Systematic Parasitology 57: 201–209, 2004. © 2004 Kluwer Academic Publishers. Printed in the Netherlands.

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Tormopsolus orientalis Yamaguti, 1934 (Digenea: Acanthocolpidae) from Seriola dumerili (Risso) (Perciformes: Carangidae) in the western Mediterranean Sea Pierre Bartoli1 , Rodney A. Bray2 & Francisco E. Montero3 1 Centre

d’Oc´eanologie de Marseille, UMR 6540 CNRS, DIMAR, Campus Universitaire de Luminy, Case 901, 13288 Marseille C´edex 9, France 2 Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK 3 Departamento de Zoolog´ıa & Instituto Cavanilles de Biodiversidad y Biolog´ıa Evolutiva, Universitat de Val` encia, PO Box 22085, Valencia 46071, Spain Accepted for publication 28th August, 2003

Abstract Tormopsolus orientalis Yamaguti, 1934, is redescribed from Seriola dumerili from off Corsica, Majorca and Águilas, SE Spain. The vitellarium is interrupted at the level of the ovary and both testes, and a bipartite seminal vesicle is found in many specimens. Oral sucker papillae are always seen. Type-specimens and voucher specimens from off Japan, Bermuda, Panama, Curaçao and the Great Barrier Reef have been compared with the Mediterranean species. Specimens of T. medius Reimer, 1983, from Mozambique have been studied and this species is synonymised with T. orientalis.

Introduction

Materials and methods

The greater amberjack Seriola dumerili (Risso) is a carangid perciform which has a circumglobal distribution (Froese & Pauly, 2003). It is well represented in the Mediterranean but absent in the Black Sea (SmithVaniz, 1986). Its digenean fauna has been well studied from a diversity of sites (Corkum, 1967; Fischthal & Thomas, 1972; Gijon-Botella et al., 1992; Manter, 1940; Parukhin, 1966, 1976; Yamaguti 1934, 1970) but has received less attention in the Mediterranean (Barbagallo & Drago, 1903; Fischthal, 1982; Fischthal et al., 1982; Grau et al., 1999; Odhner, 1911). In the course of our research in the Western Mediterranean, we have found numerous specimens of a species Tormopsolus Poche, 1926 in this host. The comparison of this form with 11 other species of this genus led us to the opinion that this form belongs to the widespread species of carangids, T. orientalis, Yamaguti, 1934.

Fish from off Corsica were collected alive and immediately necropsied after death. The digestive tract was sectioned into its various anatomical regions: stomach, pyloric caeca, duodenum, middle intestine, posterior intestine and rectum. Digenean specimens were collected from each of these segments under a dissecting microscope. They were never forcibly extracted, but left to liberate themselves and were studied both while still living and later as permanent preparations. Individuals were fixed in cold Bouin’s fluid between slide and coverglass without other pressure, stained in acetic carmine and mounted in Canada balsam. Fish from off Majorca died by anoxia and were collected and frozen 5-12 hours later, consequently the parasites were not in good condition. Fish from off Águilas were killed with benzocaine and the digestive tract removed and kept at 4 ◦ C for 24 hours; subsequently dead parasites were collected. Parasites from Spanish localities were fixed in AFA (alcoholformalin-acetic-acid) without pressure, stained with alum or iron-acetic carmines and mounted in Canada

