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DOI: 10.2478/s11686-012-0039-0 © W. Stefan´ski Institute of Parasitology, PAS Acta Parasitologica, 2012, 57(3), 247–272; ISSN 1230-2821

A review of the Lepocreadiidae (Digenea, Lepocreadioidea) from fishes of the waters around New Caledonia Rodney A. Bray1 and Jean-Lou Justine2 1 Department of Zoology, Natural History Museum, Cromwell Road, London SW7 5BD, UK; 2UMR 7138 Systématique, Adaptation, Évolution, Muséum National d’Histoire Naturelle, Case postale 51, 55 rue Buffon, 75231 Paris cedex 05, France

Abstract The lepocreadiid fauna of New Caledonia is reported and discussed and a new species and several new host and locality records presented. Hypoporus plataxi sp. nov. from Platax teira is described and distinguished from its only congener by its terminal genitalia, the structure of the anterior part of the alimentary system and other morphological features. New host and locality records and a description are given of Lepotrema cf. clavatum Ozaki, 1932 in Sufflamen fraenatum. New host and locality records are given of Lobatocreadium exiguum (Manter, 1963) in Pseudobalistes fuscus, which is also reported in the known hosts Abalistes filamentosus and Sufflamen fraenatum. New host and locality records are given of Opisthogonoporus amadai Yamaguti, 1937 in Branchiostegus wardi. A new host record is made for Holorchis plectorhynchi Durio et Manter, 1968 in Diagramma pictum. New records in New Caledonian waters are of Bulbocirrus aulostomi Yamaguti, 1965 in Aulostomus chinensis, Echeneidocoelium indicum Simha et Pershad, 1964 in Echeneis naucrates, Lepidapedoides kalikali Yamaguti, 1970 in Pristipomoides auricilla, Neomultitestis aspidogastriformis Bray et Cribb, 2003 in Platax teira, Opechona bacillaris (Molin, 1859) in Rastrelliger kanagurta, Prodistomum keyam Bray et Cribb, 1996 in Monodactylus argenteus and Pseudopisthogonoporus vitellosus (Pritchard, 1963) in Naso brevirostris and N. annulatus. New metrical data are presented for Holorchis castex Bray et Justine, 2007 in Diagramma pictum, Hypocreadium patellare Yamaguti, 1938 in Sufflamen fraenatum, Intusatrium robustum Durio et Manter, 1968 in Bodianus loxozonus and B. perditio and Lepidapedoides angustus Bray, Cribb et Barker, 1996 in Epinephelus chlorostigma, E. fasciatus, E. maculatus and E. retouti. Literature records are included and the fauna in general discussed.

Keywords New Caledonia, Digenea, Lepocreadiidae, Hypoporus plataxi sp. nov., zoogeography, host-specificity

Introduction

Materials and methods

The Lepocreadiidae is probably not a monophyletic group (Bray, et al., 2009b), but until more evidence is available and a robust classification can be developed, the system espoused by Bray (2005) is used. It is one of the largest digenean families with over 75 genera and over 200 species. Several reports of lepocreadiids have been made from New Caledonia, starting with Durio and Manter (1968), and these are listed below, interspersed with new data on lepocreadiids in this region. Only 10 species of lepocreadiids were listed for New Caledonian fishes by Justine (2010). Now, therefore, 32 New Caledonian species are known, belonging to 22 genera, and reported from 38 host species.

Digeneans were collected live, immediately fixed in nearly boiling saline and then transferred to 80% ethanol following the protocol of Cribb and Bray (2010). 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. The following abbreviations are used: BMNH, British Museum (Natural History) Collection at the Natural History Museum, London, UK; MNHN JNC, Muséum National d’Histoire Naturelle, Paris, France.

Corresponding author: [email protected]

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Table I. Measurements and ratios of Bulbocirrus aulostomi and Echeneidocoelium indicum Species Host n Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% External seminal vesicle reach into hindbody%** Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

Bulbocirrus aulostomi Aulostomus chinensis 2 9744 314 1170 0 46 50 377 73 53 117 547 3858 121 131 311 178 3869 5906 226 159 451 460 226 90 530 226 610 ? 46 21 3.22 12.0 2.62 2.61 0.94 60.6 65.5 6.3 1.20 5.61 39.6 4.63 0.92 3.19 5032 51.6 4.72 5.43

6642 223 994 0 40 48 314 60 36 119 457 2095 101 86 232 133 2114 3341 171 139 373 376 221 59 438 229 636 ? 56 27 3.35 15.0 2.55 1.78 1.34 50.3 63.3 9.6 1.79 6.89 31.5 5.61 0.89 3.50 3093 46.6 5.67 6.59

Echeneidocoelium indicum Echeneis naucrates 3 min max mean 3628 392 636 0 103 107 322 148 112 0 23 66 210 208 428 48 508 877 230 200 21 581 247 0 936 212 95 52 27 6.96 9.82 2.05 1.94 0.86 24.2 55.0 2.07 0 0.51 1.82 0 0 7.8 912 16.1 16.0 23.6

7305 672 765 11 136 159 406 191 154 28 37 183 327 317 664 106 1239 2254 495 310 529 1363 421 0 1725 339 151 108 69 35 13.7 17.5 3.00 2.27 1.04 30.9 72.1 2.61 0.76 0.67 3.71 7.24 0 13.5 1275 25.9 19.2 25.8

5284 524 706 4 114 133 368 172 139 12 31 131 283 274 554 76 893 1475 379 251 225 962 309 0 1308 278 120 79 65 31 10.5 14.3 2.48 2.07 0.95 27.1 61.7 2.32 0.31 0.60 2.51 3.72 0 11.0 1121 22.4 17.9 25.0

The % sign in the tables refers to the percentage relative to the body-length, except when accompanied by ** which means percentage of ventral sucker to ovary distance.

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Class Trematoda Rudolphi, 1808 Subclass Digenea Carus, 1863 Order Plagiorchiida La Rue, 1957 Suborder Lepocreadiata Olson, Cribb, Tkach, Bray et Littlewood, 2003 Superfamily Lepocreadioidea Odhner, 1905 Family Lepocreadiidae Odhner, 1905 Subfamily Lepocreadiinae Odhner, 1905 Genus Bianium Stunkard, 1930 Bianium arabicum Sey, 1996 Record from New Caledonia: Bray et al. (2010a). New Caledonian host: Lagocephalus sceleratus (Gmelin, 1789), Tetraodontidae. Genus Bulbocirrus Yamaguti, 1965 Bulbocirrus aulostomi Yamaguti, 1965 Host: Aulostomus chinensis (Linnaeus, 1766), Aulostomidae, Chinese trumpetfish. Site: digestive tract. Locality: Récif du Prony (22°15´869˝S, 166°19´680˝E, 26/10/2007). Prevalence: 1 of 1. Specimens: MNHN JNC2350. Previous New Caledonian records: None. Discussion: These specimens (Table I) are indistinguishable from the specimens from the same host species described from off Hawaii (Yamaguti 1965, 1970) and Heron Island, southern Great Barrier Reef (Bray and Cribb 1998). This constitutes our complete knowledge of the distribution of this species.

Diploproctodaeum rutellum (Mamaev, 1970) Bray, Cribb et Barker, 1996 Record from New Caledonia: Bray et al. (2010a). New Caledonian host: Platax teira (Forsskål, 1775), Ephippidae. Diploproctodaeum triodoni Bray, Cribb et Justine, 2010 Record from New Caledonia: Bray et al. (2010a). New Caledonian host: Triodon macropterus Lesson, 1831 (Triodontidae). Genus Echeneidocoelium Simha et Pershad, 1964 Echeneidocoelium indicum Simha et Pershad, 1964 Host: Echeneis naucrates Linnaeus, 1758, Echeneidae, live sharksucker. Site: intestine, digestive tract. Localities: Near Baie des Citrons, Nouméa (22°17´30˝S 166°25´30˝E, 10/10/2006); Off Baie des Citrons (22°17´441˝S, 166°25´446˝E, 14/06/2008); Off Récif Kué (22°34´476˝S, 166°30´364˝E, 14/10/2008); Interior Lagoon near Récif Toombo (22°32´536˝S, 166°29´069˝E, 21/04/2009); Off Pointe Bovis, Baie Maa (22°12´72˝S, 166°19´519˝E, 26/11/2009). Prevalence: 5 of 5 (100%). Specimens: MNHN JNC 2685, JNC2916, JNC3131, BMNH 2012.5.25.1. Previous New Caledonian Records: None. Discussion: This species is widely reported from echeneid fishes, in the Indian Ocean (Simha and Pershad 1964, Madhavi, 1970, Lakshmi and Rao 1978) and the Great Barrier Reef (Bray and Cribb 1998). It has also been reported (? as a Lessepsian migrant) off the Mediterranean coast of Israel (Fischthal 1982). Our specimens are not distinguishable from those described by earlier authors (Table I).

