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1.
The distribution of the acanthocephalan Pomphorhynchus laevis along the alimentary tract of several species of freshwaster fish was studied in both natural and experimental infections. The parasite exhibits a clear preference for a particular region of the alimentary canal, which differs from host species to host species, but it is capable of surviving in all regions of the intestine. In some species, but not in all, its growth rate may be reduced in regions outside of the preferred site. The parsite remains in the same site throughout the period of infection, emigrations occurring only preparatory to complete loss of the parasite from the host. The presence of cestodes in the region of the preferred site has no effect upon the distribution of the parasite. At high levels of infection the range of the parsite's distribution is extended. The feeding of larval parasites to fish by stomach tube also leads to an extension of range, whereas when fish are allowed to feed on cystacanths removed from their intermediate host the parasite establishes in a more anterior position than usual and in a site that it does not occupy to any extent in natural infections. It is concluded that the distribution of P. laevis in the alimentary canal of fish is determined primarily by the process of liberation, activation and establishment of the parasite, and that it normally attaches in the first available space and remains there. Unsuitable physico-chemical or feeding conditions affect only its growth, not the site that it occupies. It is suggested that P. laevis may be atypical in this respect.  相似文献   

2.
Gyrodactyloides bychowskii has been recorded in Scottish waters for the first time. The parasite was found on the gills of Atlantic salmon reared in seawater. An integrated morphological and molecular examination of the parasite was carried out. Prevalence of the parasite was greatest in February and declined to 0 by June in 1999. During 2000, parasites were located in March and November. An overdispersion of parasites was recorded, with intensity of infection reaching over 200 parasites per gill arch in some fish. Parasitised gill tissue showed hyperplasia and hypertrophy but not always at the site of parasite attachment. The internal transcribed spacer of the ribosomal RNA gene array was amplified by PCR and sequenced. This sequence shared greatest similarity with the internal transcribed spacer of Gyrdicotylus gallieni, followed by Gyrodactylus species. This is the first molecular analysis of this parasite and provides sequence data that may be used in comparison of G. bychowskii from other locations or in phylogenetic analysis of this group of Monogenea.  相似文献   

3.

Widespread coral mortality is leading to coral reef degradation worldwide. Many juvenile reef fishes settle on live coral, and their predator-avoidance behaviour is disrupted in seawater exposed to dead corals, ultimately increasing predation risk. Gnathiid isopods are micropredatory fish ectoparasites that occur in higher abundances in dead coral. However, the effect of seawater associated with dead coral on the susceptibility of fish to micropredators has never been investigated. We tested whether the infection rate of cultured gnathiid ectoparasites on individual damselfish, Pomacentrus amboinensis Bleeker 1868, from two different ontogenetic stages (juveniles and adults) was influenced by seawater exposed to three different treatments: dead coral, live coral, or no coral. Seawater treatments were presumed to contain different chemical properties and are meant to represent environmental changes associated with habitat degradation on coral reefs. Gnathiid infection of juvenile fish in seawater exposed to dead coral was twice as high as that of fish in live coral or no coral. Infection rates did not significantly differ between live coral and no coral treatments. In contrast to juveniles, the susceptibility of adults to gnathiids was not affected by seawater treatment. During experiments, juvenile fish mortality was relatively low, but was higher for infected fish (9.7%), compared to fish held without exposure to gnathiids (1.7%). No mortality occurred in adult fish that became infected with gnathiids. Our results suggest that chemical cues released from dead corals and/or dead coral colonisers affect the ability of juvenile, but not adult fish to avoid parasite infection. Considering increased habitat degradation on coral reefs and that gnathiids are more abundant in dead coral substrate, it is possible that wild juvenile fish may experience increased susceptibility to parasitic infection and reduced survival rate. This highlights the importance of including parasitism in ecological studies of global environmental change.

