The phylogenetic status of the Ancyrocephalidae Bychowsky, 1937 (Monogenea: Dactylogyroidea)

Phylogenetic analysis of selected subfamily and family taxa within the Dactylogyroidea indicates that the Ancyrocephalidae Bychowsky, 1937, is unnatural. The family contains both poly- and paraphyletic features. The analysis supports the previous elevation of the Pseudomurraytrematinae to family status and suggests that revision of the Ancyrocephalidae is necessary. Two options for revision are provided; that of returning the taxon to subfamily status within the Dactylogyridae is preferred, requiring a change in status of the Heterotesiidae to a subfamily within the Dactylogyridae.     

The evaluation for generic-level monophyly of Ancyrocephalinae (Monogenea, Dactylogyridae) using ribosomal DNA sequence data

To overcome the limited regional and taxonomic coverage in previous studies of the Ancyrocephalinae (Monogenea, Dactylogyridae), we present further findings on the molecular systematics and phylogeny of a broader selection of specimens, aiming to build a molecular phylogeny of the Ancyrocephalinae, and to assess the monophyly of each available genus, using D1-D2 domain of LSU rDNA and the combined LSU and partial sequence of SSU rDNA data sets. Our studies showed that 18 Haliotrema species were highly dispersive to form several clades with species from ten closely related genera. The host range of Euryhaliotrematoides species was not only restricted in butterfly fishes (Chaetodontidae), but widen to include the Lutjanidae. Euryhaliotrema was phylogenetically more closely related to Aliatrema than to Euryhaliotrematoides. Given that the species Haliotrema kurodai, Haliotrema spirotubiforum and Haliotrema anguiformis had a funnel-shaped base of the coiled male copulatory organ (MCO) lacking an accessory piece, and our molecular evidence, we proposed to transfer the three species to the Aliatrema as new combinations. We proposed to combine Bravohollisia and Caballeria into one genus, mainly based on molecular evidence and their similar MCO characters. Using SSU+ITS1 data set, Scutogyrus was robustly resolved as a polyphyletic group and its status should be questioned. Based on the present molecular evidence and their similar MCO characters, we proposed to combine Cichlidogyrus and Scutogyrus into one genus, Cichlidogyrus, by treating Scutogyrus as the synonym of Cichlidogyrus. Generally, the present study indicated that the vagina can make little contribution for understanding the generic-level monophyly, due to its high variability even among phylogenetically very closely related species, but it was useful for species determination. Given that phylogenetically closely related species from the same or closely related host species may have similar MCO characters but distinct haptoral characters, we consider that it is dangerous to erect a genus mainly based on different haptoral characters, particularly those separated from existing genera. The resultant phylogenetic reconstructions indicated that the radiation of some Haliotrema species has correlated with their hosts, because species from closely related fishes have correspondingly shown close relationships. Though the present study can not provide the comparison of host-parasite associations, phylogenetically closely related Haliotrema species which have similar morphological characters (both MCOs and haptors) but distinct/distantly related host species may indicate a speciation model of host-switching.     

A catalogue of the nominal species of the monogenean family Dactylogyridae Bychowsky, 1933 (excluding Dactylogyrus Diesing, 1850)

A list of the nominal species of the Dactylogyridae Bychowsky, 1933 (with the exception of Dactylogyrus Diesing, 1850) is presented. It consists of almost 100 specific names in 10 genera, with their hosts (to the generic level) and geographical region. A reference list to the authorities for the taxa is included.     

Phylogenetic relationships among monogenean gill parasites (Dactylogyridea, Ancyrocephalidae) infesting tilapiine hosts (Cichlidae): systematic and evolutionary implications

We studied the systematics of 14 species of monogenean (Ancyrocephalidae) gill parasites from West African tilapiine hosts (Cichlidae) using both morphological and genetic data. With these tools, we were able to: (i) confirm the validity of the previously described morphological parasite species and of the genus Scutogyrus; (ii) propose that some stenoxenous species (i.e., parasite species with more than one host) may be composed of sister species (e.g., Cichlidogyrus tilapiae); (iii) state that the use of the morphology of the haptoral sclerites is more suitable to infer phylogenetic relationships than the morphology of the genitalia (which seems to be more useful to resolve species-level identifications, presumably because of its faster rate of change). These results imply that: (i) the specificity of these monogenean parasites is greater than initially supposed (what were thought to be stenoxenous species may be assemblages of oïoxenous sister species); (ii) related species groups (i.e., “tilapiae,” “halli,” and “tiberianus”) have to be, as genus Scutogyrus, validated within the 54 ancyrocephalid species described from 18 species of tilapiine hosts in West Africa, (iii) the group “tilapiae,” due to its morphology and host range, have to be considered as being the most primitive; (iv) the occurrence of lateral transfers and parallel speciation processes are necessary to describe the repartition of the newly described parasite groups on the three host genera studied (Tilapia, Oreochromis, and Sarotherodon).     

