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The genus Cymothoa Fabricius, 1793 is revised for Australian waters. Cymothoa hermani Hadfield, Bruce & Smit, 2011, previously known from Tanzania on the host Selar crumenophthalmus (Bloch, 1793) is new to Australian waters. Cymothoa carangi Avdeev, 1979; Cymothoa epimerica Avdeev, 1979; Cymothoa parupenei Avdeev, 1979; Cymothoa propria Avdeev, 1979; Cymothoa rotunda Avdeev, 1979 and Cymothoa pulchrum Lanchester, 1902 are redescribed. Cymothoa curta Schioedte & Meinert, 1884, first described from the host Anableps anableps (Linnaeus, 1758); and Cymothoa plebeia Schioedte & Meinert, 1884, first described from Cape Verde; are redescribed and excluded from the Australian fauna. Cymothoa limbata Schioedte & Meinert, 1884 is placed into junior synonymy with Cymothoa eremita (Brünnich, 1783). A key to the Australian species of Cymothoa is presented.
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Accepted by J. Svavarsson: 16 Nov. 2015; published: 3 Jun. 2016
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2016 Magnolia Press
Zootaxa 4119 (1): 001
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Monograph
http://doi.org/10.11646/zootaxa.4119.1.1
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ZOOTAXA
Review of the fish-parasitic genus Cymothoa Fabricius, 1793
(Crustacea: Isopoda: Cymothoidae) from Australia
MELISSA B. MARTIN
1,2,3
, NIEL L. BRUCE
2,4,5
& BARBARA F. NOWAK
1
1
Institute for Marine and Antarctic Studies, University of Tasmania Launceston, Tasmania, 7250, Australia.
E-mail: Melissa.Martin@utas.edu.au; B.Nowak@utas.edu.au
2
Museum of Tropical Queensland, Queensland Museum, 70–102 Flinders Street, Townsville, Queensland, 4810, Australia.
E-mail: niel.bruce@qm.qld.gov.au
3
School of Marine and Environmental Sciences, University of Malaysia Terengganu, Kuala Terengganu, 21030, Malaysia.
4
School of Marine and Tropical Biology, James Cook University, Queensland, 4810, Australia
5
Water Research Group (Ecology), North-West University, Potchefstroom, 2520, South Africa
Magnolia Press
Auckland, New Zealand
4119
MARTIN ET AL.
2
·
Zootaxa 4119 (1) © 2016 Magnolia Press
MELISSA B. MARTIN, NIEL L. BRUCE & BARBARA F. NOWAK
Review of the fish-parasitic genus Cymothoa Fabricius, 1793 (Crustacea: Isopoda: Cymothoidae) from
Australia
(Zootaxa 4119)
72 pp.; 30 cm.
3 Jun. 2016
ISBN 978-1-77557-957-1 (paperback)
ISBN 978-1-77557-958-8 (Online edition)
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UBLISHED I
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2016 B
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Magnolia Press
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ISSN 1175-5326 (Print edition)
ISSN 1175-5334 (Online edition)
Zootaxa 4119 (1) © 2016 Magnolia Press
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3
REVIEW OF CYMOTHOA FABRICIUS
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Suborder Cymothoida Wägele, 1989 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Superfamily Cymothooidea Leach, 1814 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Family Cymothoidae Leach, 1814 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Key to the marine buccal-attaching genera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Genus Cymothoa Fabricius, 1793 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Key to the Australian species of the genus Cymothoa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Cymothoa bychowskyi Avdeev, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Cymothoa carangi Avdeev, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
Cymothoa epimerica Avdeev, 1979. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Cymothoa eremita (Brünnich, 1783). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Cymothoa frontalis Milne Edwards, 1840. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Cymothoa hermani Hadfield, Bruce & Smit, 2011. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Cymothoa indica Schioedte & Meinert, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Cymothoa parupenei Avdeev, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Cymothoa propria Avdeev, 1979. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Cymothoa pulchrum Lanchester, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Cymothoa rotunda Avdeev, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .43
Cymothoa vicina Hale, 1926 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Species excluded from the Australian fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Cymothoa curta Schioedte & Meinert, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .57
Cymothoa plebeia Schioedte & Meinert, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Abstract
The genus Cymothoa Fabricius, 1793 is revised for Australian waters. Cymothoa hermani Hadfield, Bruce & Smit, 2011,
previously known from Tanzania on the host Selar crumenophthalmus (Bloch, 1793) is new to Australian waters. Cy-
mothoa carangi Avdeev, 1979; Cymothoa epimerica Avdeev, 1979; Cymothoa parupenei Avdeev, 1979; Cymothoa pro-
pria Avdeev, 1979; Cymothoa rotunda Avdeev, 1979 and Cymothoa pulchrum Lanchester, 1902 are redescribed.
Cymothoa curta Schioedte & Meinert, 1884, first described from the host Anableps anableps (Linnaeus, 1758); and Cy-
mothoa plebeia Schioedte & Meinert, 1884, first described from Cape Verde; are redescribed and excluded from the Aus-
tralian fauna. Cymothoa limbata Schioedte & Meinert, 1884 is placed into junior synonymy with Cymothoa eremita
(Brünnich, 1783). A key to the Australian species of Cymothoa is presented.
Key words: Cymothoidae, taxonomy, Cymothoa, fish buccal-attaching parasites, Australia
Introduction
Schioedte & Meinert’s (1881, 1883, 1884) monographic series of the world’s parasitic isopods included the first
worldwide review of the genus Cymothoa Fabricius, 1793, which at that time included 17 species (Schioedte &
Meinert 1884). One hundred and thirty years later, Cymothoa had grown to contain 49 accepted species worldwide
(Smit et al. 2014 listed 51 including nomina dubia). Cymothoa and Anilocra Leach, 1818 are currently the equal
largest genera of the family Cymothoidae. Since Schioedte & Meinert’s monographs, major generic revisions and
regional knowledge for Cymothoa rests with the works of Brusca (1981) for the eastern Pacific; Brusca & Iverson
(1985) for Pacific Costa Rica; Richardson (1905), Menzies & Frankenberg (1966) and Menzies & Kruczynski
(1983) for the western Atlantic; Thatcher et al. (2003) for Brazil, and more recently Hadfield et al. (2013) for
South Africa and the south-western Indian Ocean. Cymothoa is the most speciose buccal-attaching genus and is
morphologically the most challenging genus within the family (Monod 1934; Brusca 1981; Hadfield et al. 2013)
due to inadequate historic species descriptions, species without type material, and intraspecific variability. This has
led to frequent incorrect identifications of fresh material and to the misuse of species names.
MARTIN ET AL.
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Zootaxa 4119 (1) © 2016 Magnolia Press
Documentation of Australian Cymothoa began with Hale’s (1926) review, which included three species:
Cymothoa limbata Schioedte & Meinert, 1884 [=Cymothoa eremita (Brünnich, 1783)]; Cymothoa indica Schioedte
& Meinert, 1884 and Cymothoa vicina Hale, 1926. Avdeev (1978a, b, 1979a, 1979b) recorded and described a
further ten species. To date, 13 species are known from Australia (Bruce et al. 2002). The exclusion of Cymothoa
curta Schioedte & Meinert, 1884 and Cymothoa plebeia Schioedte & Meinert, 1884 from the Australian fauna and
the addition of Cymothoa hermani Hadfield, Bruce & Smit, 2011 as a new record for Australian waters bring the
total of Cymothoa species in Australia to 12.
Material and methods
Methods follow Martin et al. (2013, 2014a, b). The four developmental stages of Cymothoa follow Sartor & Pires
(1988). Host nomenclature (references not provided) were obtained from Fishbase (Froese & Pauly 2015) and
Catalogue of Fishes (Eschmeyer 2015). Collection data presented in “Material examined” include all available
information that accompanied the specimen(s). All “colour” remarks for each species description refers to adult
females (ovigerous and non-ovigerous) only. The male description includes only those characters that differ from
the female. Type material of previously described species was not dissected to avoid damaging type specimens.
Abbreviations: AM—Australian Museum, Sydney; AMNH—American Museum of Natural History, New
Yor k; MNHN—Muséum national d’Histoire naturelle, Paris; MTQ—Museum of Tropical Queensland,
Townsville; NHMUK—Natural History Museum, London; NHMW—Naturhistorisches Museum, Vienna; QM
Queensland Museum, Brisbane; SAM—South Australian Museum, Adelaide; SAMC—South African Museum,
Cape Town; SMF—Senckenberg Research Institute and Natural History Museum, Frankfurt; SMNH—Swedish
Museum of Natural History, Stockholm; TINRO—Russian Pacific Federal Fisheries Research Institute,
Vladivostok; UMZC—University Museum of Zoology, Cambridge; USNPC—U.S National Parasite Collection,
Beltsville, Maryland; ZMHB—Zoologisches Museum, Museum für Naturkunde, Humboldt-Universität Berlin;
ZMO—Zoological Museum, University of Oslo; ZMUC—Zoological Museum, University of Copenhagen,
Denmark; ZRC—Zoological Reference Collection, Lee Kong Chian Natural History Museum, Singapore
(LKCNHM).
Taxonomy
Suborder Cymothoida Wägele, 1989
Superfamily Cymothooidea Leach, 1814
Family Cymothoidae Leach, 1814
Remarks. This contribution concludes a series (Martin et al. 2013, 2014a, b, 2015a, b) revising the buccal-
attaching genera of Cymothoidae from Australia. A key is provided to all the marine genera of buccal-attaching
Cymothoidae, which is complemented by the keys given to the external-attaching genera (Bruce 1987a, b, c, 1990)
and the gill-attaching genera (Bruce 1986, 1990).
Key to the marine buccal-attaching genera
This key is restricted to ovigerous female of the genera that are solely known to attach in the host’s buccal region.
This group is not monophyletic as it includes Smenispa Özdikem, 2009 a genus which shows closest affinities to
external-attaching genera such as Anilocra and Livoneca Leach, 1818 (see Bruce 1987a, 1990 for diagnoses) and
Catoessa a genus of uncertain affinities (see Bruce 1990). The monotypic genus Tetragonocephalon Avdeev, 1975
is morphologically similar to Smenispa (based on the dorsal view of the cephalon, pereon and pleon shape) and as
we are unable to find any characters to distinguish the two genera, Tetragonocephalon is not included in the key.
Zootaxa 4119 (1) © 2016 Magnolia Press
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REVIEW OF CYMOTHOA FABRICIUS
1. Cephalon posterior margin trisinuate; pereon and pleon strongly co-linear; brood pouch with posterior pockets, oostegites of
coxae 6 forming most of the marsupium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Smenispa Özdikem, 2009
- Cephalon posterior margin rounded or straight; pereon and pleon weakly co-linear; brood pouch without posterior pockets;
oostegites arising from coxae 1–6, all forming the marsupium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Pleotelson posterior margin subtriangular; body widest at pereonite 6 . . . . . . . . . . . . . . . . . . . . . . . . . . . Olencira Leach, 1818
- Pleotelson rounded or subtruncate; body widest at pereonite 4 or 5. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Antennula longer than antenna; pereopod 5–7 basis without well-developed carina; pleon and pleotelson axially twisted
(“rotated”) against the pereon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Catoessa Schioedte & Meinert, 1884
- Antennula shorter than antenna; pereopods 5–7 basis with well-developed carina; pleon and pleotelson are not axially twisted
against pereon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Antennae thick and expanded; antennula bases contiguous; cephalon not or weakly immersed in pereonite 1 . . . . . . . . . . . . 5
- Antennae slender; antennula bases narrowly to widely separated; cephalon deeply immersed in pereonite 1 . . . . . . . . . . . . . 6
5. Pereonite 1 anterolateral margins projecting forward, without slight recess, anterior margin concave . . Ceratothoa Dana, 1852
- Pereonite 1 anterolateral margins projecting laterally, with slight recess, anterior margin straight or convex . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Glossobius Schioedte & Meinert, 1883
6. Body weakly vaulted; anterior margin of cephalon with distinct rostrum; anterolateral margins of pereonite 1 extending beyond
rostrum, with prominent lobes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lobothorax Bleeker, 1857
- Body strongly vaulted; anterior margin of cephalon without distinct rostrum anterolateral margins of pereonite 1 minute or
reaching anterior margin of rostrum, without prominent lobes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7. Body ovate, less than 1.5 times as long as greatest width; pleopods 3–5 without fleshy folds . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cinusa Schioedte & Meinert, 1883
- Body subparallel or weakly ovate, more than 1.5 times as long as greatest width; pleopods 3–5 with fleshy folds . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa Fabricius, 1793
Genus Cymothoa Fabricius, 1793
Cymothoa Fabricius, 1793: 503.—Milne Edwards, 1840: 264.—Schioedte & Meinert, 1884: 223.—Kussakin, 1979: 289.—
Brusca, 1981: 185.—Brusca & Iverson, 1985: 45.—Trilles, 1994: 137.—Hadfield, Bruce & Smit, 2011: 58.—Hadfield,
Bruce & Smit, 2013: 153.
Type species. Oniscus oestrum Fabricius, 1793; by subsequent designation (Kussakin 1979).
Remarks. The genus diagnosis has been comprehensively revised by Hadfield et al. (2011, 2013). Cymothoa can
be identified by the strongly vaulted body; slender antennae with widely separated bases, antenna longer than
antennula; subtruncate rostrum; pereopods 5–7 basis with broad blade-like carina; coxae partially visible dorsally;
pereonite 7 posterolateral margins extending past pleonite 1; and pleopods 3–5 with large fleshy folds.
Brusca (1981) presented a phylogeny of the Cymothoidae family, proposing three evolutionary linages based
on host site attachment. Brusca (1981) suggested that cymothoids evolved first as externally-attaching parasites
and then further evolved to using the host’s buccal cavity and gills.
The molecular analyses of Jones et al. (2008) (using 16S mtDNA) and Ketmaier et al. (2008) (using 16S rRNA
and cytochrome oxidase I) did not support Brusca’s (1981) evolutionary hypothesis for the Cymothoidae. Neither
of these analyses revealed distinct clades for the buccal, gill or externally-attaching genera. Jones’s et al. (2008)
tree topology showed a basal division, with Nerocila Leach, 1818 sister to the buccal-attaching Cymothoa indica
Schioedte & Meinert, 1884 and Olencira praegustator (Latrobe, 1802); the other larger clade showed Ceratothoa
as sister to the Anilocra clade. Ketmaier’s et al. (2008) analysis showed Nerocila as sister to Ceratothoa and the
gill-attaching taxa within the larger clade. The analyses of Ketmaier et al. (2008) and Jones et al. (2008) did not
resolve the relationships between the cymothoid genera and their results are not congruent with the results of
Hadfield’s (2012) morphological analysis. However, it was emphasized that their results were based on small
datasets of 11 and 6 species (excluding outgroup) respectively, and were not regarded as conclusive.
Hadfield (2012) presented the first morphological cladistics analysis of the Cymothoidae based on 23 genera
and 40 multistate characters. The 50% majority rule tree showed that the external-attaching Anilocra group formed
a morphologically well-supported clade that also included the gill-attaching Livoneca Leach, 1818, the buccal-
attaching Smenispa and Paracymothoa Lemos de Castro, 1955 (Hadfield 2012). Sister to the Anilocra clade was
the buccal-attaching Ceratothoa clade, with the remaining genera basally unresolved (Hadfield 2012). The
Ceratothoa group formed a cohesive clade, with Cymothoa sister to the Cinusa+Lobothorax and
Ceratothoa+Glossobius upheld by pereonite 1 anterolateral margins encompassing the cephalon (developed into
lobes in Glossobius), pereopods 5–7 basis with large blade-like carina (except Lobothorax which has no carina),
and maxilla medial lobe partly fused (Hadfield 2012).
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Zootaxa 4119 (1) © 2016 Magnolia Press
Smit et al. (2014) showed that the highest cymothoid diversity is found within the tropics and rapidly drops as
latitude increases. This is relevant to Cymothoa, where nine species are each found in the central Indo-Pacific and
tropical Atlantic region and eight species in the western Indo-Pacific. Thirteen species of Cymothoa were known
from Australian waters, and this number has not changed despite the current review for the genus, suggesting that
perhaps different cymothoid genera have different levels of diversity or distribution.
Key to the Australian species of the genus Cymothoa
1. Pereonite 1 anterolateral margins broad, with round or subtruncate projections; extending more than half the length of cepha-
lon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
- Pereonite 1 anterolateral margins narrow, with minute projections; extending less than half the length of cephalon . . . . . . . 5
2. Uropod half or shorter than pleotelson length; pereopod 7 ischium without bulbous protrusion on inferior distal margin . . . 3
- Uropod more than half of pleotelson length; pereopod 7 ischium with bulbous protrusion on inferior distal margin. . . . . . . . .4
3. Pereopod 7 merus with bulbous protrusion on inferior distal margin; pleonites not subequal in width . . . . . Cymothoa eremita
- Pereopod 7 merus without bulbous protrusion on inferior distal margin; pleonites subequal in width . . . . . Cymothoa frontalis
4. Frontal lamina ventrally folded, appearing to envelop paired antennae; pereopod 7 basis more than 1.0 times longer than wide,
dactylus broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa propria
- Frontal lamina not ventrally folded, not enveloping paired antennae; pereopod basis less than 1.0 times longer than wide, dac-
tylus narrow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa indica
5. Body subparallel or elongate, more than 2.0 times as long as greatest width. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Body ovate or wide, less than 2.0 times as long as greatest width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
6. Cephalon subtruncate, pleotelson posterior margin rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
- Cephalon subtriangular, pleotelson posterior margin subtruncate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
7. Pereonite 7 posterolateral margin not extending beyond pleonite 3; uropod exopod longer than endopod, pereopod 7 inferior
margin smooth and straight on basis, ischium and merus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa bychowskyi
- Pereonite 7 posterolateral margin extending beyond pleonite 3; uropod exopod similar length to endopod, pereopod 7 inferior
margin with bulbous protrusion on basis, ischium and merus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa pulchrum
8. Labrum narrow and small, eyes absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa carangi
- Labrum fleshy, eyes partially or distinctly visible . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa vicina
9. Pereonite 1 with bulbous ornamentation; anterolateral margin extending beyond cephalon anterior margin . . . . . . . . . . . . . .10
- Pereonite 1 without bulbous ornamentation; anterolateral margin partially reaching or reaching anterior margin of cephalon 11
10. Coxae 5–7 visible from dorsal view, posteroventral margins with acute carina; pleonite 5 wider than pereonite 7 . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa hermani
- Coxae 5–7 not visible from dorsal view, posteroventral margins without acute carina; pleonite 5 wider than pereonite 7. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa rotunda
11. Pereonite 1 posterolateral margins with minute projections; pereopod 7 basis with acute carina, pleopod 1 exopod and endopod
margins smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa epimerica
- Pereonite 1 posterolateral margins without minute projections; pereopod 7 basis without acute carina; pleopod 1 exopod and
endopod margins irregular. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymothoa parupenei
Cymothoa bychowskyi Avdeev, 1979
Cymothoa bychowskyi Avdeev, 1979a: 230, pl. 6, 7; 1985: 217, fig. 1.—Trilles, 1994: 138.—Williams, Bunkley-Williams &
Pitlik, 2000: 157.—Kensley, 2001: 232.—Bruce, Lew Ton & Poore, 2002: 174.—Paulay, Kropp, Ng & Eldredge, 2003:
479.—Trilles & Bariche, 2006: 228.Rameshkumar, Ravichandran, Sivasubramanian & Trilles, 2013: 42, fig. 1(C).
Type material. Holotype: ovig. ♀ (28 mm), off northwestern Australia, from red cornetfish Fistularia petimba
Lacépède, 1803 (TINRO АGK 75011).
Paratypes: 4 ovig. ♀ (TINRO АPK 75012–75015), 1 non-ovig. ♀, (TINRO АPK 75016), 4 mature ♂ (TINRO
АPK 75017–75020), 1 immature ♂ (TINRO АPK 75021); same data as holotype.
