Variability and phenotypic structure of Proteocephalus torulosus (Cestoda: Proteocephalidea)--a parasite of cyprinid fishes

Variability of 5 qualitative features of Proteocephalus torulosus from 6 fish species belonging to 5 genera of the subfamily Leuciscinae was studied. Itraspecific diversity is represented by 3 scolex forms, 2 types of proglottid structure, 3 proglottid shapes, and 2 variations of the female reproductive system characteristics. The parasite living in different host species form separate ecological forms differing in the number of phenotypes and their ratio. However, the same scolex and strobila phenotypes dominate in the range of the parasite. Two P. torulosus groupings have been recognised the European and Asian ones, which differ in the representation of the proglottid structure types.     

Phylogenetic analysis of European species of Proteocephalus (Cestoda: Proteocephalidea): compatibility of molecular and morphological data, and parasite-host coevolution.

The phylogeny of European species of the tapeworm genus Proteocephalus was studied, based on partial 18S rDNA and morphological data. The group was found to be monophyletic. The analysis showed a low informative value of available morphological characters in comparison with molecular data. The morphological matrix resulted in a poorly resolved tree which is, however, compatible with the topology (Proteocephalus osculatus (Proteocephalus torulosus (Proteocephalus macrocephalus, Proteocephalus filicollis) (Proteocephalus tetrastomus, Proteocephalus percae, Proteocephalus longicollis))) based on the 18S rDNA data. A comparison performed by the program TreeMap showed a lack of significant congruency between parasite and host phylogenies. Therefore, the distribution of species in their hosts appears to be independent of the phylogeny and it is likely to be a result of host-switching, rather than co-speciation events.     

Seasonal dynamics of Proteocephalus sagittus in the stone loach Barbatula barbatula from the Hana River, Czech Republic

The seasonal cycle of the cestode Proteocephalus sagittus (Cestoda: Proteocephalidae) was studied for the first time in the stone loach Barbatula barbatula from the Hana River, Czech Republic. A total of 180 loaches were examined monthly from January to December 2001. The parasite occurred in loaches throughout the year but infection parameters differed significantly among seasons, with the highest values of prevalence and abundance from the late winter to the early summer. Parasite recruitment took place in the winter and early spring and the worms sexually matured in the late spring and early summer. In contrast to P. torulosus, the gravid worms of which laid eggs only at the end of the spring/beginning of the summer, gravid worms of P. sagittus were also found, although in low numbers, in the autumn and early winter. The rate of infection of loach with P. sagittus was neither dependent on the sex nor on the size of its fish host.     

Species of Proteocephalus Weinland, 1858 (Cestoda: Proteocephalidae), parasites of coregonid and salmonid fishes from North America: taxonomic reappraisal

A taxonomic study of species of Proteocephalus Weinland, 1858 (Cestoda: Proteocephalidae), from coregonid and salmonid fishes in North America, based largely on their type specimens, was carried out. None of the taxa studied possesses obvious autapomorphies that would justify its validity. On the contrary, all species share the following diagnostic features typical of Proteocephalus longicollis (Zeder, 1800) (syn. Proteocephalus exiguus La Rue, 1911), a polymorphic and widely distributed parasite in the Holarctic: (1) a spherical or club-shaped scolex, with sublaterally situated suckers and a vestigial but distinct and relatively large apical sucker; (2) the vagina possessing a well-developed vaginal sphincter and crossing mostly ventrally the proximal part of the cirrus sac; and (3) a long, thick-walled cirrus sac, overlapping vitelline follicles medially by more than its proximal half. Accordingly, Proteocephalus pusillus Ward, 1910; P. laruei Faust, 1920; "P. obtundus" La Rue, 1920 (nomen nudum); P. arcticus Cooper, 1921; P. wickliffi Hunter and Bangham, 1933; P. parallacticus MacLulich, 1943; P. californicus Haderlie, 1950; P. salmonidicola Alexander, 1951; and P. primaverus Neiland, 1952 are considered junior synonyms of P. longicollis.     

Comparative morphological and gene systematics analysis of the Proteocephalus thymalli (Cestoda: Proteocephalidae), a parasite of graylings in lakes Hovsogol and Baikal

Results of comparative morphological and gene analyses of the cestode Proteocephalus thymalli, an intestine parasite of the lakes Hovsogol (Thymallus arcticus nigrescens) and Lake Baikal graylings (Thymallus arcticus baicalensis) are presented for the first time. The results indicate that representatives of the P. thymalli from these fishes are really two different species.     

