Content uploaded by Jean-Lou Justine
Author content
All content in this area was uploaded by Jean-Lou Justine
Content may be subject to copyright.
1338
Accepted by J. Svavarsson: 22 Sept. 2006; published: 19 Oct. 2006 57
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2006 Magnolia Press
Zootaxa 1338: 57–68 (2006)
www.mapress.com/zootaxa/
Elthusa arnoglossi sp. nov. (Crustacea: Isopoda: Cymothoidae),
a branchial parasite of flatfishes (Bothidae) from the Chesterfield
Islands, New Caledonia
JEAN-PAUL TRILLES¹* & JEAN-LOU JUSTINE²
¹Équipe Adaptation écophysiologique et Ontogenèse, UMR 5171 (UM2, CNRS, IFREMER), GPIA (Génome,
Populations, Interactions, Adaptation), Université Montpellier 2, CP 092, Place Eugène Bataillon, F-34095
Montpellier cedex 05, France. E-mail: jean-paul. trilles@univ-montp2.fr
²Équipe Biogéographie Marine Tropicale, Unité Systématique, Adaptation, Évolution (CNRS, UPMC, MNHN,
IRD), Institut de Recherche pour le Développement, BP A5, 98848 Nouméa Cedex, Nouvelle Calédonie.
E-mail: justine@ird.nc
*Corresponding author
Abstract
The cymothoid isopod Elthusa arnoglossi sp. nov., a branchial parasite of Arnoglossus sp.
(Teleostei, Bothidae) from the Chesterfield Islands, New Caledonia (southwestern Pacific), is
described and figured. The new species is characterised by the asymmetric body of the female,
always twisted to the right side, antenna composed of 18 articles in both sexes, all female pereopods
with a distinctly dilated merus and uropods almost reaching posterior margin of pleotelson in both
sexes; males have the body with pereonites 1–3 distinctly wider than pereonites 4–5 and a well
developed appendix masculina. Within the genus the species is similar only to E. samariscii
(Shiino, 1951), a parasite of Samaris cristatus (Teleostei, Samaridae) from the Indo-Pacific. E.
arnoglossi is the first Elthusa reported on fish from the genus Arnoglossus.
Key words: Elthusa, Isopoda, Cymothoidae, Bothidae, Chesterfield Islands, New Caledonia,
parasites
Résumé
Elthusa arnoglossi sp. nov. (Crustacea: Isopoda: Cymothoidae), parasite branchial de poisson plat
(Bothidae) des Îles Chesterfield, Nouvelle-Calédonie. L’isopode Cymothoidae Elthusa arnoglossi
sp. nov., parasite branchial de Arnoglossus sp. (Teleostei, Bothidae) des Îles Chesterfield, Nouvelle-
Calédonie (Pacifique sud-ouest), est décrit et figuré. La nouvelle espèce est caractérisée par le corps
asymétrique de la femelle, toujours tordu du côté droit, des antennes composées de 18 articles dans
les deux sexes, tous les péréiopodes de la femelle avec un mérus nettement dilaté et des uropodes
qui atteignent presque le bord postérieur du pléotelson dans les deux sexes. Les mâles ont des
péréionites 1–3 nettement plus larges que les péréionites 4–5 et un appendix masculina bien
TRILLES & JUSTINE
58 © 2006 Magnolia Press
1338
ZOOTAXA développé. À l’intérieur du genre l’espèce est proche seulement de E. samariscii (Shiino, 1951),
parasite de Samaris cristatus (Teleostei, Samaridae) de l’Indo-Pacifique. E. arnoglossi est le
premier Elthusa trouvé chez un poisson du genre Arnoglossus.
Introduction
Elthusa Schioedte & Meinert, 1884 has most recently been redefined and revised by Bruce
(1990), and now contains at least 27 nominal species. Most species are known from the
Pacific or from the Indo-Pacific area (Kensley et al. 2006). Only four species have been
reported from the Atlantic and apparently there are no records from the Mediterranean;
several of these species still remain incompletely described and identified, and for many of
them hosts are still unknown (Trilles 1994).