Genus Diploproctodaeoides Reimer, 1981 Genus Hypocreadium Ozaki, 1936 Diploproctodaeoides longipygum (Oshmarin, Mamaev et Parukhin, 1961) Reimer, 1981 Record from New Caledonia: Bray et al. (2010a). New Caledonian host: Abalistes stellatus (Anonymous, 1798), Balistidae. Genus Diploproctodaeum La Rue, 1926 Diploproctodaeum arothroni Bray et Nahhas, 1998 Record from New Caledonia: Bray et al. (2010a). New Caledonian host: Arothron manilensis (Marion de Proce, 1822), Tetraodontidae. Diploproctodaeum haustrum (MacCallum, 1919) La Rue, 1926 Record from New Caledonia: Bray et al. (2010a). New Caledonian host: Aluterus monoceros (Linnaeus, 1758) (Monacanthidae).

Hypocreadium patellare Yamaguti, 1938 (Fig. 1) Host: Sufflamen fraenatum (Latreille, 1804), Balistidae, masked triggerfish. Site: digestive tract. Localities: near Ile Redika (22°32´S, 166°37´E, 11/06/2009); Interior Lagoon near Récif Toombo (22°32´536˝S, 166°29´069˝, 01/09/2009). Prevalence: 8 of 10 (80%). Specimens: MNHN JNC 2772B, JNC2940, JNC2978, JNC3034, BMNH 2012.5.25.12-15. Previous New Caledonian record: Bray et al. (2009a). Previously reported host from New Caledonia: Sufflamen fraenatum. Discussion: We find two forms of Hypocreadium in S. fraenatum. Some fit comfortably into the typical form of H. patellare as reported by us (Bray et al. 2009a) from this

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Genus Hypoporus Wang, 1989 Hypoporus plataxi sp. nov. (Figs 2–5)

Fig. 1. Hypocreadium patellare Yamaguti, 1938, pyriform form, in Sufflamen fraenatum. Ventral view, uterus in outline. Scale-bar = 500 μm

host off New Caledonia (see Table II). The other form (JNC 2978) appears to differ in being slightly pyriform (Fig. 1) and having, at best, a weakly developed anterior notch. In addition, the ventral sucker is more or less circular or oval with the long axis parallel to the equator (vs oval with the long axis at an angle to the equator, the usual arrangement in typical H. patellare (Bray et al. 2009a)). In the case of the JNC 2978 specimens, it has been difficult to mount the worms as well as we would have liked, and the edges are slightly folded, which may account for the slightly pyriform shape. As with typical H. patellare the genital pore is at the anterior oesophagus/pharynx level and the uterus reaches post-ovarian. We are considering JNC 2978 worms as another atypical H. patellare, but point out the differentiating features, some of which may disappear with further study: the pyriform shape, the slight or absent anterior notch and the ventral sucker shape and orientation. Molecular studies will be needed to elucidate the relationships of the atypical forms of H. patellare. Hypocreadium toombo Bray et Justine, 2006 Records from New Caledonia: 1. Bray and Justine (2006), 2. Bray et al. (2009b). New Caledonian host: Pseudobalistes fuscus (Bloch et Schneider, 1801) (Balistidae).

Host: Platax teira (Forsskål, 1775), Ephippidae, longfin batfish. Site: intestine, stomach. Locality: Nouméa Fish Market (11/03/2009). Prevalence: 1 of 2 (50%). Specimens: MNHN Holotype JNC2872A. Description: Based on 1 whole-mounted specimen, measurements in Table III. Body elongate linguiform, robust (Fig. 2). Tegument finely spined, spines reaching to level of posterior testis. Eye-spot pigment scattered at level of pharynx. Oral sucker large, subglobular, subterminal. Ventral sucker oval, smaller than oral sucker, in anterior third of body. Prepharynx short, entirely in posterior cavity of oral sucker, wall thick. Pharynx large, oval. Oesophagus short, narrow. Intestinal bifurcation in mid forebody. Two short anteriorly directed diverticula arise from bifurcal area. Caeca broad, end blindly close to posterior extremity. Testes two, oval, entire, tandem, contiguous, in mid hindbody. External seminal vesicle large, elongate oval, reaches dorsally over ovary. Cirrus-sac broadly claviform, mainly posterior to ventral sucker. Internal seminal broad, coiled, ventral in cirrus-sac (Fig. 3). Pars prostatica long, wide, coiled dorsally over seminal vesicle; apparently narrows distally passing into ejaculatory duct via narrow aperture (Fig. 4). Ejaculatory duct wide saccular proximally, with irregularly wrinkled thin wall, forming small, flask-shaped, relatively thick-walled chamber distally. Genital atrium distinct. Genital pore sinistral, at level of posterior part of the ventral sucker. Ovary trilobate, pretesticular with short gap. Seminal receptacle subglobular, postero-sinistral to ovary. Laurer’s canal and Mehlis’ gland not detected. Uterus intercaecal, preovarian, almost entirely in hindbody. Eggs tanned operculate. Metraterm full of eggs, wide, with glandular sheath, slightly shorter than cirrus-sac (Fig. 5). Vitellarium follicular, reaching from posterior margin of ventral sucker to posterior extremity, confluent in post-testicular region; lateral, ventral and dorsal to caeca, encroaching over gonads but not confluent in gonadal area. Excretory pore terminal; vesicle wide reaches to intestinal bifurcation. Discussion: We feel justified in describing this single specimen as it has many unusual features. It must be said that the cirrus-sac contents are hard to interpret and we have attempted to give an overall view of these contents, but we are not certain that all the details are correct. The contents are, however, unusual enough to be recognised when further specimens are found. This species does not fit comfortably into any known genus. As the terminal genitalia are of neither the ‘Opechona’ nor the ‘Lepidapedon’ type it is difficult to key satisfactorily to subfamily in Bray (2005). Nevertheless, attempting the

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Table II. Measurements and ratios of the two forms of Hypocreadium patellare Form Host n

Pyriform form Sufflamen fraenatum 5 min max mean

Typical form Sufflamen fraenatum 4 min max mean

Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Post-vitelline distance Post-ovarian distance Post-uterine distance Genital pore to ventral sucker External seminal vesicle length

1072 884 417 0 114 121 0 74 87 83 145 188 127 141 295 114 7 107 92 0 154 107 129 135 101 362 168 61 27 78.4 37.2 1.11 1.02 1.29 0.57 30.9 6.95 12.8 16.7 0 11.5 26.2 176 16.4 123 355 289 171 52

1042 1085 403 11 106 119 0 71 90 84 124 220 120 155 318 118 34 102 105 0 165 137 115 149 143 356 194 61 27 98.2 37.9 1.13 1.09 1.25 3.30 33.9 7.29 11.7 18.7 0.0 10.9 27.4 157 14.8 107 353 317 199 70

1191 1107 490 15 147 156 10 100 115 113 190 244 164 161 358 139 46 132 112 17 194 160 161 189 143 393 231 72 37 100 41.1 1.20 1.23 1.39 4.04 35.6 10.6 15.9 22.7 1.56 14.9 32.9 245 20.6 156 449 349 298 113

1130 1040 442 10 125 135 3 86 100 98 160 222 145 155 332 127 28 117 101 3 171 128 147 168 117 379 201 65 32 92.1 39.1 1.16 1.16 1.35 2.53 33.6 8.74 14.1 19.7 0.31 13.0 29.4 220 19.4 145 393 312 221 79

1302 1278 540 16 114 145 0 77 97 109 171 277 158 168 357 137 68 130 126 8 208 155 170 179 170 450 279 70 33 105.1 41.5 1.39 1.33 1.50 5.93 35.9 8.44 13.1 23.1 0.74 13.0 31.7 262 20.1 175 447 395 218 136

1145 1174 449 14 112 128 0 74 93 92 142 242 138 160 333 127 53 115 116 3 184 147 136 163 157 397 237 65 30 102.7 39.1 1.23 1.25 1.38 4.64 34.7 8.00 12.4 21.1 0.25 11.8 29.2 207 17.9 133 395 356 208 103

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Table II. Continuation Form Host n

Pyriform form Sufflamen fraenatum 5 min max mean

Typical form Sufflamen fraenatum 4 min max mean

External seminal vesicle width Metraterm length Post-vitelline distance% Post-ovarian distance% Post-uterine distance%

