Host–parasite interactions and the evolution of nonrandom mating

Some species mate nonrandomly with respect to alleles underlying immunity. One hypothesis proposes that this is advantageous because nonrandom mating can lead to offspring with superior parasite resistance. We investigate this hypothesis, generalizing previous models in four ways: First, rather than only examining invasibility of modifiers of nonrandom mating, we identify evolutionarily stable strategies. Second, we study coevolution of both haploid and diploid hosts and parasites. Third, we allow for maternal parasite transmission. Fourth, we allow for many alleles at the interaction locus. We find that evolutionarily stable rates of assortative or disassortative mating are usually near zero or one. However, for one case, in which assumptions most closely match the major histocompatibility complex (MHC) system, intermediate rates of disassortative mating can evolve. Across all cases, with haploid hosts, evolution proceeds toward complete disassortative mating, whereas with diploid hosts either assortative or disassortative mating can evolve. Evolution of nonrandom mating is much less affected by the ploidy of parasites. For the MHC case, maternal transmission of parasites, because it creates an advantage to producing offspring that differ from their parents, leads to higher evolutionarily stable rates of disassortative mating. Lastly, with more alleles at the interaction locus, disassortative mating evolves to higher levels.     

MHC variability, life-traits and parasite diversity of European cyprinid fish

Potential links between fish life history traits, an immune investment as measured by variability of the MHC genes and parasitism were analysed in 14 species of cyprinid fish. The hypothesis of the diversity of MHC genes being driven by high parasite diversity, i.e. species richness, was tested and a potential relationship between the MHC diversity and fish life-history traits including adult mortality rate, fecundity, longevity and maturity was investigated. Molecular techniques (SSCP and sequencing) were applied to analyse the MHC nucleotide diversity of the exons 2 and 3 of DAB genes belonging to the MHC class IIB. The comparative analyses, using phylogenetic independent contrasts, revealed a negative relationship between parasite species richness and adult fish mortality rate. We also found a positive relationship between nucleotide diversity of the exon 2 and parasite species richness. Our results suggest that fish species, of which populations are exposed to high parasite pressure, in terms of high parasite species richness, maintain a high genetic diversity of the exon 2 of the MHC genes (presenting the peptide binding regions), allowing them to decrease their natural mortality rate.     

HOST POPULATION STRUCTURE AND THE EVOLUTION OF VIRULENCE: A “LAW OF DIMINISHING RETURNS”

Structure in a population of host individuals, whether spatial or temporal, can have important effects on the transmission and evolutionary dynamics of its pathogens. One of these is to limit dispersal of pathogens and thus increase the amount of contact between a given pair or within a small group of host individuals. We introduce a “law of diminishing returns” that predicts an evolutionary decline of pathogen virulence whenever there are on average more possibilities of pathogen transmission between the same pair of hosts. Thus, the effect of repeated contact between hosts will be to shift the balance of any trade-off between virulence and transmissibility toward lower virulence.     

Some ecological aspects of fish populations in tropical ricefields

Fish yields from the captural method of rice-fish farming system were monitored from 1985 to 1987. Seven species were obtained, with Trichogaster pectoralis, Clarias macrocephalus, and Channa striatus dominant and economically important. Low number of species is due to double cropping of rice resulting in increased human activities. The maximum yield obtained, 174.6 kg ha^–1, is probably near the maximum for this type of rice-fish farming and the harvests have also stabilised when compared to those obtained in the early 1970s. Fish populations included more than one year-class in each harvest. A decline in individual weight, especially of herbivores, was recorded and correlated to the abundance of the species, and hence, to food supply.     