Remarks. No material of this species was examined. A loan request for the types was made, but the specimens
could not be located. Cymothoa bychowskyi can be identified by the body 2.4 times as long as wide; pereonite 1
with broad anterolateral margins reaching half the length of cephalon; coxae 2–4 posteroventral margins rounded,
coxae 5–7 posteroventral margins project laterally in dorsal view; pleonites subequal in length, pereonite 7
posterolateral margin extending to pleonite 4; pleotelson posterior margin rounded; uropods not extending beyond
posterior margin of the pleotelson and pleopod 2 of female holotype with an appendix masculina (0.7 times as long
as pleopod 2 exopod length).
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Avdeev (1979a) briefly compared C. bychowskyi to C. parupenei Avdeev, 1979a. Both species have a
subtruncate cephalon, and are widest at pereonite 5. Cymothoa bychowskyi is distinguished from C. parupenei by
the elongate body, 2.4 times longer than wide (compared to the 1.7 longer than wide body in C. parupenei);
anterolateral margins of pereonite 1 projecting forward (compared to the broader anterolateral margins of pereonite
1 curved mesially towards cephalon), and the rounded posterior margin of pleotelson (compared to the subtruncate
posterior margin of C. parupenei).
Cymothoa bychowskyi has high host specificity and the infrequent reports would suggest low occurrence.
Rameshkumar et al. (2013) reported 7.9% prevalence (3 of 38 hosts) of the species from Agatti Island,
Lakshadweep, southeastern India.
Distribution. Northwestern and Western Australia, precise locality not given (Avdeev 1979a; Kensley 2001).
Also reported from Guam, Micronesia (Williams et al. 2000), Okinawa, Japan (Williams et al. 2000) and
Lakshadweep, India (Rameshkumar et al. 2013).
Hosts. Known only from family Fistulariidae: Fistularia petimba, previously Fistularia villosa Klunzinger,
1871 (see Avdeev 1979a; Williams et al. 2000; Rameshkumar et al. 2013) and bluespotted cornetfish Fistularia
commersonii Rüppell, 1838 (Williams et al. 2000).
Cymothoa carangi Avdeev, 1979
Figures 1–2
Cymothoa carangi Avdeev, 1979b: 53, pl. 3.—Trilles, 1994: 138.—Bruce, Lew Ton & Poore, 2002: 174.—Trilles & Bariche,
2006: 228.
Cymothoa carangii.—Kensley, 2001: 232.
Material examined. Holotype: 1 ovig. ♀ (31 mm), northern Australia (precise locality not given), from Caranx sp.
(TINRO АGK 75022).
Ovigerous female. Length 31 mm, width 13 mm (holotype).
Body subrectangular, 2.1 times as long as greatest width, dorsal surface smooth, laterally sub-parallel, widest at
pereonite 6, most narrow at pereonite 1. Cephalon subtriangular, 0.5 times as long as wide, visible from dorsal
view, immersed in pereonite 1. Frontal margin rounded to form blunt rostrum, ventrally folded. Eyes absent.
Pereonite 1 anterolateral margins broad, reaching anterior margin of cephalon; pereonites 1–4 subequal in length;
pereonites 5–7 progressively narrower in length, posterior margin linear. Coxae 2–3 posteroventral margins
subtruncate; 4–7 with progressively acute carinae. Pleonites not subequal in width; posterolateral margins of
pleonite 2 acute; pleonites 3–5 progressively increasing in width, posterior margins slightly irregular. Pleotelson
subtruncate, 0.6 times as long as anterior width, anterior margin not linear, lateral margins weakly convex,
posterior margin straight, without median point.
Antennula comprised of 8 articles; peduncle articles 1 and 2 distinct and articulated; article 2 1.4 times as long
as article 1; article 3 0.6 times as long as combined lengths of articles 1 and 2, 1.2 times as long as wide. Antenna
comprised of 8 articles, peduncle article 3 1.5 times as long as article 2, 1.1 times as long as wide; article 4 1.0
times as long as wide; article 5 1.0 times as long as article 4, terminal article without setae, extending to anterior
margin of pereonite 1. Labrum lateral margins convex, anterior margin narrowly rounded.
Pereopod 1 basis 1.4 times as long as greatest width; ischium 0.7 times as long as basis; merus proximal
margin without bulbous protrusion; carpus with straight proximal margin; propodus 1.4 times as long as wide;
dactylus narrow, 1.5 times as long as propodus, 1.2 times as long as basal width. Pereopod 2 propodus 1.0 times as
long as wide; dactylus 1.1 times as long as propodus. Pereopods 3–5 similar to pereopod 2, gradually increasing in
size, without robust or simple setae. Pereopod 6 basis 1.3 times as long as greatest width, superior proximal margin
with acute carina; ischium 0.6 times as long as basis; propodus 0.9 times as long as wide; dactylus 2.1 times as long
as propodus. Pereopod 7 basis 1.3 times as long as greatest width, superior proximal margin with acute carina;
ischium 0.6 times as long as basis, inferior distal margin with bulbous protrusion; merus proximal margin with
slight bulbous protrusion, 0.4 times as long as ischium, 0.5 times as long as wide; carpus 0.2 times as long as
ischium, without bulbous protrusion, 0.4 times as long as wide; propodus 0.5 times as long as ischium, 0.9 times as
long as wide; dactylus stout, 2.5 times as long as propodus, 2.5 times as long as basal width.
Uropod not extending beyond posterior margin of pleotelson; peduncle lateral margin without setae, marginal
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setae absent. Exopod not extending past endopod, 4.7 times as long as greatest width, apically rounded, lateral
margin convex, terminating without setae, mesial margin concave. Endopod 5.0 times as long as greatest width,
apically blunt, lateral margin weakly convex, terminating without setae, mesial margin weakly concave.
FIGURE 1. Cymothoa carangi, ovigerous female holotype (31 mm; AGK 75022). A, dorsal view; B, dorsal view of pereonite
1 and cephalon; C, ventral view of mouthpart; D, antennula; E, antenna; F, lateral view.
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FIGURE 2. Cymothoa carangi, ovigerous female holotype (31 mm; AGK 75022). A–D, pereopods 1, 2, 6, 7 respectively; E,
uropod; F, dorsal view of pleotelson.
Colour. Yellowish–tan.
Remarks. Cymothoa carangi can be identified by the body being subparallel, 2.1 times as long as wide and
widest at pereonite 6; subtriangular cephalon immersed in pereonite 1; broad anterolateral margins of pereonite 1
anteriorly reaching rostrum; pereonites 1–4 subequal in length; pereonite 7 posterolateral margins extending to
pleonite 4; pleotelson subtruncate and 1.8 times wider than long; uropods not extending beyond posterior margin of
pleotelson, pereopods 4–7 basis with sharp carinae (see lateral view in fig. 1F) and pereopod 7 with a projection on
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the proximal superior region of the ischium. Avdeev (1979b) illustrated the maxillula with three terminal setae;
maxilla lateral lobe partly fused to the mesial lobe, with one recurved robust seta on each lobe; maxilliped weakly
segmented with three recurved robust setae on terminal article 3; and pleopod 1 exopod 1.1 times longer than wide,
weakly concave lateral margin, rounded distal margin and a strongly convex mesial margin. To date, no males have
been reported for the species.
This species resembles C. parupenei in the dorsally visible coxae, pereonite 1 anterolateral margins reaching
rostrum, pleotelson subtruncate and the subparallel body shape. Cymothoa carangi can be distinguished from C.
parupenei by the subtriangular cephalon (compared to the subtruncate cephalon of C.parupenei), pereopods 6 and
7 basis with sharp carinae (compared to the rounded and smooth basis on C. parupenei holotype) and the pleopod 1
endopod is smaller than the exopod (compared to the subequal endopod and exopod of C. parupenei pleopod 1).
Other Cymothoa species known to occur on the Carangidae are Cymothoa eremita (Brünnich, 1783);
Cymothoa indica Schioedte & Meinert, 1884; Cymothoa propria Avdeev, 1979 and Cymothoa pulchrum
Lanchester, 1902. These four species differ from C. carangi in the following characteristics: C. eremita has a
subtruncate cephalon, broad anterolateral margins of pereonite 1 which taper to a small produced point and
rounded posteroventral margins of coxae 4–7; C. indica has anterolateral margins of pereonite 1 not reaching
medial region of cephalon and pereopod 7 basis without sharp carina; C. propria has the cephalon frontal margin
ventrally folded, anterolateral margins of pereonite 1 with minute projection, posterior margin of pleonites irregular
and uropod reaching posterior margin of pleotelson; C. pulchrum has pereopods 6 and 7 basis without the distinctly
“blade-like” carinae, pereopods 6 and 7 dactyli more slender, and body lateral margins subparallel (especially
pereonites 1–5, which are progressively wider in C. carangi).
Distribution. Known only from northwestern Australia without specific locality (Avdeev 1979b).
Hosts. Only known from the family Carangidae: Caranx sp. (Avdeev 1979b).
Cymothoa epimerica Avdeev, 1979
Figures 3–8
Cymothoa epimerica Avdeev, 1979a: 225, pl. 2–3.—Trilles, 1994: 139; 2008: 23.Trilles & Bariche, 2006: 228.Bruce, Lew
Ton & Poore, 2002: 175.—Hadfield, Bruce & Smit, 2013: 157.
Cymothoa pulchra.—Yu & Li, 2003a: 228, fig. 5; 2003b: 267.
Not Cymothoa epimerica.—Trilles, 2008: 23 (part Cymothoa eremita SMF-567; part Cymothoa vicina SMF-572).
Type material. Holotype: 1 ovig. ♀, (17.5 mm), from the Indian Ocean off Australia, on Malabar blood snapper
Lutjanus malabaricus (Bloch & Schneider, 1801) (previously Lutianus malabaricus) (TINRO АGK 75023).
Paratypes: 4 mature ♂ (TINRO АPK 75024–75025); same data as holotype.
Types not examined. A request to borrow the holotype and paratypes were unsuccessful as the specimens could
not be located.
Material examined. Northern Territory material: 1 ♀ ovig. (27 mm), 2 immature ♂ (8, 9 mm), outer harbour,
Darwin, 7 October 2012, from estuary cod Epinephelus coioides (Hamilton, 1822), coll. Ben Diggles (MTQ
W34150). 1 ♀ ovig. (31 mm), 3 immature ♂ (8, 4, 4 mm), inner harbour, Darwin, 8 March 2013, coll. Ben Diggles
(MTQ W34170). 1 ♀ ovig. (19 mm), Darwin, 11 October 2012, from estuary cod Epinephelus coioides, coll. Ben
Diggles (MTQ W34151). 1 ♀ ovig. (29 mm), north of Melville Island, 10.1189ºS; 129.3192ºE, 2 October 1995,
coll. J. Lloyd (AM P89953).
Digital image from the Senckenberg Research Institute: 1 ♀ ovig. (20 mm), 1 immature ♂ (7 mm), Seychelles
(SMF-76), without additional information.
Ovigerous female Length 31 mm width 16 mm (non-dissected, MTQ W34170); length 27 mm width 14 mm
(dissected, MTQ W34150).
Body ovoid, 1.9 times as long as greatest width, dorsal surface smooth, laterally dome-shaped, widest at
pereonites 5 and 6, most narrow at pereonite 1. Cephalon subtriangular, 0.3 times longer than wide, visible from
dorsal view, deeply immersed in pereonite 1. Frontal margin rounded to form blunt rostrum, ventrally folded. Eyes
partially distinct. Pereonite 1 anterolateral margins curved towards cephalon, reaching anterior margin of cephalon,
posterior margins of pereonites 1–6 weakly trisinuate, moderately concave laterally; pereonite 7 extending to
pleonite 3, visible in dorsal view. Coxae 2–3 posteroventral margins subtruncate; 4–7 with sharp and acute carinae
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projecting laterally. Pleon widest at pereonite 5; pleonites posterior margin weakly bisinuate. Pleotelson rounded,
0.5 times as long as anterior width, anterior margin trisinuate, lateral margins convex, posterior margin round,
without median point.
Antennula comprised of 8 articles; peduncle articles 1 and 2 distinct and articulated; article 2 0.9 times as long
as article 1; article 3 0.5 times as long as combined lengths of articles 1 and 2, 1.0 times as long as wide. Antenna
comprised of 9 articles, peduncle article 3 1.2 times as long as article 2, 1.2 times as long as wide; article 4 1.2
times as long as wide; article 5 1.0 times as long as article 4, terminal article without setae, not extending to
posterior of pereonite 1. Labrum lateral margins convex, anterior margin narrowly rounded, without small median
point.
FIGURE 3. Cymothoa epimerica ovigerous female (31 mm; MTQ W34170). A, dorsal view; B, dorsal view of pereonite 1 and
cephalon; C, ventral view of mouthpart; D, antennula; E, antenna; F, dorsal view of pleotelson; G, lateral view.
Pereopod 1 basis 1.3 times as long as greatest width; ischium 0.9 times as long as basis; merus proximal
margin without bulbous protrusion; carpus with straight proximal margin; propodus 1.8 times as long as wide;
dactylus narrow, 1.1 times as long as propodus, 2.8 times as long as basal width. Pereopod 2 propodus 1.1 times as
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long as wide; dactylus 1.7 times as long as propodus. Pereopod 3 similar to pereopod 2, merus moderately
protruding, basis superior proximal margin with sharp carina. Pereopods 4 and 5 similar to pereopod 2, gradually
increasing in size, without robust or simple setae. Pereopod 6 basis 1.4 times as long as greatest width, superior
proximal margin with sharp and acute carina; ischium 0.5 times as long as basis; propodus 1.7 times as long as
wide; dactylus 1.5 times as long as propodus. Pereopod 7 basis 1.2 times as long as greatest width, superior
proximal margin with sharp and acute carina; ischium 0.9 times as long as basis, with bulbous protrusion; merus
proximal margin with slight bulbous protrusion, 0.3 times as long as ischium, 0.6 times as long as wide; carpus 0.3
times as long as ischium, without bulbous protrusion, 1.3 times as long as wide; propodus 0.5 times as long as
ischium, 0.8 times as long as wide; dactylus narrow, 2.5 times as long as propodus, 2.9 times as long as basal width.
FIGURE 4. Cymothoa epimerica ovigerous female (27 mm; MTQ W34150). A–E, pereopods 1, 2, 5, 6, 7 respectively; F,
uropod; G–K, pleopods 1–5 respectively.
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FIGURE 5. Cymothoa epimerica male (4 mm), pre-manca (2.2 mm) and manca (2.2 mm) (MTQ W34170). A, dorsal view of
male; B, dorsal view of pre-manca; C, dorsal view of pre-manca; D, male uropod; E, male antennula; F, male antenna; G, lateral
view male.
Pleopods margins irregular, lobes increasing in size from 1–5, exopods 1–5 gradually decreasing in size,
exopod size almost similar to endopod. Pleopod 1 exopod 0.8 times as long as wide, lateral margin deeply convex,
distally round and irregular, mesial margin strongly convex; peduncle 1.9 times as wide as long, without
retinaculae. Pleopod 2 similar to pleopod 1, mesial margin becoming more strongly produced, endopod with deep
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depressions. Pleopods 3–5 endopods proximal margins extending below exopod to peduncle, with large fleshy
folds and medial lobes present, increasing in size from pleopods 1–5.
Uropod not extending beyond posterior margin of pleotelson; peduncle 1.4 times as long as greatest width, 0.4
times as long as exopod, lateral margin without setae, marginal setae absent, apices narrowly acute, lateral margin
straight, mesial margin strongly convex. Exopod extending past endopod, 8.3 times as long as greatest width,
apically rounded, lateral margin straight, terminating without setae, mesial margin weakly straight.Endopod 7.0
times as long as greatest width, apically rounded, lateral margin weakly convex, terminating without setae, mesial
margin straight.
FIGURE 6. Cymothoa epimerica male (8 mm) (MTQ W34150). A–D, pereopods 1, 2, 6, 7 respectively; E–I, pleopods 1–5
respectively.
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FIGURE 7. Cymothoa epimerica, female (20 mm; SMF-76) A, dorsal view; B, lateral view; C, ventral view. (picture credit
Sven Tränkner, Senckenberg Research Institute).
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FIGURE 8. Cymothoa epimerica, male (7 mm; SMF-76) A, dorsal view; B, ventral view; C, lateral view. (picture credit Sven
Tränkner, Senckenberg Research Institute).
Males. Length 4 mm width 2 mm (non-dissected, MTQ W34170); length 8 mm width 3 mm (dissected,MTQ
W34150).
Body 2.0 times as long as greatest width. Cephalon rounded, 0.3 times longer than wide, visible from dorsal
view, not deeply immersed in pereonite 1. Eyes distinct, 0.4 times width of cephalon (combined eyes width).
Pereonite 1 anterolateral margin minute; posterior margins of pereonites 1–6 straight and smooth. Coxae 2–7
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posteroventral margins rounded. Pleonite 3 subequal in width to pleonites 4 and 5, visible in dorsal view; pleonites
posterior margin not trisinuate. Pleotelson subtruncate. Antennula comprised of 6 articles; shorter than antenna.
Antenna comprised of 10 articles.
Pereopod 1 basis 2.0 times as long as greatest width; ischium 0.3 times as long as basis; merus proximal
margin without bulbous protrusion; carpus with straight proximal margin; propodus 2.4 times as long as wide;
dactylus 1.5 times as long as propodus. Pereopods 2–7 similar to pereopod 1, moderately bigger in size, pereopods
6 and 7 basis superior proximal margin without raised carinae. Pleopods margins smooth, exopods greater than
endopods from 1–5; pleopod 2 appendix masculina with parallel margins, 0.6 times as long as endopod, distally
acute. Uropod exopod similar length to endopod, apices broadly rounded.
Manca. Length 2.2 mm width 1 mm.
Body 2.2 times as long as greatest width. Cephalon anterior margin rounded, 0.6 times longer than wide,
visible from dorsal view, not deeply immersed in pereonite 1. Eyes distinct, 0.4 times width of cephalon (combined
eyes width). Pereonite 1 anterolateral margin minute; posterior margins of pereonites 1–6 straight and smooth.
Pleonite 1 subequal in width to pleonites 3–5, visible in dorsal view; pleonites posterior margin smooth and
straight. Pleotelson 1.1 times as long as anterior width; lateral margins smooth and concaved; posterior margin
converging to caudomedial point, with setae. Antennula as long as antenna. Uropod endopod 4.9 times as long as
exopod length, apices narrowly rounded; exopod with setae.
Pre-manca. Length 2.2 mm width 1 mm.
Pre-manca similar to manca in body size ratio, eye shape, pereon and pleon morphology. Eyes 0.4 times width
of cephalon (combined eyes width).Cephalon anterior margin weakly subtriangular, 0.5 times longer than wide.
Pleotelson without setae. Uropod endopod 2.2 times as long as exopod length, apices narrowly rounded; exopod
without setae.
Colour. Ivory white.
Size. Present material: ovigerous females: 19–31 mm; immature males: 4–9 mm.
Remarks. Cymothoa epimerica has an ovoid body (1.9 times longer than wide); subtriangular cephalon deeply
immersed in pereonite 1; pereonite 1 anterolateral margins deeply curved towards cephalon; narrow uropodal rami
(exopod 8.3 times longer than wide); pereopods 5–7 superior proximal margin with acute carinae and dorsally
visible; coxae 6 and 7 posteroventral margins acute and dorsally visible; pleotelson posterior margin smooth and
rounded; pleopod morphology appearing damaged and irregular, and increased fleshy folds on endopods from
pleopods 3–5. The mouthparts of specimenMTQ W34150 is similar to Avdeev’s (1979a) illustrations, which
showed a maxillula with 3 terminal robust setae; lateral lobe of maxilla partly fused to mesial lobe, with 4 and 5
recurved setae on the mesial lobe and lateral lobe respectively; and a weakly segmented maxilliped. The male
specimens have a broader, subquadrate body; eyes distinct; straight pleonite posterior margins; coxae with broad
posteroventral margins, visible in dorsal view; pereopods slender without bulbous protrusions, and pleopods 1–5
margins smooth.
