Monogènes Dactylogyridae parasites de Cyprinidae du genreBarbus d'Afrique du Nord

Resumé Quinze nouvelles espèces de Monogènes Dactylogyridae sont décrites chez quinze espèces deBarbus (Teleostei, Cyprinidae) appartenant aux sous-genresB. (Barbus) etB. (Labeobarbus) en Afrique du Nord. Les barbeaux examinés proviennent des différents bassins hydrographiques du Maroc et d'une localité nommée Hamman Bourgiba en Tunisie. Dans cette dernière région, le genreBarbus n'est représenté que par une seule espèce:Barbus (B.) callensis. Au Maroc, on en dénombre actuellement quatorze dont quatre appartiennent au sous-genreLabeobarbus: il s'agit deBarbus (L.) fritschii; B. (L.) harteti; B. (L.) paytonii etB. (L.) reinii. Les dix espèces appartenant au sous-genreBarbus sont:Barbus (B.) figuiensis; B. (B.) ksibi; B. (B.) lepineyi; B. (B.) magniatlantis; B. (B.) massaensis; B. (B.) moulouyensis; B. (B.) nasus; B. (B.) pallaryi; B. (B.) setivemensis etB. (B.) issenensis.Chaque sous-genre possède son propre pool parasitaire, à l'exception deDactylogyrus marocanus n. sp., recontré sur des espèces appartenant aux deux sous-genres (B. (L.) fritschii, B. (L.) paytonii, B. (L.) harteti, B. (L.) reinii, B. (L.) nasus, B. (B.) setivimensis, B. (B.) ksibi). Sur les cinqDactylogyrus parasitant lesLabeobarbus, trois présentent une spécificité stricte vis à vis de leur hôte. Il s'agit deDactylogyrus reinii n. sp. surB. (L.) reinii; D. volutus n. sp. etD. zatensis n. sp. surB. (L.) fritschii. Les espècesD. oumiensis n. sp. etD. kulindrii n. sp. présentent une spécifité stenoxène et parasitent respectivementB. (L.) harteti, B. (L.) paytonii, B. (L.) reinii etB. (L.) fritschii, B. (L.) reinii.Nous avons recontré neufDactylogyrus chez les poisson-hôtes appartenent au sous-genreBarbus. Six d'entre eux ont une spécificité oïoxène; ce sont:D. guirensis n. sp.,D. atlasensis n. sp. etD. draaensis n. sp. surB. (B.) pallaryi; D. borjensis n. sp. surB. (B.) nasus etD. heteromorphus n. sp. etD. tunisiensis n. sp. surB. (B.) callensis. Les trois autres parasites ont un spectre d'hôtes plus large. Il s'agit deD. ksibii n. sp. recontré chezB. (B.) ksibi, B. (B.) setivimensis etB. (B.) magniatlantis; D. ksibioïdes n. sp. recontré chezB. (B.) setivimensis etB. (B.) moulouyensis. L'espèceD. fimbriphallus n. sp. stenoxène, se recontre chez les poisson-hôtes du versant Sud de l'Atlas et de la façade méditerranéenne à savoir:B. (B.) figuiensis, B. (B.) lepineyi, B. (B.) massaensis, B. (B.) moulouyensis, B. (B.) pallaryi etB. (B.) issenensis.Le rôle des Dactylogyridae en tant que marqueurs biogéographiques, phylogénétiques et taxonomiques est discuté à partir de la composition spécifique des communautés de Monogènes rencontrés et de leurs différents types morphologiques.