202 balsam. Ovigerous specimens only have been taken into account in the descriptions. Worms were studied with differential interference contrast microscopy and illustrations were made using a drawing tube. Measurements are given as the range in micrometres, with the mean in parentheses. The term ‘forebody’ refers to the distance between the anterior extremity of the body and the anterior margin of the ventral sucker; the term ‘hindbody’ refers to the distance between the posterior margin of the ventral sucker and the posterior extremity of the body. Abbreviations: BMNH, the BMNH collection at The Natural History Museum, London; HWML, the Harold W. Manter Laboratory, Lincoln, Nebraska, USA; MPM, Meguro Parasitological Museum, Tokyo, Japan; QM, Queensland Museum, Brisbane, Queensland, Australia; USNPC, United States National Parasite Collection at Beltsville, Maryland, USA. Family Acanthocolpidae Lühe, 1906 Genus Tormopsolus Poche, 1926 Tormopsolus orientalis Yamaguti, 1934 Syn. Tormopsolus medius Reimer, 1983 Host: Seriola dumerili (Risso), Perciformes: Carangidae. Locality: Near the external limit of the Scandola Nature Reserve, Corsica (42◦23 N; 08◦ 33 E). Additional localities: Off Majorca, Spain (39◦27 – 39◦ 39 N, 02◦14 −02◦19 E). Off Águilas, Spain (37◦ 29 –37◦34 N, 01◦ 9 –01◦ 15 W). Sites: Stomach and pyloric caeca (favoured sites); duodenum (Corsica). Stomach, pyloric caeca and (especially) rectum (Spain). Dates of collection: Corsica: September, 2002. Majorca: September, 1996. Águilas: January, 1999. Number of fish examined: Corsica: 7. Four young specimens (Standard Length = 19.2–24.7 cm) and one adult specimen (SL = 123 cm) were unparasitised; 2 specimens were parasitised (SL = 44.5–74.3 cm). Majorca: 68 specimens (SL = 26.5–42.5 cm cm; SL of the infected fish = 26.5–39.5 cm). Águilas: 10 specimens. (SL = 30.5–35.5 cm; SL of the infected fish = 30.5–35.5 cm). Prevalence: Corsica: 28.6%. Majorca: 12.5%. Águilas: 90%. Abundance: Corsica: 7.4. Majorca: 0.8. Águilas: 3.4. Mean intensity: Corsica: 26. Majorca: 2.9 ± 4.1. Águilas: 3.7 ± 3.5.

Mediterranean material studied Corsica: 36 adults and 6 immature specimens. Voucher specimens BMNH 2003.8.26.1-10. Majorca: 54 adult specimens. Águilas: 34 adult specimens. Voucher specimens: Parasitological collection of the Cavanilles Institute of Biodiversity and Evolutionary Biology, University of Valencia, Spain. Additional material studied T. orientalis Yamaguti, 1934 ex Seriola quinqueradiata Temminck & Schlegel, Inland Sea, Japan. Holotype and 2 paratypes on slide MPM 22912, 1 paratype on slide MPM 22913 and 9 voucher specimens on slide MPM 22914. T. orientalis Yamaguti, 1934 ex ‘bonito, possibly Zonichthys fasciatus (Bloch)’ [?Seriola rivoliana Valenciennes], Bermuda. 1 voucher specimen on slide HWML 1122 (see Hanson, 1950). T. orientalis Yamaguti, 1934 ex ‘Zonichthys falcatus (Cuv. & Val.)’ [?Seriola rivoliana Valenciennes], Bermuda. 2 voucher specimens on slide BMNH 1982.5.15.20 (see Rees, 1970). T. orientalis Yamaguti, 1934 ex Seriola mazatlana Steindachner [=S. lalandi Valenciennes], Saint Bartholomew Island, Archipelago de las Perles, Panama. 1 voucher specimen on slide HWML 22565 (see Sogandares-Bernal, 1959). T. orientalis Yamaguti, 1934 ex Seriola dumerili (Risso), Curaçao. 1 voucher specimen on slide HWML 124456 (see Nahhas & Cable, 1964). T. medius Reimer, 1983 ex Seriola [now Seriolina] nigrofasciata (Rüppell), Mozambique. Holotype and paratypes, 4 complete worms and 11 fragments on slide BMNH 2003.8.26.11-14, donated by Professor Reimer (see Reimer, 1983). T. orientalis Yamaguti, 1934 ex Seriola lalandi Valenciennes, Heron Island, Queensland (23◦27 S, 151◦55 E, 14.i.1992, coll. R.D. Adlard). 1 voucher specimen on slide QM G221989. Description (Figures 1–6) General morphology (Figures 1–2). Body very elongate with parallel margins, with maximum width at level of ventral sucker or between testes, very slightly constricted at level of ovary and testes; anterior body extremity rounded; posterior extremity tapering. Tegument. Tegument spinose. Spines on the dorsal body surface strong, long, acuminate, not scale-like, slightly recurved in distal region, distributed in clear rows in area of oral sucker; spines of 6 (sometimes 5)

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Figure 1. Tormopsolus orientalis Yamaguti, 1934 from Seriola dumerili, off Corsica. Dorsal view.

