New species of Dermoergasilus Ho & Do, 1982 (Copepoda: Cyclopoida: Ergasilidae) parasitizing endemic cichlid Paretroplus polyactis (Bleeker) in Madagascar.

Dermoergasilus madagascarensis n. sp. is described from the gills of Paretroplus polyactis, an endemic cichlid fish in Madagascar, using a combined morphological (light microscopy and SEM) and molecular approach (partial 18S rDNA, 28S rDNA, and COI sequences). The new species is characterized mainly by possessing: (i) roughly pentagonal cephalosome; (ii) antennal endopodal segments covered with slightly inflated membrane; (iii) maxillule bearing 2 equally long outer setae and a minute inner seta; (iv) interpodal sternites of swimming legs ornamented with 3­4 rows of spinules; (v) genital segment and first abdominal somite both barrel-shaped; and (vi) a caudal ramus projecting into a digitiform process with inconspicuous terminal seta and bearing 3 terminal setae. The obtained DNA sequences of Malagasy species represent the first molecular data for species of Dermoergasilus. The 28S rDNA phylogeny showed the affiliation of D. madagascarensis n. sp. to Ergasilidae and its sister relationship with cosmopolitan Ergasilus sieboldi von Nordmann, 1832. The first checklist for all species of Dermoergasilus is provided.

Species of Ergasilus von Nordmann, 1832 (Copepoda: Ergasilidae) from cichlid fishes in Lake Tanganyika.

Ergasilus (von Nordmann, 1832) (Ergasilidae) is a species-rich group of parasitic copepods with a wide distribution in freshwater, marine and brackish environments. Up to now, 9 species of Ergasilus are known from cichlid fishes in Africa. In this study, 5 species, including 3 new, were collected from the gills of 12 cichlid species (11 genera: Bathybates, Ctenochromis, Eretmodus, Gnathochromis, Lamprologus, Neolamprologus, Ophthalmotilapia, Perissodus, Simochromis, Spathodus and Tanganicodus) of the northeastern shore of Lake Tanganyika in Burundi, namely E. macrodactylus (Sars, 1909), E. megacheir (Sars, 1909), E. caparti n. sp., E. parasarsi n. sp. and E. parvus n. sp. All species found were identified and described on the basis of adult female specimens using an integrative taxonomy approach mixing morphological characterization and molecular analyses of 2 ribosomal DNA markers (partial 18S and 28S rDNA sequences). An identification key for Ergasilus species from Lake Tanganyika is included. This study provides the first molecular data for Ergasilus species in Africa. The phylogenetic analyses suggest that the Ergasilus species parasitizing Lake Tanganyikan cichlids form a well-supported clade within the Ergasilidae. However, their phylogenetic relationships with other congeners still remain unclear due to a lack of molecular data for this diverse genus.

Diversity, phylogeny and intraspecific variability of Paradiplozoon species (Monogenea: Diplozoidae) parasitizing endemic cyprinoids in the Middle East.

Diplozoidae are common monogenean ectoparasites of cyprinoid fish, with the genus Paradiplozoon being the most diversified. Despite recent studies on Diplozoidae from Europe, Africa and Asia, the diversity, distribution and phylogeny of this parasite group appears to be still underestimated in the Middle East. The objective of this study was to investigate the diversity, endemism and host specificity of diplozoids parasitizing cyprinoid fish from the Middle East, considering this region as an important historical interchange of fish fauna, and to elucidate the phylogenetic position of Middle Eastern Paradiplozoon species within Diplozoidae. Four Paradiplozoon species were collected from 48 out of 94 investigated cyprinoid species. Three known species, Paradiplozoon homoion, Paradiplozoon bliccae and Paradiplozoon bingolensis, were recorded on new cyprinoid host species, and a new species, Paradiplozoon koubkovae n. sp., was recorded on Luciobarbus capito and Capoeta capoeta from the Caspian Sea basin in Iran and Turkey. Paradiplozoon bliccae, exhibiting a wide host range in the Middle East, expressed both morphological and genetic intraspecific variabilities. The four Paradiplozoon species collected in the Middle East were placed in divergent clades, showing the rich evolutionary history of diplozoid parasites in the Middle East. Our study also revealed that two lineages of African diplozoids have a Middle Eastern origin. We stress the importance of applying an integrative approach combining morphological, ecological and molecular methods to reveal the real diversity of diplozoids.

