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The parasite fauna of Neotropical reptiles is poorly known, and the number of parasites described in these hosts does not seem to correspond to the actual species diversity in this zoogeographical region. This also applies to tapeworms such as proteocephalids, which are rarely found in reptiles and are strictly specific to their reptilian hosts. In the present paper, three new species of Ophiotaenia La Rue, 1911 are described from three dipsadine snake species (Squamata: Colubridae) in Ecuador, namely O. jeanmarctouzeti sp. n. from the Neotropical blunt-headed treesnake Imantodes cenchoa (Linnaeus), O. barraganae sp. n. from the beautiful calico snake Oxyrhopus formosus (Wied-Neuwied) and O. velascoae sp. n. from the forest flame snake Oxyrhopus petolarius (Linnaeus). The new species are characterised by type 1 uterine development, the number and distribution of testes, the size of the scolex and other metric features. As no molecular data are available on the specimens collected more than 35 years ago, the phylogenetic relationships of the individual taxa are not known.

Tapeworms of the genus Proteocephalus Weinland, 1858 (Cestoda: Proteocephalidae) are common and widespread intestinal parasites of whitefish (Coregonus spp., Salmonidae: Coregoninae). Previous taxonomic studies, based solely on morphology and inconsistently fixed specimens, concluded that all salmoniform fish, including whitefish, are parasitised by a single euryxenous and highly polymorphic species, Proteocephalus longicollis (Zeder, 1800). However, recent molecular phylogenetic analyses have revealed the existence of several species specific to individual genera or even species of salmoniform fish. In this study, Proteocephalus fallax La Rue, 1911 is redescribed based on newly collected and genetically characterised specimens from several Coregonus species in Switzerland, the type locality of the species, and in Norway. This cestode was previously synonymised with P. exiguus La Rue, 1911, a parasite of whitefish in North America, but the two species are not closely related. Proteocephalus fallax differs from P. exiguus in its larger body size, wider proglottids, shorter cirrus sac and broader scolex. In addition, the other Proteocephalus species described in whitefish are briefly discussed, with comments on their validity, host range and distribution.

Proteocephalid tapeworms of frogs of the family Ranidae ('true' frogs) are reviewed with emphasis on their species diversity, host specificity and geographical distribution. New molecular data (nuclear lsrDNA and mitochondrial COI sequences) are presented for tapeworms of four species of ranid frogs in North America, including the poorly known Ophiotaenia saphena Osler, 1931 of Rana clamitans Latreille and R. catesbeiana (Shaw), which is redescribed using new material from Arkansas, USA. Tapeworms of R. sphenocephala (Cope) and R. pipiens Schreber, the latter previously identified as O. saphena, represent another, putative new species, but are not formally described due to insufficient available material. Proteocephalus papuensis Bursey, Goldberg et Kraus, 2008 from Sylvirana supragrisea (Menzies) is transferred to Ophiotaenia La Rue, 1911 as a new combination. After a critical review of the literature, only nine nominal species of Ophiotaenia are recognised as valid, which is in contrast to the large number of ranid frogs (> 440 spp.). The reasons for this striking disparity are briefly discussed, and a key based on morphology is presented for the identification of all species of Ophiotaenia from the Ranidae. Molecular data are available for only two taxa from North America that form a monophyletic group. The relationships among tapeworms of ranid frogs occurring in other zoogeographical regions are not yet known. The taxonomic status of Batrachotaenia Rudin, 1917, which was erected to accommodate proteocephalids from amphibians, is also discussed. To facilitate future studies, a tabulated summary of all 32 species of proteocephalids belonging to three genera reported from amphibians (frogs and salamanders) is presented, with information on their hosts, distribution, and taxonomically important characters, including key measurements.

Parasitological examination of freshwater fishes of the Phongolo River in north-eastern KwaZulu-Natal, South Africa resulted in the discovery and morphological and molecular characterisation of a new species of Wenyonia Woodland, 1923 (Cestoda: Caryophyllidea). The new species from the plain squeaker, Synodontis zambezensis Peters (Siluriformes: Mochokidae), is morphologically most similar to Wenyonia acuminata Woodland, 1923, a species reported from three species of Synodontis in north-eastern, western and central Africa (Sudan, Nigeria, Democratic Republic of the Congo). Both these species are markedly different from congeners by having a nematoform body and a digitiform scolex. Wenyonia gracilis sp. n. differs from W. acuminata in its general body size, length and width of main body regions (testicular and uterine regions), a posterior extension of the testes into the uterine region, numerous postovarian vitelline follicles filling the entire medulla, eggs c. 1/3 larger in size, and a scolex with an apical introvert but devoid of longitudinal furrows and a well-defined base. Wenyonia gracilis is the seventh species in the genus and the first autochthonous caryophyllidean ever reported and described from southern Africa (south of the Zambezi River).

A survey of the species of the Proteocephalus-aggregate from sticklebacks (Actinopterygii: Gasterosteidae) is provided. The occurrence of three species in North America is confirmed: (i) Proteocephalus filicollis (Rudolphi, 1802), which has been reported from the three-spined stickleback, Gasterosteus aculeatus Linnaeus, in the northeastern part of North America (Newfoundland); (ii) Proteocephalus pugetensis Hoff et Hoff, 1929 occurs also in G. aculeatus, but in northwestern North America (British Columbia and Washington); and (iii) Proteocephalus culaeae sp. n., which is described from the brook stickleback, Culaea inconstans (Kirtland), in Manitoba (Canada). Another species, Proteocephalus ambiguus (Dujardin, 1845), a specific parasite of the nine-spined stickleback, Pungitius pungitius (Linnaeus), and type species of the genus, has also been found in North America (Alberta, Canada), but its vouchers are in poor condition and cannot be reliable assigned to this species. Both species reported from three-spined stickleback differ from each other by the shape of the scolex (rounded in P. filicollis versus continuously tapered towards the anterior extremity in P. pugetensis) and the apical sucker (widely oval to subspherical in frontal view in P. filicollis versus flattened in P. pugetensis). Proteocephalus culaeae sp. n. is characterised by a short body composed of a few, continuously widened proglottids, a short scolex narrower than the strobila and devoid of an apical sucker, a short, pyriform cirrus sac, no vaginal sphincter, and few testes. A key to species of the Proteocephalus-aggregate from sticklebacks is provided.