anterior rows enlarged (up to 24 ± 7 (14–32) long), longitudinally aligned, decrease in size posteriorly (in lateral field of mid-forebody, up to 14 ± 1.6 (11–17) long), becoming slender, hooked-shaped, very densely distributed from post-oral area to ovarian zone, less dense posteriorly, reaching virtually to posterior extremity. On ventral surface spines absent around oral sucker aperture, between suckers and around ventral sucker aperture, but stout on lateral body margins (Figure 3); small area with some small spines just posterior to oral sucker, and laterally at this level; between ventral sucker and ovarian zone spines are dense and hooked-shaped, but posteriorly they become slender, with density decreasing towards posterior extremity. Eye-spots. Granules widespread at mid-pharyngeal level. Glands. Numerous discrete gland-cells distributed in parenchyma from level of oral sucker to level of seminal vesicle or pars prostatica (Figures 1–2). Glandcell ducts open on anterior and lateral margins of oral sucker aperture (Figure 3). Suckers. Oral sucker muscular, oval to subcircular; aperture subterminal, becoming wider posteriorly and merging into lateral muscular outgrowths (papilla). Ventral sucker circular, with antero-ventrally directed aperture. Digestive system. Prepharynx very short, more often virtually absent. Pharynx oval. Oesophagus absent. Intestinal bifurcation immediately anterior to ventral sucker. Digestive caeca slender, lateral to gonads, open

Figure 2. Tormopsolus orientalis Yamaguti, 1934 from Seriola dumerili, off Corsica. Ventral view.

into urinary bladder forming uroproct; gastrodermis thin. Male reproductive system. Testes 2, close to posterior extremity, intercaecal, median, entire, longitudinally ovoid, sometimes slightly constricted medially, always widely separated from each other, never contiguous with ovary. Cirrus-sac long (15–26 (19)% of body length), length c. 39–54 (37)% of ventral sucker to ovary distance, with diameter regularly decreasing distally, intercaecal. Seminal vesicle in proximal end

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Figure 3. Tormopsolus orientalis Yamaguti, 1934 from Seriola dumerili, off Corsica. Oral sucker. Ventral view.

of cirrus-sac, large, clearly bipartite (with strong transverse wall and narrow medial connection) in 55.6% of Corsican specimens (Figures 1, 4), undivided in 38.9% of Corsican specimens (Figure 2) and questionable in 5.5% of Corsican specimens. Pars prostatica tubular, sinuous, surrounded by narrow sheath of prostatic cells. Ejaculatory duct thick-walled, provided with short, stout spines throughout its length (Figures 4, 5). Genital atrium very long, devoid of sclerified structures. Genital pore median, immediately anterior to ventral sucker. Female reproductive system. Ovary spherical to slightly oval, entire, slightly sinistral (41.7%) or dextral (33.3%), sometimes median (25%), intercaecal, pretesticular, widely separated from anterior testis. Oöcapt emerges on antero-dorsal face of ovary (Figure 6). Laurer’s canal short, thin, not convoluted, devoid of gland-cells, opens onto dorsal surface of body just anterior to ovary. Mehlis’ gland pre-ovarian. Uterine seminal receptacle very large (Figures 1–2), just pre-ovarian. Uterus pre-ovarian, intercaecal, coiling between ovary and proximal end of cirrus-sac. Eggs large, thin-walled, usually collapsed, operculate. Metraterm rectilinear, parallel to ejaculatory duct and of similar length, thick-walled, provided with short, stout spines (Figure 5). Vitellarium follicular; follicles in dorsal and ventral fields, filling post-testicular region, spaces between gonads and area between ovary and proximal extremity of cirrus-sac, interrupted at level of gonads (100%), contiguous with the cirrus-sac (47.2%), overlapping it (44.5% mean 225 µm) or just posterior to it (8.3%; mean 197 µm); anterior 15–28 (20%) of hindbody length devoid of follicles.

Figure 4. Tormopsolus orientalis Yamaguti, 1934 from Seriola dumerili, off Corsica. Terminal genitalia. Ventral view. Abbreviation: gp, genital pore.

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Figure 5. Tormopsolus orientalis Yamaguti, 1934 from Seriola dumerili, off Corsica. Distal extremities of cirrus-sac and metraterm. Ventral view.