Four new species of Dactylogyrus Diesing, 1850 (Monogenea: Dactylogyridae) parasitising the gills of northern Moroccan Luciobarbus Heckel (Cyprinidae): morphological and molecular characterisation.

Four new species of Dactylogyrus Diesing, 1850 are described from the gills of three species of Luciobarbus Heckel collected from various hydrographical basins in northern Morocco: Dactylogyrus scorpius n. sp. from Luciobarbus rifensis Doadrio, Casal-Lopez & Yahyaoui; D. benhoussai n. sp. from L. moulouyensis Pellegrin; and D. varius n. sp. and D. falsiphallus n. sp. from L. maghrebensis Doadrio, Perea & Yahyaoui. The descriptions of the new species are confirmed by molecular data (partial 18S rDNA, ITS1, and partial 28S rDNA sequences). All four species belong to the group of Dactylogyrus species, possessing a cross-shaped ventral bar and a male copulatory organ composed of a loosely coiled copulatory tube and an accessory piece with a capsule-like base and recurved distal portion. Given the high shape variability of the haptoral anchors reported among specimens of D. varius n. sp., three morphological forms within this species (D. varius f. vulgaris, D. varius f. magnus, and D. varius f. dromedarius) are recognised. However, specimens belonging to D. benhoussai n. sp. and D. varius f. vulgaris were morphologically very similar and were discriminated with certainty, only when using molecular data.

Annulotrema (Monogenea: Dactylogyridae) from the gills of African tetras (Characiformes: Alestidae) in Lake Turkana, Kenya, with descriptions of four new species and a redescription of A. elongata Paperna and Thurston, 1969.

Four new and four previously described species of Annulotrema were collected from the gills of four species (three genera, i.e. Alestes, Hydrocynus and Brycinus) of African tetras from Lake Turkana, Kenya: Annulotrema alestesnursi Paperna, 1973 from Brycinus nurse; Annulotrema ansatum n. sp., Annulotrema besalis Rehulková, Musilová and Gelnar, 2014, Annulotrema bipatens n. sp., Annulotrema cucullatum n. sp., Annulotrema nili Paperna, 1973, and Annulotrema pontile n. sp. from Hydrocynus forskahlii; and Annulotrema elongata Paperna and Thurston, 1969 from Alestes baremoze and Alestes dentex. A. elongata is re-described on the basis of new material from A. baremoze. The sclerotized structures of the haptor and male copulatory organ of A. alestesnursi and A. elongata are illustrated from their type material. H. forskahlii is a new host record for A. besalis. The findings of A. besalis and A. elongata in Kenya represent a new locality records for these helminths. Three Annulotrema spp., namely A. besalis, A. elongata and A. pontile n. sp., share the same type of male copulatory organ, which may indicate a close relationship among these species.

Annulotrema (Monogenea: Dactylogyridae) from Hydrocynus brevis (Characiformes: Alestidae) in Senegal, with descriptions of two new species and remarks on Annulotrema pikei.

Two new species of Annulotrema Paperna & Thurston, 1969 were collected from the gills of the African tiger fish, Hydrocynus brevis, from the Gambia River basin in the Niokolo-Koba National Park, Senegal. Annulotrema besalis n. sp. is characterized by having a male copulatory organ (MCO) composed of an arcuate copulatory tube articulated to an eight-shaped accessory piece with terminal claw. The new species resembles Annulotrema pikei (Price, Peebles & Bamford, 1969) in having morphologically similar types of haptoral sclerites and MCO. As a result of the differential diagnosis made for A. besalis n. sp., new information on taxonomically important features of A. pikei is provided based on illustrations of the sclerotized parts of the holotype from Hydrocynus vittatus. The report of A. pikei on the gills of Hydrocynus forskahlii by Paperna in 1979 is shown to be erroneous. Annulotrema uncata n. sp. is similar to Annulotrema alestesimberi Paperna, 1973 in its possession of a coiled copulatory tube with about two and a half rings. Features distinguishing the new species include the sharply curved shaft of the ventral anchor, the base of the copulatory tube extending to a sock-like structure and a leech-shaped vagina. The necessity of emending the generic diagnosis of Annulotrema is briefly discussed.

