Malformations of the gill filaments of the ruffe Gymnocephalus cernuus (L.) (Pisces) caused by echinostomatid metacercariae.

In parasite surveys of fishes from Lake Balaton and its tributaries in Hungary, infections with metacercariae of a species of the digenean genus Echinochasmus (Trematoda: Echinostomatidae) were found in seven species of fish. In ruffe, Gymnocephalus cernuus, malformations of the gill filaments apparently caused by these infections were observed. These malformations were in the form of bifurcations of the filaments at about their mid-length. At the point where the filaments bifurcate, an Echinochasmus metacercaria was always embedded in the cartilaginous ray of the gill filament. All specimens of the ruffe were found to be infected by these metacercariae, and each ruffe specimen was infected by 30-300 metacercariae. Such a bifurcation was found in all of the ruffe specimens, but, apart from these gill malformations, the metacercariae produced only local changes in the cartilage. In the other six infected fish species, only local signs were observed in the cartilage. Experimental infections of chicks with metacercariae resulted in the finding of the sexual adult (marita) of an unidentified species of Echinochasmus. ITS sequences of the adult and metacercaria corresponded with each other, and also with a cercaria isolated from a gravel snail (Lithoglyphus naticoides), with a 99.5-100% similarity.

Synopsis of the species of Chloromyxum Mingazinni, 1890 (Myxozoa: Myxosporea: Chloromyxidae).

A synopsis of the species of Chloromyxum Mingazinni, 1890 (Myxozoa: Myxosporea: Chloromyxidae) is presented, including 140 nominal species. For each species the most relevant morphological and morphometric characteristics are indicated. Included are data on the site of infection within the host, the original host and the host locality, plus a full bibliography of the original records for these species. A diagrammatic illustration of a spore of each species is also provided.

Cryptic species of Didymobothrium rudolphii (Cestoda: Spathebothriidea) from the sand sole, Solea lascaris, off the Portuguese coast, with an analysis of their molecules, morphology, ultrastructure and phylogeny.

Didymobothrium rudolphii (Cestoda: Spathebothriidea) was collected seasonally from the sand sole, Solea lascaris, off the northern, central and southern areas of the Portuguese coast. Morphological and molecular analyses were conducted in order to examine the possible existence of cryptic species and to facilitate the circumscription of their morphological boundaries. Data were compared between D. rudolphii specimens from each of the 3 geographical areas and 4 seasons, and principal components analysis of 18 morphological characters was used to detect differences. Two distinct genotypes were present with sequence divergences of 1.9% and 2.1% in the large subunit (lsrDNA) and second internal transcribed spacer (ITS-2) of ribosomal DNA (rDNA), respectively. The less common 'central' genotype was present only off the central area from summer to winter, whereas the 'common' genotype was present throughout the year off the northern and southern areas, but only during spring in the central area. No sequence variation was found within each genotype. The presence of 2 distinct genetic entities was supported by morphological analyses, which showed the 'central' genotype specimens to be more slender and elongate, although morphometric ranges overlapped considerably for most characters. Molecular phylogenetic analysis of 4 of the 5 known genera of the Spathebothriidea showed Spathebothrium to be the earliest branching lineage and the 2 genotypes of Didymobothrium formed a sister group to Cyathocephalus. The concordance of genetic differences with variation in host diet according to season and locality could account for sympatric speciation occurring in the central region of the Portuguese coast.

Diplectanocotyla Yamaguti, 1953 (Monogenea: Diplectanoidea) from Megalops cyprinoides (Broussonet) (Teleostei: Megalopidae) off Peninsular Malaysia.