31 224 11.0 33.1 25.6

57 239 10.1 33.5 30.3

keys in both subfamilies whose members have a cirrus-sac, Lepocreadiinae and Lepidapedinae, this form appears closest to Hypoporus Wang, 1989. The type, and only known species, of this genus is H. phylloides Wang, 1989 reported in the Pacific rudderfish Psenopsis anomala (Temminck et Schlegel, 1844) (Perciformes, Centrolophidae) from off Fujian Province, China (Wang 1989). The details of the terminal genitalia of H. phylloides are not well known, but the genital pore is postero-sinistral to the ventral sucker, as in H. plataxi sp. nov. The ovary is described as round or lobed, and illustrated as lobed, as in H. plataxi. The species differ, however, in that the prepharynx and oesophagus are distinct in H. phylloides, and anteriorly directed intestinal diverticula are absent. The internal seminal vesicle of H. phylloides is illustrated as small and subglobular. The external seminal vesicle in H. phylloides reaches only to the middle of the ventral sucker to ovary distance. Despite several reports of digeneans from this host, this parasite has not been reported from other parts of the IndoPacific Ocean (Madhavi 1972, Gu and Shen 1979, Dyer et al. 1988, Hafeezullah 1990, Bray and Cribb 2003, Bray et al. 2009b, Bray et al. 2010b). Genus Intusatrium Durio et Manter, 1968 Intusatrium robustum Durio et Manter, 1968 (Figs 6–8) Hosts: Bodianus loxozonus (Snyder, 1908), Labridae, blackfin hogfish; Bodianus perditio (Quoy et Gaimard, 1834), Labridae, golden-spot hogfish. Site: Intestine, digestive tract. Locality: B. loxozonus, Grand récif Aboré, côté Passe de Dumbéa (22°21´30˝S, 166°15´30˝E, 22/10/2004); B. perditio, Grand récif Aboré, côté Passe de Dumbéa (22°21´30˝S, 166°15´30˝E, 22/10/2004); Passe de Boulari (22°32´S, 166°28´E, 09/11/2005); Récif Le Sournois (22°31´30˝S, 166°26´30˝E, 03/10/2006); Fish Market (09/07/2008); Off Récif Kué (22°35´238S, 166°29´838E, 22/10/2008); Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 20/05/2009). Prevalence: B. loxozonus, 2 of 3 (67%); B. perditio, 8 of 8 (100%). Specimens: B. loxozonus, MNHN JNC 1219, 1407, BMNH 2006.8.23.11–13; B. perditio, MNHN JNC 1404, 1417, 1418,

134 296 14.6 37.7 29.3

64 264 12.8 34.7 27.6

93 292 13.4 36.1 33.1

79 268 11.5 34.5 31.0

1633, JNC1969, JNC2619, JNC 2707B, JNC2939, BMNH 2006.8.23.21–26, 2012.5.25.19. Previous New Caledonian records: 1. Durio and Manter (1968), 2. Bray and Justine (2006), 3. Bray et al. (2009b). Previously reported New Caledonian hosts: Bodianus loxozonus (2), B. perditio (1, 2, 3). Discussion: This species is known only from the waters off New Caledonia. Durio and Manter (1968) reported the species in Bodianus [as Lepidaplois] perditio and Bray and Justine (2006) reported the species in B. perditio and B. loxozonus. There is some confusion over the detail of the structure of the male terminal genitalia. Durio and Manter (1968) described the cirrus-sac as ‘containing in its posterior half a cellular-walled tube (= prostatic vesicle ?), straight or sinuous, followed by a prostatic tube of about same length lined with microvilli, and terminally a thin-walled expansion followed by very short, thick-walled cirrus. Prostatic cells few, near middle of cirrus sac; darker staining cells in posterior part of sac.’ They also described the genital atrium as having an ‘ovoid outpocketing (atrial sac) extending within cirrus sac alongside prostatic tube almost to middle of cirrus sac, often containing eggs.’ Yamaguti (1971) thought that the atrial sac described by Durio and Manter was, in fact, a ‘swollen pars prostatica’. Bray et al. (1997) illustrated the terminal genitalia and found no sperm in the internal male duct, the walls of which varied in thickness and the definition adopted by Bray (2005) was based on this observation. Having examined several specimens from the type host, B. perditio we find a thick-walled internal male duct, expanded about centrally to form a small, vesicular pars prostatica (Figs 7–8). Sperm was not seen in the internal male duct. In several cases a cirrus is formed, protruding from the genital pore and in this case the internal male duct is stretched to a more or less rectilinear course. Durio and Manter (1968) detected similarities to the Haploporidae in this species, but molecular evidence (Bray et al. 2009b) indicates that it is a lepocreadioid, and clusters with a morphologically variable group of labrid parasites. Measurements of our specimens are given in Table IV. Two species of Lepocreadium Stossich, 1903 from labrids show similarities to Intusatrium. Lepocreadium bimarinum Manter, 1940 was first reported in Semicossyphus pulcher (Ayres, 1854) (as Pime-

Lepocreadiidae (Digenea) from fishes of the New Caledonia

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Figs 2–5. Hypoporus plataxi sp. nov. in Platax teira. 2. Ventral view, uterus in outline. 3. Ventral plane of cirrus-sac. Abbreviations: ed – ejaculatory duct, ga – genital atrium, gp – genital pore, isv – internal seminal vesicle, pp – pars prostatica, vs – outline of ventral sucker. 4. Dorsal plane of cirrus-sac, external seminal vesicle, ovary and seminal receptacle. Abbreviations: ed – ejaculatory duct, esv – external seminal vesicle, ov – outline of ovary, pp – pars prostatica, sr – seminal receptacle. 5. Metraterm and glandular sheath. Abbreviations: cs – outline of cirrus-sac, mt – metraterm. Scale-bars = 2, 1,000 μm, 3, 5 = 200 μm, 4 = 500 μm

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Table III. Measurements and ratios of Hypoporus plataxi sp. nov. Host n

Platax teira 1

Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% External seminal vesicle reach into hindbody%** Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

3306 530 801 15 339 311 0 274 166 0 221 0 231 240 430 191 549 608 244 234 19 320 271 0 423 284 630 64–77 62–77 (69) 28–36 (32) 16.0 24.2 0.68 0.77 1.88 18.4 90.2 19.0 0 6.67 0 0.58 0 13.0 1033 31.2 9.67 12.8

lometopon sp.), Labridae, California sheephead and Bodianus diplotaenia (Gill, 1862) (Labridae), Mexican hogfish off Cerros Island, Mexico (type locality) and Socorro Island, Mexico and the hogfish Lachnolaimus maximus (Walbaum, 1792) (Labridae) off Tortugas, Florida (Manter 1940). Manter stated that the ‘specimens from Bodianus were dead when collected and partly macerated’. Montgomery (1957) and Sogandares-Bernal and Hutton (1959) reported, but did not describe, this species from S. pulcher off La Jolla, California. Nahhas and Cable (1964) reported, but did not describe, this species from the Spanish hogfish Bodianus rufus (Linnaeus, 1758) (Labridae) off Jamaica. Fischthal (1977) reported, but did not describe, this species from L. maximus off Drowned Cays, Belize. Fernandes and Kohn (1984) briefly described and illustrated this species from the planehead filefish Stephanolepis hispidus (Linnaeus, 1766) (Monacanthidae) off Angra dos Reis, Rio de Janeiro State, Brazil. Based mainly on the specimens from S. pulcher and L. maximus, Manter (1940) described the cirrus-sac as ‘containing a short muscular tube near its base, a wider pars prostatica in its middle half and a cirrus (straight or slightly sinuous)’ which is similar to that of Intusatrium. The illustration (Manter 1940, fig. 20) shows a coiled internal seminal vesicle with sperm. Further observations may find this species to belong in Intusatrium. Lepocreadium bodiani Machida, 2004 in Bodianus bilunulatus (Lacepède, 1801), Labridae, tarry hogfish off Koniya, Kagoshima Prefecture, Japan (Machida 2004) has an internal seminal vesicle described as ‘oval to elongate-oval’, the pars prostatica is ‘globular’ and the ejaculatory duct is ‘short and narrow’. This may also be a species of Intusatrium, but the vitellarium reaches into the forebody in contrast to the condition in I. robustum and L. bimarinum. Genus Lepidapedoides Yamaguti, 1970 Lepidapedoides angustus Bray, Cribb et Barker, 1996 Host: Epinephelus chlorostigma (Valenciennes, 1828), Serranidae, brownspotted grouper; Epinephelus fasciatus (Forsskål, 1775), Serranidae, blacktip grouper; Epinephelus maculatus (Bloch, 1790), Serranidae, highfin grouper; Epinephelus retouti Bleeker, 1868, Serranidae, red-tipped grouper. Site: Digestive tract, caecum, gallbladder. Locality: E. chlorostigma, Off Récif Toombo, deep-sea, 200– 300 m (22°34´187S, 166°26´292E, 01/12/2009); E. fasciatus, Récif Le Sournois (22°31´30˝S, 166°26´30˝E, 28/11/2005); Near Passe de Dumbéa (22°21´111S, 166°15´446E, 08/09/2009); E. maculatus, Off Ever Prosperity, external slope, depth 60–80 m (22°27´S, 166°21´E, 17/04/2007); Récif Kué, External slope (22°34´892S, 166°29´673E, 19/06/2007); E. retouti, Off Barrier Reef near Passe de Dumbéa (22°21´090S, 166°14´0000E, 03/11/2009). Prevalence: E. chlorostigma, 1 of 3 (33%); E. fasciatus, 3 of 23 (13%); E. maculatus, 2 of 26 (7.7%); E. retouti, 1 of 3 (33%).

