Host specificity of adult versus larval cestodes of the elasmobranch tapeworm order Trypanorhyncha

Host specificity between the adult and final larval stages (plerocercus, plerocercoid, or merocercoid) of a diversity of trypanorhynch species was compared using the host specificity index (HS s). Index values were generated for a total of 63 species representing all five trypanorhynch superfamilies and 11 families. Host specificity of both adults and final larvae was found to be widely variable among species, ranging from very high (oioxenous) to very low (euryxenous) for both stages. However, in general, host specificity was highest for the adult stage in the definitive host (mean HS s=3.86) and lowest for the final larval stage in the second intermediate host (mean HS s=6.29). This difference was found to be significant using the Wilcoxon signed-rank test. Limited data available for procercoids in the first intermediate host suggest that this stage exhibits a degree of specificity intermediate between that of the former two stages (mean HS s=4.23). No taxonomic trend was seen. Species with a plerocercoid final larval stage (mean HS s=8.62) were significantly less host-specific than those with plerocerci or merocercoids (mean HS s=5.56). This result may reflect the use of paratenic hosts by species possessing the relatively more resilient plerocercoid as their final larval stage. These results provide an example of how HS s can be used to compare levels of host specificity, in this instance, among stages of polyxenous life cycles. They also emphasise the importance of articulating the life cycle stage under consideration when general statements are made about host specificity.     

Cestodes of cestodes of Peruvian freshwater stingrays

Adult tetraphyllidean tapeworms (Platyhelminthes: Eucestoda) from the spiral intestines of 3 species of potamotrygonid stingrays (Paratrygon aiereba, Potamotrygon castexi, and Portamotrygon motoro) in the Madre de Dios river in Peru were found to host numerous cysts embedded in their parenchymal tissues. Histological sections of the cysts revealed the presence of a scolex bearing 4 suckers and an unarmed apical organ consistent with larval stages of both Cyclophyllidea and Proteocephalidea. To further elucidate their identities, partial 28S ribosomal DNA (rDNA) sequences were characterized from 3 cysts and 4 adult Rhinebothrium spp. 'host' worms and screened against all available cestode 28S rDNA data. Initial BLAST screening and subsequent alignment ruled out the possibility that the cysts were cyclophyllidean, and the cyst and adult sequences were thus aligned together with all available lecanicephalidean, litobothriidean, proteocephalidean, and tetraphyllidean sequences. Sequences from all 3 cysts were identical, and phylogenetic analysis clearly placed them among derived members of the Proteocephalidea, although no exact match was found. Sequences from the adult host worms formed 2 identical pairs and grouped together with other tetraphyllidean species from rays. These results are compared with records of hyperparasites of South American catfish cestodes. This is the first confirmed record of a proteocephalidean cestode parasitizing a tetraphyllidean cestode.     

Seasonal and ontogenetic dynamics in trophic transmission of parasites

Transmission rates from the intermediate (amphipods) to the definitive hosts (fish) were quantified for two helminth species ( Cyathocephalus truncatus , Cestoda, and Cystidicola farionis , Nematoda) both seasonally and through the ontogeny of the final hosts (arctic charr, Salvelinus alpinus , and brown trout, Salmo trutta ). Amphipods ( Gammarus lacustris ) were important prey for both fish species, especially in the autumn. Both parasite species had low infection levels in amphipods compared to high abundance in fish. The seasonal variations in transmission rate of C. truncatus procercoids from amphipods to fish were in accordance with the observed abundance in fish hosts, being highest in the autumn and lowest during late winter and early summer. During summer, however, the estimated monthly transmission rates of C. truncatus were higher than the observed infection levels in the fish, suggesting restricted establishment success and shorter development time and longevity of the cestode in fish at higher temperatures. The accumulated transmission of C. farionis over the ontogeny of arctic charr was similar to the observed age-specific infection levels, reflecting a high establishment success and longevity of this parasite in charr. In contrast, brown trout exhibited an infection level that was much lower than the estimated transmission rates, suggesting a high resistance against C. farionis in these fish. The magnitudes of the estimated transmission rates were sufficient to explain the paradoxical contrast between low infection levels in the intermediate and high in the final hosts.     