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4.
Nested species subsets are a common pattern in many types of communities found in insular or fragmented habitats. Nestedness occurs in some communities of ectoparasites of fish, as does the exact opposite departure from random assembly, anti-nestedness. Here, we looked for nested and anti-nested patterns in the species composition of communities of internal parasites of 23 fish populations from two localities in Finland. We also compared various community parameters of nested and anti-nested assemblages of parasites, and determined whether nestedness may result simply from a size-related accumulation of parasite species by feeding fish hosts. Nested parasite communities were characterised by higher prevalence (proportion of infected fish) and intensities of infection (number of parasites per fish) than anti-nested communities; the two types of non-random communities did not differ with respect to parasite species richness, however. In addition, the correlation between fish size and the number of parasite species harboured by individual fish was much stronger in nested assemblages than in anti-nested ones, where it was often nil. These results were shown not to be artefacts of sampling effort or host phylogeny. They apply to both assemblages of adult and larval parasites, which were treated separately. Since species of larval parasites are extremely unlikely to interact with one another in fish hosts, the establishment of nestedness appears independent of the potential action of interspecific interactions. The species composition of these parasite communities is not determined from within the community, but rather by the extrinsic influence of host feeding rates and how they amplify differences among parasite species in probabilities of colonisation or extinction. Nested patterns occur in parasite communities whose fish hosts accumulate parasites in a predictable fashion proportional to their size, whereas anti-nested communities occur in parasite communities whose fish hosts do not, possibly because of dietary specialisation preventing them from sampling the entire pool of parasite species available locally. Thus, nestedness in parasite communities may result from processes somewhat different from those generating nested patterns in free-living communities.  相似文献   

5.
In gregarious animals, group size correlates negatively with infection levels by some kinds of parasites and positively with infection by others. Conflicting selection pressures can be exerted simultaneously on a host species by different parasite species. Among stationary, mixed-species shoals of juvenile threespine sticklebacks, Gasterosteus aculeatus , and blackspotted sticklebacks, Gasterosteus wheatlandi , shoal size correlated differently with levels of infection by two species of ectoparasites. Stickleback shoal size correlated positively with infection levels by the copepod Thersitina gasterostei , which is transmitted among fish by short-lived planktonic larvae. In contrast, infection levels by the highly mobile crustacean parasite Argulus funduli did not decrease as shoal size increased, as predicted from an earlier laboratory experiment. The species composition of the different stickleback shoals also had an influence on some aspects of infection by these two parasite species. The contrasting mode of transmission of the two parasites results in one parasite species having a higher transmission rate among fish within large shoals, whereas the success of the other parasite species is independent of fish shoal size. The two ectoparasites may thus exert different selection pressures on stickleback shoal sizes.  相似文献   

6.
Parasites may have strong eco‐evolutionary interactions with their hosts. Consequently, they may contribute to host diversification. The radiation of cichlid fish in Lake Victoria provides a good model to study the role of parasites in the early stages of speciation. We investigated patterns of macroparasite infection in a community of 17 sympatric cichlids from a recent radiation and 2 older species from 2 nonradiating lineages, to explore the opportunity for parasite‐mediated speciation. Host species had different parasite infection profiles, which were only partially explained by ecological factors (diet, water depth). This may indicate that differences in infection are not simply the result of differences in exposure, but that hosts evolved species‐specific resistance, consistent with parasite‐mediated divergent selection. Infection was similar between sampling years, indicating that the direction of parasite‐mediated selection is stable through time. We morphologically identified 6 Cichlidogyrus species, a gill parasite that is considered a good candidate for driving parasite‐mediated speciation, because it is host species‐specific and has radiated elsewhere in Africa. Species composition of Cichlidogyrus infection was similar among the most closely related host species (members of the Lake Victoria radiation), but two more distantly related species (belonging to nonradiating sister lineages) showed distinct infection profiles. This is inconsistent with a role for Cichlidogyrus in the early stages of divergence. To conclude, we find significant interspecific variation in parasite infection profiles, which is temporally consistent. We found no evidence that Cichlidogyrus‐mediated selection contributes to the early stages of speciation. Instead, our findings indicate that species differences in infection accumulate after speciation.  相似文献   