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255

Figs 6–8. Intusatrium robustum Durio et Manter, 1968 in Bodianus perditio. 6. Ventral view, uterus in outline. 7. Male terminal genitalia. 8. Cirrus-sac. Scale-bars = 6, 1,000 μm, 7, 200 μm, 8, 100 μm

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Table IV. Measurements and ratios of Intusatrium robustum Host n min Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% External seminal vesicle reach into hindbody%** Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

1027 390 289 6 136 120 0 102 69 24 0 0 157 154 132 32 5 38 53 71 0 127 99 0 144 106 42 49 21 21.2 21.7 1.04 1.12 1.21 3.34 9.32 15.7 1.89 0 0 0 0 12.8 289 28.1 12.4 14.1

Bodianus perditio 16 max

Bodianus loxozonus 2 mean

2660 799 739 35 261 245 72 201 158 89 189 0 493 498 318 107 183 243 140 204 0 343 289 0 385 282 436 115 56 29 42.8 32.7 2.01 2.22 2.05 12.66 75.4 22.1 4.53 7.12 0 0 0 12.0 739 27.8 12.9 14.5

1699 500 484 17 185 177 25 146 112 51 68 0 266 274 204 62 43 108 95 146 0 213 220 0 247 216 311 69 53 24 30.2 28.5 1.39 1.52 1.61 6.10 31.8 18.5 2.99 3.72 0 0 0 12.0 484 28.5 12.6 14.6

3157 920 912 14 273 246 73 193 202 100 283 0 352 368 387 82 88 272 217 247 0 407 0 507 406 525 243 51 30 29.1 28.9 1.29 1.49 1.22 8.63 32.4 16.6 3.16 8.97 0 0 0 12.3 912 28.9 12.9 16.0

3178 842 841 14 245 267 43 167 198 107 287 0 340 368 292 112 17 265 244 228 0 427 396 0 479 387 638 79 54 28 26.5 26.5 1.39 1.38 1.34 8.34 6.45 20.1 3.35 9.02 0 0 0 9.2 841 26.5 13.4 15.1

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Lepocreadiidae (Digenea) from fishes of the New Caledonia

Table V. Measurements and ratios of Lepidapedoides angustus Host n Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% External seminal vesicle reach into hindbody%** Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

Epinephelus chlorostigma 2 1729 389 551 5 96 100 46 57 53 171 174 218 102 96 177 41 237 298 128 109 37 175 146 0 192 128 313 168 70 33 22.5 31.9 1.06 0.96 1.87 17.2 79.5 18.1 9.90 10.0 12.6 2.12 0 10.3 334 19.3 10.1 11.1

Epinephelus fasciatus 2 2001 455 623 0 102 112 52 69 55 226 174 306 110 105 190 40 205 320 137 125 58 177 156 0 184 159 412 241 69 34 22.8 31.1 1.07 0.94 2.05 16.0 64.1 20.6 11.3 8.71 15.3 2.88 0 9.50 318 15.9 8.85 9.19

1580 372 481 2 107 111 18 68 62 139 148 237 121 114 137 53 140 208 119 95 23 155 119 0 164 144 336 155 74 33 23.5 30.5 1.13 1.03 1.79 13.1 67.4 21.3 8.81 9.36 15.0 1.48 0.00 8.68 244 15.5 9.79 10.4

Epinephelus Epinephelus maculatus retouti 1 1 1932 429 615 0 120 125 43 66 63 178 208 295 112 107 179 51 211 321 126 122 75 143 156 0 209 199 388 240 66 37 22.2 31.8 0.94 0.85 1.97 16.6 65.7 20.1 9.23 10.8 15.3 3.90 0.00 9.29 320 16.6 7.43 10.8

1944 730 547 0 107 110 48 52 61 169 180 247 106 103 166 44 266 299 120 121 58 134 136 14 157 148 499 240 70 37 37.5 28.1 0.99 0.94 1.80 15.4 88.9 25.7 8.69 9.26 12.7 2.98 0.74 8.53 299 15.4 6.88 8.08

1791 460 584 0 122 130 19 70 71 207 166 287 113 113 172 49 168 248 100 93 77 126 113 12 150 106 409 272 67 35 25.7 32.6 0.92 0.87 1.84 13.8 67.7 22.8 11.5 9.28 16.0 4.30 0.69 9.63 297 16.6 7.04 8.38

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Specimens: E. chlorostigma, MNHN JNC3142B, BMNH 2012.5.25.5; E. fasciatus, MNHN JNC1666, JNC1667, JNC3040, BMNH 2006.8.23.6, 2012.5.25.6; E. maculatus, MNHN JNC2157E, JNC2187B; E. retouti, MNHN JNC3083. Previous New Caledonian records: 1. Rigby et al. (1997, erroneously as L. kerapu), 2. Bray and Justine (2006), 3. Justine et al. (2010a). Previously reported New Caledonian hosts: Epinephelus fasciatus (2, 3), E. merra (1), E. maculatus (3). Discussion: Justine et al. (2010a) included these records in their paper on the parasites of serranids in New Caledonian waters. These specimens all conform to the L. angustus morphology as described by Bray et al. (1996) apart from one specimen from the gall-bladder of E. maculatus (JNC 2178B) which is broader than the others, with a width 38% of the length, which is a similar width to that quoted for L. dollfusi (Durio et Manter, 1968) (Bray et al. 1996) (Table V). Other features such as the position of the genital pore at the anterior margin of the ventral sucker, the entire ovary and the distinct prepharynx indicate a similarity to L. angustus. Bray et al. (2009b) have presented molecular evidence that Lepidapedoides is a typical lepocreadiid, close to genera such as Prodistomum Linton, 1910 and Opechona Looss, 1907, and is not close to Lepidapedon Stafford, 1904. Lepidapedoides dollfusi (Durio et Manter, 1968) Bray, Cribb et Barker, 1996 Syn. Neolepidapedon dollfusi (Durio et Manter, 1968) Records from New Caledonia: Durio and Manter (1968). New Caledonian Hosts: Epinephelus spp. ‘red cod’ and ‘spotted grouper’. Lepidapedoides kalikali Yamaguti, 1970 (Fig. 9) Syn. Lepidapedon (Lepidapedoides) kalikali Yamaguti, 1970 Host: Pristipomoides auricilla (Jordan, Evermann et Tanaka, 1927), Lutjanidae, goldflag jobfish. Site: Digestive tract. Localities: Off Passe de Dumbéa, deep-sea (150–250 m depth, 22°20´995S, 166°13´773E, 19/02/2008); West of Passe de Dumbéa (22°19´S, 166°13´E, 02/04/2008). Prevalence: 2 of 2 (100%). Specimens: MNHN JNC2457-1, JNC2468-1, BMNH 2012.2.15.6. Previous New Caledonian records: None. Discussion: This species is known from its original description in P. [as Arnillo] auricilla and the lavender jobfish Pristipomoides sieboldii (Bleeker, 1854) off Hawaii (Yamaguti 1970) and reports from the crimson jobfish P. filamentosus (Valenciennes, 1830) off Chichijima Island, Japan and P. auricilla off Koror Island, Palau (Kuramochi 2011). Our worms differ slightly from the original description in being slightly narrower, with a shorter forebody, but in other ways resemble Yamaguti’s original description closely (Fig. 9, Table

Fig. 9. Lepidapedoides kalikali Yamaguti, 1970 in Pristipomoides auricilla. Ventral view, uterus in outline. Scale-bar = 500 μm

VI). In a cladistic study based on morphology (Bray et al. 1998, Cribb et al. 2002), L. kalikali was resolved as a member of a small clade of species mostly restricted to lutjanids (including caesionines – Miller and Cribb 2007). Genus Lepotrema Ozaki, 1932 Lepotrema cf. clavatum Ozaki, 1932 (Fig. 10) Host: Sufflamen fraenatum (Latreille, 1804), Balistidae, masked triggerfish.

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Table VI. Measurements and ratios of Lepidapedoides kalikali, distinctive characters emboldened Hosts Locality n

Pristipomoides auricilla New Caledonia 11 min max mean

Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% External seminal vesicle reach into hindbody%** Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

1402 518 449 0 104 126 0 70 90 100 98 165 125 98 127 92 61 50 129 138 0 193 165 0 217 179 306 68 58 27 32.3 29.7 0.91 0.77 1.21 3.15 75.5 20.0 6.59 6.81 11.5 0 0 8.25 229 15.1 12.7 15.4

1741 703 553 12 148 174 30 112 114 167 160 262 178 186 199 125 109 163 196 0 307 271 0 349 289 435 278 71 38 43.5 34.5 1.48 1.29 1.66 6.80 297 26.9 11.0 11.4 16.9 0 0 13.7 316 20.3 19.2 23.0

1561 593 492 3 136 145 9 95 103 129 134 223 153 145 158 110 111 77 145 162 0 265 224 0 306 229 373 155 62 34 38.0 31.6 1.14 1.00 1.41 4.93 153 23.9 8.23 8.62 14.3 0 0 10.1 270 17.3 17.0 19.6

