Accumulation and exchange of parasites during adaptive radiation in an ancient lake

In the ancient Lake Baikal, Russia, amphipod crustaceans have undergone a spectacular adaptive radiation, resulting in a diverse community of species. A survey of microsporidian parasites inhabiting endemic and non-endemic amphipod host species at the margins of Lake Baikal indicates that the endemic amphipods harbour many microsporidian parasite groups associated with amphipods elsewhere in Eurasia. While these parasites may have undergone a degree of adaptive radiation within the lake, there is little evidence of host specificity. Furthermore, a lack of reciprocal monophyly indicates that exchanges of microsporidia between Baikalian and non-Baikalian hosts have occurred frequently in the past and may be ongoing. Conversely, limitations to parasite exchange between Baikalian and non-Baikalian host populations at the margins of the lake are implied by differences in parasite prevalence and lack of shared microsporidian haplotypes between the two host communities. While amphipod hosts have speciated sympatrically within Lake Baikal, the parasites appear instead to have accumulated, moving into the lake from external amphipod populations on multiple occasions to exploit the large and diverse community of endemic amphipods in Lake Baikal.     

Fauna of parasites of burbot <Emphasis Type="Italic">Lota lota</Emphasis> (L.) (Gadiformes: Lotidae) from reservoirs of Baikal Rift Zone

Fauna of parasites of burbot from the Baikal Rift Zone (Lake Baikal, Lake Khuvsgul, and Tsipo- Tsipikan and Kuanda-Chara lakes) includes 46 species of 11 classes. The species composition of fauna of parasites of burbot decreases in the following sequence: Lake Baikal (37 species) > lakes Tsipo-Tsipikan (18 species) > Lake Khuvsgul (15 species) > Kuanda-Chara lakes (11 species). The burbot populations of the studied lakes form two large groups according to the results of cluster analysis of species composition of the parasite fauna. The first group includes the host populations of Tsipo-Tsipikan and Kuanda-Chara lakes; the second combines the burbot populations of the gilfs and estuaries of tributaries of Lake Baikal and Lake Khuvsgul. The use of parasitological data in addition to molecular genetic information and paleontological information brings new arguments to the discussion about the evolutionary history of burbot.     

Component communities of parasites in the grayling Thymallus thymallus (L.) (Salmoniformes, Thymallidae) and the minnow Phoxinus phoxinus (L.) (Cypriniformes, Cyprinidae) from the Pechora River

Component communities of parasites of Thymallus thymallus and Phoxinus phoxinus are described. The communities differ in the number of species and in the number of dominate species, which is one in P. phoxinus and usually two in T. thymallus. In the first case the dominate species is allogenic, while in the second case dominate species are autogenic. Values of the species diversity indexes are also different for these communities. The differences between parasite communities of T. thymallus and P. phoxinus underline positions of these hosts in hydrobiocenose. Species specialists take first place in the parasite communities both of the hosts, that is characteristic for the parasite communities of the fishes from the boreal submountain faunistic complex. Parasite communities in both of the hosts consist of three groups of species discriminated by the ratio of their biomasses, that suggests forming of the groups by a coordination of the species biomasses.     

The parasite fauna of the big and little Baikal oil fishes

The parasite fauna of two endemic fish species of the genus Comephorus (Cottoidei: Comephoridae) includes 11 species. Six species were found in Comephorus baicalensis and 9--in C. dybowskii. Two species only are endemic for Baikal: Myxidium perniciosum and Gyrodactylus comephori. Other 9 species are distributed throughout Palaearctic region. Ecology of parasites and their hosts is briefly discussed.     

Nested assemblages resulting from host size variation: the case of endoparasite communities in fish hosts

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.     

