Potential Parasite Transmission in Multi-Host Networks Based on Parasite Sharing

Epidemiological networks are commonly used to explore dynamics of parasite transmission among individuals in a population of a given host species. However, many parasites infect multiple host species, and thus multi-host networks may offer a better framework for investigating parasite dynamics. We investigated the factors that influence parasite sharing – and thus potential transmission pathways – among rodent hosts in Southeast Asia. We focused on differences between networks of a single host species and networks that involve multiple host species. In host-parasite networks, modularity (the extent to which the network is divided into subgroups of rodents that interact with similar parasites) was higher in the multi-species than in the single-species networks. This suggests that phylogeny affects patterns of parasite sharing, which was confirmed in analyses showing that it predicted affiliation of individuals to modules. We then constructed “potential transmission networks” based on the host-parasite networks, in which edges depict the similarity between a pair of individuals in the parasites they share. The centrality of individuals in these networks differed between multi- and single-species networks, with species identity and individual characteristics influencing their position in the networks. Simulations further revealed that parasite dynamics differed between multi- and single-species networks. We conclude that multi-host networks based on parasite sharing can provide new insights into the potential for transmission among hosts in an ecological community. In addition, the factors that determine the nature of parasite sharing (i.e. structure of the host-parasite network) may impact transmission patterns.     

Identifying hotspots of parasite diversity from species–area relationships: host phylogeny versus host ecology

Interspecific variation in parasite species richness among host species has generated much empirical research. As in comparisons among geographical areas, controlling for variation in host body size is crucial because host size determines resource availability. Recent developments in the use of species–area relationships (SARs) to detect hotspots of biodiversity provide a powerful way to control for host body size, and to identify ‘hot’ and ‘cold hosts’ of parasite diversity, i.e. hosts with more or fewer parasites than expected from their size. Applying SAR modelling to six large datasets on parasite species richness in vertebrates, we search for hot and cold hosts and assess the effect of other ecological variables on the probability that a host species is hot/cold taking body size (and sampling effort) into account. Five non‐sigmoid SAR models were fitted to the data by optimisation; their relative likelihood was evaluated using the Bayesian information criterion, before deriving an averaged SAR function. Overall, the fit between the five SAR models and the actual data was poor; there was substantial uncertainty surrounding the fitted models, and the best model differed among the six datasets. These results show that host body size is not a strong or consistent determinant of parasite species richness across taxa. Hotspots were defined as host species lying above the upper limit of the 80% confidence interval of the averaged SAR, and coldspots as species lying below its lower limit. Our analyses revealed (1) no apparent effect of specific ecological factors (i.e. water temperature, mean depth range, latitude or population density) on the likelihood of a host species being a hot or coldspot; (2) evidence of phylogenetic clustering, i.e. hosts from certain families are more likely to be hotspots (or coldspots) than other species, independently of body size. These findings suggest that host phylogeny may sometimes outweigh specific host ecological traits as a predictor of whether or not a host species harbours more (or fewer) parasite species than expected for its size.     

Compositional turnover in host and parasite communities does not change network structure

Decreasing similarity between ecological communities with increasing geographic distance (i.e. distance‐decay) is a common biogeographical observation in free‐living communities, and a slightly less common observation for parasite communities. Ecological networks of interacting species may adhere to a similar pattern of decreasing interaction similarity with increasing geographic distance, especially if species interactions are maintained across space. We extend this further, examining if host–parasite networks – independent of host and parasite species identities – become more structurally dissimilar with increasing geographic distance. Utilizing a global database of helminth parasite occurrence records, we find evidence for distance‐decay relationships in host and parasite communities at both regional and global scales, but fail to detect similar relationships in network structural similarity. Host and parasite community similarity were strongly related, and both decayed rapidly with increasing geographic distance, typically resulting in complete dissimilarity after approximately 2500 km. Our failure to detect a decay in network structural similarity suggests the possibility that different host and parasite species are filling the same functional roles in interaction networks, or that variation in network similarity may be better explained by other geographic variables or aspects of host and parasite ecology.     