A comparative study of the scoleces of monozoic tapeworms (Cestoda: Caryophyllidea), parasites of catostomid and cyprinid fishes (Teleostei: Cypriniformes) in the Nearctic Region, was carried out using light and scanning electron microscopy. Scoleces of 22 genera of North American caryophyllideans were characterised and their importance for taxonomy, classification and phylogenetic studies was critically reviewed. Nearctic genera exhibit a much higher variation in the shape and form of scoleces compared with taxa in other biogeographical regions. The following basic scolex types can be recognised in Nearctic caryophyllideans: monobothriate (Promonobothrium Mackiewicz, 1968), loculotruncate (Promonobothrium, Dieffluvium Williams, 1978), bothrioloculodiscate (Archigetes Leuckart, 1878, Janiszewskella Mackiewicz et Deutsch, 1976, Penarchigetes Mackiewicz, 1969, Pseudoglaridacris Oros, Uhrovič et Scholz, 2018), fixomegabothriate (Capingens Hunter, 1927), bulbate and bulboacuminate (Atractolytocestus Anthony, 1958), cuneiloculate (Hypocaryophyllaeus Hunter, 1927, Rowardleus Mackiewicz et Deutsch, 1976, Spartoides Hunter, 1929), biacetabulate, bulboloculate, bothrioloculodiscate (Biacetabulum Hunter, 1927), tholate (Hunterella Mackiewicz et McCrae, 1962), cuneifimbriate (Khawia Hsü, 1935), cuneiform (Calentinella Mackiewicz, 1974, Caryophyllaeides Nybelin, 1922, Edlintonia Mackiewicz, 1970), hastate (Pseudolytocestus Hunter, 1929), loculotholate (Bialovarium Fischthal, 1953, Pliovitellaria Fischthal, 1951), and cuneiformoloculate (Glaridacris Cooper, 1920, Isoglaridacris Mackiewicz, 1965). The same type of scolex may be shared by species of different genera or families and species of the same genus can have a scolex of conspicuously different morphology, e.g. in Promonobothrium. Scolex morphology may be therefore of limited use in generic designation.

The first molecular assessment of phylogenetic relationships of cladorchiid digeneans (superfamily Paramphistomoidea Fischoeder, 1901) from freshwater fishes based on 28S rDNA, ITS2 and cox1 sequences reveals the subfamilies Dadayiinae Fukui, 1929 and Kalitrematinae Travassos, 1933 as non-monophyletic, whereas Dadaytrema Travassos, 1931 represented by three species is monophyletic. Fourteen species of cladorchiids were found in characiform, perciform and siluriform fishes in the Neotropical Region (Brazil and Peru), with numerous new host and geographical records. The first scanning electron micrographs of seven species are presented. Two new species of dadayiine and one new species of kalitrematine paramphistomes are described. Microrchis macrovarium sp. n. from Pimelodella cristata (Müller et Troschel) (type host), Tetranematichthys quadrifilis (Kner) and Pterodoras granulosus (Valenciennes) in Brazil and Peru differs from all three congeners in that the testes are directly tandem, not oblique, and that the ovary, between the caecal ends, is widely separated from the testes. Pronamphistoma philippei sp. n. from Heros sp. in Brazil is distinguished from the type and only species, Pronamphistoma cichlasomae Thatcher, 1992, by the absence of the anterior collar-like expansion present in the type species, the presence of extramural rather than intramural pharyngeal sacs, and the unusual development of the dorsal and ventral exterior circular muscle fibre series in the acetabulum. Pseudocladorchis romani sp. n. from P. granulosus (type host), Brachyplatystoma vaillantii (Valenciennes), Calophysus macropterus (Lichtenstein), Megalodoras uranoscopus (Eigenmann et Eigenmann) and Oxydoras niger (Valenciennes) in Brazil and Peru, is most similar to Pseudocladorchis nephrodorchis Daday, 1907 but differs in the shape of the testes (irregular, versus reniform in the latter species) and the size of the ovary (as large as, or larger than, the testes in the new species). The generic diagnosis of Pronamphistoma Thatcher, 1992 is amended. Dadaytremoides parauchenipteri Lunaschi, 1989 is transferred to Doradamphistoma Thatcher, 1979 as D. parauchenipteri (Lunaschi, 1989) comb. n. based on morphological and molecular evidence.

Nanayella gen. n. is proposed to accommodate four new species of dactylogyrid monogeneans found on the gills of two species of pimelodid catfishes (Siluriformes) from the Amazon River basin in Peru and the Upper Paraná River basin in Brazil. Nanayella gen. n. is mainly characterised by the presence of modified hook pairs 5 and 6, which have a slender shank with slightly enlarged base, inconspicuous thumb, and straight point. Nanayella aculeatrium gen. n. et sp. n. (type species) from Sorubim lima (Bloch et Schneider) (type host) differs from its congeners mainly by possessing seven sclerotised spines on one side of circular ornamentation of the vaginal atrium. Nanayella fluctuatrium gen. n. et sp. n. is characterised by the presence of sclerotised undulations surrounding the circular ornamentation of the vaginal atrium. Nanayella amplofalcis gen. n. et sp. n. is typified by possessing hook pairs 1 and 2 with a robust, wide shank and a round, weakly sclerotised subunit at the base. Nanayella processusclavis gen. n. et sp. n. differs from congeners by the presence of a conspicuous posterior medial process on the dorsal bar. Urocleidoides megorchis Mizelle et Kritsky, 1969 described from the gills of S. lima in Brazil is transferred to Nanayella gen. n. as Nanayella megorchis comb. n. based on morphological and molecular evidence. Analyses of partial sequences of the 28S rRNA gene revealed all five species of the new genus to form a monophyletic clade within a bigger clade comprised of dactylogyrids of South American pimelodids.

