Acanthocephala from lake fishes in Wisconsin: ecology and host relationships of Pomphorhynchus bulbocolli (Pomphorhynchidae)

Pomphorhynchus bulbocolli Linkins in Van Cleave, 1919, was considerably more common in fishes of the river-connected Tichigan Lake than of the landlocked Silver Lake, southeastern Wisconsin. It is reported from 17 species of principal, accessory, and occasional definitive hosts (new record in Moxostoma carinatum) and from 13 species of paratenic hosts (new records in Amia calva, Ictalurus punctatus, Lepomis cyanellus, and Pomoxis nigromaculatus). Infection patterns were influenced by fish species, feeding behavior, temperature, availability of intermediate host, type of water body, fish movement, and changes in fish host community. Host roles are not fixed but are often interchangeable. A seasonal cycle in prevalence, intensity, and maturation was evident, with greatest abundance and maturation during summer and recruitment during summer and autumn. Recruitment of new infections, development, and release of eggs, however, occurred all year. Sex ratio changed from near equal in new infections to one more highly in favor of females in older adults. Female fish were considerably more frequently and heavily infected than males. No relationship with fish age (size) was evident. Worms were mostly attached in posterior intestinal locations but initial establishment sites correlated with temperature. Translocation of P. bulbocolli due to competitive exclusion in concurrent infections was not observed. The significance of extraintestinal larval forms in the cycle of transmission was noted.     

Effects of coinfection with Pomphorhynchus bulbocolli on development of leptorhynchoides thecatus (Acanthocephala) in amphipods (Hyalella azteca)

The effect of the presence of Pomphorhynchus bulbocolli on the development of Leptorhynchoides thecatus (Acanthocephala) in amphipod intermediate hosts (Hyalella azteca) was examined. Two groups of amphipods were exposed either to eggs of both species (experimental) or only to L. thecatus eggs (control). Amphipods of both groups were dissected 39 days postexposure. The percentage of L. thecatus at the cystacanth stage and mean abundance of cystacanths and precystacanths per amphipod were determined. Contingency table analysis and Fisher's exact tests demonstrated that a significantly smaller proportion of L. thecatus reached the cystacanth stage in coinfected amphipods of the experimental group than either worms in the control group or worms in L. thecatus-only infections of the experimental group. Interspecific interaction between acanthocephalans in intermediate hosts has not been reported previously. Coinfection reduces the number of L. thecatus in amphipods available for transmission to definitive hosts. Habitat separation by eggs of L. thecatus and P. bulbocolli might reduce this effect if amphipods are less likely to become coinfected when eggs are in different microhabitats than when they are not.     

Population dynamics of Pomphorhynchus bulbocolli in Gammarus pseudolimnaeus

The population dynamics of larval Pomphorhynchus bulbocolli were studied in the amphipod intermediate host, Gammarus pseudolimnaeus. Amphipods were collected monthly from April 1982 to March 1983 from a study site at the West Fork of Drake's Creek, Simpson Co., Kentucky. The number of infected amphipods per m2 was lowest (0.6) in May and highest (27.3) in November. A total of 2,502 amphipods was examined; 1,575 (62.9%) were infected with P. bulbocolli. Infection intensity ranged from 1 to 12 with a mean of 1.83 +/- 0.03. Single and multiple worm infections were found in 800 (50.8%) and 775 (49.2%) amphipods, respectively. Female amphipods had higher prevalence and intensity of infections than males but differences were not significant. Prevalence in the monthly samples was lowest in May (5.1%) and highest in October (88.1%). The monthly mean intensity of infection was lowest in July (1.10 +/- 0.04) and highest in November (2.02 +/- 0.08). Amphipods in length classes 3 through 15 were infected with P. bulbocolli. In late summer and fall, amphipods in all length classes were infected, while in late winter and spring only large amphipods were infected. Recruitment of P. bulbocolli into amphipods was seasonal and the total infrapopulation within an amphipod was usually acquired at one time. There was little correlation between size and number of larvae present.     