New species of Dactylogyridae Bychowsky, 1933 infecting the gills of Myloplus schomburgkii (Jardine) and Colossoma macropomum (Cuvier) in the Peruvian Amazon

Systematic Parasitology - Four new dactylogyrid species are described, three species (Anacanthorus camposbacae n. sp., Anacanthorus carmenrosae n. sp. and Notozothecium nanayensis n. sp.) from the...     

Molecular phylogeny of species of Ligophorus (Monogenea: Dactylogyridae) and their affinities within the Dactylogyridae

The taxonomic framework of Ligophorus, monogenean specialists of the gills of grey mullets (Mugilidae), is evaluated and its interspecific relationships are assessed for the first time using molecular data. The position of Ligophorus within the paraphyletic Ancyrocephalinae is re-assessed based on newly sequenced species. Furthermore, the relationship between morphometric and genetic interspecific similarities is evaluated. Partial 28S and complete ITS1 rDNA sequences from representatives of 14 of the 16 nominal species of Ligophorus from the Mediterranean, Black and Azov Seas were analysed together with published sequences of members of the Dactylogyridae. The phylogenetic analyses of the Dactylogyridae (i) confirmed the position of Ligophorus within the marine Ancyrocephalinae; (ii) revealed a sister relationship between Ergenstrema and Ligophorus, whose species are all exclusive parasites of grey mullets; and (iii) substantiated the affinities of Ergenstrema with the marine Ancyrocephalinae. The phylogenetic analysis restricted to Ligophorus confirmed the distinct status of the included species. The ITS1 region provided the highest divergence between species and phylograms with the strongest branch support. Both the 28S and ITS1 phylograms revealed two main clades. One included species from hosts with Mediterranean and NE Atlantic distribution and another was formed by species parasitising several Liza spp., including Lz. haematocheilus from the Northwestern Pacific, and Mugil cephalus, which suggests an origin outside the Mediterranean for the latter clade. The phylogenetic evidence presented herein indicated that a combination of host-switching and lineage duplication events accounted for the diversification of this genus in the Mediterranean basin. The agreement between molecular and morphological interspecific similarities observed in Ligophorus supports the validity of morphometric characters used for species identification.     

Phylogenetic relationships of sponges with placochelae or related spicules (Poecilosclerida, Guitarridae) with a systematic revision

All the currently known sponge species bearing placochelae or placochelae-like spicules (i.e. belonging to the genera Guitarra, Coelodischela, Tetrapocillon, Euchelipluma , and Hoplakithara ) have been reviewed and their relationships delineated by cladistic analysis. A matrix of 18 taxa and 14 characters is included. The species Isodictya palmata and Esperiopsis fucorum were used as an outgroup, because they shared either monactines or diactines and smooth palmate isochelae with different members of the ingroup. Cladistic analysis using PAUP produced two equally parsimonious trees of 33 steps (CI = 0.758). Strict, Semistrict and Majority Rule consensus trees displayed the same topology. The phylogeny of the trees was not totally resolved. The bootstrap 50% majority-rule consensus tree supported, to a greater or lesser extent, the previously detected monophyletic groups. A common linkage for the ingroup was found in 72% of instances. The genus Euchelipluma appeared as monophyletic in 75% of instances while the group which included the genus Guitarra did so in 66%. The monophyly of the species with sigmoid microscleres and without spiny isochelae (G. isabellae/sigmatifera/antarctica/dendyi and C. diatomorpha/massa ) received 56% support as did the group of species with spiny isochelae, whereas monophyly of the group laplani/ bipocillifera received 64% support and the genus Tetrapocillon 56%. According to our cladistic analysis, all the species bearing placochelae or derived forms should be allocated to Guitarridae. Within this family, the genus Euchelipluma appeared as monophyletic while Guitarra was paraphyletic. The single species of the genus Hoplakithara clearly belongs to the Guitarra ( sigmatifera ) group and thus becomes synonymous with Guitarra. G. solorzanoi is considered here a synonymy of G. laplani. A diagnosis for the 10 valid species of Guitarra known to date as well as for the two species of Coelodischela and the two of Euchelipluma is given.     