Marine isopods are among the most poorly studied of Crustacea groups in many
regions of the world, particularly in tropical regions. Unsurprisingly, the cymothoid fauna
of New Caledonia which includes the Chesterfield Islands is still largely unstudied. During
the EBISCO cruise (October 2005) arranged by IRD (Institut de Recherche pour le
Développement, France) in the Chesterfield Islands, several specimens of a new species of
Elthusa were collected from flatfishes of the genus Arnoglossus Bleeker, 1862 (Teleostei,
Bothidae).
In this article, we describe all stages (ovigerous and non-ovigerous female, male and
manca larvae 1) of the new species.
Materials and methods
Host fishes were collected by trawls on RV Alis; hosts infected with cymothoids were
selected and immediately fixed in 70% ethanol. Isopods were removed later from the gill
chamber of host fish, measured for total length (TL) and maximum width, and preserved
in 70% ethanol. All measurements are in millimetres. Mouthparts and appendages were
carefully dissected and figures were drawn using a camera lucida.
All specimens are deposited in the collection of the Muséum national d’Histoire
naturelle, Paris (MNHN) as well as the infected fishes. Identification of the hosts was
performed by Dr. M. Desoutter (MNHN). Other fish taxonomy is according to Froese &
Pauly (2006).
Family Cymothoidae Leach, 1814
Genus Elthusa Schioedte & Meinert, 1884
Elthusa Schioedte & Meinert, 1884: 337; Bruce, 1990: 254
© 2006 Magnolia Press 59
ELTHUSA
1338
ZOOTAXA
Type species
Livoneca emarginata Bleeker, 1857, by monotypy (Schioedte & Meinert 1884). One
of Bleeker's type specimens reported and figured by Schioedte & Meinert from the
Rijksmuseum von Natuurlijke Historie, Leiden, is apparently lost and not available (Trilles
1979). One female syntype is held in this museum (Bruce 1990) and one representative
specimen (an ovigerous female probably observed by Schioedte & Meinert (1884) from
Ambon under the gill-cover of an upeneid fish) is held in the MNHN, Paris, under
registration number 241 (Trilles 1976).
Species now included in the genus Elthusa
From the Pacific: Elthusa californica (Schioedte & Meinert, 1884), E. caudata
(Schioedte & Meinert, 1884), E. foveolata (Hansen, 1897), E. frontalis (Richardson,
1910), E. intermedia (Nierstrasz, 1931), E. menziesi (Brusca, 1981), E. myripristae Bruce,
1990, E. neocytta (Avdeev, 1975), E. ochotensis (Kussakin, 1979), E. philippinensis
(Richardson, 1910), E. parabothi Trilles & Justine, 2004, E. puhi (Bowman, 1960), E.
sacciger (Richardson, 1909), E. samoensis (Schioedte & Meinert, 1884), E. sigani Bruce,
1990, E. splendida (Sadowsky & Moreira, 1981), E. turgidula (Hale, 1926); also the new
species described in this work Elthusa arnoglossi sp. nov.
From the Indo-Pacific: Elthusa nanoides (Stebbing, 1905), E. propinqua
(Richardson, 1904), E. raynaudii (Milne-Edwards, 1840), E. samariscii (Shiino, 1951), E.
vulgaris (Stimpson, 1857).
From the Atlantic: Elthusa alvaradoensis Rocha-Ramirez et al., 2005, E. atlantniroi
(Kononenko, 1988), E. methepia (Schioedte & Meinert, 1884) and E. tropicalis (Menzies
& Kruczynski, 1983).
Elthusa arnoglossi sp. nov.
(Figs 1–4)
Material examined
Holotype.
&
(ovig, 11.0 mm), MNHN–Is 5893. Chesterfield Islands, Banc Nova
Nord, EBISCO cruise, RV Alis, station CP 2539, 10 October 2005, 22°20'S, 159°24' E,
depth 315–320 m.