Two known and two new species of Diplectanocotyla Yamaguti, 1953 (D. gracilis Yamaguti, 1953, D. megalopis Rakotofiringa & Oliver, 1987, D. langkawiensis n. sp. and D. parva n. sp.) were collected from Megalops cyprinoides (Megalopidae) off Langkawi, Kedah and Matang, Perak, Peninsular Malaysia. All four species possess similar types of sclerotised male and female reproductive structures and similar soft anatomical features. The squamodisc sclerites of all four species have spine-like projections with varying degrees of visibility and shapes (sharp-pointed to triangular). In D. megalopis and D. langkawiensis n. sp. the spines are sharp-pointed and distinct on sclerites from rows 5-6 onwards. In D. gracilis and D. parva n. sp. the sclerite spines are triangular, lightly sclerotised and occur on almost all of the sclerites. D. parva n. sp. has comparatively the smallest set of anchors, bars, squamodiscs and squamodisc suckers. The anchors and bars of the other three species are almost similar in overall size, and the main distinguishing feature is the relative lengths of the inner and outer roots of the ventral anchors. In D. gracilis the outer root is very much smaller than the inner root and they are disposed almost at a right angle to each other. In D. megalopis the outer root is usually about half the length of the inner root and the roots are inclined at c.60 degrees to each other. In D. langkawiensis n. sp. the roots are inclined at c.40 degrees degrees and the outer root is of a similar length or only slightly shorter than the inner root. The openings of the two squamodisc suckers of all four Diplectanocotyla species are surrounded by tiny scale-like spines. Bifid tegumental spines are found in the posterior region of all four species, differing only in their extent: in D. parva n. sp. the tegumental spines are only distributed in the peduncular region and not beyond, whilst in the other three species the tegumental spines extend from the posterior level of the testis to the end of the peduncle. An amended diagnosis of Diplectanocotyla and a key to its species are appended.

Saturnius minutus n. sp. and S. dimitrovi n. sp. (Digenea: Hemiuridae) from Mugil cephalus L. (Teleostei: Mugilidae), with a multivariate morphological analysis of the Mediterranean species of Saturnius Manter, 1969.

Three species of the bunocotyline genus Saturnius Manter, 1969 are described from the stomach lining of mugilid fishes of the Mediterranean and Black Seas. Two of the species are new: S. minutus n. sp. occurs in Mugil cephalus off the Mediterranean coast of Spain; and S. dimitrovi n. sp., a parasite of M. cephalus off the Bulgarian Black Sea coast and the Spanish Mediterranean coast, was originally described as S. papernai by Dimitrov et al. (1998). In addition, S. papernai Overstreet, 1977 is redescribed from M. cephalus off the Spanish Mediterranean coast and from Liza aurata and L. saliens off the Bulgarian Black Sea coast. The three species are distinguished morphometrically using univariate and multivariate analyses. These results were verified using Linear Discriminant Analysis which correctly allocated all specimens to their species designations based on morphology (i.e. 100% successful classification rate) and assigned almost all specimens to the correct population (locality). The following variables were selected for optimal separation between samples: the length of the forebody, ventral sucker and posterior testis, the length and width of the posteriormost pseudosegment, and the width of the muscular flange at ventral sucker level.

Robinia aurata n. g., n. sp. (Digenea: Hemiuridae) from the mugilid Liza aurata with a molecular confirmation of its position within the Hemiuroidea.

Robinia aurata n. g., n. sp. is described from Liza aurata (Mugilidae), the golden grey mullet, from the Ebro Delta, Spanish Mediterranean. The new genus differs from all other hemiurid genera in the combined possession of muscular flanges and a vestigial ecsoma. Within the Bunocotylinae, which currently accommodates 2 genera, Bunocotyle and Saturnius, the new genus exhibits a unique combination of blind caeca, Juel's organ, post-ovarian bulk of the uterus in the hind-body, and tegumental papillae surrounding the oral and ventral sucker apertures. Furthermore, Robinia n. g. differs from both Bunocotyle and Saturnius in the nature of the muscular extensions around the oral sucker, with the shape of a muscular belt in the latter and numerous muscular papillae in the former. The phylogenetic hypothesis for the Bunocotylinae developed from sequence data analyses based on partial lsrDNA and complete ssrDNA combined (22 species) and V4 domain of the ssrRNA gene (37 species) supports the erection of the new genus and confirms its position within the Hemiuroidea. Both molecular analyses confirmed the monophyly of the Hemiuroidea, its division into 2 major clades and the polyphyly of the Derogenidae, as in previous studies, and suggest that the Gonocercinae (with 2 genera, Gonocerca and Hemipera), may require a distinct familial status. Finally, there was poor support for the distinct status of the Lecithasteridae and Hemiuridae, following previous suggestions based on different sequence data sets. A key to genera of the Bunocotylinae is presented.

