(Digenea: Apocreadiidae) in the Yellowfin Goatfish

Homalometron moraveci n. sp. (Digenea: Apocreadiidae) in the yellowfin goatfish, Mulloidichthys vanicolensis (Valenciennes, 1831) (Perciformes: Mullidae), from New Caledonia and the Great Barrier Reef, with a checklist of digeneans of Mulloidichthys species

RODNEY A. BRAY1, JEAN-LOU JUSTINE2 & THOMAS H. CRIBB3 1Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, UK. Email: [email protected] 2Équipe Biogéographie Marine Tropicale, Unité Systématique, Adaptation, Évolution (CNRS, UPMC, MNHN, IRD), Institut de Recherche pour le Développement, BP A5, 98848 Nouméa Cedex, Nouvelle-Calédonie. E-mail: [email protected] 3Centre for Marine Studies and Department of Microbiology and Parasitology, The University of Queensland, Brisbane, Queensland 4072, Australia. E-mail: [email protected]

Abstract

Homalometron moraveci n. sp. is described from the yellowfin goatfish, Mulloidichthys vanicolensis, from the waters off New Caledonia and from the Swain Reefs of the Great Barrier Reef. Its unique combination of narrow, elongate body and long hermaphroditic duct reaching well into the hindbody, distinguish this species from all other species of Homa- lometron. A checklist of species of digeneans reported in species of Mulloidichthys shows that most of the records are of members of the Opecoelidae and this is the first record of an apocreadiid from this host genus.

Key words: Digenea, Apocreadiidae, Homalometron moraveci n. sp., Mullidae, Mulloidichthys, New Caledonia, Great Barrier Reef

Résumé

Homalometron moraveci n. sp. est décrit de Mulloidichthys vanicolensis de Nouvelle-Calédonie et des Swain Reefs du Récif de la Grande Barrière, Australie. L’espèce est distinguée de tous les autres Homalometron par sa combinaison unique de caractères, son corps allongé et étroit et son long canal hermaphrodite continuant bien dans l’arrière du corps. Une liste des espèces de digènes mentionnées chez les espèces de Mulloidichthys montre que la plupart sont des Ope- coelidae, et que ceci constitue la première mention d’un Apocreadiidae dans ce genre d’hôte.

Introduction

Members of the mullid genus Mulloidichthys Whitley, 1929 are widespread in tropical reef waters of the Atlantic, Indian and Pacific Oceans. We have recovered, and report on here, specimens of the apocreadiid genus Homalometron Stafford, 1904 in the yellowfin goatfish Mulloidichthys vanicolensis (Valenciennes, 1831) from off New Caledonia and from the Swain Reefs in the southern Great Barrier Reef. As far as we are aware this is the first record of an apocreadiid in a member of the family Mullidae (Cribb & Bray 1999). The only previous record of a digenean from M. vanicolensis is that of a sanguinicolid blood-fluke Phthinomita sp. B, from Lizard Island in the northern Great Barrier Reef (Nolan & Cribb 2006). Some confusion exists in the identity of these goatfishes and it is likely that some records of digeneans from other named species of Mul- loidichthys actually refer to records from M. vanicolensis.

Accepted by N. Dronen: 17 May 2007; published: 12 Jul. 2007 41 Materials and methods

Digeneans collected from freshly killed fish were fixed by being pipetted into hot boiling seawater or saline and immediately preserved in ethanol or formalin. Whole-mounts were stained with Mayer's paracarmine, cleared in beechwood creosote and mounted in Canada balsam. Measurements were made through a drawing tube on an Olympus BH-2 microscope, using a Digicad Plus digitising tablet and Carl Zeiss KS100 software adapted by Imaging Associates, and are quoted in micrometres, with the range and the mean in parentheses (Table 1). The following abbreviations are used: BMNH, the British Museum (Natural History) Collection at the Natural History Museum, London, UK; MNHN, Muséum National d’Histoire Naturelle, Paris, France; QM, Queensland Museum Collection, Brisbane, Australia.