Paratypes. Allotype,
%
(7.0 mm), MNHN–Is 5894; 23
&
(16 ovig, TL 9.0–13.0 mm; 7
non-ovig, TL 10.5–13.0 mm); 2 intermediate stages (8.5–9.0 mm); 4
%
( 6.5, 7.0, 7.5 and
8.5 mm), MNHN–Is 5895, 5896, 5897, 5898. Parasite collection number JNC1639.
Type-host
Arnoglossus sp. The fish specimens could not be identified at the species level and are
either Arnoglossus polyspilus (Günther, 1880) or A. japonicus Hubbs, 1915
(Pleuronectiformes, Bothidae). They are deposited in the collections of the MNHN under
number 2006–0759.
TRILLES & JUSTINE
60 © 2006 Magnolia Press
1338
ZOOTAXA Site of attachment
The female occurs only in the lower gill chamber, attached to the branchial arches,
with the head facing the buccal cavity; the male usually settles beneath the female.
FIGURE 1. Elthusa arnoglossi sp. nov. A–C, holotype,
&
, ovigerous. A, dorsal view; B, lateral
view; C, ventral view; D, allotype,
%
. E, manca larva 1. Scale bars: 2 mm (A–D); 0.5 mm (E).
© 2006 Magnolia Press 61
ELTHUSA
1338
ZOOTAXA
FIGURE 2. Elthusa arnoglossi sp. nov., paratype
&
, ovigerous. A, antennule; B, antenna; C,
maxilliped; D, maxilliped apex; G, maxillule; H, maxilla; J, mandible; K–L, pereopods 1–2; M–O,
pereopods 5–7; paratype
&
, non-ovigerous. E, maxilliped; F, maxilliped, apex; I, maxilla. Scale
bars: 0.5 mm (A, B, C, K, L, M, N, O); 0.2 mm (E); 0.1 mm (F, G, H, I, J); 0.05mm (D).
TRILLES & JUSTINE
62 © 2006 Magnolia Press
1338
ZOOTAXA
FIGURE 3. Elthusa arnoglossi sp. nov., paratype
%
. A, antennule; B, antenna; C, maxilliped; D,
maxillule, apex; E, maxilla; F, maxilla, apex; G, mandible; H–I, mandible palp, apex; J–P,
pereopods 1–7. Scale bars: 0.5 mm (A, B, J, K, L, M, N, O, P); 0.2 mm (G); 0.1 mm (C, E); 0.05
mm (D, F, H, I).
© 2006 Magnolia Press 63
ELTHUSA
1338
ZOOTAXA
FIGURE 4. Elthusa arnoglossi sp. nov., paratype
&
, ovigerous. A–E, pleopods 1–5; F, uropod;
paratype
%
: G–K, pleopods 1–5; L, uropod. Scale bar: 0.5 mm (A–L).
Etymology
The species name is derived from that of the host genus, Arnoglossus.
TRILLES & JUSTINE
64 © 2006 Magnolia Press
1338
ZOOTAXA Description of ovigerous and non-ovigerous females (Figs 1A–C, 2A–O, 4A–F)
Body about 1.5 times as long as wide, widest at pereonite 4, asymmetrical, deeply
twisted to the right side. Pereon slightly vaulted at the level of pereonites 2–4.
Cephalon about 1.6 times as wide as long, roughly triangular, anterior margin acute in
dorsal view, with minute rostral point, posterior margin broadly rounded, not enclosed in
the pereonite 1. Eyes about 0.5 times width of cephalon, ovate, black. Coxal plates of
pereonites 2–7 more or less visible in dorsal view.
Pereon broad, about 1.15 as wide as long, always twisted to the right side. Pereonite 1
longest, 2–4 progressively decreasing in length, 5–7 shortest and subequal in length.
Posterior margin of pereonite 7 deeply curved. Pereonites 3–4 are expanded on the right
side compared to the left.
Pleon about 1.8 as wide as long; all pleonites visible; pleonite 1 shorter and narrower
than others, partially covered by pereonite 7; pleonites 2–5 progressively narrower
towards posterior, 2–4 subequal in length, 5 longest with posterior margin widely
bisinuate. Pleotelson hemispherical, about 1.6 times as wide as long, posterior margin
rounded.