Two new species of Cichlidogyrus (Monogenea: Dactylogyridae) parasitizing the gills of African cichlid fishes (Perciformes) from Senegal: morphometric and molecular characterization.

Two new species of Cichlidogyrus are described from the gills of three species of African cichlids collected from the Gambia River basin in the Niokolo-Koba National Park, Senegal: Cichlidogyrus dracolemma n. sp. from Hemichromis letourneuxi and Cichlidogyrus nageus n. sp. from Sarotherodon galilaeus and Tilapia guineensis. The new species are recognized on the basis of morphological differences in the male copulatory organs (MCOs) and apparent differences in the SSU rDNA and ITS1 sequence data. C. dracolemma n. sp. resembles specimens of Cichlidogyrus from Hemichromis bimaculatus, probably misidentified as Cichlidogyrus bychowskii by Paperna in 1965, in having morphologically similar types of haptoral sclerites and MCO. The new species is characterized by possessing an MCO with a kite-shaped termination of the accessory piece and a vagina distally surrounded by a circular superficial sclerotization of the body surface. Molecular phylogenetic analyses showed a strongly supported clade including C. dracolemma n. sp. and two species collected from Hemichromis fasciatus, namely Cichlidogyrus longicirrus Paperna, 1965 and Cichlidogyrus falcifer Dossou and Birgi, 1984. Based on both morphology and molecular phylogeny, C. nageus n. sp. appears to be closely related to Cichlidogyrus acerbus Dossou, 1982 obtained from S. galilaeus. Morphologically, C. nageus n. sp. is readily separated from all known congeners by the characteristic pestle shaped termination of the accessory piece of the MCO. Molecular phylogenetic reconstruction showed that C. dracolemma n. sp. and C. nageus n. sp. cluster with the Cichlidogyrus species possessing the same haptoral configuration and host preferences (on subfamily and tribe level, respectively).

Nearctic Dactylogyrus species (Platyhelminthes, Monogenea) parasitizing cypriniform fishes in the context of morphology and phylogeny, with descriptions of seven new species.

Ribosomal DNA sequences are currently available for 32 morphologically recognized species of Dactylogyrus parasitizing Nearctic cypriniforms, but only 16 of them are assigned to nominal species. Herein, morphological data on 28 of the 32 species are provided, together with comments on their phylogenetic relationships in the context of the morphology of taxonomically important structures. Seven new species of Dactylogyrus are described from five species of leuciscids and one species of catostomids, as follows: D. aduncus n. sp. from Campostoma spadiceum, D. cloutmani n. sp. from Luxilus chrysocephalus isolepis, D. cornifrons n. sp. from Cyprinella venusta, D. fimbratus n. sp. from Rhinichthys cataractae, D. mcallisteri n. sp. from Semotilus atromaculatus, and D. chieni n. sp. and D. haneki n. sp. from Hypentelium nigricans. Four previously described species of Dactylogyrus, D. atromaculatus from S. atromaculatus, D. eos from C. neogaeus, D. parvicirrus from Notemigonus crysoleucas and D. perlus from L. c. isolepis, are redescribed and/or figured. As for the remaining 17 species, only the male copulatory organs (MCOs) taken from the respective hologenophores are illustrated. On the basis of phylogenetic analyses, two main clades of Nearctic Dactylogyrus were recognized and supported by the different morphology of the MCO. The first one included 22 strictly Nearctic species sharing the same MCO type with Dactylogyrus spp. parasitizing cyprinids likely of Asian origin. The second clade encompassed Dactylogyrus spp. with diverse MCO morphology and was placed in the sister position to Dactylogyrus spp. parasitizing European leuciscids and North-West African cyprinids.