Phellinus coronadensis: a new species from southern Arizona, USA

Phellinus coronadensis is characterized and described morphologically as a new species from southern Arizona, USA. This fungus was previously reported as P. torulosus based on morphological similarities of the basidiomes and type of wood decay. However, P. coronadensis is restricted to two mountain ranges in southern Arizona and found almost exclusively on living southwestern white pine (Pinus strobiformis). Phellinus torulosus is found primarily in Europe and parts of Asia and is primarily associated with hardwood hosts. Based on sequence analysis of small subunit mitochondrial ribosomal DNA (mt-SSU), we determined that P. coronadensis is in a different lineage from P. torulosus and apparently more closely related to the P. pini complex. The taxon associated with southwestern white pine, being distinct and not yet having been validly named, is proposed as a new species here.     

Infectivity of Ornithodiplostomum ptychocheilus and Posthodiplostomum minimum (Trematoda: Diplostomidae) cercariae following exposure to cadmium

The effect of cadmium (Cd) exposure on infectivity of cercariae of 2 trematode species (Ornithodiplostomum ptychocheilus and Posthodiplostomum minimum) to their second intermediate fish host was evaluated. Individual fathead minnows (Pimephales promelas) were exposed to cercariae that had been treated with Cd solutions at concentrations of 0, 2, 20, or 200 microg/L for 2 hr. Two weeks later, the numbers of encysted metacercariae in the brain (O. ptychocheilus) and body cavity (P. minimum) of the fish were evaluated. ANOVA analyses indicated a strong negative effect of Cd concentration on cercaria infectivity. The species x Cd concentration interaction was not significant, indicating that the magnitude of Cd-induced reduction in infectivity was similar between O. ptychocheilus and P. minimum. The results show that short-term exposure to Cd, even at low concentrations, interferes with transmission processes that affect the recognition and penetration of cercariae, the migration and survival of metacercariae within the second intermediate host, or both.     

A key to the species of cestodes (tapeworms) parasitic in British and Irish freshwater fishes

Methods for the collection, fixation, preservation and examination of the cestodes parasitic in British freshwater fishes are described. An illustrated key to the species is provided, with the exception of the members of the genus Proteocephalus , for which the information is tabulated.     

The helminth parasites of the bullhead Cottus gobio (L.) and the stone loach Noemacheilus barbatulus (L.) from the River Avon, Hampshire

Specimens of Cottus gobio and Noemacheilus barbatulus from the River Avon, Hampshire, were examined between July 1970 and July 1971. Both fish were found to act as hosts for Nicolla gallica (adult), Pomphorhynchus laevis (adult) and Triaenophorus nodulosus (plerocercoid). The incidence and intensity of infection were lower in N. barbatulus than in C. gobio , and this was related to differences in host diet. A small number of N. barbatulus were also found to harbour Proteocephalus torulosus . Seasonal cycles of incidence and intensity were evident in N. gallica , with a peak occurring in the winter. P. laevis and T. nodulosus showed no such cycles. Increasing host size was associated with a higher infection of both N. gallica and P. laevis , but not T. nodulosus . The diet of the host was considered to be important in influencing this increase. Differences in host reproductive condition affected the population levels of N. gallica only, where an increased parasite burden was evident in gravid female fish. This was considered to influence the observed increase in incidence and intensity of this parasite during the winter. None of the parasites showed any seasonal maturation cycles. There was no evidence of any interactions between the parasite populations. Host diet and the availability of infective larvae were considered to be the most important features influencing the number of parasites which became established, and the rate at which this occurred. Temperature probably acted indirectly on the establishment of N. gallica , by controlling the availability of the larval stages, and also, by influencing the host reproductive condition, on the length of time for which the parasite was retained by the host.     

Enteric helminths of perch (Perca fluviatilis L.) and yellow perch (Perca flavescens Mitchill): stochastic or predictable assemblages?