Brood pouch (filled with eggs, embryos or manca larvae) of the ovigerous female is
prominent in lateral view and made up of five pairs of alternatively overlapping oostegites
arising from sternites 1–5, anterior pair partly overlapping mouth parts.
Antennule much shorter than antenna, with 8 articles, reaching posterior of cephalon,
basal articles nearly in contact; Antenna with bases wide apart, composed of 18 articles,
extending to or beyond posterior margin of pereonite 1, proximal article 6 sometimes with
2 plumose setae respectively on the anterodistal and posterodistal angles. Mandible incisor
acute but reduced, molar prominent; palp slender, article 1 widest and article 2 longest,
sometimes with only a single setae on distolateral margin of article 3. Maxillule with 1
large and 3 small spines. Maxilla with 3 blunt spines on lateral lobe in ovigerous female
(Fig. 2H), 2 of which longer, recurved and acute in non-ovigerous female (Fig. 2I).
Maxilliped with oostegial lobe bearing some plumose setae and distal article of palp with 2
blunted spines in ovigerous female (Fig. 2C–D), recurved and acute in non-ovigerous (Fig.
2E–F).
Pereopods almost similar with distinctly dilated merus, humped, decreasing slightly in
size from pereopod 1 to 7, dactyli short and basis without carina. Propodus of all
pereopods short, about as long as combined lengths of merus and carpus; ischium of
pereopods 1–3 about 0.6–0.7 length of basis; ischium and basis of pereopods 4–7 nearly
subequal.
Pleopods with all rami lamellar, decreasing progressively in size, exopod slightly
larger than endopod; all pleopods without coupling hooks on peduncle medial margin and
folding or accessory lobes on endopod.
Uropods short, almost reaching posterior margin of pleotelson, biramous, rami
unequal, exopod shorter than endopod, apices blunted pointed.
© 2006 Magnolia Press 65
ELTHUSA
1338
ZOOTAXA
Description of male (Figs 1D, 3A–P, 4G–L)
Smaller than the female. Body roughly bilaterally symmetrical, about 2.1 as long as
wide, widest at pereonite 3.
Cephalon subtriangular about 1.7 as wide as long, slightly immersed in pereonite 1,
anterior margin in blunted point. Eyes visible, about 0.6 times width of cephalon.
Pereonites 1–3 distinctly longer and wider than 4–7 subequal in length but decreasing
progressively in width. Coxae of pereonite 2–7 more or less visible in dorsal view.
Pleon about 1.6 as wide as long and about 0.45 width of pereon, lateral margins sub-
parallel. All pleonites visible, pleonite 1 shorter than others, 2–4 subequal in length, fifth
slightly longest with posterior margin widely bisinuate. Pleotelson about 0.5 times as long
as wide, subtriangular to posteriorly round.
Antennule and antenna similar to female, but antennule with some very little setae on
distal margin of articles 1–5 and antenna extending to the pereonite 3. Mouthparts similar
to female but mandible palp article 3 sometimes with more setae.
All pereopods sub-similar, without dilated merus, but dactyli 1–3 relatively longest.
Pleopod 2 with appendix masculina, approximately 0.7 as long as endopod.
Uropod rami subequal, bluntly rounded, exopod larger than endopod, extending to
posterior of pleotelson.
Manca larvae 1 (Fig. 1E)
The specimen reproduced in Figure 1E is at the intramarsupial stage 1 (manca larvae 1
or pulli 1) with 7 pereonites but only 6 pereopods and characterized by: enlarged cephalon,
sub-triangular; ovate pereon and narrowed pleon with sub-parallel margins; acute
pleotelson with posterior margin rounded; uropods short, both rami broadly rounded, not
reaching posterior margin of pleotelson.
Colour
Female and male yellowish.
Size range
Ovig
&
: 9.0–13 mm in body length; non-ovig
&
: 10.5-13.0 mm; intermediate stage:
8.5–9.0 mm;
%
: 6.5–8.5 mm.