Phylogenetic analysis of the Dactylogyroides longicirrus (Monogenea: Dactylogyridae) based on the 18S and ITS 1 ribosomal genes

The present study describes the molecular phylogenetic analysis of Dactylogyroides longicirrus (Monogenea: Dactylogyridae) infecting the gill filaments of fish Puntius sophore from the site Guwahati, Assam, India. The parasite Dactylogyroides longicirrus (Tripathi, 1959) Gusev, 1976 from Northeast Indian region is presented based on sequence data of a 738 base-pair fragment of ribosomal 18S small subunit and first internal transcribed spacer (ITS 1). Phylogenetic relationships were inferred using neighbour joning and maximum parsimony methods and the results support the validation of D. longicirrus. The study is also supported by secondary structure model prediction by using minimum free energy which can be considered a promising tool for monogenean species identification. This is the first report of this parasite from Northeast region of India, with this, the 18S and ITS 1 rDNA region amplified in the study is also the first sequence of the genus Dactylogyroides.     

Phylogeny and a revised classification of the Monogenoidea Bychowsky, 1937 (Platyhelminthes)

A hypothesis (CI=57.3%) on the evolutionary relationships of families comprising the class Monogenoidea is proposed based on 141 character states in 47 homologous series and employing phylogenetic systematics. Based on the analysis, three subclasses, the Polyonchoinea, Polystomatoinea and Oligonchoinea, are recognised. The analysis supports independent origins of the Montchadskyellidae within the Polyonchoinea and of the Neodactylodiscidae and Amphibdellatidae within the order Dactylogyridea (Polyonchoinea); the suborder Montchadskyellinea is raised to ordinal status and new suborders Neodactylodiscinea and Amphibdellatinea are proposed to reflect these origins. The Gyrodactylidea (Polyonchoinea) is supported by three synapomorphies and comprises the Gyrodactylidae, Anoplodiscidae, Tetraonchoididae and Bothitrematidae. The analysis supports recognition of the Polystomatoinea comprising Polystomatidae and Sphyranuridae. Evolutionary relationships within the Oligonchoinea indicate independent origins of three ordinal taxa, the Chimaericolidea (monotypic), Diclybothriidea (including Diclybothriidae and Hexabothriidae) and Mazocraeidea (with five suborders). The suborder Mazocraeinea comprises the Plectanocotylidae, Mazocraeidae and Mazoplectidae, and is characterised by two synapomorphies. The suborder Gastrocotylinea, characterised by presence of accessory sclerites in the haptoral sucker, is divided into two infraorders, the monotypic Anthocotylina infraorder novum and Gastrocotylina. Two superfamilies of the Gastrocotylina are recognised, the Protomicrocotyloidea and Gastrocotyloidea; the Pseudodiclidophoridae is considered incertae sedis within the Gastrocotylina. The suborder Discocotylinea comprises the Discocotylidae, Octomacridae and Diplozoidae and is supported by four synapomorphies. The monotypic Hexostomatinea suborder novum is proposed to reflect an independent origin of the Hexostomatidae within the Mazocraeidea. The terminal suborder Microcotylinea comprises four superfamilies, the Microcotyloidea, Allopyragraphoroidea, Diclidophoroidea and Pyragraphoroidea. The analysis supports incorporation of the Pterinotrematidae in the Pyragraphoroidea and rejection of the monotypic order Pterinotrematidea. The following taxa are also rejected for reasons of paraphyly and/or polyphyly: Articulonchoinea, Bothriocotylea, Eucotylea, Monoaxonematidea, Tetraonchidea, Gotocotyloidea, Anchorophoridae and Macrovalvitrematidae. The Sundanonchidae, Iagotrematidae and Microbothriidae were not included in the analysis because of lack of pertinent information regarding character states.     

Certain problems of the taxonomy of monogeneans of the order Dactylogyridea (Platyhelminthes: Monogenea)

Different taxonomic problems of the Dactylogyridea Bychowsky, 1937 are discussed. From the recent point of view, all old genera need a careful revision. Taxa of higher ranks should be analysed taking into account their host phylogeny and zoogeography. The order Tetraonchidea Bychovsky should be disband and all its families should be moved to the Dactylogyridea. The aberrant families Montchadskyellidae and Anoplodiscidae also belong to the Dactylogyridea. Dactylogyrideans are most similar to the primitive monogeneans.     