Component communities of perch (Perca fluviatilis L) in Eurasia and the North American yellow perch (Perca flavescens Mitchill) were examined to determine the nature of their parasite communities. The scale of this investigation is continental and includes data collected across the distribution of each host species. Data were compiled from the literature and from 5 sample sites in North America. Four parasite species were found to occur frequently in the helminth component communities of P. flavescens. The cestodes Bothriocephalus cuspidatus and Proteocephalus pearsei, the digenean Crepidostomum cooperi, and the nematode Dichelyne cotylophora comprised a suite of species of which some or all occurred in most samples. Similarly, a group of 4 predictable parasite species was identified for P. fluviatilis in Eurasia, the digenean Bunodera luciopercae, the nematode Camallanus lacustris, the cestode Proteocephalus percae, and the acanthocephalan Acanthocephalus lucii. Specificity was not a requirement for predictability. Despite geographical isolation for millions of years, and different fish species interactions within and between continents, the predictability of these parasite assemblages indicates they are shaped by a biology, especially feeding patterns, common to both perch species. This is evidence that parasite assemblages comprised of nonhost-specific parasites in freshwater fishes are not merely stochastic assemblages but have key components that are predictable at this broad continental scale.     

Diapause of copepods as an element for stabilizing the parasite system of some fish helminths

The analysis of infection dynamics of copepods, which are the intermediate hosts of three helminth species of freshwater fishes (Triaenophorus crassus, Proteocephalus exiguus and P. percae), has shown that diapause in the life cycle of the copepods is favourable for preserving the infection in the waterbody until physiological prerequisites for successful infection of the final host are acquired. The peculiarity of a copepod's life cycle may determine the strategy of a parasite in its preimaginal phase as a waiting stage, and the duration of the residence of helminth larvae in copepods that have an obligatory diapause, is one of the elements the provide stability in parasite systems.     

A Redescription of the Cestodes Proteocephalus filicollis (Rudolphi) from Gasterosteus aculeatus L., and P. ambiguus (Dujardin) from Pungitius pungitius (L.)

Based on material from two stickleback species living in the same biotope, Proteocephalus filicollis (Rudolphi, 1802) and Proteocephalus ambiguus (Dujardin, 1845) are regarded as two distinct species. The most useful character for their discrimination is the presence and absence of an apical scolex of P. filicollis and P. ambiguus , respectively. Furthermore, transverse sections revealed that the longitudinal muscles consist of approximately 5 layers of muscle fibres in P. filicollis , as opposed to a single layer in P. ambiguus . The two species differ also in total length, width, number of segments and in the state (progress) of maturation of the proglottids along the strobila.     

The evolution of the Proteocephalidea (Platyhelminthes,Eucestoda) based on an enlarged molecular phylogeny,with comments on their uterine development

We present a molecular phylogeny of the Proteocephalidea based on 28S rDNA sequence data that is a follow-up to the paper by Zehnder & Mariaux (1999). Twenty-three new sequences, including three outgroups are added in our new data-set. The Gangesiinae Mola, 1929 and the Acanthotaeniinae Freze, 1963 appear to be the most primitive clades. They are followed by a robust clade comprising the Palaearctic Proteocephalinae Mola, 1929 from freshwater fishes. The structure of the more derived clades, comprising most Neotropical and Nearctic species, is less resolved. At the nomenclatural level, we erect a new genus, Glanitaenia n. g. for G. osculata (Goeze, 1782) n. comb., previously Proteocephalus osculatus, and define an aggregate for the Palaearctic Proteocephalus Weinland, 1858. After a re-examination of all of the studied taxa, we identify two types of uterine development and show the importance of this character for the systematics of the order. Our phylogeny does not support the classical view of a Neotropical origin for the Proteocephalidea but rather favours an Old World origin of the group either in saurians or Palaearctic Siluriformes.     

Comparative life histories of two diplostomid trematodes, Ornithodiplostomum ptychocheilus and Posthodiplostomum minimum

A lack of comparative studies limits our understanding of interspecific variation in parasite life histories, especially for species that incorporate asexual and resting stages into their life cycles. Ornithodiplostomum ptychocheilus and Posthodiplostomum minimum are sympatric diplostomid trematodes that share the same first (pond snail, Physa gyrina) and second (fathead minnow, Pimephales promelas) intermediate hosts within lakes in Alberta, Canada. Interspecific differences in the body sizes of adults, cercariae, and metacercariae were evaluated from parasites recovered from laboratory-raised chickens, snails, and minnows, respectively. Differences in adult worm fecundity and cercariae production were estimated from chickens and snails exposed to known numbers of parasite larvae. Posthodiplostomum minimum adults, metacercariae, and eggs were 97, 86, and 5% larger than those of 0. ptychocheilus, respectively. The average numbers of eggs produced per day by P. minimum was 3 times higher than for O. ptychocheilus, and P. minimum produced approximately twice as many cercariae per day. Although the larger of the 2 species had higher egg and cercariae production, conclusions regarding covariation between adult worm size and the reproductive rates of other life cycle stages require further studies on related species.     