A new species of Nasicola Yamaguti, 1968 (Monogenea: Capsalidae) from the nasal cavities of Thunnnus obesus and a redescription of N. klawei (Stunkard, 1962) from T. albacares off Brazil.

Two species of Nasicola Yamaguti, 1968 are described from the nasal cavities of tunas ( Thunnus spp.) from off the coast of Brazil: N. brasiliensis n. sp. from T. obesus (Lowe) and the type-species, N. klawei (Stunkard, 1962), from T. albacares (Bonn.). The new species is differentiated from N. klawei on the basis of the large number of testes and from N. hogansi Wheeler & Beverley-Burton, 1986 by its greater body-size, proportionately smaller haptor and smaller number of marginal spines. The host-specificity of Nasicola spp. is commented upon.

Dactylogyridean monogeneans of the siluriform fishes of the Old World.

This is a catalogue and discussion of the known dactylogyridean monogenean genera of siluriform fishes of the Old World. Of a total of 38 nominal genera, only 19 are considered valid. Seventeen of these 19 genera are currently in the Ancyrocephalidae (containing the Ancyrocephalinae and Ancylodiscoidinae), whilst the other two (Neocalceostoma and Neocalceostomoides) are in the Neocalceostomatidae. The 17 genera are Anchylodiscus, Ancylodiscoides, Bagrobdella, Bifurcohaptor, Bychowskyella, Chauhanellus, Cornudiscoides, Hamatopeduncularia, Mizelleus, Paraquadriacanthus, Pseudancylodiscoides, Protoancylodiscoides, Quadriacanthus, Schilbetrema, Schilbetrematoides, Synodontella and Thaparocleidus. Clariotrema Long, 1981 and Neobychowskyella Ma, Wang & Li, 1983 are considered synonyms of Bychowskyella Akhmerov, 1952, Anacornuatus Dubey, Gupta & Agarwal, 1992 is considered a synonym of Quadriacanthus Paperna, 1961, Mizellebychowskia Gupta & Sachdeva, 1990 is considered a synonym of Neocalceostoma Tripathi, 1959 and Hargitrema Tripathi, 1959 is treated as a synonym of Hamatopeduncularia Yamaguti, 1953. It is proposed that the Ancylodiscoidinae be raised to family status within the order Dactylogyridea to accommodate these 17 'ancyrocephalid' genera from siluriforms, together with Malayanodiscoides and Notopterodiscoides from notopterids. A key and the diagnostic characteristics of the 19 recognised dactylogyridean genera from catfishes plus two from notopterids, together with a list of species and synonyms, are included. New combinations made in this work are Thaparocleidus avicularia (Chen, 1987) n. comb., T. calyciflorus (Chen, 1987) n. comb., T. choanovagina (Luo & Lang, 1981) n. comb., T. dissimilis (Chen, 1988) n. comb., T. leiocassis (Reichenbach-Klinke, 1959) n. comb., T. meticulosa (Chen, 1987) n. comb., T. parasoti (Zhao & Ma, 1999) n. comb., T. persculpus (Chen, 1987) n. comb., T. valga (Chen, 1987) n. comb. and T. wulingensis (Yao & Wang, 1997) n. comb. [all from Silurodiscoides] and Bychowskyella glyptothoraci (Ma, Wang & Li, 1983) n. comb. [from Neobychowskyella].

Metadena phoceae n. sp. (Digenea: Cryptogonimidae), a rectal parasite of the shore rockling Gaidropsarus mediterraneus (Teleostei: Lotidae) in the western Mediterranean.