7.
Gyrodactylus salaris is a freshwater, monogenean ectoparasite of Baltic strains of Atlantic salmon Salmo salar on which it generally causes no clinical disease. Infection of other strains of Atlantic salmon in Norway has resulted in high levels of juvenile salmon mortality and highly significant reductions in the population. The parasite is a major exotic disease threat to wild Atlantic salmon in the UK. This paper qualitatively assesses the risk of introduction and establishment of G. salaris into the UK. The current UK fish health regime prevents the importation of live salmonids from freshwater in territories that have not substantiated freedom from G. salaris. The importation of other species, e.g. eels Anguilla anguilla and non-salmonid fish, represents a low risk because the likelihood of infection is very low and the parasite can only survive on these hosts for less than 50 d. Importation of salmon carcasses presents a negligible risk because harvested fish originate from seawater sites and the parasite cannot survive full strength salinity. The importation of rainbow trout Oncorhynchus mykiss carcasses from G. salaris infected freshwater sites might introduce the parasite, but establishment is only likely if carcasses are processed on a salmonid farm in the UK. A number of mechanical transmission routes were considered (e.g. angling equipment, canoes, ballast water) and the most important was judged to be the movement of live fish transporters from farms on mainland Europe direct to UK fish farms. In the future, territories may have to substantiate freedom from G. salaris and economic drivers for live salmonid imports may strengthen. Under these circumstances, legal or illegal live salmonid imports would become the most significant risk of introduction.  相似文献   

8.
Two different genetic types of tilapia, Mozambique tilapia Oreochromis mossambicus (MT), and Pargo-UNAM (PU; a synthetic hybrid whose genetic composition is 50% Florida red tilapia, 25% Rocky Mountain tilapia, and 25% red variant Oreochromis niloticus), were acclimatized to salinity and exposed to seawater from the Gulf of Mexico off the port of Veracruz, Mexico. Both fish types were infected by the monogenean ectoparasite Neobenedenia sp. and were killed within 2 to 3 wk. A crude worm extract was prepared from whole specimens collected during the original outbreak and used to immunize naive hosts of the same 2 types of tilapia. Immunized fish were then exposed to seawater, which resulted in Neobenedenia sp. infection. Immunization did not confer any protection against Neobenedenia sp. infection. However, the experiment enabled detailed analysis of the dynamics of infection and comparison of the effects of the parasite on the 2 host types. Although both tilapia types exhibited similar resistance to infection (as they harbored similar parasite burdens in the early phase of infection), PU is less tolerant to Neobenedenia sp., as a mean parasite abundance of ca. 50 worms fish-' killed all hosts within a fortnight, while 22% of MT survived up to 3 wk, harboring a mean parasite abundance of ca. 900 worms fish-'. Our results suggest that, as reported elsewhere, Neobenedenia sp. could negatively affect mariculture off the Mexican coast of the Gulf of Mexico.  相似文献   

9.
Incidence, intensity and frequency distributions of infections in perch, Perca fluviatilis , with the eyeflukes Tylodelphys clavata and Diplostomum gasterostei were studied in a lake at monthly intervals over a period of two years. T. clavata had appeared in the lake immediately prior to the study, and its levels of infection continued to rise throughout the period of investigation whereas those of D. gasterostei remained relatively steady. The main period of infection of fish by both species of parasites was late summer, but a second and minor infection took place in spring. In the first year after its introduction T. clavata exhibited a seasonal cycle in incidence and intensity of infection and disappeared from fish in summer, but in the second year this cycle was masked by increased variation in development times and lack of synchrony throughout the parasite population. It is concluded that T. clavata has a life span of one year or less and that the parasites die within the fish, mainly in summer. D. gasterostei in contrast has a life span of at least one year, and disappearance of the parasite from the population is probably due to death of some heavily infected fish, although it was not possible to determine if this was due directly to the presence of the parasite.  相似文献   

