Benedenia ovata ( Goto, 1894 ) Johnston, 1929

Whittington, I. D., Deveney, M. R. & Wyborn, S. J., 2001, A revision of Benedenia Diesing, 1858 including a redescription of B. sciaenae (van Beneden, 1856) Odhner, 1905 and recognition of Menziesia Gibson, 1976 (Monogenea: Capsalidae), Journal of Natural History 35 (5), pp. 663-777 : 720-725

Benedenia ovata ( Goto, 1894) Johnston, 1929

(®gure 26)

Synonyms. Epibdella ovata Goto, 1894 ; Epibdella (Phylline) ovata ( Goto, 1894) Parona and Perugia, 1895 ; Epibdella (Benedenia) ovata ( Goto, 1894) Monticelli, 1902 ; Tristoma ovata ( Goto, 1894) Heath, 1902 ; Benedenia (Parabenedenia) ovata ( Goto, 1894) Johnston, 1929 ; B. akaisaki Iwata, 1990 (new synonym).

Material studied. FU: nos. M-0001 (holotype of B. akaisaki ), M-0002, M-0003 (paratypes of B. akaisaki ) (3 slides, 3 individuals) ex gills of Caprodon schlegelii (GuÈnther) (Serranidae) , Coast of Kagosima Prefecture, Japan. No type material for B. ovata from the collection of Goto could be located and we nominate FU M-0001 as the neotype for B. ovata .

Observations. Our analysis of B. ovata is based on the original description by Goto (1894) and on specimens of B. akaisaki prepared and described by Iwata (1990) which we consider to be synonymous with B. ovata . How many specimens Goto (1894) used in his description of B. ovata is unknown but Iwata’s description (as B. akaisaki ) was based on three whole-mounts, all of which we examined. Although Goto’s description of B. ovata lacks comprehensive measurements, the description by Iwata (1990) of B. akaisaki is thorough and should be consulted for details. It is unclear whether or not Iwata was familiar with B. ovata or any other Benedenia species because B. akaisaki was not compared with other congeners. Our observations indicate that the specimens of B. akaisaki display a combination of characters that identify them clearly as B. ovata . Benedenia ovata is distinctive because it has a transversely broad haptor. The small accessory sclerites are located posteriorly on the haptor and have a distinctive shape (®gure 26B±D). Goto depicted a pair of tendons passing close to the proximal ends of the accessory sclerites and we have determined that these tendons pass through the bi®d notch of the accessory sclerites (®gure 26A). In addition, Goto described the muscles which insert at the anterior and posterior edge of the haptor which created marginal notches. Despite an appearance which suggests that Iwata heavily ¯attened the specimens prepared in his study, these notches were visible in our observations of his material. The anterior hamuli are recurved (®gure 26C) and the posterior hamuli possess a broad root and a strongly hooked distal tip (®gure 26D). The haptor bears a marginal valve which is scalloped and wider anteriorly (®gure 26A). No other details of the marginal valve could be determined because of external damage and the heavily ¯attened condition of the specimens. Few other features of B. ovata require special mention because the original description by Goto is accurate. We con®rmed in Iwata’s specimens that the vaginal pore is located in the mid-body between the germarium and the common genital pore. Parasites recovered from ®sh markets, as Iwata’s benedeniines were, are often in poor condition and are di cult to process to produce museum quality whole-mounts. Iwata, however, has prepared remarkably good specimens, especially considering these parasites were most probably dead when collected. Good preparations are of vital importance for future taxonomic work on benedeniines and the quality of Iwata’s material helped our study enormously.

Type-host and locality. Anthias (5 Caprodon, Temminck and Schlegel , see Paxton et al., 1989) schlegelli ( Serranidae ), Misaki, Japan.

Published records and descriptions. Goto (1894); Iwata (1990).

Published host records. Serranidae : Anthias Schlegelii (see Goto, 1894; sic in breach of binomial nomenclature); Caprodon schlegelii (see Iwata, 1990).

Site . Gills.

Distribution. Misaki, Japan ( Goto, 1894); Kagosima Prefecture, Japan ( Iwata, 1990).