P. sieboldi & P. auricilla Hawaii after Yamaguti (1970) 17 min max 1300 600

1850 900

80 100

150 180

60 60 100

100 100 150

100 100 120 30

170 170 230 70

70 70

220 180

80 80

320 250

80 80

320 250

48 28 46.2 38.6 0.80 0.80 1.67 2.79 100 23.3 7.69 15.8 21.4 0 0 9.23

60 35 48.6

16.7 6.15 6.15

0.82 0.82 1.80

8.11

12.4

17.3 17.3

260

Fig. 10. Lepotrema cf. clavatum Ozaki, 1932 in Sufflamen fraenatum. Ventral view, uterus in outline. Scale-bar = 500 μm

Site: Digestive tract. Localities: Inside Lagoon, facing Récif Toombo (22°32´361S, 166°26´992E, 06/11/2007); Shallow, Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 20/11/2008); Shallow, Interior Lagoon near Récif Toombo (22°32´606S, 166°28´991E, 25/11/2008). Prevalence: 3 of 11 (27%). Specimens: MNHN JNC2372, JNC2763, JNC2772B, BMNH 2012.5.25.18. Previous New Caledonian records: None. Description: Based on 6 whole-mounted specimens, measurements in Table VII. Body pyriform, widest in hindbody (Fig. 10). Tegument finely spines, spines reaching close to posterior extremity. Eye-spot pigment scattered around pharynx/oral sucker region in some, but not all, specimens. Oral

Rodney A. Bray and Jean-Lou Justine

sucker large, subglobular, subterminal. Ventral sucker oval, of similar size to oral sucker, just pre-equatorial. Prepharynx short, in posterior cavity of oral sucker, wall thick. Pharynx large, oval. Oesophagus short, narrow. Intestinal bifurcation in posterior forebody. Caeca broad, reach to about middle of post-testicular region. Testes two, oval, entire, symmetrical or slightly oblique, in mid hindbody. External seminal vesicle large, but usually obscured by eggs. Cirrus-sac claviform, mainly dorsal to ventral sucker. Internal seminal vesicle oval. Pars prostatica vesicular. Ejaculatory duct long, thick-walled. Genital atrium distinct. Genital pore sinistral, ventral to sinistral caecum at the level of the anterior margin of the ventral sucker. Ovary trilobate, immediately pretesticular, close to ventral sucker. Laurer’s canal not detected. Seminal receptacle dorsal or dorso-lateral to ovary. Mehlis’ gland dorsal to ovary. Uterus intercaecal, pretesticular, passes ventrally to ovary, overlaps posterior edge of ventral sucker. Eggs tanned operculate. Metraterm shorter than cirrus-sac, distal extremity with large folded muscular pad. Vitellarium follicular, reaching from pharynx to posterior extremity, confluent in forebody and post-testicular region; lateral, ventral and dorsal to caeca. Excretory pore dorsal, in anterior post-testicular region; vesicle reaches to testes, not traced further. Discussion: This form is similar to the type species Lepotrema clavatum Ozaki, 1932, keying to that species in the key presented by Bray and Cribb (1996b), but it differs from L. clavatum in being broader, in the testes being symmetrical or nearly so, in the shorter cirrus-sac and slightly more anteriorly extensive vitellarium. Subsequent to Bray and Cribb (1996b) one further species has been described, Lepotrema monile Bray et Cribb, 1998 from Ward’s damsel Pomacentrus wardi Whitley, 1927 (Pomacentridae) off Heron Island in the southern Great Barrier Reef (Bray and Cribb 1998). It differs from other species of Lepotrema, including our specimens, in lacking the large folded muscular metraterm pad, which is replaced by a relatively weakly developed sphincter and in the ovary showing only the slightest indications of trilobation. Bray and Cribb (1996b) listed two further species which they thought could possibly belong in Lepotrema, namely Preptetos cylindricus Wang, 1989 and Lepocreadium navodoni Shen, 1986, both from off China. P. cylindricus from the monacanthids Thamnaconus [as Navodon] septentrionalis (Günther, 1874) and the fan-bellied leatherjacket Monacanthus chinensis (Osbeck, 1765) has a subglobular ovary and oblique testes (Wang 1989). L. navodoni from Thamnaconus [as Navodon] modestus (Günther, 1877) has oblique testes but a distinctly trilobate ovary (Shen 1986). Machida and Kuramochi (1999) reported L. clavatum from T. modestus off Japan, and mentioned that their specimens were large (2,100–2,500 long). The published records of Lepotrema or putative Lepotrema species indicate that they are found equally in Balistidae, Monacanthidae and Pomacentridae (each with 31% of records each), with occasional single reports in Bothidae, Chaetodontidae and Pomacanthidae.

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Lepocreadiidae (Digenea) from fishes of the New Caledonia

Table VII. Measurements and ratios of Lepotrema? clavatum Host n Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

Sufflamen fraenatum 6 min max mean 722 460 307 17 100 111 0 76 75 7 54 167 109 98 97 30 0

1326 681 483 39 214 240 17 176 172 35 103 236 264 240 268 83 12

1069 552 399 26 165 180 3 133 122 23 80 204 201 172 195 58 3

0 97 82 0 171 99 0 133 94 129 50 44 27 43.2 32.7 1.05 0.86 1.36 0 15.9 0.85 4.52 14.0 0 0 13.5 107 14.9 14.4 15.7

7 141 192 0 255 167 0 251 185 237 137 62 33 71.0 42.6 1.31 1.06 1.73 1 20.2 3.13 10.73 27.7 0 0 20.8 247 19.6 23.7 19.9

1 122 142 0 194 130 0 193 130 185 96 57 31 53.1 37.8 1.19 0.94 1.48 0 17.5 2.18 7.82 19.9 0 0 17.7 194 17.9 18.7 18.1

Genus Lobatocreadium Madhavi, 1972 Lobatocreadium exiguum (Manter, 1963) Madhavi, 1972 (syn. Lepocreadium exiguum Manter, 1963; Pseudocreadium exiguum (Manter, 1963) Yamaguti, 1970; Pseudocreadium hainanensis Shen, 1990) Hosts: Abalistes filamentosus Matsuura et Yoshino, 2004, Balistidae; Pseudobalistes fuscus (Bloch et Schneider, 1801), Balistidae, yellow-spotted triggerfish; Sufflamen fraenatum (Latreille, 1804), Balistidae, masked triggerfish. Sites: Intestine, digestive tract. Localities: A. filamentosus, Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 21/04/2009); P. fuscus, Récif Toombo (22°26´10S, 166°33´00˝E, 13/12/2005); Ilôt Signal, off Nouméa (22°17´S, 166°17´E, 30/05/2006); Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 20/05/2009); Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 30/04/2009); S. fraenatum, Shallow, Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 20/11/2008); Shallow, Interior Lagoon near Récif Toombo (22°32´606S, 166°28´991E, 25/11/2008); Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 01/09/2009); Inside Lagoon, facing Récif Toombo (22°32´361S, 166°26´992E, 06/11/2007); near Ile Redika (22°32´S, 166°37´E, 11/06/2009). Prevalences: A. filamentosus, 1 of 2 (50%); P. fuscus, 4 of 5 (80%); S. fraenatum, 7 of 11 (64%). Specimens: A. filamentosus, JNC2913; P. fuscus, MNHN JNC1680, 1844, JNC2940, JNC2927, BMNH 2006.8.23. 7–10; S. fraenatum, JNC2761, JNC2762, JNC2763, JNC2771A, JNC2771C, JNC3034, JNC2374, JNC2978, BMNH 2012.5.25.7-11. Previous New Caledonian records: 1. Bray and Justine (2006), 2. Bray et al. (2009b). Previously reported New Caledonian host: Pseudobalistes fuscus (1, 2). Discussion: These specimens fit comfortably into the descriptions of L. exiguum (see Bray and Cribb 1996a, 2002; Manter 1963; Shen 1990; Yamaguti 1970) (Table VIII). Abalistes filamentosus is a newly reported host, although L. exiguum has been reported in the starry triggerfish Abalistes stellatus (Anonymous, 1798) from the Swain Reefs, Great Barrier Reef (Bray and Cribb 2002). Bray and Cribb (1996a) considered Pseudocreadium hainanensis as synonym of L. exiguum, and it should be noted that Shen (1990) described it from ‘Balistes capistratus’, off Hainan Island, southern China. This host is, according to Froese and Pauly (2012), a synonym of Sufflamen fraenatum. Lobatocreadium exiguum was originally described from an unidentified trigger fish (Balistidae) off Fiji (Manter 1963) and all subsequent records have been from members of this family throughout the Indo-West Pacific Region, reaching as far east as Moorea in French Polynesia and Hawaii (Bray and Cribb 1996a, 2002) and as far west as the Maldives in the Indian Ocean (Lorber et al. 2006).

