Dendromonocotyle species (Monogenea: Monocotylidae) are the only monocotylids to parasitize the skin of chondrichthyan hosts. Currently 11 species are recorded from the skin of ray species in the Dasyatidae, Myliobatidae and Urolophidae. There have been increasing reports of Dendromonocotyle outbreaks on rays kept in public aquaria. This paper provides a broad review of Dendromonocotyle that should assist taxonomists and aquarists with species identification and help decisions on potential control methods for Dendromonocotyle infections. The taxonomy and host-specificity of Dendromonocotyle are discussed and a key to current species is provided. We summarise what little is known about the biology of Dendromonocotyle including egg embryonation and hatching, feeding, camouflage and reproduction. The efficacy of freshwater baths, chemical treatments and biological control measures such as the use of cleaner fish for Dendromonocotyle is also discussed. We demonstrate that effective control of Dendromonocotyle on captive rays is hampered by the lack of basic biological data on the life cycle of the parasites. A case history is provided outlining the success of a public aquarium (Underwater World, Mooloolaba, Queensland, Australia) in managing D. pipinna infections on captive Taeniura meyeni without chemical intervention simply by taking measures to reduce host stress.
Over a 7-year period, parasites have been collected from 28 species of groupers (Serranidae, Epinephelinae) in the waters off New Caledonia. Host-parasite and parasite-host lists are provided, with a total of 337 host-parasite combinations, including 146 parasite identifications at the species level. Results are included for isopods (5 species), copepods (19), monogeneans (56), digeneans (28), cestodes (12), and nematodes (12). When results are restricted to those 14 fish species for which more than five specimens were examined and to parasites identified at the species level, 109 host-parasite combinations were recorded, with 63 different species, of which monogeneans account for half (32 species), and an average of 4.5 parasite species per fish species. Digenean records were compared for 16 fish species shared with the study of Cribb et al. (2002); based on a total of 90 parasite records identified at the species level, New Caledonia has 17 new records and only seven species were already known from other locations. We hypothesize that the present results represent only a small part of the actual biodiversity, and we predict a biodiversity of 10 different parasite species and 30 host-parasite combinations per serranid. A comparison with a study on Heron Island (Queensland, Australia) by Lester and Sewell (1989) was attempted: of the four species of fish in common and in a total of 91 host-parasite combinations, only six parasites identified at the species level were shared. This suggests strongly that insufficient sampling impairs proper biogeographical or ecological comparisons. Probably only 3% of the parasite species of coral reef fish are already known in New Caledonia.
Branchotenthes octohamatus sp. n. (Monogenea: Hexabothriidae) is described from the gills of the southern fiddler ray, Trygonorrhina fasciata Müller et Henle (Elasmobranchii: Rhinobatidae), off Adelaide, South Australia. It is distinguished from the type species, Branchotenthes robinoverstreeti Bullard et Dippenaar, 2003, by producing eggs that are joined end to end forming a chain, in the morphology of the male copulatory organ that has a pronounced constriction in duct diameter between proximal and distal regions, the possession of a thin muscular layer surrounding the proximal part of the male copulatory organ and distal region of the vaginae, and by the absence of a raised process on the shaft of the hamulus. An amended generic diagnosis is provided and the reliability of sperm duct number as a generic character is discussed. The oncomiracidium of B. octohamatus is also described and is the first monogenean to be described with only eight hooklets in the larval haptor. This discovery of eight hooklets may be important for higher-level monogenean evolutionary hypotheses.
Dendromonocotyle colorni sp. n. (Monogenea: Monocotylidae) is described from the dorsal skin surface of two specimens of Himantura uarnak (Forsskål) kept at the Eilat Underwater Observatory in Israel. Dendromonocotyle colorni is distinguished from the other eight species in the genus by the morphology of the terminal papillar sclerite on the haptor, the distal portion of the male copulatory organ and the morphology of the vagina. The development of the male copulatory organ is detailed for D. colorni and the adaptations of species of Dendromonocotyle to life on the dorsal skin surface of rays are discussed. Dendromonocotyle octodiscus Hargis, 1955 was identified from the dorsal skin surface of the southern stingray Dasyatis americana Hildebrand et Schroeder off Bimini, Bahamas and represents a new host record.
Empruthotrema quindecima sp. n. (Monogcnea: Monocotylidae) is described from the nasal fossae of the blue-spotted fantail ray Taeniura lymma (Forsskâl, 1775) collected from the reef flats of Heron Island and Lizard Island located at the southern and northern sections, respectively, of the Great Barrier Reef in Queensland, Australia. Empruthotrema quindecima has 15 marginal loculi on the haptor which distinguishes it from the other five members of the genus which have either 13 or 14 marginal loculi. The generic diagnosis of Empruthotrema Johnston ct Ticgs, 1922 is amended to accommodate the new species, a key to species is provided and the evolution of the different configurations of the haptoral loculi within the genus is discussed.