Drivers of plant individual-based pollinator visitation network topology in an arid ecosystem

Interactions with pollinators underlie the structure and function of plant communities. Network analysis is a valuable tool for studying plant-pollinator interactions, but these networks are most frequently built by aggregating interactions at the species level. Interactions are between individuals and an advantage of individual-based networks is the ability to integrate inter-individual variation in traits and environmental context within complex ecological networks. We studied the influence of inter-individual variation on pollinator sharing among foundation shrubs and cactus in a desert ecosystem using plant individual-based pollinator visitation networks. We hypothesized that the traits that alter attractiveness of plants to pollinators will also influence an individual plant's role within the visitation network. Foundation plants growing with higher densities of nearby blooming shrubs had higher pollinator visitation rates and had greater access to the conspecific mating pool, suggesting widespread and diffuse pollination facilitation within this community. Further, shrub density influenced the role of betweenness centrality and the effective number of partners (e^H). Floral display size also influenced the effective number of interaction partners but did not directly influence the centrality measures for individual plants or other measures of network structure despite increasing visitation rates. The individual-based visitation networks were significantly modular and module membership was predicted by species identity and pollinator visitation rates. Ecological and individual context mediate the outcome of pollinator-mediated interactions and are fundamental drivers of whole community structure. This study shows that the density of immediate neighbours can influence the overall structure of plant-pollinator interaction networks. Exploring the contribution of intraspecific variation to community interaction networks will improve our understanding of drivers of community-level ecological dynamics.     

Host dispersal as the driver of parasite genetic structure: a paradigm lost?

Understanding traits influencing the distribution of genetic diversity has major ecological and evolutionary implications for host–parasite interactions. The genetic structure of parasites is expected to conform to that of their hosts, because host dispersal is generally assumed to drive parasite dispersal. Here, we used a meta‐analysis to test this paradigm and determine whether traits related to host dispersal correctly predict the spatial co‐distribution of host and parasite genetic variation. We compiled data from empirical work on local adaptation and host–parasite population genetic structure from a wide range of taxonomic groups. We found that genetic differentiation was significantly lower in parasites than in hosts, suggesting that dispersal may often be higher for parasites. A significant correlation in the pairwise genetic differentiation of hosts and parasites was evident, but surprisingly weak. These results were largely explained by parasite reproductive mode, the proportion of free‐living stages in the parasite life cycle and the geographical extent of the study; variables related to host dispersal were poor predictors of genetic patterns. Our results do not dispel the paradigm that parasite population genetic structure depends on host dispersal. Rather, we highlight that alternative factors are also important in driving the co‐distribution of host and parasite genetic variation.     

Patterns of interactions of a large fish-parasite network in a tropical floodplain

1.?Describing and explaining the structure of species interaction networks is of paramount importance for community ecology. Yet much has to be learned about the mechanisms responsible for major patterns, such as nestedness and modularity in different kinds of systems, of which large and diverse networks are a still underrepresented and scarcely studied fraction. 2.?We assembled information on fishes and their parasites living in a large floodplain of key ecological importance for freshwater ecosystems in the Paraná River basin in South America. The resulting fish-parasite network containing 72 and 324 species of fishes and parasites, respectively, was analysed to investigate the patterns of nestedness and modularity as related to fish and parasite features. 3.?Nestedness was found in the entire network and among endoparasites, multiple-host life cycle parasites and native hosts, but not in networks of ectoparasites, single-host life cycle parasites and non-native fishes. All networks were significantly modular. Taxonomy was the major host's attribute influencing both nestedness and modularity: more closely related host species tended to be associated with more nested parasite compositions and had greater chance of belonging to the same network module. Nevertheless, host abundance had a positive relationship with nestedness when only native host species pairs of the same network module were considered for analysis. 4.?These results highlight the importance of evolutionary history of hosts in linking patterns of nestedness and formation of modules in the network. They also show that functional attributes of parasites (i.e. parasitism mode and life cycle) and origin of host populations (i.e. natives versus non-natives) are crucial to define the relative contribution of these two network properties and their dependence on other ecological factors (e.g. host abundance), with potential implications for community dynamics and stability.     

Integrating phylogenetic and ecological distances reveals new insights into parasite host specificity

The range of hosts a pathogen infects (host specificity) is a key element of disease risk that may be influenced by both shared phylogenetic history and shared ecological attributes of prospective hosts. Phylospecificity indices quantify host specificity in terms of host relatedness, but can fail to capture ecological attributes that increase susceptibility. For instance, similarity in habitat niche may expose phylogenetically unrelated host species to similar pathogen assemblages. Using a recently proposed method that integrates multiple distances, we assess the relative contributions of host phylogenetic and functional distances to pathogen host specificity (functional–phylogenetic host specificity). We apply this index to a data set of avian malaria parasite (Plasmodium and Haemoproteus spp.) infections from Melanesian birds to show that multihost parasites generally use hosts that are closely related, not hosts with similar habitat niches. We also show that host community phylogenetic ß‐diversity (Pßd) predicts parasite Pßd and that individual host species carry phylogenetically clustered Haemoproteus parasite assemblages. Our findings were robust to phylogenetic uncertainty, and suggest that phylogenetic ancestry of both hosts and parasites plays important roles in driving avian malaria host specificity and community assembly. However, restricting host specificity analyses to either recent or historical timescales identified notable exceptions, including a ‘habitat specialist’ parasite that infects a diversity of unrelated host species with similar habitat niches. This work highlights that integrating ecological and phylogenetic distances provides a powerful approach to better understand drivers of pathogen host specificity and community assembly.     

