The life cycle and transmission dynamics of the larval stages of Hypoderaeum conoideum

The morphology of the different larval stages and life cycle of Hypoderaeum conoideum (Trematoda: Echinostomatidae) are described. The freshwater snail species Lymnaea peregra (Gastropoda: Lymnaeidae) serves as the natural first intermediate host and this and L. corvus serve as experimental first intermediate hosts. These and other freshwater snails, such as Physella acuta and Gyraulus chinensis, in turn serve as second intermediate hosts. Adult worms were obtained from chicks and ducks, but not from rats, mice and golden hamsters. The morphology of the larval stages is compared with previous work on H. conoideum. Several aspects of the biology of the life history stages are described with emphasis on the transmission dynamics of the free-living stages. Differential suitability of the snail species that may act as first and/or second intermediate hosts is studied and discussed.     

Temporal variation in parasite loading in relation to life history patterns of Littorina obtusata and L. fabalis

Parasitic infection of Littorina obtusata (L.) and Littorina fabalis Turton by larval digeneans was studied in summer (August) and winter (February) on a sheltered shore at St Michael's Island, Isle of Man. Sampling was timed to investigate parasite loading at different stages of the littorinids' life history. Nine species of digeneans were identified and infection rates were similar between the species, with a slightly higher prevalence in females for L. obtusata and males in L. fabalis. Temporal differences in infection were small, although metacercariae and degenerate infections were greater in female L. obtusata in February. Analysis of shell parameters and size of the penis in males and the pallial oviduct in females revealed morphological differences between parasitised and uninfected individuals. Parasitic stunting of the penis was evident in both species and the pallial oviduct was also smaller in parasitised females. There was also a tendency for parasitised individuals to have larger shells; although size also varied with season for L. fabalis.     

Variations in the number of metacercariae in the daughter sporocysts of Microphallus pygmaeus (Trematoda: Microphallidae)

The daughter sporocystes of Microphallus pygmaeus from the snails of Littorina saxatilis and L. obtusata were investigated. The average quantity of metacercariae in sporocysts is influenced by host species, its syze and by the season. There are differences in the average number of larvae from parthenites of L. saxatilis, collected in the Barents and White seas.     

Impact of trematode parasitism on the fauna of a North Sea tidal flat

The impact of larval trematodes on the fauna of a North Sea tidal flat is considered at the individual and at the population level, depicting the digenean parasites of the common periwinkle,Littorina littorea, and their life cycles, as an example. On the German North Sea coast,L. littorea is first intermediate host for 6 larval trematodes representing 6 digenean families —Cryptocotyle lingua (Heterophyidae),Himasthla elongata (Echinostomatidae),Renicola roscovita (Renicolidae),Microphallus pygmaeus (Microphallidae),Podocotyle atomon (Opecoelidae) andCercaria lebouri (Notocotylidae). All exceptP. atomon utilize shore birds as final hosts; adultP. atomon parasitize in the intestine of teleosts, mainly pleuronectid flatfish. Second intermediate hosts ofC. lingua are various species of fish; the cercariae ofH. elongata encyst in molluscs and polychaetes, those ofR. roscovita in molluscs;M. pygmaeus has an abbreviated life cycle;C. lebouri encysts free on solid surfaces; and the fish trematodeP. atomon utilizes benthic crustaceans, mainly amphipods, as second intermediate hosts. On the tidal flats of the Königshafen (Sylt), up to 77% of the periwinkles have been found to be infested by larval trematodes. Maximum infestations in individual samples were 23% forC. lingua, 47% forH. elongata and 44% forR. roscovita. The digeneans cause complete parasitic castration of their carriers and hence exclude a considerable proportion of the snails from the breeding population. Infestation reduces the longevity of affected hosts, and size-related, trematode-induced differential mortality causes changes in the normal size-frequency distribution of individual snail-age classes. Young flatfishPleuronectes platessa from the Königshafen are 100% infested with metacercariae ofC. lingua. Heavy infestation of the gills causes obstruction of blood vessels and respiratory impairment; metacercariae in the eyes and optic nerves cause visual and neurological disturbances. A single metacercaria is sufficient to kill a larval fish.Mytilus edulis andCardium (Cerastoderma) edule are 100% infested with metacercariae ofH. elongata. Heavy infestation impairs the byssus-thread production in mussels and affects the burrowing ability of cockles. Longevity and resistance to environmental — particularly thermal — stress are reduced in bivalves infested withH. elongata andR. roscovita. There is evidence that, in the study area, population size and age composition of the molluscs discussed are (indirectly) controlled by trematode parasites employing sea birds as final hosts, rather than directly by the predatory activities of these birds.     