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Fifteen new species of the Dactylogyridae (Monogenea) parasitic on fifteen species of barbels (Barbus) from North Africa (Teleostei, Cyprinidae) are described. The fishes studied belong to two subgenera,B. (Labeobarbus) andB. (Barbus), collected from various hydrographical basins of Morocco and from the Hamman Bourgiba locality in Tunisia. In the latter area, the genusBarbus is represented by onlyBarbus (Barbus) callensis. In Morocco, fourteen species are listed, four of which belong to the subgenusLabeobarbus; these areBarbus (L.) fritschii; B. (L.) harteti; B. (L.) paytonii andB. (L.) reinii. The other ten species belong to the subgenusBarbus: these areBarbus (B.) figuiensis; B. (B.) ksibi; B. (B.) lepineyi; B. (B.) magniatlantis; B. (B.) massaensis; B. (B.) moulouyensis; B. (B.) nasus; B. (B.) pallaryi; B. (B.) setivimensis andB. (B.) issenensis. Each of the two subgenera has its unique parasitic fauna, except forDactylogyrus marocanus n. sp. collected on species belonging to both subgenera (B. (L.) fritschii, B. (L.) paytonii, B. (L.) harteti, B. (L.) reinii, B. (B.) nasus, B. (B.) setivimensis andB. (B.) ksibi). Of the five monogeneans found onLabeobarbus, three appear to be specific to one host: they areDactylogyrus reinii n. sp. onB. (L.) reinii, andD. volutus n. sp. andD. zatensis n. sp. onB. (L.) fritschii. D. kulindrii n. sp. parasitisedB. (L.) reinii andB. (L.) fritschii; andD. oumiensis n. sp. occurred onB. (L.) reinii, B. (L.) paytonii andB. (L.) harteti. NineDactylogyrus species were found in fishes belonging to the subgenusBarbus. Six of them have an oïoxenous specificity: these areD. guirensis n. sp.,D. atlasensis n. sp. andD. draaensis n. sp. onB. (B.) pallaryi; D. borjensis n. sp. onB. (B.) nasus andD. heteromorphus n. sp. andD. tunisiensis n. sp. on(B.) callensis. These other three have a wider range of hosts: they areD. ksibii n. sp. collected fromB. (B.) ksibi, B. (B.) setivimensis andB. (B.) magniatlantis, andD. ksibioïdes n. sp. found onB. (B.) setivimensis andB. (B.) moulouyensis. D. fimbriphallus n. sp. is a characteristic parasite of fishes from the southern side of the Atlas mountains and the Mediterranean coast (B. (B.) figuiensis, B. (B.) lepineyi, B. (B.) massaensis, B. (B.) moulouyensis, B. (B.) pallaryi andB. (B.) issenensis).The role of dactylogyrids as biogeographical phylogenetic and taxonomic indicators is discussed in relation to the specific structure of the monogenean communities and the different morphological types found.

     

New species of Cichlidogyrus Paperna, 1960 (Monogenea,Ancyrocephalidae) from the gills of Sarotherodon occidentalis (Daget) (Osteichthyes,Cichlidae) in Guinea and Sierra Leone (West Africa)

The examination of gill parasites from èSarotherodon occidentalisè (Daget) (Pisces, Cichlidae) in Guinea and Sierra Leone (West Africa) revealed the presence of seven species of Monogenea. Two of them have previously been described: ichlidogyrus halli; (Price & Kirk, 1967) and Scutogyrus ecoutini Pariselle & Euzet, 1995. Five are considered as new species, all belonging to Cichlidogyrus Paperna, 1960. They are C. bouvii n. sp., C. fontanai n. sp., C. guirali n. sp., C. paganoi n. sp. and C. sanjeani n. sp.Résumé L'examen des parasites branchiaux de Sarotherodon occidentalis (Daget) (Pisces, Cichlidae) en Guinée et en Sierra Leone (Afrique de l'Ouest) a révélé la présence de sept espèces de Monogènes. Deux ont déjàè été décrites: Cichlidogyrus halli (Price & Kirk, 1967) et Scutogyrus ecoutini Pariselle & Euzet, 1995. Cinq, appartenant toutes au genre Cichlidogyrus Paperna, 1960, sont considérées comme nouvelles, C. bouvii n. sp., C. fontanai n. sp., C. guirali n. sp., C. paganoi n. sp. et C. sanjeani n. sp.     

New and known species of Dactylogyrus Diesing, 1850 (Monogenea,Dactylogyridae) from cyprinid fishes of the River Tigris,Iraq

Four new Dactylogyrus species are described and two redescribed from cyprinids of the River Tigris, Iraq. These are as follows: Dactylogyrus barbioides n. sp. from Barbus grypus; D. orbus n. sp. from Barbus lacerta; D. barbuli n. sp. from Barbus barbulus; D. macrostomi n. sp. from Cyprinion macrostomi; D. pavlovskyi Bychowsky, 1949 from Barbus grypus and Barbus sharpeyi; and D. inutilis Bychowsky, 1949 from Barbus xanthopterus. A phylogenetic and zoogeographical analysis is presented.     