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Table VIII. Measurements and ratos of Lobatocreadium exiguum Host n Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

Abalistes Pseudobalistes filamentosus fuscus 1 1 771 580 341 52 145 188 0 131 160 0 118 254 182 174 292 108 0 149 277 0 142 174 0 138 168 69 54 66 32 75.3 44.3 1.26 0.93 1.17 0 9.0 0 15.29 32.9 0 0 37.8 87 11.3 18.5 17.9

695 472 324 24 143 167 0 122 122 0 139 229 173 165 216 102 0 93 200 0 129 180 0 136 141 60 52 59 23 68.0 46.6 1.21 0.99 1.37 0 8.7 0 20.05 32.9 0 0 31.1 95 13.7 18.6 19.6

Sufflamen fraenatum

min

8 max

mean

641 448 283 14 131 145 0 103 109 0 66 215 137 135 166 90 0 140 126 0 91 114 0 105 99 31 63 56 29 68.9 42.8 0.99 0.84 1.13 0 3.3 0 9.96 31.3 0 0 25.8 48 7.2 14.1 15.2

917 695 443 69 189 241 0 163 213 0 150 308 189 203 272 130 0 204 268 0 161 194 0 158 157 104 161 70 44 75.8 48.3 1.16 1.06 1.33 0 12.1 0 20.25 36.5 0 0 35.8 152 16.6 17.8 21.4

743 531 340 47 150 178 0 124 148 0 110 252 156 168 220 108 0 163 218 0 118 148 0 133 128 55 82 64 34 71.3 45.6 1.04 0.95 1.21 0 7.4 0 14.79 33.9 0 0 29.7 88 11.7 15.8 18.0

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Lepocreadiidae (Digenea) from fishes of the New Caledonia

Genus Multitestis Manter, 1931 Multitestis magnacetabulum Mamaev, 1970 and Multitestis pyriformis Manter, 1963 New Caledonian record: Bray et al. (2010b). New Caledonian host: Platax teira. Genus Neohypocreadium Machida et Uchida, 1987 Neohypocreadium dorsoporum Machida et Uchida, 1987 Hosts: Chaetodon auriga Forsskål, 1775, Chaetodontidae, threadfin butterflyfish; Chaetodon flavirostris Günther, 1874, Chaetodontidae, black butterflyfish; Chaetodon lineolatus Cuvier, 1831, Chaetodontidae, lined butterflyfish; Chaetodon melannotus Bloch et Schneider, 1801, Chaetodontidae, blackback butterflyfish; Chaetodon ulietensis Cuvier, 1831, Chaetodontidae, Pacific double-saddle butterflyfish. Site: Intestine. Locality: Lagoon (November 1997). Prevalence: C. auriga, 14 of 20 (70%); C. flavirostris, 24 of 34 (71%); C. lineolatus, 6 of 11 (55%); C. melannotus, 1/23 (4%); C. ulietensis, 4/19 (21%). Previous New Caledonian records: none. Discussion: A study of the genus Neohypocreadium from the Indo-Pacific region is at present under way. Neohypocreadium longisaccatum Machida et Uchida, 1987 Host: Chaetodon flavirostris Günther, 1874, Chaetodontidae, black butterflyfish. Site: Intestine. Locality: Lagoon (November 1997). Prevalence: 1/34 (3%). Previous New Caledonian records: None. Discussion: A study of the genus Neohypocreadium from the Indo-Pacific region is at present under way. Genus Neomultitestis Machida, 1982 Neomultitestis aspidogastriformis Bray et Cribb, 2003 Host: Platax teira (Forsskål, 1775), Ephippidae, longfin batfish. Site: intestine, stomach. Locality: Nouméa Fish Market, allegedly from “off Bouraké” (11/03/2009). Prevalence: 1 of 2 (50%). Specimens: MNHN JNC2872B, BMNH 2012.5.25.4. Previous New Caledonian records: None. Discussion: This species is unmistakeable due to its highly distinctive ventral sucker divided into three loculi (Table IX). Its

Table IX. Measurements and ratios of Neomultitestis aspidogastriformis Host n

Platax teira 6 min max mean

Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width Ovary length Ovary width Testicular field length Testicular field width Post-testicular distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Testicular field length%

718 786 283 0 131 147 0 95 108 0 64 128 201 429 311 98 89 123 245 561 195 52 24 110 39.4 1.38 2.91 1.24 23.6 0 8.85 17.9 43.3 111 12.5 26.0

1044 1204 452 14 203 222 0 134 162 37 185 284 292 685 519 212 154 189 392 1037 364 58 29 127 44.5 1.54 3.29 1.60 36.4 4.28 20.8 31.8 54.6 208 21.9 45.5

900 1063 383 9 175 196 0 124 140 14 123 224 253 598 443 162 126 152 313 772 278 55 26 118 42.4 1.45 3.06 1.39 30.6 1.56 13.5 24.6 49.0 159 17.7 35.0

only previous report is from the same host species off Heron Island, southern Great Barrier Reef (Bray and Cribb 2003). Genus Opechona Looss, 1907 Opechona bacillaris (Molin, 1859) Dollfus, 1927 (Fig. 11) Syns. see Bray and Gibson (1990) Host: Rastrelliger kanagurta (Cuvier, 1816), Scombridae, Indian mackerel. Site: Digestive tract. Locality: Nouméa Fish Market (20/06/2007, 16/12/2008).

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Prevalence: 4 of 15 (27%) Specimens: MNHN JNC2192, JNC2194, JNC2196, JNC2855, BMNH 2012.5.25.16-17. Previous New Caledonian records: None. Discussion: The Opechona species mostly reported from Rastrelliger kanagurta (and its synonyms) is O. bacillaris. First re-

Rodney A. Bray and Jean-Lou Justine

ported by Madhavi (1972) from the Bay of Bengal, it has subsequently been reported from off NW Australia (Korotaeva 1974), Saya-de Malya Bank, Indian Ocean (Parukhin 1985, 1988) and again from the Bay of Bengal (Madhavi et al. 1986, Madhavi and Triveni Lakshmi 2011). It may well be that the Indo-Pacific form is distinct from the commonly reported northern Atlantic form, which has been reported numerous times in the Atlantic mackerel Scomber scombrus Linnaeus, 1758 and chub mackerel Scomber japonicus Houttuyn, 1782 in addition to many other host-species (Bray and Gibson 1990). Only three of our 21 undamaged specimens were extended so that useful measurements could be taken (Fig. 8, Table X). Fourteen were immature and in the remaining four the oral sucker was deeply retracted into the forebody, a common feature of the Atlantic form. Bray and Gibson (1990) considered Lepocreadium guptai Gupta et Gupta, 1987, reported from R. kanagurta (as Scomber microlepidotus) from the Bay of Bengal (Gupta and Gupta 1987), as a synonym of O. bacillaris. Available descriptions and illustrations of Opechona from this host (Gupta and Gupta 1987, Madhavi and Triveni Lakshmi 2011) show that L. guptai differs distinctly from O. bacillaris of Madhavi and Triveni Lakshmi (2011) and our specimens in sucker ratio (ventral sucker about 150–178% of oral sucker width, vs 70 or 78–88%). The other Opechona species reported from R. kanagurta is O. siddiqi Ahmad, 1984 (nec 1986) from the Arabian Sea (Ahmad 1984). The caecal ending are described as blind and its status in the genus is, therefore, equivocal, and it is also differentiated from O. bacillaris by its huge eggs (100–120 long) and the vitelline field reaches into the forebody. Genus Opisthogonoporus Yamaguti, 1937 Opisthogonoporus amadai Yamaguti, 1937 (Fig. 12)

Fig. 11. Opechona bacillaris (Molin, 1859) in Rastrelliger kanagurta. Ventral view. Scale-bar = 500 μm

Host: Branchiostegus wardi Whitley, 1932, Malacanthidae, Ward’s tilefish. Site: Digestive tract. Locality: Off Récif Kué, deep-sea (150–300 m depth, 22°35´707S, 166°30´387E, 12/02/2008). Prevalence: 1 of 2 (50%). Specimens: MNHN JNC2454. Previous New Caledonian records: None. Discussion: One specimen is available (Table X). Although the condition is not ideal, the general characteristics of the worm are very similar to the specimens described by Yamaguti (1937) from the horsehead tilefish Branchiostegus japonicus (Houttuyn, 1782) from the Sea of Japan. A few dimensions differ slightly, e.g. the eggs are generally slightly larger in our specimen (33–38 × 11–15 vs 26–33 × 15–18), and details of the cirrus-sac contents are not visible, but we have little doubt that our specimen is conspecific with the Japanese form. Machida et al. (1970) later reported this species from B. japonicus from north of the Tsushima Islands, Japan.