南极鱼类后生动物寄生虫研究进展: 线虫、绦虫与桡足类

极端的环境造就了南极独特的生物群体, 其中鱼类是南大洋生态系统中最具多样性的脊椎动物, 也是许多寄生虫的中间或终末宿主。南极鱼类寄生虫种类丰富, 是南大洋海洋生物多样性的重要组成部分。探究南极鱼类及其寄生虫的营养关系可为阐释南极海洋生态系统功能及其变动提供重要的生态数据。虽然关于南极鱼类寄生虫的研究已有一百多年的历史, 但这些研究主要集中在寄生虫的种类鉴定、区系调查和组织病理等方面。由于南极鱼类寄生虫研究跨度时间长、地域范围广, 相关研究较为零散。文章综述了南极鱼类寄生线虫、绦虫以及桡足类的种类组成、宿主范围和地理分布等方面的研究, 并对今后开展南极鱼类寄生虫研究工作提出了展望。     

Analysis of parasitic communities in fishes from Lake Baikal

Analysis of infracommunities and component communities of fish parasites in Lake Baikal has been conducted for the first time. It has been revealed that parasite infracommunities for the majority of Baikal fishes are weakly balanced and impoverished (the Berger-Parker Index is > 0.5; Evension is < 0.5; the Brillouin Index is < 1). The highest diversity and balance of the communities are characteristic for carnivorous fishes (Brachymystax lenok, Hucho taimen, Thymallus arcticus, Esox lucius, and Percafluviatilis). The component parasitic communities of Leuciscus leuciscus baicalensis, Rutilus rutilus, and Leocottus kesslerii are the most diverse in Lake Baikal since the Shennon index for L. leuciscus baicalensis, R. rutilus, and L. kesslerii is 2.4, for Paracotlus knerii--2.2, Limnocoitus godlewskii--2.3, Phoxinus phoxinus--2.1, Lota lota and Limnocuttus pallidus--1.9, P. fluviatilis--1.8, Leuciscus idus--1.8. The component parasitic communities of other fishes in Lake Baikal have low indices of biological diversity (H = 0.5-1.05, Smp is close to 1). A classification of mature and immature components of parasitic communities based on the ratio of specialist species and generalist species has been proposed. It is established that the component parasitic communities in sublitoral, profundal, and pseudoabyssal zones are mature, while in the littoral zone they are immature (impoverished and weakly balanced). The component parasitic communities in benthophagous fishes and predators are mature, in planktivorous fishes they are immature. The component parasitic communities are mature in the family Cyprinidae and immature in the families Coregonidae and Cottidae. The component parasitic communities of the Boreal Plain and Boreal Submountain faunal complexes are mature, but they are immature in Lake Baikal and Arctic freshwater complexes.     

Seasonal dietary shifts enhance parasite transmission to lake salmonids during ice cover

Changes in abiotic and biotic factors between seasons in subarctic lake systems are often profound, potentially affecting the community structure and population dynamics of parasites over the annual cycle. However, few winter studies exist and interactions between fish hosts and their parasites are typically confined to snapshot studies restricted to the summer season whereas host‐parasite dynamics during the ice‐covered period rarely have been explored. The present study addresses seasonal patterns in the infections of intestinal parasites and their association with the diet of sympatric living Arctic charr (Salvelinus alpinus) and brown trout (Salmo trutta) in Lake Takvatn, a subarctic lake in northern Norway. In total, 354 Arctic charr and 203 brown trout were sampled from the littoral habitat between June 2017 and May 2018. Six trophically transmitted intestinal parasite taxa were identified and quantified, and their seasonal variations were contrasted with dietary information from both stomachs and intestines of the fish. The winter period proved to be an important transmission window for parasites, with increased prevalence and intensity of amphipod‐transmitted parasites in Arctic charr and parasites transmitted through fish prey in brown trout. In Arctic charr, seasonal patterns in parasite infections resulted mainly from temporal changes in diet toward amphipods, whereas host body size and the utilization of fish prey were the main drivers in brown trout. The overall dynamics in the community structure of parasites chiefly mirrored the seasonal dietary shifts of their fish hosts.     

Evolutionary implications of the adaptation to different immune systems in a parasite with a complex life cycle

Many diseases are caused by parasites with complex life cycles that involve several hosts. If parasites cope better with only one of the different types of immune systems of their host species, we might expect a trade-off in parasite performance in the different hosts, that likely influences the evolution of virulence. We tested this hypothesis in a naturally co-evolving host-parasite system consisting of the tapeworm Schistocephalus solidus and its intermediate hosts, a copepod, Macrocyclops albidus, and the three-spined stickleback Gasterosteus aculeatus. We did not find a trade-off between infection success in the two hosts. Rather, tapeworms seem to trade-off adaptation towards different parts of their hosts' immune systems. Worm sibships that performed better in the invertebrate host also seem to be able to evade detection by the fish innate defence systems, i.e. induce lower levels of activation of innate immune components. These worm variants were less harmful for the fish host likely due to reduced costs of an activated innate immune system. These findings substantiate the impact of both hosts' immune systems on parasite performance and virulence.     