Cymothoa epimerica pre-mancae (fig. 5B) and mancae (fig. 5C) have unique uropod morphology, with the
endopod twice the length of the exopod long in pre-mancae (four times the exopod length in mancae). The
pleotelson and uropods of pre-mancae are without setae (setae present in mancae). Both stages of pre-mancae and
mancae were found within the same brood pouch of the ovigerous female specimen from MTQ W34170. The pre-
mancae develops into free-swimming mancae which is released from the female marsupium in search of a suitable
host (Smit et al. 2014).
Trilles (2008) recorded C. epimerica from Seychelles (SMF-76) and the Red Sea (SMF-567 and SMF-572)
without host association or illustrations, and briefly mentioned the similarities of C. epimerica and C. curta. After
examining the digital images from the Senckenberg Research Institute, specimen SMF-76 from the Seychelles
agrees withC. epimerica, whereas SMF-567 from the Red Sea is here re-identified as C. eremita and SMF-572 re-
identified as Cymothoa vicina Hale, 1926. Cymothoa epimerica and C. curta have a similar ovoid body, pleotelson
posterior margin rounded and cephalon immersed in pereonite 1. Cymothoa curta differs from C. epimerica in the
small and acute anterolateral margins of pereonite 1; posterolateral margins of coxae rounded and not dorsally
visible; pereopod 6 basis superior proximal margin rounded; pereopod 7 basis highly raised and broad; uropodal
rami broader thanC. epimerica, uropod apices rounded; and the smooth mesial and lateral margins of pleopods 1–
5.
Yu & Li’s (2003a, b) illustrations of their specimens identified as C. pulchra agree with C. epimerica in having
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a wide and oval-shaped body; dorsally visible and posteriorly acute coxae; rounded posterior margin of pleotelson
and in dorsal view, the uropods not extending beyond pleotelson posterior margin. Cymothoa pulchrum has a more
subrectangular and subparallel body shape, rostrum dorsally appearing more subtruncate compared to the rounded
rostrum of C. epimerica, subequal and stout uropodal rami and pereopod 7 basis superior proximal margin not
dorsally visible.
Distribution. Northern Territory, Australia.
Hosts. Known from the family Serranidae: Epinephelus coioides (present study) and family Lutjanidae:
Lutjanus malabaricus (Avdeev 1979a).
Cymothoa eremita (Brünnich, 1783)
Figures 9–13
Oniscus oestrum.—Spengler 1775: 312, pl. 7 (figs. i–k).
Oniscus eremita Brünnich, 1783: 319.
Cymothoa Leschenaultii Leach, 1818: 352.—Desmarest, 1825: 309.
Cymothoa Mathoei Leach, 1818: 353; Desmarest, 1825: 309.
Cymothoa mathoei.—Milne Edwards, 1840: 270.—Hilgendorf, 1869: 114.—Gerstaecker, 1901: 182, 258.
Cymothoa leschenaultii.—White, 1847: 109.—Miers, 1880: 461.—Ellis, 1981: 124.
Cymothoa matthaei [sic].—White, 1847: 110.—Bleeker, 1857: 22.
Cymothoa mathaei.—Lucas, 1850: 248.—Kossmann, 1880: 117, pl. 10 (figs. 1–3).
Cymothoa edwardsii Bleeker, 1857: 21, 33, tab. II, fig. 12.—Miers, 1880: 461.—Gerstaecker, 1901: 261.
Cymothoa stromatei Bleeker, 1857: 21, 33, 35 tab II, fig. 13.—Miers, 1880: 461.—Gerstaecker, 1901: 181.—Lanchester, 1902:
377.—Richardson, 1910: 22.—Hale, 1926: 214, fig. 9h.—Brian & Dartevelle, 1949: 184.—Sachlan, 1952: 41, 50 photo
28.—Pillai, 1954: 15.
Cymothoa eremita.—Schioedte & Meinert, 1884: 259, tab. VII (Cym. XXV), figs. 3–13.—Stebbing, 1893: 354; 1910: 102.—
Gerstaecker, 1901: 182.—Thielemann, 1910: 39, figs. 37, 38, tab. 4.—Nierstrasz, 1915: 90; 1931: 135, pl. 10, fig. 9.—
Monod, 1924: 100; 1933: 195; 1976: 859, figs. 23–25.—Boone, 1935: 215, pl. 63.—Shiino, 1951: 81 figs. 2 (b–c).—
Avdeev, 1978a: 30; 1982b: 69.—Trilles, 1975: 987, pl. II (12–13); 1979b: 261; 1986: 627, tab. 1; 1994: 139; 2008: 23.—
Bowman & Tareen, 1983: 25, fig. 20.—Radhakrishnan & Nair, 1983: 96, 105, 107.—Saito, Itani & Nunomura, 2000:
65.—Shireen, 2000: 21, figs. 1–3.—Kensley, 2001: 232.—Yu & Li, 2003a: 228, fig. 4; 2003b: 267.—Trilles & Bariche,
2006: 228.—Williams & Bunkley-Williams, 2009: 557.—Trilles, Ravichandran & Rameshkumar, 2011: 446.—
Rameshkumar, Ravichandran & Trilles, 2012: 191.—Hadfield, Bruce & Smit, 2013: 158, figs. 3–5.
Cymothoa limbata Schioedte & Meinert, 1884: 248, tab. VII (Cym. XXV), figs. 1, 2.—Hale, 1926: 214.—Nierstrasz, 1931:
136.—Bruce, Lew Ton & Poore, 2002: 175 [new synonymy].
Cymothoa edwardsi.—Nierstrasz, 1931: 135.
Cymothoa sp. (an. eremita Brünnich, 1783) [sic].—Monod, 1934: 13, pl. 27 (a–b), pl. 30 (b).
Cymothoa erimitae (typographical error?).—Sachlan, 1955: 31.
Cymothoa cinerea Bal & Joshi, 1959: 567, pl. 2, figs. 1–5.—Kensley, 2001: 232.
Cymothoa cinerius.—Joshi & Bal, 1960: 446.
Cymothoa mathieui.—Ellis, 1981: 124.
Cymothoa leaschenaultii [sic].—Kensley, 2001: 232.
Cymothoa epimerica.—Trilles, 2008: 23 (SMF-567).
Type material. Holotype: 1 non-ovig. ♀ (26 mm), from the buccal cavity of black pomfret Parastromateus niger
(Bloch, 1795), (ZMUC-CRU-10078). Not examined, see Hadfield et al. 2013 for further details.
Material examined. 1 non-ovig. ♀ (34 mm), near Bathurst Island, Northern Territory, 18 April 1988, from
gold-banded jobfish Pristipomoides multidens (Day, 1871), coll. M. Pearce (MTQ W30413).
1 ovig. ♀ (31 mm), no additional information (AM P8689). 1 ovig. ♀, 30 mm, provenance presumed to be
New Guinea; no additional information (AM P9609). The specimens (AM P8689 and AM P9609) were previously
examined by Herbert M. Hale without additional data (see remarks for further discussion).
Digital image from the Senckenberg Research Institute: 1 ♀ ovig. (18 mm), 1 immature ♂ (11 mm), Red Sea
(SMF-567), without additional information.
Diagnosis and description. Hadfield et al. (2013) provided a detailed diagnosis and description of the type
material. We include here illustrations and a brief diagnosis of the Australian female specimen only (MTQ
W30413).
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FIGURE 9. Cymothoa eremita, male (34 mm; MTQ W30413) A, dorsal view; B–E, pereopods 1, 2, 6, 7 respectively; F, ventral
view.
Body subrectangular, 1.7 times as long as greatest width, dorsal surface rugose, laterally sub-parallel, widest at
pereonite 6, most narrow at pereonite 1. Cephalon subtruncate, 0.4 times longer than wide, visible from dorsal
view. Eyes partially visible. Pereonite 1 anterolateral margins minute, not reaching anterior margin of cephalon;
pereonites 1–4 subequal in length and width length, posterior margin linear; pereonites 5–7 subequal in length.
Coxae 2–3 posteroventral margins subtruncate; 4–7 without acute carinae. Pleonite 1 narrower than pleonites 2–5,
visible in dorsal view; pleonites 1–5 with irregular posterior margin; pleonites 3–5 progressively increasing in
width. Pleotelson subtruncate, 0.3 times as long as anterior width, anterior margin irregular, lateral margins
concave, posterior margin straight, without median point. Pereopods 1 and 2 dactyli slender; carpus proximal
margin straight; merus proximal margin with slight bulbous protrusion; basis superior proximal basis without acute
carinae. Pereopods 3–5 similar to pereopod 2, gradually increasing in size, without robust or simple setae.
Pereopods 6 and 7 superior proximal basis with broad and acute carinae; merus proximal margin with slight
bulbous protrusion; dactyli slender.
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FIGURE 10. Cymothoa eremita ovigerous female (30 mm; AM P9609). A, dorsal view; B, front view of pereonite 1 and
cephalon; C, ventral view of mouthpart; D, antennula; E, antenna; F, lateral view.
Size. Ovigerous females: 19–44 mm; non-ovigerous females: 17–35; males: 9–24 mm; second pullus: 2 mm
(Schioedte & Meinert 1884; Bal & Joshi 1959; Monod 1976; Trilles 1979b; Shireen 2000).
Remarks. Hadfield et al. (2013) redescribed and diagnosed C. eremita from the female holotype (ZMUC-
CRU-10078). C. eremita can be identified by pereonite 1 anterolateral margins extending nearly half the length of
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cephalon; cephalon subtruncate; pleon as wide as pereon; uropods not extending to pleotelson posterior margin;
pereopod 7 with a bulbous protrusion on ischium; and small horn-like structures on the posterolateral margins of
pereonite 1.
FIGURE 11. Cymothoa eremita ovigerous female (30 mm; AM P9609). A–D, pereopods 1, 2, 6, 7 respectively; E, uropod.
Cymothoa eremita is morphologically variable (see Hadfield et al. 2013). The holotype of C. eremita (ZMUC-
CRU-10078) has a subtriangular body, 2.7 times longer than wide, body widest at pereonite 6; pleon not immersed
in pereonite 7; and the smooth carina on pereopod 7 basis (1.7 times longer than its greatest width). The female
syntype of Cymothoa mathoei (NHMUK 1979.407.2) has a subparallel body, 2.4 times longer than wide; a raised
dorsal median longitudinal ridge; pleon immersed in pereonite 7; and pereopod 7 basis with an irregular carina,
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which is 1.3 times longer than its greatest width. The holotype of Cymothoa leschenaultii differs from the previous
mentioned specimens by having a more ovoid body shape, 2.1 times longer than wide, body widest at pereonite 5;
pleonite posterior margins noticeably more trisinuate, pereonite 7 moderately laterally encompasses pleon and a
highly raised carina on pereonite 7 basis, 1.0 times longer than its greatest width.
FIGURE 12. Cymothoa eremita, female (18 mm; SMF-567) A, dorsal view; B, lateral view; C, ventral view. (picture credit
Sven Tränkner, Senckenberg Research Institute).
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FIGURE 13. Cymothoa eremita, male (11 mm; SMF-567) A, dorsal view; B, ventral view; C, lateral view. (picture credit Sven
Tränkner, Senckenberg Research Institute).
The extent to which variation in this species is either regional, host mediated or both is still not apparent, as
indicated by variations in the specimens allocated to C. eremita by Hadfield et al. (2013). The Australian specimen
is most similar to the syntype specimen of C. mathoei (illustrated by Hadfield et al. 2013). The Australian
specimen has a wider body, 1.7 times longer than wide (as opposed to 2.4 times longer than wide); irregular
posterior margins of the pleonites; more visible cephalon with anterolateral margins of pereonite 1 minute; and a
slightly raised carina on pereopod 7 basis.
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Among Hale’s specimens held at the South Australian Museum, we discovered a specimen of “C. stromatei
without host and locality data (AM P9609). Hale (1926) indicated he examined a specimen from New Guinea and
briefly compared C. stromatei to C. vicina Hale, 1926 stating that both species have a subtruncate cephalon anterior
margin. Hale (1926) did not mention examining the type specimens of Cymothoa stromatei Bleeker, 1857 (from
“Batavia” = Java). The close agreement of Hale’s (1926) figures of C. stromatei to our Northern Territory material
figured here suggests that the specimen is that which he examined from New Guinea. The specimens from New
Guinea and Australia are very similar, the only variation noted is the more prominent anterolateral margins of
pereonite 1 in Hale’s New Guinea specimen.
Trilles (1994) originally proposed the synonymy of Cymothoa limbata Schioedte & Meinert 1884 with C.
eremita. Hadfield et al. (2013) also noted the strong resemblance of pereonite 1 and the pleon morphology of the
two species. Cymothoa limbata resembles our examined specimens in cephalon, pereon, pleon and pleotelson
morphology, and we maintain the synonymy of C. limbata with C. eremita.
Distribution. Known from the Indian Ocean and Indo-Pacific regions. For locality details see Hadfield et al.
(2013) and Trilles (1994, 2008).
Hosts. Present study from Lutjanidae. Known from 10 host families (family Aulopidae, Carangidae,
Haemulidae, Mugilidae, Psettodidae, Serranidae, Siganidae, Sphyraenidae, Stromateidae, Tetraodontidae). For fish
host species details, see Hadfield et al. (2013).
Cymothoa frontalis Milne Edwards, 1840
Cymothoa frontale (Cymothoé frontal) Milne Edwards, 1840: 271.
Cymothoa frontalis.—White, 1847: 110.—Heller, 1868: 146.—Schioedte & Meinert, 1884: 226, tab. VI (Cym. XXIV) figs. 1,
2.—Gerstaecker, 1901: 261.—Nierstrasz, 1931: 136.—Avdeev, 1978b: 282; 1990: 32, figs. 1–6.—Trilles, 1975: 980, pl. I
(3–5).—Bruce, Lew Ton & Poore, 2002: 175.—Trilles & Bariche, 2006: 228: figs. 1–3.
Identity uncertain
Cymothoa frontalis.—Dana, 1853: 750, pl. 49, fig. 12 (a–b).—Trilles, 1994: 143.
Cymothoa frontalis.—Rameshkumar, Ravichandran, Sivasubramanian & Trilles, 2013: 42, fig. 1(B).
Type material. Syntypes: 2 ovig. ♀ (17, 22 mm, not examined), from the Indian Ocean; precise location and host
not known (MNHN–IU–2007–4059).
Diagnosis and description. We provide a brief diagnosis of the species. For a detailed description of the type
material see Hadfield (2012).
Body subparallel, 2.0 times as long as greatest width, smooth and polished in appearance, laterally sub-parallel,
widest at pereonite 5, most narrow at pereonite 1. Cephalon subtriangular, 0.7 times longer than wide, visible from
dorsal view. Frontal margin subacute. Eyes partially visible. Pereonite 1 anterolateral margins minute, not reaching
anterior margin of cephalon; pereonites 1–4 subequal in length, posterior margin moderately concave, with small
medial point; pereonites 5–7 subequal in length. Coxae 2–3 posteroventral margins subtruncate; 6 and 7 with
moderately acute carinae. Pleonites subequal in width, visible in dorsal view; pleonites posterior margin irregular.
Pleotelson subtruncate, 0.7 times as long as anterior width, anterior margin irregular, lateral margin straight,
posterior margin irregular, without median point. Antennula comprised of 8 articles. Antenna comprised of 9
articles. Pereopods 1 and 2 dactyli slender; carpus with straight proximal margin; merus proximal margin without
bulbous protrusion; basis superior proximal basis broad, without acute carinae. Pereopods 3–5 similar to pereopod
2, gradually increasing in size, without robust or simple setae. Pereopods 6 and 7 superior proximal basis with
broad and raised carinae; merus proximal margin without bulbous protrusion; dactyli stout. Uropod not extending
beyond pleotelson posterior margin, peduncle 0.9 times longer than exopod rami, apices narrowly rounded.
Size. Ovigerous females: 24–28 mm; non-ovigerous female: 20 mm (Schioedte & Meinert 1884).
Remarks. Cymothoa frontalis has a narrow and elongate body; subtriangular cephalon not immersed in
pereonite 1; pereonite 1 with minute anterolateral margins; posterior margins of pereonites weakly trisinuate;
pereonite 7 deeply arched and appearing to overlap pleonites 1–3; pleon narrow; pleonites subequal in width;
uropods half the length of the pleotelson; and pleotelson posterior margin subtruncate. Cymothoa frontalis also
appears to have the posterolateral margins of pereonites more curved than Trilles’s (1975) specimen.
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Cymothoa frontalis closely resembles C. indica in the subtriangular cephalon and pereonite 1with minute
anterolateral margins that do not reach beyond the anterior half of the cephalon. Cymothoa indica differs from C.
frontalis in having a moderately broad and weakly subparallel body [2.2 times longer than wide in Hale’s (1926)
illustration and 2.3 times longer than wide in Trilles & Bariche’s (2006) illustrations of C. indica]; uropod reaching
or extending beyond pleotelson posterior margin and pereopod 7 basis without raised carina.
Dana’s (1853) illustration of C. frontalis showed a rounded rostrum, coxae on pereonites 3–7 visible from
dorsal view, and posterior margin of pereonites evenly curved. Nearly all of Dana’s collections were lost with the
wreck of the USS Peacock on the bar of the Colombia River (Dana 1852b), so the identity of his specimen(s)
cannot be established and we have excluded the record from the synonymy.
A single female specimen of C. frontalis from one of 14 Strongylura leiura (Bleeker, 1850) was recorded from
Mathupettai, India (Rameshkumar et al. 2013). The Indian specimen is very similar to the syntype in the
subtriangular cephalon not being immersed in pereonite 1 and pereonite 1 with minute anterolateral margins that do
not reach beyond the anterior half of the cephalon, but differs from the syntype in having a more rounded
pleotelson posterior margin, uropodal rami not reaching half of pleotelson length, and a subtruncate cephalon
anterior margin. This is the only specimen reported from Strongylura (family Belonidae) and without further
description of the Indian specimen, we regard this record as of uncertain identity.
Distribution. Known from the Indian Ocean (White 1847; Schioedte & Meinert 1884; Trilles 1975);
Singapore (Heller 1868); Bangkok (Schioedte & Meinert 1884; Trilles 1975); and off west coast of Australia
(Avdeev 1978b).
Hosts. Only known from the family Cyclopteridae: on the gills of Cyclopterus (see White 1847).
Cymothoa hermani Hadfield, Bruce & Smit, 2011
Figures 14–15
Cymothoa hermani Hadfield, Bruce & Smit, 2011: 57, figs. 1–7.—Hadfield, Bruce & Smit, 2013: 163.
Type material. Holotype: 1 ♀ (ovig. 28 mm; not examined), Miwi Island, Kiwani Bay, Zanzibar, Tanzania,
06°21’S, 39°20’E, 26 April 2008, host marbled parrotfish Leptoscarus vaigiensis (Quoy & Gaimard, 1824), coll.
H. Van der Bank (SAMC A47890).
Paratypes: 1 ovig. ♀ (23 mm), 2 mature ♂ (14, 15 mm; not examined), same data as holotype (SAMC
A47891).
Material examined. Southeastern Queensland material. 1 ♀ ovig. (37 mm), off Gladstone, March 1979, from
parrotfish, coll. A. Kilvert (MTQ W8961). 1 ♂ (18 mm), off Gladstone, March 1971 or 1979, from parrotfish, coll.
A. Kilvert (MTQ W34275).
Diagnosis and description. Hadfield et al. (2011, 2013) provided figures and a detailed description of the type
material. We include here illustrations and a brief diagnosis of the Australian female specimen (MTQ W8961).