Tapeworms of the genus Caryophyllaeus Gmelin, 1790 (Caryophyllidea: Caryophyllaeidae), common parasites of cyprinid fishes, are reviewed and taxonomic status of 42 nominal taxa that have been placed in the genus during its long history is clarified. The following seven species occurring in the Palaearctic Region are recognised as valid: C. laticeps (Pallas, 1781), C. auriculatus (Kulakovskaya, 1961), C. balticus (Szidat, 1941) comb. n. (syn. Khawia baltica Szidat, 1941), C. brachycollis Janiszewska, 1953, C. fimbriceps Annenkova-Chlopina, 1919, C. syrdarjensis Skrjabin, 1913, and newly described Caryophyllaeus chondrostomi sp. n. (= C. laticeps morphotype 4 of Bazsalovicsová et al., 2014) from common nase, Chondrostoma nasus (Linnaeus), found in Austria and Slovakia. The new species differs by the paramuscular or cortical position of preovarian vitelline follicles, a large, robust body (up to 64 mm long), conspicuously long vas deferens, flabellate scolex with small wrinkles on the anterior margin, and anteriormost testes located in a relatively short distance from the anterior extremity. Caryophyllaeus kashmirenses Mehra, 1930 and Caryophyllaeus prussicus (Szidat, 1937) comb. n. are considered to be species inquirendae, C. truncatus von Siebold in Baird, 1853 and C. tuba von Siebold in Baird, 1853 are nomina nuda. Data on the morphology, host spectra, distribution and known life-cycles of valid species are provided. Phylogenetic interrelations of four species of the genus including its type species and newly described C. chondrostomi were assessed based on an analysis of sequences of lsrDNA and cox1. A key to identification of all valid species of Caryophyllaeus is also provided.

Fourteen species (twelve adult and two larval) of nematodes belonging to the Ascaridoidea, Camallanoidea, Cosmocercoidea, Habronematoidea, Oxyuroidea, Seuratoidea and Trichinelloidea were collected from fishes in the Sudan (River Nile and Atbara Dam Lake), whereas only four species (one adult and three larval) of the Ascaridoidea, Camallanoidea, Dioctophymatoidea and Trichinelloideafrom fishes in Ethiopia (Lake Tana). The Sudanese material also contained two previously unknown species, Cucullanus mormyri sp. n. from Mormyrus caschive Linnaeus (type host), Mormyrus sp. and Marcusenius cyprinoides (Linnaeus) (all Mormyridae), and Procamallanus (Spirocamallanus) pseudospiralis sp. n. from Synodontis schall (Bloch et Schneider) (type host), S. frontosus Vaillant and S. nigrita Valenciennes (all Mochokidae), which are described based on light and scanning electron microscopical studies. Cucullanus baylisi Campana-Rouget, 1961, a little-known parasite of Synodontis spp., is redescribed in detail. A key to Procamallanus (Spirocamallanus) spp. from African inland fishes is provided. Falcaustra guiersi Vassiliadès, 1973 is considered a junior synonym of F. hexapapillata (Khalil, 1962). The findings represent several new host and geographical records. Cucullanus baylisi Lakshmi, 2000 (= a homonym to C. baylisi Campana-Rouget, 1961) is re-named as C. dubius nom. n. and is considered a species inquirenda.

Parasitological examinations of 102 specimens of the lesser great cormorant Phalacrocorax carbo sinensis (Blumenbach) from two nesting regions in the Czech Republic (South Bohemia and South Moravia) were carried out at the Institute of Parasitology, Czech Academy of Sciences (previously the Czechoslovak Academy of Sciences) in the years 1987-1992. In them, a total of 19 species of helminth parasites was found, including Trematoda (11 species), Cestoda (2), Nematoda (4) and Acanthocephala (2), which can be divided into three main groups regarding their host specificity: parasites specific for cormorants (Phalacrocorax spp.) (37%), those parasitic mainly in cormorants (16%) and non-specific parasites (47%). Of the 19 species recorded, 100% were found in South Moravia, but only 47% of these 19 species in South Bohemia. The higher number of helminth species in cormorants from South Moravia and a higher proportion of non-specific species may be associated with the presence of the large Nové Mlýny water reservoir, in addition to better ecological and environmental conditions in this warmer region. Scanning electron microscopical examination of three common nematode species parasitising cormorants, Contracaecum rudolphii Hartwich, 1964, Desmidocercella incognita Solonitsin, 1932 and Syncuaria squamata (von Linstow, 1883), revealed some taxonomically important, previously unreported morphological features, such as the cephalic structures, numbers and distribution of male caudal papillae or the shapes of spicules.

Monozoic cestodes of the recently amended genus Promonobothrium Mackiewicz, 1968 (Cestoda: Caryophyllidea), parasites of suckers (Cypriniformes: Catostomidae) in North America, are reviewed, with information on their host specificity, distribution and data on the scolex morphology of seven species studied for the first time using scanning electron microscopy (SEM). Evaluation of type and voucher specimens from museum collections and newly collected material of most species indicated the following valid nominal species: Promonobothrium minytremi Mackiewicz, 1968 (type species); P. ingens (Hunter, 1927); P. hunteri (Mackiewicz, 1963); P. ulmeri (Calentine et Mackiewicz, 1966); P. fossae (Williams, 1974) and P. mackiewiczi (Williams, 1974). Rogersus Williams, 1980 with its only species R. rogersi is transferred to Promonobothrium based on morphological and molecular data. Promonobothrium currani sp. n. and P. papiliovarium sp. n. are described from Ictiobus bubalus (Rafinesque) and Ictiobus niger (Rafinesque), and Erimyzon oblongus (Mitchill), respectively. The newly described species can be distinguished from the other congeners by the morphology of the scolex, the position of the anteriormost vitelline follicles and testes, the presence of postovarian vitelline follicles and the shape of the ovary. Molecular phylogenetic analyses of six species based on sequences of the small and large subunits of the nuclear ribosomal RNA genes (ssrDNA, lsrDNA) confirmed the monophyletic status of the genus and supported the validity of the species analysed. A key to identification of all species of Promonobothrium based on morphological characteristics is provided.