Some aspects of Artemia biology affected by cestode parasitism

Artemia individuals from a wild population, bearing cestode cysticercoids (Hymenolepididae) were kept in the laboratory during four months to observe influences of the parasite on the host biology. No differences were found between parasitized and unparasitized shrimps regarding final adult size. The higher survival of parasitized animals suggests that parasitism is not pathogenic. The most dramatic effect was host castration. Parasitized individuals showed higher total lipid levels, probably linked to carotenoid pigments conveying a red color to parasitized shrimps.     

Differences in pyloric caeca morphology between Arctic charr Salvelinus alpinus ecotypes: adaptation to trophic specialization or parasite‐induced phenotypic modifications?

Gut-tract morphology differed between benthivorous and zooplanktivorous Arctic charr Salvelinus alpinus caught in the littoral and pelagic zones of a sub-Arctic lake. The differences were related to trophic niche and infection with the cestodes Cyathocephalus truncatus and Eubothrium salvelini . Measurements that were unrelated to cestode infection, including intestinal length, numbers of pyloric caeca and posterior pyloric caeca morphology, differed between S. alpinus that had consumed different prey types, suggesting an adaptive response to divergent trophic niches. The anterior pyloric caeca of S. alpinus with high numbers of cestodes were significantly wider and shorter than those in fish with lower levels of infection. The differences were strongly associated with infection by C. truncatus , which was site-selective and attached to the anterior caeca with a large scolex. Differences in caecal measurements were probably the result of mechanical damage caused by scolex attachment and host responses to infection with C. truncatus . The differences in anterior caeca were also an indirect indicator of trophic specialization. Benthivorous S. alpinus were more exposed to, and more heavily infected with C. truncatus than were pelagic zooplanktivorous fish, because this cestode had Gammarus lacustris as its intermediate host. Nevertheless, care should be exercized when using gut measurements for making inferences about the trophic ecology of fish because a history of infection with cestodes may have profound effects on caecal morphology.     

Hymenolepis nana: immunity against oncosphere challenge in mice previously given viable or non-viable oncospheres of H. nana, H. diminuta, H. microstoma and Taenia taeniaeformis

When mice, previously given oral inoculation with viable oncospheres of the heterologous cestode species (Hymenolepis diminuta, H. microstoma, Taenia taeniaeformis) and the homologous one (H. nana), were challenged with oncospheres of H. nana 4 days after the primary inoculation, they showed strong and complete resistance to H. nana challenge, respectively. However, the resistance was not evoked in mice given either infective eggs of Toxocara canis or non-viable oncospheres of all cestode species examined. Congenitally athymic nude mice given viable oncospheres did not show any resistance to H. nana either. Eosinophil infiltration around cysticercoids of H. nana in the intestinal villi appeared to be more prominent in mice previously given viable oncospheres of H. diminuta than in mice given non-viable oncospheres or PBS only. Some of the eosinophils in the villus harboring cysticercoid(s) of H. nana invaded the epithelia in the former, whereas all eosinophils remained in the lamina propria in the latter. There was almost no eosinophil infiltration in nude mice. Microscopic observations revealed that oncospheres of H. diminuta, which require beetles as the intermediate host like H. microstoma, could invade the mouse intestinal tissue. Therefore, it is strongly suggested that the strong cross resistance to H. nana in mice, induced by oncospheres of all heterologous cestode species, is thymus-dependent and due to oncospheral invasion into the intestinal tissue of mice.     

Ecological causes of sex-biased parasitism in threespine stickleback

Males and females can differ in levels of parasitism and such differences may be mediated by the costs of sexual selection or by ecological differences between the genders. In threespine stickleback, Gasterosteus aculeatus , males exhibit paternal care and territorial nest defence and the costs of reproduction may be particularly high for males relative to females. We monitored levels of parasitism for 15 years in a population of stickleback infected by four different parasite species. Consistent with general predictions, overall parasite prevalence (total parasitism) was greater in males than in females. However, this excess did not occur for each species of parasite. Males had higher prevalence of a cestode Cyathocephalus truncatus and a trematode Bunodera sp. relative to females, while females had higher prevalence of a cestode Schistocephalus solidus and nematodes. This suggested ecological sources to differences in parasitism rather than reproductive costs and therefore we examined diet of unparasitized stickleback, predicting that differences in dietary niche would influence relative parasitism. This was partially confirmed and showed that female stomach contents had increased frequency of pelagic items, the major habitat for the primary host of S. solidus whereas males exhibited increased frequency of benthic items, the dominant habitat of C. truncatus and Bunodera. Temporal shifts in the extent and direction of differential parasitism among years between the sexes were associated with temporal shifts in dietary differences. Our results, combined with those in the literature, suggest that ecological differences between genders could be a more important component to patterns of parasitic infection in natural populations than currently appreciated.     