10.
Infections of brown trout Salmo trutta L. by larvae of the nematode Eustrongylides sp. were studied over a period of two years in Fernworthy Reservoir, Devon. The parasite, here as elsewhere in Britain, is acquired only in the lake itself and not in the streams feeding it. Both native and introduced brown trout were infected but not rainbow trout which did not survive sufficiently long in the reservoir. The parasite showed a preference for female fish and for fish over 20 cm in length. Since these latter were approaching the end of their natural life span in the lake, there was insufficient time for the parasites to accumulate in older fish. No seasonal changes in infection levels or size composition of the parasite population were recorded. The larvae occurred in capsules, the majority of which were attached to the stomach wall. No local pathological effects associated with the capsules were observed and infested fish were of similar length, weight and condition to uninfected ones. The absence of harmful effects upon the trout in Fernworthy Reservoir is compared and contrasted with the effects of the parasite in other localities. It is concluded that although Eustrongylides can occasionally cause damage to some species of fish, there is no conclusive evidence that it does so to trout. but its unsightly appearance and harmful effects on birds renders it an undesirable parasite in any fishery.  相似文献   

11.
Native and exotic fishes were collected from 29 sites across coastal and inland New South Wales, Queensland and Victoria, using a range of techniques, to infer the distribution of Bothriocephalus acheilognathi (Cestoda: Pseudophyllidea) and the host species in which it occurs. The distribution of B. acheilognathi was determined by that of its principal host, carp, Cyprinus carpio; it did not occur at sites where carp were not present. The parasite was recorded from all native fish species where the sample size exceeded 30 and which were collected sympatrically with carp: Hypseleotris klunzingeri, Hypseleotris sp. 4, Hypseleotris sp. 5, Phylipnodon grandiceps and Retropinna semoni. Bothriocephalus acheilognathi was also recorded from the exotic fishes Gambusia holbrooki and Carassius auratus. Hypseleotris sp. 4, Hypseleotris sp. 5, P. grandiceps, R. semoni and C. auratus are new host records. The parasite was not recorded from any sites in coastal drainages. The only carp population examined from a coastal drainage (Albert River, south-east Queensland) was also free of infection; those fish had a parasite fauna distinct from that of carp in inland drainages and may represent a separate introduction event. Bothriocephalus acheilognathi has apparently spread along with its carp hosts and is so far restricted to the Murray-Darling Basin. The low host specificity of this parasite is cause for concern given the threatened or endangered nature of some Australian native freshwater fish species. A revised list of definitive hosts of B. acheilognathiis presented.  相似文献   

12.
The epizootiology and histopathology of the myxosporean Parvicapsula sp. was studied during monthly health surveys of 4 groups of coho salmon Oncorhynchus kisutch at a commercial farm in Puget Sound, Washington, USA, from 1984 to 1986. No Parvicapsula sp. was detected in histological samples taken from juvenile fish in fresh water, but the parasite was detected in fish from all groups 2 to 8 mo after transfer to seawater net pens. Groups placed in seawater net pens in November and December had a higher prevalence of infection, and a longer period of continuous detected infection, than those introduced into net pens in May. For the groups transferred to seawater in November and December, the highest infection prevalence (45 to 90%) was detected during the following March and April. Among 13 tissues examined histologically, only the pseudobranch and kidney were positive for Parvicapsula sp., with 26 (62%) of 42 positive fish showing infections only in the pseudobranch, 5 (12%) showing infections only in the kidney, and 11 (26%) showing infections in both organs. Both the pseudobranch and kidney were apparent primary infection sites, but pseudobranch infections appeared to persist longer in a population. Pseudobranch infections were frequently heavy and associated with extensive inflammation and necrosis of filament and lamellar tissues. The kidney had been the only infection site reported for Parvicapsula sp. in previous studies of coho salmon.  相似文献   

13.
Parasite infection patterns were compared with the occurrence of their intermediate hosts in the diet of nine sympatric fish species in a New Zealand lake. Stomach contents and infection levels of three gastrointestinal helminth species were examined from the entire fish community. The results highlighted some links between fish host diet and the flow of trophically transmitted helminths. Stomach contents indicated that all but one fish species were exposed to these helminths through their diet. Host feeding behaviour best explained infection patterns of the trematode Coitocaecum parvum among the fish community. Infection levels of the nematode Hedruris spinigera and the acanthocephalan Acanthocephalus galaxii, however, were not correlated with host diets. Host specificity is thus likely to modulate parasite infection patterns. The data indicate that host diet and host-parasite compatibility both contribute to the distribution of helminths in the fish community. Furthermore, the relative influence of encounter (trophic interactions between prey and predator hosts) and compatibility (host suitability) filters on infection levels appeared to vary between host-parasite species associations. Therefore, understanding parasite infection patterns and their potential impacts on fish communities requires determining the relative roles of encounter and compatibility filters within and across all potential host-parasite associations.  相似文献   