This paper summarises the results of parasitological examinations of the European eel Anguilla anguilla (Linnaeus) in the Czech Republic, carried out at the Institute of Parasitology, Czech Academy of Sciences (previously the Czechoslovak Academy of Sciences) within the period of 50 years (1958-2008). Even though this survey is limited to the Czech Republic, it provides extensive data probably incomparable with any other study anywhere regarding the number of eels examined and parasites found. A total of 723 eels was examined from 42 localities that belong to all of the three main river drainage systems in the country, i.e. the Elbe, Danube and Oder river basins. Of the 31 species of adult and larval macroparasites including Monogenea (4 species), Trematoda (3), Cestoda (3), Nematoda (11), Acanthocephala (5), Hirudinea (1), Bivalvia (1), Copepoda (1), Branchiura (1) and Acariformes (1), most of them (30) were recorded from the Elbe River basin. These parasites can be divided into three main groups regarding their host specificity: parasites specific for eels (26%), non-specific adult parasites occurring also in other fishes (61%) and non-specific larvae (13%). The highest number (19) of parasite species was recorded in the Mácha Lake fishpond system in northern Bohemia. The parasite communities in eels from the individual localities exhibited large differences in their species composition and diversity depending on local ecological conditions. The parasite fauna of A. anguilla in the Czech Republic is compared with that in other European countries. The nematode Cucullanus egyptae Abdel-Ghaffar, Bashtar, Abdel-Gaber, Morsy, Mehlhorn, Al Quraishy et Mohammed, 2014 is designated as a species inquirenda.

Using scanning and transmission electron microscopy, ultrastructure of the anterior organ and posterior funnel-shaped canal of Gyrocotyle urna Wagener, 1852 (Cestoda: Gyrocotylidea) from ratfish, Chimaera monstrosa (Holocephali), was studied for the first time. The proper anterior organ is localised at a short distance (about 170 µm) from an apical pore surrounded by a receptor field, whereas its distal end is marked by a muscular sphincter. The tegumental surface of this organ is covered with short filitriches of irregular length; large area of muscle layers traverse beneath the tegumental layer. The funnel-shaped canal of G. urna (2.5-3.0 mm long) is a specialised, muscular part of the posterior attachment organ; it opens on the rounded elevation on the dorsal body surface. The tegumental layer bears conical sclerite-like structures (up to 1.5 µm long). It produces electron-dense bodies that are transported into a canal lumen and surrounded thick muscle area mixed with numerous nerve fibres. The present ultrastructural study of G. urna indicates that gyrocotylideans share some ultrastructural characters of the anterior organ with spathebothriidean cestodes with a single anterior attachment sucker-like organ. In contrast, the unique posterior rosette attachment organ with funnel-shaped canal of the Gyrocotylidea resembles the haptor of polyopisthocotylean monogeneans in its position at the posterior end of the body and presumed origin. The above-mentioned features add more clarity to support the basal position of the Gyrocotylidea Poche, 1926 among cestodes. In addition, they also indicate a possible relationship of gyrocotylidean ancestors with monogeneans.

Frezella gen. n. is proposed to accommodate Frezella vaucheri sp. n. from poorly known auchenipterid fish, Tocantinsia piresi (Miranda Ribeiro), from the Xingú River, one of the principal tributaries of the lower Amazon River in Brazil. The new genus belongs to the Proteocephalinae because of the medullary position of the testes, ovary (yet some follicles penetrate to the cortex on the dorsal side), vitelline follicles and uterus. It differs from other proteocephaline genera in the morphology of the scolex, which includes a metascolex composed of two distinct zones: anterior, strongly wrinkled part posterior to the suckers, and posterior, sparsely folded zone. Frezella can also be differentiated by having the internal longitudinal musculature hypertrophied laterally on both sides, the presence of some ovarian follicles in the cortex on the dorsal side and the presence of additional pair of tiny, thin-walled osmoregulatory canals situated slightly dorsomedian to ventral canals. Frezella vaucheri is the first helminth parasite reported from T. piresi, which occurs in the lower reaches of the Amazon and Tocantins River basins in Brazil.

Molecular analysis of an extensive specimen collection of morphologically distinct tapeworms of the genus Caryophyllaeus Gmelin, 1790 (Cestoda: Caryophyllidea), parasites of cyprinid fishes in the Palaearctic Region, brought evidence of host-related plasticity in critical morphological characters widely used for species circumscription and classification of these tapeworms. The results obtained here do not support the traditionally used morphological concept of species-defining characters of the order Caryophyllidea Carus, 1863, especially due to high morphological plasticity of the scolex and the anterior end of the body. Several morphotypes within both Caryophyllaeus laticeps (Pallas, 1781) and Caryophyllaeus brachycollis Janiszewska, 1953 generally corresponding to different hosts and geographical regions, were recognised likely suggesting host- and geography-related intraspecific morphological variability. Genetic data confirmed euryxenous host specificity of both species, most profoundly C. laticeps, but did not support the existence of cryptic species. In fact, some of the fish hosts may harbour both of the congeneric species including several of their respective morphotypes. The pattern of morphological and genetic divergence observed in both cestode species studied indicates a scenario of possible host range extensions and subsequent parasite diversification. If molecular sequence variability and host-related morphological polymorphism are confirmed in other groups of monozoic cestodes, it may complicate species identification and straightforward classification of these parasites.

Tapeworms of the order Spathebothriidea Wardle et McLeod, 1952 (Cestoda) are reviewed. Molecular data made it possible to assess, for the first time, the phylogenetic relationships of all genera and to confirm the validity of Bothrimonus Duvernoy, 1842, Diplocotyle Krabbe, 1874 and Didymobothrium Nybelin, 1922. A survey of all species considered to be valid is provided together with new data on egg and scolex morphology and surface ultrastructure (i.e. microtriches). The peculiar morphology of the members of this group, which is today represented by five effectively monotypic genera whose host associations and geographical distribution show little commonality, indicate that it is a relictual group that was once diverse and widespread. The order potentially represents the earliest branch of true tapeworms (i.e. Eucestoda) among extant forms.