Proximate factors affecting the larval life history of Acanthocephalus lucii (Acanthocephala)

The growth and eventual size of larval helminths in their intermediate hosts presumably has a variety of fitness consequences. Therefore, elucidating the proximate factors affecting parasite development within intermediate hosts should provide insight into the evolution of parasite life histories. An experimental infection that resulted in heavy intensities of an acanthocephalan (Acanthocephalus lucii) in its isopod intermediate host (Asellus aquaticus) permitted the examination of parasite developmental responses to variable levels of resource availability and intraspecific competition. Isopods were infected by exposure to egg-containing fish feces, and larval infrapopulations were monitored throughout the course of A. lucii development. The relative rate of parasite growth slowed over time, and indications of resource constraints on developing parasites, e.g., crowding effects, were only observed in late infections. Consequently, the factors likely representative of resource availability to larval parasites (host size and molting rate) primarily affected parasite size in late infections. Moreover, at this stage of infection, competitive interactions, gauged by variation in worm size, seemed to be alleviated by greater resources, i.e., larger hosts that molted more frequently. The relatively rapid, unconstrained growth of young parasites may be worse for host viability than the slower, resource-limited growth of larger parasites.     

Life history and population biology of adult Acanthocephalus lucii (Acanthocephala: Echinorhynchidae)

The life history and population biology of adult A. lucii in perch, Perca fluviatilis L., from the Forth and Clyde canal, Scotland, was investigated during May 1979-September 1981. There was an annual cycle in the size of the parasite population; prevalence and abundance (+/- SE) were highest during late spring and summer (70-90% and 14 +/- 4.3 to 16 +/- 5.6 worms/fish, respectively) but declined during late autumn and reached a minimum during winter (50-60% and 2.1 +/- 0.9 to 3.2 +/- 0.6 worms/fish). Parasite maturation was associated with higher water temperatures during spring and summer and most shelled acanthors were probably produced during summer and fall. There was only 1 generation of A. lucii per year, although generations tended to overlap and individuals within each generation did not develop synchronously. The sex ratio of adults was initially near unity but favoured females in the later stages of the infection. The distribution of A. lucii among perch was highly aggregated and stomach content analysis suggested that this was partly due to heterogeneity in perch feeding behaviour. The negative binomial and Poisson lognormal models fitted the data on worm distribution. Seasonal changes in the degree of parasite aggregation were detected, but no conclusive evidence of density-dependent controls on parasite population growth was obtained.     

Acanthocephalus bufonis (Acanthocephala) from Bufo marinus (Bufonidae: Amphibia) in Hawaii.

During a survey of the helminth parasites of the introduced toad, Bufo marinus, on O'ahu, Hawaii, an acanthocephalan corresponding to Acanthocephalus bufonis (Shipley, 1903) was found in the intestinal tract. This is a new host and locality record for A. bufonis which has only previously been recorded from amphibians in the Orient. Possible mechanisms for the introduction of A. bufonis to Hawaii, and its transmission to the toad, are discussed. Almost 98% of toads were infected with a mean intensity of 28.6 acanthocephalans per infected toad. There was a significant negative correlation between host length and intensity of infection with subadult toads having significantly higher infection levels than adult male and female toads. Trunk length of both male and female acanthocephalans was significantly related to host length.     

A new species of Pomphorhynchus (Acanthocephala: Palaeacanthocephala) in freshwater fishes from central Chile

This study describes a new species of Pomphorhynchus collected from Percilia gillissi Girard, 1855 from the Za?artu canal, between the sister basins of the Itata and Laja rivers, in central Chile. Pomphorhynchus moyanoi n. sp. is characterized by an asymmetrical, well-differentiated subspherical bulb and 12-14 longitudinal rows of 13-14 hooks; the third and the fourth hook in each row are stout. Among South American species, P. moyanoi n. sp. shows some similarities to the Chilean species P. yamagutii Schmidt & Hugghins, 1973, but it differs in having a longer neck, larger bulb, and different proboscis armature arrangement. Pomphorhynchus moyanoi n. sp. differs from P. patagonicus Ortubay, Ubeda, Semenas & Kennedy 1991, in the bulb shape (protuberances), number of rows, fourth hook size and basal hook size. Pomphorhynchus moyanoi n. sp. also differs from P. sphaericus in the arrangement of hooks (number of rows and hooks per row), length and width of the proboscis, neck width, and symmetry of the bulb.     

