Natuurlijke Historie, Leiden, as are specimens of Bleeker's other isopod species, and Dr. L.B. Holthuis, who kindly furnished this information, pointed out that N. trivittata was not included in Nierstrasz's (1915) account of the Cymothoidae in the Leiden Museum. It seems likely that the type of N. trivittata is not extant, and the question of the identity of N. trivittata and N. serra cannot be resolved now.

FIGURE 9.—Nerocila sigani, new species, 18.6 mm holotype from Siganus oramin: a, habitus, dorsal; b, habitus, lateral; c, head, ventral; d, antenna 1; e, distal margin of segment 5 of antenna 1 showing cluster of esthetes; f, antenna 2; g–m, pereopods 1–7; n–r, pleopods 1–5; s, right uropod.

FIGURE 10.—Nerocila arres, new species (a, e, f, i, j, k, n, o, p = 30.5 mm holotype from Nemipterus japonicus; b, g = 29.0 mm paratype from Acanthopagrus latus; c, d, h, l, m = 26.2 mm paratype from Epinephalus tauvina): a–c, habitus, dorsal; d, e, head and pereon, lateral; f–h, head, ventral; i, left antenna 1; j, left antenna 2; k, l, pereopod 1; m, n, pereopod 2; o, p, pereopod 3.

FIGURE 11.—Nerocila arres, new species (a, c, e, h, i, k = 30.5 mm holotype from Nemipterus japonicus; b, d, f, g, j, l = 26.2 mm paratype from Epinephalus tauvina): a, b, pereopod 4; c, d, pereopod 5; e, f, pereopod 6; g, pereopod 7, distal segments; h, pereopod 7; i, j, pleopod 1; k, l, pleopod 2.

FIGURE 12.—Nerocila arres, new species (a, c, e, j, k = 26.2 mm paratype from Epinephalus tauvina; b, d, f, h, i = 30.5 mm holotype from Nemipterus japonicus; g = 29.0 mm paratype from Acanthopagrus latus): a, b, pleopod 3; c, d, pleopod 4; e, f, pleopod 5; g–k, uropod.

FIGURE 13.—Nerocila serra, syntype from Bangka Strait, Indonesia, Evertebratavd. Riksmuseum Stockholm, Isopoda 4974: a, habitus, dorsal; b, head, ventral; c, left pereopod 1; d, e, endopods of left and right uropods.

The syntype of N. serra is very similar to N. arres. Swellings are present on the dactyls of pereopods 1, 4, and 5, but not pereopod 3 (pereopod 2 lacks dactyls). Pereopod 3 has a single distal spine on the propus. It differs from N. arres in the uropodal endopod, which is longer and narrower, with a shallower notch on the medial margin, and with smaller serrations on the lateral margin. The body has 3 longitudinal dark stripes, which have persisted in preservative for more than a century. Three of our Kuwait specimens of Nerocila lack pigmentation; the specimen from Nemipterus tolu has 3 stripes as in N. serra, but lighter in shade.

Catoessa Schioedte and Meinert, 1884

Catoessa Schioedte and Meinert, 1884:309, 310. [Type-species, by monotypy, Catoessa scabricauda Schioedte and Meinert, 1884, Adonara I, Lesser Sunda Is., Indonesia. Host unknown. Deposition, Berlin Museum.]

TABLE 1.—Comparison of species of Catoessa

The genus Catoessa was proposed for C. scabricauda, represented by a single ovigerous from Adonara Island, one of the Lesser Sunda Islands east of Flores Island. Schioedte and Meinert's (1884) Latin diagnosis of Catoessa contains almost nothing of generic value, and there is little justification for separating Catoessa from Lironeca on the basis of their diagnosis. However, the narrow pleonite 1 and the form of the pleura of pleonites 2–5 distinguish C. scabricauda and the new species described below from all of the more than 50 species of Lironeca, hence we continue to recognize the little known genus Catoessa as valid.