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.     

Network analysis of host–virus communities in bats and rodents reveals determinants of cross‐species transmission

Bats are natural reservoirs of several important emerging viruses. Cross‐species transmission appears to be quite common among bats, which may contribute to their unique reservoir potential. Therefore, understanding the importance of bats as reservoirs requires examining them in a community context rather than concentrating on individual species. Here, we use a network approach to identify ecological and biological correlates of cross‐species virus transmission in bats and rodents, another important host group. We show that given our current knowledge the bat viral sharing network is more connected than the rodent network, suggesting viruses may pass more easily between bat species. We identify host traits associated with important reservoir species: gregarious bats are more likely to share more viruses and bats which migrate regionally are important for spreading viruses through the network. We identify multiple communities of viral sharing within bats and rodents and highlight potential species traits that can help guide studies of novel pathogen emergence.     

Keystone species in seed dispersal networks are mainly determined by dietary specialization

A central issue in ecology is the definition and identification of keystone species, i.e. species that are relatively more important than others for maintaining community structure and ecosystem functioning. Network theory has been pointed out as a robust theoretical framework to enhance the operationality of the keystone species concept. We used the concept of centrality as a proxy for a species’ relative importance for the structure of seed dispersal networks composed of either frugivorous bats or birds and their food‐plants. Centrality was expected to be determined mainly by dietary specialization, but also by body mass and geographic range size. Across 15 Neotropical datasets, only specialized frugivore species reached the highest values of centrality. Furthermore, the centrality of specialized frugivores varied widely within and among networks, whereas that of secondary and opportunistic frugivores was consistently low. A mixed‐effects model showed that centrality was best explained by dietary specialization, but not by body mass or range size. Furthermore, the relationship between centrality and those three ecological correlates differed between bat– and bird–fruit networks. Our findings suggest that dietary specialization is key to understand what makes a frugivore species a keystone in seed dispersal networks, and that taxonomic identity also plays a significant role. Specialized frugivores may play a central role in network structuring and ecosystem functioning, which has important implications for conservation and restoration.     

Host functional and phylogenetic composition rather than host diversity structure plant–herbivore networks

Declining plant diversity alters ecological networks, such as plant–herbivore interactions. However, our knowledge of the potential mechanisms underlying effects of plant species loss on plant–herbivore network structure is still limited. We used DNA barcoding to identify herbivore–host plant associations along declining levels of tree diversity in a large‐scale, subtropical biodiversity experiment. We tested for effects of tree species richness, host functional and phylogenetic diversity, and host functional (leaf trait) and phylogenetic composition on species, phylogenetic and network composition of herbivore communities. We found that phylogenetic host composition and related palatability/defence traits but not tree species richness significantly affected herbivore communities and interaction network complexity at both the species and community levels. Our study indicates that evolutionary dependencies and functional traits of host plants determine the composition of higher trophic levels and corresponding interaction networks in species‐rich ecosystems. Our findings highlight that characteristics of the species lost have effects on ecosystem structure and functioning across trophic levels that cannot be predicted from mere reductions in species richness.     

Could diffuse coevolution explain the generic eggshell color of the brown-headed cowbird?

The brown-headed cowbird (hereafter cowbird) is an avian brood parasite that produces an egg dissimilar to those produced by the majority of its diverse host community. The cowbird’s generic egg may result from a Jack-of-all-trades strategy; however, the evolutionary mechanisms that select for their generic eggs are unclear. Here we propose that the cowbird’s eggshell phenotypes have evolved via diffuse coevolution, which results from community-level selective pressures, rather than via pairwise coevolution that occurs between a particular host species and its brood parasite. Under diffuse coevolution the cowbird’s host community, with varying eggshell phenotypes and recognition abilities, would select for a cowbird eggshell phenotype intermediate to those of its host community. This selection is exerted by hosts that reject cowbird eggs, rather than those that accept them; therefore, we expect cowbird eggshell colors can be approximated by both the phenotypes and rejection abilities of their host community. Here we use eggshell reflectance data from 43 host species to demonstrate that the cowbird eggshell phenotypes are reasonably predicted (within 2 just noticeable differences) by the eggshell phenotypes and rejection rates of their hosts. These findings suggest that cowbird eggshell phenotypes, and potentially those of other some generalist parasites, may evolve via diffuse coevolution. Importantly, this research provides insight into the underlying evolutionary processes that explain observed phenotypic variation and provides a framework for studying selection on both specialist and generalist parasites’ traits.     