Body ovoid, 1.8 times as long as greatest width, dorsal surface rugose, laterally convex, widest at pereonite 4,
most narrow at pereonite 1. Cephalon subtriangular, 1.2 times longer than wide, moderately visible from dorsal
view. Frontal margin rounded, ventrally folded. Eyes absent. Pereonite 1 anterolateral margins broad, reaching
anterior margin of cephalon; pereonites 1–4 subequal in length, posterior margin moderately linear; pereonites 5–7
subequal in length. Coxae 2–3 posteroventral margins subtruncate; 5 and 6 with broadly acute carinae. Pleonites 1–
5 progressively wider, visible in dorsal view; pleonites posterior margin irregular. Pleotelson 0.6 times as long as
anterior width, anterior margin irregular, lateral margin convex, posterior margin rounded, without median point.
Antennula comprised of 6 articles. Antenna comprised of 7 articles. Pereopods 1 and 2 dactyli slender; carpus with
straight proximal margin; merus proximal margin without bulbous protrusion; basis superior proximal basis
smooth, not raised, without acute carinae. Pereopods 3–5 similar to pereopod 2, gradually increasing in size,
without robust or simple setae. Pereopods 6 and 7 superior proximal basis without broad and raised carinae; merus
proximal margin without bulbous protrusion; dactyli narrow. Uropod half as long as pleotelson length; peduncle
1.1 times as long as exopod; exopod 1.3 times as long as endopod, apices narrowly rounded.
Colour. Ivory white to yellowish-tan.
Size. Ovigerous females: 23–37 mm; adult males 14–18 mm (Hadfield et al. 2011 and present study).
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FIGURE 14. Cymothoa hermani ovigerous female (37 mm; MTQ W8961). A, dorsal view; B, ventral view of mouthpart; C,
front view of pereonite 1 and cephalon; D, dorsal view of pleotelson; E, lateral view.
Remarks. Cymothoa hermani can be identified by the unique bulbous ornamentation on pereonite 1, pereonite
1 anterolateral margins rounded and extending past rostrum, long and slender dactyli and the numerous lobes on
pleopods 4 and 5 in the ovigerous female (Hadfield et al. 2011, 2013).
The Australian specimens and Tanzanian holotype are similar in having a subtriangular cephalon, pereonite 1
anterolateral margins rounded and extending past rostrum and pleonites subparallel. Differences include the
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Australian specimens having a more ovoid body shape (compared to the subparallel lateral margins), pereonite 1
anterolateral margins reaching rostrum (compared to pereonite 1 producing past rostrum), the more acute
posteroventral margins of coxae 4–7 (compared to the straight and smooth margins), pereopods 6 and 7 with a less
developed carinae basis, and the rounded pleotelson posterior margin (compared to the subquadrate pleotelson
posterior margin). Male specimens from Australia (MTQ W34275) and Hadfield’s et al. (2011) paratype are
similar.
Cymothoa borbonica Schioedte & Meinert, 1884 and C. epimerica are the most similar species to C. hermani
in the subtriangular cephalon, pereonite 1 anterolateral margins reaching rostrum, and the ovate body shape.
Cymothoa borbonica has no bulbous ornamentation on pereonite 1, the lengths of pereonites 6 and 7 are more than
half the length of pereonite 4, acute posterior margins on coxae 4–7, sharp carinae basis on pereopods 6 and 7 and
the body is dorsomedially flattened (compared to the medial ridge on pereonites 2–4 on C. hermani, visible from
the lateral view). Cymothoa epimerica has posterior margins of coxae acute, basis of pereopods 6 and 7 acute and
dorsally visible, pleon trisinuate and pleopod margins appearing irregular.
FIGURE 15. Cymothoa hermani ovigerous female (37 mm; MTQ W8961). A, antennula; B, antenna; C–F, pereopods 1, 2, 6, 7
respectively; G, uropod.
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Distribution. Known from southeastern Queensland and the Northern Territory, Australia (present study) and
Tanzania, Africa.
Hosts. Known from the host from the family Scaridae (Hadfield et al. 2011; present study). The host
Leptoscarus vaigiensis is known to occur from the northern Red Sea to South Africa and eastwards to Japan and
New Zealand (Hadfield et al. 2011).
Cymothoa indica Schioedte & Meinert, 1884
Cymothoa Indica Schioedte & Meinert, 1884: 250, tab. VIII (Cym. XXVI) figs. 1–4.
Cymothoa indica.—Chilton, 1924: 887.—Hale, 1926: 212, fig. 8 (a–k).—McNeill, 1926: 318.—Nierstrasz, 1931: 133, fig. 1,
pl. 10 (figs. 5–8).—Avdeev, 1978b: 282; 1982b: 69.—Trilles, 1975: 981, pl. I (6, 7); 1994: 144; 2008: 23.—Veerapan &
Ravichandran, 2000: 1.—Kensley, 2001: 232.—Bruce, Lew Ton & Poore, 2002: 175.—Rajkumar, Santhanam & Perumal,
2004: 113.—Rajkumar, Perumal & Trilles, 2005a: 87, figs. 1, 2.—Rajkumar, Vasagam, Perumal & Trilles, 2005b: 269.—
Trilles & Bariche, 2006: 223, figs. 1–3.—Ravi & Rajkumar, 2007: 251, fig. 2.—Jones, Miller, Grutter & Cribb, 2008:
477.— Trilles, Ravichandran & Rameshkumar, 2011: 446.
Uncertain identity
Cymothoa sp. Monod, 1934: 14, pl. 28 (a–b), pl. 30 (d).
Cymothoa indica.—Panikkar & Aiyar, 1937: 429.—Rameshkumar & Ravichandran, 2010: 67, fig. 1.—El-Shahawy &
Desouky, 2010: 107, fig. 1 (a, b).—Al-Zubaidy & Mhaisen, 2014: 58, figs. 1, 2.
Type material. The syntypes (1 ovig. ♀, 20 mm; 1 immature ♂, 9 mm) were collected from Bangkok from an
unknown host (Schioedte & Meinert 1884).
Requests were made for the material but it is not held in the collection of the Zoological Museum, Museum für
Naturkunde, Berlin (Oliver Coleman, personal communication) nor the Zoological Museum of Copenhagen,
Denmark (Jørgen Olesen, personal communication).
Material examined. Northwestern and northeastern Queensland material: 1 ovig. ♀ (22 mm), 1 immature ♂
(11 mm), Stn. 506, Gulf of Carpentaria, 12°36.8’S, 141°18.4’E, December 1990, trawled J. Smith, FRV “Southern
Surveyor”, 36.5 m depth, from silvermouth trevally Ulua aurochs (Ogilby, 1915), coll. CSIRO (MTQ W17481). 1
ovig. ♀ (20 mm), Stn. 470, southeastern Gulf of Carpentaria, 16°77.83’S, 139°52.17’E, 7 m depth, 14 December
1963, from Cynoglossus sp., Cat. # 470, coll. CSIRO Fisheries on trawler “Rama” (AM P89834). 2 ovig. ♀ (20, 26
mm), Bowen Harbour, Port Denison, 20°02.0’S, 148°25.0’E, year 1926, coll. E. H. Rainford (AM P8574). 2
mature ♂ (10, 12 mm), Bowen, Port Denison, 20°05.0’S, 148°25.0’E, coll. E. H. Rainford (AM P9587). 1 ♀ (15
mm), Bowen Harbour, Port Denison, 20°02.0’S, 148°25.0’E, year 1925, coll. E. H. Rainford (AM P8484). 2 ovig.
♀ (16, 18 mm), 1 mature ♂ (15 mm), Bowen, from whiting, coll. E. H. Rainford (MTQ W197). 1 ovig. ♀ (20 mm),
1 immature ♂ (10 mm), Bowen Harbour, Port Denison, 20°02.0’S, 148°25.0’E, year 1926, from Sillago sp., coll. E.
H. Rainford (AM P8573). 1 mature ♂ (15 mm), off Cooktown, November 1975, from trevally, coll. S. Tanner
(MTQ W5745). 1 non-ovig. ♀ (16 mm), 1 mature ♂ (12 mm), Townsville, from inside fish (MTQ W10273).
Central Queensland material: 1 nonͲovig. ♀ (19 mm), 1 immature ♂ (10 mm), Yeppoon, 23°08.0’S,
150°41.0’E, 24 June 1991, from bumpnose trevally Carangoides hedlandensis (Whitley, 1934), coll. G. Monteith
(MTQ W17483). 1 immature ♂ (10 mm), Northwest Island, 23°03.0’S, 151°07.0’E, December 1925, coll. G. P.
Whitley (AM P8559). 1 ovig. ♀ (30 mm), Northwest Island, 23°03.0’S, 151°07.0’E, 27.4 m depth, December
1929–January 1930, coll. Mel Ward (AM P89954). 1 immature ♂ (6 mm), Port Curtis, 23°04.0’S, 150°05.0’E (AM
P9600).
Southeastern Queensland material: 2 ovig. ♀ (15, 17 mm), 1 immature ♂ (6 mm), Stn. 4, Mary River, 12
October 1981, from tongue of Sillago sp. (MTQ W10429). 1 immature ♂ (8 mm), Mary River, 14 April 1982
(MTQ W11015). 1 ovig. ♀ (23 mm), 1 immature ♂ (10 mm), Moreton Bay, 27°17.0’S, 153°15.0’E, year 2005,
from sand sillago Sillago ciliata Cuvier, 1829, GENBANK # E422801, coll. Mieke Burger (MTQ W28288)(see
Jones et al. 2008). 1 ovig. ♀ (20 mm), No. 77, Southport, year 1920, from the gills of long tom, coll. R. Pohlman
(MTQ W5761). 2 ovig. ♀ (18, 24 mm), 1 ♂ (10 mm), Hollywell, Southport, 8 April 1953, from gill slits of eel,
coll. Hale (MTQ W6005).
Non-Australian material: 1 ♀ (15 mm), Vanuatu, Erromango, Port Narvin, 18°74.1’S, 169°21.1’E, 28 May
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1996, coll. M. M Grouther and J. Williams (AM P89837). 1 ♀ (23 mm), examined by Hale without other available
data (AM P89832).
Colour. Present material ranges from pale yellow to dark brown. Some of the ovigerous female specimens
from Sillago ciliata are darker at the anterior region of the body and are progressively lighter posteriorly.
Size. Ovigerous females: 15–30 mm; males: 6–15 mm (present material).
Remarks. Cymothoa indica has a subtriangular cephalon; pereonite 1 anterolateral margins minute and not
reaching the middle of the cephalon; pereopod 7 ischium inferior distal margin with a distinctive lobe; pleotelson
posterior margin subtruncate; and uropodal rami of similar length as pleotelson posterior margin (for figure
examples see Hale 1926; Trilles 1975; Trilles & Bariche 2006). The present study showed that the eyes vary from
being either absent or moderately visible, pleotelson similar or wider than pereonite 7, cephalon deeply embedded
in pereonite 1 or dorsally visible, and pereopod 7 basis with moderately or highly raised carina.
Cymothoa indica resembles Cymothoa plebeia Schioedte & Meinert 1884 and C. frontalis in the body size,
cephalon and pleotelson morphology. Cymothoa frontalis is identified by the non-linear posterior margins of
pereonites; pereonites posterolateral margins convex; and dactyli of pereopods 1 and 2 slender and long, and nearly
touching the merus. Cymothoa plebeia is recognised by the lack of the prominent ischium lobe of C. indica and the
smooth pereon posterior margins.
We regard the two records of C. indica from the Red Sea (El-Shahawy & Desouky 2010; Al-Zubaidy &
Mhaisen 2014) as doubtful. El-Shahawy & Desouky (2010) provided figures of only pereopod 7, “showing the
characteristic lobe on the postero-angle of the ischium”, however, the figures portray a distinct protrusion on the
merus and not the ischium, which is not seen in C. indica. Al-Zubaidy & Mhaisen’s (2014) specimens [reported
from blue spot mullet Moolgarda seheli (Forsskål, 1775)] would appear similar to C. eremita in the subtruncate
cephalon and pereonite 1 reaching half the length of cephalon.
Trilles et al. (2011) mentioned that Panikkar & Aiyar (1937) collected 51 Cymothoa indica specimens from
two localities near the Madras coast (the Adyar backwaters and four miles up the river from Adyar) from several
host species (mainly the freshwater orange chromide cichlid Etroplus maculatus (Bloch, 1795) and pearlspot
cichlid Etroplus suratensis (Bloch, 1790)). Without a description or access to the specimens, we exclude this record
from synonymy. We also agree withTrilles et al. (2011) that the record of C. indica on tilapia Oreochromis
mossambicus (Peters, 1852) by Rameshkumar & Ravichandran (2010) is a misidentification and that record is also
removed from synonymy.
Distribution. Known from southeastern Pacific and Indian Ocean: Bangkok (Schioedte & Meinert 1884);
Beirut (see Trilles & Bariche 2006); India (Chilton 1924; Veerapan & Ravichandran 2000; Rajkumar et al. 2004,
2005a, 2005b; Ravi & Rajkumar 2007; Trilles & Bariche 2006; Trilles et al. 2011); Australia (Hale 1926; McNeill
1926; Jones et al. 2008); Indonesia (Nierstrasz 1931; Trilles 2008); Vietnam (Trilles 1975).
Hosts. Current studies are from host families Carangidae, Cynoglossidae, Sillaginidae and an unknown eel.
Previously reported from host of the family Bagridae: long whiskers catfish, Mystus gulio (Hamilton, 1822);
Belonidae: spottail needlefish, Strongylura strongylura (van Hasselt, 1823) (see Rajkumar et al. 2004); Clupeidae:
Bloch's gizzard shad Nematolosa nasus (Bloch, 1795); Gobiidae: maned gobi, Oxyurichthys macrolepis (Bleeker,
1849) (see Ravi & Rajkumar 2007), goby Glossogobius giuris (Hamilton, 1822) (see Chilton 1924); Latidae:
larvae of barramundi, Lates calcarifer (Bloch, 1790) (see Rajkumar et al. 2005a); Siganidae: streaked spinefoot
Siganus javus (Linnaeus, 1766) (see Rajkumar et al. 2005b); Sparidae:common pandora Pagellus erythrinus
(Linnaeus, 1758) (see Trilles & Bariche 2006); Sphyraenidae: yellowstripe barracuda Sphyraena chrysotaenia
Klunzinger, 1884 (see Trilles & Bariche 2006), obtuse barracuda Sphyraena obtusata Cuvier, 1829 (see Veerapan
& Ravichandran 2000; Trilles & Bariche 2006); Synodontidae: snakefish Synodus myops (Forster, 1801)
(previously Trachinocephalus myops) (see Veerapan & Ravichandran 2000; Trilles & Bariche 2006).
Cymothoa parupenei Avdeev, 1979
Figures 16–17
Cymothoa parupenei Avdeev, 1979a: 228, pl. 4–5.—Trilles, 1994: 147.—Kensley, 2001: 233.—Bruce, Lew Ton & Poore,
2002: 175.—Trilles & Bariche, 2006: 228.Rameshkumar, Ravichandran, Sivasubramanian & Trilles, 2013: 42, fig.
1(A).
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Material examined. Paratype: 1 mature ♂ (13 mm), from the Indian Ocean off Australia, from blackspot goatfish
Parupeneus spilurus (Bleeker, 1854) (TINRO АPK 75027). A loan request was made for the holotype (TINRO
AGK 75026) but the specimen could not be located.
Male Length 13.5 mm width 7 mm (paratype).
FIGURE 16. Cymothoa parupenei, male paratype (13 mm; APK 75027). A, dorsal view; B, front view of pereonite 1 and
cephalon; C, dorsal view of pleotelson; D, lateral view.
Body subrectangular, 1.8 times as long as greatest width, dorsal surface smooth, widest at pereonite 5, most
narrow at pereonite 1. Cephalon 0.5 times longer than wide, visible from dorsal view, semi-circular. Frontal
margin rounded to form blunt rostrum. Eyes partially visible. Pereonite 1 anterolateral margins broad, nearly
reaching anterior margin of cephalon; pereonites 1–6 posterior margin smooth, laterally concave; pereonites 5–7
subequal in length. Coxae 2–7 posteroventral margins rounded. Pleonite 1 similar length to pleonites 2–5, visible in
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dorsal view; pleonites posterior margin moderately irregular; pleonites 3–5 progressively wider. Pleotelson 0.5
times as long as anterior width, anterior margin not trisinuate, lateral margin weakly concaved, posterior margin
subtruncate and irregular, without median point.
Antennula comprised of 8 articles; peduncle articles 1 and 2 distinct and articulated; article 2 1.3 times as long
as article 1; article 3 0.6 times as long as combined lengths of articles 1 and 2, 0.9 times as long as wide. Antenna
comprised of 9 articles, peduncle article 3 2.0 times as long as article 2, 1.0 times as long as wide; article 4 1.1
times as long as wide; article 5 0.8 times as long as article 4, terminal article without setae.
FIGURE 17. Cymothoa parupenei, male paratype (13 mm; APK 75027). A, antennula; B, antenna; C–F, pereopods 1, 2, 6, 7
respectively; G, uropod.
Pereopod 1 basis 1.7 times as long as greatest width, superior proximal margin smooth, without raised carina;
ischium 0.6 times as long as basis; merus proximal margin without bulbous protrusion; carpus with straight
proximal margin; propodus 1.4 times as long as wide; dactylus narrow, 1.5 times as long as propodus, 2.6 times as
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long as basal width. Pereopod 2 basis 1.6 times as long as greatest width, superior proximal margin smooth,
without raised carina; propodus 1.1 times as long as wide; dactylus 1.4 times as long as propodus. Pereopods 3–5
similar to pereopod 2, gradually increasing in size, without robust or simple setae. Pereopod 6 basis 1.2 times as
long as greatest width, superior proximal margin smooth, with sharp and raised carina; ischium 0.6 times as long as
basis, propodus 1.3 times as long as wide, dactylus 2.1 times as long as propodus. Pereopod 7 basis 1.3 times as
long as greatest width; ischium 0.9 times as long as basis, inferior margin with slight bulbous protrusion; merus
proximal margin without bulbous protrusion, merus 0.3 times as long as ischium, 0.5 times as long as wide; carpus
0.3 times as long as ischium, without bulbous protrusion, 0.6 times as long as wide; propodus 0.7 times as long as
ischium, 0.6 times as long as wide; dactylus stout, 1.8 times as long as propodus, 2.3 times as long as basal width.
Uropod reaching posterior margin of pleotelson, peduncle 1.1 times as long as greatest width, 0.8 times as long
as exopod, lateral margin without setae, marginal setae absent, apices broadly blunt, lateral margin convex, mesial
margin convex. Exopod extending past endopod, 3.6 times as long as greatest width, apically rounded, lateral
margin straight, terminating without setae, mesial margin weakly straight. Endopod 3.0 times as long as greatest
width, apically rounded, lateral margin weakly convex, terminating without setae, mesial margin straight.
Colour. Light brown in ethanol (present study), ivory white in Rameshkumar et al. (2013).
Remarks. Cymothoa parupenei can be identified by the wide anterolateral margins on pereonite 1 extending
beyond the anterior margins of cephalon; semi-circular cephalon anterior margin (subtruncate cephalon in
holotype); pleonite 1 almost entirely overlapped by pereonite 7; uropods reaching posterior margin of the
pleotelson; pereopod 7 basis with smooth and moderately raised carina, ischium inferior distal margin with slight
bulbous projection; and pleotelson 2.1 times wider than long (2.6 times wider than long in holotype). Avdeev’s
(1979a) illustration of the ovigerous female showed a maxillula with 3 terminal setae; lateral lobe of maxilla partly
fused to mesial lobe, with 1 recurved seta on each mesial and lateral lobe; a weakly segmented maxilliped, terminal
article 3 with 3 recurved robust setae; pleopod 1 rami of subequal size; endopod mesial margin deeply oblique and
distally straight, lateral margins deeply convex, with proximal margins extending beyond peduncle. The male
pleopod 2 (illustrated by Avdeev 1979a, fig. 5) has an appendix masculina half the exopod length; exopod lateral
margin weakly convex, with medial margin distally rounded and weakly convex.