Distribution and occurrence of <Emphasis Type="Italic">Ligula intestinalis</Emphasis> (L.) plerocercoids (Cestoda,Ligulidae) in the fishes of Lake Tana,Ethiopia

The infection rate by Ligula intestinalis has been studied in 14 fish species (Varicorhinus beso, Garra dembecha, Labeobarbus intermedius, L. crassibarbis, L. tsanensis, L. megastoma, L. brevicephalus, L. nedgia, L. acutirostris, L. gorgorensis, L. dainellii, L. macrophthalmus, Barbus humilis, and B. tanapelagius) from Lake Tana (Ethiopia). Plerocercoids have been found only in the fishes of gg. Labeobarbus and Barbus. Ligula has been found in seven out of ten studied large barbels. Barbels larger than 23 cm are not infected with ligula. It is revealed that a specific “synergism” exists between Ligula intestinalis and cestode Khawia sp. parasitizing the intestine of barbels. This synergism is manifested as a regular cooccurrence of these parasites. In juvenile Labeobarbus, fish coparasitized by Khawia sp. and L. intestinalis occurred more often than fish infected with each of this parasite separately.     

Bravohollisia Bychowsky & Nagibina, 1970 and Caballeria Bychowsky & Nagibina, 1970 (Monogenea: Ancyrocephalidae) from Pomadasys hasta (Bloch) (Pomadasyidae), with the description of a new attachment mechanism

Nine species of monogeneans were collected from the fish Pomadasys hasta (Bloch): Bravohollisia magna Bychowsky & Nagibina, 1970; B. rosetta n. sp.; B. reticulata n. sp.; B. gussevi n. sp.; B. kritskyi n. sp.; Caballeria pedunculata Bychowsky & Nagibina, 1970; C. robusta Bychowsky & Nagibina, 1970; C. liewi n. sp.; and C. intermedius n. sp. The anchors of these species contain a canal beginning in the shaft and ending near the point; ducts of haptoral glands enter the canal in the shaft and are associated with net-like structures near the anchor tip. The net-like structures of Bravohollisia spp. are extensive, while those of Caballeria spp. are smaller and more compact. These nets probably assist the monogeneans in attaching to the gill filaments. Caballeria spp. possess four pairs of extensible haptoral digits. Species of Bravohollisia and Caballeria posses two seminal vesicles.     

New evidence of hexaploidy in' large' African Barbus with some considerations on the origin of hexaploidy

Three species of' large' African Barbus from the Republic of Guinea (West Africa), were characterized by the chromosome numbers (2N) of 148 (B. bynni occidentalis and B. wurtzi ) and 150 (B. petitjeani Daget, 1962). All these species have a karyotype which corresponds to the evolutionary hexaploid level. The karyotype of B. petitjeani is composed of chromosomes clearly grouped into morphologically homomorphic sextets which may document the origin of hexaploidy via an autopolyploidic event. Present findings extend the known distribution of evolutionary hexaploidy in' large' African Barbus to West Africa and show evidence of the pan-African distribution of this phenomenon.     

Papuadessus pakdjoko – a new genus and species of rheobiont diving beetle from New Guinea of potential use for environmental impact assessments (Coleoptera,Dytiscidae)

Papuadessus pakdjoko n.gen., n.sp. is described from New Guinea. This species is well characterised by its coloration and the shape of the male genitalia. It is not yet possible to identify closer relatives of this species based on morphological evidence. The new species is a rheobiont, and only occurs at the edge of medium-sized to large rivers in gravelly and sandy substrates. Sites with mud or other finer substrates were unihabited by P. pakdjoko. It is concluded that this species could, therefore, be a good indicator to use in environmental impact assessments, especially where logging and large scale mining strongly increase the input of fine substrates to rivers.     