Title: Espèce néarctique de Dactylogyrus (Platyhelminthes, Monogenea) parasitant des poissons cypriniformes dans un contexte de morphologie et phylogénie, avec description de sept nouvelles espèces. Abstract: Des séquences d'ADN ribosomal sont actuellement disponibles pour 32 espèces morphologiquement reconnues de Dactylogyrus parasitant les cypriniformes néarctiques, mais seulement 16 d'entre elles sont attribuées à des espèces nominales. Ici, des données morphologiques sur 28 des 32 espèces sont fournies avec des commentaires sur leurs relations phylogénétiques dans le contexte de la morphologie des structures taxonomiquement importantes. Sept nouvelles espèces de Dactylogyrus sont décrites à partir de cinq espèces de leuciscidés et d'une espèce de catostomidés, comme suit : D. aduncus n. sp. de Campostoma spadiceum, D. cloutmani n. sp. de Luxilus chrysocephalus isolepis, D. cornifrons n. sp. de Cyprinella venusta, D. fimbratus n. sp. de Rhinichthys cataractae, D. mcallisteri n. sp. de Semotilus atromaculatus et D. chieni n. sp. et D. haneki n. sp. de Hypentelium nigricans. Quatre espèces de Dactylogyrus précédemment décrites, D. atromaculatus de S. atromaculatus, D. eos de C. neogaeus, D. parvicirrus de Notemigonus crysoleucas et D. perlus de L. c. isolepis, sont redécrites et/ou figurées. Comme pour les 17 espèces restantes, seuls les organes copulateurs mâles (OCM) prélevés sur les hologénophores respectifs sont illustrés. Sur la base d'analyses phylogénétiques, deux clades principaux de Dactylogyrus néarctiques ont été reconnus et soutenus par des morphologies différentes de l'OCM. Le premier comprenait 22 espèces strictement néarctiques partageant le même type d'OCM que les Dactylogyrus spp. parasitant des cyprinidés probablement d'origine asiatique. Le deuxième clade comprenait des Dactylogyrus spp. avec diverses morphologies d'OCM et était placé en position sœur des Dactylogyrus spp. parasitant les leuciscidés européens et les cyprinidés d'Afrique du Nord-Ouest.

Host-Specific Parasites Reveal the History and Biogeographical Contacts of Their Hosts: The Monogenea of Nearctic Cyprinoid Fishes.

Host-specific parasites exhibit close co-evolutionary associations with their hosts. In the case of fragmented/disjunct host distribution, host-specific parasites may reflect the biogeographical history of regions and/or the role played by contacts of hosts. The present study was focused on Dactylogyrus (Monogenea) species almost exclusively parasitizing cyprinoid fishes. We investigated the phylogenetic relationships between Dactylogyrus parasites of Nearctic cyprinoids (Leuciscidae) and Dactylogyrus parasites of Palearctic cyprinoids and used Dactylogyrus phylogeny to explore the biogeography of fish hosts in Europe and North America. Phylogenetic analyses revealed that two Nearctic clades of Dactylogyrus spp. have different origins. Historical contacts between European and North American leuciscids were accompanied by the host switching of Dactylogyrus species. In the Nearctic region, Dactylogyrus parasites also colonized non-leuciscid fishes. Dactylogyrus spp. of three Nearctic leuciscid clades were included in the phylogenetic reconstruction; only Dactylogyrus spp. of the Plagopterinae had a common origin. Dactylogyrus species did not reflect the phylogenetic relationships among leuciscid clades, suggesting that past co-diversification was overshadowed by colonization events mediated by paleogeographic and climatological changes and extensive drainage reorganization. Host-specific monogeneans serve as a supplementary tool to reveal the historical biogeographical contacts between freshwater fish from the North America and Europe and also contemporary contacts of leuciscids in North America.

Integrating morphological and molecular approaches for characterizing four species of Dactylogyrus (Monogenea: Dactylogyridae) from Moroccan cyprinids, with comments on their host specificity and phylogenetic relationships.