Remarks
The genus Elthusa Schioedte & Meinert, 1884, briefly defined in Latin by the Danish
authors, was recently clearly redefined by Bruce (1990) who at that time recognized 25
species, including several incompletely described and characterized. Twenty eight species
are now recognized, most species (24) being known from the Pacific or Indo-Pacific area.
Only 4 species are reported from the Atlantic Ocean.
Host identity remains unknown for several species of Elthusa but five species are
TRILLES & JUSTINE
66 © 2006 Magnolia Press
1338
ZOOTAXA known to parasitize pleuronectiform fishes, either specifically [E. parabothi from
Bothidae (Trilles & Justine 2004), E. samariscii from Samaridae (Shiino 1951; Biju
Kumar & Bruce 1997), E. methepia from Achiridae (Schioedte & Meinert 1884)] or
occasionally [E. raynaudii from Pleuronectidae (Chilton 1911), E. vulgaris from
Paralichthyidae (Ho 1975; Brusca 1978)]. E. parabothi was previously reported from
Parabothus kiensis (Tanaka, 1918). So, E. arnoglossi is the second species of Elthusa
described from the host family Bothidae. However, this species is the first cymothoid
collected on gills of flatfishes of the genus Arnoglossus.
Elthusa arnoglossi can be easily distinguished from most of the species of the genus
that have a bilaterally symmetrical body (E. alvaradoensis, E. atlantniroi, E. caudata, E.
menziesi, E. neocytta, E. ochotensis, E. puhi, E. raynaudii, E. sacciger, E. samoensis, E.
splendida, E. tropicalis, E. turgidula, E. vulgaris) or only weakly asymmetrical (E.
californica, E. foveolata, E. frontalis, E. intermedia, E. methepia, E. myripristae, E.
philippinensis and E. parabothi). Four species (E. nanoides, E. propinqua, E. samariscii
and E. sigani) are distinctly asymmetrical. From them, only one species, E. samariscii,
first described by Shiino (1951) and recently redescribed by Biju Kumar & Bruce (1997)
show some similarity to E. arnoglossi. However, E. arnoglossi can be separated from this
species by: female body more asymmetrical, dissimilar in shape and always deeply twisted
only to the right side while E. samariscii is slightly twisted to one side or another (dextral
or sinistral); antenna composed of 18 articles (only 12 for E. samariscii) extending to or
beyond margin of pereonite 1(instead of posterior margin of cephalon); all pereopods with
a distinctly dilated merus lacking in E. samariscii; uropods longer and almost reaching
posterior margin of pleotelson (instead barely reaching beyond anterior quarter of
pleotelson); male body with pereonite 1–3 relatively wider and longer beside others
(character not related to the isopods moulting process); antenna with 18 articles instead of
12; pleopod 2 with appendix masculina distinctly longer and uropods reaching posterior
margin of pleotelson
Acknowledgements
We thank Bertrand Richer de Forges (IRD, Nouméa) who collected the fish, the members
of the EBISCO cruise on RV Alis, Dr. Martine Desoutter, who kindly identified the hosts
and Dr. Danielle Defaye (MNHN, Paris) for their help. Also, we wish to thank anonymous
reviewers and Dr. Jörundur Svavarsson (University of Iceland) for helpful and
constructive comments on the manuscript.
© 2006 Magnolia Press 67
ELTHUSA
1338
ZOOTAXA
References
Avdeev, V.V. (1975) Two representatives of parasitic isopods of the genus Lironeca (Cymothoidae)
from the region of Australia and New Zealand. Parasitologia, 3, 247–251.
Biju Kumar, A. & Bruce, N.L. (1997) Elthusa samariscii (Shiino, 1951) (Isopoda, Cymothoidae)
parasitizing Samaris cristatus Gray, 1831, off the Kerala coast, India. Crustaceana, 70 (7),
780–787.