Catoessa gruneri, new species

FIGURES 14b–n, 15

MATERIAL (all specimens except the holotype are paratypes).—From Leiognathus fasciatus (Lacépède), gills: North of platform, 30 Aug 1976, USNM 191069, with empty marsupium 14.4, transitional with small appendix masculina 15.0; Ras-al-Ard, 30 Aug 1976, USNM 191067, 2 7.6, 8.0, WL, 6.4. From Ilisha indica (Swainson), gills: Doha, 18 Dec 1976, USNM 191066, 3 10.6, 11.5, 12.8. From Therapon puta (Cuvier and Valenciennes), WL, USNM 191070, ovigerous 15 (holotype). From Leiognathus daura: SG 611 (trawl), Apr 1982, USNM 191064, 2 ovigerous 13.2, 13.4, 3 non-ovigerous 11.2, 12.5, 13.0; SG 611 (trawl), Apr 1982, 3 ovigerous 12.7, 13.8, 15.0, 2 non-ovigerous 14.0, 14.2. From unknown host: SG 714 (seine), 31 Oct 1981, USNM 191065, ovigerous 13.5, non-ovigerous 10.8; SG 605 (seine), Sep 1981, USNM 191063, 2 ovigerous 9.3, 10.4, 2 non-ovigerous 9.6, 10.4, 15 4.4, 5.6, 5.6, 5.8, 6.0, 6.0, 6.2, 6.3, 6.4, 6.5, 6.7, 6.7, 7.0, 7.1, 8.0.

ETYMOLOGY.—Named for Hans-Eckard Gruner, Zoologishes Museum der Humboldt-Universität zu Berlin, in appreciation of his noteworthy contributions to the biology of the Isopoda.

DESCRIPTION.—Body symmetrical, sides of pereon nearly parallel. Head pyriform, narrowed anterior to eyes, reflexed ventrally into rounded lobe separating antenna 1. Eyes well developed. Pereonites 1–6 subequal in length; pereonite 7 apparently shorter because of overlapping by pereonite 6. Coxae 2–6 much shorter than their pereonites, coxa 7 only slightly shorter. Pleonite 1 partly to nearly completely overlapped by pereonite 7, narrower than rest of pleon. Pleonites 2–5 about as wide as pereonite 7; epimera directed laterally, narrower than central parts of pleonites, more or less separated from one another by gaps. Telson subtriangular, slightly wider than long.

Antenna 1 reaching anterior margin of pereonite 1, broader than antenna 2, basal segments of antennae 1 widely separated; 3 segments of peduncle successively longer, 5 flagellar segments each with distal row of esthetes. Antenna 2 subequal in length to antenna 1, more slender, 8–10-segmented.

Incisors of right and left mandibles differing in form (Figure 14i,j); 3rd segment of palp with a few distal serrate spines. Maxilla 1 with 4 apical spines. Maxilla 2 with 2 spines on inner lobe and 1 on outer lobe. Maxilliped with 3 spines on apical segment.

FIGURE 14.—Joryma sawayah, 12.5 mm paratype from Ilisha indica: a, pleon and telson, dorsal. Catoessa gruneri (b–f = 14.4 mm paratype from Leiognathus fasciatus; g–n = 15 mm holotype from Therapon puta): b, habitus, dorsal; c, habitus, lateral; d, head and pereonite 1, dorsal; e, pleon and telson, dorsal; f, head, ventral; g, right antenna 1; h, right antenna 2; i, left mandible; j, incisor of right mandible; k, maxilla 1; l, maxilla 2; m, maxilliped; n, pereopod 1.

FIGURE 15.—Catoessa gruneri, new species, 15 mm holotype from Therapon puta: a, pereopod 2; b, pereopod 4; c, pereopod 7; d, pleopod 1; e, pleopod 2; f, pleopod 5.

