BENEDENIINAE Johnston, 1931

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 : 666-667

BENEDENIINAE Johnston, 1931

Amended subfamilial diagnosis. Capsalidae sensu Yamaguti (1963) . Haptor aseptate, usually bearing three pairs of large median sclerites (in anterior to posterior succession): accessory sclerites (modi®ed hooklets I?; Kearn, 1963) usually nearest haptor centre; anterior hamuli; posterior hamuli; in some genera, one (e.g. Oligoncobenedenia Yamaguti, 1965 ) or both (e.g. Dioncopseudobenedenia Yamaguti, 1965 ) pairs of hamuli may be absent; accessory sclerites and one pair of hamuli may be absent in some genera (e.g. Calicobenedenia Kritsky and Fennessy, 1999 ); 14 peripheral hooklets (pairs II±VIII), pair II between hamuli. Marginal valve usually present, often scalloped. Ventral surface of haptor usually apapillate, rarely papillate (e.g. Trimusculotrema Whittington and Barton, 1990 ). Anterior attachment organs, one on each side of anterior end, disc-like (®gure 1B, C) or elongate (®gure 1A) usually bearing anterior areas where glandular secretions may be released. Pharynx protrusible, round or lobulate, with intrinsic gland cells. Testes two, juxtaposed. Vas deferens may or may not expand to form seminal vesicle immediately anterior to testes. Male copulatory organ a penis (e.g. Benedenia Diesing, 1858 ) or cirrus (e.g. Trimusculotrema ), usually enclosed in muscular sac (e.g. in Neobenedenia Yamaguti, 1963 ) or within canal with weakly muscularized wall (e.g. Benedenia and Menziesia Gibson, 1976 ); rarely provided with terminal sclerite(s) or papilla(e); rarely entirely sclerotized (e.g. Allometabenedeniella Velasquez, 1982 ). Speci®c details of anatomy of male copulatory organ and associated structures are important, are often characteristic of, and help to determine, particular genera. Male accessory gland extensive, with reservoir. Glands of Goto, if present, usually in posterior angle between testes, but similar cells noted in anterior angle between testes (e.g. some Benedenia spp. and Trimusculotrema spp. ). Germarium with internal chamber. Male and female genital apertures usually common, sometimes separate (e.g. Trimusculotrema ), generally opening on left body margin at level of pharynx. Vagina single, rarely absent (e.g. Neobenedenia ); communicates proximally with vitelline reservoir; position of vaginal pore variable, but usually near left side of body, very occasionally opening with common genital pore. Vaginal seminal receptacle (cf. oviductal seminal receptacle in Entobdella soleae (van Beneden and Hesse, 1864) Johnston, 1929 ; see Kearn, 1985) present or not. Single vitelline reservoir usually reported. Eggs tetrahedral, often with appendage. Parasites of external surfaces, buccal cavity, branchial chamber and gills of marine teleosts and elasmobranchs.

Type genus. Benedenia Diesing, 1858 .

Other genera. Allobenedenia Yamaguti, 1963 ; Allometabenedeniella Velasquez, 1982 ; Ancyrocotyle Parona and Monticelli, 1903 ; Benedeniella Johnston, 1929 ; Calicobenedenia Kritsky and Fennessy, 1999 ; Dioncopseudobenedenia Yamaguti, 1965 ; Lagenivaginopseudobenedenia Yamaguti, 1966 ; Menziesia Gibson, 1976 ; Metabenedeniella Yamaguti, 1958 ; Neobenedenia Yamaguti, 1963 ; Oligoncobenedenia Yamaguti, 1965 ; Pseudallobenedenia Yamaguti, 1966 ; Trimusculotrema Whittington and Barton, 1990 .

Remarks. Whittington and Kearn (1993) recognized 16 genera in the Benedeniinae . Egorova (1997) recognized Entobdellinae Bychowsky, 1957 and placed in it Entobdella Blainville in Lamarck, 1818 and Pseudoentobdella Yamaguti, 1963 . A separate paper on the Entobdellinae is in preparation by the senior author. Menziesia was rejected by Egorova (1997) and its species were incorporated into Benedenia . Our investigations have led us to recognize Menziesia , albeit with a diOEerent species composition, but we reject Tareenia Hussey, 1986 for reasons discussed at length below.