Cyprinid fishes are known to harbour highly host-specific gill-associated parasites of Dactylogyrus. High similarity in the morphology of sclerotized structures among Dactylogyrus species, especially those parasitizing congeneric cyprinoids, makes their identification difficult. In this paper, four previously known species of Dactylogyrus are characterized and illustrated under a reliable taxonomic framework integrating morphological and molecular evidence, and their phylogenetic relationships are investigated using molecular data. The species are as follows: D. borjensis from Luciobarbus zayanensis; D. draaensis from Luciobarbus lepineyi; D. ksibii from Luciobarbus ksibi and Luciobarbus rabatensis; and D. marocanus from Carasobarbus fritschii, L. ksibi, L. zayanensis and Pterocapoeta maroccana. Our results revealed intraspecific genetic variability among specimens of D. ksibii collected from two different hosts and geographically distant basins. Phylogenetic reconstruction showed that Dactylogyrus spp. parasitizing Moroccan cyprinids are representatives of three main lineages corresponding to morphological differences and host specificity. Our records of D. marocanus on L. zayanensis and P. maroccana increase the range of available host species i.e.,eight species of four cyprinid genera representing two phylogenetic lineages (i.e., Barbinae and Torinae).

Five new monogenean species from the gills of Mulloidichthys vanicolensis (Perciformes: Mullidae) off New Caledonia, with the proposal of Volsellituba n. g. and Pennulituba n. g. (Monogenea: Dactylogyridae).

Three species of Volsellituba n. g. and two species of Pennulituba n. g. are described from the gills of the yellowfin goatfish Mulloidichthys vanicolensis (Valenciennes) off New Caledonia (South Pacific). Volsellituba and Pennulituba resemble dactylogyrid genera whose species lack eye-spots and possess tandem or slightly overlapping gonads, a single prostatic reservoir, a male copulatory organ without an accessory piece, a dextroventral non-sclerotised vagina, ventral and dorsal anchor/bar complexes, and hooks with shanks comprising a single slender unit. Volsellituba n. g. is characterised by its species having ventral tegumental folds at the level of the vaginal duct, a male copulatory organ with a subterminal basal opening and a two-piece dorsal bar. The following new species of Volsellituba are described: V. orchidea n. sp. (type-species), V. nabla n. sp. and V. elephantina n. sp. Pennulituba n. g. is distinguished by its species possessing a male copulatory organ with a wing-shaped membrane and a dorsal bar with a poorly to non-sclerotised medial part. This genus includes P. piratifalx n. sp. (type-species) and P. cymansis n. sp. A tabular summary of the species of Haliotrema Johnston & Tiegs, 1922 reported from mullids is provided.

Uncovering the diversity of monogeneans (Platyhelminthes) on endemic cypriniform fishes of the Balkan Peninsula: new species of Dactylogyrus and comments on their phylogeny and host-parasite associations in a biogeographic context.

Seven new species of Dactylogyrus Diesing, 1850 (Dactylogyridae) are described from the gills of seven endemic species of cyprinoids (Cyprinidae, Leuciscidae) inhabiting the Balkan Peninsula: Dactylogyrus romuli n. sp. from Luciobarbus albanicus (Greece), Dactylogyrus remi n. sp. from Luciobarbus graecus (Greece), Dactylogyrus recisus n. sp. from Pachychilon macedonicum (Greece), Dactylogyrus octopus n. sp. from Tropidophoxinellus spartiaticus (Greece), Dactylogyrus vukicae n. sp. from Delminichthys adspersus (Bosnia and Herzegovina), Dactylogyrus leptus n. sp. from Chondrostoma knerii (Bosnia and Herzegovina), and Dactylogyrus sandai n. sp. from Telestes karsticus (Croatia). To delineate species boundaries, we used an integrative taxonomic approach combining morphological and genetic data. With these tools, we illustrate that some species of monogeneans considered as cryptic might be designated as pseudocryptic (morphologically similar, not easily differentiated) after a posteriori detailed morphological examination, as happened with D. romuli n. sp. and D. remi n. sp. Thus, for accurate species characterization, it is particularly important to acquire both morphological and molecular data from the same individual specimens, ideally along with illustrations of taxonomically important structures directly taken from hologenophores. Using phylogenetic reconstruction, we investigated the phylogenetic position of newly described Dactylogyrus species within Dactylogyrus species from Balkan cyprinoids with regard to morphological characteristics, host range, and geographical distribution.