Excretory system. Excretory vesicle I-shaped, thinwalled, long, reaches to level of posterior extremity of ovary [36-59 (43%) of hindbody length]; posterior extremity receives extremities of digestive caeca. Excretory pore terminal. Measurements (see Tables 1, 2). Supplementary data below (based on 20 adult specimens from Corsica): Body breadth at level of: – ovary: 200–520 (320) – anterior testis: 210–476 (310) – posterior testis: 197–444 (301) Eggs (collapsed): 62–74 × 25–36 (67 × 29) (n=60) Eggs (not collapsed): 62–71×32–40 (66×36) (n=20)

Discussion At the present time, 11 species of Tormopsolus are recognised (see Bray & Cribb, 2001). Among them, only three have their vitellarium described as interrupted at the level of the ovary and both testes: T. orientalis Yamaguti, 1934, T. filiformis Sogandares-

Figure 6. Tormopsolus orientalis Yamaguti, 1934 from Seriola dumerili, off Corsica. Proximal female genital system. Dorsal view.

Bernal & Hutton, 1959 and T. medius Reimer, 1983. We have examined specimens of the latter species and include measurements of four laterally mounted worms in our Table 2. We can detect no substantial differences between these worms and T. orientalis and, therefore, consider them synonymous. Further evidence for this synonymy is included in the following discussion. T. filiformis is the only species of Tormopsolus that has been described with a bipartite seminal vesicle (Sogandares-Bernal & Hutton, 1959; Hafeezullah, 1991). This condition occurs in 55.6% of Corsican specimens, but the seminal vesicle is undivided in 38.9% and indeterminable in 5.5%. In specimens with a bipartite vesicle, the strong transverse septum is obvious; in other specimens, the septum could be widely open and forced against the lateral wall when the seminal vesicle is full of spermatozoa. Hafeezullah (1991, p. 109) pointed out that in T. filiformis the septum of seminal vesicle is ‘sometimes . . . not clearly seen’, but considered that, even if the septum is not visible in some specimens, the seminal vesicle is fundamentally bipartite. We have detected a bipartite seminal

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Table 1. Measurements, ratios and features of Tormopsolus specimens. Unless otherwise stated the ratios are given as percentages of the body-length. Species Host Locality n

T. orientalis Seriola dumerili Corsica 20

T. orientalis Seriola quinqueradiata Inland Sea, Japan 13

T. orientalis Seriola rivoliana (?) Bermuda (Rees) 2

T. orientalis Seriola rivoliana (?) Bermuda (Hanson) 1

Length Width Forebody Hindbody (HB) Oral sucker (OS) Prepharynx Pharynx Ventral sucker (VS) Genital atrium (GA) length Genital atrium into hindbody Cirrus sac (CS) VS to post end CS (VS-CS) VS to Ant. Vit. Extent VS to Ovary Ovary Ovary to AT Anterior testis (AT) Between testes Posterior testis (PT) Post-testicular region (PTR) Eggs Width% Forebody% Hindbody% VS% of OS (length) VS% of OS (width) CS length% VS-CS% of VS-Ovary VS-Vit% VS-Ovary% Ovary to AT% Between testes% PTR% OS length% VS length% Prepharynx% Pharynx length% AT length% PT length% Ov length% GA length% GA into HB% of VS–Ov % of hindbody devoid of vit. Vit. gap at ovary% Vit. gap at AT% Vit. gap at PT%

4,961–11,305 (7,412) 327–474 (394) 272–448 (330) 4,334–10,627 (6,756) 168–227 × 176–227 (194 × 203) 0–86 (26) 86–139 × 67–134 (118 × 105) 288–368 × 291–381 (326 × 332) 474–1,344 (771) 96–838 (373) 832–1, 376 × 64–189 (1, 099 × 124) 998–2,144 (1,391) 896–2,368 (1,357) 2,416–6,144 (3,730) 192–349 × 122–235 (250 × 174) 416–1,632 (846) 422–730 × 154–337 (552 × 234) 166–736 (401) 374–739 × 163–349 (574 × 241) 246–794 (490) 62–74 × 32–40 (67 × 29) 3.6–7.3 (5.4) 3.0–6.8 (4.6) 87.4–94.0 (90.9) 162–171 (168) 147–192 (168) 12.2–16.8 (14.8) 27.4–49.3 (37.8) 18.0–20.9 (18.3) 46.3–56.0 (50.2) 6.4–18.0 (11.2) 3.0–7.8 (5.3) 4.1–9.6 (6.5) 2.1–3.4 (2.7) 3.0–5.8 (4.6) 0–1.1 (04) 1.0–2.1 (1.6) 5.8–8.9 (7.5) 5.7–9.2 (7.8) 2.3–4.2 (3.4) 5.4–14.8 (10.0) 3.8–16.5 (9.9) 15.2–27.7 (20.1) 100 100 100