Pereopods with rather broad bases. Pereopods 1–6 subequal. Pereopod 7 distinctly longer, with elongate ischium; ischium and merus notably broader than carpus and propus. Pleopods without accessory gills or surface folds; peduncles of pleopods 1–4 each with 4 retinacula.

Uropods reaching slightly beyond telson; rami sublinear, narrowing slightly distally, exopod slightly longer than endopod.

RELATIONSHIPS.—Until now the genus Catoessa has been known only from the type-species, C. scabricauda, based on a single ovigerous . The most obvious differences between C. scabricauda and C. gruneri are summarized in Table 1.

Joryma, new genus

Agarna Schioedte and Meinert 1884:328–329 [partim].—Barnard 1936:169.—Pillai 1964:211.

DIAGNOSIS.—Pereonite 1 produced into lobe along 1 or both lateral margins of head. Coxae of pereonites 2 and 3 inflated, much larger than remaining coxae; area medial to coxa may be inflated also, resembling dorsolateral bosses (“ovarian bosses”) of epicarideans. Pleon nearly as wide as pereon; pereonite 1 partly or completely covered by pereonite 7. Antennae 1 well separated at bases. Mandibular palp enlarged, very stout, indistinctly or non-segmented.

TYPE-SPECIES.—Joryma sawayah, new species.

COMPOSITION.—Agarna engraulidis Barnard, 1936, Agarna tartoor Pillai, 1954, Agarna brachysoma Pillai, 1954.

ETYMOLOGY.—In the tradition of William Elford Leach, who formed a number of flabelliferan generic names from anagrams of Caroline and Carolina, the name of our new genus is an anagram of Mary Jo, wife of the first author. Gender feminine.

RELATIONSHIPS.—The inflated anterior coxae and especially the mandibular palps will immediately identify Joryma. In Agarna the posterior pereon is much elevated mid-dorsally, and pereonites 4–7 are greatly expanded laterally; antennae 1 are narrowly separated at their bases, and the mandibular palp is not enlarged. Richardson (1905) made the curious statement that pereonite 4 has 2 coxae and 2 pereopods and pereonite 7 has no appendages. This statement is erroneous; pereonites 4 and 7 each have 1 pair of coxa and 1 pair of pereopods, although overlapping of coxae and lateral pereonite expansions tends to obscure the arrangement. Pillai (1954) described Agarna tartoor as lacking coxa 7, and in 1964 he stated that coxa 7 was absent and pereonite 4 had 2 coxae.

Despite our respect for the quality of Pillai's taxonomic contributions, and although we have not seen the specimens he described, we are firmly convinced that Pillai made the same mistake as Richardson, and that his Agarna tartoor and A. brachysoma have normal arrangements of coxae. Furthermore, Pillai considered the coxal arrangement to have generic value, for he transferred Agarna malayi Tiwari, 1953, to Indusa (lapsus for Idusa as in Richardson 1904, 1905) saying, “That Tiwari's species does not show this [distribution of coxae] is sufficient proof that it does not belong to Agarna.” The important generic feature of Agarna, however, is the hunched asymmetrical pereon, with the posterior pereonites flattened and expanded laterally on one side. In this feature A. malayi is similar to the type-species, A. carinata Schioedte and Meinert 1884, and should be returned to its original genus.

The remaining nominal species of Agarna, A. engraulidis Barnard, 1936, was described from off the mouth of the Devi River (∼ 20°N), Orissa State, India. Pillai (1964) transferred it to Livoneca, but it agrees in all respects with our diagnosis of Joryma and is transferred herein to the latter genus.