Parasites as biological tags for stock discrimination in marine fish from South American Atlantic waters

The use of parasites as biological tags in population studies of marine fish in the south-western Atlantic has proved to be a successful tool for discriminating stocks for all species to which it has been applied, namely: Scomber japonicus, Engraulis anchoita, Merluccius hubbsi and Cynoscion guatucupa, the latter studied on a broader geographic scale, including samples from Uruguayan and Brazilian waters. The distribution patterns of marine parasites are determined mainly by temperature-salinity profiles and by their association with specific masses of water. Analyses of distribution patterns of some parasite species in relation to gradients in environmental (oceanographic) conditions showed that latitudinal gradients in parasite distribution are common in the study area, and are probably directly related to water temperature. Indeed, temperature, which is a good predictor of latitudinal gradients of richness and diversity of species, shows a latitudinal pattern in south-western Atlantic coasts, decreasing southwards, due to the influence of subtropical and subantarctic marine currents flowing along the edge of the continental slope. This pattern also determines the distribution of zooplankton, with a characteristic specific composition in different water masses. The gradient in the distribution of parasites determines differential compositions of their communities at different latitudes, which makes possible the identification of different stocks of their fish hosts. Other features of the host-parasite systems contributing to the success of the parasitological method are: (1) parasites identified as good biological tags (i.e. anisakids) are widely distributed in the local fauna; (2) many of these species show low specificity and use paratenic hosts; and (3) the structure of parasite communities are, to a certain degree, predictable in time and space.     

Food borne parasites as indicators of trophic segregation between Arctic charr and brown trout

The habitat and diet choice and the infection (prevalence and abundance) of trophically transmitted parasites were compared in Arctic charr and brown trout living sympatrically in two lakes in northern Norway. Arctic charr were found in all main lake habitats, whereas the brown trout were almost exclusively found in the littoral zone. In both lakes the parasite fauna reflected the niche segregation between trout and charr. Surface insects were most common in the diet of trout, but transmit few parasites, and accordingly the brown trout had a relatively low diversity and abundance of parasites. Parasites transmitted by benthic prey such as Gammarus and insect larva, were common in both salmonid host species. Copepod transmitted parasites were much more common in Arctic charr, as brown trout did not include zooplankton in their diets. Parasite species that may use small fish as transport hosts, were far more abundant in piscivorous fish, especially brown trout. The seasonal dynamics in parasite infection were also consistent with the developments in the diet throughout the year. The study demonstrates that the structure of parasite communities of charr and the trout is highly dependent on shifts in habitat and diet of their hosts both on an annual base and through the ontogeny, in addition to the observed niche segregation between the two salmonid species.     

Ecological determinants of parasite acquisition by exotic fish species

Disease‐mediated threats posed by exotic species to native counterparts are not limited to introduced parasites alone, since exotic hosts frequently acquire native parasites with possible consequences for infection patterns in native hosts. Several biological and geographical factors are thought to explain both the richness of parasites in native hosts, and the invasion success of free‐living exotic species. However, the determinants of native parasite acquisition by exotic hosts remain unknown. Here, we investigated native parasite communities of exotic freshwater fish to determine which traits influence acquisition of native parasites by exotic hosts. Model selection suggested that five factors (total body length, time since introduction, phylogenetic relatedness to the native fish fauna, trophic level and native fish species richness) may be linked to native parasite acquisition by exotic fish, but 95% confidence intervals of coefficient estimates indicated these explained little of the variance in parasite richness. Based on R²‐values, weak positive relationships may exist only between the number of parasites acquired and either host size or time since introduction. Whilst our results suggest that factors influencing parasite richness in native host communities may be less important for exotic species, it seems that analyses of general ecological factors currently fail to adequately incorporate the physiological and immunological complexity of whether a given animal species will become a host for a new parasite.     