Spontaneous infection of Hymenolepis nana in hamsters (author's transl)]

Hymenolepis nana (von Siebold, 1852), the dwarf tapeworm causing hymenolepiasis, has been reported to be the common intestinal cestode of rodents and man throughout the world. The authors found spontaneously occurred hymenolepiasis in conventional laboratory hamsters with mass and heavy infections. Some individuals were infected with as many as 188 to 290 worms and in addition, numerous cysticercoids were found in the intestinal villi from the same hamsters. According to the early investigations it is said that there are two ways of infection to rodents. In this study the authors considered a natural autoinfection is to be the case because cysticercoids and immature worms were abundant in the intestines of hamsters. The infection rate of the hamsters was 15% as 6 hamsters were found infected out of 40. All the cases of the heavily infected hamsters showed showed loss of weight and some of them died. Particular damage of the intestinal mucosa was not seen by post-mortem examinations, however, chronic inflammation and destruction of intestinal mucosa caused by the worms observed microscopically (the given pictures). Heavy infection of Hymenolepis nana occurred spontaneously in hamsters has rarely been found in Japan. The case reported here seems to be very peculiar one.     

Niche partitioning in the cestode communities of two elasmobranchs

Several randomization methods have been used to investigate the influence of competitive interactions in shaping parasite community structure. Marine fish parasite communities have often been regarded as unstructured assemblages with little or no resource limitation and, therefore, not prone to competitive influences. In this study, null models were used to assess the niche overlap of cestode communities of two distinct but closely related elasmobranch hosts: the round stingray, Urobatis halleri , Cooper (1863) and the skate, Leucoraja naevus , Muller and Henle (1841). Cestode species distributions were analyzed for randomness with respect to two niche axes: host size class and location within host spiral intestine. Niche overlap of cestode species was calculated for each niche type using MacArthur and Levins' and Pianka's indices, and compared to overlap values obtained from randomly generated communities. Cestodes of both host species were distributed heterogeneously among valves and host size-classes. The majority of parasite species (including all common ones) within U. halleri and all species within L. naevus had significantly nonrandom distributions with respect to at least one niche category. Cestode species pairs tended to overlap significantly less than expected in spiral intestine valves and significantly more than expected among host size classes. We conclude that cestode communities are structured deterministically in a way consistent with expectations based on competition.     

Parasites of freshwater resident and anadromous Arctic charr (Salvelinus alpinus) in Greenland

Twenty-one metazoan parasite species were found in Arctic charr at three coastal sites in west Greenland, including Neascus sp. and the nematode Pseudoterranova decipiens , new records for Arctic charr. Twelve species are new records for Arctic charr in Greenland. The quantitative composition of the parasite communities varied, but the freshwater species ( Diphyllobothrium ditremwn, Eubothriutn salvelini, Proteocephalus longicollis, Crepidostotnutn farianis ) were usually numerically dominant, even in anadromous fish. The absence from Greenland of Mysidacea, Amphipoda, Ephemeroptera and Odonata excludes parasites requiring such organisms as intermediate hosts, including the cestode Cyathocephalus truncatus , nematodes of the genus Cystidicola , and freshwater acanthocephalans.     