Joryma sawayah, new species

FIGURES 14a, 16–18

MATERIAL.—From Ilisha indica (Swainson), gills: South of Faylaka, 29 Jul 1976, USNM 191060, ovigerous 21.5, non-ovigerous 21.0, 12.5 (paratypes). Doha, 29 Jul 1976, USNM 191058, ovigerous 20.8 (holotype), USNM 191059, non-ovigerous 17.7 (paratype); North of platform, 29 Jul 1976, USNM, 191061, ovigerous 22.3, 13.3 (paratypes), USNM 191057, ovigerous 22.3, 13.4 (paratypes). “Gord of Oha,” WL, 22 Jan 1978, ovigerous 21.8. From Therapon puta (Cuvier and Valenciennes): WL, ovigerous 19.0.

ETYMOLOGY.—“Sawayah” is the local name and the host fish, Ilisha indica.

DESCRIPTION.—Body slightly asymmetrical, curving slightly to one side, flattened dorsoventrally. Head pyriform, posterior part covered by pereonite 1, anteriorly not reflexed but produced into short shelf in front of antenna 1. Eyes moderately well developed, but usually covered by pereonite 1 and not visible dorsally. Pereonite 1 produced on each side into inflated bilobed process reaching well beyond anterior margin of head. Coxa 2 much enlarged and inflated, produced anteriorly into lobe underlying process of pereonite 1. Coxa 3 inflated, shorter than lateral margin of pereonite 3. Coxae 4–7 occupying anterior halves of pereonite margins; anterior halves of pereonites 4–7 overlapped laterally by preceding pereonites, giving impression that coxa belongs to pereonite anterior to its own. Posterolateral parts of pereonites somewhat inflated. Anterior 1, 2, or 3 pleonites overlapped laterally by pereonite 7. Pleonites as wide as pereon, all subequal in width and length, rather short, lateral parts curving posteriorly. Telson a little wider than long, subtriangular, with distinct caudomedial lobe.

FIGURE 16.—Joryma sawayah, new species (a, b, j, k, l = 20.8 mm holotype from Ilisha indica; c = 12.5 mm paratype from Ilisha indica; d, h = 21.8 mm from Ilisha indica; e, g = 22.3 mm paratype from Ilisha indica; f, i = 19 mm from Therapon puta): a, habitus, dorsal; b, habitus, lateral; c–f, head and pereonite 1, dorsal; g–i, pleon and telson, dorsal; j, antenna 1; k, antenna 2; l, mandible.

FIGURE 17.—Joryma sawayah, new species 20.8 mm holotype from Ilisha indica; a, labrum, dorsal; b, labrum, ventral; c, maxilla 1; d, maxilla 2; e, maxilliped; f–h, pereopods 1–3; i–m, pleopods 1–5; n, uropod.

FIGURE 18.—Joryma sawayah, new species, 13.3 mm from Ilisha indica: a, pleopod 2. Catoessa gruneri, new species (b = 15.0 mm transitional from Leiognathus fasciatus; c = 6.3 mm from unknown host): b, pleon and telson, dorsal, c, pleopod 2.

TABLE 2.—Comparison of species of Joryma

Antennae 1 well separated at base, rather short, reaching midlength of eye, 8-segmented. Antenna 2 slightly longer and distinctly thicker than antenna 1, 8-segmented.

Mandible with very thick palps forming lateral boundaries to other mouthparts, no segmentation evident in palps. Maxilla 1 with 4 strong apical spines. Maxilla 2 inner and outer lobes with minute scales, each with 2 spines. Maxilliped with 7 spines on apical segment.

Pereopods rather short, bases not expanded. Pereopods 1–3 subequal; pereopods 4–7 subequal, longer than pereopods 1–3.

Peduncles of pleopods without retinacula, expanded laterally into rounded lobes. Exopods of pleopods 3–5 with proximolateral triangular lobes. Appendix masculina of pleopod 2 shorter than endopod, nearly straight, tapering gradually to narrowly rounded apex.

Uropods much shorter than telson; rami longer than peduncle, pyriform, exopod longer than endopod.

COMPARISONS.—The most obvious differences between J. sawayah and the other species assigned to Joryma are given in Table 2.