TITLE: Découverte de la diversité des Monogènes (Plathelminthes) des poissons cypriniformes endémiques de la péninsule balkanique : nouvelles espèces de Dactylogyrus et commentaires sur leur phylogénie et les associations hôte-parasite dans un contexte biogéographique. ABSTRACT: Sept nouvelles espèces de Dactylogyrus Diesing, 1850 (Dactylogyridae) sont décrites à partir des branchies de sept espèces endémiques de cyprinoïdés (Cyprinidae, Leuciscidae) habitant la péninsule balkanique : Dactylogyrus romuli n. sp. de Luciobarbus albanicus (Grèce), Dactylogyrus remi n. sp. de Luciobarbus graecus (Grèce), Dactylogyrus recisus n. sp. de Pachychilon macedonicum (Grèce), Dactylogyrus octopus n. sp. de Tropidophoxinellus spartiaticus (Grèce), Dactylogyrus vukicae n. sp. de Delminichthys adspersus (Bosnie-Herzégovine), Dactylogyrus leptus n. sp. de Chondrostoma knerii (Bosnie-Herzégovine) et Dactylogyrus sandai n. sp. de Telestes karsticus (Croatie). Pour la délimitation des espèces, nous avons utilisé une approche taxonomique intégrative combinant des données morphologiques et génétiques. Avec ces outils, nous illustrons que certaines espèces de Monogènes considérées comme cryptiques pourraient être désignées comme pseudocryptiques (morphologiquement similaires, pas facilement différenciées) après un examen morphologique détaillé a posteriori, comme cela s'est produit avec D. romuli n. sp. et D. remi n. sp. Ainsi, pour une caractérisation précise des espèces, il est particulièrement important d'acquérir des données morphologiques et moléculaires à partir des mêmes spécimens individuels, idéalement avec des illustrations de structures taxonomiquement importantes directement prises à partir d'hologénophores. En utilisant la reconstruction phylogénétique, nous avons étudié la position phylogénétique des espèces de Dactylogyrus nouvellement décrites au sein des espèces de Dactylogyrus des cyprinoïdés des Balkans en ce qui concerne les caractéristiques morphologiques, la gamme d'hôtes et la distribution géographique.

Diversity of monogeneans and tapeworms in cypriniform fishes across two continents.

Cypriniformes, which exhibit a wide geographical distribution, are the most species-rich group of freshwater fishes. Despite considerable research on their parasites, no reliable estimates of their parasite diversity on a large geographical scale are available. In the present review, we analyse species richness of two parasitic flatworm groups (monogeneans and tapeworms) reported from cypriniform fishes in the two most intensively studied parts of the Holarctic region, Europe and North America. We also review knowledge on parasite speciation and host-parasite coevolution, and emphasise the risk of parasite co-introduction resulting from transfers of cypriniforms among different continents. As parasite diversity in European cypriniforms has been more intensively explored, we predicted a lower level of knowledge on parasite diversity in North American fishes, despite North America having a higher diversity of cypriniforms than Europe. Our data revealed a higher mean species richness of monogeneans and tapeworms per cypriniform species in Europe compared with North America. We showed that species richness of both parasite taxa in both continents is strongly affected by sample size, but that fish traits also play an important role in determining monogenean and tapeworm species richness in European cyprinoids. We recorded higher host specificity for cypriniform parasites in North America, even within parasite genera shared by cypriniforms on both continents. The host range of monogeneans parasitising cyprinoids on both continents was affected by phylogeny, indicating an effect of parasite life history on host specificity. The difference in parasite host range between the two continents could potentially be explained by either the low overall level of sampling activity in North America or an underestimation of parasite diversity in Europe. We suggest that future research efforts be focussed on cypriniforms in order to obtain reliable data for robust assessments of parasite species richness and phylogenies, to assess host-parasite coevolution and to reveal fish biogeography.