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Lepocreadiidae (Digenea) from fishes of the New Caledonia

Table X. Measurements and ratios of lepocreadiids Species

Host

Opechona bacillaris

Rastrelliger kanagurta

n Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% External seminal vesicle reach into hindbody%** Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance

min

3 max

mean

1149 277 347 0 130 112 0 104 93 20 52 41 101 98 166 52 154 206 53 67 6 104 122 0 120 120 195 37 76 37 20.5 30.2 0.64 0.78 1.20 17.9 64.8 14.0 1.78 3.88 3.57 0 0 14.5 306

315 449 14 159 127 74 120 98 69 67 86 109 99 233 70 205 289 76 77 47 114 130 0 137 133 240 52 84 41 25.3 32.3 0.82 0.88 1.29 21.3 74.8 17.8 5.11 5.00 6.36 4 0 16.7 394

1299 294 411 5 147 121 47 111 97 46 59 61 105 99 203 62 182 261 67 73 31 109 127 0 127 126 210 49 81 38 22.8 31.6 0.72 0.82 1.25 20.0 70.3 16.2 3.43 4.57 4.62 3 0 15.5 350

Opisthogono- Prodistomum Pseudopisthoporus amadai keyam gonoporus vitellosus Branchiostegus Monodactylus Naso wardi argenteus brevirostris 1 1 1 1326 251 304 0 67 91 3 58 71 181 0 196 110 117 341 120 232 315 109 125 0 159 129 0 231 140 113 46 33–38 (35) 11–15 (14) 19.0 23.0 1.63 1.28 1.28 23.8 73.7 8.54 13.6 0 14.8 0 0 25.7 108

831 284 401 0 48 74 81 55 52 24 188 154 78 79 ? ? ? 48 68 79 0 92 87 0 131 111 62 34 62 27 34.2 48.2 1.63 1.06 1.44 5.79 ? 7.49 3 22.7 19 0 0 ? 246

2248 374 950 11 140 151 88 104 95 85 539 511 393 220 384 99 0 203 155 129 0 181 213 0 198 177 141 49 64 34 16.6 42.2 2.80 1.46 1.58 9.02 0 6.28 3.77 24.0 22.7 0 0 17.1 439

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Table X. Continuation Species

Host

Opechona bacillaris

Rastrelliger kanagurta

n Pre-vitelline distance% Anterior testis length% Posterior testis length%

min

3 max

mean

25.9 7.71 9.13

28.2 9.89 10.5

26.9 8.50 9.81

Genus Postlepidapedon Zdzitowiecki, 1993 Postlepidapedon secundum (Durio et Manter, 1968) Bray, Cribb et Barker, 1997 Syn. Intusatrium secundum Durio et Manter, 1968 New Caledonian records: 1. Durio and Manter (1968), 2. Bray et al. (1997), 3. Bray and Justine (2006). New Caledonian hosts: ‘brown-blotched parrot fish’ (1, possibly C. graphicus – 2), Choerodon graphicus (De Vis, 1885) (Labridae) (2, 3). Preptetos Pritchard, 1960 Preptetos cannoni Barker, Bray et Cribb, 1993 Hosts: Siganus argenteus (Quoy et Gaimard, 1825), Siganidae, streamlined spinefoot; Siganus canaliculatus (Park, 1797), Siganidae, white-spotted spinefoot; Siganus doliatus GuérinMéneville, 1829–1838, Siganidae, barred spinefoot; Siganus lineatus (Valenciennes, 1835), Siganidae, golden-lined spinefoot; Siganus spinus (Linnaeus, 1758), Siganidae, little spinefoot. Site: Intestine. Localities: S. argenteus, New Caledonia (21°30´S, 165°30´E, Nov., 1997); S. canaliculatus, New Caledonia (21°30´S, 165°30´E, Nov., 1997); S. doliatus New Caledonia (21°30´S, 165°30´E, Nov., 1997), Récif de Crouy, New Caledonia (22°20´957S, 166°21´375E, 20/05/2008); S. lineatus, Nouméa Fish Market, New Caledonia (25/07/ 2008); S. spinus, New Caledonia (21°30´S, 165°30´E, Nov., 1997). Prevalence: S. argenteus, 1 of 1; S. canaliculatus, 1 of 3 (33%); S. doliatus, 3 of 10 (30%); S. lineatus, 1 of 7 (14%); S. spinus, 3 of 3 (100%). Previous New Caledonian records: None. Discussion: A study of the genus Preptetos from the IndoPacific region is at present under way. Preptetos sp. Host: Acanthurus xanthopterus Valenciennes, 1835, Acanthuridae, yellowfin surgeonfish.

Opisthogono- Prodistomum Pseudopisthoporus amadai keyam gonoporus vitellosus Branchiostegus Monodactylus Naso wardi argenteus brevirostris 1 1 1 46.1 12.0 17.4

29.7 11.0 15.8

19.5 8.05 8.81

Site: Intestine. Localities: Nouméa Fish Market, New Caledonia (11/03/2009). Discussion: A study of the genus Preptetos from the IndoPacific region is at present under way. Genus Prodistomum Linton, 1910 Prodistomum keyam Bray et Cribb, 1996 Host: Monodactylus argenteus (Linnaeus, 1758), Monodactylidae, silver moony. Site: Digestive tract. Locality: Nouméa Fish Market (23/07/2010; 15/06/2007). Prevalence: 3 of 3 (100%). Specimens: MNHN JNC3215. Previous New Caledonian records: None. Discussion: These specimens, one of which was in good enough condition for measurement (see Table X), appear to be practically identical to the description by Bray and Cribb (1996c) from the same host from Moreton Bay, Queensland, Australia. Slight, but insignificant, differences occur between the range of measurements of 9 type-specimens and our measured specimen include width, prepharynx and pharynx dimensions and post-caecal length. Genus Pseudopisthogonoporus Yamaguti, 1970 Pseudopisthogonoporus vitellosus (Pritchard, 1963) Yamaguti, 1970 (syn. Opisthogonoporus vitellosus Pritchard, 1963) Hosts: Naso brevirostris (Cuvier, 1829), Acanthuridae, spotted unicornfish; Naso annulatus (Quoy et Gaimard, 1825), Acanthuridae, white-margin unicornfish. Site: Digestive tract. Locality: N. brevirostris, Récif de Crouy (22°21´825S, 166°21´961E, 31/10/2008); Naso annulatus Nouméa Fish Market (22/04/2011). Prevalence: 1 of 1 (100%). Specimens: N. brevirostris, MNHN JNC2732; Naso annulatus MNHN JNC3376C, BMNH 2012.5.25.3.

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Previous New Caledonian records: None. Discussion: Our specimens are indistinguishable from those described by Pritchard (1963), Yamaguti (1970) and Bray & Cribb (1998), and those whose morphology was commented on by Machida and Uchida (1990) and Toman (1989) (Table X). The species has been reported almost exclusively from Naso spp., including records from N. brevirostris by each of the authors listed above, from Hawaii, The Great Barrier Reef, Japan and the Seychelles. Genus Stegodexamene Macfarlane, 1951 Stegodexamene anguillae Macfarlane, 1951 New Caledonian record: Moravec and Justine (2007). New Caledonian hosts: Anguilla obscura Günther, 1872, A. reinhardtii Steindachner, 1867 (Anguillidae). Subfamily Lepidapedinae Yamaguti, 1958 Genus Holorchis Stossich, 1901 Holorchis castex Bray et Justine, 2007 Host: Diagramma pictum (Thunberg, 1792), Haemulidae, painted sweetlips. Site: Digestive tract. Locality: Ilôt Lebris, off Ouano (21°50´108S, 166°45´155E, 25/10/2007), Nouméa Fish Market (16/09/2010). Prevalence: 3 of 9 (33%). Specimens: MNHN JNC2342, JNC3253A, B. Previous New Caledonian record: Bray and Justine (2007). Previously reported New Caledonian hosts: Diagramma pictum. Discussion: We include new data as one new specimen is much larger (Table XI) than any of the 5 specimens previously described (Bray and Justine 2007). Holorchis plectorhynchi Durio et Manter, 1968 (Fig. 13)

Fig. 12. Opisthogonoporus amadai Yamaguti, 1937 in Branchiostegus wardi. Ventral view, uterus in outline. Scale-bar = 500 μm

Host: Diagramma pictum (Thunberg, 1792), Haemulidae, painted sweetlips. Site: digestive tract. Locality: Between Larégnière et Récif Crouy (22°20´702S, 166°19´295E, 05/05/2008); Interior Lagoon near Récif Toombo (22°32´536S, 166°29´069E, 25/08/2009). Prevalence: 3 of 8 (37.5%). Specimens: MNHN JNC2511, JNC2510, JNC3023, BMNH 2012.5.25.20. Previous New Caledonian record: Durio and Manter (1968). Previously reported New Caledonian hosts: Plectorhinchus lineatus (Linnaeus, 1758) (Haemulidae), Lethrinus miniatus (Forster, 1801) (Lethrinidae). Discussion: Durio and Manter (1968) described H. plectorhynchi from the yellowbanded sweetlips Plectorhinchus

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Table XI. Measurements and ratios of Holorchis spp. Species Host n Length Width Forebody Pre-oral lobe Oral sucker length Oral sucker width Prepharynx Pharynx length Pharynx width Oesophagus Intestinal bifurcation to ventral sucker Vitellarium to ventral sucker Ventral sucker length Ventral sucker width Cirrus-sac length Cirrus-sac width External seminal vesicle reach into hindbody Ventral sucker to ovary Ovary length Ovary width Ovary to anterior testis Anterior testis length Anterior testis width Distance between testes Posterior testis length Posterior testis width Post-testicular distance Post-caecal distance Egg length Egg width Width% Forebody% Sucker length ratio 1: Sucker width ratio 1: Oral sucker: pharynx width 1: Ventral sucker to ovary% External seminal vesicle reach into hindbody%** Post-testicular distance% Oesophagus% Intestinal bifurcation to ventral sucker% Vitellarium to ventral sucker% Ovary to anterior testis% Distance between testes% Cirrus-sac length% Pre-vitelline distance Pre-vitelline distance% Anterior testis length% Posterior testis length%