Khawia abbottinae sp. n. is described from the Chinese false gudgeon, Abbottina rivularis (Basilewsky) (Cyprinidae: Gobioninae), from the Yangtze River basin in China. The new species can be distinguished from the congeneric species mainly by the arrangements of the testes, which form two longitudinal bands (other congeneric species have the testes irregularly scattered throughout the testicular region) and their number (at maximum 85 testes versus at least 160 in the other Khawia spp.), and the morphology of the scolex, which varies from cuneiform to widely bulbate scolex, being separated from the remaining body by a short neck and possessing a smooth, blunt or rounded anterior margin. Other typical features of K. abbottinae are its small size (total length less than 1.5 cm) and body shape, with the maximum width at its first third. The distinct status of the new species was confirmed by molecular data (ssrDNA and ITS1 sequences). Phylogenetic analyses revealed a close relationship of the new species with K. rossittensis (Szidat, 1937) and K. parva (Zmeev, 1936), parasites of crucian carp and goldfish (Carassius spp.), but both species markedly differ from K. abbottinae in their morphology. Until now, five valid species of Khawia (K. abbottinae, K. japonensis, K. rossittensis, K. saurogobii and K. sinensis) have been reported from China.

Australotaenia de Chambrier et de Chambrier, 2010 has been proposed to accommodate two species of proteocephalidean cestodes from hylid frogs (Litoria spp.) in Australia. Recently, apparently congeneric cestode, for which the name A. bunthangi sp. n. is proposed, was found in the homalopsid snake Enhydris enhydris (Schneider) (Serpentes: Homalopsidae) from South-East Asia (Cambodia). This finding indicates a much wider range of definitive hosts of species of this genus, i.e. amphibians and reptiles, which is exceptional among proteocephalideans. Postcyclic parasitism, i.e. predation of the definitive host infected with sexually mature parasites, cannot be excluded but does not seem to be probable. In addition, the occurrence of A. bunthangi in the former Indochina extends the range of the geographical distribution of the genus to another zoogeographical region. The new species differs from both species of Australotaenia in the relative size of an apical organ, the diameter of which equals to that of suckers (versus much smaller in the remaining species, in which the width of the apical organ represents less than 2/3 of the diameter of the suckers), much smaller scolex and suckers (width 150 microm and diameter of suckers 50-55 microm versus 245-420 microm and 100-140 microm, respectively), and longer body (224 mm versus 57-121 mm). In addition, A. bunthangi differs from A. hylae (Johnston, 1912) (type-species of the genus) by the number of testes (46-64 versus 74-106 in A. hylae) and by the ovary width/proglottis width ratio (55-65% versus 68-71% in A. hylae). Australotaenia bunthangi differs from A. grobeli de Chambrier et de Chambrier, 2010 by relative size of the cirrus-sac (its length represents 18-24% of the width of the proglottis versus 27-33% in A. grobeli) and by the diameter of the embryophore (25-27 microm versus 18-23 microm in A. grobeli).

Spermiogenesis and ultrastructure of mature spermatozoon of the caryophyllidean cestode Hunterella nodulosa, a parasite of suckers (Catostomidae), have been studied by transmission electron microscopy. This monozoic tapeworm is unique in its mode of attachment and represents the second North American species studied. The process of spermiogenesis of H. nodulosa follows the general pattern already described in other caryophyllideans. The most characteristic feature is the presence of a slight rotation of the flagellar bud, which seems to be a typical character of spermiogenesis in this cestode group. The mature spermatozoon of H. nodulosa is characterized by the presence of one axoneme of 9 + "1" type of the trepaxonematan flatworms surrounded by a semi-arc of cortical microtubules in its anterior extremity, parallel nucleus and cortical microtubules arranged in a parallel pattern, which corresponds to the Type III pattern of cestode spermatozoa according to Levron et al. (2010). Comparison of the present data with those available for other caryophyllideans did not reveal substantial differences, even though they belong to different families, infect different hosts (catostomid, cyprinid and siluriform fishes) and occur in distant zoogeographical regions. This indicates uniformity of the process of sperm formation and spermatozoon ultrastructure in one of the evolutionarily most ancient groups of tapeworms.

Specimens of the nematode genus Rhabdochona Railliet, 1916 (Rhabdochonidae) were collected during helminthological examination of four species of cyprinid fishes in two rivers of the Amur River basin in the Russian Far East (Primorsky Region) in June 2011. Detailed light microscopical (LM) and scanning electron microscopical (SEM) examinations (the latter used for the first time for the reported nematode species) of the available material revealed the presence of three inadequately described nominal species of this genus: R. (Rhabdochona) denudata (Dujardin, 1845) from the spotted steed Hemibarbus maculatus Bleeker (Gobioninae), and R. (Rhabdochona) longispicula Belous in Roytman, 1963 and R. (Globochonoides) coronacauda Belous, 1965 from Culter alburnus Basilewsky (Cultrinae) in the Ilistaya River. Detailed morphological study of these worms, especially SEM examination, made it possible to reveal some previously unreported morphological features (e.g., the presence of sublabia or the character of ventral precloacal ridges) and to confirm other taxonomically important characters such as the shape of deirids, number of anterior prostomal teeth, number and situation of lateral preanal and postanal papillae or the detailed structure of the crown-like formation on the female tail tip in R. coronacauda. Unidentifiable Rhabdochona (Rhabdochona) gravid females were recorded from the humpback Chanodichthys dabryi (Bleeker) (Cultrinae) in the Ilistaya River and from the Amur minnow Rhynchocypris lagowskii (Dybowski) (Leuciscinae) in the Komissarovka River.

A new genus, Ritacestus, is proposed to accommodate Ritacestus ritaii (Verma, 1926) comb. n. (syn. Proteocephalus ritaii), a parasite of the catfish Rita rita (Hamilton) in India. The new genus, which is placed in the Gangesiinae, is characterized by (i) a small, subspherical scolex formed by four large lobes separated from one another by longitudinal grooves, with a large, widely oval to pyriform rostellum-like apical organ, larger than suckers and possessing an apical hemispherical depression; (ii) paramuscular and cortical position of some vitelline follicles (most follicles are situated medullary); (iii) ventral and dorsal bands of vitelline follicles usually uninterrupted ventral to terminal genitalia and reaching to the posterior margin of proglottides; (iv) the vagina always anterior to the cirrus-sac; (v) a large size of the body (length up to 51 cm); and (vi) development of the uterus of type 2. In its morphology, especially shape of the scolex and apical organ, and paramuscular and cortical position of some vitelline follicles, Ritacestus resembles Postgangesia Akhmerov, 1969, but differs in the presence of a genital atrium (both genital pores of Postgangesia are separate), the anterior position of the vagina (almost always posterior in the latter genus), position of vitelline follicles in cross sections (dorsal and ventral bands in Ritacestus versus only a lateral band in the latter genus), and dorsal excretory canals indistinguishable in mature and gravid proglottides of R. ritaii (well developed in Postgangesia spp.). The type and only species of the genus, R. ritaii, is redescribed on the basis of new material from the type host from the Ganges River basin in India and its neotype is designated.