New names for homonyms in the genus Dactylogyrus Diesing, 1850 (Monogenea, Dactylogyridae)

The following new replacement names for homonyms in the genus Dactylogyrus Diesing are proposed: Dactylogyrus acrossocheili nom. nov. for D. forcipatus Wu & Wang, 1983; D. birgii nom. nov. for D. simplex Birgi & Lambert, 1987; D. cheni nom. nov. for D. orientalis Hu & Chen, 1979; D. hui nom. nov. for D. xenocypris Hu & Chen, 1979; D. limae nom. nov. for D. hamatus Chinabut & Lim, 1994; D. mikailovi nom. nov. for D. gracilis Mikailov, 1974; D. pakistanensis nom. nov. for D. mrigali Rizvi, 1978; D. papernai nom. nov. for D. magnum Paperna, 1973; D. saurogobii nom. nov. for D. falcatus Wu, Wang & Song, 1983; and D. semifasciolatae nom. nov. for D. lineatus Luo & Lang, 1982.     

Chauhanellus Bychowsky & Nagibina, 1969 (Monogenea) from ariid fishes (Siluriformes) of Peninsular Malaysia

Twelve new species of Chauhanellus Bychowsky & Nagibina, 1969 have been found on six species of ariid from Peninsular Malaysia: Chauhanellus trifidus n. sp., C. digitalis n. sp., C. malayanus n. sp., C. forcipis n. sp. and C. intermedius n. sp. from Arius sagor; C. aspinous n. sp. from Arius venosus; C. caelatus n. sp. from Arius caelatus; C. auriculatum n. sp., C. poculus n. sp. and C. pulutanus n. sp. from Arius maculatus; C. duriensis n. sp. from Arius thalassinus; and C. osteogeneiosi n. sp. from Osteogeneiosus militaris. Some of these Chauhanellus species possess characteristics that are not commonly observed in the genus. C. aspinous n. sp., C. intermedius n. sp. and C. digitalis n. sp. exhibit features found in both Chauhanellus and Hamatopeduncularia: these include absence of spines on the mainpart of the dorsal anchors in C. aspinous n. sp. and C. intermedius n. sp. and presence of haptoral digitation in C. digitalis n. sp. Other features are the five transverse rows of peduncular spines in C. duriensis n. sp., ear-like projections on the anchors in C. auriculatum n. sp., and thin sclerotised plates that partly envelope the ventral anchors in C. forcipis n. sp. Mid-dorsal appendices occur on the dorsal bars of seven of the present species.     

A revision of the Calicotylinae Monticelli, 1903(Monogenea: Monocotylidae)

A total of 153 elasmobranchs (46 species), either freshly collected from the Gulf of Mexico, USA and Tasmania, Australia or museum specimens collected from various localities worldwide, were examined for calicotyline (Monocotylidae) monogeneans. Thirty-five elasmobranchs, representing 17 species, were infected with Calicotyle spp. which we identified as the following previously described species: C. asterii (Szidat, 1970) Timofeeva, 1985, C. kroyeri Diesing, 1850, C. macrocotyle Cordero, 1944, C. similis (Szidat, 1972) Timofeeva, 1985, C. splendens (Szidat, 1970) Timofeeva, 1985, C. stossichi Braun, 1899 and C. urolophi Chisholm, Beverley-Burton & Last, 1991. The Calicotylinae, which comprises the genera Calicotyle and Dictyocotyle, is revised based on supplementary material as well as deposited type-material. We consider 14 of the 17 nominal Calicotyle spp. to be valid. C. rosinae Kusnetzova, 1970 is synonymised with C. macrocotyle, C. sjegi Kusnetzova, 1970 is considered a species inquirenda and C. inermis Woolcock, 1936 a species incertae sedis. Additional data and illustrations to show the morphological features of the hamuli and male copulatory organ, the form of the intestinal caeca, vaginae and ovary and the distribution of the vitellarium are provided for all valid species. The distribution of the 14 hooklets in the adult haptor of Dictyocotyle coeliaca Nybelin, 1941 is illustrated for the first time. We provide new host and locality records for C. asterii, C. kroyeri, C. macrocotyle and C. stossichi and new locality records for C. similis and C. splendens. A key to species of the Calicotylinae is also included. Host-specificity, geographical distribution and the need for information regarding the development of individuals from juvenile to adult are discussed.     