Metadena phoceae n. sp. is described and figured in detail from the rectum of the shore rockling Gaidropsarus mediterraneus (Lotidae) in the Gulf of Marseilles and off Corsica. It is compared with other cryptogonimids occurring in Mediterranean and Black Sea fishes and is distinguished from its closest relatives, M. depressa (Stossich, 1883) and M. pauli (Vlassenko, 1931), which occur in sparid and sciaenid fishes, respectively, by both morphometrical and biological features. The host-specificity of cryptogonimids occurring in Mediterranean and Black Sea fishes is commented upon.

Redescriptions of two echinostomes from freshwater fishes, with comments on Singhia Yamaguti, 1958 and Caballerotrema Prudhoe, 1960 (Digenea: Echinostomatidae).

The type-species of Singhia and Caballerotrema (Digenea: Echinostomatidae), parasites of freshwater fishes which have previously been assigned to the subfamily Singhiinae Yamaguti, 1958, are redescribed. One of the specimens of the type-series of Caballerotrema brasiliense Prudhoe, 1960 was identified as C. aruanense Thatcher, 1980. C. arapaimense Thatcher, 1980 is considered a species inquirenda. Himasthla piscicola Stunkard, 1960 is transferred to Caballerotrema as C. piscicola n. comb. The generic diagnoses of Singhia and Caballerotrema are redefined and Singhia is re-allocated to the Echinostomatinae Looss, 1899, the Singhiinae becoming a synonym of the latter.

Three little known acanthostomine digeneans from Uranoscopus scaber L. in the western Mediterranean.

Three species of acanthostomine digeneans, Anisocoelium capitellatum from the gall-bladder and Anisocladium fallax and A. gracilis from the intestine, are redescribed from the teleost Uranoscopus scaber in the western Mediterranean. The latter two species, which have in the past been confused and synonymised, are distinguished clearly on the basis of morphology, size and site.

Pseudoterranova decipiens species A and B (Nematoda, Ascaridoidea): nomenclatural designation, morphological diagnostic characters and genetic markers.

Five genetically distinct and reproductively isolated species have been detected previously within the morphospecies Pseudoterranova decipiens from the Arctic-Boreal, Boreal and Antarctic. Morphological analysis was carried out on male specimens identified by genetic (allozyme) markers, allowing the detection of significant differences at a number of characters between two members of the P. decipiens complex, namely P. decipiens A and B. On the basis of such differences, the nomenclatural designation for the two species is discussed. The names Pseudoterranova krabbei n. sp. and P. decipiens (sensu stricto) are proposed for species A and B, respectively. Morphological and genetic differentiation between the two species is shown using multivariate analysis. Allozyme diagnostic keys for routine identification of the four members of the P. decipiens complex, namely P. decipiens (s.s.), P. krabbei, P. bulbosa and P. azarasi, irrespective of sex and life-history stage, are provided.

Re-validation of Echinostoma miyagawai Ishii, 1932 (Digenea: Echinostomatidae) on the basis of the experimental completion of its life-cycle.

The life-cycle of Echinostoma miyagawai, a Eurasian species closely related to E. revolutum, was completed in the laboratory, and the morphology of the larval stages and the adults obtained experimentally was studied. Planorbis planorbis and Anisus vortex were the first intermediate hosts in the brackish Lake Durankulak on the Bulgarian Black Sea coast. Characteristic features of the cercaria include: a prominent collar with 37 spines; a tail as long as the body and with seven conspicuous fin-folds, the two ventral fin-folds being very close to each other; and a specific number and distribution of both the para-oesophageal gland-cell outlets and sensilla. The adult is characterised by: a very elongate body with a constriction at the posterior border of the ventral sucker; a large head collar with relatively small spines; a spherical ventral sucker which is only about half the maximum body width; a long cirrus-sac reaching posteriorly dorsal to the middle of the ventral sucker; indented subglobular testes; and a vitellarium forming two lateral fields of follicles which are almost confluent in the post-testicular space. The species described in this study resembles E. miyagawai, as described by Kosupko, in the morphology of larval stages and both the site and the general morphology of the adults. It differs from both E. revolutum, as described by both Kanev and Nasincová, and E. echinatum (also referred to as E. lindoense and E. barbosai by Kanev). The re-examination of Kanev's voucher specimens from his experimental studies used in his delimitation of E. revolutum and E. echinatum showed that the specimens identified by him as E. revolutum represent two distinct forms which consistently differ both from each other and from the redescription of E. revolutum which was based upon them. It also revealed that a number of specimens were wrongly identified and erroneously treated as E. echinatum by Kanev and co-workers; these include members of different genera (Hypoderaeum and Echinoparyphium) and an Echinostoma species of the group possessing 47 collar spines. The relative merits of the features used by Kanev and co-workers in discriminating the closely related Echinostoma spp. are discussed in detail with respect to the experimental evidence provided by these authors.

