Benedeniella Johnston, 1929

Whittington, Ian D., 2010, Revision of Benedeniella Johnston, 1929 (Monogenea: Capsalidae), its assignment to Entobdellinae Bychowsky, 1957 and comments on subfamilial composition, Zootaxa 2519, pp. 1-30 : 5-6

Benedeniella Johnston, 1929

Amended generic diagnosis: With features of the Capsalidae sensu Yamaguti (1963) and Entobdellinae (amended above). Median haptoral sclerites large. Ventral haptor surface apapillate. Marginal valve conspicuous, wider anteriorly. Tendons entering haptor from body musculature conspicuous; passing through notch at proximal end of accessory sclerites before dividing, with part of tendon attaching to proximal ends of anterior hamuli and part attaching to ventral haptor tissue. Anterior attachment organs comprising ventral pair of circular, saucer-like discs, one on each ventrolateral border, bearing series of narrow complete and some incomplete ‘grooves’ anterolaterally. Dorsal protuberances present, in anterior to posterior succession, in form of horn-like papillae near junction of anterior attachment organs with body, excretory papillae and conical structures at peduncle where body joins haptor. Glands of Goto absent. Male copulatory organ a long cirrus. Cirrus sac elongate with muscular walls containing single male accessory gland reservoir. Long terminal male genital duct containing cirrus separate from long uterus for most of its length, uniting close to common genital pore, opening dorsally near left body margin at level of posterior region of left anterior attachment organ. Vagina long; vaginal pore opening dorsally near left body margin at different levels (anteriorly in B. macrocolpa ; posteriorly in B. posterocolpa ). Proximal region of vagina unspecialised, communicating directly with dorsal part of vitelline reservoir. Parasites of ventral skin of myliobatid elasmobranch rays.

Type species: Benedeniella macrocolpa ( Lühe, 1906) Yamaguti, 1963 [synonyms: Epibdella (Benedenia) macrocolpa Lühe, 1906 ; Epibdella macrocolpa ( Lühe, 1906) MacCallum, 1927 ; Benedenia (Benedeniella) macrocolpa ( Lühe, 1906) Johnston, 1929 ; Benedenia macrocolpa ( Lühe, 1906) Meserve, 1938 ].

Other species: Benedeniella posterocolpa ( Hargis, 1955) Yamaguti, 1963 [synonym: Benedenia posterocolpa Hargis, 1955 ].

Remarks: Johnston (1929) presented a history and examined the validity of some ‘tristomatid’ (= capsalid) genera and divided Entobdella and Benedenia into subgenera based on the position of the vaginal pore in relation to the common genital pore. The subgenus Benedeniella was proposed for species of Benedenia in which the vagina opened beside the common genital pore but where the vagina travelled posteriorly behind the testes and then anteriorly to the vitelline reservoir. Johnston (1929) included Epibdella (Benedenia) macrocolpa Lühe, 1906 as the type and only species of his new subgenus Benedeniella as Benedenia (Benedeniella) macrocolpa . Yamaguti (1963) raised Benedeniella to generic rank and transferred Benedenia posterocolpa Hargis, 1955 to Benedeniella because it also possessed a long vagina but which opened posteriorly some distance from the common genital pore. Yamaguti’s (1963) amended generic diagnosis perpetuated errors made by Lühe (1906) and Hargis (1955) regarding the anatomy of the uterus in B. macrocolpa and B. posterocolpa , respectively. Lühe (1906) referred to a ‘short uterus’ but a ‘common genital pore’ in B. macrocolpa inferring that this species has a long common genital duct along which eggs pass but which also accommodated the male copulatory organ. For B. posterocolpa, Hargis (1955) described the uterus as short, opening dorsally close to the bifurcation of the gut and separate from the marginal male pore, located posterolateral to the left anterior attachment organ. The arrangement of the terminal male and female genital ducts in both Benedeniella species was resolved and clarified by Kearn & Whittington (1992). The amended generic diagnosis presented above reflects the fact that the terminal male genital duct containing the cirrus and the uterus are each long ducts that remain parallel but separate from each other for most of their length and unite close to the common genital pore ( Kearn & Whittington 1992; see also redescriptions below). Two other Benedeniella species have been described, but neither is consistent with the concept for the genus as revised here. The report of Benedeniella congeri Yamaguti, 1958 [synonyms: Neobenedeniella congeri ( Yamaguti, 1958) Yamaguti, 1963 ; Benedenia congeri ( Yamaguti, 1958) Byrnes, 1986 ] from the gills of Conger myriaster (Brevoort) (Teleostei: Anguilliformes : Congridae ; conger and garden eels), Inland Sea of Japan was dealt with by Ogawa et al. (1995) when this taxon was considered a junior synonym of Benedenia epinepheli ( Yamaguti, 1937) Meserve, 1938 . This action was considered sound by Whittington et al. (2001).

Benedeniella unnithani Gupta & Chanana, 1976 was reported from the gills of a Caranx sp. (Teleostei: Perciformes : Carangidae ; jacks and pompanos) at Kavaratti, Laccadive Islands, Arabian Sea, India. No details about deposition of material in a museum were provided by Gupta & Chanana (1976) and Whittington & Barton (1990) attempted to locate and borrow specimens without success. I have made two further unsuccessful efforts to find their material. Based on their description, the three capsalid specimens they reported are clearly not a species of Benedeniella . Benedeniella unnithani is here considered a species incertae sedis.