Spatial ecology of a root parasite – from pattern to process

Abstract Occurrence patterns of parasitic plants are constrained by the distribution of suitable hosts and movement patterns of seed vectors and, accordingly, represent a simplified system to study many aspects of spatial ecology and determinants of distribution. Previous work has focused on the aerially hemiparasitic mistletoes, and it is unclear whether root parasites are affected by similar factors. Here, we evaluate spatial patterns in the root parasitic Santalum lanceolatum in an arid shrubland in north‐western New South Wales, central Australia. In this region, the principal host is a long‐lived nitrogen fixing shrub Acacia tetragonophylla closely associated with ephemeral creek‐lines. The location of 765 individuals of both species was mapped along a 250‐m section of creek‐line using a total survey station, and occurrence patterns of the root parasite related to host distribution and landscape context. We used Ripley's K‐function and the O‐ring statistic to determine whether the distribution of S. lanceolatum was random, aggregated or regular; the spatial scales at which these patterns occurred; and to quantify any spatial associations between the parasite and its host, A. tetragonophylla. While acacias were closely associated with the creek‐line, S. lanceolatum plants were more tightly clustered, displaying significant clustering at two spatial scales (1.2 m and 8.8 m). We suggest that host quality may act as an important constraint, with only those acacias growing in or near the creek‐line being physiologically capable of supporting a parasite to maturity. Insights gained from spatial analysis are used to guide ongoing research in this system, and highlight the utility of the O‐ring statistic for understanding patterns of distribution affected by multiple processes operating at critical scales.     

Conserving large populations of lions – the argument for fences has holes

Packer et al. reported that fenced lion populations attain densities closer to carrying capacity than unfenced populations. However, fenced populations are often maintained above carrying capacity, and most are small. Many more lions are conserved per dollar invested in unfenced ecosystems, which avoid the ecological and economic costs of fencing.     

A checklist of parasites of non-native populations of the fish rotan Perccottus glenii (Odontobutidae): Communication II

The fish rotan, Perccottus glenii Dybowski, 1877 (Odontobutidae) is one of the most successful fish invaders during 20th and beginning of 21st century in European and Asian fresh waters. The diversity of its parasites within its invaded range was first reviewed by Sokolov, Reshetnikov, et al. (2014). The current paper analyzes new data from 2013 to 2019 on protistan and metazoan parasites of the non-native populations of this fish. We used information from peer-reviewed journals, difficult to access grey literature and our own recently obtained original data. 45 literature sources were included in the analysis. The newly-available data originate from 95 water bodies of 13 river basins, isolated lakes in West Siberia, and Gulf of Finland, spanning a total of eight countries. A total of 132 recently detected parasite species/taxa are listed in this paper. Each species is presented with its location in/on the host, geographical localities, countries, and according literature sources. When available, number of examined fish as well as prevalence and range of intensity of infection are cited from the original source. This review enlarges contemporary knowledge on parasites of non-native populations of the studied fish species up to 167 species/taxa. We highlight perspectives for further parasitological investigations of this invasive fish.     

Spatial variation in parasite abundance: evidence of geographical population structuring in southern garfish Hyporhamphus melanochir

Southern garfish Hyporhamphus melanochir were examined for metazoan parasites from nine sites in three regions (Spencer Gulf, Gulf St Vincent and northern Kangaroo Island) in South Australia to document parasite assemblages, identify candidate species suitable for use as biological tags and investigate spatial variation in parasite abundance. Four ectoparasite and 10 endoparasite species were identified representing Cestoda, Trematoda, Monogenea, Nematoda, Acanthocephala, Copepoda and Isopoda. Lernaeenicus hemirhamphi, Micracanthorhynchina hemirhamphi, Mothocya halei and Philometra sp. were suggested for 'permanent' biological markers. Multivariate discriminant function analysis showed that most sites could be distinguished based on differences in parasite abundance. Four endoparasites (Conohelmins sp., Hysterothylacium sp., M. hemirhamphi and Philometra sp.) were most important for site characterization. Limited spatial variation in permanent endoparasite abundance among localities in northern Spencer Gulf provided evidence for a distinct northern Spencer Gulf population with little interregional mixing. In contrast, considerable spatial variation in permanent endoparasite abundance between localities sampled off Kangaroo Island implied limited local movement and suggested H. melanochir may comprise a metapopulation structure. These results largely align with recent evidence from otolith chemistry that indicates fine-scale geographical population structuring in South Australian waters.     