Cymothoa parupenei resembles Cymothoa oestrum (Linnaeus, 1758) in the semi-circular cephalon, pereonites
1–6 posterior margin smooth and pleonites 3–5 becoming progressively wider. Cymothoa oestrum differs from C.
parupenei in having anterolateral margins of pereonite 1 that project forward (compared to the more broad and
concave anterolateral margins of C. parupenei) and pereopods 5–7 basis have sharp carinae visible in lateral view
(see Thatcher et al. 2003) compared to blunt carina of Avdeev’s (1979a) specimens. Cymothoa parupenei is host
specific to Mullidae and has an Indo-Pacific distribution, whereas C. oestrum is known from a variety of fishes and
has a western Atlantic distribution (Thatcher et al. 2003).
Rameshkumar et al. (2013) recorded a prevalence of 5.9% (1 in 17 hosts) captured at Nagapattinam, India.
Distribution. Reported from northwestern Australia (Avdeev 1979a; Kensley 2001) and India (Rameshkumar
et al. 2013).
Hosts. Known from family Mullidae: Parupeneus spilurus (see Avdeev 1979a) and sulphur goatfish Upeneus
sulphureus Cuvier, 1829 (see Rameshkumar et al. 2013).
Cymothoa propria Avdeev, 1979
Figures 18–21
Cymothoa propria Avdeev, 1979b: 50, pl. 1, 2.—Trilles, 1994: 147.—Bruce, Lew Ton & Poore, 2002: 176.—Trilles & Bariche,
2006: 228.
Material examined. Paratypes: 2 ovig. ♀ (15, 20 mm), collected from northern Australia, on yellow-stripe scad
Selaroides leptolepis (Cuvier, 1833) (TINRO АPK 75002–75005).
Note: The female holotype (TINRO AGK 75001) and male paratypes (TINRO APK 75006–75008) could not
be located, and our illustrations are based on female paratypes.
Ovigerous female Length 20 mm width 8 mm (paratype).
Body rhomboid, 2.1 times as long as greatest width, dorsal surface smooth, widest at pereonite 5 and 6.
Cephalon subtriangular, 1.1 times longer than wide, slightly visible from dorsal view, not immersed in pereonite 1.
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Frontal margin rounded to form blunt rostrum, ventrally folded. Eyes partially distinct. Pereonite 1 anterolateral
margins minute, reaching posterior margins of eyes; posterior margins of pereonites 3–6 irregular; pereonites 5–7
subequal in length, posterolateral margins arched. Coxae 2–7 posteroventral margins rounded. Pleonites subequal
in width, visible in dorsal view, pleonites posterior margins irregular. Pleotelson 0.5 times as long as anterior width,
anterior margin irregular, lateral margins weakly concave, posterior margin subtruncate, without median point.
Antennula comprised of 8 articles; peduncle articles 1 and 2 distinct and articulated; article 2 1.3 times as long
as article 1; article 3 0.8 times as long as combined lengths of articles 1 and 2, 1.2 times as long as wide. Antenna
comprised of 7 articles, peduncle article 3 1.4 times as long as article 2, 1.8 times as long as wide; article 4 1.2
times as long as wide; article 5 0.9 times as long as article 4, terminal article without setae, not extending to
posterior of pereonite 1.
FIGURE 18. Cymothoa propria, ovigerous female paratype (20 mm; 75002-75005). A, dorsal view; B, front view of pereonite
1 and cephalon; C, dorsal view of pleotelson; D, lateral view.
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FIGURE 19. Cymothoa propria, ovigerous female paratype (20 mm; 75002-75005). A, antennula; B, antenna; C, uropod; D–
G, pereopods 1, 2, 6, 7 respectively.
Pereopod 1 basis 1.1 times as long as greatest width, superior proximal margin smooth, with moderately raised
carina; ischium 0.6 times as long as basis; merus proximal margin without bulbous protrusion; carpus with straight
proximal margin; propodus 1.0 times as long as wide; dactylus narrow, 1.6 times as long as propodus, 2.4 times as
long as basal width. Pereopod 2 basis 1.4 times as long as greatest width, superior proximal margin smooth,
without raised carina; propodus 1.3 times as long as wide; dactylus 1.5 times as long as propodus. Pereopods 3–5
similar to pereopod 2, gradually increasing in size, without robust or simple setae. Pereopod 6 basis 1.0 times as
long as greatest width, superior proximal margin smooth, without raised carina; ischium 0.7 times as long as basis,
inferior distal margin with slight protrusion; propodus 0.9 times as long as wide, dactylus 2.0 times as long as
propodus. Pereopod 7 basis 0.9 times as long as greatest width, superior proximal margin smooth, with raised
carina; ischium 0.7 times as long as basis, with bulbous protrusion; merus proximal margin with slight bulbous
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protrusion, 2.9 times as long as ischium, 2.0 times as long as wide; carpus 0.2 times as long as ischium, with slight
bulbous protrusion, 0.4 times as long as wide; propodus 0.5 times as long as ischium, 1.0 times as long as wide;
dactylus stout, 2.0 times as long as propodus, 2.5 times as long as basal width.
Uropod not extending beyond posterior margin of pleotelson; peduncle 2.2 times as long as greatest width, 0.7
times as long as exopod, lateral margin without setae, marginal setae absent, lateral margin straight, mesial margin
straight. Exopod extending past endopod, 6.0 times as long as greatest width, apically rounded, lateral margin
straight, terminating without setae, mesial margin weakly straight. Endopod 2.8 times as long as greatest width,
apices narrowly acute, lateral margin weakly convex, terminating without setae, mesial margin straight.
Colour. Chestnut brown.
FIGURE 20. Cymothoa propria, ovigerous female paratype (15 mm; APK 75002-75005). A, dorsal view; B, front view of
pereonite 1 and cephalon; C, ventral view of mouthpart and pereopod 1 (dactylus and propodus); D, lateral view.
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FIGURE 21. Cymothoa propria, ovigerous female paratype (15 mm; APK 75002-75005). A–E, pereopods 1, 2, 3, 6, 7
respectively.
Remarks. Cymothoa propria has a rhomboid body that is widest at pereonites 5 and 6, pereonites 5–7
subequal in length; cephalon subtriangular and weakly immersed in pereonite 1; anterolateral margins on pereonite
1 minute; pleonites subequal in width; pleotelson posterior margin subtruncate, 2.2 times wider than long; uropodal
rami reaching posterior margin of pleotelson; pereopod 7 ischium inferior distal margin with bulbous protrusion
and superior proximal basis of pereopod 7 with raised carina. Avdeev’s (1979b) illustration of the female holotype
included a simple maxillula, with 3 terminal setae; maxilla with 8 and 3 recurved setae on mesial lobe and lateral
lobe respectively; maxilliped weakly segmented, terminal article 3 with five recurved setae; pleopod 1 exopod and
endopod subequal in size; endopod mesial and distal margin straight, with deeply convex lateral margins, and
proximal margins not extending beyond peduncle. Avdeev’s (1979b) male illustration differs from the female by
having visible eyes, subequal pereonite length, minute anterolateral margins of pereonite 1, rounded posterior
margin of pleotelson and appendix masculina present on pleopod 2.
Cymothoa propria resembles C. indica and C. plebeia Schioedte & Meinert, 1884 in the subtriangular
cephalon and pleonites subequal in width. Cymothoa propria differs from both of those species in having a highly
raised carina on pereopod 7 basis, 1.1 times wider than long (compared to the 0.9 times wider than long basis in C.
indica and C. plebeia); pereopod 7 dactylus extending almost to the distal margin of carpus (compared to dactylus
of C. indica and C. plebeia that does not extend to distal margin of carpus) and the ventrally folded frontal lamina,
appearing to immerse the paired antennae.
Distribution. Northern Australia (Avdeev 1979b).
Hosts. Only known from Carangidae: Selaroides leptolepis (see Avdeev 1979b).
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Cymothoa pulchrum Lanchester, 1902
Figures 22–26
Cymothoa pulchrum Lanchester, 1902: 377, pl. 35, figs. 8, 8a.—Monod, 1924: 100.—Trilles, 1975: 991, pl. II (16); 1994:
148.—Galzin & Trilles, 1979: 257, figs. 1–52.—Avdeev, 1982b: 69.—Williams, Bunkley-Williams & Dyer, 1996: 1, fig.
5.—Kensley, 2001: 233.—Trilles & Bariche, 2006: 228.
Cymothoa pulchra.—Nierstrasz, 1915: 92, pl. 3 (fig. 11), pl. 4 (figs. 12, 13); 1931: 133, pl. 10 (figs. 1–4).—Monod, 1934: 12,
pl. 26 (a–b), 30 (a).—Shiino, 1951: 81, 85, figs. 4 (a–h).—Avdeev, 1978b: 281.—Saito, Itani & Nunomura, 2000: 65.—
Bruce, Lew Ton & Poore, 2002: 176.Nagasawa & Uyeno, 2012: 139, fig. 1.
Not Cymothoa pulchra.—Yu & Li, 2003a: 228, fig. 5; 2003b: 267 [=Cymothoa epimerica].
Type material. Lanchester (1902) reported a 35 mm female specimen from Pulau Bidan, northern Straits of
Malacca, Malay Peninsula, from an unknown host. The marine isopods identified by Lanchester (1902) were part
of the collection from the Skeat Expedition to the Malay Peninsula, of which most (if not all) are deposited at the
University Museum of Zoology, Cambridge (UMZC) (University of Cambridge 2014). The University Museum of
Zoology’s online catalogue shows two Cymothoa specimens: Cymothoa sp. [UMZC I.51104] and Cymothoa
stromatei [UMZC I.51106]. Cymothoa sp. was collected from the Baram River, (Sarawak; Borneo) by C. Hose, and
registered on the 11 June 1898, without additional information. Cymothoa stromatei had been described by
Lanchester’s (1902) and is registered as “Skeat Collection” on the 30
th
November 1899 in the museum’s catalogue.
There is no indication from the catalogue that the type for Cymothoa pulchrum is there, thus a loan request was
made to investigate all other available Cymothoa sp., but as UMZC is undergoing re-development, the zoological
collections were currently not accessible (Richard Preece, personal communication).
Material examined. Singaporean material: 1 immature ♂ (9 mm) (ZRC 2015.0042); 1 ovig. ♀ (19 mm), 1
immature ♂ (9 mm) (ZRC 2015.0043); 1 ovig. ♀ (23 mm), 1 immature ♂ (9 mm) (ZRC 2015.0044); 1 ovig. ♀ (16
mm), 1 immature ♂ (6 mm) (ZRC 2015.0045); 1 ovig. ♀ (20 mm), 1 immature ♂ (8 mm) (MTQ W34289); 1 ovig.
♀ (21 mm) (ZRC 2015.0046); all St. SW 77, mouth of Sungei Teris, OBS Camp 2, gill net sampling, 1°24.54’N,
103°56.18’E, 24 October 2012, from spotted green pufferfish Tetraodon nigroviridis Marion de Procé, 1822, coll.
Ng Heok Hee and Ronald Huys.
1 ovig. ♀ (17 mm), St. SW 116, off Pulau Tekong, gill net and tangle net sampling, 1°25.615’N, 104°04.316’E,
29 October 2012, from Tetraodon nigroviridis, coll. Tan Heok Hui and fisherman (MTQ W34290).
Ovigerous females Length 17 mm width 9 mm (non-dissected, MTQ W34290); length 23 mm width 9 mm
(dissected, ZRC 2015.0044).
Body subparallel, 2.1 times as long as greatest width, dorsal surface smooth, widest at pereonites 3–5, most
narrow at pereonite 1. Cephalon 0.3 times longer than wide, visible from dorsal view, subtruncate. Frontal margin
rounded to form blunt rostrum. Eyes partially visible. Pereonite 1 anterolateral margins broad, nearly reaching
cephalon anterior margin; pereonites 1–4 posterior margin irregular, subequal in length; pereonites 5–7 subequal in
length, pereonite 7 posterolateral margin arched. Coxae visible from dorsal view, coxae 2–4 posteroventral margins
subtruncate; 5–7 with moderately acute carinae. Pleon partially overlapped by pereonite 7; pleonites 1–4 subequal
in length and width, visible in dorsal view; pleonites posterior margin smooth; pleonite 5 posterior margin straight.
Pleotelson 0.6 times as long as anterior width, anterior margin moderately irregular, lateral margin weakly concave,
posterior margin rounded, without median point.
Antennula comprised of 7 articles; peduncle articles 1 and 2 distinct; article 2 0.6 times as long as article 1;
article 3 0.5 times as long as combined lengths of articles 1 and 2, 1.0 times as long as wide; extending to middle of
cephalon. Antenna comprised of 7 articles, peduncle article 3 0.8 times as long as article 2, 1.0 times as long as
wide; article 4 0.8 times as long as wide; article 5 0.8 times as long as article 4, terminal article without setae.
Labrum fleshy, lateral margins convex, anterior margin broadly acute, without small median point.
Pereopod 1 basis 1.8 times as long as greatest width, superior proximal margin smooth, without raised carina;
ischium 0.6 times as long as basis; merus proximal margin without bulbous protrusion; carpus with straight
proximal margin; propodus 2.0 times as long as wide; dactylus slender, 1.1 times as long as propodus, 4.4 times as
long as basal width. Pereopod 2 basis 1.3 times as long as greatest width, superior proximal margin smooth,
without raised carina; propodus 1.7 times as long as wide; dactylus 1.6 times as long as propodus. Pereopod 6 basis
2.2 times as long as greatest width, superior proximal margin smooth, without raised carina; ischium 0.6 times as
long as basis, propodus 2.1 times as long as wide, dactylus 1.3 times as long as propodus. Pereopod 7 basis 1.1
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times as long as greatest width, superior proximal margin moderately raised carina; ischium 0.5 times as long as
basis, with bulbous protrusion; merus proximal margin with slight bulbous protrusion, merus 0.3 times as long as
ischium, 0.4 times as long as wide; carpus 0.5 times as long as ischium, without bulbous protrusion, 0.7 times as
long as wide; propodus 0.9 times as long as ischium, 1.3 times as long as wide; dactylus slender, 2.0 times as long
as propodus, 4.2 times as long as basal width.
FIGURE 22. Cymothoa pulchrum, ovigerous female Singapore (17 mm; MTQ W34290). A, dorsal view; B, ventral view of
mouthpart; C, front view of pereonite 1 and cephalon; D, dorsal view of pleotelson; E, lateral view.
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FIGURE 23. Cymothoa pulchrum, ovigerous female Singapore (23 mm; ZRC 2015.0044). A, antennula; B, antenna; C,
uropod; D–G, pereopods 1, 2, 6, 7 respectively.
Pleopods without setae, depression present on central dorsal surface of pleopods exopods 2, 3 and 5; basal
projections present, increasing in size from pleopods 1–5; exopod proximal mesial margins extending near
peduncle, increasing in size from pleopods 1–5; exopod larger than endopod. Pleopod 1 exopod 1.0 times as long
as wide, lateral margin convex, distally broadly rounded, mesial margin straight; peduncle 3.0 times as wide as
long, without retinaculae. Pleopods 2–5 lateral margins becoming strongly convex, large fleshy folds present.
Uropod not extending beyond posterior margin of pleotelson; peduncle 1.1 times as long as greatest width, 1.0
times as long as exopod rami, lateral margin straight, mesial margin straight. Exopod subequal length to endopod,
3.3 times as long as greatest width, apically rounded, lateral margin convex, terminating without setae, mesial
margin concave. Endopod 3.2 times as long as greatest width, apically rounded, lateral margin weakly straight,
terminating without setae, mesial margin straight.
Male. Length 8 mm, width 4 mm (non-dissected, MTQ W34289); length 9 mm, width 4 mm (dissected, ZRC
2015.0042).
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FIGURE 24. Cymothoa pulchrum, ovigerous female Singapore (23 mm; ZRC 2015.0044). A–E, dorsal view of pleopods 1–5
respectively; F–I, ventral view of pleopods 2–5 respectively.
Body 2.2 times as long as greatest width. Cephalon subtriangular, 0.4 times longer than wide, visible from
dorsal view, not deeply immersed in pereonite 1. Eyes partially visible. Pereonite 1 anterolateral minute, not
reaching half cephalon length; posterior margins of pereonites 1–5 straight and smooth. Coxae 2–7 posteroventral
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margins rounded. Pleonites subequal in width, not overlapped by pereonite 7; pleonites 1–4 posterior margins
linear, pleonite 5 posterior margin weakly bisinuate. Pleotelson weakly subtruncate. Antennula comprised of 9
articles; subequal in length to antenna. Antenna comprised of 8 articles.
Pleopods exopod and pleopod margins smooth; pleopod 2 appendix masculina with parallel margins, 0.8 times
as long as endopod, distally acute, pleopod 5 endopod with thick fleshy folds.
FIGURE 25. Cymothoa pulchrum, male Singapore (8 mm; MTQ W34289). A, dorsal view; B, ventral view of mouthpart; C,
antennula; D, antenna; E, front view of pereonite 1 and cephalon; F, uropod; G, dorsal view of pleotelson; H, lateral view.
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FIGURE 26. Cymothoa pulchrum, male Singapore (9 mm; ZRC 2015.0042). A–D, pereopods 1, 2, 6, 7 respectively; E–I,
pleopods 1–5 respectively.
Colour. Pale tan.
Size. Ovigerous females: 34–39 mm; non-ovigerous females: 21–29 mm; males: 7–20 mm; pullus stage: 10
mm (Lanchester 1902; Nierstrasz 1915; Monod 1934; Trilles 1975; Galzin & Trilles 1979; present study).
Remarks. Cymothoa pulchrum has a subparallel body, widest at pereonites 3–5; cephalon anteriorly
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subtruncate; wide and subtruncate pereonite 1 anterolateral margins which is nearly reaching rostrum; pleon
partially overlapped by pereonite 7; pleotelson posterior margin round; basis of pereopods 6 and 7 superior
proximal margin with raised carinae; pereopod 7 with prominent lobe on ischium; coxae visible from dorsal view
and uropods reaching half the pleotelson length (visible from ventral view). The male specimens differ from the
females in having a subtriangular cephalon that is not deeply immersed in pereonite 1, eyes partially visible
(compared to the females with eyes absent), pereonite 1 anterolateral margins minute and not reaching half
cephalon length, pleonites not subequal in width and not overlapped by pereonite 7, pleopod 2 appendix masculina
with parallel margins, and pleopod 5 endopod with thick fleshy folds.
Galzin & Trilles (1979) described and illustrated C. pulchrum at different life stages. Galzin & Trilles’s (1979)
female specimens differ from the Australian females in having a more highly raised carina on pereopod 7 basis,
straight inferior distal margin on pereopod 7 ischium, setae present on segments 5–7 on antennula and appendix
masculina present on pleopod 2. The males from Galzin & Trilles’s (1979) illustration varies from the Singaporean
material in having marginal setae on uropod peduncle; appendix masculina present on pleopod 2 and extending
beyond the endopod margin; and setae present on segments of antenna and antennula.
Cymothoa epimerica differs from C. pulchrum by the oval body, uropodal rami more narrow and slender,
cephalon subtriangular, coxae 5–7 posteroventral margins acute and visible in dorsal view, and pereopods 6 and 7
basis with sharp and acute carinae (compared to the raised carina without an acute edge). Cymothoa eremita differs
from C. pulchrum by having anterolateral margins of pereonite 1 more tapered, pleotelson more subtruncate, coxae
posteroventral margins rounded and not visible from dorsal view. Yu & Li (2003b) misidentified “C. pulchra” from
Hainan Island, South China Sea, their drawings allowing identification of their material as C. epimerica (see
remarks on C. epimerica).
Distribution. Known from the central and western Indo-Pacific region: Singapore (current material); Malaysia
(Lanchester 1902); Indonesia (Nierstrasz 1915, 1931); Sri Lanka (Monod 1924); Vietnam (Monod 1934; Trilles
1975); Japan (Shiino 1951; Saito et al. 2000; Nagasawa & Uyeno 2012); Australia (Avdeev 1978b); French
Polynesia and Fiji (Galzin & Trilles 1979).