Remarks. Neither Goto (1894) nor Iwata (1990) compared B. ovata with congeneric species but this taxon remains, with B. jaliscana and B. rohdei , among the most distinctive of Benedenia species because of its transversely broad haptor and the position of the vaginal pore which opens in the mid-body between the germarium and the common genital pore. We have concluded that the dorsal or ventral position of the vaginal pore is di cult to discriminate in material currently available and further study is required to fully assess this feature for B. ovata . We believe B. ovata should be retained in Benedenia for the same reasons and arguments outlined for those species with a vaginal pore anterior to the common genital pore (see p. 682±683).

Bychowsky (1957) noted septa on the haptor of B. ovata , Megalobenedenia (as Benedenia ) derzhavini and B. sebastodis . Although M. derzhavini does bear septa ( Egorova, 1994), we did not observe septa on the haptor of B. ovata . We were not in a position, however, to prepare any specimens of B. ovata in glycerine gelatin as suggested by Bychowsky.

Benedenia pompatica Buhrnheim, Gomes and Varela, 1973 (®gure 27)

Material studied. IOC: Nos 30.813a, 30.813b, 30.813c, 30.815a, 30.815b, 30.816 and 30.817 (paratypes) (each comprising 1 slide, 1 individual5 a total of 7 specimens). Benedenia pompatica was described from two host species, Argyrosoma regium (probably Argyrosomus regius Asso , see Gri ths and Heemstra, 1995 for a complete discussion) and Seriola sp. but we are unable to attribute a particular host species to the types because none of the slides bear host information. However, of 13 specimens collected by Buhrnheim et al. (1973), only one was reported from Seriola sp. All specimens collected were found on the gills of their hosts (Buhrnheim et al., 1973).

Observations. Most of the paratypes are of poor quality and IOC do not lend holotypes. Our study of B. pompatica is, therefore, based on the paratypes and the original description. There is a large size range among the material studied but B. pompatica can be a large species with a range in total length of 3224 to 10 527 and a mean of 6283 (n 5 4 specimens). The anterior attachment organs are large and always exceed the size of the relatively small pharynx. The haptor on which the three pairs of median sclerites are positioned posteriorly is relatively large and circular (®gure 27A). The large accessory sclerites are broad, slightly curved, taper to a pointed distal tip and bear a notch at their proximal ends through which tendons pass (®gure 27A). The narrow anterior hamuli are about two-thirds the length of the accessory sclerites; their distal ends are rounded but bear a small spike subterminally at right angles to the longitudinal axis of the hamuli (®gure 27C). The proximal ends of the anterior hamuli overlap the proximal ends of the accessory sclerites and the distal ends of the anterior hamuli overlap completely the small posterior hamuli (®gure 27A). We counted 14 hooklets. Details of what could be determined for the marginal valve are: four to ®ve lobes between hooklets of pair II; two lobes between hooklets of pairs II and III; one lobe between hooklets III and IV, and between IV and V; 11 small lobes between hooklets V and VI; 10±11 small lobes between hooklets VI and VII. Some of the small lobes may be fused. Details of the marginal valve anteriorly could not be resolved (®gure 27A).

Few details of the reproductive system could be determined because of the poor quality of the material. We can con®rm that glands of Goto are present. Buhrnheim et al. (1973, ®gure 5) depicted a long narrow uterus running parallel for much of the length of the penis canal and they show and describe a common genital pore. None of the material available was su ciently good to clarify this character. Buhrnheim et al. described the vagina as opening some distance from the common genital pore and we can con®rm this feature.

Type-host and locality. Buhrnheim et al. (1973) did not specify which of the two host species was the type. We list, therefore, both host species as presented by Buhrnheim et al. (1973): Argyrosoma regium (5 Argyrosomus regius ) ( Sciaenidae ); Seriola sp. (Carangidae) , Cabo Branco, Portugese Coast of Africa, Atlantic Ocean.

Published records and description. Buhrnheim et al. (1973).