Host specificity and molecular phylogeny of larval Digenea isolated from New Zealand and Australian topshells (Gastropoda: Trochidae)

The maintenance of strict host specificity by parasites when several closely related host species live in sympatry is poorly understood. Species of intertidal trochid snails in the genera Diloma, Melagraphia and Austrocochlea often occur together and are parasitised by a single digenean morphotype (Platyhelminthes, Trematoda), tentatively placed in Opecoelidae. Of the 10 trochid species (6 from New Zealand, 4 from Australia) we examined, six were found to be infected, and the prevalence of infection was as high as 17.5%. We obtained molecular data (mitochondrial 16S and nuclear rDNA internal transcriber spacer 2 sequences, representing 774 bp), to infer phylogenetic relationships amongst these Digenea. Our phylogeny separated the single morphotype into three clearly defined clades (which are almost certainly separate biological species): (i) those infecting two species of Austrocochlea from Tasmania, (ii) those infecting Diloma subrostrata in Otago and Southland, New Zealand, and (iii) those infecting all the parasitised New Zealand topshells (Melagraphia aethiops, D. subrostrata, Diloma nigerrima and Diloma arida) throughout the country. This last group comprised two subclades, one infecting only D. subrostrata and one infecting the other three species. Two D. subrostrata populations were each found to be infected by genetically distinct parasites, yet sympatric populations of the other snails were not necessarily infected. This study is thus the first to reveal cryptic species of digeneans in a single population of a molluscan first intermediate host. We point out also that the degree of host specificity would have been grossly underestimated if, in the absence of our genetic analysis, we had only considered digenean morphology. Our results shed light on the conditions that may favour switching among intermediate hosts in digeneans, and on the presence/absence of host specificity in these parasites.     

Spatial and temporal variation of trematode infection in coexisting populations of intertidal gastropods Littorina saxatilis and L. obtusata in the White Sea

Trematode infection was studied in sympatric populations of the periwinkles Littorina saxatilis and L. obtusata in 2 regions of Kandalaksha Bay of the White Sea to assess host-parasite interactions at the population level. Twenty-seven spatially separated populations were each surveyed in 1984-1994; 2 heavily infected populations were investigated annually over a 16 yr period. Ten trematode species were found in the periwinkle populations. The closest association in spatial distribution and temporal dynamics was observed between 3 ecologically and morphologically similar trematodes of the 'pygmaeus' group: Microphallus piriformes, M. pygmaeus and M. pseudopygmaeus. For these 3 species, the prevalences were closely associated in the 2 host species when spatially separated sites from the 2 studied regions were considered, while in the 2 populations studied over the 16 yr period, a correlation was only observed between the infection levels of L. saxatilis and L. obtusata by either M. piriformes and immature microphallids. Likewise, within each host species, significant correlations were revealed between the prevalence of the different microphallids of the 'pygmaeus' groups. However, they were fewer and weaker when the long-term dynamics of infection in the 2 heavily infected populations were considered. Most other trematodes did not show significant association in prevalence either within or between the 2 host species on spatial and temporal scales.     