Hosts. Present material from family Tetraodontidae. Previously reported from Carangidae: Caranx sp. (see
Monod 1924); Tetraodontidae: stellate puffer Arothron stellatus (Anonymous, 1798) see Monod 1934; Trilles
1975; Avdeev 1978b; Galzin & Trilles 1979; Nagasawa & Uyeno 2012) and guineafowl puffer Arothron meleagris
(Anonymous, 1798) (see Galzin & Trilles 1979); Diodontidae: longspined porcupinefish Diodon holocanthus
Linnaeus, 1758 (see Shiino 1951; Williams et al. 1996; Nagasawa & Uyeno 2012); spot-fin porcupinefish Diodon
hystrix Linnaeus, 1758 (see Williams et al. 1996; Galzin & Trilles 1979); black-blotched porcupinefish Diodon
liturosus Shaw, 1804 (see Williams et al. 1996) and spotfin burrfish Chilomycterus reticulatus (Linnaeus, 1758)
(see Nagasawa & Uyeno 2012).
Cymothoa rotunda Avdeev, 1979
Figures 27–28
Cymothoa rotunda Avdeev, 1979a: 223, pl. 1.—Trilles, 1994: 148.—Kensley, 2001: 233.—Bruce, Lew Ton & Poore, 2002:
176.—Trilles & Bariche, 2006: 228.
Material examined. Holotype: 1 ovig. ♀ (31 mm), from the Indian Ocean off Western Australia (precise locality
not given), from long-tailed catfish Euristhmus lepturus (Günther, 1864) (TINRO АGK 75010).
Ovigerous female Length 31 mm width 20 mm (holotype).
Body ovoid, 1.4 times as long as greatest width, dorsal surface rough, laterally dome-shaped, widest at
pereonite 4, most narrow at pereonite 1. Cephalon subtriangular, 0.9 times longer than wide, visible from dorsal
view. Frontal margin rounded to form blunt rostrum, ventrally folded. Eyes absent. Pereonite 1 anterolateral
margins broad, unique bulbous ornamentation; posterior margins of pereonites irregular; pereonites 5–7 subequal
in length, posterolateral margins arched. Coxae 2–7 posteroventral margins rounded. Pleonite 1 visible in dorsal
view; pleonites 3–5 posterior margin bisinuate, subequal in width. Pleotelson 0.4 times as long as anterior width,
lateral margin weakly convex (right lateral margin damaged exposing pleopods), posterior margin rounded,
without median point.
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FIGURE 27. Cymothoa rotunda, ovigerous holotype (31 mm; AGK 75010). A, dorsal view; B ventral view of mouthpart; C,
antennula; D, antenna; E, uropod; F, front view of pereonite 1 and cephalon; G, lateral view.
Antennula comprised of 7 articles; peduncle articles 1 and 2 distinct and articulated; article 2 1.3 times as long
as article 1; article 3 0.6 times as long as combined lengths of articles 1 and 2, 1.3 times as long as wide. Antenna
comprised of 8 articles, article 3 1.1 times as long as article 2, 1.0 times as long as wide; article 4 1.0 times as long
as wide; article 5 1.0 times as long as article 4, terminal article without setae, extending half of pereonite 1. Labrum
anterior margin broad and convex.
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FIGURE 28. Cymothoa rotunda, ovigerous holotype (31 mm; AGK 75010). A–D, pereopods 1, 2, 6, 7 respectively; E,
pleopod 1; F, dorsal view of pleotelson.
Pereopod 1 basis 1.7 times as long as greatest width, superior proximal margin smooth, without raised carina;
ischium 0.5 times as long as basis; merus proximal margin without bulbous protrusion; carpus proximal margin
straight; propodus 1.1 times as long as wide; dactylus narrow, 1.7 times as long as propodus, 3.1 times as long as
basal width. Pereopod 2 basis 1.6 times as long as greatest width, superior proximal margin smooth, without raised
carina; propodus 1.0 times as long as wide; dactylus 1.2 times as long as propodus. Pereopods 3–5 similar to
pereopod 2, gradually increasing in size, without robust or simple setae. Pereopod 6 basis 1.2 times as long as
greatest width, superior proximal margin smooth, with slightly raised carina; ischium 0.6 times as long as basis,
propodus 1.0 times as long as wide, dactylus 1.6 times as long as propodus. Pereopod 7 basis 1.6 times as long as
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greatest width, superior proximal margin smooth, with raised carina; ischium 0.5 times as long as basis, with
bulbous protrusion; merus proximal margin with slight bulbous protrusion, merus 0.5 times as long as ischium, 0.6
times as long as wide; carpus 0.2 times as long as ischium, with slight bulbous protrusion, 0.2 times as long as
wide; propodus 0.6 times as long as ischium, 1.2 times as long as wide; dactylus narrow, 1.8 times as long as
propodus, 2.8 times as long as basal width.
Pleopod 1 exopod 1.0 times as long as wide, lateral margin convex, distally broadly rounded, mesial margin
straight; endopod 1.1 times as long as wide, lateral margin weakly convex, distally broadly rounded, mesial margin
straight; peduncle 2.0 times as wide as long.
Uropod not extending beyond posterior margin of pleotelson; peduncle 1.7 times as long as greatest width, 0.7
times as long as exopod, lateral margin without setae, marginal setae absent, apices narrowly acute, lateral margin
straight, mesial margin straight. Exopod subequal in length to endopod, 2.9 times as long as greatest width, apically
rounded, lateral margin straight, terminating without setae, mesial margin weakly straight. Endopod 2.9 times as
long as greatest width, apically rounded, lateral margin weakly convex, terminating without setae, mesial margin
straight.
Colour. Greenish yellow.
Remarks. Cymothoa rotunda can be distinguished by the ovoid body (1.4 times longer than wide), widest and
longest at pereonite 4; cephalon subtriangular; frontal margin ventrally folded; pereonite 1 anterolateral margins
broad and tapers to a narrow point that reaches the rostrum, with unique bulbous ornamentation; pleon shape
appearing horizontally suboval; uropods not reaching pleotelson posterior margin; and pleotelson posterior margin
rounded. Avdeev’s (1979a) illustration of the ovigerous female also included: simple maxillula, with 3 terminal
setae; lateral lobe of maxilla partly fused to mesial lobe, with 1 and 2 recurved setae on mesial lobe and lateral lobe
respectively; weakly segmented maxilliped, terminal article 3 with 2 recurved setae; pleopod 2 with appendix
masculina present, exopod bigger than endopod, convex endopod mesial margin, distally rounded, deeply convex
lateral margins, without proximal margins extending beyond peduncle.
Cymothoa rotunda is similar to C. hermani and C. borbonica from the subtriangular cephalon and rounded
pleotelson posterior margin (and bulbous ornamentation in C. hermani). Cymothoa hermani differs from C.
rotunda from the less rounded body, 1.8 to 1.9 times longer than wider (compared to the 1.4 times longer than wide
body), dorsally visible coxae and trisinuate posterior margin of pleonites 2–5. The female paratype of C. hermani
has 4 robust setae on the maxillula and pleopod 2 without an appendix masculina (Hadfield et al. 2011) compared
to the 3 robust setae on maxillula and pleopod with an appendix masculina of C. rotunda (Avdeev 1979a).
Cymothoa borbonica does not have the unique bulbous ornamentation on pereonite 1, body sub-parallel, pleonites
not subequal in width, and coxae 2–7 posteroventral margins acute and dorsally visible.
Distribution. Western Australia (Avdeev 1979a); known only from the original description.
Hosts. Only known from the family Plotosidae: long-tailed catfish Euristhmus lepturus (Günther, 1864) (see
Avdeev 1979a).
Cymothoa vicina Hale, 1926
Figures 29–38
Cymothoa vicina Hale, 1926: 214, fig. 9 (a–g).—Nierstrasz, 1931: 136.—Avdeev, 1978b: 282.—Springthorpe & Lowry, 1994:
64.—Trilles, 1994: 149.—Bruce, Lew Ton & Poore, 2002: 174.—Trilles & Bariche, 2006: 228.
Cymothoa epimerica.—Trilles, 2008: 23 (SMF-572).
Material examined. Holotype: 1 ovig. ♀ (15 mm), Tweed River, NSW, 28°21.0’S, 153°17.0’E, from a mullet,
coll. Franks (AM P8590).
Australian material: 1 mature ♂ (15 mm), Mandurah, WA, 1984, from gills of cobbler Cnidoglanis
macrocephalus (Valenciennes, 1840), coll. C. M. A. Williams (MTQ W30410). 2 ovig. ♀ (23, 25 mm), 4 immature
♂ (4, 4, 9, 10 mm), Moreton Bay, QLD, 3 May 1977, from mullet Mugil sp. (MTQ W7302).
Digital images from the Senckenberg Research Institute: 3 ♀ ovig. (19,22, 23 mm), 1 immature ♂ (11 mm),
Red Sea (SMF-572).
Ovigerous female Length 15 mm, width 7 mm (holotype).
Body oblong, 2.1 times as long as greatest width, dorsal surface smooth, laterally subparallel, widest at
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pereonite 4 and pereonite 5, most narrow at pereonite 1, Cephalon subtruncate, 0.4 times longer than wide, visible
from dorsal view. Frontal margin without median point. Eyes oval with distinct margin, 0.4 times width of head.
Pereonite 1 anterior margin moderately produced medially, anterolateral margins broad, reaching past eyes;
pereonites 1–4 posterior margin smooth, progressively wider; pereonites 5–7 subequal in length, posterior margin
moderately arched. Coxae 2–3 posteroventral margins rounded; 4–7 with acute carinae. Pleonites 1–4 subequal in
length and width, visible in dorsal view; pleonites posterior margin smooth; pleonite 5 posterior margin straight.
Pleotelson 0.4 times as long as anterior width, anterior margin linear, lateral margins weakly concave, posterior
subtruncate, without median point.
FIGURE 29. Cymothoa vicina ovigerous female holotype (15 mm; AM P8590). A, dorsal view; B, front view of female
pereonite 1 and cephalon; C, antennula; D, antenna; E, uropod; F, dorsal view of pleotelson; G, lateral view.
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FIGURE 30. Cymothoa vicina ovigerous female holotype (15 mm; AM P8590). A, maxilliped; B, mandible; C–F, pereopods
1, 2, 6, 7 respectively; G–H, pleopods 1 and 2 respectively.
Antennula comprised of 8 articles; peduncle articles 1 and 2 distinct; article 2 0.9 times as long as article 1;
article 3 0.7 times as long as combined lengths of articles 1 and 2, 1.1 times as long as wide; extending to middle of
eye. Antenna comprised of 9 articles, peduncle article 3 0.9 times as long as article 2, 1.1 times as long as wide;
article 4 1.0 times as long as wide; article 5 1.0 times as long as article 4, terminal article without setae. Labrum
lateral margins convex, anterior margin rounded, without small median point. Mandibular process present, ending
with an acute incisor; articles of mandibular palp distinct. Maxilliped comprised of 3 articles, well segmented,
article 3 with 1 apical spine and 4 setae.
Pereopod 1 basis 1.7 times as long as greatest width, superior proximal margin smooth, without raised carina;
ischium 1.2 times as long as basis; merus proximal margin without bulbous protrusion; carpus with straight
proximal margin; propodus 1.4 times as long as wide; dactylus slender, 1.7 times as long as propodus, 2.4 times as
long as basal width. Pereopod 2 basis 1.4 times as long as greatest width, superior proximal margin smooth,
without raised carina; propodus 1.6 times as long as wide; dactylus 1.3 times as long as propodus. Pereopods 3–5
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similar to pereopod 2, gradually increasing in size, without robust or simple setae. Pereopod 6 basis 1.3 times as
long as greatest width, superior proximal margin smooth, without raised carina; ischium 1.0 times as long as basis,
propodus 1.2 times as long as wide, dactylus 1.7 times as long as propodus. Pereopod 7 basis 1.4 times as long as
greatest width, superior proximal margin smooth, with moderately raised carina; ischium 0.6 times as long as basis,
with slight bulbous protrusion; merus proximal margin without slight bulbous protrusion, merus 0.5 times as long
as ischium; carpus 0.3 times as long as ischium, without bulbous protrusion, 0.5 times as long as wide; propodus
0.7 times as long as ischium, 1.7 times as long as wide; dactylus stout, 1.5 times as long as propodus, 2.8 times as
long as basal width.
FIGURE 31. Cymothoa vicina ovigerous female (23 mm; MTQ W7302). A, dorsal view; B, ventral view of mouthpart; C,
antennula; D, antenna; E, view pereonite 1 and cephalon; F, uropod; G, lateral view.
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FIGURE 32. Cymothoa vicina ovigerous female (23 mm; MTQ W7302). A–D, pereopods 1, 2, 6, 7 respectively; E–H,
pleopods 1, 2, 4, 5 respectively; I, dorsal view of pleotelson.
Pleopods without depression on central dorsal surface of each pleopod rami; basal projections present; exopod
proximal mesial margins extending past peduncle; exopod larger than endopod. Pleopod 1 exopod 1.1 times as
long as wide, lateral margin weakly concave, distally narrowly rounded, medial margin weakly oblique, mesial
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margin strongly convex; peduncle 0.3 times as wide as long, without retinaculae; pleopod 2 appendix masculina
with straight margins, 0.6 times as long as endopod, distally broadly rounded.
Uropod not extending beyond posterior margin of pleotelson; peduncle 1.3 times as long as greatest width, 0.8
times as long as exopod, lateral margin without setae, marginal setae absent, apices broadly acute, lateral margin
straight, mesial margin strongly convex. Exopod extending past endopod, 3.3 times as long as greatest width,
apically rounded, lateral margin convex, terminating without setae, mesial margin concave. Endopod 3.2 times as
long as greatest width, apically rounded, lateral margin weakly straight, terminating without setae, mesial margin
straight.
FIGURE 33. Cymothoa vicina male (9 mm; MTQ W7302). A, dorsal view; B, ventral view of mouthpart; C, antennula; D,
antenna; E, uropod; F, lateral view.
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FIGURE 34. Cymothoa vicina male (9 mm; MTQ W7302). A–C, pereopods 1, 2, 7 respectively; D–F, pleopods 2, 3, 5
respectively.
Males. Body 2.2 times as long as greatest width. Cephalon subtriangular, 0.4 times longer than wide. Pereonite
1 anterolateral minute, reaching posterior margin of eyes; pereonites 1–6 posterior margins straight and smooth.
Coxae 2–7 dorsally visible, posteroventral margins rounded. Pleonites subequal in width, similar to female.
Pleotelson semi-rounded. Antennula similar size to antenna. Antenna comprised of 8 articles.
Pereopod 1 basis 1.7 times as long as greatest width; ischium 0.5 times as long as basis; merus proximal
margin without bulbous protrusion; carpus with straight proximal margin; propodus 2.0 times as long as wide;
dactylus 1.2 times as long as propodus. Pereopods 2–6 similar to pereopod 1, moderately bigger in size. Pereopod
7 basis superior proximal margin without raised carina, inferior margin of ischium, merus and carpus straight.
Pleopods margins smooth, exopods greater than endopods from 1–5. Pleopod 2 appendix masculina with parallel
margins, 0.7 times as long as endopod, distally broad. Pleopods 3 and 5 endopods proximal margins extending
below peduncle. Uropod exopod similar length to endopod, apices broadly rounded.
Colour. Holotype dark brown, non-type material whitish to yellowish tan.
Size. Ovigerous females: 15–25 mm; males: 4–10 mm.
Remarks. The female holotype has shrivelled since Hale’s (1926) illustration of the specimen. Cymothoa
vicina has a suboval body [appearing wider in Hale’s (1926) illustration]; eyes distinct (0.3 times length of
cephalon for combined eyes width); subtruncate cephalon [rostrum appearing acute in Hale’s (1926) illustration];
anterolateral margins of pereonite 1 almost reaching anterior margin of the cephalon; subequal in length and width
of pereonites 1–4; pleonites of subequal width; smooth and raised carina on pereopod 7 basis; and 2.5 times wider
than long pleotelson. Hale (1926) also noted that the appendix masculina was retained on pleopod 2 of the female.
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FIGURE 35. Cymothoa vicina, female (22 mm; SMF-572) A, dorsal view; B, ventral view; C, lateral view. (picture credit Sven
Tränkner, Senckenberg Research Institute).
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FIGURE 36. Cymothoa vicina, male (11 mm; SMF-572) A, dorsal view; B, ventral view; C, lateral view. (picture credit Sven
Tränkner, Senckenberg Research Institute).
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FIGURE 37. Cymothoa vicina, female (23 mm; SMF-572) A, dorsal view; B, ventral view; C, lateral view. (picture credit Sven
Tränkner, Senckenberg Research Institute).
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FIGURE 38. Cymothoa vicina, female (19 mm; SMF-572) A, dorsal view; B, ventral view; C, lateral view. (picture credit Sven
Tränkner, Senckenberg Research Institute).
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Two ovigerous females from Moreton Bay (MTQ W7302) occurred on the host Mugil sp., a host-association
similar to that of the holotype. Both the Moreton Bay specimens and the holotype have a subtriangular cephalon;
subtruncate pleotelson; uropods not extending beyond pleotelson; pleonites subequal in width; smooth and raised
carina on pereopod 7 basis. The Moreton Bay specimens differ from the holotype in having a more ovate rather
than suboval body (1.7 times longer than wide); eyes partially visible; pereonites 1–4 anterolateral margins broad
and more pronounced; inferior distal margin of pereopod 7 ischium more produced (compared to holotype); and
pleopods 1 and 2 margins irregular (compared to the simpler endopod and pleopod margins in the female holotype)
The males differ from the females (both males and females from Moreton Bay) by having a subparallel body,
rounded pleotelson posterior margin; and the minute anterolateral margins of pereonite 1.
Trilles (2008) recorded C. epimerica from the Seychelles (SMF-76) and the Red Sea (SMF-567 and SMF-572)
without host association or illustrations. Digital images obtained from the Senckenberg Research Institute shows
all the female Red Sea specimens (19, 22 and 23 mm) from SMF-572 to be Cymothoa vicina Hale, 1926. These
specimens are the same as the specimens from Moreton Bay, particularly the female’s ovate body; eyes partially
visible; pereonites 1–4 anterolateral margins broad and more pronounced and the inferior distal margin of pereopod
7 ischium more produced.
Cymothoa vicina resembles C. oestrum in the pereonite 1 anterolateral margins broad and reaching past eyes,
cephalon anterior margin subtruncate, pleotelson posterior margin subtruncate, and pereopod 7 basis with raised
carina. The differences noted in C. oestrum are the dorsally visible coxae and the rhomboidal body morphology
(compared to the oval-shaped body in female specimens from MTQ W7302). Thatcher’s et al. (2003) illustration
of C. oestrum showed antennula articles 1–3 has small spinules, which are not present in C. vicina.
Distribution. Reported from western and eastern Australia and the Red Sea (Trilles 2008, specimens SMF-
572).
Hosts. Known from hosts of families Plotosidae and Mugilidae.
Species excluded from the Australian fauna
Cymothoa curta Schioedte & Meinert, 1884
Figures 39–43
Cymothoa curta Schioedte & Meinert, 1884: 228, tab. VI (Cym. XXIV) figs. 3–6.—Trilles, 1994: 139.—Trilles & Bariche,
2006: 228.
Excluded
Cymothoa curta.—Avdeev, 1978b: 282; 1982a: 65.—Bruce, Lew Ton & Poore, 2002: 175 [=Cymothoa sp.].
Material examined. Lectotype (here designated): 1 ovig. ♀ (14 mm), locality unknown, from largescale foureyes
Anableps anableps (Linnaeus, 1758) (previously Anableps tetrophthalmi), (NHMW 6142).
Paralectotype (here designated): 1 immature ♂ (7 mm), same data as lectotype (NHMW 25653).
Brazilian material: 3 ovig. ♀ [14 mm (MTQ W34285), 15 mm (MTQ W34286), 19 mm (MTQ W34288)], 1
immature ♂ [7 mm (MTQ W34288)], lagoon, Ajuruteua, Bragança, Pará, 0.806782°S, 46.632843°W, 8 December
2012, from four-eyed fish Anableps microlepis Müller & Troschel, 1844, coll. Han Xiao and Jeferson Carneiro.