Acanthocephalus saurius n. sp. (Acanthocephala: Echinorhynchidae) and other helminths from the lizard Norops limifrons (Sauria: Polychrotidae) from Costa Rica

Acanthocephalus saurius n. sp. from the small intestine of Norops limifrons collected in Costa Rica is described and illustrated. Acanthocephalus saurius n. sp. represents the third acanthocephalan species in which gravid individuals have been found in lizard hosts. In addition to A. saurius; 1 species of Trematoda, Mesocoelium monas; 3 species of Nematoda, Piratuba digiticauda, Strongyluris panamaensis, and Acuariidae gen. sp. (larvae in cysts) were found.     

Histopathology in Leuciscus cephalus (Pisces: Cyprinidae) resulting from infection with Pomphorhynchus laevis (Acanthocephala).

The histopathology present in Leuciscus cephalus Risso 1826 from the River Po, naturally infected with the acanthocephalan Pomphorhynchus laevis Müller 1776, was studied and described. The parasite was only found within the host's digestive tract, and a density of 10 P. laevis per square centimeter was common. On histological examination it was noticed that both male and female parasites penetrated deeply through all the layers of the host digestive tract by means of their slender neck, bulb and proboscis; thus the bulb and proboscis become encapsulated beneath the serosa and mesentery. As a result, the presence of a capsule was noticed on the external surface of the host digestive tract. P. laevis caused local damage to the intestinal wall, and the parasite elicited an intense host reaction. The response of L. cephalus at the attachment site of the acanthocephalan was hyperplasia of host connective tissue, with cellular infiltration. In a few hosts, the presence of a fibrous nodule on the outer surface of the alimentary canal was observed.     

Seasonality of recruitment and reproduction of Acanthocephalus tumescens (Acanthocephala) in fishes from Lake Moreno (Patagonia, Argentina)

The seasonality of recruitment and reproduction of Acanthocephalus tumescens at the component population level was investigated in Lake Moreno (Patagonia, Argentina). Seasonal samples of the principal fish host species were taken between spring 1999 and spring 2000. Prevalence, mean intensity, coefficient of dispersion, sex ratio, proportion of maturity stages of females, and percentages of the 2 sources of infection in the stomach of fishes were calculated. Overall prevalence, mean intensity, and coefficient of dispersion showed the same pattern of seasonal changes. The seasonal feeding patterns of fishes affect the occurrence of A. tumescens, producing 1 peak in spring and the other peak in autumn. The lower temperature in winter delay reproductive processes after the autumn period of recruitment.     

Movement of Pomphorhynchus bulbocolli larvae from the hemocoel to the peripheral circulation of Gammarus pseudolimnaeus

Acanthors and early acanthellae of Pomphorhynchus bulbocolli were observed in the peripheral circulation of experimentally infected laboratory-reared Gammarus pseudolimnaeus. This has not previously been reported for larval P. bulbocolli during development in the intermediate host. The timing of larval movement from hemocoel to vessels and sinuses was related to intensity of infection. In an experimental group with a low mean intensity of infection, the movement of larval P. bulbocolli into the peripheral circulation occurred later (9 days postfeeding) than in a group with a high mean intensity of infection (2 days postfeeding). Larvae were observed in the peripheral circulation for a longer period of time in amphipods from the group with the high mean intensity of infection. Dead and slightly melanized acanthors and early acanthellae were present in the hemocoel at the time living larvae moved into the peripheral circulation. Only 1 incidence of a hemocytic response to the larvae was observed in the first 20 days postfeeding.     

Differential influence of Pomphorhynchus laevis (Acanthocephala) on the behaviour of native and invader gammarid species.

Although various species of acanthocephalan parasites can increase the vulnerability of their amphipod intermediate hosts to predation, particularly by altering their photophobic behaviour, their influence on the structure of amphipod communities and the success of invader species has so far received little attention. We compared the prevalence and behavioural influence of a fish acanthocephalan parasite, Pomphorhynchus laevis, in two species of amphipods, Gammarus pulex and Gammarus roeseli in sympatry in the river Ouche (Burgundy, eastern France). There, G. pulex is a resident species, whereas G. roeseli is a recent coloniser. Both uninfected G. pulex and G. roeseli were strongly photophobic, although less so in the invading species. However, there was no significant difference in reaction to light between infected and uninfected G. roeseli, whereas infected G. pulex were strongly photophilic. We discuss our results in relation to the parasite's ability to manipulate invading host species, the possibility that resistant individuals have been selected during the invasion process, and the role that acanthocephalan parasites can play in shaping the structure of amphipod communities.     