Interaction of bass tapeworm, Proteocephalus ambloplitis, and Neoechinorhynchus sp. (Acanthocephala) in largemouth bass, Micropterus salmoides

The number of plerocercoids of the bass tapeworm, Proteocephalus ambloplitis, in wild largemouth bass was negatively correlated (r = -0.94) with the number of Neoechinorhynchus sp. Competitive inhibition between the 2 parasites appeared to exist. Similarly, the numbers of Neoechinorhynchus sp. in wild bass decreased when adult bass tapeworms were present in the intestine. Proteocephalus ambloplitis plerocercoids used to challenge bass vaccinated with either P. ambloplitis adult or Neoechinorhynchus sp. antigens were smaller (P less than 0.05) when recovered than those used to challenge control bass. Based on preliminary results, both antigens might have enabled the bass to limit growth and/or development of the invading bass tapeworm, plerocercoids. Cross-protective immunity may be the reason for this occurrence, in which case, it could offer an explanation for competitive inhibition existing between P. ambloplitis and Neoechinorhynchus sp.     

High intensity and prevalence of two species of trematode metacercariae in the fathead minnow (Pimephales promelas) with no compromise of minnow anti-predator competence

Opportunity for parasites to manipulate host behavioral phenotype may be influenced by several factors, including the host ecology and the presence of cohabiting parasites in the same host. Metacercariae of Ornithodiplostomum ptychocheilus and "black spot" Crassiphiala bulboglossa have similar life cycles. Each parasite uses a littoral snail as a first intermediate host, fathead minnows as a second intermediate host, and a piscivorous bird as a final host. Metacercariae of black spot encyst in the dermal and epidermal tissues, while metacercariae of O. ptychocheilus encyst on the brain over a region that coordinates optomotor responses. Because of site differences within the host, we predicted that O. ptychocheilus metacercariae might manipulate the behavioral phenotype of minnows to facilitate transmission to the final host, but metacercariae of black spot would not. In our study population, prevalence was 100% for O. ptychocheilus , with an overall median intensity of 105 metacercariae per minnow. Prevalence of black spot was 60%, with a median abundance and intensity of 12 and 20 metacercariae per minnow for the overall sample and for infected fish, respectively. Minnows accumulated both parasites over time, producing significant correlations between intensity and minnow body length and between intensities of the 2 parasites. Minnows infected with black spot had on average twice as many O. ptychocheilus metacercariae as similar-sized minnows without any black spot cercariae. We found no correlation between body condition of minnows and intensity for either parasite. We measured 2 aspects of anti-predator competence to test for effects linked to parasite intensity. We found no correlation between intensity of either species of parasite and latency to behavioral response to attack from a mechanical model heron, nor was there any effect of parasite intensity on a measure of shoaling affinity. The absence of any detectable effect of metacercariae on anti-predator competence in minnows may reflect selection against parasite pathology from predation by non-hosts of the parasites and overwinter mortality due to low dissolved oxygen.     

Estimation of the endohelminth parasite species richness in freshwater fishes from La Mintzita Reservoir, Michoacán, Mexico

In total, 9 endohelminth species were found to parasitize 7 fish species (2 cyprinids, 4 goodeids, and 1 poeciilid) from La Mintzita Reservoir, Michoacán, in central Mexico; 5 were larvae, including 3 allogenic species (Clinostomum complanatum, Tylodelphys sp., Posthodiplostomum minimum) and 2 autogenic species (Serpinema trispinosum, Spiroxys sp.). Four were enteric autogenic adults, i.e., Margotrema bravoae, Bothriocephalus acheilognathi, Proteocephalus longicollis, and Rhabdochona lichtenfelsi. The metacercariae of P. minimum reached the highest levels of prevalence and mean abundance among host species. Our results confirm the depauperate nature of the helminth communities of freshwater fishes from central Mexico. On the basis of this data set, we estimated the total endohelminth species richness for each component community by using 7 nonparametric estimators whose performance was evaluated with the unscaled measures of bias, precision, and accuracy. We found that Chaol and Bootstrap are the most precise and least biased methods for the 7 component communities: however, species richness was consistently underestimated. The underestimation was an unavoidable consequence of the patchy distribution of helminth species among different component communities, particularly at the small sample size used in our study.