Comparison of digenean faunas of sillaginid fishes from inshore and offshore waters of Visakhapatnam Coast, Bay of Bengal

The sillaginid fishes of Visakhapatnam Coast, Bay of Bengal, are found to be hosts for 16 species of digeneans of which seven are adult digeneans belonging to families Opecoelidae, Zoogonidae and Hemiuridae and nine are larval digeneans representing families Bucephalidae, Heterophyidae, Cryptogonimidae, Acanthocolpidae, Hemiuridae and Didymozoidae. The digenean fauna of sillaginid fishes from shallow coastal waters (< 10 m depth) and deeper offshore waters (30–50 m depth) are compared and results are discussed in the light of existing information on the migratory movements of different species of sillaginids and their occurrence in coastal or offshore waters. The usefulness of metacercariae of these fishes as 'biological tags' is considered.     

The nature and evolution of the association among digeneans, molluscs and fishes

Patterns of association of digenean families and their mollusc and vertebrate hosts are assessed by way of a new database containing information on over 1000 species of digeneans for life-cycles and over 5000 species from fishes. Analysis of the distribution of digenean families in molluscs suggests that the group was associated primitively with gastropods and that infection of polychaetes, bivalves and scaphopods are all the results of host-switching. For the vertebrates, infections of agnathans and chondrichthyans are apparently the result of host-switching from teleosts. For digenean families the ratio of orders of fishes infected to superfamilies of molluscs infected ranges from 0.5 (Mesometridae) to 16 (Bivesiculidae) and has a mean of 5.6. Individual patterns of host association of 13 digenean families and superfamilies are reviewed. Two, Bucephalidae and Sanguinicolidae, are exceptional in infecting a range of first intermediate hosts qualitatively as broad as their range of definitive hosts. No well-studied taxon shows narrower association with vertebrate than with mollusc clades. The range of definitive hosts of digeneans is characteristically defined by eco-physiological similarity rather than phylogenetic relationship. The range of associations of digenean families with mollusc taxa is generally much narrower. These data are considered in the light of ideas about the significance of different forms of host association. If Manter's Second Rule (the longer the association with a host group, the more pronounced the specificity exhibited by the parasite group) is invoked, then the data may suggest that the Digenea first parasitised molluscs before adopting vertebrate hosts. This interpretation is consistent with most previous ideas about the evolution of the Digenea but contrary to current interpretations based on the monophyly of the Neodermata. The basis of Manter's Second Rule is, however, considered too flimsy for this interpretation to be robust. Problems of the inference of the evolution of patterns of parasitism in the Neodermata are discussed and considered so intractable that the truth may be presently unknowable.     

The role of Bithynia tentaculata in the transmission of larval digeneans from a gravel pit in the Lower Thames Valley

The freshwater snail, Bithynia tentaculata (Gastropoda: Prosobranchia), is frequently found in gravel pits and its role in the transmission of larval digeneans was assessed in one of these unstable water bodies from the Lower Thames Valley (UK) from June 1982 to December 1984. Eight species of cercarial and five species of metacercarial infections were reported from B. tentaculata, with up to 7.7% and 4.7% prevalence of infections, respectively, occurring in the snail, making it the most important host at this site. Seasonal peaks in cercarial infections occurred in late autumn/early winter but little seasonality was apparent in the occurrence of metacercarial infections. The life history characteristics of both hosts and parasites, which can affect intramolluscan digenean dynamics, are discussed.     