Effects of Acanthocephalus lucii (Acanthocephala) on intermediate host survival and growth: implications for exploitation strategies

Intermediate host exploitation by parasites is presumably constrained by the need to maintain host viability until transmission occurs. The relationship between parasitism and host survival, though, likely varies as the energetic requirements of parasites change during ontogeny. An experimental infection of an acanthocephalan (Acanthocephalus lucii) in its isopod intermediate host (Asellus aquaticus) was conducted to investigate host survival and growth throughout the course of parasite development. Individual isopods were infected by exposure to fish feces containing parasite eggs. Isopods exposed to A. lucii had reduced survival, but only early in the infection. Mean infection intensity was high relative to natural levels, but host mortality was not intensity dependent. Similarly, a group of naturally infected isopods harboring multiple cystacanths did not have lower survival than singly infected isopods. Isopods that were not exposed to the parasite exhibited sexual differences in survival and molting, but these patterns were reversed or absent in exposed isopods, possibly as a consequence of castration. Further, exposed isopods seemed to have accelerated molting relative to unexposed controls. Infection had no apparent effect on isopod growth. The effects of A. lucii on isopod survival and growth undermine common assumptions concerning parasite-induced host mortality and the resource constraints experienced by developing parasites.     

Life history and population biology of larval Acanthocephalus lucii (Acanthocephala: Echinorhynchidae) in the isopod Asellus aquaticus

Hemocoels of 8,731 Asellus aquaticus collected from the Forth and Clyde canal in Glasgow, Scotland, from January 1980 to March 1981 were examined for larvae of Acanthocephalus lucii. Prevalence and mean intensity were generally low (1.5-8.3% and 1.0-1.6, respectively), but there was a slight seasonal infection pattern with fewer infected isopods during summer, reflecting the appearance of a new isopod generation. Although there were no distinct seasonal trends in the proportions of each larval stage, recruitment of larvae probably occurred mainly during summer and autumn. Some larvae reached the cystacanth stage by late summer or autumn; others overwintered as acanthors or acanthellae and completed development the following spring. The maximum life span of larvae was limited to 1 yr by annual turnover of the isopod population. The distribution of larval A. lucii among isopods was slightly aggregated. There was a peaked pattern in the relationship between isopod length and the prevalence, abundance, and degree of parasite aggregation. The rate of parasite development in laboratory-infected isopods was linearly related to temperature between 9 and 22 C; the temperature threshold was 5.7 C, and the larval parasite required 598 degree-days above threshold to complete development. Among laboratory-infected isopods, 2 mechanisms that could regulate the larval parasite population were detected: intraspecific competition and direct, parasite-induced isopod mortality. However, the intensity of infection in the natural habitat was consistently low and may have remained below the level at which these mechanisms operated.     

Pomphorhynchus patagonicus n. sp. (Acanthocephala: Pomphorhynchidae) from freshwater fishes of Patagonia, Argentina

Pomphorhynchus patagonicus n. sp. is described from Lake Rosario, Chubut Province, Argentina. It is characterized by a long neck forming an asymmetrical bulb with 2 well differentiated dorsal protruberances and 14 alternating rows of 13 and 14 proboscis hooks, each row with a stout fourth hook. It most closely resembles Pomphorhynchus sebastichthydis Yamaguti, 1939, from Japan, but differs in the bulb protruberances and in having more rows of hooks and more hooks per row and in the shapes of the fourth and basal hooks. Among American species P. patagonicus shows some similarities to Pomphorhynchus yamagutii Schmidt and Hugghins, 1973, from Chile, but differs with respect to the longer neck, bulb protruberances, and proboscis armature. Pomphorhynchus patagonicus is endemic to Patagonia, where its definitive type host is the endemic fish Patagonina hatcheri (Atherinidae) and its intermediate host the endemic freshwater amphipod Hyalella patagonica. It has been found also in autochthonous fishes belonging to the families Galaxiidae and Percichthyidae and in introduced salmonid fishes.