Bleeker, P. (1857) Recherches sur les Crustacés de l’Inde Archipélagique. II. Sur les Isopodes
Cymothoadiens de l’Archipel Indien. Verhandelingen der NatuurkundigeVereeniging in Neder-
landsch-Indië, Batavia (Acta Societatis Scientiarum Indo-Neêrlandicae, Batavia), 2, 20–40,
pls I–II.
Bleeker, P. (1862) Sur quelques genres de la famille des Pleuronectoïdes. Verslagen Akademie
Amsterdam, 13, 422–429.
Bowman, T.E. (1960) Description and notes on the biology of Lironeca puhi n.sp. (Isopoda:
Cymothoidae), parasite of the Hawaiian moray eel, Gymnothorax eurostus (Abbot). Crusta-
ceana, 1 (2), 84–91, pl.II.
Bruce, N.L. (1990) The genera Catoessa, Elthusa, Enispa, Ichthyoxenus, Idusa, Livoneca and Nori-
leca n. gen. (Isopoda, Cymothoidae), crustacean parasites of marine fishes, with description of
eastern Australian species. Records of the Australian Museum, 42 (3), 247–300.
Brusca, R. (1978) Studies on the cymothoid fish symbionts of the Eastern Pacific (Crustacea, Iso-
poda, Cymothoidae). II. Systematics and Biology of Lironeca vulgaris. Occasional papers of
the Allan Hancock Foundation, New Series, 2, 1–19.
Brusca, R. (1981) A monograph of the Isopoda Cymothoidae (Crustacea) of the eastern Pacific.
Zoological Journal of the Linnean Society, 73, 117–199.
Chilton, C. (1911) Scientific Results of the New Zealand Government Trawling Expedition, 1907.
Crustacea. Records of the Canterbury Museum, 1, 285–312, pl. 58.
Froese, R. & Pauly, D. (2006) FishBase: a global information system on fishes. World Wide Web
electronic publication. Available from http://www. fishbase.org (accessed April 2006).
Günther, A. (1880) Report on the shore fishes procured during the voyage of H. M. S. Challenger in
the years 1873–1876. In: Report on the scientific results of the voyage of H. M. S. Challenger
during the years 1873–76. Challenger Reports, Zoology, 1(6), 1–82.
Hale, H.M. (1926) Review of Australian isopods of the cymothoid group. Part. II. Transactions of
the Royal Society of South Australia, 50, 201–234, pls XXXVI–XXXVII.
Hansen, H.J. (1897) Report on the dredging operations off the West coast of Central America to the
Galapagos, to the West coast of Mexico, and in the Gulf of California, in charge of Alexander
Agassiz, carried on by the U.S. Fish Commission Steamer “Albatross” during 1891, Lieut.
Commander Z.L. Tanner U.S.N., commanding. Bulletin of the Museum of Comparative Zool-
ogy at Harvard College, in Cambridge, 31 (1897–1898), 95–129, pls 1–7.
Hubbs, C.L. (1915) Flounders and soles from Japan collected by the United States Bureau of Fish-
eries steamer "Albatross" in 1906. Proceedings of the United States National Museum, 48, no.
2082, 449–496.
Ho, J-S. (1975) Parasitic Crustacea. Fish Bulletin, 165, 69–72.
Kensley, B., Schotte, M. & Schilling, S. (2006) World list of marine, freshwater and terrestrial iso-
pod crustaceans. Smithsonian Institution, Washington, D.C. Available from http://
www.nmnh.si.edu/iz/isopod (accessed April 2006).
Kononenko, A.F. (1988) Lironeca atlantniroi sp. n. (Isopoda, Cymothoidae), a parasitic Isopoda
from fishes of the North Atlantic. Parazitologiya (St. Petersburg), 22 (5), 425–428.
Kussakin, O.G. (1979) Marine and Brackish Water Isopod Crustacea. Suborder Flabellifera, Akad-
emy of Science, U.S.S.R. Leningrad, 470 pp. (in Russian)
Menzies, R.J. & Kruczynski, W.L. (1983) Isopoda Crustacea (exclusive of Epicaridae). Memoirs of
TRILLES & JUSTINE
68 © 2006 Magnolia Press
1338
ZOOTAXA the Hourglass Cruises, 6, 1–126.