1 ovig. ♀ (17 mm), 1 immature ♂ (9 mm), Raposa Beach, São Luiz, Maranhão, 2.454693°S, 44.080582°W, 27
October 2012, from four-eyed fish Anableps microlepis, coll. Han Xiao and Jeferson Carneiro (MTQ W34284).
1 ovig. ♀ (17 mm), 1 immature ♂ (7 mm), Salinópolis, Pará, 0.6059535°S, 47.366180°W, 20 October 2012,
from four-eyed fish Anableps microlepis coll. Han Xiao and Jeferson Carneiro (MTQ W34283).
Ovigerous female Length 14 mm width 7 mm (lectotype).
Body ovoid, 1.8 times as long as greatest width, dorsal surface smooth, widest at pereonite 5, most narrow at
pereonite 1, lateral margins moderately convex. Cephalon 0.2 times longer than wide, visible from dorsal view,
semi-circular. Frontal margin rounded to form blunt rostrum. Eyes partially visible. Pereonite 1 anterior margin
linear, anterolateral margins minute; pereonites 1–5 posterior margins smooth and linear, progressively wider;
pereonites 2–6 subequal in length. Coxae 2–4 posteroventral margins rounded, 4–7 narrowly produced points.
Pleonites 1–4 subequal in length and width, visible in dorsal view; pleonites 2–4 posterior margin bisinuate;
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pleonite 5 trisinuate. Pleotelson 0.4 times as long as anterior width, lateral margin weakly convex, posterior margin
rounded.
Antennula comprised of 8 articles; peduncle articles 1 and 2 distinct; article 2 0.9 times as long as article 1;
article 3 0.3 times as long as combined lengths of articles 1 and 2, 1.0 times as long as wide; extending to middle of
eye. Antenna comprised of 9 articles, peduncle article 3 0.8 times as long as article 2, 1.0 times as long as wide;
article 4 1.1 times as long as wide; article 5 1.0 times as long as article 4, terminal article without setae, extending
to posterior of pereonite 1. Labrum lateral margins convex, anterior margin rounded.
FIGURE 39. Cymothoa curta, ovigerous female holotype (14 mm; NHMW 6142). A, dorsal view; B, ventral view of
mouthpart; C–F, pereopods 1, 2, 6, 7 respectively; G, lateral view.
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FIGURE 40. Cymothoa curta, ovigerous male paratype (7 mm; NHMW 6142). A, dorsal view; B, ventral view of mouthpart
and pereopod 1 (dactylus and propodus); C–F, pereopods 1, 2, 6,7 respectively; G, uropod; H, lateral view.
Pereopod 1 basis 1.7 times as long as greatest width, superior proximal margin smooth, without raised carina;
ischium 0.7 times as long as basis; merus proximal margin without bulbous protrusion; carpus with straight
proximal margin; propodus 1.1 times as long as wide; dactylus slender, 1.7 times as long as propodus, 2.4 times as
long as basal width. Pereopod 2 basis 1.7 times as long as greatest width, superior proximal margin smooth,
without raised carina; propodus 1.2 times as long as wide; dactylus 1.5 times as long as propodus. Pereopods 3–5
similar to pereopod 2, gradually increasing in size, without robust or simple setae. Pereopod 6 basis 1.0 times as
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long as greatest width, superior proximal margin smooth, with moderately raised carina; ischium 1.3 times as long
as basis, propodus 1.0 times as long as wide, dactylus 1.8 times as long as propodus. Pereopod 7 basis 1.1 times as
long as greatest width, superior proximal margin smooth, with raised carina; ischium 0.7 times as long as basis,
with bulbous protrusion; merus proximal margin without bulbous protrusion, merus 0.3 times as long as ischium,
0.4 times as long as wide; carpus 0.2 times as long as ischium, without bulbous protrusion, 0.4 times as long as
wide; propodus 0.5 times as long as ischium, 1.2 times as long as wide; dactylus slender, 1.7 times as long as
propodus, 2.4 times as long as basal width.
FIGURE 41. Cymothoa curta, ovigerous female Brazilian (17 mm; MTQ W34283). A, dorsal view; B, front view of cephalon;
C, antennula; D, antenna; E, uropod; F, dorsal view of pleotelson; G, lateral view.
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FIGURE 42. Cymothoa curta, ovigerous female Brazilian (17 mm; MTQ W34283). A–D, pereopods 1, 2, 6, 7 respectively; E–
H, pleopods (dorsal view) 1–4 respectively; I–L, pleopods (ventral view) 2–5 respectively.
Uropod not reaching posterior margin of pleotelson, peduncle 0.8 times as long as exopod, peduncle lateral
margin without setae; marginal setae absent, apices broadly rounded, 1.6 times as long as greatest width, lateral
margin straight, mesial margin strongly convex. Exopod extending past endopod, 7.0 times as long as greatest
width, mesial margin weakly convex.
Male. Length 7 mm, width 3 mm (paralectotype).
Body subtriangular, 2.1 times as long as greatest width, dorsal surface smooth, widest at pereonite 6. Cephalon
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0.6 times longer than wide, visible from dorsal view, subtruncate, not deeply immersed in pereonite 1. Eyes
partially visible. Pereonite 1 anterolateral minute, posterior margins of pereonites 6 and 7 not deeply arched. Coxae
2–4 anteroventral and posteroventral margins rounded. Pleon not overlapped by pereonite 7; pleonites subequal in
width, posterior margin not trisinuate. Pleotelson subtruncate. Antennula comprised of 8 articles.
Pereopod 1 basis 1.5 times as long as greatest width; ischium 0.5 times as long as basis; merus proximal
margin without bulbous protrusion; carpus with straight proximal margin; propodus 1.3 times as long as wide;
dactylus 2.3 times as long as propodus. Pereopods 2–7 similar to pereopod 1, moderately bigger in size, pereopods
6 and 7 basis superior proximal margin without raised carinae. Uropod exopod and endopod apices broadly
rounded.
FIGURE 43. Cymothoa curta, male Brazilian (7 mm; MTQ W34283). A, dorsal view; B, antennula; C, antenna; D–G,
pereopods 1, 2, 6, 7 respectively; H, lateral view; I, uropod.
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Colour. Female and male type appearing yellow–tan. Brazilian female and male specimens with blackish grey
colouration.
Size. Ovigerous females: 14–17 mm; males: 7–9 mm.
Remarks. Cymothoa curta has a stout body (1.8 times longer than wide); cephalon rounded; pereonite 1
anterolateral margin minute, and not extending anteriorly beyond half of cephalon length; pereonites 1–4 subequal
in length; pleonites 2–4 posterior margin bisinuate; pleonite 5 trisinuate; pereopod 7 basis with raised carina and
uropods which do not extend past the pleotelson posterior margin. The male paralectotype differs from the
lectotype in having a subtriangular body morphology, cephalon dorsally more visible, pleon not overlapped by
pereonite 7, anterolateral and posterolateral margins of coxae 2–7 smooth and linear, and pereopod 7 basis without
a sharp carina.
The Brazilian female specimens are similar to the lectotype in the rounded cephalon, pereonites 1–4 subequal
in length; pleonites 2–4 posterior margin bisinuate; and uropods which do not extend past the pleotelson posterior
margin. The Brazilian specimens differ from the lectotype in the moderately subtruncate pleotelson posterior
margin, and the pleon partially overlapped by pereonite 7. Pleopods of the Brazilian female specimens have the
exopods 1–5 similar size to endopod, depression on the central dorsal surface of pleopods 2–4; pleopod 5 endopod
has large fleshy folds present; endopods 2–5 lateral margin strongly convex, distally broadly round, medial margin
oblique, mesial margin strongly convex; and peduncle lobes from pleopods 1–5 present and becoming
progressively larger. The Brazilian males differ from the paralectotype male in the more subparallel body
morphology.
Schioedte & Meinert (1884) identified the type host, Anableps anableps, without mention of the type locality.
Anableps anableps is a South American fish that does not occur in the Indo-Pacific (Froese & Pauly 2015). The
present material of C. curta fromBrazilian Anableps microlepis verifies that the host species does not occur in
Australia. Avdeev (1978b) apparently found C. curta in the buccal cavity of sailfin velifer Velifer hypselopterus
Bleeker, 1879 (family Veliferidae) from the Arafura Sea. Velifer hypselopterus is known to occur in the Indo-
Pacific region (Froese & Pauly 2015). It is highly unlikely, given the host and geographic difference, that Schioedte
& Meinert’s (1884) C. curta from Brazil and Avdeev’s (1978b) record are of the same species, and as Avdeev
(1978b) did not indicate where the material was deposited, we exclude the species from the Australian fauna.
Cymothoa epimerica is similar toC. curta from the ovoid body shape (1.8 and 1.9 times longer than wide
respectively), pleonites posterior margins trisinuate and uropods which do not extend past the pleotelson posterior
margin. Cymothoa epimerica differs from C. curta by the more developed anterolateral margins of pereonite 1 that
reaches the anterior margin of cephalon; posterolateral margin of pereonite 1 with minute projection; dorsally
visible coxae, with acute posterolateral margins; acute carinae of pereopods 6 and 7 basis (compared to the highly
raised carina in C. curta), long and narrow uropods (compared to the more stout uropodal rami), and the irregular
margins of all pleopods.
Distribution. South America: Brazil (present material).
Hosts. Anablepidae: Anableps anableps (see Schioedte & Meinert 1884) and Anableps microlepis (present
study).
Cymothoa plebeia Schioedte & Meinert, 1884
Figures 44–45
Cymothoa plebeia Schioedte & Meinert, 1884: 236, tab. IX (Cym. XXVII) figs. 1, 2; Van Name, 1920: 43, 59, figs. 12, 13.—
Monod, 1927: 694: 1931: 4.—Brian & Dartevelle, 1949: 84, 127, 169, 172, 176, 179, 186, figs. 92–106.—Trilles, 1975:
978, pl. I, figs. 1, 2; 1979a: 522; 1986: 627, tab 1; 1994: 147.—Rokicki, 1984a: 71, fig. 5.—Bruce, Lew Ton & Poore,
2002: 175.—Trilles & Bariche, 2006: 228.
Excluded
Cymothoa plebeia.—Avdeev, 1978b: 282 [=Cymothoa epimerica].
Cymothoa plebeia.—Rokicki, 1985: 95, tabs. 1–3, fig. 8; 1986: 254 [=Cymothoa epimerica].
Not Cymothoa plebeia.—Rokicki, 1984b: 60, figs. 13–16 [=Nerocila armata].
Material examined. Holotype: 1 ovig. ♀ (21 mm), Cape Verde (type locality as “ad Promontorium viride” by
Schioedte & Meinert) unknown host (ZMO F15977).
Ovigerous female Length 21 mm, width 9 mm (holotype).
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FIGURE 44. Cymothoa plebeia, ovigerous female holotype (21 mm; F15977). A, dorsal view; B, ventral view of mouthpart
and pereopod 1 (dactylus and propodus); C, antennula; D, antenna; E, lateral view.
Body subparallel, 2.2 times as long as greatest width, dorsal surface smooth, widest at pereonite 5, most narrow
at pereonite 1. Cephalon 0.2 times longer than wide, visible from dorsal view, anterior margin semi-circular.
Frontal margin rounded to form blunt rostrum, ventrally folded. Eyes absent. Pereonite 1 anterior margin
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moderately raised medially, anterolateral margins minute; pereonites 1–4 posterior margin smooth, progressively
wider; pereonites 5–7 subequal in length, posterior margin moderately arched. Coxae 2–7 posteroventral margins
rounded. Pleon not overlapped by pereonite 7, visible in dorsal view; pleonites posterior margin smooth; pleonites
1–4 subequal in length and width; pleonite 5 posterior margin straight. Pleotelson 0.5 times as long as anterior
width, anterior margin moderately bisinuate, lateral margin weakly convex, posterior margin emarginate, without
median point.
FIGURE 45. Cymothoa plebeia, ovigerous female holotype (21 mm; F15977). A–D, pereopods 1, 2, 6, 7 respectively.
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Antennula comprised of 9 articles; peduncle articles 1 and 2 distinct, not articulated; article 2 1.0 times as long
as article 1; article 3 0.6 times as long as combined lengths of articles 1 and 2, 1.4 times as long as wide; extending
to middle of cephalon. Antenna comprised of 9 articles, peduncle article 3 1.4 times as long as article 2, 1.6 times
as long as wide; article 4 0.9 times as long as wide; article 5 0.9 times as long as article 4, terminal article without
setae, extending to middle of cephalon. Labrum fleshy, lateral margins convex and produced, anterior margin
rounded.
Pereopod 1 basis 1.5 times as long as greatest width, superior proximal margin smooth, without raised carina;
ischium 0.7 times as long as basis; merus proximal margin without bulbous protrusion; carpus with straight
proximal margin; propodus 1.8 times as long as wide; dactylus slender, 1.9 times as long as propodus, 3.2 times as
long as basal width. Pereopod 2 basis 1.5 times as long as greatest width, superior proximal margin smooth,
without raised carina; propodus 1.6 times as long as wide; dactylus 1.4 times as long as propodus. Pereopods 3–5
similar to pereopod 2, gradually increasing in size, without robust or simple setae. Pereopod 6 basis 1.4 times as
long as greatest width, superior proximal margin smooth, with moderately raised carina; ischium 0.5 times as long
as basis, propodus 1.0 times as long as wide, dactylus 1.9 times as long as propodus. Pereopod 7 basis 1.2 times as
long as greatest width, superior proximal margin irregular, with raised carina; ischium 0.6 times as long as basis,
without bulbous protrusion; merus proximal margin without bulbous protrusion, merus 0.2 times as long as
ischium, 0.5 times as long as wide; carpus 0.3 times as long as ischium, without bulbous protrusion, 0.5 times as
long as wide; propodus 0.8 times as long as ischium, 1.2 times as long as wide; dactylus stout, 1.6 times as long as
propodus, 2.9 times as long as basal width.
Uropod not extending past posterior margin of pleotelson, peduncle 0.9 times as long as exopod rami, peduncle
lateral margin without setae; marginal setae absent, lateral margin concave.
Colour. Light brown.
Size. Females: 13.5–25 mm; males: 6–19 mm; second pulli: 2.6–12.5 mm (Van Name 1920; Brian &
Dartevelle 1949; Trilles, 1975, 1979a; Rokicki 1984a).
Remarks. Cymothoa plebeia can be distinguished by the fleshy labrum; rounded rostrum; small and acute
anterolateral margins of pereonite 1, anterior margin of pereonite 1 medially raised; pleon not immersed by
pereonite 7; pereopods 6 and 7 superior proximal margin of dactyli slightly raised; pereopod 7 basis with raised
carina; uropods not extending past posterior margin of pleotelson and the emarginate pleotelson that is 1.7 times
wider than long.
Van Name (1920) identified and described a single male specimen (19 mm, AMNH 3236) that allowed
comparison to Schioedte & Meinert’s (1884) single female specimen. Both male and female have similar body
morphology, the noted differences are that the male cephalon is more visible in dorsal view, eyes partially visible
(absent in female), and cephalon appearing more subtruncate.
Cymothoa indica is most similar to C. plebeia in pereonite 1, pleon, pleotelson and pereon morphology. The
differences noted include C. indica having a subtriangular cephalon (compared to the more rounded anterior
margins and straight lateral margins of C. plebeia cephalon); pereonite 1 anterolateral margin projecting forward
(anterolateral margin curved inward to cephalon in C. plebeia); pereopod 7 ischium with a prominent lobe on the
inferior distal region (not visible in C. plebeia); and pleotelson posterior margin rounded (emarginate in C.
plebeia).
Avdeev’s (1978b) and Rokicki’s (1985, 1986) records of C. plebeia from the sparid Pagrosomus sp. are
questionable. The two possible host species from the genus are Pagrus major (Temminck & Schlegel, 1843) with a
northwestern Pacific distribution, and Pagrus auratus (Foster, 1801) known from the Western-Pacific. Neither host
species conform to the known distribution of C. plebeia from the eastern Atlantic Ocean and therefore both
Avdeev’s and Rokicki’s records of C. plebeia are here excluded from synonymy.
Distribution. Known from the eastern Atlantic Ocean and northwestern African coast: Cape Verde (Schioedte
& Meinert 1884; Van Name 1920); Congo (Van Name 1920; Brian & Dartevelle 1949); Cameroon (Monod 1931;
Trilles 1975); Angola (Brian & Dartevelle 1949); Benin and Ghana (Trilles 1975); and Senegal (Trilles 1979a;
Rokicki 1984a).
Schioedte & Meinert (1884) designated the type locality as “ad Promontorium viride” (Latin for Cape Verde).
Other authors (Brian & Dartevelle 1949; Trilles 1975, 1994; Rokicki 1984b) have refered to the location as “from
Cap Vert” or “Senegal”.
Avdeev (1978b) recorded C. plebeia from the western coast of Australia, but did not indicate where the
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specimens were deposited. Like several other of Avdeev’s species, the specimens could not be located at TINRO.
As there are no specimens of C. plebeia in Australian collection, and given with the restricted distribution of C.
plebeia in the eastern Atlantic Ocean, we here exclude C. plebeia from the Australian fauna.
Hosts. Reported from families of Haemulidae and Sciaenidae; bigeye grunt Brachydeuterus auritus
(Valenciennes, 1832) (see Monod 1931; Trilles 1975, 1979a; Rokicki 1984a).
Acknowledgements
We would like to thank curators and researchers from their respective museums and universities for access and/or
information on of cymothoid material: Barbara Done and Dr Robyn Cumming (Museum of Tropical Queensland,
Townsville); Dr Ben Diggles (DigsFish Services Pty Ltd, Queensland); Dr Stephen Keable (Australian Museum,
Sydney); the late Dr Ian Whittington (South Australian Museum, Adelaide); Han Xiao (Leibniz Centre for Marine
Tropical Ecology, Germany); Dr Marina Malyutina (Institute of Marine Biology, Vladivostok, Russia) for hand-
carrying Avdeev’s particularly important type material; Helen Wong Pei San (National University of Singapore,
Tropical Marine Science Institute); Dr Richard Preece (University Museum of Zoology, Cambridge); Dr Oliver
Coleman (Zoological Museum, Museum für Naturkunde, Berlin); and Dr Jørgen Olesen (Zoological Museum of
Copenhagen, Denmark). Special thanks for images go out to Dr Peter Dworschak (Natural History Museum,
Vienna); Åse Ingvild Wilhelmsen and Karsten Sund (Natural History Museum, University of Oslo) for type
specimens; and Prof. Dr. Micheal Tuerkay, Dr Moritz Sonnewald and Sven Tränkner (Senckenberg Research
Institute, Department of Marine Zoology, Crustacean Section) for non-type specimens. Cymothoa pulchrum
specimens were collected during the Comprehensive Marine Biodiversity Survey of Singapore, a project organized
and primarily supported by the National Parks Board of Singapore (NParks), together with the Tropical Marine
Science Institute (TMSI) and the Zoological Reference Collection (ZRC) of the Lee Kong Chian Natural History
Museum (LKCNHM), National University of Singapore. The survey is co-sponsored by Shell Singapore, Asian
Pacific Breweries and the Hong Kong & Shanghai Bank (Singapore). We thank Dr Kerry Hadfield from the North-
West University South Africa, who has generously shared her preliminary cladistics results; description of the type
material and helped resolved a number of the complex synonymies for Cymothoa. Gratitude is also extended to The
Museum of Tropical Queensland–Queensland Museum for provision of facilities during MBM’s internship. MBM
acknowledges the travel cost and proof reading services of University of Tasmania; the Australian museum for the
Geddes Postgraduate Award and the Public Service Department of the Malaysian Government for a postgraduate
scholarship.
