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. amurensis is synonymized with T. nodulosus, whereas T. orientalis is considered to be a synonym of T. crassus. Previous synonymisation of T. meridionalis with T. crassus, first proposed by Dubinina (1987), is also accepted.
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.
Larvae (metacestodes) of tapeworms of the cyclophyllidean family Gryporhynchidae (previously included in the Dilepididae) occur in different internal organs of fresh- and brackish water fish (110 fish species of 27 families in 12 orders reported), which serve as the second intermediate hosts. The species composition, spectrum of fish hosts, sites of infection, and geographical distribution of gryporhynchids recorded from fish are reviewed here on the basis of literary data and examination of extensive material from helminthological collections. Metacestodes of the following genera have been found in fish: Amirthalingamia Bray, 1974 (1 species), Ascodilepis Guildal, 1960 (1), Cyclustera Fuhrmann, 1901 (4), Dendrouterina Fuhrmann, 1912 (1), Glossocercus Chandler, 1935 (3), Neogryporhynchus Baer et Bona, 1960 (1), Paradilepis Hsü, 1935 (5), Parvitaenia Burt, 1940 (2), and Valipora Linton, 1927 (3). However, most published records concern only three species, namely Neogryporhynchus cheilancristrotus (Wedl, 1855) from the intestinal lumen, Paradilepis scolecina (Rudolphi, 1819) from the liver and mesenteries, and Valipora campylancristrota (Wedl, 1855) from the gall bladder of cyprinids and other fish in the Palaearctic Region. Data on other species as well as reports from other regions are very scarce and almost no information is available from Australia, tropical Asia and South America. A recent study of gryporhynchid metacestodes from Mexico (Scholz and Salgado-Maldonado 2001), which reported 13 species, suggested that they may be more common than indicated by records in the literature. Although only a few cases of pathogenic influence of larvae on fish hosts have been reported, the veterinary importance of gryporhynchids remains to be assessed on the basis of more detailed studies. The data available indicate a strict host and site specificity of some species whereas others occur in a wide spectrum of fish hosts and are not strictly site-specific. Evaluation of Paradilepis larvae from the liver of salmonid fish from British Columbia, Canada, identified as P. simoni Rausch, 1949 by Ching (1982), has shown that they probably belong to two species, P. simoni and P. rugovaginosus Freeman, 1954. Metacestodes of the latter species and those of Cyclustera magna (Baer, 1959) from the intestinal wall of Tilapia zillii (Gervais) from Kenya are reported from fish for the first time.
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 (Šrámek, 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.
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.
Two fish cestodes, the little-known Eubothrium fragile (Rudolphi, 1802) and E. rugosum (Batsch, 1786), the type species of the genus Eubothrium Nybelin, 1922, are redescribed on the basis of new material from twaite shad, Alosa fallax (Lacépède, 1803), from England and burbot, Lota lota (Linnaeus, 1758), from Russia, respectively. The tapeworms are compared with two other species of the genus, E. crassum (Bloch, 1779) and E. salvelini (Schrank, 1790), common parasites of salmonid fish in the Holarctic. The most notable differential characters are the size and the shape of the scolex (smaller and oval in E. fragile), the shape of the apical disc (four or more indentations in E. crassum), the number and size of the testes (the largest and least numerous in E. rugosum), and the position and size of the vitelline follicles (almost entirely cortical in distribution in E. fragile and E. crassum versus largely medullary in E. rugosum and E. salvelini). A comparison of species has also shown the morphological similarity of the freshwater species (E. rugosum and E. salvelini) on one hand and those of marine origin, E. fragile and E. crassum, on the other, with the latter species occurring also in fresh waters. A key to the identification of the species studied is also provided.
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.
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.