Species of Characidotrema Paperna & Thurston, 1968 (Monogenea: Dactylogyridae) from fishes of the Alestidae (Characiformes) in Africa: new species, host-parasite associations and first insights into the phylogeny of the genus.

BACKGROUND: African tetras (Alestidae) belonging to Brycinus Valenciennes are known to be parasitized with monogeneans attributed to two genera, Annulotrema Paperna & Thurston, 1969 and Characidotrema Paperna & Thurston, 1968 (Dactylogyridae). During a survey of monogeneans parasitizing alestids, species of Characidotrema were collected in Cameroon, D. R. Congo, Senegal, South Africa, Sudan and Zimbabwe. This paper provides new morphological data and the first molecular analysis broadening our knowledge on the diversity of these parasites. RESULTS: Seven species (four known and three new) of Characidotrema are reported from two species of Brycinus: C. auritum n. sp. and C. vespertilio n. sp. from B. imberi (Peters); and C. brevipenis Paperna, 1969, C. nursei Ergens, 1973, C. pollex n. sp., C. spinivaginus (Paperna, 1973) and C. zelotes Kritsky, Kulo & Boeger, 1987 from B. nurse (Rüppell). Species identification was based on morphological analysis of the sclerotized structures supported by nuclear ribosomal DNA (partial 18S rDNA, ITS1, and 28S rDNA) sequence data. Morphological analysis confirmed that the most apparent character distinguishing species in the genus is the morphology of the male copulatory organ and vagina. Observations on the haptoral sclerotized elements of these parasites by means of phase contrast microscopy revealed the presence of a sheath-like structure relating to the ventral anchor, a feature that supplements the generic diagnosis of Characidotrema. Maximum Likelihood and Bayesian analyses of the large subunit (28S) rDNA sequences recovered Characidotrema species isolated from the two Brycinus hosts as monophyletic, and indicated a closer relationship of this group to monogeneans parasitizing African cyprinids (Dactylogyrus spp.) and cichlids (species of Cichlidogyrus Paperna, 1960, Scutogyrus Pariselle & Euzet, 1995, and Onchobdella Paperna, 1968) than to those from catfishes (species of Quadriacanthus Paperna, 1961, Schilbetrema Paperna & Thurston, 1968 and Synodontella Dossou & Euzet, 1993). The overall agreement between the morphological diversification of the MCOs and the molecular tree observed in this study indicates that significant phylogenetic signals for clarifying relationships among species of Characidotrema are present in the characteristics of the MCO. CONCLUSIONS: It seems that intra-host speciation is an important force shaping the present distribution and diversity of Characidotrema but further studies are necessary to confirm this hypothesis and assess questions related to the phylogeny of these parasites. To identify potential co-speciation events, co-phylogenetic analyses of these monogeneans and their alestid hosts are required.

Quadriacanthus species (Monogenea: Dactylogyridae) from catfishes (Teleostei: Siluriformes) in eastern Africa: new species, new records and first insights into interspecific genetic relationships.