3,352–8,816 (4,475) 319–690 (404) 249–448 (326) 2,772–7,990 (3,885) 129–215 × 157–262 (162 × 194) 43–105 (63) 90–170 × 78–160 (112 × 109) 173–378 × 215–430 (264 × 294) 286–620 (421) 0–126 (29) 631–1, 418 × 84–174 (904 × 144) 539–1,359 (766) 319–1,044 (573) 1,219–3,642 (1,754) 104–332 × 134–305 (167 × 185) 225–1,002 (402) 227–602 × 185–337 (362 × 252) 40–386 (171) 254–909 × 168–327 (448 × 244) 280–1,116 (534) 71–87 × 31–45 (76 × 38) 7.7–11.4 (9.2) 5.1–10.6 (7.7) 81.8–90.6 (86.1) 128–184 (163) 138–171 (156) 14.9–27.0 (20.5) 28.2–60.7 (43.9) 9.3–19.4 (12.6) 35.6–44.2 (38.7) 6.7–12.1 (8.8) 1.2–6.1 (3.6) 8.0–17.5 (11.6) 2.4–4.8 (3.8) 4.0–7.9 (6.2) 0.5–2.3 (1.5) 1.8–3.6 (2.6) 5.2–11.0 (8.4) 5.8–12.2 (10.2) 2.7–5.0 (3.8) 6.6–12.3 (9.5) 0–4.8 (1.3) 11.0–21.6 (14.6) 83 100 100

4,827–4,993 485–512 342–455 4,149–4,163 217–193 × 215 ? 66 × 105 336–375 × 265 590–749 34–160 1,322–1, 772 × 132–200 918–951 863–866 1,901–1,926 253–276 × 202–214 280–318 355–365 × 263–347 102–234 423–439 × 256–328 652–698 66–72 × 33 10.0–10.3 7.1–9.1 83.4–86.0 155–194 124 27.4–35.5 47.7–50.0 17.3–17.9 38.6–39.4 5.8–6.4 2.0–4.8 13.5–14.0 3.9–4.5 7.0–7.5 ? 1.4 7.1–7.6 8.5–9.1 5.1–5.7 12.2–15.0 1.8–8.4 20.7–20.9 100 100 100

5,662 364 411 4,957 179 × 188 83 157 × 109 294 × 278 788 157 2, 106 × 151 1,904 1,709 2,878 248 × 187 247 452 × 209 143 414 × 216 508 72 × 34 6.4 7.3 87.5 148 164 37.2 66.2 30.2 50.8 4.4 2.5 9.0 3.2 5.2 1.5 2.8 8.0 7.3 4.4 13.9 5.4 34.5 100 100 100

207 Table 2. Measurements, ratios and features of Tormopsolus specimens. Unless otherwise stated the ratios are given as percentages of the body-length. Species Host Locality n

T. medius Seriola nigrofasciata Mozambique 4

T. orientalis Seriola lalandi Heron Island 1

T. orientalis Seriola lalandi Gulf of Panama 1

T. orientalis Seriola dumerili Curaçao 1

Length Width Forebody Hindbody (HB) Oral sucker (OS) Prepharynx Pharynx Ventral sucker (VS) Genital atrium (GA) length Genital atrium into hindbody Cirrus sac (CS) VS to post end CS (VS–CS) VS to Ant. Vit. Extent VS to Ovary Ovary Ovary to AT Anterior testis (AT) Between testes Posterior testis (PT) Post-testicular region (PTR) Eggs Width% Forebody% Hindbody% VS% of OS (length) VS% of OS (width) CS length% VS-CS% of VS-Ovary VS-Vit% VS-Ovary% Ovary to AT% Between testes% PTR% OS length% VS length% Prepharynx% Pharynx length% AT length% PT length% Ov length% GA length% GA into HB% of VS-Ov % of hindbody devoid of vit. Vit. gap at ovary% Vit. gap at AT% Vit. gap at PT%