Pseudorhabdosynochus spp. (Monogenea: Diplectanidae) from the gills of Epinephelus spp. in Brazilian waters.

Pseudorhabdosynochus sulamericanus n. sp. from the gills of Epinephelus niveatus has a reniform proximal region of the cirrus-bulb which is divided in four chambers and contains a large, round reservoir of the male accessory glands, a partly sclerotised vagina which is enclosed in a muscular funnel cap and squamodiscs with 15-16 open concentric rows of elements. P. beverleyburtonae (Oliver, 1984) is redescribed from E. marginatus with additional morphological data. These are the first reports of Pseudorhabdosynochus spp. in South American Atlantic waters, where the potentiality for the mariculture of Epinephelus spp. is currently being evaluated. Pseudorhabdosynochus hargisi (Oliver & Paperna, 1984) n. comb. is proposed for Diplectanum hargisi.

A quantitative approach to the evaluation of the morphological variability of two echinostomes, Echinostoma miyagawai Ishii, 1932 and E. revolutum (Frölich, 1802), from Europe.

A comparative morphometric analysis was conducted on two European species of Echinostoma in order to examine the degree of the variability in the metrical characteristics of the adults and to assess their value in discriminating species. Adult E. miyagawai and E. revolutum, obtained experimentally, were compared using univariate and multivariate statistical analyses of 35 and 25 metrical characters, respectively. All subsets of worms of different ages represented homogeneous samples with respect to their morphometric characteristics; however, univariate analyses revealed significant differences in 22 and 23 variables between the corresponding age subsets of the two species, and it was found that the different allometric growth patterns contribute to this. The variables, body width at the posterior border of the ventral sucker, pharynx length and width, ovary length, testes length and width and length of the pre-ovarian region, exhibited isometric or positive allometric growth in E. miyagawai and negative allometry in E. revolutum. A cluster analysis based on 61 specimens and 25 variables separated E. revolutum and E. miyagawai unambiguously, producing an exact ordering of the specimens with respect to their identity and age. A forward stepwise discriminant analysis identified five variables (body width at the posterior border of ventral sucker, head collar width, length of oesophagus, width of ventral sucker and length of the pre-ovarian region) which yielded a 100% accurate classification of the two species. We suggest, therefore, that the morphometric characteristics of the adult worms should be used in studies attempting the identification of species or isolates of Echinostoma spp. More comparative data need to be gathered in order that the species boundaries within the 'revolutum' group be defined more accurately.

Cercarial emergence of Ichthyocotylurus erraticus (Rudolphi, 1809), I. variegatus (Creplin, 1825) and Apatemon gracilis (Rudolphi, 1819) (Digenea: Strigeidae): contrasting responses to light:dark cycling.

Many digenean cercariae have been shown to emerge from their molluscan hosts with distinct shedding patterns that have enabled the discrimination of morphologically similar species, or even strains. In this study the cercarial emission patterns of three strigeid species, Ichthyocotylurus erraticus, I. variegatus and Apatemon gracilis, from experimentally infected natural hosts were found to exhibit rhythms that correlated with the light:dark cycle. Both Ichthyocotylurus spp. exhibited a diurnal pattern of release in which cercariae emerged during the light period. Each demonstrated a latent period before the liberation of large numbers of cercariae and yielded similar numbers of cercariae daily. These rhythms offered no means for the discrimination of these two morphologically similar species. A. gracilis cercariae demonstrated a very different circadian rhythm in which the majority emerged at the onset of darkness with no latent period, whereas the cercarial numbers released daily were far greater. Differences could be related to piscine host behaviour.