A review and evaluation of study design considerations for site-specifically assessing the health of fish populations

Since there are not any well developed procedures for site-specifically evaluating the health of fish populations, most field surveillance programs have been restricted to collecting information on chemistry, toxicity, bioaccumulation, biochemical alterations orin situ benthic community structure. Identification of the mechanism and significance of contaminant effects on fish populations depends on the ability to identify changes and to distinguish changes in survival, food availability or food conversion efficiency, from coincidental changes associated with alterations in habitat or natural variability. Preliminary identification of the characteristics of adult fish can be used to cost-effectively focus financial resources on the alterations which are of relevance to interpretation of impacts and identification of causal factors. Proper interpretation requires that appropriate attention be given to monitoring level and strategy, selection of species and timing of sampling, sample size requirements and choice of reference site.     

Biology of parasite populations: population dynamics of bothriocephalids (Cestoda-Pseudophyllidea) in teleostean fish

Relationships between sympatric populations of four species of Cestoda belonging to the genus Bothriocephalus ( B. barbarus. B. gregarius, B. funiculus and B. scorpii ) and populations of their respective hosts Scophthalmus rhombus, Pserra maxima, Ciliata mustela and Myoxocephalus scorpius were investigated. The parasites were overdispersed within their host populations. The mean intensities of B. burbarus were 1.59 ( s.d. =0.83) in the Mediterranean (Gulf of Lions) and 1.25 ( s.d. =0.53) in the Atlantic (coasts of Brittany). The mean intensities of B. gregarius were 62.3 ( s.d. =49.7) in the Mediterranean and 14.06 ( s.d. =18.48) in the Atlantic, compared to 1.78 ( s.d. =0.3) for B. funiculus and 21.6 ( s.d. =17.3) for B. scorpii .
The distribution of prevalence of B. gregarius corresponded to a lognormal distribution while those of B. barbatus and B. fwticulus corresponded to negative binomials. No differences in infection could be related to host sex. The parasitization of B. gregariusdepended on host size. In brill, infection by E. barbatus was independent of host size and remained stable after the fish reached 180mm in length. Parasitization of turbot by E. gregarius was lower in young fish (≤110–150 mm) and increased with the host size. Compared to the Atlantic populations of B. gregarius and B. barbatus, the Mediterranean populations were more frequently and intensively infected. The differences in the population dynamics of B. gregarius and B. barbarus would appear to depend on whether or not the life cycle involves a paratenic host.     

The importance of fish stocking in the dissemination of parasites throughout a group of reservoirs

In view of the known importance of anthropochore movements of fish in disseminating their parasites, it was hypothesized that local movements of fish between neighbouring water bodies for stocking and management purposes would similarly disseminate their parasites, resulting in a high degree of similarity in the parasite communities of fish in water bodies that have been linked by fish transfers. This hypothesis was tested by comparing the helminth parasite communities in brown trout, Salmo trutta Linnaeus and rainbow trout, Oncorhynchus mykiss (Walbaum), in reservoirs in south west England. In one group of six reservoirs there is a recent history of transfers of rainbow trout, and in the second group of four there have been no transfers over the last decade. There was no difference in mean similarity, using Jaccard's Coefficient and a Percentage Similarity Index, between the helminth faunas of trout in the reservoirs linked by transfers and those not so linked, and the distribution of several individual species of helminths was not consistent with their having been disseminated in the course of fish stockings. The hypothesis is thus rejected and it is concluded that anthropochore transfers of trout between reservoirs have been of little or no significance in the dispersal of parasites throughout the reservoirs or in the transfer of helminths from one reservoir to another. The reasons for this unexpected conclusion are discussed in the light of indications that factors unrelated to fish transfers were important determinants of helminth community composition in individual reservoirs.     

The functional traits of host fish can act as good predictors for parasite composition in a neotropical floodplain

Parasite diversity can be influenced by the interaction of environmental factors and host traits, but understanding which traits can be decisive for the establishment of the parasite may provide subsidies for a better understanding of the host-parasite relationship. In this study, we investigated whether functional traits, diet, and host phylogeny can predict the similarity of the endoparasite composition of a fish assemblage in a Brazilian floodplain. Of the three evaluated components, the host's diet was the factor that showed the greatest influence on the composition and similarity of endoparasites, demonstrating the highest value of the explanation. The functional traits and phylogeny, despite presenting significant values (unique effect and global effect), showed low explainability in the composition of the endoparasites. When analyzing the joint effects, all components showed significant influence. Hosts that live in the same environment that are phylogenetically related and have a similar ecology have a certain degree of homogeneity in their parasite assemblages and, because they are endoparasites (which are acquired trophically along the chain), diet is the main driver of parasite richness and similarity. Overall, host traits can be one of the main determinants of parasite composition, so studies that address the functional traits of the host provide a representation of local diversity and define the possible patterns of these parasite communities.     