4,730–5,377 (5,076) lateral 247–389 (334) 4,165–4,675 (4,500) 192–214 (205) long 27–115 (50) 134–150 (143) long 317–333 (323) long 576–1,202 (809) 84–330 (243) 678–860 × 86–113 (750 × 99) 758–944 (823) 734–982 (898) 2,312–2,460 (2,409) 144–178 × 93–131 (166 × 117) 273–405 (337) 367–429 × 148–197 (400 × 164) 148–220 (179) 378–461 × 138–179 (414 × 157) 224–570 (397) 63–74 × 28–33 (69 × 31) lateral 5.2–7.2 (6.6) 86.4–93.5 (88.7) 153–157 (155) ? 13.0–18.2 (14.9) 30.8–38.6 (34.3) 15.5–19.0 (17.6) 45.7–48.9 (47.5) 5.3–8.1 (6.7) 3.1–4.4 (3.5) 4.7–10.6 (7.7) 3.8–4.4 (4.1) 5.9–6.7 (6.3) 0.5–2.2 (1.0) 2.6–3.1 (2.8) 7.3–8.5 (7.9) 7.6–8.9 (8.2) 2.9–3.7 (3.3) 10.7–21.6 (16.1) 3.4–13.8 (10.2) 17.6–21.0 (19.9) 100 100 100

6,255 451 555 5473 161 × 152 202 96 × 88 227 × 291 602 307 1, 005 × 134 1,173 1,147 2,574 230 × 176 657 422 × 279 380 531 × 312 645 72 × 36 7.2 8.9 87.5 141 191 16.1 45.6 18.3 41.1 10.5 6.1 10.3 2.6 3.6 3.2 1.5 6.7 8.5 3.7 9.6 11.9 21 100 100 100

10,979 455 660 9,997 203 × 214 348 146 × 105 322 × 359 1,241 870 1, 964 × 116 2,606 2,417 6,299 167 × 128 1,268 358 × 144 631 430 × 126 658 83 × 37 4.1 6.0 91.1 158 168 17.9 41.4 22.0 57.4 11.6 5.8 6 1.9 2.9 3.2 1.3 3.6 3.9 1.5 11.3 13.8 24.2 100 100 100

11,489 615 554 10,515 258 × 284 100 244 × 136 420 × 420 1,021 338 2, 561 × 154 2,581 2,475 6,321 335 × 269 693 666 × 301 544 693 × 326 1,088 74 × 32 5.4 4.8 91.5 163 148 22.3 40.8 21.5 55 6 4.7 9.5 2.2 3.7 0.9 2.1 5.8 6 2.9 8.9 5.3 23.5 100 100 100