14.
Seasonal and ontogenetic dynamics in trophic transmission of parasites   总被引:1,自引:0,他引:1  
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.  相似文献   

15.
Fish host range of seven isolates of Cryptocaryon irritans (Ciliophora)   总被引:2,自引:0,他引:2  
Host specificity of the marine ciliate Cryptocaryon irritans was investigated using seven parasite isolates and six experimental fish species. The low degree of host specificity of C. irritans to teleost fish has been confirmed. Furthermore, five fish species occurring outside the geographical range of this parasite were also susceptible to infection; three of these could be at risk to Cryptocaryon through aquaculture practices.  相似文献   

16.
Duong  B.  Blomberg  S. P.  Cribb  T. H.  Cowman  P. F.  Kuris  A. M.  McCormick  M. I.  Warner  R. R.  Sun  D.  Grutter  A. S. 《Coral reefs (Online)》2019,38(2):199-214

The pelagic larval stage is a critical component of the life cycle of most coral reef fishes, but the adaptive significance of this stage remains controversial. One hypothesis is that migrating through the pelagic environment reduces the risk a larval fish has of being parasitised. Most organisms interact with parasites, often with significant, detrimental consequences for the hosts. However, little is known about the parasites that larval fish have upon settlement, and the factors that affect the levels of parasitism. At settlement, coral reef fishes vary greatly in size and age (pelagic larval duration), which may influence the degree of parasitism. We identified and quantified the parasites of pre-settlement larvae from 44 species of coral reef fishes from the Great Barrier Reef and explored their relationship with host size and age at settlement, and phylogeny. Overall, less than 50% of the larval fishes were infected with parasites, and over 99% of these were endoparasites. A Bayesian phylogenetic regression was used to analyse host-parasite (presence and intensity) associations. The analysis showed parasite presence was not significantly related to fish size, and parasite intensity was not significantly related to fish age. A phylogenetic signal was detected for both parasite presence and intensity, indicating that, overall, closely related fish species were likely to have more similar susceptibility to parasites and similar levels of parasitism when compared to more distantly related species. The low prevalence of infection with any parasite type and the striking rarity of ectoparasites is consistent with the ‘parasite avoidance hypothesis’, which proposes that the pelagic phase of coral reef fishes results in reduced levels of parasitism.

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17.
The helminth parasites present in 412 lesser sandeels (Ammodytes tobianus) taken from June 1996 to May 1997 from the Aran Islands on the west coast of Ireland were examined. Ten helminth parasite species were recorded, and more than 92% of the sandeels were infected with at least 1 helminth species. Seven of the species were digeneans, including Brachyphallus crenatus, Hemiurus communis, Derogenes varicus, Lecithaster gibbosus, Opechona bacillaris, Cryptocotyle lingua, and Galactosomum lacteum; 2 nematodes, including Hysterothylacium sp. and Contracaecum sp.; and 1 cestode, Scolex pleuronectis. Three of the 7 digenean species were either larvae or immature. Only 2 species, the digeneans G. lacteum and H. communis, had prevalences greater than 50%. The dominant species was G. lacteum, accounting for 67% of all parasites present. The relationship between spawned groups, host length, and season versus the abundance, prevalence, species richness, and the total number of parasites in the infracommunities was investigated. No difference was found between the parasite communities of the 2 spawning races of the host population. Mean abundance and prevalence of the different parasite species showed seasonal variation. Numbers of parasite species and numbers of parasites increased with fish length. The role of A. tobianus as an intermediate host for helminths was assessed; it was determined that most were infectious to birds or mammals, with the majority of the parasite species being autogenic (infectious to fish). The mean number of parasites per fish was nearly a quarter of the value recorded for A. tobianus in the North Sea, where a much higher intensity of infection was recorded.  相似文献   