BACKGROUND: African catfishes of the families Bagridae and Clariidae are known to be parasitized with monogeneans of Quadriacanthus Paperna, 1961 (Dactylogyridae). The genus remains taxonomically challenging due to its speciose nature and relatively wide host range representing two fish orders, i.e. Siluriformes and Osteoglossiformes, in Africa and Asia. Here, we investigated diversity of Quadriacanthus spp. parasitizing Clarias gariepinus (Burchell), Heterobranchus bidorsalis Geoffroy Saint-Hilaire, and Bagrus docmak (Forsskål) collected in the Lake Turkana (Kenya) and Nile River Basin (Sudan). The interspecific relationships among Quadriacanthus spp. parasitizing catfishes inferred from ribosomal DNA sequences were investigated for the first time. METHODS: A combined morphological and molecular approach was used for description of the new species and for a critical review of the previously described Quadriacanthus spp., by means of phase contrast microscopic examination of sclerotized structures, and assessing the genetic divergence among the species found using rDNA sequences. RESULTS: Seven species (including four new) of Quadriacanthus were identified. These were as follows: Quadriacanthus aegypticus El-Naggar & Serag, 1986, Quadriacanthus clariadis Paperna, 1961, Quadriacanthus fornicatus n. sp., Quadriacanthus pravus n. sp., and Quadriacanthus zuheiri n. sp. from Clarias gariepinus (Clariidae); Quadriacanthus mandibulatus n. sp. from Heterobranchus bidorsalis (Clariidae); and Quadriacanthus bagrae Paperna, 1979 from Bagrus docmak (Bagridae). For both 18S-ITS1 and 28S rDNA regions, Q. clariadis from a clariid fish was found to be most closely related to Q. bagrae from a bagrid host. Quadriacanthus mandibulatus n. sp. was observed to be the most distant species from the others. The separation of Q. mandibulatus n. sp. from the other species corresponds with the different morphology of its copulatory tube. The copulatory tube is terminally enlarged in Q. mandibulatus n. sp., while the tube in all other congeners studied is comparatively small and with an oblique tapering termination. CONCLUSIONS: This study contributes to a better understanding of African dactylogyrid diversity and provides the first molecular characterization of Quadriacanthus spp. The observed interspecific genetic relationships among Quadriacanthus spp. from clariids and Q. bagrae from a bagrid host suggest a possible host-switching event in the evolutionary history of the genus. Our records extend the currently known geographical range for Quadriacanthus spp. to Kenya and Sudan.

A revised diagnosis of Thylacicleidus (Monogenea: Dactylogyridae) with a redescription of the type species, Thylacicleidus serendipitus, and descriptions of two new species from Southeast Asian pufferfishes (Tetraodontiformes: Tetraodontidae).

Two new and 1 previously described species of Thylacicleidus Wheeler and Klassen, 1988, are reported from 2 species of brackish to freshwater puffers (Tetraodontidae) from Thailand. The generic diagnosis of Thylacicleidus is emended to incorporate the following characters: presence of a dextral muscular vagina, a vas deferens looping the left intestinal cecum, 2 prostatic reservoirs, a sclerotized tubular copulum without an accessory piece, and dorsal anchors with reduced outer root. The type species, Thylacicleidus serendipitus, is redescribed based on new information obtained from specimens found on the gills of the type host, Tetraodon nigroviridis Marion de Procé, 1822 (spotted green puffer). Two new species, Thylacicleidus latus n. sp. and Thylacicleidus brunensis n. sp., are described from the gills of T. nigroviridis and Tetraodon biocellatus Tirant, 1885 (eyespot puffer), respectively. The occurrence of common generic characters in species of Thylacicleidus and Haliotrema Johnston and Tiegs, 1922, another dactylogyrid genus recorded from tetraodontiform hosts, signifies the close relationship between the 2 genera and raises the question of synonymy. Nevertheless, because of morphological variability within Haliotrema spp., extremely wide host spectrum and the probable polyphyly of the genus, it is suggested that Thylacicleidus is a valid genus.

Three new species of Dactylogyrus Diesing, 1850 (Monogenea: Dactylogyridae) from the gills of the bala sharkminnow Balantiocheilos melanopterus (Cyprinidae) from Thailand.

Three new species of Dactylogyrus are described from the gills of the cyprinid fish Balantiocheilos melanopterus (Bleeker) imported from Thailand via the aquarium trade. They are D. coartatus n. sp., D. macrocolpius n. sp. and D. melanopteri n. sp. The double-bar species, D. coartatus n. sp. and D. melanopteri n. sp., are characterised by hooks that are markedly large in relation to the size of the anchors. Clip-like anchors with a sharply recurved, long point are a unique feature of D. coartatus n. sp. The single-bar species, D. macrocolpius n. sp., is characterised by having a copulatory organ composed of a spiral copulum and a medial accessory piece with thorns.