Potential hosts of Prostephanus truncatus (Coleoptera: Bostrichidae) among native and agroforestry trees in Kenya

This study was carried out to establish the extent to which an invasive stored products pest, Prostephanus truncatus Horn exploits wood as a feeding and breeding resource in Kenya. Twenty seven out of 84 native and agroforestry trees and shrubs supported the breeding of P. truncatus under laboratory conditions. Adult survival (<1% to 18%) for 8 weeks was recorded on 51 tree species that did not support breeding. The breeding success of this beetle varied widely between host tree species and showed no obvious trends based on tree systematics or wood hardness. Studies of P. truncatus survival on two host tree species found that the greatest increase in the P. truncatus population occurred on stem sapwood, whilst the smallest population increase was on twigs. Breeding also varied greatly from season to season; studies on ten selected host species showed that reproduction was often greatest in wood samples collected and tested during the dry season in September. The implications of these findings for the management of P. truncatus infestations in farm stored maize are discussed.     

Host specificity of two species of Gnathia (Isopoda) determined by DNA sequencing blood meals

Host specificity data for gnathiid isopods are scarce because the parasitic stages are difficult to identify and host-parasite contact is often brief. We examined two common nocturnal species, Gnathia falcipenis and Gnathia sp. C, collected in light traps from two locations at Lizard Island on the Great Barrier Reef, Australia. Engorged third stage gnathiids were photographed and permitted to moult into adults to allow identification. We compared approximately 580 bp sequences of 16S mtDNA from blood meals with host sequences available on GenBank using BLASTn. Where homology was <98%, familial identity was investigated with neighbour-joining trees. All blood meal sequences (n=60) and homologous fish sequences (n=87) from GenBank were used in a Bayesian analysis, which identified all but three sequences to family. The host frequency distributions used by each species were significantly different; only four host families were shared. No gnathiids fed on elasmobranchs, blennies or apogonids, and most fed on host families whose representatives are typically large. Gnathia sp. C showed a distinct predilection for nemipterids. Gnathia falcipenis often parasitised sand-dwelling families, and unlike sympatric diurnal gnathiid species, it also frequently parasitised pomacentrids. We conclude that G. falcipenis and Gnathia sp. C operate as generalist micropredators with preferences.     

Linking larvae and adults of Apharyngostrigea cornu, Hysteromorpha triloba, and Alaria mustelae (Diplostomoidea: Digenea) using molecular data

Because the taxonomy of trematodes is based on adults, the larval stages of most digeneans cannot be identified to species based on morphology alone. Molecular data provide a means of linking larval stages to known adults. We obtained sequences from the barcode region of cytochrome oxidase I (CO1) from adult and larval parasites of fish, frogs, birds, and mammals across North America. Sequences from adult Apharyngostrigea cornu, Hysteromorpha triloba, and Alaria mustelae (Diplostomoidea: Digenea) from definitive hosts matched those of meta- and mesocercariae from fish and frogs. These data provided new information on the distributions of all 3 parasite species. Metacercariae of A. cornu, which have not been previously reported in North American hosts, were found in Notemigonus crysoleucas, Pimephales notatus, and Catostomus commersonii in the St. Lawrence River. Metacercariae of H. triloba are reported in Canadian waters and in N. hudsonius for the first time. Alaria mustelae is reported for the first time in frogs from Quebec, Canada, and an additional species of Alaria was detected in California. Sequences of internal transcribed spacer rDNA from a subset of specimens support the same species boundaries indicated by CO1 divergence. There was little divergence in CO1 sequences from an unidentified diplostomid species sampled at a large spatial scale.     

Determinants and consequences of interspecific body size variation in tetraphyllidean tapeworms

Tetraphyllidean cestodes are cosmopolitan, remarkably host specific, and form the most speciose and diverse group of helminths infecting elasmobranchs (sharks, skates and rays). They show substantial interspecific variation in a variety of morphological traits, including body size. Tetraphyllideans represent therefore, an ideal group in which to examine the relationship between parasite body size and abundance. The individual and combined effects of host size, environmental temperature, host habitat, host environment, host physiology, and host type (all likely correlates of parasite body size) on parasite length were assessed using general linear model analyses using data from 515 tetraphyllidean cestode species (182 species were included in analyses). The relationships between tetraphyllidean cestode length and intensity and abundance of infection were assessed using simple linear regression analyses. Due to the contrasting morphologies between shark and batoid hosts, and contrasting physiologies between sharks of the Lamnidae family and other sharks, analyses were repeated in different subsets based on host morphology and physiologies (“sharks” vs. batoids) to determine the influence of these variables on adult tetraphyllidean tapeworm body size. Results presented herein indicate that host body size, environmental temperature and host habitat are relatively important variables in models explaining interspecific variations in tetraphyllidean tapeworm length. In addition, a negative relationship between tetraphyllidean body size and intensity of infection was apparent. These results suggest that space constraints and ambient temperature, via their effects on metabolism and growth, determine adult tetraphyllidean cestode size. Consequently, a trade-off between size and numbers is possibly imposed by external forces influencing host size, hence limiting physical space or other resources available to the parasites.     