First report of metazoan parasites from the cichlid Pseudocrenilabrus philander and the cyprinid Enteromius paludinosus in a South African Ramsar wetland

Parasites of two small fish species from a Ramsar wetland in South Africa were studied in 2014–2015. The cichlid Pseudocrenilabrus philander (Weber, 1897) was parasitised by the copepod Lernaea cyprinacea Linnaeus, 1758, the monogenean Gyrodactylus thlapi Christison, Shinn & van As, 2005 and four gryporhynchid metacestode (Cyclophyllidea) species: Paradilepis scolecina (Rudolph, 1819), Paradilepis maleki Khalil, 1961, Neogryporhynchus lasiopeius Baer & Bona, 1960 and Valipora campylancristrota (Wedl, 1855). The cyprinid Enteromius paludinosus (syn. Barbus paludinosus) (Peters, 1852) was infected with the monogenean parasites Dogielius intorquens Crafford, Luus-Powell & Avenant-Oldewage, 2012, Dactylogyrus teresae Mashego, 1983, and three Dactylogyrus spp. These results represent several new locality as well as host records and further contribute information on the parasitic diversity in the Barberspan Ramsar wetland.     

New and known species of Dactylogyrus Diesing, 1850 (Monogenea,Dactylogyridae) from Iranian freshwater cyprinid fishes

Three new Dactylogyrus species, D. rectotrabus n. sp. and D. acinacus n. sp. from Garra rufa plus D. carassobarbi n. sp. from Carassobarbus luteus are described from River Dez (Persian Gulf Basin) of Iran. Three known species, D. haplogonus from Rutilus frisii kutum, D. chalcalburni from Alburnus alburnus and Alburnoides bipunctatus, and D. pavlovskyi from Barbus sharpeyi are recorded from the Rivers Sefid (Caspian Basin), Zayandeh (Central Iran) and Dez (Tigris-Euphrates Basin), respectively. Comments on the monogenean fauna of Iranian freshwater fishes are presented.     

Remarkable Rhinophoridae in a growing generic genealogy (Diptera: Calyptratae,Oestroidea)

Four new genera (Apomorphyto gen.n. from Costa Rica, Bixinia gen.n. from Australia, Rhinodonia gen.n. from New Caledonia, Rhinopeza gen.n. from Papua New Guinea) and nine new species (Apomorphyto inbio sp.n. , Bixinia collessi sp.n. , B. solitaria sp.n. , B. spei sp.n. , B. variabilis sp.n. , B. winkleri sp.n. , Rhinodonia antiqua sp.n. , R. flavicera sp.n. , Rhinopeza gracilis sp.n.) of Rhinophoridae (Diptera: Calyptratae, Oestroidea) are described. All new species were included in a morphology‐based phylogenetic analysis to provide arguments for the justification and monophyly (when nonmonotypic) of the new genera and for including these in the Rhinophoridae. The New Caledonian Rhinodonia is a candidate sister taxon to all other rhinophorids, and the Australasian ‘axiniine’ species emerge inside a clade of all Neotropical taxa thus suggesting migration from South America across Antarctica into Australia. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:51C1F448‐DDD0‐4F14‐8173‐B8C687F7E841 .     

Reproductive biology of four large riverine fishes (Cyprinidae) in a man-made lake,Orange River,South Africa

The reproductive biology ofBarbus holubi, B. kimberleyensis, Labeo capensis andL. umbratus was examined in a large reservoir on the Orange River, South Africa. The findings are integrated into the existing knowledge on largeBarbus andLabeo species, which coexist in most river systems in Africa and Asia.LargeBarbus spawn on gravel beds within the river channel during floods in spring or summer. In continuously flowing regulated rivers, time of spawning is governed by water temperatures. They have moderate fecundity; large eggs, incubation time of several days and the larvae are initially immobile with large yolk sacs. In the impoundment, they spawn in the inflowing regulated river withB. kimberleyensis spawning four to six weeks later than the more cold-tolerantB. holubi, the dominant largeBarbus. Survival is generally good and the juveniles disperse throughout the lake, but unseasonal release of cold water from an upstream impoundment may cause poor reproductive success.In contrast,Labeo species generally spawn on newly flooded ground, usually leaving the main river channel. Spawning may or may not be preceded by a longitudinal migration. Labeos are relatively fecund with small (30%Barbus size) eggs which hatch quickly and the larvae swim in bursts up into the water column before sinking down again.L. capensis does not require a longitudinal spawning migration and breeds throughout the lake, depending on local conditions. Large temporal variation in gonadal development within the population can result in more than one spawning. Dispersal within the lake is poor.L. umbratus uses larger inflowing tributaries for spawning thanL. capensis and its juveniles have a much greater power of dispersal. Early dependence on external feeding and undependable occurrence of conditions for spawning and juvenile feeding makes for variable reproductive success.     

