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)

Benedenia Diesing, 1858, a genus of capsalid (benedeniine) monogeneans, is redefined. The generic diagnosis is amended to include: the path of tendons in the haptor from extrinsic muscles in the body; presence and form of the marginal valve; a penis occupying a penis canal with weakly muscular wall; a weakly muscular accessory gland reservoir proximal to the penis and enclosed by a proximal extension of the wall of the penis canal; male and female genital apertures usually common, rarely separate; vagina with pore usually close to the common genital pore but may open in mid body between the germarium and the common genital pore, or anterior to the common genital pore. A conservative approach is adopted and the generic diagnosis is clarified and broadened to accommodate species that display some variation in reproductive anatomy, especially of the female system. We argue against potential alternative actions such as defining Benedenia strictly to contain species with separate male and female genital apertures and against recognition of a separate genus, Tareenia Hussey, 1986, for species with a vaginal pore anterior to the common genital pore. Under our conception, Benedenia comprises 21 species: B. sciaenae (van Beneden, 1856) Odhner, 1905 (type species); B. acanthopagri (Hussey, 1986) comb. nov.; B. anticavaginata Byrnes, 1986; B. bodiani Yamaguti, 1968; B. elongata (Yamaguti, 1968) Egorova, 1997; B. epinepheli (Yamaguti, 1937) Meserve, 1938; B. hawaiiensis Yamaguti, 1968; B. hendorffi(von Linstow, 1889) Odhner, 1905; B. hoshinai Ogawa, 1984; B. innobilitata Burhnheim Gomes and Varela, 1973: B. jaliscana Bravo-Hollis, 1952; B. lolo Yamaguti, 1968; B. lutjani Whittington and Kearn, 1993: B. monticellii (Parona and Perugia, 1895) Johnston, 1929; B. ovata (Goto, 1894) Johnston. 1929: B. pompatica Burhnheim, Gomes and Varela, 1973; B. rohdei Whittington, Kearn and Beverley-Burton, 1994; B. scari Yamaguti, 1968; B. sekii (Yamaguti, 1937) Meserve, 1938; B, seriolae (Yamaguti, 1934) Meserve, 1938; and B. synagris Yamaguti, 1953. The type species, B. sciaenae, is redescribed based on new material from Australia. No types for this taxon were designated and we have assigned a series of voucher specimens. Tareenia acanthopagri Hussey, 1986 becomes B. acanthopagri (Hussey, 1986) comb. nov. and T. anticavaginata (Byrnes, 1986) Egorova, 1997 and T. lutjani (Whittington and Kearn, 1993) Egorova, 1997 are returned to Benedenia as B. anticavaginata and B. lutjani Benedenia akaisaki Iwata, 1990 is considered a synonym of B. ovata and B. kintoki Iwata, 1990 is considered a synonym of B. elongata. Two species, B, madai Ishii and Sawada, 1938 and B. pagrosomi Ishii and Sawada, 1938, are considered species inquirendae. Based on the redefinition of Benedenia, the diagnosis for the Benedeniinae is amended. Tareenia is synonymized with Benedenia but Menziesia Gibson, 1976 is recognized and its generic diagnosis amended to include: anterior attachment organs tending to form a 'hooded' appearance; prominent anterior gland cells between the pharynx and the anterior margin of the body: long penis, tapering proximally, occupying a penis canal with weakly muscular wall: penis canal and penis describe a sigmoid; accessory gland reservoir dorsal and alongside, or posterior and lateral to, proximal end of the penis and enclosed by a proximal extension of the wall of the penis canal. Under this conception. Menziesia comprises: M. noblei (Menzies. 1946) Gibson, 1976 (type species); M. malaboni (Velasquez. 1982) comb. nov.: M. merinthe (Yamaguti, 1968) Gibson. 1976: M. ovalis (Yamaguti, 1968) Gibson, 1976: and M. sebastodis (Yamaguti, 1934) comb, nov. A key to valid species of Benedenia and Menziesia is provided and a list is presented of published records of undescribed or unattributed species of Benedenia. Some protocols are suggested for preparation of benedeniine material to enhance future taxonomic studies and comparisons. The host-specificity and geographic distribution of species in these revised genera are discussed. The composition of the Capsalidae is discussed and some difficulties in defining and distinguishing between its different subfamilies are considered.

First published record of the pathogenic monogenean parasite Neobenenenia melleni (Capsalidae) from Australia

The monogenean Neobenedenia melleni (Mac- Callum, 1927) Yamaguti 1963 is a well-known and virulent pathogen in culture conditions recorded from the skin of many teleost fish species worldwide. Until now, N. melleni has not been reported from wild or cultured fish in Australian waters. This study documents a recent outbreak of N. melleni that occurred on Lates calcarifer (barramundi) cultivated in sea cages in Hinchinbrook Channel between Hinchinbrook Island and mainland Queensland, Australia, which resulted in the loss of 200 000 fish (50 tonnes). The origin of this outbreak is unclear because N. melleni has not been recorded from any wild host species in Australia and strict quarantine regulations exclude the possibility of its introduction on imported fish. We propose that N. melleni occurs naturally on wild populations of some teleost species in Australian waters and that the few surveys of wild fish conducted along the eastcoast have failed to report this species. The possibility that uncharacteristically low water temperatures led to the outbreak is discussed.