Upstream effects of a reservoir on fish assemblages 45 years following impoundment

Fish assemblage structure, rarefied species richness, species diversity and evenness of assemblages upstream of a reservoir in Oklahoma, U.S.A., were compared pre and post‐impoundment as well as in contemporary collections from streams above and below the reservoir. There were significant shifts in assemblage structure between historical and contemporary collections above the reservoir but not between contemporary assemblages above and below the impoundment. Indicator species analysis revealed that the sand shiner Notropis stramineus and fathead minnow Pimephales promelas have declined, whereas largemouth bass Micropterus salmoides and western mosquitofish Gambusia affinis have increased in relative abundance in assemblages upstream of the impoundment. Species richness was lower in contemporary assemblages compared with historical assemblages. Furthermore, contemporary assemblages below the dam had lower species richness, diversity and evenness compared with contemporary collections above the dam. These results highlight the spatial and temporal extent of reservoirs altering fish assemblages upstream of impoundments.     

Global spread of helminth parasites at the human–domestic animal–wildlife interface

Changes in species distributions open novel parasite transmission routes at the human–wildlife interface, yet the strength of biotic and biogeographical factors that prevent or facilitate parasite host shifting are not well understood. We investigated global patterns of helminth parasite (Nematoda, Cestoda, Trematoda) sharing between mammalian wildlife species and domestic mammal hosts (including humans) using >24,000 unique country‐level records of host–parasite associations. We used hierarchical modelling and species trait data to determine possible drivers of the level of parasite sharing between wildlife species and either humans or domestic animal hosts. We found the diet of wildlife species to be a strong predictor of levels of helminth parasite sharing with humans and domestic animals, followed by a moderate effect of zoogeographical region and minor effects of species’ habitat and climatic niches. Combining model predictions with the distribution and ecological profile data of wildlife species, we projected global risk maps that uncovered strikingly similar patterns of wildlife parasite sharing across geographical areas for the different domestic host species (including humans). These similarities are largely explained by the fact that widespread parasites are commonly recorded infecting several domestic species. If the dietary profile and position in the trophic chain of a wildlife species largely drives its level of helminth parasite sharing with humans/domestic animals, future range shifts of host species that result in novel trophic interactions may likely increase parasite host shifting and have important ramifications for human and animal health.     

Population structure of a vector‐borne plant parasite

Parasites are among the most diverse groups of life on Earth, yet complex natural histories often preclude studies of their speciation processes. The biology of parasitic plants facilitates in situ collection of data on both genetic structure and the mechanisms responsible for that structure. Here, we studied the role of mating, dispersal and establishment in host race formation of a parasitic plant. We investigated the population genetics of a vector‐borne desert mistletoe (Phoradendron californicum) across two legume host tree species (Senegalia greggii and Prosopis velutina) in the Sonoran desert using microsatellites. Consistent with host race formation, we found strong host‐associated genetic structure in sympatry, little genetic variation due to geographic site and weak isolation by distance. We hypothesize that genetic differentiation results from differences in the timing of mistletoe flowering by host species, as we found initial flowering date of individual mistletoes correlated with genetic ancestry. Hybrids with intermediate ancestry were detected genetically. Individuals likely resulting from recent, successful establishment events following dispersal between the host species were detected at frequencies similar to hybrids between host races. Therefore, barriers to gene flow between the host races may have been stronger at mating than at dispersal. We also found higher inbreeding and within‐host individual relatedness values for mistletoes on the more rare and isolated host species (S. greggii). Our study spanned spatial scales to address how interactions with both vectors and hosts influence parasitic plant structure with implications for parasite virulence evolution and speciation.     