Holorchis castex Diagramma pictum 1 13 333 2314 3377 130 653 691 0 386 351 176 2113 840 579 560 746 393 0 163 571 376 1536 735 633 74 938 709 594 80 34 17.4 25.3 0.89 0.81 1.96 1.22 0 41.2 1.32 15.9 6.30 11.5 0.55 5.60 2537 19.0 5.51 7.03

min

Holorchis plectorhynchi Diagramma pictum 3 max

mean

3073 521 940 42 199 251 0 123 132 71 525 0 190 198 150 74 29 38 153 146 577 254 215 67 270 224 539 68 63 28 12.0 21.3 0.95 0.71 1.82 1.08 76.4 17.6 1.74 11.8 0 18.8 2.19 4.19 940 21.3 7.23 8.39

6663 802 1421 52 270 306 0 155 168 150 789 0 326 322 317 103 196 208 344 264 1683 482 395 161 628 340 1384 112 67 39 17.0 30.6 1.21 1.07 1.91 3.12 210 20.8 3.56 17.1 0 25.3 3.85 4.87 1421 30.6 8.58 9.43

4605 639 1140 46 238 286 0 139 155 110 645 0 250 246 213 88 106 96 248 198 1017 362 285 128 413 269 914 89 65 32 14.5 26.0 1.04 0.86 1.85 1.81 127 19.5 2.52 14.7 0 21.1 2.82 4.61 1140 26.0 8.03 8.86

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Lepocreadiidae (Digenea) from fishes of the New Caledonia

lineatus (Linnaeus, 1758) (as Plectorhinchus goldmani (Bleeker, 1853) [Haemulidae] and the trumpet emperor Lethrinus miniatus (Forster, 1801) [Lethrinidae]. We recovered 3 complete specimens and some fragments in D. pictum in 3 fishes from 2 localities. One of our specimens is larger than any reported originally, but otherwise our material is similar to the original description (Table XI). We found no lepocreadiids in L. miniatus in spite of significant sampling (27 fish) (Justine et al. 2010b).

Zoogeography

Fig. 13. Holorchis plectorhynchi Durio et Manter, 1968 in Diagramma pictum. Ventral view, uterus in outline. Scale-bar = 1,000 μm

Thirty-two lepocreadiid species are now reported from New Caledonian waters, although it is likely that many more are awaiting discovery. Of these 6 (19%) are possibly endemic: Diploproctodaeum triodoni, Hypocreadium toombo, Hypoporus plataxi, Intusatrium robustum, Holorchis castex, Holorchis plectorhynchi. Shared only with Heron Island: 5 (16%): Lepidapedoides angustus, Lepidapedoides dollfusi, Neomultitestis aspidogastriformis, Postlepidapedon secundum, Preptetos cannoni. Shared only with New Zealand: 1 freshwater: Stegodexamene anguillae. Shared only with West Pacific (as far east as Hawaii and French Polynesia, including Heron Island): 11 (34%): Bulbocirrus aulostomi, Opisthogonoporus amadai, Diploproctodaeum rutellum, Lepidapedoides kalikali, Lepotrema clavatum, Multitestis magnacetabulum. Shared with Indo-West Pacific Region total, including West Pacific: 23 (72%): Bianium arabicum, Diploproctodaeoides longipygum, Diploproctodaeum arothroni, Diploproctodaeum macracetabulum, Hypocreadium patellare, Lobatocreadium exiguum, Multitestis pyriformis, Neohypocreadium dorsoporum, Neohypocreadium longisaccatum, Prodistomum keyam, Pseudopisthogonoporus vitellosus. Indo-West Pacific species, which apparently has passed into Mediterranean Sea (Lessepsian migration): 1: Echeneidocoelium indicum. Circum-tropical species: 1: Diploproctodaeum haustrum. Cosmopolitan species: 1: Opechona bacillaris. The fish-digenean fauna of New Caledonia is of a typical Indo-West Pacific type, with the majority of its fauna shared only with parts of this region (72%). The next largest component of the fauna is the endemic species (19%). The southern Great Barrier Reef (Heron Island, Swain Reefs) shares the most species (16, i.e. 50%), and the Great Barrier Reef as a whole shares 19 (59%) species. The remainder are individual species whose distributions vary. One is found in eels in freshwater and is shared just with New Zealand. One species is essentially Indo-West Pacific, but has apparently invaded the Mediterranean Sea via the Suez Canal. One species is found not only in the tropical Indo-Pacific, but also in the tropical Atlantic. One species is cosmopolitan, and is the only species reported in both New Caledonia and the eastern Pacific.

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These data should be considered provisional as there are considerable biases in sampling in all oceans, particularly the Pacific Ocean. Some localities, e.g. Japan, the Great Barrier Reef and, now, New Caledonia, are fairly well sampled, whereas other regions are poorly sampled or unexplored. Similar caveats may be advanced in consideration of the techniques used to identify these species. Hitherto, these have mostly been morphological, and molecular study alerts us to the fact that many cryptic species occur and that fragmented sampling along clines may indicate spurious specific distinctions. Host-specificity Our findings indicate that lepocreadiids in this region are mainly parasites of perciforms (72% of records) and tetraodontiforms (22%), with occasional reports in anguilliforms (4%) and gasterosteiforms (2%). Within the major host orders, the parasites are spread widely among families, but it is clear that sampling bias has affected these results to some extent. The tetradontiform family Balistidae has the most reports (14%), closely followed by the perciformes Chaetodontidae and Serranidae (both 12%), Siganidae (10%) and Ephippidae and Haemulidae (both 8%). Considering those species reported more than once, the following types of specificity are exhibited (see Miller et al. 2011). Oioxenous: Bulbocirrus aulostomi, Diploproctodaeoides longipygum, Hypocreadium toombo, Neomultitestis aspidogastriformis, Postlepidapedon secundum, Prodistomum keyam, Holorchis castex. Stenoxenous (to host genus): Bianium arabicum, Diploproctodaeum arothroni, Intusatrium robustum, Lepidapedoides dollfusi, Lepidapedoides kalikali, Multitestis pyriformis, Opisthogonoporus amadai, Stegodexamene anguillae. Stenoxenous (to host family): Diploproctodaeum haustrum, Diploproctodaeum rutellum, Echeneidocoelium indicum, Lepidapedoides angustus, Lobatocreadium exiguum, Pseudopisthogonoporus vitellosus. Stenoxenous (to host order): Diploproctodaeum macracetabulum, Preptetos cannoni, Holorchis plectorhynchi, Multitestis magnacetabulum. Euryxenous: Hypocreadium patellare, Lepotrema clavatum, Opechona bacillaris. Miller et al. (2011), having surveyed the patterns of hostspecificity of digeneans of Great Barrier Reef fishes stated that ‘no host euryxenous host distribution should be accepted on the basis of morphology alone’ and argued that true euryxenicity is relative rare. One case they recognised was Lepotrema clavatum. Of the other euryxenous species listed above, Opechona bacillaris is a cosmopolitan species, with over 100 reports, mainly in scombrids and gadids. It is likely that this species is genuinely euryxenous, but also, it is likely that many records are erroneous. Of the 17 records we have of Hypocreadium patellare, 14 are from balistids, two from monacanthids and one from a haemulid (Perciformes) (Paru-

Rodney A. Bray and Jean-Lou Justine

khin 1978). As stated above this species is variable and the significance of the variability is poorly understood, but it is likely that the perciform record is either erroneous or of an accidental infection. Similarly some the species listed as stenoxenous (to host order) are probably also more specific than this designation implies. Preptetos cannoni is found in members of the perciform families Siganidae (mainly) and Pomacentridae and Miller et al. (2011) accepted this host distribution. On the other hand, Holorchis plectorhynchi is reported from haemulids and once (Durio and Manter 1968) in a lethrinid. As mentioned above, we have examined many specimens of the lethrinid in question and not recovered H. plectorhynchi, indicating that the record is likely to be of an accidental infection, or possibly an error. Of the 16 records of Diploproctodaeum macracetabulum, 15 are from the balistid Abalistes stellatus and one (Parukhin 1989) from a triacanthid, indicating that this species is practically oioixenous. Acknowledgements. Many students and colleagues were involved in the fishing expeditions and the parasitological survey, especially Aude Sigura, Adeline Grugeaud, Charlotte Schoelinck, Eva Řehulková and Bernard Marchand. The identification of several fishes was confirmed, from photographs, by ichthyologists: Bernard Séret (MNHN, Paris); Ronald Fricke (Staatliches Museum für Naturkunde, Stuttgart), and John E. Randall (Bishop Museum, Hawaii). Worms from chaetodontids and some siganids were collected in November 1997 by Tom Cribb, Sylvie Pichelin, Serge Morand, Betty Faliex, Claude Chauvet, Rene Galzin, Mark Rigby, Cedrik Lo and Pierre Sasal.

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(Accepted May 28, 2012)

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