What makes a species central in a cleaning mutualism network?

Mutualisms often form networks of interacting species, characterized by the existence of a central core of species that potentially drive the ecology and the evolution of the whole community. Centrality measures allow quantification of how central or peripheral a species is within a network, thus informing about the role of each species in network organization, dynamics, and stability. In the present study we addressed the question whether the structural position of species in the network (i.e. their topological importance) relates to their ecological traits. We studied interactions between cleaner and client reef fishes to identify central and peripheral species within a mutualistic network, and investigated five ecological correlates. We used three measures to estimate the level of centrality of a species for distinct structural patterns, such as the number of interactions and the structural proximity to other species. Through the use of a principal component analysis (PCA) we observed that the centrality measures were highly correlated (92.5%) in the studied network, which indicates that the same species plays a similar role for the different structural patterns. Three cleaner and ten client species had positive values of centrality, which suggests that these species are modulating ecological and evolutionary dynamics within the network. Higher centralities were related to higher abundances and feeding habits for client fishes, but not for cleaners. The high correlation between centrality measures in the present study is likely related to the nested structure of the cleaning network. The cleaner species’ set, by having central species that are not necessarily the most abundant ones, bears potentially more vulnerable points for network cohesiveness. Additionally, the present study generalizes previous findings for plant–animal mutualisms, as it shows that the structure of marine mutualisms is also related to a complex interplay between abundance and niche‐related features.     

Parasite species richness in carnivores: effects of host body mass, latitude, geographical range and population density

Aim  Comparative studies have revealed strong links between ecological factors and the number of parasite species harboured by different hosts, but studies of different taxonomic host groups have produced inconsistent results. As a step towards understanding the general patterns of parasite species richness, we present results from a new comprehensive data base of over 7000 host–parasite combinations representing 146 species of carnivores (Mammalia: Carnivora) and 980 species of parasites.
Methods  We used both phylogenetic and non-phylogenetic comparative methods while controlling for unequal sampling effort within a multivariate framework to ascertain the main determinants of parasite species richness in carnivores.
Results  We found that body mass, population density, geographical range size and distance from the equator are correlated with overall parasite species richness in fissiped carnivores. When parasites are classified by transmission mode, body mass and home range area are the main determinants of the richness of parasites spread by close contact between hosts, and population density, geographical range size and distance from the equator account for the diversity of parasites that are not dependent on close contact. For generalist parasites, population density, geographical range size and latitude are the primary predictors of parasite species richness. We found no significant ecological correlates for the richness of specialist or vector-borne parasites.
Main conclusions  Although we found that parasite species richness increases instead of decreases with distance from the equator, other comparative patterns in carnivores support previous findings in primates, suggesting that similar ecological factors operate in both these independent evolutionary lineages.     

Competition and host size mediate larval anuran interactions with trematode parasites

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Predicting the similarity of parasite communities in freshwater fishes using the phylogeny,ecology and proximity of hosts

Parasite communities tend to be dissimilar in hosts that are geographically, phylogenetically, ecologically and developmentally distant from one another. The decay of community similarity is a powerful and increasingly common method of studying parasite beta diversity, but most studies have examined only a single type of distance. Here, we evaluate distances based on the phylogeny, ecology, spatial proximity and size of hosts, as predictors of the similarity of parasite communities in individual hosts, host populations and host species. We surveyed parasites in six species of fish collected simultaneously from six localities in the St. Lawrence River, Canada, and species in a common group of larval parasites were discriminated using DNA sequences from barcode region of cytochrome c oxidase I. Distances based on the habitat use patterns of host species were good predictors of short‐term, ecological similarity of parasite communities, such as that operating at the scale of the individual host. The genetic distance between host species was associated with almost all types of similarity at all scales, particularly qualitative and phylogenetic similarity of parasite communities at the level of populations and meta‐populations of hosts. The trophic level, diet, spatial proximity and size of hosts were poor predictors of parasite community similarity. The increased taxonomic resolution provided by molecular data increased the explanatory power of regression models, and different factors were implicated when parasite species were distinguished with DNA barcodes than when larval parasites were lumped into morphospecies, as is commonly practiced.     