Mothocya Costa, 1851

Mothocya species

FIGURE 19

MATERIAL.—From gill chamber of Strongylura strongylura (van Hasselt): SG, 6 Oct 1981, immature 18.5 mm, 13.7 mm.

Monod (1971) showed convincingly that the supposed differences between Mothocya Costa, 1851, and Irona Schioedte and Meinert, 1884, cannot be substantiated. Hence we consider Irona a junior synonym of Mothocya.

The difficulty in assigning specimens of Mothocya to a species has been pointed out by Monod (1971, 1976). Clear-cut distinctions among the 10 nominal species are rare, and the limits of variability within the species are poorly known. Changes in body proportion associated with sexual maturation further complicate the picture. Since neither Kuwait specimen is fully differentiated, identification to species with confidence is not possible now. When revisionary studies improve our knowledge of the taxonomy of Mothocya, we hope the illustrations given herein (Figure 19) will aid in specific identification of the Kuwait specimens.

Cymothoa Fabricius, 1793

Cymothoa eremita? (Brünnich, 1783)

FIGURE 20

Oniscus eremita Brünnich, 1783:319–325.

Cymothoa eremita.—Trilles, 1975:987–989 [synonymy, distribution, hosts].

MATERIAL.—From Pampus argenteus (Euphrasen): 28 Feb 1979, USNM 191043, with empty marsupium, 39. From Sphyraena obtusata Cuvier: Doha, Mar 1977, USNM 191045, without oostegites, 22.

DISCUSSION.—Several nominal species of Cymothoa, characterized by large size, nearly quadrate head, moderately wide amphicephalic processes of pereonite 1, and relatively short and wide telson, cannot be distinguished from one another at present with confidence. These species include Cymothoa eremita Brünnich, 1783; C. stromatei Bleeker, 1857; C. borbonica Schioedte and Meinert, 1884; C. eximia Schioedte and Meinert, 1884; and C. truncata Schioedte and Meinert, 1884. Cymothoa pulchrum Lanchester, 1902, is similar to the above species, but appears to be distinguishable by the broader amphicephalic processes. The Kuwait specimens are provisionally assigned to the oldest of the above species. The validity of the other species listed above must be considered questionable until revisionary studies based on adequate material can be carried out. Details of their morphology are poorly known or unknown, and a detailed description of the Kuwait specimen would contribute little in support of our provisional identification, hence our account is limited to a few illustrations of salient features. As in Schioedte and Meinert's (1884) specimens, the amphícephalic processes are relatively larger in the larger specimen.

FIGURE 19.—Mothocya species from Strongylura strongylura (a–p = 18.5 mm ; q–r, 13.7 mm ): a, habitus, dorsal; b, habitus, lateral; c, head and pereonite 1, dorsal; d, pleon and telson, dorsal; e–k, pereopods 1–7; l-p, pleopods 1–5; q, pleon and telson, dorsal; r, pleopod 2.

FIGURE 20.—Cymothoa eremita? (a–e = 39 mm from Pampus argenteus; f–j, 22 mm from Sphyraena obtusata): a, habitus, dorsal; b, head and pereopod 1, ventral; c, pereopod 2; d, pereopod 4; e, pereopod 7; f, habitus, dorsal; g, pereon, lateral; h, antennae 1 and 2; i, pereopod 1; j, pereopod 7.

DISTRIBUTION.—Cymothoa eremita is very widely distributed in the Indo-Pacific. Accepting Trilles’ (1975) synonymy, this species ranges from Japan through the Pescadores, the Philippines and Indonesia to the Cape York Peninsula, Australia, and east to the Society Islands. To the west it has been reported from Singapore, Malaysia and Bangkok, Ceylon, the Indian Peninsula from Madras to Bombay, Mauritius, the Seychelles, and the Red Sea. If C. recta Dana (1853) is a synonym, as has been reported, the range is extended to Hawaii. It has not been reported previously from the Arabian Gulf.