Speciation of Protopolystoma Bychowsky, 1957 (Monogenea: Polystomatidae) in hosts of the genus shape Xenopus (Anura: Pipidae)

The taxonomy, geographical distribution and hostrange of the polystomatid genus ProtopolystomaBychowsky, 1957 are reviewed. P. xenopodis(Price, 1943) and five new species are recognised,which occur in clawed toads ( Xenopus spp.)throughout subsaharan Africa. Of the two clawed toadsubgenera, Xenopus and Silurana, only theformer is infected. Protopolystoma spp. aredifferentiated by morphological variation of the gut,large hamulus and penis armature. P.xenopodis is found in Xenopus laevis subspeciesin South Africa, Transkei, Zimbabwe, DemocraticRepublic of Congo (D.R.C.), Rwanda, Uganda, Kenya andCameroon ( X. l. poweri and X. l.sudanensis are new host records). It also occurs inintroduced populations of X. l. laevis in theUnited States (southern California) and United Kingdom(South Wales). In subsaharan Africa the speciesdisplays significant, but continuous, geographicalvariation of penis spine size between southernpopulations in X. l. laevis and those in morenortherly host subspecies. Data on the natural hostrange of this parasite were complemented by anexperimental study of host-specificity in the southernform. This can produce patent infections in X.l. victorianus and X. gilli, but not X.wittei nor X. (Silurana) tropicalis. P.simplicis n. sp. is endemic to central and eastAfrican areas, infecting X. laevis subspecies ineastern D.R.C., Rwanda, Uganda and western Kenya, X. wittei-like hosts in eastern D.R.C., westernUganda, Rwanda and Burundi, X. vestitus inwestern Uganda and Xenopus sp. at Nairobi,Kenya. P. ramulosus n. sp. occurs in X.fraseri-like toads in eastern D.R.C. (Gabon andCameroon are also possible literature records), and P. fissilis n. sp. is found in X. fraseri-and X. wittei-like species in Cameroon andeastern D.R.C., and in southern Rwanda, respectively. Two Protopolystoma taxa are found in X.muelleri populations now suspected to representdistinct species: P. occidentalis n. sp. occursin X. muelleri (western form) in Ghana, Togo,Nigeria and Cameroon, while P. orientalis n. sp.is found in X. muelleri (eastern form) in SouthAfrica, Zimbabwe and Tanzania. The allopatricallydistributed species P. ramulosus, P.simplicis, P. occidentalis and P.orientalis form a relatively homogenous grouping withsome interspecific morphological overlap. These taxaare distinguished from P. xenopodis by penisspine morphology and from P. fissilis by hamulusroot form and aspects of gut morphology. Unidentified Protopolystoma sp. have been recorded in X. clivii in Ethiopia, X. fraseri aff. inCameroon and Xenopus sp. in Kenya and Tanzania. At some localities, single host species were infectedby two representatives of Protopolystoma. P. fissilis was recorded in eastern D.R.C. with P. ramulosus, with Protopolystoma sp. inCameroon in X. fraseri-like hosts and with P. simplicis in X. wittei-like hosts in Rwanda. P. xenopodis co-occurred with P. simplicisin X. laevis subspecies through central and eastAfrica.     

Phylogenetic analyses of the family Tetraonchidae (Platyhelminthes: Monogenea)

A phylogenetic reconstruction of the monogenean family Tetraonchidae was carried out by methods of parsimony-based cladistics. The analysis included 20 species of tetraonchids and two out-groups (Sundanonchus tomanorum and Dactylogyrus amphibothrium) and was based on 34 morphofunctional characters. Software PAUP 4.0 and Winclada were used for the phylogenetic reconstructions. Obtained results allow proposing a preliminary phylogenetic hypothesis of the family Tetraonchidae along with the discussion of host-parasite association. According to the current taxonomic view, the family Tetraonchidae included two genera. Cladistic analysis showed a monophyly of the family and the genus Tetraonchus Diesing, 1858. Two representative of the former genus, Tetraoncus monenteron and T. borealis, parasitize the pikes (Esocoformes: Esocidae) and the grayling (Salmonidae: Thymallinae) respectively. The genus Salmonchus Spassky et Roytman, 1958 has a complicated structure and its intrageneric relationships were not completely resolved; in general, the analysis allows to recognise several species groups: Salmonchus oncorhynchi--the parasite of the Oncorhynchus masou smolt living during the first year of life in fresh water; four species (S. variabilis, S. gussevi, S. grumosus, S. alaskensis) inhabiting specifically the whitefishes (Salmonidae: Coregoninae); all reminder of Salmonchus species occurring on the salmons (Salmonidae: Salmoninae). The bootstrap test gives a support only for the following clades: family Tetraonchidae (75%), genus Tetraonchus (88%); a group of Salmonchus species associated with the whitefishes (93%) and grouping of four species (S. huhonis, S. pseudolenoki, S. skrjabini and S. lenoki) from the lenoks (Brachymystax) and taimens (Hucho) (61%).