Published host records. Carangidae : Seriola sp. ; Sciaenidae : Argyrosoma regium (5 Argyrosomus regius ).

Site . Gills.

Distribution. Cabo Branco, Portuguese Coast of North Africa, Atlantic Ocean.

Remarks. Despite the poor quality of type material and of the original description, our analysis of B. pompatica indicates it is a valid species distinguished clearly from others in the genus by a combination of the following characters: its large size; the large circular haptor; the posterior position of the median sclerites on the haptor; the large curved accessory sclerites with tapering distal tips; the rounded distal tips of the anterior hamuli which bear a small spike-like projection subterminally; large anterior attachment organs. Further examination of new material should show whether B. pompatica has a long uterus as described by Buhrnheim et al. (1973) and clarify features of the male system.

Buhrnheim et al. (1973) suggested that B. pompatica was most similar to B. sciaenae . Both species are large (e.g. ®gure 3A), have large anterior attachment organs and have their large sclerites located posteriorly on the haptor. However, from data presently available, B. sciaenae is a much larger species and can attain a total length of up to 25 mm. Furthermore, these species can be distinguished by the shapes of their accessory sclerites, anterior hamuli and the arrangement of the marginal valve. Benedenia sciaenae possesses a long uterus which opens separately and not via a common genital pore (see p. 674). Further study of specimens of B. pompatica is needed to determine whether or not this is a feature of this species. Moreover, B. sciaenae and B. pompatica can be found on members of the Sciaenidae and, indeed, on the same host species, although B. pompatica was also recorded from a carangid, Seriola sp. Benedenia sciaenae , however, is mostly reported from the external surfaces of its hosts (see p. 676) whereas B. pompatica is recorded only from the gills (Buhrnheim et al., 1973).

Benedenia rohdei Whittington, Kearn and Beverley-Burton, 1994 (®gure 28)

Material studied. QM: Nos GL18312±18316 (holotype and paratypes) (5 slides, 5 individuals) ex gills of Lutjanus carponotatus (Lutjanidae) from Heron Island, Queensland, Australia; IDW: Nos CAP322, CAP328 (paratypes) (2 slides, 2 individuals) ex gills of L. carponotatus (Lutjanidae) from Heron Island, Queensland, Australia and CAP125 ±127 (vouchers) (3 slides, 3 individuals) from Lizard Island, Queensland, Australia.

Observations. An extensive study of B. rohdei was made using live and preserved, mounted specimens ( Whittington et al., 1994). Original measurements were based on 16 whole-mounts. This study also involved an examination of the oncomiracidium. Benedenia rohdei is distinguished from other species by possessing a sclerite on the distal tip of the penis (®gure 28F). It also has a longitudinally elongate haptor (®gure 28A) and vitelline follicles displaced from the median region of the body posterior to the testes by large muscles ( Whittington et al., 1994).

Type-host and locality. Lutjanus carponotatu s ( Lutjanidae ), Heron Island, Great Barrier Reef, Queensland, Australia.

Published records. Whittington et al. (1994); Ernst and Whittington (1996); Whittington and Cribb (1999); Whittington and Ernst (in preparation); Deveney et al. (in preparation).

Description. Whittington et al. (1994).

Published host record. Lutjanidae : Lutjanus carponotatus .

Sites. Gills.

Distribution. Heron and Lizard Islands, Great Barrier Reef, Queensland, Australia.

Remarks. Examination of the holotype and paratypes of this species shows that the original account by Whittington et al. (1994) is accurate. In the original description, B. rohdei was compared with B. jaliscana which has also been described as having an`armed cirrus’. Closer examination of B. jaliscana showed it to have three elongate papillae with conical tips on the end of its penis (®gure 24A) as noted by Whittington et al. (1994) rather than the three`stout hooks’ mentioned in the original description by Bravo-Hollis (1952). This arrangement is distinctly diOEerent from the single penis sclerite (®gure 28F) in B. rohdei . As no other described Benedenia species has any sclerotized elements associated with the reproductive systems, this feature alone clearly discriminates B. rohdei in this genus.