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.
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.