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., Daniel Barčák, Mikuláš Oros, Vladimíra Hanzelová, Tomáš Scholz., and Obsahuje bibliografii
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., Mikuláš Oros, Jan Brabec, Roman Kuchta, Anindo Choudhury, Tomáš Scholz., and Obsahuje bibliografii
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.
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.
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 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 Hsü, 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.
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., Mikuláš Oros, Dalibor Uhrovič, Anindo Choudhury, John S. Mackiewicz and Tomáš Scholz., and Obsahuje bibliografii
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.