Monospecific helminth and arthropod infections in an urban population of brown rats from Doha, Qatar

Parasitic infections were studied for the first time in an urban population of brown rats (Rattus norvegicus) from Doha. Only one species of helminth was found, the cestode Hymenolepis diminuta, and one ectoparasite, the flea Xenopsylla astia, from a sample size of 136 rats (52 males and 84 females). The prevalence of H. diminuta was 17.6%, increasing with host age but not in relation to host sex nor season of capture. Host age was a key factor in influencing abundance of infection, although there was a significant three-way interaction with season and host sex arising through heavy infections in juvenile male rats in the summer. The prevalence of X. astia was 45.6%, although both prevalence and abundance of infestations were season and host age dependent. In the winter prevalence and abundance were similar in both host age and sex groups, but in the summer both parameters of infestation were markedly higher among juveniles compared with adults. We found evidence for some association between these two species: H. diminuta was more prevalent among rats with fleas than among those without, although this association was season-, and independently sex- and age-dependent. There were no quantitative interactions and reasons for this are discussed in relation to the foraging and breeding behaviour of the brown rat in Qatar.     

Description and genetic characterisation of Hysterothylacium (Nematoda: Raphidascarididae) larvae parasitic in Australian marine fishes

Nematodes belonging to the genus Hysterothylacium (family Raphidascarididae) infect various species of marine fish in both the larval and adult stages. Humans can be accidentally infected upon eating infected seafood. In spite of their importance, relatively little is known of their occurrence and systematics in Australia. An examination of various species of marine teleosts in Australian waters revealed a high prevalence of Hysterothylacium larval types. In the present study, seven previously undescribed Hysterothylacium larval morphotypes (V to VII and IX to XII) were discovered. In total we found 10 different morphotypes and we genetically characterised nine morphotypes identified. A morphological dichotomous identification key has been established to differentiate these morphotypes. Since some larvae of Hysterothylacium from marine fishes cannot be differentiated morphologically from other nematode larvae, such as Paraheterotyphlum, Heterotyphlum, Iheringascaris and Lapetascaris, the first and second internal transcribed spacers (ITS-1 and ITS-2) of nuclear ribosomal DNA (rDNA) of these larvae were characterised to confirm their taxonomic status. This genetic characterisation implied that some distinct morphotypes belong to different developmental stages of the same species. In addition, it revealed that some morphotypes can comprise distinct genotypes. No match was found between ITS-1 and ITS-2 sequences obtained from larvae in the present study and those from adults available in the GenBank, highlighting the lack of knowledge on occurrence of adult nematodes infecting Australian fish.     

Complete resistance to challenges with Hymenolepis nana cysticercoids derived from mouse, rat and beetle in mice

and 1986. Complete resistance to challenges with Hymenolepis nana cysticercoids derived from mouse, rat and beetle in mice. International Journal for Parasitology 16: 623–628. When BALB/c and dd strains of mice were given eggs of Hymenolepis nana, they all became completely resistant not only to challenge with mouse-derived cysticercoids but also to challenges with rat-derived and beetle-derived cysticercoids. Serum IgG antibodies at 47–60 days post egg inoculation reacted strongly with these three different host-derived cysticercoids when examined by IFA test, but IgA and IgM isotypes reacted very weakly. Antibodies of infected mouse sera (IgG, IgM and IgA were examined) reacted not only with the protoscolex (scolex of the excysted juvenile) but also with the outer cyst wall. By contrast, uninfected mouse sera and immune sera prepared seven days post cysticercoid inoculation did not react at all. Antigens of both cyst wall and protoscolex appeared to be of parasite origin and not of host origin, and appeared similar in parasites from the different host species.