The mallard Anas platyrhynchos is the most abundant water bird species in Austria, but there is no record of its helminth community. Therefore, this work aimed to close that gap by recording and analysing the parasite community of a large number of birds from Austria for the first time. A total of 60 specimens shot by hunters in autumn were examined for intestinal parasites. The following taxa were recovered (prevalence given in parentheses): Cestoda: Diorchis sp. (31.7%) and Fimbriarioides intermedia (1.7%); Acanthocephala: Filicollis anatis (5%), Polymorphus minutus (30%) and one cystacanth unidentified (1.7%); Trematoda: Apatemon gracilis (3.3%), Echinostoma grandis (6.7%), Echinostoma revolutum (6.7%) and Notocotylus attenuatus (23.3%); Nematoda: Porrocaecum crassum (1.7%) and one not identified (1.7%). The frequency distribution of parasites showed a typical pattern in which 39 birds (65%) were either not parasitized or were harbouring up to five worms, whereas more intense infestations occurred in a lesser number of hosts. Compared to other studies from central and eastern Europe, an extremely depauperate helminth community, particularly of the cestodes and nematodes, was found. Polymorphus minutus was observed as having highly variable morphology and, therefore, molecular genetic characterization by DNA barcoding was carried out. Species identification was confirmed by comparing data with the reference cytochrome c oxidase subunit 1 gene sequence from P. minutus available in GenBank.

The bird-infecting acanthocephalan Polymorphus minutus has been suggested to comprise different lineages or even cryptic species using different intermediate hosts. To clarify this open question, we investigated Polymorphus cf. minutus cystacanths originating from amphipod intermediate hosts from 27 sites in Germany and France. Parasites and hosts were identified using integrated datasets (COI and/or morphology for hosts and COI + ITS1-5.8S-ITS2 for parasites).

Mitochondrial and nuclear data (ITS1) strongly support the existence of three cryptic species in Polymorphus cf. minutus (type 1-3). These three types reveal a high degree of intermediate host specificity, with Polymorphus type 1 only encountered in Gammarus fossarum type B, Polymorphus type 2 in Echinogammarus sp. and Echinogammarus berilloni, and Polymorphus type 3 in Gammarus pulex and Gammarus roeselii. Our results point to a so far neglected cryptic diversity of the genus Polymorphus in Central Europe. Furthermore, Polymorphus type 2 is most likely a non-native parasite in Germany that co-invaded with E. berilloni from the Mediterranean area. Potentially, type 3 originates from South-East Europe and migrated to Germany by G. roeselii, where it might have captured G. pulex as an intermediate host. Therefore, our findings can be seen in the context of ecological globalization in terms of the anthropogenic displacement of intermediate hosts and its impact on the genetic divergence of the parasites.

Behavioural alterations induced by parasites in their intermediate hosts can spatially structure host populations, possibly resulting in enhanced trophic transmission to definitive hosts. However, such alterations may also increase intermediate host vulnerability to non-host predators. Parasite-induced behavioural alterations may thus vary between parasite species and depend on each parasite definitive host species. We studied the influence of infection with 2 acanthocephalan parasites (Echinorhynchus truttae and Polymorphus minutus) on the distribution of the amphipod Gammarus pulex in the field. Predator presence or absence and predator species, whether suitable definitive host or dead-end predator, had no effect on the micro-distribution of infected or uninfected G. pulex amphipods. Although neither parasite species seem to influence intermediate host distribution, E. truttae infected G. pulex were still significantly more vulnerable to predation by fish (Cottus gobio), the parasite's definitive hosts. In contrast, G. pulex infected with P. minutus, a bird acanthocephalan, did not suffer from increased predation by C. gobio, a predator unsuitable as host for P. minutus. These results suggest that effects of behavioural changes associated with parasite infections might not be detectable until intermediate hosts actually come in contact with predators. However, parasite-induced changes in host spatial distribution may still be adaptive if they drive hosts into areas of high transmission probabilities.

Trophically transmitted parasites are likely to strongly influence food web-structure. The extent to which they change the trophic ecology of their host remains nevertheless poorly investigated and field evidence is lacking. This is particularly true for acanthocephalan parasites whose invertebrate hosts can prey on other invertebrates and contribute to leaf-litter breakdown. We used a multiple approach combining feeding experiments, neutral lipids and stable isotopes to investigate the trophic ecology of the freshwater amphipod Gammarus roeseli parasitized by the bird acanthocephalan Polymorphus minutus. Infected compared to uninfected amphipods consumed as many dead isopods, but fewer live isopods and less leaf material. Infection had no influence on the total concentration of neutral lipids. Contrary to what we expected based on laboratory findings, the nitrogen isotope signature, which allows for the estimation of consumer's trophic position, was not influenced by infection status. Conversely, the carbon isotope signature, which is used to identify food sources, changed with infection and suggested that the diet of infected G. roeseli includes less perilithon (i.e. fixed algae on rocks, stones) but more terrestrial inputs (e.g. leaf material) than that of uninfected conspecifics. This study shows evidence of changes in the trophic ecology of P. minutus-infected G. roeseli and we stress the need to complement feeding experiments with field data when investigating top-down effects of infection in an opportunistic feeder which adapts its diet to the available food sources.