General response patterns of fish populations to stress: an evaluation using an individual-based simulation model

General response patterns of fish populations tostress, originally proposed by Colby for fisheriesrehabilitation and later adapted by Munkittrick forcontaminants, were evaluated using an individual-basedsimulation model. General response patterns relatechanges in population-level variables to the type ofstress. The model follows the daily growth,mortality, and spawning of individual yellow perch andwalleye through their lifetime, and was corroboratedusing Oneida Lake data. Two versions of the model wereused: population (yellow perch only) and community(dynamic predation on yellow perch by walleye). Eightstresses were imposed on the population and communityversions of the model and 100-year simulations wereperformed. Response patterns were defined by changesin predicted yellow perch mean population abundance,mean age of adults, and mean adult growth (representedby mean length at age-7). Proposed response patternswere similar to those predicted using the populationversion of the model. Simulations using the communityversion of the model distorted the response patterns,either causing amplification, dampening, or reversalof many of the patterns. Predicted response patternsbecame unique when additional variables were included.Our model results suggest that caution is appropriatein interpreting general response patterns based onmean age, or when the population of interest plays amajor role in a relatively simple food web. The responsepattern approach may be better at identifying the lifestage impacted rather than the mechanism of the stress.     

Reconciling genetic expectations from host specificity with historical population dynamics in an avian brood parasite, Horsfield's Bronze-Cuckoo Chalcites basalis of Australia

Mitochondrial DNA (mtDNA) is being used increasingly to explore the evolution of host specificity in avian brood parasites. A stable coevolutionary equilibrium between multiple phylogenetically unrelated hosts and a brood parasitic species predicts that mtDNA diversity in the parasite should be relatively deep and phylogenetically structured. Also, the different intraspecific clades resulting from parasitism to multiple sympatric hosts should themselves occur sympatrically. However, mtDNA diversity in brood parasites is as susceptible to effects of historical population dynamics as in any species. We demonstrate the relevance of these dynamics to the use of mtDNA in understanding coevolution between an Australian brood-parasite, Horsfield's Bronze-Cuckoo Chalcites basalis and its hosts, Malurus fairy-wrens and Acanthiza thornbills. Previous ecological and behavioural analyses argue that Malurus- and Acanthiza-specific host races exist in C. basalis. Yet mtDNA diversity in C. basalis is low and phylogenetically unstructured (mean sequence divergence 0.15 +/- 0.07%, range 0.00%-0.31%) and tests of mtDNA neutrality and range expansion vs. population stability (Tajima's D, Fu & Li's F* and D*, Fu's F(S), mismatch analyses) all indicate that C. basalis has expanded its range very recently, probably within the last few tens of thousands of years following climatic amelioration after a peak of aridity in the late Pleistocene. The low mtDNA diversity and its lack of phylogenetic structure in C. basalis deny the existence of evolutionarily long-term stable host races in C. basalis but not the possibility of recently evolved ones. They highlight the need for renewed behavioural and ecological study of the relationship between C. basalis and its hosts. Our findings illustrate the need to understand the evolutionary context in which a brood parasite and its hosts have evolved if mtDNA data are to be used in testing hypotheses concerning the origin and maintenance of host specificity. They also add to the growing body of work illustrating the use of mismatch analyses and Fu's F(S) in detecting range expansions.     

Competing for blood: the ecology of parasite resource competition in human malaria–helminth co‐infections

Ecological theory suggests that co‐infecting parasite species can interact within hosts directly, via host immunity and/or via resource competition. In mice, competition for red blood cells (RBCs) between malaria and bloodsucking helminths can regulate malaria population dynamics, but the importance of RBC competition in human hosts was unknown. We analysed infection density (i.e. the concentration of parasites in infected hosts), from a 2‐year deworming study of over 4000 human subjects. After accounting for resource‐use differences among parasites, we find evidence of resource competition, priority effects and a competitive hierarchy within co‐infected individuals. For example reducing competition via deworming increased Plasmodium vivax densities 2.8‐fold, and this effect is limited to bloodsucking hookworms. Our ecological, resource‐based perspective sheds new light into decades of conflicting outcomes of malaria–helminth co‐infection studies with significant health and transmission consequences. Beyond blood, investigating within‐human resource competition may bring new insights for improving human health.