TABLE 1. Dimensions of Homalometron moraveci n. sp. from two localities. Locality New Caledonia Swain Reefs n72 Length 3,399–5,881 (4,359) 3,631–3,947 Width 235–396 (319) 307–315 Forebody 904–1,469 (1,104) 958–1,072 Pre-oral lobe 0–36 (8) 7–10 Oral sucker 91–169 x 94–174 (130 x 138) 133–134 x 133–140 Prepharynx 179–326 (237) 233–253 Pharynx 64–117 x 58–81 (91 x 67) 84–86 x 78 Oesophagus 40–129 (80) 40–98 Intestinal bifurcation to ventral sucker 453–748 (565) 463–516 Vitellarium to ventral sucker 302–620 (454) 307–342 Ventral sucker (VS) 151–188 x 137–203 (171 x 175) 159–164 x 171–177 Hermaphroditic duct 308–396 (355) 301–372 Encroachment of hermaphroditic duct into hindbody 119–163 (159) 101–191 Pars prostatica 102–310 (234) 214–243 Seminal vesicle (SV) 122–275 x 64–137 (194 x 114) 99–200 x 84–153 Extent of SV into hindbody 426–722 (559) 490–504 Ventral sucker to ovary 628–1,119 (838) 707–722 Ovary 100–189 x 86–175 (149 x 142) 125–130 x 114–125 Ovary to Anterior testis 0–7 (1) 9–10 Anterior testis 222–367 x 122–261 (287 x 204) 234–253 x 171–174 Distance between testes 0–94 (32) 5–56 Posterior testis 268–457 x 141–281 (337 x 221) 264–275 x 171–178 Post-testicular region 924–2,068 (1,437) 1,126–1,291 Post-caecal region 192–320 (264) 136–213 (168) Eggs 59–80 x 34–54 (74 x 45) non-ovigerous Width as % of body-length 6.38–8.53 (7.40) 7.77–8.68 Forebody as % of body-length 22.3–28.5 (25.6) 26.4–27.2 Sucker-length ratio 1:1.04–2.00 (1.37) 1:1.19–1.23 Sucker-width ratio 1:1.10–1.97 (1.32) 1:1.27–1.29 Oral sucker:pharynx 1:1.56–2.49 (2.05) 1:1.71–1.80 VS to ovary as % of body-length 16.4–21.2 (19.1) 18.3–19.5 VS to SV as % of VS to ovary 58.7–72.2 (67.2) 67.9–71.3 Post-testicular region as % of body-length 27.2–38.9 (32.7) 31.0–32.7

Results

Family Apocreadiidae Skrjabin, 1942

Subfamily Apocreadiinae Skrjabin, 1942

Genus Homalometron Stafford, 1904

Homalometron moraveci n. sp. (Figs 1–3)

Type-host: Mulloidichthys vanicolensis (Valenciennes, 1831), Mullidae, yellowfin goatfish. Site: Digestive tract. Type-locality: Fish market, Nouméa, allegedly from off Bouraké, Commune de Boulouparis, New Cale- donia (21°58’S, 166°00’E) (21/09/2006). Other locality: Swain Reefs, Great Barrier Reef (21°53’S, 152°21’E, Feb., 2003). Prevalence: New Caledonia, 25% (1 of 4); Swain Reefs, 100% (1 of 1). Type-specimens: New Caledonia: Holotype MNHN JNC1966-1, paratypes MNHN JNC1966-2–8, BMNH 2007.5.1.1–5, QM G223078; Swain Reefs: paratypes QM G223079, BMNH 2007.5.1.6. Etymology: The species is named for Dr Frank Moravec (České Budějovice, Czech Republic) in recogni- tion of his major contributions to fish parasitology, and as he was involved in the collection of the specimens. Description. Based on 11 complete worms and several fragments from New Caledonia and 2 well-devel- oped, but non-ovigerous, worms from Swain Reefs. Measurements in Table 1. Body-elongate, narrow. Tegu- ment spinous, spines reach to posterior extremity, sparse in hindbody. Pre-oral lobed short or absent. Oral sucker subglobular, mouth subterminal. Ventral sucker rounded to oval, slightly to distinctly larger than oral sucker. Forebody relatively short. Prepharynx long. Pharynx oval, small. Oesophagus distinct. Intestinal bifurcation in mid-forebody. Caeca long, narrow, terminate blindly, unequal, slightly separated from posterior extremity. Testes two, oval with long axis longitudinally, entire, tandem, contiguous to distinctly separate, in mid- hindbody. Seminal vesicle oval, about halfway between ventral sucker and ovary. Pars prostatica long, narrow, sheath of gland cells difficult to distinguish from other gland-cells scattered in parenchyma, some evi- dence in some specimens of slightly denser packing of cells in this region, enters hermaphroditic duct distinctly posterior to ventral sucker. No ejaculatory duct or cirrus detected. Hermaphroditic duct long, narrow, passes dorsally to ventral sucker, distinctly into hindbody (Table 1). Genital pore opens in median line immediately anterior to ventral sucker. Ovary subglobular to oval, pretesticular, contiguous with or slightly separated from anterior testis. Semi- nal receptacle canalicular, oval, mostly pretesticular. Laurer’s canal opens dorsally to ovary. Mehlis’ gland preovarian. Uterus preovarian, intercaecal. Metraterm not strongly differentiated, starts at about posterior level of seminal receptacle, walls narrow. Eggs few, large, operculate, with thin, tanned shells. Vitellarium fol- licular, anterior extent in region of seminal vesicle, ventral, dorsal and lateral to caeca, not overlapping gonads, may have gaps at or around ovarian level, confluent dorsally and ventrally in post-testicular region. Excretory pore terminal. Vesicle I-shaped, reaches almost to posterior testis. Lymphatic vessels not detected.