Monozoic cestodes of the genus Khawia Hsü, 1935 (Caryophyllidea: Lytocestidae), parasites of cyprinid fish in Europe, Asia, Africa and North America, are revised on the basis of taxonomic evaluation of extensive materials, including recently collected specimens of most species. This evaluation has made it possible to critically assess the validity of all 17 nominal species of the genus and to provide redescriptions of the following seven species considered to be valid: Khawia sinensis Hsü, 1935 (type species); K. armeniaca (Cholodkovsky, 1915); K. baltica Szidat, 1941; K. japonensis (Yamaguti, 1934); K. parva (Zmeev, 1936); K. rossittensis (Szidat, 1937); and K. saurogobii Xi, Oros, Wang, Wu, Gao et Nie, 2009. Several new synonyms are proposed: Khawia barbi Rahemo et Mohammad, 2002 and K. lutei Al-Kalak et Rahemo, 2003 are synonymized with K. armeniaca; K. coregoni Kritscher, 1990 with Caryophyllaeus laticeps (Pallas, 1781) (family Caryophyllaeidae); K. cyprini Li, 1964 and K. iowensis Calentine et Ulmer, 1961 with K. japonensis; K. dubia (Szidat, 1937) (syn. Bothrioscolex dubius Szidat, 1937) with K. rossittensis; and Tsengia neimongkuensis Li, 1964 and T. xiamenensis Liu, Yang et Lin, 1995 with K. sinensis. Khawia prussica (Szidat, 1937) (syn. Bothrioscolex prussicus Szidat, 1937) is considered to be species incertae sedis, but its morphology indicates it may belong to Caryophyllaeus Gmelin, 1790 (Caryophyllaeidae). The molecular analysis of all seven valid species, based on comparison of sequences of two nuclear ribosomal and two mitochondrial genes, has shown that the species form three major groups clustered according to their fish hosts. Five species from common and crucian carp and goldfish were grouped together, whereas K. armeniaca from barbels (Barbinae) and K. baltica from tench (Tinca) formed separate clades. In contrast, geographical distribution does not seem to play a crucial role in grouping of individual taxa. A phylogenetic tree based on morphological characters was incongruent with that inferred from molecular data, which indicates that some morphological traits may be homoplastic. A key to identification of all species of Khawia based on morphological characteristics is provided.

Specimens of three little-known species of Rhabdochona (Nematoda: Rhabdochonidae) were collected during occasional examinations of some freshwater fishes in India: R. (Rhabdochona) hellichi turkestanica (Skryabin, 1917) in Schizothorax sp. (Cyprinidae, Cypriniformes) from the Lodhomakhola and Rangit Rivers, West Bengal and Sikkim, respectively; R. (R.) hospeti Thapar, 1950 in Tor sp. (Cyprinidae) from the Rangit River; and R. (Globochona) mazeedi Prasad et Sahay, 1965 in Clupisoma garua (Hamilton) (Schilbeidae, Siluriformes) from the Farakka Dam Lake, West Bengal. Their detailed light and electron microscopical studies revealed some taxonomically important, previously not observed features and made possible their redescription. Fourth-stage larvae of R. hospeti are described for the first time. Rhabdochona hellichi turkestanica (syns. R. denudata filamentosa Bykhovskaya-Pavlovskaya, 1936, R. kashmirensis Thapar, 1950, R. schizothoracis Siddiqi et Khattak, 1984) is proposed as a subspecies, differing from the nominotypical subspecies R. hellichi hellichi (Srámek, 1901) mainly in the shape of the distal end of the left spicule, molecular data and geographical distribution. Rhabdochona moraveci Katoch et Kalia, 1991 (a homonym to R. moraveci Duggal et Kaur, 1987) is renamed R. indica nom. n. The following six species are considered new junior synonyms of R. hospeti: Comephronema [sic] mackiewiczi Malhotra et Rautela, 1984, Rhabdochona moraveci Duggal et Kaur, 1987, R. bifidum Kakar et Bilqees, 2007, R. uvaginus Kakar et Bilqees, 2007, R. bolani Kakar, Bilqees et Ahmad, 2008 and R. cephalodiverticula Kakar, Bilqees et Ahmad, 2008. Rhabdochona edentati Paul et Majumdar, 1994 is considered a species incertae sedis.

Ophiotaenia georgievi sp. n. (Proteocephalidea: Proteocephalinae) is described from the intestine of endemic colubrid snake Leioheterodon geayi Mocquard (Colubridae) from Antananarivo in Madagascar. The new species is the first species of Ophiotaenia La Rue, 1911 reported from Madagascar. It differs from all Ophiotaenia species parasitic in African snakes in the possession of a three-layered embryophore of eggs (other African species have two-layered embryophore). Furthermore, O. georgievi can be distinguished by the number of testes (92-140), width of the scolex (225-235 jm), total body length (57 mm), cirrus-sac length/proglottis width ratio (19-32%), and number of lateral uterine diverticula (23-28 on each side). Ophiotaenia georgievi represents the second proteocephalidean cestode reported from Madagascar, the first one being Deblocktaenia ventosaloculata (Deblock, Rosé et Broussart, 1962), a parasite from Ithycyphus miniatus. A list of Ophiotaenia species parasitic in venomous and non-venomous snakes is provided and possible existence of other new congeneric species in snakes from Madagascar is discussed.