Urotrematidae Poche, 1926 (Platyhelminthes: Digenea) in Chinese freshwater fishes.

The genera Sinineobucephalopsis and Sinogastromyzontrema are diagnosed and placed in the family Urotrematidae. Sinineobucephalopsis macrocirrus is redescribed from Glyptothorax fokiensis in Guangdong and Hunan provinces. Urotrema glyptothoraci is considered synonymous with Sinineobucephalopsis macrocirrus. The new combinations Sinineobucephalopsis postlecitha, Sinineobucephalopsis proeilecitha and Sinineobucephalopsis sinipercae are formed for species originally placed in Urotrema and Urotrematulum. Sinogastromyzontrema guangxiensis is redescribed from Sinogastromyzon wui in Guangxi Province. Urotrema parallelorchis is considered a synonym of Sinogastromyzontrema guangxiensis. A key to the species or species-complexes in the family Urotrematidae is given.

Chalcinotrema thatcheri n. sp. (Digenea: Haploporidae) from Brazilian freshwater fishes, a redescription of C. ruedasueltensis Thatcher, 1978 and comments on the validity of the genus.

Chalcinotrema thatcheri n. sp. from the intestine of Schizodon knerii is described. It is most closely related in general morphology to C. salobrensis Freitas, 1947 and C. ruedasueltensis Thatcher, 1978, from which it differs, respectively, in: testis-size, the distribution of the vitelline follicles and the distance between the gonads; and body-size, egg-size, the extent of the caeca and the shape of the seminal vesicle. The holotype of C. ruedasueltensis is redescribed with original figures. Chalcinotrema Freitas, 1947, is considered a good genus distinct from Carassotrema Park, 1938, on the basis of the presence of uterine loops between the ovary and testis and its totally different geographical distribution. Carassotrema tilapiae Nasir & Gomez, 1976 is transferred to the former genus as Chalcinotrema tilapiae (Nasir & Gomez, 1976) n. comb.

Echinorhynchus brayi n. sp. (Acanthocephala: Echinorhynchidae) from Pachycara crassiceps (Roule) (Zoarcidae), a deep-sea fish.

Echinorhynchus brayi n. sp. (Palaeacanthocephala: Echinorhynchidae) is described from Pachycara crassiceps (Roule) (Zoarcidae) from the Porcupine Seabight, Northeast Atlantic. The new species closely resembles E. canyonensis Huffman & Kliever, 1977, a parasite of a Pacific zoarcid, but has longer lemnisci, larger eggs and larger testes. E. brayi n. sp. can be readily differentiated from the ten other Echinorhynchus spp. recorded from deep-sea fishes (E. abyssicola, E. gadi, E. longiproboscis, E. malacocephali, E. melanoglaeae, E. muraenolepisi, E. petrotschenkoi, E. sebastolobi, E. trachyrinci and E. truttae), because it has fewer hooks per longitudinal row.

A redescription of Pseudobacciger harengulae (Yamaguti, 1938) (Digenea: Faustulidae) from Sprattus sprattus phalericus (Risso) and Engraulis encrasicholus ponticus Alexandrov off the Bulgarian Black Sea coast, with a review of the genus Pseudobacciger Nahhas & Cable, 1964.

Pseudobacciger harengulae is recorded from the caeca of Sprattus sprattus phalericus and Engraulis encrasicholus ponticus (new host records) in the Black Sea (new geographical record) off the Bulgarian coast. This is the first record of a species of the genus Pseudobacciger in the Black Sea. An illustrated redescription on the basis of the Bulgarian material is presented. The specimens from the two hosts differ from one another in the shape of the arms of the excretory vesicle. A review of Pseudobacciger is presented; for each of the three species of the genus, data on their hosts, distribution, morphology and life-cycle (if known) are summarised.