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1. Tracking the assembly of nested parasite communities: Using β ‐diversity to understand variation in parasite richness and composition over time and scale

   https://doi.org/10.1111/1365-2656.13204  

   Moss, Wynne E. ; McDevitt‐Galles, Travis ; Calhoun, Dana M. ; Johnson, Pieter T. J. ; Cattadori, ed., Isabella ( March 2020 , Journal of Animal Ecology)

   Community composition is driven by a few key assembly processes: ecological selection, drift and dispersal. Nested parasite communities represent a powerful study system for understanding the relative importance of these processes and their relationship with biological scale. Quantifyingβ‐diversity across scales and over time additionally offers mechanistic insights into the ecological processes shaping the distributions of parasites and therefore infectious disease.

   To examine factors driving parasite community composition, we quantified the parasite communities of 959 amphibian hosts representing two species (the Pacific chorus frog,Pseudacris regillaand the California newt,Taricha torosa) sampled over 3 months from 10 ponds in California. Using additive partitioning, we estimated how much of regional parasite richness (γ‐diversity) was composed of within‐host parasite richness (α‐diversity) and turnover (β‐diversity) at three biological scales: across host individuals, across species and across habitat patches (ponds). We also examined howβ‐diversity varied across time at each biological scale.

   Differences among ponds comprised the majority (40%) of regional parasite diversity, followed by differences among host species (23%) and among host individuals (12%). Host species supported parasite communities that were less similar than expected by null models, consistent with ecological selection, although these differences lessened through time, likely due to high dispersal rates of infectious stages. Host individuals within the same population supported more similar parasite communities than expected, suggesting that host heterogeneity did not strongly impact parasite community composition and that dispersal was high at the individual host-level. Despite the small population sizes of within‐host parasite communities, drift appeared to play a minimal role in structuring community composition.

   Dispersal and ecological selection appear to jointly drive parasite community assembly, particularly at larger biological scales. The dispersal ability of aquatic parasites with complex life cycles differs strongly across scales, meaning that parasite communities may predictably converge at small scales where dispersal is high, but may be more stochastic and unpredictable at larger scales. Insights into assembly mechanisms within multi‐host, multi‐parasite systems provide opportunities for understanding how to mitigate the spread of infectious diseases within human and wildlife hosts.

    

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2. Contrasting drivers of diversity in hosts and parasites across the tropical Andes

   https://doi.org/10.1073/pnas.2010714118  

   McNew, Sabrina M. ; Barrow, Lisa N. ; Williamson, Jessie L. ; Galen, Spencer C. ; Skeen, Heather R. ; DuBay, Shane G. ; Gaffney, Ariel M. ; Johnson, Andrew B. ; Bautista, Emil ; Ordoñez, Paloma ; et al ( March 2021 , Proceedings of the National Academy of Sciences)

   Geographic turnover in community composition is created and maintained by eco-evolutionary forces that limit the ranges of species. One such force may be antagonistic interactions among hosts and parasites, but its general importance is unknown. Understanding the processes that underpin turnover requires distinguishing the contributions of key abiotic and biotic drivers over a range of spatial and temporal scales. Here, we address these challenges using flexible, nonlinear models to identify the factors that underlie richness (alpha diversity) and turnover (beta diversity) patterns of interacting host and parasite communities in a global biodiversity hot spot. We sampled 18 communities in the Peruvian Andes, encompassing ∼1,350 bird species and ∼400 hemosporidian parasite lineages, and spanning broad ranges of elevation, climate, primary productivity, and species richness. Turnover in both parasite and host communities was most strongly predicted by variation in precipitation, but secondary predictors differed between parasites and hosts, and between contemporary and phylogenetic timescales. Host communities shaped parasite diversity patterns, but there was little evidence for reciprocal effects. The results for parasite communities contradicted the prevailing view that biotic interactions filter communities at local scales while environmental filtering and dispersal barriers shape regional communities. Rather, subtle differences in precipitation had strong, fine-scale effects on parasite turnover while host–community effects only manifested at broad scales. We used these models to map bird and parasite turnover onto the ecological gradients of the Andean landscape, illustrating beta-diversity hot spots and their mechanistic underpinnings.