Global patterns in helminth host specificity: phylogenetic and functional diversity of regional host species pools matter

Host specificity has a major influence on a parasite's ability to shift between human and animal host species. Yet there is a dearth of quantitative approaches to explore variation in host specificity across biogeographical scales, particularly in response to the varying community compositions of potential hosts. We built a global dataset of intermediate host associations for nine of the world's most widespread helminth parasites (all of which infect humans). Using hierarchical models, we asked if realised parasite host specificity varied in response to regional variation in the phylogenetic and functional diversities of potential host species. Parasites were recorded in 4–10 zoogeographical regions, with some showing considerable geographical variation in observed versus expected host specificity. Parasites generally exhibited the lowest phylogenetic host specificity in regions with the greatest variation in prospective host phylogenetic diversity, namely the Neotropical, Saharo‐Arabian and Australian regions. Globally, we uncovered notable variation in parasite host shifting potential. Observed host assemblages for Hydatigera taeniaeformis and Hymenolepis diminuta were less phylogenetically diverse than expected, suggesting limited potential to spillover into unrelated hosts. Host assemblages for Echinococcus granulosus, Mesocestoides lineatus and Trichinella spiralis were less functionally diverse than expected, suggesting limited potential to shift across host ecological niches. By contrast, Hyd. taeniaeformis infected a higher functional diversity of hosts than expected, indicating strong potential to shift across hosts with different ecological niches. We show that the realised phylogenetic and functional diversities of infected hosts are determined by biogeographical gradients in prospective host species pools. These findings emphasise the need to account for underlying species diversity when assessing parasite host specificity. Our framework to identify variation in realised host specificity is broadly applicable to other host–parasite systems and will provide key insights into parasite invasion potential at regional and global scales.     

Impact of a social parasite on ant host populations depends on host species,habitat and year

Parasites often affect the abundance and life‐history traits of their hosts. We studied the impact of a social parasite – a slavemaking ant – on host ant communities using two complementary field manipulations. In the first experiment, we analysed the effect of social parasite presence on host populations in one habitat. In a second experiment, conducted in two habitats, we used a cross‐fostering design, analysing the effect of sympatric and allopatric social parasites. In the first experiment, host colonies benefited to some extent from residing in parasite‐free areas, showing increased total production. Yet, in the second experiment, host colonies in plots containing social parasites were more productive, and this effect was most evident in response to allopatric social parasites. We propose several explanations for these inconsistent results, which are related to environmental variability. The discrepancies between the two habitats can be explained well by ecological variation as a result of differences in altitudes and climate. For example, ant colonies in the colder habitat were larger and, for one host species, colonies were more often polygynous. In addition, our long‐term documentation – a total of four measurements of community structure in 6 years – showed temporal variation in abundance and life‐history traits of ant colonies, unrelated to the manipulations. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 559–570.     

Functional and phylogenetic uniqueness of helminth and flea assemblages of two South African rodents

The loss of a particular species from a community may have different effects on its functioning, depending on the presence or absence of functionally similar or phylogenetically close species in that community (redundancy). Redundancy is thus defined as the fraction of species diversity not expressed by functional or phylogenetic diversity. We assessed functional and phylogenetic alpha- and beta-redundancy in helminth and flea assemblages of two species of South African rodents, Rhabdomys dilectus and Rhabdomys pumilio, using community uniqueness as the inverse indicator of redundancy. We asked whether patterns of functional and phylogenetic alpha- and beta-uniqueness differed between (i) parasite groups (endo- versus ectoparasites), (ii) host species within parasite groups, and (iii) biomes within host species. We found differences between the two hosts in the functional and phylogenetic alpha-uniqueness (but not beta-uniqueness) of flea, but not helminth, assemblages. Significant correlations between the alpha-uniqueness of parasite assemblages and the total parasite prevalence were found only for phylogenetic uniqueness and only in helminths. Pairwise site-by-site dissimilarities in uniqueness (beta-uniqueness) and pairwise dissimilarity in prevalence were significantly associated (positively) in helminths but not in fleas. A between-biome difference in functional (but not phylogenetic) alpha-uniqueness was found in both helminth and flea assemblages harboured by R. pumilio. We conclude that the resilience of parasite assemblages in terms of the effect on hosts depends not only on their transmission strategy but also on traits of host species and environmental factors.