Experimental susceptibility of some reef fish species to Benedenia lutjani (Monogenea : Capsalidae), a parasite of Lutjanus carponotatus (Pisces : Lutjanidae)

The susceptibility of species of lutjanid, lethrinid and serranid fish to infection by either larval or post-larval (juvenile and adult) specimens of the capsalid monogenean Benedenia lutjani Whittington and Kearn (1993) was examined experimentally. Four species of lutjanids became infected when exposed to larvae of B. lutjani, but three species of lethrinids and four species of serranids were not susceptible to larvae under the same conditions. Variability in the intensity of infection by larvae occurred within and between lutjanid species. Few post-larval specimens of B. lutjani transferred between individuals of the specific host Lutjanus carponotatus (Richardson 1842) in 60-l aquaria and none transferred between specimens of L. carponotatus in a 7,500-l concrete tank. These results indicate that transfer of post-larval B. lutjani between individuals of the specific host is unlikely to occur in the wild. Other lutjanid species did not become infected when exposed to specimens of L. carponotatus infected heavily by post-larval B. lutjani, but two lethrinid species were susceptible to infection under the same conditions. These data indicate that different factors may mediate host-specificity for larval and post-larval B. lutjani.

Alguns trematódeos monogenéticos da família Capsalidae Baird, 1853, em peixes do Oceano Atlântico - Costa Continental Portuguesa e Costa do Norte da África

No presente trabalho é feito um estudo de cinco trematódeos monogenéticos pertencentes à família Capsalidae Baird, 1853. Para a subfamília Benedeniinae Johnston, 1931, foram criadas duas espécies pertencenteso ao gênero Benedenia Diesing, 1858: Benedenia pompatica sp. n., que muito se assemelha a Benedenia sciaenae (Beneden, 1852), diferindo desta principalmente pela estrutura do 2º e 3º pares de macroganchos e Benedenia innobilitata sp. n. que mais se aproxima de Benedenia jaliscana Bravo-Hollis, 1951, afastando-se desta espécie pela ausência do cirro armado e também pela estrutura e localização da vagina. Da subfamília Trochopodiane (Price, 1936) são apresentados novos hospedadores: Lepidotrigla cavillone para Trochopus pini (Beneden & Hesse, 1863) e Serranus cabrilla para Megalocotyle grandiloba Paperna e Kohn, 1964, sendo apresentadas com novas descrições e ilustrações. Ainda desta subfamília é criada uma nova espécie do gênero Allomegalocotyle (Robinson, 1961) Yamaguti, 1963. A. gabbari sp. n. diferencia-se de A. johnstoni (Robinson, 1961), única espécie do gênero, pela morfologia dos macroganchos, localização do poro vaginal e vesícula seminal bem desenvolvida e externa.

A preliminary phylogenetic analysis of the Capsalidae (Platyhelminthes : Monogenea : Monopisthocotylea) inferred from large subunit rDNA sequences

Phylogenetic relationships within the Capsalidae (Monogenea) were examined Using large subunit ribosomal DNA sequences from 17 capsalid species (representing 7 genera, 5 subfamilies), 2 outgroup taxa (Monocotylidae) plus Udonella caligorum (Udonellidae). Trees were constructed using maximum likelihood, minimum evolution and maximum parsimony algorithms. An initial tree, generated from sequences 315 bases long, Suggests that Capsalinae, Encotyllabinae, Entobdellinae and Trochopodinae are monophyletic, but that Benedeniinae is paraphyletic. Analyses indicate that Neobenedenia, currently in the Benedeniinae, should perhaps be placed in 2 separate subfamily. An additional analysis was made which omitted 3 capsalid taxa (for which only short sequences were available) and all outgroup taxa because of alignment difficulties. Sequence length increased to 693 bases and good branch support was achieved. The Benedeniinae was again paraphyletic. Higher-level classification of the Capsalidae, evolution of the Entobdellinae and issues of species identity in Neobenedenia are discussed.

Effects of temperature, salinity, desiccation and chemical treatments on egg embryonation and hatching success of Benedenia seriolae (Monogenea : Capsalidae), a parasite of farmed Seriola spp.