HOMALOMETRON MORAVECI N. SP. FROM GOATFISH Zootaxa 1525 © 2007 Magnolia Press · 43 FIGURES 1–3. Homalometron moraveci n. sp. 1. Holotype, ventral view, New Caledonia. 2. Ventral view, immature specimen, Swain Reefs. 3. Terminal genitalia, lateral view, New Caledonia. 4. Proximal female system in immature specimen, Swain Reefs, dorsal view. Abbreviations: LC, Laurer’s canal; M, metraterm; O, ovary; PP, pars prostatica; SR, seminal receptacle; SV, seminal vesicle; T, testis; U, uterus; V, vitellarium; VR, vitelline reservoir. Scale-bars: 1, 2, 1000μm; 3, 200μm; 4, 100μm.

This species fits into the concept of Homalometron Stafford, 1904 espoused by Cribb & Bray (1999) and Cribb (2005). Unusual or unique features are the narrow, elongate body and the long hermaphroditic duct reaching well into the hindbody. Cribb & Bray (1999) listed the then recognised species of Homalometron, and divided them into 8 mor- phologically distinguishable groups. One further species has subsequently been recognised, Homalometron papilliferum (Szidat, 1956), which has twice been redescribed recently (Ostrowski de Nuñez et al. 2000; Shi- mazu et al. 2000). It fits into Group 2 of Cribb & Bray (1999), having three pairs of papillae on the ventral sucker. Groups 1–7 are clearly distinguishable from H. moraveci n. sp., leaving only Group 8, the ‘Typical species’, as a possible venue for this new form. This group contains 12 species, 7 of which can immediately be distinguished from H. moraveci by their relatively short oval body-shape, by the seminal vesicle being dorsal to or overlapping the ventral sucker and with the vitellarium reaching to the ventral sucker: H. pallidum Stafford, 1904, the type-species, H. armatum (MacCallum, 1895), H. dowgialloi Dyer, Williams & Bunkley-Williams, 1992, H. manteri (Overstreet, 1970), H. pseudopallidum Martorelli, 1986, H. sophiae (Stossich, 1886) and H. vinodae (Ahmad, 1985) (Stossich 1886; Hunter & Bangham 1932; Miller 1959; Stunkard 1964; Overstreet 1970; Ahmad 1985; Martorelli 1986; Dyer et al. 1992). Other species in group 8 differ from H. moraveci n. sp. in the following characteristics: i. H. longisinosum (Manter, 1937) from tetraodontids in the tropical eastern Pacific differs in its distinctly fusiform shape, its relatively short hermaphroditic duct, reaching dorsally to the anterior part of the ventral sucker (and originally also called a genital sinus in the figure legend) and its ‘slightly lobed, squarish’ testes (Manter 1937). ii. H. mexicanum (Manter, 1937) from balistids, labrisomids and monacanthids from the Pacific coast of Mexico and the Gulf of Mexico and adjacent waters, is somewhat fusiform and flattened with thin, frill-like marginal folds, the hermaphroditic duct is short, reaching to near the anterior margin of the ventral sucker and the testes are ‘more or less rectangular’ and lobed (Manter 1937). iii. H. stradbrokensis Cribb & Bray, 1999 from the common silver belly Gerres subfasciatus Cuvier, 1830 [Gerreidae] from off Stradbroke Island, southern Queensland, Australia, is a similarly elongate form, but has a relatively shorter hermaphroditic duct and prepharynx, a short oval pars prostatica and greater distances between the gonads (Cribb & Bray 1999). iv. H. synagris (Yamaguti, 1953) from nemipterids in the Indo-West Pacific is elongate, but slightly lan- ceolate, with a hermaphroditic duct reaching dorsally to the middle of the ventral sucker, a short oval pars prostatica, a seminal vesicle that overlaps the ventral sucker, a very short distance between the ventral sucker and the ovary and a relatively longer post-testicular region (Yamaguti 1953; Cribb & Bray 1999). v. H. verrunculi Cribb & Bray, 1999 (replacement name for Homalometron caballeroi Lamothe- Argumedo, 1965 nec (Bravo-Hollis, 1953)) from the finescale triggerfish Balistes polylepis Steindachner, 1876 [Balistidae] in the Gulf of California is distinctly relatively wider, the hermaphroditic duct reaches dorsally to the ventral sucker and the distance between the ovary and ventral sucker is relatively small (Lamothe- Argumedo 1965).