洞庭湖外睾吸虫新种及其生活史

本文报告洞庭湖区鲶鱼肠道寄生的洞庭湖外睾吸虫Exorchis dongtinghuensis sp.nov(新种)及其全程生活史,其第一中间宿主为湖北钉螺Oncomelania hupensis;第二中间宿主为鲤鱼、鲫鱼和金鱼;终宿主为鲶鱼Parasilurus asotus。作者对各期宿主作了人工感染试验和现场自然感染调查。对其发育过程作了观察比较。     

Effect of the digenean parasites of fish on the fauna of Mediterranean lagoons

Attention is drawn to the effects of parasites on their hosts, taking as a model the digenean parasites of teleosts (hereafter: fish) from lagoons along the French Mediterranean coast. Because digeneans have a heteroxenic life cycle, their impact is not limited to the definitive host, which harbours the sexual adults, but is extended to the first host (mollusc) and to the second host ("invertebrate" or fish). Adult parasites, in order to ensure efficient sexual reproduction, never cause excessive damage to their definitive host, usually only exploiting the intestinal fluids; however, the host must intensify its search for prey, which results in a diminished fitness. Within the first host, 'larval' stages of digenean parasites invade the gonads, resulting in its castration, then exhaustion and eventually death. The diversion of energy from the second hosts towards the parasites forces them to intensify their search for food, resulting in decreased fitness and an increased risk of being eaten; in addition, manipulation of the host's behaviour by parasites drives this host into the food chain of the definitive host. In lagoons, many individuals of almost all species of fish and invertebrates act as first, second and/or definitive hosts for digeneans. Obviously, parasites have a severe impact on the population dynamics of key taxa, on the food web and therefore also on the functioning of the whole lagoon ecosystem. Yet this impact has been largely overlooked or underestimated in functioning models, by ecologists, who tend to prioritize more apparent trophic relationships.     

Redescription and life cycle of the monorchiid Postmonorcheides maclovini Szidat, 1950 (Digenea) from the Southwestern Atlantic Ocean: Morphological and molecular data

The adult monorchiid, Postmonorcheides maclovini Szidat, 1950, digenean parasite of the Patagonian blennie Eleginops maclovinus (Cuvier) (Eleginopidae) from Puerto Deseado (47° 45′ S, 65° 55′ W), Argentina, was characterized and its life cycle elucidated. P. maclovinus is the only species of the genus Postmonorcheides, proposed by Szidat (1950) from Tierra del Fuego province (~ 54° S), Argentina. This digenean uses the Patagonian blennie as definitive host, and the intertidal bivalve Lasaea adansoni (Gmelin) (Lasaeidae) as both first and second intermediate hosts (metacercariae encyst inside sporocysts), being the first record of this clam as intermediate host of trematode parasites. The cercaria may, in addition to encysting in the sporocyst, emerge and presumably infect other intermediate hosts. This is the second report of a monorchiid species with metacercariae encysting inside the sporocyst. Adults were found parasitizing the fish stomach, pyloric caeca and intestine with a prevalence of 100%; sporocysts with cercariae and/or metacercariae were found parasitizing the gonad of the bivalve with a prevalence of 2.78%. The cercariae possess a well-developed tail and eye-spots are absent. The ITS1 sequence from the adult digeneans found in the Patagonian blennie, identified as P. maclovini, was found to be identical to the ITS1 sequences obtained both from sporocysts containing cercariae and encysted metacercariae found in L. adansoni.     

Does a facultative precocious life cycle predispose the marine trematode Proctoeces cf. lintoni to inbreeding and genetic differentiation among host species?

Intraspecific variability in parasite life cycle complexity (number of hosts and species of hosts in the life cycle) may have an impact how parasite genetic variation is partitioned among individual parasites, host individuals or host species within a given area. Among digenean trematodes, a three-host life cycle is common. However, a few species are precocious and may reach sexual maturity in what is typically regarded as the second intermediate host. The objective of this study was to determine whether a precocious life cycle predisposes digeneans to possible inbreeding or genetic subdivision among host species. As a study system, we used the digenean Proctoeces cf. lintoni whose metacercariae precociously mature (facultative) without a cyst wall in the gonads of multiple sympatric species of keyhole limpets (Fissurella spp.), typically regarded as the second intermediate hosts. Genotyped parasites were collected from four species of limpets and the clingfish Sicyases sanguineus, the third and final host where sexual maturity occurs. We found very high microsatellite diversity, Hardy–Weinberg equilibrium over all genotyped individuals, and little to no genetic structuring among parasites collected from the different host species. The fact that metacercariae do not encyst in the keyhole limpets, coupled with the high mixing potential of an aquatic environment, likely promote panmixia in local populations of P. cf. lintoni.     