HOSTS.—The list of recorded hosts given by Trilles (1975) includes 10 species in 6 genera. Neither Pampus argenteus nor Sphyraena obtusata has been reported as a host for C. eremita, but 7 of the Philippine specimens of Richardson (1910) were accompanied by a label reading, “Parasitic in mouths (gill chamber?) of a barracuda-like fish.”

Discussion

It may seem surprising that 6 of the 9 species of Cymothoidae from Kuwait have not been previously described, considering that about 135 nominal species of this family have been reported from Indo-Pacific waters. However, a recent study by Williams and Williams (1981), demonstrating that what had been thought to be a single species of Anilocra was actually at least 9 species, suggests that there must be many Indo-Pacific species of Cymothoidae yet to be discovered, some of them incorrectly identified as described species.

The Arabian Gulf is so shallow (average depth 35 m) that any exchange of water between it and the adjacent Indian Ocean through the narrow (100 km) Strait of Hormuz is modest (Sverdrup, Johnson, and Fleming, 1946). Nevertheless, the Arabian Gulf has no endemic species of fishes (Blegvad, 1944), and we predict that the new cymothoids described herein will be found in the Indian Ocean proper as well as in the Arabian Gulf. The Arabian Gulf was completely emptied several times during the Pleistocene (Kassler, 1973), and there has not been sufficient time for it to develop endemic species.

Literature Cited

Ahmed, Mahmoud M.

1970a. New Isopoda (Flabellifera) from Iraq and Arabian Gulf, III: Ichthyoxenus asymmetrica sp. nov. Bulletin of the Iraq Natural History Museum (University of Baghad), 4(3):33–36.

1970b. New Isopoda (Flabellifera) from Iraq and Arabian Gulf, IV: Nerocila heterozota sp. nov. Bulletin of the Iraq Natural History Museum (University of Baghdad), 4(3):55–58.

Barnard, K.H.

1925.   Contributions to the Crustacean Fauna of South Africa, 9: Further Additions to the List of Isopoda. Annals of the South African Museum, 20(1):381–412.

1936.   Isopods Collected by the R.I.M.S. “Investigator.” Records of the Indian Museum, 38(2):147–191.

Belon, Pierre

1553.   De Aquatilibus, libri duo: Cum conibus ad viuam effigiem, quoad eius fieri potuit, expressis. 448 pages. Paris.

Bleeker, Pieter

1857.   Recherches sur les Crustacés de l'Inde Archipélagique, II: Sur les isopodes cymathoadiens de l'Archipel Indien. Verhandelingen Natuurkunde Vereeniging Nederlandsch-Indië, 2(5):20–40, plates 1, 2.

Blegvad, H.

1944.   Fishes of the Iranian Gulf. In K. Jessen and R. Spärck, editors, Danish Scientific Investigations in Iran, 3:1–247. Copenhagen.

Bowman, Thomas E.

1978.   Restoration of the Subgenus Emphylia Koelbel for the Parasitic Isopod Nerocila sundaica Bleeker (Flabellifera: Cymothoidae). Crustaceana, 34(1):33–44.

Bruce, Niel L.

1982.   On a Small Collection of Marine Isopoda (Crustacea) from Hong Kong. In B.S. Morton and C.K. Tseng, editors, Proceedings of the First International Marine Biological Workshop: The Marine Flora and Fauna of Hong Kong and Southern China, pages 315–324. Hong Kong: Hong Kong University Press.

Brünnich, Th.

1783.   Den barbugede Pampelfish (Coryphaena apus) en nye Art, og dens Giaest, Strukketrolden (Oniscus eremita) en Opdagelse af Dr. Konig paa Madrass. Nye Samling af det Kongelige Danske Videnskabersnes Selskabs Skrifter, 2:319–325.

Chilton, Charles

1926.   Zoological Results of a Tour in the Far East: The Tanaidacea and Isopoda of Talé Sap. Records of the Indian Museum, 28(3):173–185.