208 vesicle in one of the 13 T. orientalis specimens from Japan, which are rather poorly preserved and heavily flattened. The three specimens of T. orientalis from off Bermuda show evidence of a bipartite seminal vesicle and four of five specimens (whole worms and fragments where the feature is seen) of T. medius have distinctly bipartite seminal vesicles. The unflattened worm from off Heron Island has an entire seminal vesicle, as has the worm from off Panama. The worm from off Curaçao has what appears to be a multipartite seminal vesicle, presumably due to flattening at fixation. It seems clear to us that the bipartite nature of the seminal vesicle varies between worms and may sometimes be affected by fixation, particularly heavy flattening. Although possibly sharing a bipartite seminal vesicle, T. filiformis and T. orientalis differ conspicuously in that the former possesses a very long forebody and a very long prepharynx, an intestinal bifurcation located in the mid-forebody, no oral sucker papillae and definitive hosts which are not carangids. The oral sucker papillae, although not described in Tormopsolus spp. prior to Bray & Cribb (2001), are seen clearly in 9 of the 13 Japanese specimens, including the holotype, and probably obscured in the others by flattening. This feature is also clearly seen in two of the specimens from off Bermuda (Hanson’s specimen and one of Rees’), all the specimens from off Mozambique and the specimens from off Heron Island, Curaçao and Panama. Sogandares-Bernal (1959) detected differences to the types in his specimen from off Panama, but considered them intraspecific variations, and pointed out that Hanson’s (1950) material from off Bermuda agreed with his ‘in all details’. Rees (1970) reported specimens of T. orientalis in ‘Zonichthys falcatus (Cuv. & Val.), the bonito’ from off Bermuda, two of which we have studied (Table 1), saying that Hanson (1950) reported it from the same host. In fact, Hanson says ‘bonito, possibly Zonichthys fasciatus (Bloch)’. According to Eschmeyer (2003) Zonichthys Swainson, 1839 is a synonym of Seriola Cuvier, 1816 and Froese & Pauly (2003) listed both Seriola fasciata and S. falcata. S. fasciata (Bloch), the lesser amberjack, is a valid species, whereas S. falcata Valenciennes is a synonym of S. rivoliana Valenciennes, the almaco jack. In the long lists of common names given for these species by Froese & Pauly (2003), ‘bonito’ does not occur, whereas Z. falcatus (Cuvier & Valeciennes) is known as the bonito in Beebe & Tee-Van (1933). Thus, it is not entirely clear in which host species Rees (1970) ac-

tually found her specimens, but judging from the name given by Rees (1970), it should be S. rivoliana and almost certainly a species of Seriola. The distribution given by Froese & Pauly (2003) for both S. fasciata and S. rivoliana includes Bermuda. The measurements, ratios and features given on Tables 1 and 2 indicate the variability exhibited by this species. A note of caution should be sounded, however, in that the material has clearly been treated in different ways, from the heavily flattened (Japanese material) to unflattened (Heron Island material). The variation detected in the larger samples, i.e. our new material and the Japanese material, encompasses practically all of the variation seen elsewhere. T. orientalis has been reported in Seriola dumerili off Curaçao by Nahhas & Cable (1964). The only other record of Tormopsolus from S. dumerili is that of T. hawaiiensis Yamaguti, 1970 from off Hawaii (Yamaguti, 1970). This species differs from T. orientalis in lacking a confluence of the vitelline follicles between the gonads and apparently does not always have vitelline gaps at the level of the gonads. Predilection for sites among species of Tormopsolus may vary, in that T. orientalis from off Corsica apparently has a preference for the stomach and pyloric caeca, although also occurring in the duodenum, whereas most other findings of T. orientalis, including those from Spanish waters are reported in the intestine. Post-mortem migration may account for some of these findings. It should be noted, however, that T. orientalis was reported in the stomach of carangids by Lebedev (1970) and Rees (1970), and the hosts from off Corsica were dissected within five minutes of the death of the host. Bray & Cribb (2001) produced a cladistic tree where T. orientalis and T. medius are sister taxa so, as they are herein considered synonymous, the tree stands but with these two branches pruned to one. Sister to the resulting branch is (T. asiatica Parukhin, 1976, T. attenuatus Bray & Cribb, 2001). These forms differ from T. orientalis in lacking vitelline gaps at the level of the testes, which seem to virtually always occur in T. orientalis (Tables 1 and 2).

Acknowledgements Fieldwork on Corsica was carried out under the auspices of the Comité Scientifique de la Réserve Naturelle de Scandola, and PB was funded by the French Ministère de l’Environnement and by the Ré-

209 gional Authorities of Corsica. Research in Spain was supported by project OCYT No. GV-01-9 from the Valencian Government and MCYT No BOS2002-878. We acknowledge Fernando de la Gándara and Antonio García (Instituto Español de Oceanografía) for technical support. Thanks are also due to Dr Juan Antonio Raga (University of Valencia) for his comments and assistance and to Dr David I. Gibson (The Natural History Museum) for reading the manuscript. We are most grateful to Dr Eric Hoberg and Dr Patricia Pillitt, United States National Parasite Collection, Beltsville, Maryland, USA, Dr Jun Araki of the Meguro Parasitological Museum, Tokyo, Japan and Dr Scott Gardner and Dr Ina van der Veen, Harold W. Manter Laboratory, Lincoln, Nebraska, USA, for the loan of specimens and to Professor Lothar Reimer for the donation of specimens.

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