Digenean parasites of Mulloidichthys spp.

Mulloidichthys is a small genus of bottom-dwelling coral reef fishes, with about six recognised species (Michael 2004). There is some confusion over the identity of some of the hosts. M. auriflamma (Forsskål, 1775) has been reported as host of digeneans from various localities. According to Froese & Pauly (2007) this

HOMALOMETRON MORAVECI N. SP. FROM GOATFISH Zootaxa 1525 © 2007 Magnolia Press · 45 name has been replaced by Parupeneus forsskali (Fourmanoir & Guézé, 1976), the Red Sea goatfish, which occurs in the Red Sea and Gulf of Aden. Apparently, the use of the name M. auriflamma elsewhere probably refers to the yellowfin goatfish M. vanicolensis or the yellowstripe goatfish M. flavolineatus (Lacepède, 1801). M. samoensis Günther, 1874 is now considered a synonym of M. flavolineatus. The records of digeneans from Mulloidichthys species are included in table 2. Twenty-six species are reported as adult worms, 50% of them being opecoelids and 23% hemiurids. In terms of the number of records 61% are of opecoelids and 23% hemiurids. No apocreadiids have previously been reported from this or any other mullid. This new family for the genus only extends its pattern of low host-specificity. Of the 25 families of fishes now known to harbour species of Homalometron, only three (Balistidae, Gerreidae and Soleidae) harbour more than three species; 16 families harbour only one species of Homalometron. No other digenean was found in M. vanicolensis from New Caledonia.

TABLE 2. Records of digeneans from Mulloidichthys species.