Barbus xanthos,a new barbel from the Southern Aegean basin (Teleostei: Cyprinidae)

Barbus xanthos, a new species, is described from the Eşen, Dalaman, Tersakan and Büyük Menderes rivers in south-western Anatolia. It differs from other Barbus species in the adjacent basins by having 53–60 lateral line scales, a weakly ossified last unbranched dorsal-fin ray (about 33–50%), numerous small irregular-shaped black or dark-brown spots smaller than scales, often forming large, black or dark-brown blotches on back and flank in juveniles and adults, and a straight or slightly convex posterior dorsal-fin margin. B. xanthos differs from its most closely related congener, B. pergamonensis, by nine nucleotide substitution sites in the mitochondrial DNA cytochrome oxidase I barcode region.     

Cophylogenetic relationships between Dactylogyrus (Monogenea) ectoparasites and endemic cyprinoids of the north-eastern European peri-Mediterranean region

The study of host–parasite coevolution is one of the cornerstones of evolutionary biology. The majority of fish ectoparasites belonging to the genus Dactylogyrus (Monogenea) exhibit a high degree of host specificity. Therefore, it is expected that their evolutionary history is primarily linked with the evolutionary history of their cyprinoid fish hosts and the historical formation of the landmasses. In the present study, we used a cophylogenetic approach to investigate coevolutionary relationships between endemic Cyprinoidea (Cyprinidae and Leuciscidae) from selected regions in southern Europe and their respective Dactylogyrus species. A total of 49 Dactylogyrus species including endemic and non-endemic species were collected from 62 endemic cyprinoid species in the Balkan and Apennine Peninsulas. However, 21 morphologically identified Dactylogyrus species exhibited different genetic variants (ranging from 2 to 28 variants per species) and some of them were recognized as cryptic species on the basis of phylogenetic reconstruction. Phylogenetic analyses revealed several lineages of endemic and non-endemic Dactylogyrus species reflecting some morphological similarities or host affinities. Using distance-based and event-based cophylogenetic methods, we found a significant coevolutionary signal between the phylogenies of parasites and their hosts. In particular, statistically significant links were revealed between Dactylogyrus species of Barbini (Cyprinidae) and their hosts belonging to the genera Aulopyge, Barbus and Luciobarbus. Additionally, a strong coevolutionary link was found between the generalist parasites D. alatus, D. sphyrna, D. vistulae, and their hosts, and between Dactylogyrus species of Pachychilon (Leuciscidae) and their hosts. Cophylogenetic analyses suggest that host switching played an important role in the evolutionary history of Dactylogyrus parasitizing endemic cyprinoids in southern Europe. We propose that the high diversification of phylogenetically related cyprinoid species in the Mediterranean area is a process facilitating the host switching of specific parasites among highly diverse congeneric cyprinoids.     

Gyrodactylus medaka n. sp. (Monogenea: Gyrodactylidae) parasitic on wild and laboratory-reared medaka Oryzias latipes (Beloniformes: Adrianichthyidae) in Japan