Costa, Achille

1851.   Caratteri di alcuni de'generi e specie nuove segnete nel presente catalogo. In F.W. Hope, Catalogo dei crostacei italiani e di molto altri del Mediterraneo, pages 41–48. Naples.

Dana, James D.

1853.   Crustacea, Part 2. United States Exploring Expedition during the Years 1838, 1839, 1840, 1841, 1842, under the Command of Charles Wilkes, U.S.N., 13:685–1618. Atlas (1855), pages 1–27, plates 1–96. Philadelphia.

Gerstaecker, Adolf

1882.   Isopoda: Wood Lice [part]. In H.G. Bronn, editor, Klassen und Ordnungen des Thier-Reichs…, volume 5 (Arthropoda), part 2, fascicles 4–8, pages 97–278, plates 9–22.

Hale, Herbert M.

1940.   Report on the Cymothoid Isopoda Obtained by the F.I.S. “Endeavour” on the Coasts of the Queensland, New South Wales, Victoria, Tasmania, and South Australia. Transactions of the Royal Society of South Australia, 64(2):283–304, plate 18.

Kassler, P.

1973.   The Structural and Geomorphic Evolution of the Persian Gulf. In B.H. Purser, editor, The Persian Gulf, Holocene Carbonate Sedimentation and Diagenesis in a Shallow Epicontinental Sea, pages 11–32. New York: Springer-Verlag.

Kensley, Brian

1978.   Guide to the Marine Isopods of Southern Africa. 173 pages. Cape Town: South Africa Museum.

Koelbel, Karl

1879.   Über einige neue Cymothoiden. Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Klasse der Kaiserlichen Akademie der Wissenschaften, 78(1):401–416, plates 1, 2.

Miers, Edward J.

1880.   On a Collection of Crustacea from the Malaysian Region, Part IV: Penaeidea, Stomatopoda, Isopoda, Suctoria, and Xiphosura. Annals and Magazine of Natural History, series 5, 5(29):457–472, plate 15.

Monod, Théodore

1934.   Isopodes marins des campagnes du “de Lanessan.” Institut Océanographique de l'Indochine, 23:1–22, plates 1–45.

1971.   Sur quelques Crustacés de Tuléar. Téthys, supplément 1:165–192.

1976.   Crustacés Isopodes, I: Corallanidae, Anilocridae, Cymothoidae. In Th. Monod and R. Serene, editors, Expédition Rumphius II (1975): Crustacés parasites, commensaux,…Bulletin du Muséum National d'Histoire Naturelle (Paris), series 3, 391 (Zoologie 273):853–870.

Morton, Brian

1974.   Host Specificity and Position on the Host in Nerocila phaeopleura Bleeker (Isopoda, Cymothoidae). Crustaceana, 26(2): 143–148, plate 1.

Nierstrasz, H.F.

1915.   Die Isopoden-Sammlung im Naturhistorischen Reichsmuseum zu Leiden, I: Cymothoidae. Zoologische Mededelingen, 1:71–108.

1918.   Alte und Neue Isopoden. Zoologische Mededelingen, 4(2): 103–142, plates 9, 10.

1931.   Die Isopoden der Siboga-Expedition, 3: Isopoda Genuina, 2: Flabellifera. Siboga-Expeditie, 32c:123–232, plates 10, 11.

Pillai, N. Krishna

1954.   A Preliminary Note on the Tanaidacea and Isopoda of Travancore. Bulletin of the Central Research Institute, University of Travancore, Trivandrum, 3C(1): 1–21.

1964.   Parasitic Isopods of the Family Cymothoidae from South Indian Fishes. Parasitology, 54:211–223.

Richardson, Harriet R.

1904.   Contributions to the Natural History of the Isopoda. Proceedings of the United States National Museum, 27(1359):1–89.