An ultrastructural study of the ovarian follicles and their associated oviducts of the cestode Gyrocotyle urna Grube et Wagener, 1852, a parasite from the spiral valve of the rabbit fish, Chimaera monstrosa L., was undertaken. Each follicle gives rise to follicular oviduct, which opens into one of the five collecting ducts, through which pass mature oocytes. These collecting ducts open into an ovarian receptacle which, in turn, opens via a muscular sphincter (the oocapt) to the main oviduct. The maturation of oocytes surrounded by the syncytial interstitial cells within the ovarian follicles of G. urna follows a pattern similar to that in Eucestoda. The ooplasm of mature oocytes contain lipid droplets (2.0 x 1.8 microm) and cortical granules (0.26 x 0.19 microm). The cytoplasm of primary and secondary oocytes contains centrioles, indicating the presence of the so-called "centriole cycle" during oocyte divisions. A morphological variation between different oviducts was observed. The luminal surface of the follicular and the collecting oviducts is smooth. The zones of the septate junctions are present within the distal portion of the net-like epithelial wall of the collecting ducts close to the ovarian receptacle. The syncytial epithelial lining of the ovarian receptacle, oocapt and main oviduct is covered with lamellae and cilia. Cortical granules secreted from mature oocytes occur freely within the lumen of the main oviduct that functions as a fertilisation canal. A division of the ovary into separated parts with their own collecting ducts as that typical of Gyrocotyle has been observed in neodermates, basal monogenean family Chimaericolidae, and Neoophora (some Proseriata and Fecampiidae). Ultrastructural data thus reveal several unique morphological characteristics of gyrocotylideans, the most basal taxon of tapeworms (Cestoda).

A comparative study of the scoleces of caryophyllidean tapeworms (Cestoda: Caryophyllidea), parasitic in cypriniform fishes in the Palaearctic Region, was carried out using light and scanning electron microscopy. Three-dimensional pictures of the scoleces of 18 species of caryophyllidean cestodes of the Capingentidae (1 species), Caryophyllaeidae (7) and Lytocestidae (10), and outlines of the scoleces and anterior extent of the testes and vitelline follicles of 19 Palaearctic taxa were documented. Both species of Atractolytocestus Anthony, 1957 possess a bulboacuminate scolex, whereas species of Archigetes Leuckart, 1876 have fossate scoleces of the bothrioloculodiscate type, with loculi, bothrium-like depressions and an apical disc. Breviscolex orientalis Kulakovskaya, 1962, the only member of the Capingentidae, has a cuneiform scolex, as do both taxa of the lytocestid genus Caryophyllaeides Nybelin, 1922. The scoleces of two species of Caryophyllaeus Gmelin, 1790 are flabellate, whereas that of the congeneric C. fimbriceps Annenkova-Chlopina, 1919 is cuneicrispitate. Khawia Hsü, 1935, the most specious Palaearctic genus, with seven taxa that we consider to be valid, has the highest diversity in scolex morphology: semi-bulbate, flabellate, cuneiform, cuneifimbriate, truncated cuneiform-flabellate and festoon-like. Species of Monobothrium Nybelin, 1922 have either a digitiform scolex with widened posterior part or cuneiform, with lateral auricular extensions. Paracaryophyllaeus gotoi (Motomura, 1927) is characteristic in its possessing a bulbate scolex, whereas Paraglaridacris limnodrili (Yamaguti, 1934) has a fossate scolex of the bulboloculate type with bothrium-like depressions and feebly developed lateral loculi. Anterior extent of the testes and vitelline follicles and their mutual position show a somewhat higher variability than scolex shape, with intraspecific variation in some taxa, such as Atractolytocestus sagittatus (Kulakovskaya et Akhmerov, 1965), B. orientalis, Khawia armeniaca (Cholodkovsky, 1915) and K. sinensis Hsii, 1935. Based on scolex morphology and relative position of the anterior testes and vitelline follicles, a key is provided to facilitate the routine identification of 20 Palaearctic caryophyllidean taxa.

The scolex surface of the mature spathebothriidean Cyathocephalus truncatus (Pallas, 1781), a parasite of the brown trout Salmo trutta fario L., was studied using scanning and transmission electron microscopy. A particular attention was paid to microtriches, unique structure on the surface of the Cestoda. The scolex of C. truncatus is covered with two types of filiform microtriches (filitriches): aciculate (approximately 3 microm long) and capillate (approximately 10 microm long). Capillate microtriches, which have never been reported in any other spathebothriideans, are described for the first time using transmission electron microscopy. The tegument covered with filiform microtriches only (no spiniform microtriches are present) is typical of cestode groups supposed to be the most basal, e.g., Gyrocotylidea, Spathebothriidea, and Caryophyllidea.

The surface structures and gland cells of the posterior rosette organ of Gyrocotyle urna Grube et Wagener, 1852, a member of the group presumed to be the most basal of the tapeworms (Cestoda: Gyrocotylidea), was studied by scanning electron and transmission electron microscopy. Surface structures on the outer (oriented away from the intestinal wall) and inner (in contact with the intestinal wall) rosette surfaces differ from each other and represent a transitional form between microvilli and microtriches typical of tapeworms (Eucestoda). The inner surface of the rosette possesses numerous glands. On the basis of the size and electron-density of their secretory granules, three types of unicellular gland cells can be distinguished. The least common type (Type I) is characterized by the production of small, round, electron-dense granules of about 0.3 microm in diameter, whereas another type of secretion (Type II) is formed from homogenous, moderately electron-dense, spheroidal granules of about 0.7 microm in diameter. The most common type of glands (Type III) is recognized by a secretion comprising large, elongate, electron-dense granules of about 1 microm long and 0.5 microm broad. The secretory granules of the three types of the glands are liberated by an eccrine mechanism and the gland ducts open via small pores on the inner rosette surface. The complex of secretory glands of the posterior rosette of G. urna is similar to those in the anterior attachment glands of monogeneans (as opposed to the types of glands present in other helminth groups). However, the tegumental surface structures of Gyrocotyle are supporting evidence for the relationship between the Gyrocotylidea and Eucestoda.