18.
In aquatic ecosystems, fish play a key role in parasite accumulation and transmission to predacious animals. In the present study, realized on seven populations of a small cyprinid fish species, the European bitterling Rhodeus amarus, we investigated (1) the role of the European bitterling as a potential intermediate or paratenic host, (2) the ability of the fish to accumulate parasites with similar final host group, and (3) its significance as a potential source of parasite infection in the ecosystem in respect to habitat characteristics. A total of 36 parasite species were recorded; 31 species (90% of all parasite specimens) were classified as endoparasites. Most of the endoparasites were found in the larval life stage, using bitterling as an intermediate or paratenic host. In particular, parasite community structure showed significantly higher proportions of allogenic parasites in comparison with autogenic. The supposed co-occurrence of parasite species with identical final host groups showed only a weak association. The adjacent reservoir areas were a significant determinant of both the total and infracommunity parasite species richness and for the mean parasite abundance. No relationship between the distance of sampling site from the adjacent reservoir and parasite community characteristics was found. As a small-sized fish with a wide distribution range and high local abundances, the European bitterling can represent a natural prey for a wide range of piscivorous predators. Due to its susceptibility to the number of larval endoparasites, this fish species may therefore fulfill the role as important transmitter of parasites to their final hosts.  相似文献   

19.
A total of 668 specimens representing 18 species of meso- and bathypelagic fishes collected from the western North Atlantic were examined for parasites. Seventeen species and 39.1% of the specimens harboured at least one type of parasite. The highest number of parasite taxa recovered from a single fish species was seven. Host species had overall infection prevalences ranging from 10.0 to 88.9% of the specimens examined. Cestodes were most common (22.8% of all specimens examined), followed by fungi (6.6%), nematodes (6.1%) and digenetic trematodes (4.9%). Mesopelagic fishes showed a greater prevalence of infection (49.1%) than bathypelagic fishes (28.9%). Most parasites recovered were immature; however, based on the presence of adult and postlarval stage parasites, definitive and second intermediate host status is suggested for at least three fish species (Nemichthys scolopaceus, Nessorhamphus ingolfianus and Eurypharynx pelecanoides ). The presence of the cestode Nybelinia and Anisakis-lype nematodes among a number of host species may have been due to predation on the euphausiid Nematoscelis , which was found in the stomachs of all host species infected by these two parasite taxa. Higher overall infection prevalences among host species were found than have been previously reported for mid-water fishes and it is possible that this may be a function of near-bottom presence of the fishes over the continental slope. It is suggested that the low prevalence of parasites within meso- and bathypelagic fishes when compared to benthic and shallow-water species reflects the lower overall energy of, and reduced probability of host-to-host transfer in, the deep-sea pelagic ecosystem.  相似文献   

20.
Gobbin  Tiziana P.  Tiemersma  Ron  Leone  Giulia  Seehausen  Ole  Maan  Martine E. 《Hydrobiologia》2021,848(16):3817-3831

Parasite-mediated selection may initiate or enhance differentiation between host populations that are exposed to different parasite infections. Variation in infection among populations may result from differences in host ecology (thereby exposure to certain parasites) and/or intrinsic immunological traits. Species of cichlid fish, even when recently diverged, often differ in parasite infection, but the contributions of intrinsic and extrinsic causes are unknown. Here, we compare infection patterns between two closely related host species from Lake Victoria (genus Pundamilia), using wild-caught and first-generation laboratory-reared fish, as well as laboratory-reared hybrids. Three of the commonest ectoparasite species observed in the wild were also present in the laboratory populations. However, the infection differences between the host species as observed in the wild were not maintained in laboratory conditions. In addition, hybrids did not differ in infection from either parental species. These findings suggest that the observed species differences in infection in the wild might be mainly driven by ecology-related effects (i.e. differential exposure), rather than by intrinsic species differences in immunological traits. Thus, while there is scope for parasite-mediated selection in Pundamilia in the wild, it has apparently not yet generated divergent evolutionary responses and may not enhance assortative mating among closely related species.

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