Larval digenean community parasitizing the freshwater snail, Chilina dombeyana (Pulmonata: Chilinidae) in Patagonia, Argentina, with special reference to the notocotylid Catatropis chilinae

The objective of this research was to describe the structure of larval digenean communities in the freshwater snail Chilina dombeyana in Lake Mascardi, an oligotrophic Andean Patagonian lake. In total, 1,923 snails were collected during 2 sampling periods. Specimens were examined, and 8 digenean species were identified, all with allogenic life cycles. The snail infracommunities nearly always occurred as single-species infections, distributed mainly in the hepatopancreas. The double infections (0.2%) were always prepatent, and involved a schistosome and the notocotylid Catatropis chilinae. The overall prevalence, the prevalence of C. chilinae, and total species prevalence (without C. chilinae) significantly increased with snail size. Catatropis chilinae was the dominant species in all months and across all snail size classes. Maximum richness was reached in the size class between 16 and 18 mm. Diversity indices, i.e., the Shannon-Wiener Index and the reciprocal of the Simpson Index, increased with snail size only during the second sampling period. No robust evidence of antagonistic interactions was found in the digenean community of C. dombeyana from Lake Mascardi.     

Digeneans of fishes from the Adriatic Sea with a description of Lecithaster atherinae n. sp. from Atherina (Hepsetia) boyeri

A collection of whole mount preparations of digeneans was studied. The parasites were collected from several species of fishes caught in a fishing area of the South Adriatic Sea. A host-parasite list is given. Twenty species of fish were analysed and twenty-seven species of digeneans were found, twenty of which had previously been described in the same host, and six of which were observed in new hosts and/or localities. One digenean, hitherto undescribed, is illustrated in this paper and described as Lecithaster atherinae n.sp. The new species can be distinguished from L. bombayensis by the testes not being lobed; from L. extralobus by having four rather than five ovarian lobes; from L. indicus and L. maeoticus by the ovarian lobes not being finger-like; from L. leiostomi by having definitely smaller eggs.     

Mechanisms underlying digenean-snail specificity: role of miracidial attachment and host plasma factors

Digenetic trematodes usually show a high degree of specificity for their molluscan intermediate hosts. A panel of 4 digenean species (Echinostoma paraensei, E. trivolvis, Schistosoma mansoni, and Schistosomatium douthitti) and 5 snail species (Biomphalaria glabrata, Helisoma trivolvis, Lymnaea stagnalis, Stagnicola elodes, and Helix aspersa representing 3 gastropod families) was used to assess the relative contributions of miracidial behavior, host plasma osmolality, and host plasma factors in dictating specificity. Additional experiments were undertaken with a fifth digenean, Echinoparyphium sp. Expected patterns of compatibility were first confirmed; each parasite species produced patent infections in its known snail host, but not in the other snail species. One exception was S. douthitti, which unexpectedly did not infect L. stagnalis. As judged by direct observation and by noting their disappearance after exposure to snails, miracidia were generally less likely to attach to or penetrate incompatible than compatible hosts. However, over half of the miracidia of each parasite species attached to or attempted penetration of both compatible and incompatible hosts, suggesting that under the experimental conditions used, miracidial host location and attachment behaviors were not of overriding importance in dictating observed patterns of specificity. For each digenean species, the percentage of larvae that became immobile, rounded, showed tegumental damage, or died over a 6-hr interval in plasma of the various snails was assessed. In no case was plasma from a compatible host harmful to sporocysts or rediae. In contrast, in 8 of 16 (50%) incompatible combinations, snail plasma had a significant negative effect on sporocyst condition. In 4 of 12 (33%) incompatible combinations, plasma had a significant negative effect on rediae. In 9 of 10 combinations tested, lymnaeid plasma was toxic for the parasites of planorbid snails and in 2 of 4 combinations, planorbid plasma was toxic for the parasites of lymnaeid snails. Toxicity was not attributable to differences in plasma osmolality between snail species. The ability of plasma from incompatible snails to affect viability of both sporocysts and rediae was surprisingly strong, suggesting that humoral factors play a greater role in dictating patterns of digenean-snail specificity than previously appreciated.     