The effects of temperature and salinity on the embryonation period and hatching success of eggs of Benedenia seriolae were investigated. Temperature strongly influenced embryonation period; eggs first hatched 5 days after laying at 28 degreesC and 16 days after laying at 14 degreesC. The relationship between temperature and embryonation period is described by quadratic regression equations for time to first and last hatching. Hatching success was >70% for B. seriolae eggs incubated at temperatures from 14 to 28 degreesC. However, no B. seriolae eggs embryonated and hatched at 30 degreesC and

The effect of the cleaner fish Labroides dimidiatus on the capsalid monogenean Benedenia lolo parasite of the labrid fish Hemigymnus melapterus

The cleaner fish Labroides dimidiatus affected the pigmented monogenean parasite Benedenia lolo on the fish Hemigymnus melapterus (Labridae) held in aquaria. The effect of cleaner fish varied with the size class of fish; only small fish [a posteriori size class < 11-5 cm standard length (L-S)] exposed to cleaner fish had fewer monogeneans compared with fish not exposed to cleaner fish. The abundance of monogeneans on large fish (a posteriori size class > 11-5 cut L-S) was not affected by cleaner fish. The size-frequency distributions of monogeneans on both size-classes of H. melapterus were affected by cleaner fish. Fish exposed to cleaner fish had fewer large (> 3 mm) and more small (< 1 mm) monogeneans than fish not exposed to cleaner fish, suggesting cleaner fish selectively removed larger monogeneans. This difference was more pronounced on large fish. In the absence of cleaner fish, small fish had almost as many monogeneans as large fish; they also had more small monogeneans than the large fish, suggesting small fish were more vulnerable to infection by monogeneans than larger fish. This suggests that the cleaner fish L. dimidiatus has the potential to control benedeniine monogeneans on captive fish and highlights the importance of taking into account fish size in studies of the effect of cleaner fish on ectoparasites. (C) 2002 The Fisheries Society of the British Isles. Published by Elsevier Science Ltd. All rights reserved.

Preliminary characterisation and extraction of anterior adhesive secretion in monogenean (platyhelminth) parasites

Secreted anterior adhesives, used for temporary attachment to epithelial surfaces of fishes (skin and gills) by some monogenean (platyhelminth) parasites have been partially characterised. Adhesive is composed of protein. Amino acid composition has been determined for seven monopisthocotylean monogeneans. Six of these belong to the Monocotylidae and one species, Entobdella soleae (van Beneden et Hesse, 1864) Johnston, 1929, is a member of the Capsalidae. Histochemistry shows that the adhesive does not contain polysaccharides, including acid mucins, or lipids. The adhesive before secretion and in its secreted form contains no dihydroxyphenylalanine (dopa). Secreted adhesive is highly insoluble, but has a soft consistency and is mechanically removable from glass surfaces. Generally there are high levels of glycine and alanine, low levels of tyrosine and methionine, and histidine is often absent. However, amino acid content varies between species, the biggest differences evident when the monocotylid monogeneans were compared with E. soleae. Monogenean adhesive shows similarity in amino acid profile with adhesives from starfish, limpets and barnacles. However, there are some differences in individual amino acids in the temporary adhesive secretions of, on the one hand, the monogeneans and, on the other hand, the starfish and limpets. These differences may reflect the fact that monogeneans, unlike starfish and barnacles, attach to living tissue (tissue adhesion). A method of extracting unsecreted adhesive was investigated for use in further characterisation studies on monogenean glues.

Some monogenoidea parasitic on peruvian marine fishes, with description of Anoplocotyloides chorrillensis new species and new records

The presence of four Monogenoidea parasitic on marine fishes from the central Peruvian coast is recorded. One of them, Anoplocotyloides chorrillensis (Monocotylidae) described from the gills of Rhinobatos planiceps (Rhinobatidae) is considered a new species. The three other species are: Caballerocotyla autralis Oliva, 1986 (Capsalidae); Callorhynchocotyle marplatensis Suriano & Incorvaia, 1982 (Hexabothriidae) and Anoplocotyloides papillatus (Doran, 1953) (Monocotylidae) parasitic on Sarda chiliensis chiliensis (Scombridae), Callorhinchus callorhinchus (Callorhinchidae) and Rhinobatos planiceps (Rhinobatidae) respectively.

Monogenean parasitic on marine fishes from Perú and Chile: three new species and two new combinations

Five species of monogeneans collected from marine fishes of the Pacific coast of Chile and Perú were studied. Three of them are new species: Interniloculus chilensis n. sp. (Capsalidae), Neoheterobothrium insalaris n. sp. (Diclidophoridae) and Loxura peruensis n. sp. (Axinidae) parasitic on Sebastes capensis (Scorpaenidae), Paralichthys sp. (Bothiidae) and Belone scapularis (Belonidae), respectively. Two new combinations are proposed, Intracotyle neghmei (Microcotylidae) for Neobivagina neghmei Villalba, 1987 and Hargicotlyle conceptionensis (Diclidophoridae) for Choricotyle conceptionensis Villalba, 1987.

Caballerocotyla lenti n. sp., a capsalid monogenean from Auxis thazard (Scombridae) from off the southeastern coast of Brazil

Caballerocotyla lenti n. sp. (Monogenea: Capsalidae), recovered from the gills of Auxis thazard (Lacépède) captured off the coast of Rio de Janeiro, Brazil, is described using light and scanning electron microscopy. The new species is characterized by: a tegument with 2-5 rows of dorso-marginal, unicuspid spines; 53-54 round testes; a constricted pharynx with numerous papillae on its border; and a haptor with a plicate marginal border, a central polygonal area and seven complete septa. C. manteri (Price, 1951) and C. gouri Chauhan, 1953 sensu Murugesh (1995) are figured and commented upon.