Heliocotyle ewingi sp. n. (Monogenea: Monocotylidae) is described from the gills of Myliobatis australis Macleay, 1881 (Myliobatididae) collected from Norfolk Bay near Hobart, Tasmania, Australia. Heliocotyle ewingi can be distinguished readily from the only other species in the genus, Heliocotyle kartasi Neifar, Euzet et Ben Hassine, 1999, by the presence of a single pseudoseptum on each of the peripheral loculi except the posteriormost, eyespots and by the morphology of the male copulatory organ which is a short, straight sclerotised tube which lacks a sclerotised accessory piece. The generic diagnosis is revised to accommodate the new species and the anterior glands are discussed.
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.
The Capsalidae are monogeneans parasitizing ''skin'', fins and gills of marine fishes. Some capsalids are pathogenic to cultivated fish and a few have caused epizootic events. It is a cosmopolitan family with broad host associations (elasmobranchs and teleosts, including sturgeons). Approximately 200 capsalid species are placed in nine subfamilies and 44-46 genera, some of which are well known (Benedenia, Capsala, Entobdella, Neobenedenia). Sturgeons host two capsalid species (Nitzschiinae) and 15 species in five genera are reliably reported from elasmobranchs. The combination of ancient (shark, ray, sturgeon) and modern (teleost) host fish lineages indicates that capsalid evolution is likely a blend of coevolution and host-switching, but a family phylogeny has been lacking due to deficient knowledge about homologies. The current phenetic subfamilial classification is discussed in detail using a preliminary phylogeny generated from large subunit ribosomal DNA sequence data from representatives of five subfamilies. Monophyly of the Capsalidae is supported by possession of accessory sclerites. Hypotheses are proposed for the possible radiation of capsalids. A suggestion that Neobenedenia melleni, a pathogenic species atypical due to its broad host-specificity (>100 host teleost species from >30 families in five orders), may be a complex of species is supported from genetic evidence. This may explain peculiarities in biology, taxonomy, host associations and geographic distribution of N. 'melleni' and has implications for fish health. Holistic studies using live and preserved larval and adult capsalid specimens and material for genetic analysis are emphasised to further determine identity, phylogeny and details of biology, especially for pathogenic species.
The elongated encased spermatophores of the capsalid (entobdelline) monogeneans Neoentobdella diadema (Monticelli, 1902) Kearn et Whittington, 2005 and N. apiocolpos (Euzet et Maillard, 1967) Kearn et Whittington, 2005 have been found attached by their proximal ends to the region of the vaginal opening, with the bulk of the spermatophore projecting from the vagina and therefore lying outside the body. In spite of previous reports, no spermatophores were found projecting from the common genital opening and if spermatophore exchange is as rapid as it is in the related entobdelline Entobdella soleae, then the chances of finding a spermatophore in this location are small. In N. diadema and N. apiocolpos it is likely that sperm enters the vagina through the open proximal end of an attached spermatophore, after which the empty spermatophore case is probably discarded. There is no evidence for a previous proposal that the whole spermatophore is engulfed by the vagina followed by digestion of the case to release the sperm. Three specimens of N. diadema were found each with two spermatophore cases projecting from the vagina and a specimen of N. apiocolpos carried three cases. Assuming that each parasite is able to donate or receive only one spermatophore at each mating, then the presence of one spermatophore does not prevent a further mating and acceptance of a fresh spermatophore. In spite of differences between the spermatophores of E. soleae and N. diadema/N. apiocolpos, the events of spermatophore exchange may be similar.
Myliocotyle borneoensis sp. n. and M. multicrista sp. n. (Monocotylidae: Heterocotylinae) are described from the gills of the mottled eagle ray, Aetomylaeus maculatus (Gray), and the banded eagle ray A. nichofii (Bloch et Schneider) (Myliobatidae), respectively, collected from the northern coast of Malaysian Borneo. These are the first monogeneans to be described on elasmobranchs from Borneo. The formerly monotypic Myliocotyle (for M. pteromylaei) was distinguished from other monocotylids by the distribution and morphology of the eight sclerotised dorsal haptoral accessory structures and the morphology of the male copulatory organ. However, we have determined that M. pteromylaei has ten structures on the dorsal surface of the haptor. Myliocotyle borneoensis is distinguished from M. pteromylaei by the morphology of the male copulatory organ and its accessory piece. Myliocotyle multicrista has 12 sclerotised dorsal haptoral accessory structures and a male copulatory organ with two accessory pieces. Additional sclerotised ridges across the ventral surfaces of each loculus (except the posterior-most pair) are also present in M. multicrista. The diagnosis for Myliocotyle is revised given our discovery of additional dorsal haptoral accessory structures in the type species and to accommodate other new characters of the two new species. Anterior secretions of Myliocotyle are discussed.