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... Historic cymothoid records specific to the Malaysian region are those of Lanchester (1902) collected during his Skeat Expedition to Peninsular Malaysia and Anand Kumar et al. (2015Kumar et al. ( , 2017 for cymothoids off the coast of Miri in Borneo. Unfortunately, the type specimen for Cymothoa pulchrum Lanchester, 1902 could not be located (see Martin et al. 2016) and material examined by Anand Kumar et al. (2015Kumar et al. ( , 2017 was not deposited in any museum or repository. Miers (1880) reported several species "from the Malaysian" region, but Miers' use of the term 'Malaysian' included present day Indonesia; Nerocila laevinota Miers, 1880 was the only species explicitly recorded from Malaysia but is considered a junior synonym of Nerocila sundaica Bleeker, 1857. ...
... Cymothoa eremita is one of the most common and widely reported species of Cymothoa, and interestingly one of the few species to display low host specificity (11 known host families) and wide geographical distribution from the western Indian Ocean (e.g., Mozambique) to the Central Indo-Pacific (e.g., Australia) (see Hadfield et al. 2013;Martin et al. 2016). The first report of this species for Malaysian waters was by Anand et al. (2015Anand et al. ( , 2017 in Sarawak, where the species was collected from the buccal cavity of Psettodes erumei. ...
... Cymothoa pulchrum has an anteriorly subtruncate cephalon; wide, subtruncate pereonite 1; anterolateral margins nearly reaching rostrum; pleon partially overlapped by pereonite 7; rounded pleotelson posterior margin; raised carinae on pereopods 6 and 7 bases; prominent lobe on ischium pereopod 7; coxae visible from dorsal view; uropods reaching half pleotelson length, visible from ventral view. For further comparisons on gender and comparatively similar species, refer to Martin et al. (2016). Lanchester (1902) reported a female specimen of the species from the northern Straits of Malacca, Malay Peninsula as part of the collection from the Skeat Expedition to the Malay Peninsula. ...
Article
A checklist of parasitic cymothoids from Malaysian waters is presented based on available literature and material collected from 2010 to 2020. Most of the collected specimens were recorded from waters of Terengganu, east coast of Peninsular Malaysia (facing the South China Sea), whereas literature records were included from Sarawak, along the Miri coast of northwest Borneo. The checklist comprises 19 species in ten genera, seven of which are new records from Malaysia: Anilocra nemipteri Bruce, 1987; Ceratothoa barracuda Martin, Bruce & Nowak, 2015; Ceratothoa carinata (Bianconi, 1869); Cymothoa epimerica Avdeev, 1979; Elthusa sigani Bruce, 1990; Joryma engraulidis (Barnard, 1936) and Renocila richardsonae Williams & Bunkley-Williams, 1992. Eight new host records are based on collected specimens: Anilocra nemipteri was dorsally attached on Nemipterus nemurus (Bleeker, 1857), Nemipterus nematophorus (Bleeker, 1854), Nemipterus tambuloides (Bleeker, 1853) and Nemipterus thosaporni Russell, 1991 (family Nemipteridae); Ceratothoa carinata was found in the buccal cavity of Decapterus macrosoma Bleeker, 1851 (family Carangidae); Cymothoa eremita (Brunnich, 1783) was attached in the buccal cavity of Nemipterus tambuloides (Bleeker, 1853) and Nemipterus furcosus (Valenciennes, 1830); Elthusa sigani was found attached on the gills of Pterois russelli Bennett, 1831 (family Scorpaenidae), and Renocila richardsonae was attached on the caudal fin of Upeneus japonicus (Houttuyn, 1782) (family Mullidae). All cymothoid species listed here are known to have a Central Indo-Pacific distribution, with some ranging as far as the Western Indian Ocean. A cymothoid–host association is here listed from 28 fish families, with the most commonly reported from Carangidae (pompanos, jack mackerels, runners, scads), Engraulidae (anchovies), and Leiognathidae (ponyfishes, slipmouths). This paper is the first comprehensive review of both verified literature records and deposited specimens. A key for the family of Cymothoidae in Malaysian waters is given.
... The genus diagnosis has been well revised by Hadfield et al. (2011Hadfield et al. ( , 2013. The species of Cymothoa can be identified by the strongly vaulted body; slender antennae with widely separated bases, cephalon deeply immersed in pereonite 1, antenna longer than antennula; subtruncate rostrum; bases of pereopods 5-7 with broad blade-like carina; coxae partially visible dorsally; pereonite 7 posterolateral margins extending past pleonite 1; and pleopods 3-5 with large fleshy folds (Martin et al. 2016). The genus Cymothoa can be well separated from other similar buccal-attaching cymothoid genera. ...
... Avdeev (1979) minimally described Cymothoa bychowskyi based on the materials collected from the red cornetfish Fistularia petimba off north-western Australia, and the holotype and paratypes are deposited in the museum of the Russian Pacific Federal Fisheries Research Institute, Vladivostok. A recent museum enquiry and a loan request for the types by Martin et al. (2016) revealed that the specimens could not be located. Following the original description the species has not been well studied, and a redescription is required based on the fresh specimens. ...
... . 1.-Trilles 1994: 138. - Williams et al. 2000: 157. -Kensley 2001: 232. -Bruce et al. 2002: 174. -Paulay et al. 2003: 479. -Trilles and Bariche 2006: 228. -Rameshkumar et al. 2013 fig. 1(C).-Martin et al. 2016: 6. -Ravichandran et al. 2019 ...
Article
The present study reports two poorly known buccal-attaching fish-parasitic cymothoids, Ceratothoa carinata (Bianconi, 1869) and Cymothoa bychowskyi Avdeev, 1979, from Indian waters. Ceratothoa carinata (Bianconi, 1869) infesting the shortfin scad, Decapterus macrosoma (Bleeker, 1851) collected from the Malabar Coast, India, is redescribed and illustrated based on the ovigerous female. It appears to be the first record of this species from Indian waters. Further, the poorly known Cymothoa bychowskyi Avdeev, 1979 infesting the red cornetfish Fistularia petimba Lacepède, 1803 collected from the south-west coast of India is redescribed and illustrated from female and male stages. An updated checklist of buccal-attaching fish-parasitic cymothoids is also provided.
... Minimal cymothoid records specific to the Malaysian region are attributed to records from Lanchester (1902) during his Skeat Expedition to Peninsular Malaysia and Anand for cymothoids off the coast of Miri in Borneo. Unfortunately, the type specimen for Cymothoa pulchrum Lanchester, 1902 could not be located (see Martin et al. 2016) and material examined by Anand were not deposited in any established museums or repository, and one could only rely on their published photographs and have been noted to have misidentification in their record. Bruce and Wong (2015) have strongly indicated that there have been few historic accounts for isopods from South East Asia. ...
... pereonite 1 anterolateral margins deeply curved towards cephalon; pereopods 5-7 superior proximal margin with acute carinae and dorsallyvisible; and coxae 6 and 7 posteroventral margins acute and dorsally visible (Martin et al. 2016). Our female specimens are very much similar to the illustrations and photographs provided by Martin et al. (2016), with the distinctively visible pereonite 1 margins and acute coxae 6 and 7 posteroventral margins, which is not as prominent in Cymothoa eremita or Cymothoa pulchrum. ...
... pereonite 1 anterolateral margins deeply curved towards cephalon; pereopods 5-7 superior proximal margin with acute carinae and dorsallyvisible; and coxae 6 and 7 posteroventral margins acute and dorsally visible (Martin et al. 2016). Our female specimens are very much similar to the illustrations and photographs provided by Martin et al. (2016), with the distinctively visible pereonite 1 margins and acute coxae 6 and 7 posteroventral margins, which is not as prominent in Cymothoa eremita or Cymothoa pulchrum. ...
Preprint
A checklist of parasitic cymothoids from Malaysian waters is presented based on available literature and material collected from 2010 to 2020. Most of the collected specimens were recorded from waters of Terengganu, east coast of Peninsular Malaysia (facing South China Sea), whereas literature records were represented from Sarawak, along the Miri coast of northwest Borneo. The checklist comprises 18 species under 10 genera, seven of which are new records from Malaysia, which includes Anilocra nemipteri Bruce, 1987; Ceratothoa barracuda Martin, Bruce and Nowak, 2015; Ceratothoa carinata (Bianconi, 1869); Cymothoa epimerica Avdeev, 1979; Elthusa sigani Bruce, 1990; Joryma engraulidis (Barnard, 1936) and Renocila richardsonae Williams and Bunkley-Williams, 1992. Eight new host records are based on collected specimens: Anilocra nemipteri was dorsally attached on Nemipterus nemurus (Bleeker 1857), Nemipterus nematophorus (Bleeker 1854), Nemipterus tambuloides (Bleeker 1853), and Nemipterus thosaporni Russell 1991 (family Nemipteridae); Ceratothoa carinata was found in the buccal cavity of Decapterus macrosoma Bleeker 1851 (family Carangidae); Cymothoa eremita (Brunnich, 1783) was attached in the buccal cavity of Nemipterus tambuloides and Nemipterus furcosus (Valenciennes 1830); Elthusa sigani was found attached on Pterois russelli Bennett 1831 (family Scorpaenidae); and Renocila richardsonae was attached on the caudal fin of Upeneus japonicus (Houttuyn 1782) (family Mullidae). All cymothoid species listed here are known to have a Central Indo Pacific distribution, with some ranging as far as the western Indian Ocean. The cymothoid-host association is here listed from 28 fish families, with the most common reported from Carangidae (pompanos, jack mackerels, runners, scads), Engraulidae (anchovies) and Leiognathidae (ponyfishes, slipmouths). This paper is the first comprehensive treatment to update both verified literature data and deposited specimens, with a key for the family Cymothoidae in Malaysian waters.
... The genus Cymothoa Fabricius, 1793 includes 49 valid species across the globe and it is equal largest with the genus Anilocra Leach, 1818 under the family cymothoidae (Martin et al. 2016). Only 5 species of the genus Cymothoa are known from Indian water viz., Cymothoa bychowskyi Avdeev, 1979, C. eremita (Brünnich, 1783, C. frontalis Milne Edwards, 1840, C. indica Schiöedte &Meinert 1884, andC. ...
Article
The parasitic isopod Cymothoa indica Schiöedte and Meinert, 1884 was recorded from the buccal cavity of the host Glossogobius giuris (Hamilton, 1822) collected from Chilka Lake, Odisha in 1924, but its identification was doubtful as mentioned by the author in his book on “Fauna of Chilika Lake: Tanaidacea and Isopoda”. The present report of parasitic isopod C. indica from the buccal cavity of the host Sphyraena obtusata Cuvier, 1833 collected from the water of Bay of Bengal, Gopalpur-on-Sea confirms its occurrence along the Odisha coast. The record of the isopod parasite Norileca indica (H. Milne Edwards, 1840) from the branchial cavity of the host Atule mate (Cuvier, 1833) collected from the water of Bay of Bengal, Gopalpur-on-Sea is the first record of this parasite from the coastal water of Odisha, India. The record of Nerocila arres Bowman and Tareen, 1983 from the caudal peduncle of the host fish species Terapon puta Cuvier, 1829, and the isopod N. depressa Milne Edwards, 1840 from the host fish species Megalaspis cordyla (Linnaeus, 1758) are the new host records for these respective parasites.
... In recent decades cymothoid generic diagnoses have become increasingly precise and restrictive (e.g., Hadfield et al. 2013Hadfield et al. 2014. Recently, Martin et al. (2016) gave a world key to the marine buccal-attaching genera of the Cymothoidae. The description of the new species given here together with the descriptive data given by Yu and Bruce (2009) allows for a detailed generic description and diagnosis comparable to diagnoses for related genera. ...
Article
The genus Lobothorax Bleeker, 1857 is revised with the description of a new species collected from the gempylidaen fish Promethichthys prometheus (Cuvier, 1832) from the southwestern coast of India. A revised generic diagnosis is provided based on the redescription of the type species. Lobothorax aurita (Schioedte and Meinert, 1883) is here synonymised with Lobothorax typus Bleeker, 1857 based on the original description. Lobothorax nicosmiti Aneesh, Bruce and Kumar sp. nov. is described from the female stage and it is characterized by: pereonite 1 anterolateral expansion not extending to the anterior margin of cephalon; coarsely pitted pereonites dorsal surfaces; pereonites without dorsal median longitudinal ridges; anteriorly truncate cephalon; pleotelson about 0.6 times as long as wide, posterior margin weakly emarginate, broadly sub-truncate, lateral margins convex; maxilliped palp article 3 with three RS; pereopods basis much wider with prominent carina. A key to the species of Lobothorax Bleeker, 1857 is presented.
... Ambulatory pereopods excludes Obtusotelson from the Aegidae White, 1850 and Cymothoidae Leach, 1814 (e.g. see Bruce, 2009;Hadfield et al., 2014;Martin et al., 2016), while the Anuropidae Stebbing, 1893 (undifferentiated uropods), Protognathiidae Wägele and Brandt, 1988 (pereopod 7 absent) and Gnathiidae Leach, 1814 (only five functional pereopods) are all excluded on the basis of diverse and different characters. The remaining families for consideration are the Cirolanidae Leach, 1814, Corallanidae Hansen, 1890 and Tridentellidae Bruce, 1984. ...
Article
One new species of Plakolana Bruce, 1993 and three new species of Cirolana Leach (1818), from the Lower Cretaceous (Aptian) Sierra Madre Formation dolomites at El Espinal quarries (Central-East Chiapas, SE Mexico) are reported and described and compared with other fossil species Cirolanidae from around the world. The specimens consist of the body and a few moults, with showing various degrees of decay but most of them with preserved taxonomic features, including mandibles, pereonites, pereopods, pleonites, pleotelson and uropods. This is the third report of fossil marine or estuarine isopods from the Cretaceous of Mexico and the second on the Peracarida of El Espinal quarries, as well as the first record of fossil Cirolana and Plakolana from Mexico.
... Most species of Cymothoidae attach to the integument, fins, tongue, gills and buccal cavity (e.g. those in the genera Anphira, Asotana, Braga, Cymothoa, Livoneca, Paracymothoa and Telotha) (Thatcher, 1988;Araujo et al., 2009;Gomiero et al., 2012;Martin et al., 2016;Murrieta-Morey et al., 2016;Oliveira et al., 2017). Other species drill holes in the host integument (e.g. ...
Article
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Most freshwater species of Cymothoidae are distributed in South America. They have mainly been recorded in the eastern and western regions of the Amazon River basin. However, in this ecosystem, the biodiversity of this group may be greater if the entire Amazon basin is considered. In this regard, the aim of the present study was to provide an updated list of isopod species of the family Cymothoidae that are found in fish in the Brazilian Amazon region and to report on new fish host occurrences and expanded geographical distributions for cymothoid isopods that parasitize fish in the southwestern Brazilian Amazon region. The parasites found in fish specimens were collected, fixed and identified later. We found eight species of Cymothoidae parasitizing different host fish species in the southwestern Amazon region. However, we found 14 species of Cymothoidae throughout the Brazilian Amazon region. Three additional species are thus reported here, which increases the number of species of Cymothoidae in this region to 17. These additional species are also new records for Brazil. Therefore, this study has contribute to expand the knowledge about the distribution and diversity of Cymothoidae in the Amazon basin.
Article
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Cymothoa ianuarii Schioedte & Meinert, 1884 is rediscovered almost 136 years after its original description and redescribed from specimens collected in the state of São Paulo. This species is mainly characterized by adult females with cephalon with conspicuous eyes and not deeply immersed in pereonite 1, pereonites 5-6 much wider than 4, pleotelson twice as wide as long and pleopods 1-5 decreasing in size; Cymothoa excisa Perty, 1833 and C. oestrum (Linnaeus, 1758) are recorded from the state of Bahia. Illustrations and an updated distribution map for these species in Brazil are provided.
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The majority strand of mitochondrial genomes of crustaceans usually exhibits negative GC skews. Most isopods exhibit an inversed strand asymmetry, believed to be a consequence of an inversion of the replication origin (ROI). Recently, we proposed that an additional ROI event in the common ancestor of Cymothoidae and Corallanidae families resulted in a double-inverted skew (negative GC), and that taxa with homoplastic skews cluster together in phylogenetic analyses (long-branch attraction, LBA). Herein, we further explore these hypotheses, for which we sequenced the mitogenome of Asotana magnifica (Cymothoidae), and tested whether our conclusions were biased by poor taxon sampling and inclusion of outgroups. (1) The new mitogenome also exhibits a double-inverted skew, which supports the hypothesis of an additional ROI event in the common ancestor of Cymothoidae and Corallanidae families. (2) It exhibits a unique gene order, which corroborates that isopods possess exceptionally destabilized mitogenomic architecture. (3) Improved taxonomic sampling failed to resolve skew-driven phylogenetic artefacts. (4) The use of a single outgroup exacerbated the LBA, whereas both the use of a large number of outgroups and complete exclusion of outgroups ameliorated it.
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Bambalocra intwalagen. et sp. nov . is described from Sodwana Bay, north-eastern South Africa. The monotypic genus is characterised by the broadly truncate anterior margin of the head with a ventral rostrum, coxae 2–5 being ventral in position not forming part of the body outline and not or barely visible in dorsal view, and the posterolateral margins of pereonites 6 and 7 are posteriorly produced and broadly rounded. The antennulae bases are widely separated, with both antennula and antenna slender. The species is known only from the type locality and the known hosts are species of Pomacanthidae (Angelfish). A revised key to the externally attaching genera of Cymothoidae is provided.
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The parasite fauna of Okinawan coral reef fishes was surprisingly depauperate. Our results differ from the generally accepted greater diversity of Pacific. vs. Atlantic parasite faunas. Previous comparisons have not been based identical-direct examinations of fishes. The rarity of casual parasites (leeches and ageid isopods) suggested that this life strategy may be relatively unsuccessful on Okinawan reefs. Nudibranch and tongue worm associations may be unique to this coral reef system. Most parasites either occurred at levels approximately equivalent to that seen on Caribbean reefs (cymothoid isopods, gnathiid isopods, copepods, barnacles, nematodes, tapeworms), orat lower levels (thorny-headed worms, flukes, gill worms). Only turbellarians were more abundant onOkinawan than Caribbean fishes. Overall the parasites in Okinawan coral reef fishes seem less diverse and less numerous than those in the Caribbean. Tumors, lymphocystis lesions and abnormalities were muchmorerare in Okinawan fishes. Williams, E. H., Jr., L. Bunkley-Williams and W. G. Dyer. 1996. Metazoan parasites of some Okinawan coral reef fishes with a general comparison to the parasites of Caribbean coral reef fishes. Galaxea 13: 1-13. Google Scholar+ [197]
Book
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The Catalog of Fishes covers more than 61,700 species and subspecies, over 11,000 genera and subgenera, and includes in excess of 34,000 bibliographic references. Entries for species, for example, consist of species/subspecies name, genus, author, date, publication, pages, figures, type locality, location of type specimen(s), current status (with references), family/subfamily, and important publication, taxonomic, or nomenclatural notes. Nearly all original descriptions have been examined, and much effort has gone into determining the location of type specimens. Online version: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp
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Hitherto, more than 300 species of isopod crustaceans have been reported in Japan and its neighboring waters, but no comprehensive catalogue has been published in this country. Therefore, in order to promote studies on the group, Saito has prepared a tentative catalogue on isopod crustaceans in Japan. Nunomura (on all the order but the suborder Epicaridea) and Itani (on the suborder Epicaridea) revised the draft. In the present paper, we listed 644 species (29 species of Anthuridea, 19 species of Gnathiidea, 161 species of Asellota, 100 species of Epicaridea. 75 species of Valvifera. 120 species of Flabellifera and 139 species of Oniscidea and 1 species of Tyloidea).
Article
The distribution of the parasites on the host, their mode of attachment to the host tissue, and the incidence and intensity of infestation of 17 species of copepods belonging to eight families and of four species of isopods belonging to the family Cymothoidae infesting 12 species of marine fishes along the South-west (Trivandrum) coast of India are described. The parasites show host specificity and they are highly selective as to the of attachment on the host. Incidence and intensity of infestation of the majority of the parasites examined are higher in the female fish than in the male.