Among the potential effects of parasitism on host condition, the ‘increased host abilities’ hypothesis is a counterintuitive pattern which might be predicted in complex-life-cycle parasites. In the case of trophic transmission, a parasite increasing its intermediate host's performance facing non-host predators improves its probability of transmission to an adequate, definitive host. In the present study, we investigated the cost of infection with the acanthocephalan Polymorphus minutus on the locomotor/escape performance of its intermediate host, the crustacean Gammarus roeseli. This parasite alters the behaviour of its intermediate host making it more vulnerable to predation by avian definitive hosts. We assessed the swimming speeds of gammarids using a stressful treatment and their escape abilities under predation pressure. Despite the encystment of P. minutus in the abdomen of its intermediate host, infected amphipods had significantly higher swimming speeds than uninfected ones (increases of up to 35%). Furthermore, when interacting with the non-host crustacean predator Dikerogammarus villosus, the highest escape speeds and greatest distances covered by invertebrates were observed for parasitized animals. The altered behaviour observed among the manipulated invertebrates supported the ‘increased host abilities’ hypothesis, which has until now remained untested experimentally. The tactic of increasing the ability of infected intermediate hosts to evade potential predation attempts by non-host species is discussed.

Phenotypic alterations induced by parasites in their intermediate hosts often result in enhanced trophic transmission to appropriate final hosts. However, such alterations may also increase the vulnerability of intermediate hosts to predation by non-host species. We studied the influence of both infection with 3 different acanthocephalan parasites (Pomphorhynchus laevis, P. tereticollis, and Polymorphus minutus) and the availability of refuges on the susceptibility of the amphipod Gammarus pulex to predation by 2 non-host predators in microcosms. Only infection with P. laevis increased the vulnerability of amphipods to predation by crayfish, Orconectes limosus. In contrast, in the absence of refuges, the selectivity of water scorpions, Nepa cinerea, for infected prey was significant and did not differ according to parasite species. When a refuge was available for infected prey, however, water scorpion selectivity for infected prey differed between parasite species. Both P. tereticollis- and P. laevis-infected gammarids were more vulnerable than uninfected ones, whereas the reverse was true of P. minutus-infected gammarids. These results suggest that the true consequences of phenotypic changes associated with parasitic infection in terms of increased trophic transmission of parasites deserve further assessment.

Larval helminths often share individual intermediate hosts with other larval worms of the same or different species. In the case of immature acanthocephalans capable of altering the phenotype of their intermediate hosts, the benefits or costs of host sharing can be evaluated in terms of increased or decreased probability of transmission to a suitable definitive host. Competitive interactions among the immature stages of acanthocephalans within the intermediate host could create additional costs of host sharing, however. The effects of intraspecific and interspecific interactions were measured in 3 sympatric species of acanthocephalans exploiting a population of the amphipod Echinogammarus stammeri in the River Brenta, Italy. The strength of interactions was assessed from differences in the size achieved by infective cystacanths in the intermediate host. The size of Pomphorhynchus laevis cystacanths was not correlated with host size, whereas the size of Acanthocephalus clavula and Polymorphus minutus cystacanths increased with host size. Reductions in cystacanth size caused by intraspecific competition were only detected in P. laevis, but may also occur in both A. clavula and P. minutus. When co-occurring in the same amphipod with cystacanths of A. clavula, cystacanths of P. laevis attained a smaller size than when they occurred on their own. This effect was not reciprocal, with the size of A. clavula cystacanths not being affected. This supports earlier suggestions that it is adaptive for A. clavula to associate with P. laevis in amphipod intermediate hosts, with both species going to the same fish definitive hosts. In contrast, cystacanths of P. laevis achieved their largest size when they co-occurred in an amphipod with a cystacanth of P. minutus, which has a different definitive host (i.e. birds). These findings suggest that the net benefits of sharing an intermediate host can only be estimated by taking into account both the effects on transmission success and the consequences for cystacanth development.

When two parasite species are manipulators and have different definitive hosts, there is a potential for conflict between them. Selection may then exist for either avoiding hosts infected with conflicting parasites, or for hijacking, i.e. competitive processes to gain control of the intermediate host. The evidence for both phenomena depends largely on the study of the relative competitive abilities of parasites within their common intermediate host. We studied the effects of simultaneous infection by a fish acanthocephalan parasite, Pomphorhynchus laevis, and a bird acanthocephalan parasite, Polymorphus minutus, on the behaviour of their common intermediate host, the amphipod Gammarus pulex. We compared the reaction to light and vertical distribution of individuals infected with both parasites to those of individuals harbouring a single parasite species and uninfected ones under controlled conditions. Compared to uninfected gammarids that were photophobic and tended to remain at the bottom of the water column, P. laevis-infected gammarids were attracted to light, whereas P. minutus-infected individuals showed a modified vertical distribution and were swimming closer to the water surface. The effects of both P. laevis and P. minutus appeared to be dependent only on their presence, not on their intensity. Depending on the behavioural trait under study, however, the outcome of the antagonism between P. laevis and P. minutus differed. The vertical distribution of gammarids harbouring both parasites was half-way between those of P. laevis- and P. minutus-infected individuals, whereas P. laevis was able to induce altered reaction to light even in the presence of P. minutus. We discuss our results in relation to the occurrence of active avoidance or hijacking between conflicting manipulative parasites and provide some recommendations for future research.