Parasites of forage fishes in the vicinity of Steller sea lion (Eumetopias jubatus) habitat in Alaska

Fish serve as intermediate hosts for a number of larval parasites that have the potential of maturing in marine mammals such as Steller sea lions (Eumetopias jubatus). We examined the prevalence of parasites from 229 fish collected between March and July 2002 near two islands used by Steller sea lions in Southeast Alaska and island habitats in the Aleutian Islands. Sea lion populations have remained steady in Southeast Alaska but have been declining over the last 30 yr in the Aleutian Islands. Even though the fish samples near the Southeast Alaska haul-outs were composed of numerous small species of fish and the Aleutian Islands catch was dominated by juveniles of commercially harvested species, the parasite fauna was similar at all locations. Eleven of the 20 parasite taxa identified were in their larval stage in the fish hosts, several of which have been described from mammalian final hosts. Four species of parasite were more prevalent in Southeast Alaska fish samples, and seven parasite species, including several larval forms capable of infecting marine mammals, were more prevalent in fish from the Aleutian Islands. Nevertheless, parasites available to Steller sea lions from common fish prey are not likely to be a major factor in the decline of this marine mammal species.     

A review of the role of parasites in the ecology of reptiles and amphibians

A great diversity of parasites, from viruses and bacteria to a range of remarkable eukaryotic organisms, exploit reptile and amphibian hosts. Recent increases in the emergence of infectious disease have revealed the importance of understanding the effects of interactions between hosts and their parasites. Here we review the effects of parasite infection on a range of demographic, behavioural, genomic and physiological factors in reptile and amphibian species. Reviewing these parasite roles collectively, and prioritising areas for research, advances our ecological understanding and guides direction for conservation in a time of rapid species decline. Poorly resolved systems include Gymnophionan amphibians and Crocodilian hosts, in addition to viral and bacterial parasites. Future research should seek to understand processes enabling population recovery and examining synergistic interactions of parasites with fragmentation, climate change and other processes that threaten species persistence.     

Climate change: what will it do to fish–parasite interactions?

Climate change‐related factors are predicted to affect aquatic environments in many ways. Fish physiology, immunology, behaviour, and parasite‐avoidance strategies are likely to be affected by climate change and this may lead to ecosystem‐level changes. Parasitic organisms that exploit fish are also likely to be affected by climate change, both directly and via climate effects on their hosts. It is possible that climate change will alter the prerequisites for parasite transfer, for example, through changes in phenological relationships, and/or change the direction and pressure of selection in host–parasite relationships. Our review indicates strong multifactorial effects of climate change on fish–parasite systems. Increased water temperature is, on the one hand, predicted to enhance parasite metabolism, resulting in more rapid spread of parasites; on the other hand, the occurrence of some parasites could also decrease if the optimal temperature for growth and transmission is exceeded.     

Diversity,consistency, and seasonality in parasite assemblages of two sympatric marine fish Pagrus pagrus (Linnaeus, 1758) and Pagellus bogaraveo (Brünnich, 1768) (Perciformes: Sparidae) off the coast of Algeria in the western Mediterranean Sea

Various host characteristics (i. e., feeding habits, geographic distribution) and habitat characteristics (i.e., seasonality) influence the structure of parasite assemblages. To compare the parasite assemblages of hosts representatives of two genera of the same fish family, simultaneously occupying a geographic region, and to examine if seasonal variations influence parasite occurrence and abundance, we examined the parasite assemblages of two sympatric marine fish, Pagrus pagrus (n = 308) and Pagellus bogaraveo (n = 315) off the coast of Algeria in the western Mediterranean. Specimens were collected during summer and autumn over three consecutive years (2014–2016). Parasite assemblages were high in species richness and abundance. We compiled an inventory of 40 parasite taxa, including ectoparasitic monogeneans and crustaceans, and endoparasitic trematodes, cestodes, acanthocephalans, and nematodes. Endoparasite taxa primarily consisted of adult gastro-intestinal parasites and long lived larval helminths. Information on the parasite community structure and seasonal variations in parasite populations of these two hosts from the Mediterranean is here provided. Observed patterns of composition, diversity, dominance, and similarity indicate an overall consistency in assemblage structure. Although each host species harbored distinct parasite communities, they shared a high proportion of parasite species suggesting similar use of a common local pool of parasites. However, most shared species did not contribute to structuring the assemblages. Seasonal patterns in parasite abundance were observed for both hosts, with peak prevalence, abundance, and diversity in autumn. Results suggest that, regardless of a common pool of parasites being available to sympatric species, several ecological filters over time, led to distinct, independent variations in the parasite assemblages in each species.     