Gyrodactylus medaka n. sp. (Monogenea: Gyrodactylidae) is described from the skin, fins, and gills of medaka Oryzias latipes (Beloniformes: Adrianichthyidae) from Japan. This new species was collected from wild medaka in Hiroshima, Aichi, Saga, and Kumamoto prefectures, and laboratory-reared medaka in Chiba and Aichi prefectures. The small marginal hook sickle (≤4?μm) and the length of the marginal hook of the new species are the diagnostic morphological characters differentiated from other gyrodactylids reported from Asia. The pairwise sequence divergences for the interspecific variation in ITS regions and the phylogenetic analysis suggest that the populations of G. medaka n. sp. may have a similar genetic variation as the medaka populations in Japan. Gyrodactylus medaka n. sp. and Dactylogyrus oryziasi (Monogenea: Dactylogyridae) can maintain their populations in laboratory aquaria using medaka as their hosts, and these monogeneans and medaka have the potential as experimental model animals for clarifying various aspects of their host-parasite relationships. In addition, we report the composition of modified ammonium picrate-glycerin (APG) and show it is advantageous for monogenean taxonomy.     

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.     

The distribution of monogenean parasites on aquatic vertebrates inhabiting Chinese inland waters

The general distribution of monogenean parasites of aquatic vertebrates inhabiting Chinese inland waters is summarised. Five hundred and seventy-two out of a total of 581 species of monogeneans were discovered on fish, while only nine species were found on Amphibia and Reptilia. Most dactylogyrids and diplozoids parasitise cyprinids, while ancyrocephalids occur on silurids and cyprinids, and gyrodactylids are found mainly on cyprinids and cobitids. Analyses of host-specificity and host-diversity suggests that the family Ancyrocephalidae should be divided into several families parallel with the Dactylogyridae. Of 12 subfamilies of the Cyprinidae, only the Gobiobotinae was found to be free of infection with monogeneans, and the genus Dactylogyrus has more species than any other monogenean genus associated with every cyprinid subfamily.     

Co‐evolutionary patterns in congeneric monogeneans: a review of Dactylogyrus species and their cyprinid hosts

Patterns associated with the evolution of parasite diversity, speciation and diversification were analysed using Dactylogyrus species (gill monogeneans) and their cyprinid hosts as a model. The aim of this study was to use this highly specific host–parasite systems to review: (1) the diversity and distribution of Dactylogyrus species, (2) the patterns of organization and structure of Dactylogyrus communities, (3) the evolution and determinants of host specificity and (4) the mode of Dactylogyrus speciation and co‐evolutionary patterns in this Dactylogyrus–cyprinid systems. Dactylogyrus are a highly diverse group of parasites, with their biogeography and distribution clearly linked to the evolutionary history of their cyprinid hosts. The coexistence of several Dactylogyrus species on one host is facilitated by increasing niche distances and the differing morphology of their reproductive organs. The positive interspecific and intraspecific interactions seem to be the most important factors determining the structure of Dactylogyrus communities. Host specificity is partially constrained by parasite phylogeny. Being a strict specialist is an ancestral character for Dactylogyrus, being the intermediate specialists or generalists are the derived characters. The evolution of attachment organ morphology is associated with both parasite phylogeny and host specificity. Considering larger and long‐lived hosts or hosts with several ecological characters as the measures of resource predictability, specialists with larger anchors occurred on larger or longer‐living fish species. Intra‐host speciation, a mode of speciation not often recorded in parasites, was observed in Dactylogyrus infecting sympatric cyprinids. Sister parasite species coexisting on the same host occupied niches that differed in at least one niche variable. Intra‐host speciation, however, was not observed in Dactylogyrus species of congeneric hosts from geographically isolated areas, which suggested association by descent and host‐switching events.     

Identification of Monogenea made easier: a new statistical procedure for an automatic selection of diagnostic linear measurements in closely related species

We introduce a new statistical method to select which morphological characters are most useful to identify monogenean species. The method estimates the average size overlap of candidate diagnostic structures among a set of species to individuate those that mostly differ between the species. To demonstrate our approach, we report a comprehensive analysis conducted on two of the most species‐rich monogenean genera: Dactylogyrus Diesing, 1850 and Gyrodactylus von Nordmann, 1832. We demonstrate that, in contrast to common taxonomic practice, very few but highly diagnostic measurements are necessary to correctly identify a specimen. In particular, we found that most of Dactylogyrus and Gyrodactylus species can be classified on the basis of the width of the supplementary connecting bar and of the length of the hook sickle, respectively.