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
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.
The morphology of the scoleces of 11 Proteocephalus species, parasites of freshwater fish in the Palaearctic Region, was compared using light and scanning electron microscopy. The following taxa were evaluated: Proteocephalus ambiguus (Dujardin, 1845); P. cernuae (Gmelin, 1790); P. exiguus La Rue, 1911; P. filicollis (Rudolphi, 1802); P. macrocephalus (Creplin, 1825); P. osculatus (Goeze, 1782); P. percae (Müller, 1780); P. pollunicola Gresson, 1952; P. sagitlus (Grimm, 1872); P. thymalli (Annenkova-Chlopina, 1923); and P. torulosus (Batsch, 1786). Some features as overall shape of the scolex, its size, shape and size of an apical sucker were found to be fairly stable and species-specific. The taxa more easily distinguishable from congeners on the basis of their scolex morphology were P. cernuae, P. macrocephalus, P. osculatus, P. percae and P. torulosus. The taxonomic importance of the scolex is discussed.
The male genitalia of the fritillary butterfly Issoria lathonia (L.) were examined and reconstructed based on sagittal and horizontal sections. Nine intrinsic muscles were identified consistent with previous results. The retractor of the anal tube probably operates the "rectal plate", a large, sclerotised, arched plate present ventral to the rectum and dorsal to the phallus in all Issoria s. str. species. The function of the rectal plate is still largely unknown, but it has presumably an important function during copulation. The retractor of the phallus inserts on the phallus, and also on a small, ventral sclerite in the anellus. The retractor of the vesica is smaller in I. lathonia than its counterpart in other Argynnini and originates more centrally inside the phallus. The tergal sclerite, common in most Argynnini, has no attaching muscle and its evolutionary origin remains unclear. The presence of an intrinsic muscle (i3) originating on the tegumen and inserting on the valve in Argynnini cannot be confirmed here. Though generally absent in butterflies, this muscle has been reported once in the North American Argynnis subgenus Speyeria.
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 × 1.8 µm) and cortical granules (0.26 × 0.19 µm). 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).