Family Parasite Species Host Species Reference Locality Acanthocolpidae Stephanostomum sp. metacercaria M. flavolineatus (El-Naffar et al. 1992) Arabian Gulf Stephanostomum sp. metacercaria II M. flavolineatus (Kardousha 2005) Arabian Gulf Apocreadiidae Homalometron moraveci n. sp. M. vanicolensis Present study New Caledonia Present study Great Barrier Reef Bucephalidae Dollfustrema sp. juvenile M. pfluegeri (Yamaguti 1970) Hawaii Prosorhynchus sp. juvenile 'M. auriflamma' (Yamaguti 1970) Hawaii Didymozoidae Epithelionematobothrium mulloidich- M. pfluegeri (Yamaguti 1970) Hawaii thydis Yamaguti, 1970 Hemiuridae Aphanurus stossichii (Monticelli, 'M. auriflamma' (Nahhas & Sey 2002) Arabian Gulf 1891) Clupenurus srivastavai Ramadan, 'M. auriflamma' (Ramadan 1986) Red Sea 1986 Elytrophallus mulloidichthydis M. pfluegeri (Yamaguti 1970) Hawaii Yamaguti, 1970 Lecithochirium parvum Manter, 1947 M. martinicus (Nahhas & Cable 1964) Jamaica Lecithochirium priacanthi Yamaguti, 'M. auriflamma' (Yamaguti 1970) Hawaii 1970 M. pfluegeri (Yamaguti 1970) Hawaii Sterrhurus monticellii (Linton, 1898) M. martinicus (Siddiqi & Cable 1960) Puerto Rico Lecithasteridae Weketrema hawaiiensis (Yamaguti, M. flavolineatus (Yamaguti 1970) Hawaii 1970) M. pfluegeri (Yamaguti 1970) Hawaii Lepocreadiidae Neolepidapedon (Neolepidape- 'M. auriflamma' (Ramadan 1987) Red Sea doides) yamagutii Ramadan, 1987 Monorchiidae Lasiotocus delicatus Manter & Prit- 'M. auriflamma' (Manter & Pritchard Hawaii chard, 1961 1961) Lasiotocus weke Yamaguti, 1970 M. flavolineatus (Yamaguti 1970) Hawaii M. pfluegeri (Yamaguti 1970) Hawaii Opecoelidae Dactylostomum pfluegeri (Yamaguti, M. pfluegeri (Yamaguti 1970) Hawaii 1970) to be continued.

46 · Zootaxa 1525 © 2007 Magnolia Press BRAY ET AL. TABLE 2. (continued) Family Parasite Species Host Species Reference Locality Dactylostomum woolcocki Ramadan, 'M. auriflamma' (Ramadan 1985) Red Sea 1985 Dactylostomum yamagutii Ramadan, 'M. auriflamma' (Ramadan 1985) Red Sea 1985 Helicometrina trachinoti Siddiqi & M. martinicus (Siddiqi & Cable 1960) Puerto Rico Cable, 1960 Opecoeloides brachyteleus Manter, M. martinicus (Manter 1947) Florida 1947 (Siddiqi & Cable 1960) Puerto Rico (Nahhas & Cable 1964) Curacao (Nahhas & Cable 1964) Jamaica (Nahhas & Carlson Jamaica 1994) Opecoeloides elongatus Manter, 1947 M. martinicus (Siddiqi & Cable 1960) Puerto Rico (Nahhas & Cable 1964) Curacao (Nahhas & Cable 1964) Jamaica (Dyer et al. 1985) Puerto Rico (Nahhas & Carlson Jamaica 1994) Opecoeloides vitellosus (Linton, M. martinicus (Siddiqi & Cable 1960) Puerto Rico 1900) (Dyer et al. 1992) West Indies (Dyer et al. 1998) Puerto Rico Opecoelus adelongatus Nagaty, 1954 'M. auriflamma' (Nagaty 1954) Red Sea Opecoelus mulloidichthydis M. flavolineatus (Yamaguti 1970) Hawaii Yamaguti, 1970 M. pfluegeri (Yamaguti 1970) Hawaii 'M. auriflamma' (Yamaguti 1970) Hawaii Opegaster hawaiiensis Yamaguti, M. pfluegeri (Yamaguti 1970) Hawaii 1970 Paropecoelus lanceolatus (Martin, M. flavolineatus (Martin 1960) Hawaii 1960) 'M. auriflamma' (Pritchard 1966) Hawaii Pseudopecoelina sp. M. flavolineatus (El-Naffar et al. 1992) Arabian Gulf Pseudopecoeloides wekeula 'M. auriflamma' (Yamaguti 1970) Hawaii Yamaguti, 1970 M. flavolineatus (Yamaguti 1970) Hawaii Sanguinicolidae Phthinomita sp. B M. vanicolensis (Nolan & Cribb 2006) Lizard Island Zoogonidae Neozoogonus longicecus Siddiqi & M. martinicus (Siddiqi & Cable 1960) Puerto Rico Cable, 1960 (Dyer et al. 1998) Puerto Rico

Frank Moravec (České Budějovice, Czech Republic) and Aude Sigura, student, participated to the parasito- logical survey in New Caledonia. Fishes from New Caledonia were identified from photographs by Ronald Fricke (Staatliches Museum für Naturkunde, Stuttgart, Germany).

References

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