Trematode fauna of prosobranch snails of the genus Semisulcospira in Lake Biwa and the connected drainage system

The parasite fauna of prosobranch snails of the genus Semisulcospira was surveyed in Lake Biwa and the adjacent water system. One aspidogastrean and 28 digenetic trematode taxa were detected in 19209 snails consisting of 10 morphological species. There was no trematode species peculiar to members of the subgenus Biwamelania that is endemic to the Lake Biwa water system. However, one species, Notocotylus magniovatus, was found only in the non-endemic subgenus Semisulcospira. Of 23 digenean taxa detected in more than one host, 13 were distributed in both the lake and the tributaries. Seven of these had host taxa, more than 1% of which were infected with the parasite in both the lake and the tributaries, four had such hosts only in the tributaries, and two had no such hosts. Three species detected only in Lake Biwa were previously reported from other rivers in Japan. In the seven species detected only in the tributaries, two species had life cycles that could be maintained only in rivers. These results indicate that the core areas for the distribution of parasites of Semisulcospira are tributaries, and the lake is a sink for these species. These results contradict the expectation that the parasite fauna should be richer in the lake than in tributaries because the lake is a stable habitat over a geological time scale and has more divergent freshwater animals than the adjacent water system.     

The association of Zygocotyle lunata and Echinostoma trivolvis with Chaetogaster limnaei, an ectosymbiont of Helisoma trivolvis

Helisoma trivolvis snails collected from a lake in Warren County, New Jersey, in June 2007 possessed the ectosymbiont Chaetogaster limnaei (Annelida). Some of these snails were also infected with larval stages of Zygocotyle lunata and Echinostoma trivolvis. Chaetogaster limnaei associated with the infected snails fed on the cercariae of both digeneans. Zygocotylae lunata cercariae were observed in the stomach of C. limnaei and whole cercariae were loosely attached to the ventral surface of the chaetogasters. Cercariae in the stomachs were digested within 48 hr and probably served as a source of nutrient for the annelids. Whole cercariae and 1 nonviable metacercaria of E. trivolvis were seen in the stomachs of the chaetogasters. The protective action of the chaetogasters on the transmission of the cercariae of E. trivolvis and Z. lunata to second intermediate hosts in the wild awaits further study.     

Modification of the structure of penial glands in males of the intertidal molluscs Littorina saxatilis and L. obtusata under the influence of the infestation by trematode parthenites

Sporocycts of the "pygmaeus" microphallides (Microphallus piriformes) are localized in hepatopancreas and gonads of Littorina molluscs causing total parasitic castration. A histological study of penial glands in Littorina saxatilis and L. obtusata males infested with trematodes M. piriformes has been made. Copulatory organs of noninfested molluscs, molluscs after recent contamination (with not completely formed daughter sporocysts), and molluscs containing mature metacercariae inside daughter sporocysts were examined. Based on the data obtained, probable dynamics of the histological structure of infested glandular apparatus was established. It was shown, that the trematode infestation have an influence on the muscular and secretory parts of penial glands. The wall of the penial gland muscular capsule becomes more fine in infested L. saxatilis. On the contrary, this wall is vastly thicker in infested L. obtusata, as compared with noninfested individuals. Glandular cells of the molluscs' penial glands decreases the amount of granular secret in both species. In L. obtusata the number of secretory cells is shown to be reduced up to their total disappearance. The above pathological changes probably prevent normal function of penial glands.