Body size,trophic level,and the use of fish as transmission routes by parasites

Within food webs, trophically transmitted helminth parasites use predator–prey links for their own transfer from intermediate prey hosts, in which they occur as larval or juvenile stages, to predatory definitive hosts, in which they reach maturity. In large taxa that can be used as intermediate and/or definitive hosts, such as fish, a host species’ position within a trophic network should determine whether its parasite fauna consists mostly of adult or larval helminths, since vulnerability to predation determines an animal’s role in predator–prey links. Using a large database on the helminth parasites of 303 fish species, we tested whether the proportion of parasite species in a host that occur as larval or juvenile stages is best explained by their trophic level or by their body size. Independent of fish phylogeny or habitat, only fish body length emerged as a significant predictor of the proportion of parasites in a host that occur as larval stages from our multivariate analyses. On average, the proportion of larval helminth taxa in fish shorter than 20 cm was twice as high as that for fish over 100 cm in length. This is consistent with the prediction that small fishes, being more vulnerable to predation, make better hosts for larval parasites. However, trophic level and body length are strongly correlated among fish species, and they may have separate though confounded effects on the parasite fauna exploiting a given species. Helminths show varying levels of host specificity toward their intermediate host when the latter is the downstream host involved in trophic transmission toward an upstream definitive host. Given this broad physiological compatibility of many helminths with fish hosts, our results indicate that fish body length, as a proxy for vulnerability to predators, is a better predictor of their use by helminth larvae than their trophic level based on diet content.     

Species composition and pattern of fish distribution in Siberia

At the end of the first half of the 20th century, 61 species and subspecies of fish, including one invader (bleak Alburnus alburnus) and 17 species—endemics of Baikal—were reported for Siberia (Berg, 1933, 1949). At the present time, in rivers, lakes, and reservoirs of Siberia, 96 fish species and subspecies are described, of which 80 are aboriginal and 16 are invaders; 33 species are endemics of Baikal. Each landscapegeographic zone of Siberia has its specific ichthyofauna: in rivers and lakes of the south of Siberia, species of boreal piedmont complex dominate; in water bodies of the steppe, forest-steppe, taiga, and southern parts of the forest-steppe zone boreal plain dominates; in the zone of tundra, arctic freshwater dominates; and in Baikal, the most part (35) of species belongs to the Baikal autochtonous complex. Most researchers, as before, distinguish in Siberia two ichthyogeographic sites—West Siberian and East Siberian—that enter the Arctic Province of the Circumpolar subregion of the Holarctic region. Baikal is considered in the rank of the suboblast of the Holarctic.     

Temperature and multiple parasites combine to alter host community structure

Predicting the effects of climate change requires understanding complex interactions among multiple abiotic and biotic factors. By influencing key interactions among host species, parasites can affect community and ecosystem structuring. Yet, our understanding of how multiple parasites and abiotic factors interact to alter ecosystem structure remains limited. To empirically test the role of temperature variation and parasites in shaping communities, we used a multigenerational mesocosm experiment composed of four sympatric freshwater crustacean species (isopods and amphipods) that share up to four parasite species. Mesocosms were assigned to one of four different treatments with contrasting seasonal temperatures (normal and elevated) and parasite exposure levels (continuous and arrested (presence or absence of parasite larvae in mesocosm)). We found that parasite exposure and water temperature had interactive effects on the host community. Continuous exposure to parasites altered the community structure and differences in water temperature altered species abundance. The abundance of the amphipod Paracalliope fluviatilis decreased substantially when experiencing continuous parasite exposure and elevated water temperatures. Elevated temperatures also led to parasite-induced mortality in another amphipod host, Paracorophium excavatum. Contrastingly, isopod hosts were affected much less, suggesting increasing temperatures in conjunction with higher parasite exposure might increase their relative abundance in the community. Changes in invertebrate host populations have implications for other species such as fish and birds that consume crustaceans as well as having impacts on ecosystem processes, such as aquatic primary production and nutrient cycling. In light of climate change predictions, parasite exposure and rise in average temperatures may have substantial impacts on communities and ecosystems, altering ecosystem structure and dynamics.