1905.   A Monograph on the Isopods of North America. Bulletin of the United States National Museum, 54:i–liii, 1–727.

1910.   Marine Isopods Collected in the Philippines by the U.S. Fisheries Steamer “Albatross” in 1907–08. U.S. Bureau of Fisheries Document, 736:1–44.

Schioedte, J.C. and F. Meinert

1881.   Symbolae ad monographiam cymothoarum crustaceorum isopodum familiae, II: Anilocridae. Naturhistorisk Tidsskrift, series 3, 13:1–166, plates 1–10.

1884.   Symbolae ad monographiam cymothoarum crustaceorum isopodum familiae, IV: Cymothoidae, Trib. II: Cymothoinae, Trib. III: Livonecinae. Naturhistorisk Tidsskrift, series 3, 14:221–454, plates 6–18.

Serène, R.

1937.   Inventaire des invertébrés marins de l'Indochine (ler liste). Notes Institut Océanographique de l'Indochine, 30:1–83.

Sverdrup, H.U., Martin W. Johnson, and Richard H. Fleming

1942.   The Oceans, Their Physics, Chemistry, and General Biology. x + 1087 pages. New York: Prentice-Hall.

Tiwari, Krishna Kant

1953.   On a New Species of the Rare Cymothoid Genus Agarna Schi. & Mein., Parasitic on the Clupeid Fish Nematalosa nasus (Bl.) in the Bay of Bengal. Records of the Indian Museum, 50:295–300, plate 6.

Trilles, Jean-Paul

1975.   Les Cymothoidae (Isopoda, Flabellifera) des collections du Muséum National d'Histoire Naturelle de Paris, III: Les Cymothoinae Schioedte et Meinert, 1884, genre Cymothoa Fabricius, 1787. Bulletin du Muséum National d'Histoire Naturelle (Paris), series 3, 318 (Zoologie 225):977–993.

Williams, Lucy Bunkley, and Ernest H. Williams, Jr.

1981.   Nine New Species of Anilocra (Crustacea: Isopoda:Cymothoidae), External Parasites of West Indian Coral Reef Fishes. Proceedings of the Biological Society of Washington, 94(4):1005–1047.

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Legends for illustrations must be submitted at the end of the manuscript, with as many legends typed, double-spaced, to a page as convenient.

Illustrations must be submitted as original art (not copies) accompanying, but separate from, the manuscript. Guidelines for preparing art may be secured from Series Section, SI Press. All types of illustrations (photographs, line drawings, maps, etc.) may be intermixed throughout the printed text. They should be termed Figures and should be numbered consecutively as they will appear in the monograph. If several illustrations are treated as components of a single composite figure, they should be designated by lowercase italic letters on the illustration; also, in the legend and in text references the italic letters (underlined in copy) should be used: “Figure 9b.” Illustrations that are intended to follow the printed text may be termed Plates, and any components should be similarly lettered and referenced: “Plate 9b.” Keys to any symbols within an illustration should appear on the art rather than in the legend.

Some points of style: Do not use periods after such abbreviations as “mm, ft, USNM, NNE.” Spell out numbers “one” through “nine” in expository text, but use digits in all other cases if possible. Use of the metric system of measurement is preferable; where use of the English system is unavoidable, supply metric equivalents in parentheses. Use the decimal system for precise measurements and relationships, common fractions for approximations. Use day/month/year sequence for dates: “9 April 1976.” For months in tabular listings or data sections, use three-letter abbreviations with no periods: “Jan, Mar, Jun,” etc. Omit space between initials of a personal name: “J.B. Jones.”

Arrange and paginate sequentially every sheet of manuscript in the following order: (1) title page, (2) abstract, (3) contents, (4) foreword and/or preface, (5) text, (6) appendixes, (7) notes section, (8) glossary, (9) bibliography, (10) legends, (11) tables. Index copy may be submitted at page proof stage, but plans for an index should be indicated when manuscript is submitted.