Bothriocephalidean tapeworms parasitic in the blackfish, Centrolophus niger (Gmelin) (Perciformes: Centrolophidae), are redescribed on the basis of the evaluation of freshly collected specimens and museum material. This evaluation enabled us to supplement species diagnoses by new morphological characters of potential use for phylogenetic analyses, including the data from scanning electron microscopical observations, and to provide a key to identification of the following four species occurring in this fish: Amphicotyle heteropleura (Diesing, 1850); Milanella familiaris Kuchta et Scholz, 2008 (both Triaenophoridae); Bothriocotyle solinosomum Ariola, 1900; and Echinophallus wageneri (Monticelli, 1890) (both Echinophallidae). Large spiniform microtriches were observed on the surface of the posterodorsal margin of segments of B. solinosomum, E. wageneri and M. familiaris. The invalidity of Atelemerus Guiart, 1935, first proposed by Bray et al. (1994), is supported by the present data and its type species, A. acanthodes Guiart, 1935, is newly synonymised with E. wageneri.

A survey of proteocephalidean cestodes found in the firewood catfish Sorubimichthys planiceps (Spix et Agassiz) from the Amazon River is provided. The following taxa parasitic in S. planiceps are redescribed on the basis of their type specimens and material collected recently in the Amazon River, near the type localities in Brazil, and in Iquitos, Peru: Monticellia lenha Woodland, 1933; Nomimoscolex lenha (Woodland, 1933) (syn. Proteocephalus lenha Woodland, 1933); and Monticellia megacephala Woodland, 1934, for which a new genus, Lenhataenia, is proposed, with L. megacephala (Woodland, 1934) comb. n. as its type and only species. The new genus is a member of the Monticelliinae, i.e. has all genital organs in the cortex, and is most similar to Chambriella in possessing biloculate suckers and lacking a metascolex. It differs in the morphology of the cirrus-sac that contains a strongly coiled, thick-walled internal sperm duct (vas deferens) and a muscular cirrus of the appearance typical of most proteocephalideans, whereas that of Chambriella is sigmoid, with voluminous, tightly sinuous thick-walled internal sperm duct. In addition, Lenhataenia possesses a well developed internal musculature, whereas the internal musculature of Chambriella is weakly developed, formed by a low number of muscle fibres. The scolex morphology and distribution of microtriches of Peltidocotyle lenha (Woodland, 1933) (syn. Othinoscolex lenha Woodland, 1933 and Othinoscolex myzofer Woodland, 1933), Chambriella sp. and Choanoscolex sp. are described using scanning electron microscopy. The two latter taxa may be new for science and are reported from S. planiceps for the first time.

A comparative study of the scolex hook morphology of five species of tapeworms of the genus Triaenophorus Rudolphi, 1793 (Cestoda: Pseudophyllidea), parasites of pikes (Esox lucius L. and E. reichertii Dybowski) in the Palaearctic Region, was carried out. Measurements of scolex hooks of 81 plerocercoids and 492 adults from different hosts and regions were compared using basic statistics and forward stepwise linear discriminant analysis. The shape of the scolex and that of tridental hooks were found to be suitable only for differentiation of the taxa with a similar shape of hooks, i.e. Triaenophorus nodulosus (Pallas, 1781) from T. amurensis Kuperman, 1968, and T. crassus Forel, 1868 from T. meridionalis Kuperman, 1968 and T. orientalis Kuperman, 1968, respectively. In contrast, discriminant analysis did not enable reliable separation of specimens of individual taxa of these two morphological groups due to high intraspecific variability and overlaps between species. This was reflected in low classification efficiencies (average 83%) of all species of the T. crassus group, whereas all T. amurensis specimens were misidentified as T. nodulosus. The new data also considerably enlarge (up to twofold) the size range of the species described by Kuperman in 1968, which invalidates suitability of the most important discriminant characteristic, the width of the basal plate, for delimitation of Triaenophorus species. Based on the present data, all Kuperman's taxa are considered to represent only distinct geographical populations of T. nodulosus and T. crassus. As a result, T. anmurensis is synonymized with T. nodulosus, whereas T. orientalis is considered to be a synonym of T. crassus. Previous synonymisation of T. ineridionalis with T. crassus, first proposed by Dubinina (1987), is also accepted.

The vitellogenesis of Paraechinophallus japonicus (Yamaguti, 1934), the first pseudophyllidean tapeworm of the family Echinophallidae studied using transmission electron microscope, is described on the basis of ultrastructural observations of specimens from the benthopelagic fish Psenopsis anomala (Temminck et Schlegel, 1844) (Perciformes: Centrolophidae). The process of vitellogenesis in P. japonicus follows the same general pattern observed in other tapeworms. Five stages of vitellocyte development have been distinguished. The first stage corresponds to immature cells containing ribosomes and mitochondria. The second stage of development is characterized by the appearance of granular endoplasmic reticulum and Golgi complexes, formation of shell globules and lipid droplets at the periphery of the cell cytoplasm. Vitellocyte of the third stage presents accumulation of shell globules and lipid droplets. During the fourth stage, shell globule clusters are formed, and lipid droplets and rosettes of alpha-glycogen are accumulated. Mature vitelline cells are characterized by a great number of lipid droplets with glycogen in the centre of the cytoplasm, whereas shell globule clusters are situated more peripherally. The interstitial tissue of vitelline follicles of P. japonicus is syncytial with long cytoplasmic projections extending between vitelline cells. The presence of a large amount of lipid droplets in the vitelline cytoplasm within the eggs of P. japonicus may be related to egg accumulation in the uterine sac.

Holotype and paratype of Ascocotyle (Phagicola) rara Arruda, Muniz-Pereira et Pinto, 2002, a heterophyid trematode recently described on the basis of two worms collected by Lauro Travassos in 1921 in the intestine of Ixobrychus exilis (Gmelin) from Brazil, were studied. The morphology of the worms revealed their conspecificity with Ascocotyle (Phagicola) angeloi Travassos, 1928 found in the same host. Both the taxa have a similar length (between 600 and 900 microm) and shape of the body (long pyriform), the long intestinal caeca reaching to the ovarian level, a long posterior muscular prolongation of the oral sucker and the prepharynx, transverse uterine loops situated between the ventral sucker and testes, and the gonotyl with more than 20 digitiform pockets. Consequently, A. (P.) rara is proposed as a junior synonym of Ascocotyle (Phagicola) angeloi.