Milne Edwards, H. (1840) Histoire Naturelle des Crustacés comprenant l’anatomie, la physiologie
et la classification de ces animaux. Librairie Encyclopédique Roret, Paris, III, 605 pp.
Nierstrasz, H.F. (1931) Die Isopoden der Siboga-Expedition III. Isopoda Genuina II. Flabellifera.
Siboga-Expeditie, 32c, 123–232, pls X–XI.
Richardson, H. (1904) Contributions to the natural history of the Isopoda. Proceedings of the
United States National Museum, 27, no 1350, 1–89.
Richardson, H. (1909) Isopods collected in the North West Pacific by the U.S. Bureau of Fisheries
Steamer “Albatross” in 1906. Proceedings of the United States National Museum, 37, 75–129.
Richardson, H. (1910) Marine isopods collected in the Philippines by the U.S. Fisheries Steamer
“Albatross” in 1907–1908. Deparment of Commerce and Labor, Bureau of Fisheries , n° 736,
1–44.
Rocha-Ramírez, A., Chávez-López, R. & Bruce, N.L. (2005) Elthusa alvaradoensis n. sp (Isopoda,
Cymothoidae) from the gill chamber of the lizardfish, Synodus foetens (Linnaeaus, 1766).
Crustaceana, 78 (6), 701–707.
Sadowsky, V. & Moreira, P.L. (1981) Occurrence of Squalus cubensis Rivero, 1936, in the Western
South Atlantic Ocean, and incidence of its parasitic isopod Lironeca splendida sp. n. Studies on
Neotropical Fauna and Environment, 16, 137–150.
Shiino, S.M. (1951) On the cymothoid Isopoda parasitic in Japanese fishes. Bulletin of the Japanese
Society of Scientific Fisheries, 16, 81–89.
Schioedte, J.C. & Meinert, F. (1884) Symbolae ad Monographiam Cymothoarum Crustaceorum
Isopodum familiae. IV. Cymothoidae. Trib. II. Cymothoinae. Trib. III. Livonecinae. Naturhis-
torisk Tidsskrift, series III, XIV, 221–454, pls VI (Cym. XXIV)–XVIII(Cym. XXXVI).
Stebbing, T.R.R. (1905) Report on the isopods collected by Professor Herdman at Ceylon, in 1902.
In: W.A. Herdman (Ed.), Report to the Government of Ceylon on the Pearl Oyster Fisheries in
the Gulf of Manaar. Royal Society, London, 4, Supplements report no 23, 1–64.
Stimpson, W. (1857) The Crustacea and Echinodermata of the Pacific shores of North America.
Boston Journal of Natural History, 6, 503–513.
Tanaka, S. (1918) Twelve new species of Japanese fishes. Dobutsugaku Zasshi [Zool. Mag. Tokyo],
223–227 (in Japanese)
Trilles, J.P. (1976) Les Cymothoidae (Isopoda, Flabellifera) des collections du Museum National
d'Histoire Naturelle de Paris. IV. Les Lironecinae Schioedte & Meinert, 1884. Bulletin du
Muséum National d'Histoire Naturelle, Paris, 3ème série, 390 (Zool. 272), 773–800.
Trilles, J.P. (1979) Les Cymothoidae (Isopoda, Flabellifera; parasites de poissons) du Rijksmuseum
Van Natuurlijke Historie, Leiden. II. Afrique, Amérique et Régions Indo-Ouest-Pacifiques.
Zoologische Mededelingen, Leiden, 54, 245–275.
Trilles, J.P. (1994) Les Cymothoidae (Crustacea, Isopoda) du Monde. Prodrome pour une Faune.
Studia Marina, 21/22 (1991), 1–288.
Trilles, J.P. & Justine, J-L. (2004) Une nouvelle espèce de Cymothoidae et trois Aegidae (Crusta-
cea, Isopoda) récoltés sur des poissons de profondeur au large de la Nouvelle-Calédonie. Zoo-
systema, 26 (2), 211–233.