    

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3. Detecting parasite associations within multi-species host and parasite communities

   https://doi.org/10.1098/rspb.2019.1109  

   Dallas, Tad A. ; Laine, Anna-Liisa ; Ovaskainen, Otso ( October 2019 , Proceedings of the Royal Society B: Biological Sciences)

   Understanding the role of biotic interactions in shaping natural communities is a long-standing challenge in ecology. It is particularly pertinent to parasite communities sharing the same host communities and individuals, as the interactions among parasites—both competition and facilitation—may have far-reaching implications for parasite transmission and evolution. Aggregated parasite burdens may suggest that infected host individuals are either more prone to infection, or that infection by a parasite species facilitates another, leading to a positive parasite–parasite interaction. However, parasite species may also compete for host resources, leading to the prediction that parasite–parasite associations would be generally negative, especially when parasite species infect the same host tissue, competing for both resources and space. We examine the presence and strength of parasite associations using hierarchical joint species distribution models fitted to data on resident parasite communities sampled on over 1300 small mammal individuals across 22 species and their resident parasite communities. On average, we detected more positive associations between infecting parasite species than negative, with the most negative associations occurring when two parasite species infected the same host tissue, suggesting that parasite species associations may be quantifiable from observational data. Overall, our findings suggest that parasite community prediction at the level of the individual host is possible, and that parasite species associations may be detectable in complex multi-species communities, generating many hypotheses concerning the effect of host community changes on parasite community composition, parasite competition within infected hosts, and the drivers of parasite community assembly and structure. 

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4. Rodent Parasite Data for the Sevilleta National Wildlife Refuge, New Mexico (1990-1998)

   https://doi.org/10.6073/pasta/b739d2c75e142f5742b7bfe74c97d5c1  

   Duszyinski, Don ( January 2016 , Environmental Data Initiative)

   The 100,000 ha Sevilleta National Wildlife Refuge (SNWR) in central New Mexico lies in a transition zone that straddles several major biomes of the Southwest, including Great Basin Shrub-Steppe, Mogollon Pinon-Juniper Woodland, Great Plains Grassland and Chihuahuan Desert. During 9 years, (1990-1998), collaborating with the University of New Mexico's Long Term Ecological Research (LTER) program, 3,235 rodents (28 species in 4 families) were collected and identified from permanent collecting sites on the 3 major habitat types (grassland, desert/creosote, woodland) on the SNWR. Hosts were necropsied for endoparasites (protozoa [coccidia], helminths) and some ectoparasites. We identified and analyzed all the parasites found in these hosts. By 1998, we had in place the means to easily identify and moniter the parasites from all mammalian hosts caught on the LTER Phase II grant.This is not just another parasite survey; the data we collected was unique for several reasons: 1) This was the first complete inventory of a natural assemblage of parasites from all mammalian (rodent) hosts in 3 different communities, each from a distinctly defined geographic locality (habitat type) over the period of a decade, and beyond; 2) This study was part of a multidisciplinary approach to address conceptual issues of climate change on ecosystem structure and function at multiple scales (individuals, communities, etc) and correlative data from these related studies will strengthen and contribute to the robustness of this data set; 3) As the only parasite study on any of the LTER projects nationwide, it provided an ideal model, and perhaps incentive for parallel longterm studies of parasite communities to be examined in a variety of other habitat types, and from a variety of different perspectives, and other LTER sites in the network.Upon completing the work, we were able to use these long-term data to try to understand the dynamics of natural host-parasite assemblages. Hypotheses were then erected to test/address at least these questions: How do the different parasite communities colonize, mature, climax and senesce over time (or do they?), Do they vary in response to abiotic (climate change) and/or biotic (dispersal, colonization) factors? What temporal and spatial scales, and among what kinds of organisms, do coevolutionary processes influence the community organization of these parasites? Studies of the dynamics of multiple, coexisting species are confined primarily to microtine rodents and have hinted that multiannual cycles tend to be synchronous (Brown and Heske 1990). Are similar patterns seen for the parasites of our desert rodents? Answers to these questions relating to community structure, as well as to questions concerning parasite biodiversity on the SNWR, can be answered paritially or completely by the information we gathered on the parasite species infecting rodents collected on the SNWR. Initial emphasis of our work was on identifying all the parasites collected, by processing 8 consecutive years of parasite data, and on training the undergraduate and graduate students involved in the art of taxonomy and nomenclature of parasitic protozoans and helminths,  to supply some of these answers. 

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5. Linking community assembly and structure across scales in a wild mouse parasite community

   https://doi.org/10.1002/ece3.5785  

   Rynkiewicz, Evelyn C. ; Fenton, Andy ; Pedersen, Amy B. ( December 2019 , Ecology and Evolution)

   Understanding what processes drive community structure is fundamental to ecology. Many wild animals are simultaneously infected by multiple parasite species, so host–parasite communities can be valuable tools for investigating connections between community structures at multiple scales, as each host can be considered a replicate parasite community. Like free‐living communities, within‐host–parasite communities are hierarchical; ecological interactions between hosts and parasites can occur at multiple scales (e.g., host community, host population, parasite community within the host), therefore, both extrinsic and intrinsic processes can determine parasite community structure. We combine analyses of community structure and assembly at both the host population and individual scales using extensive datasets on wild wood mice (Apodemus sylvaticus) and their parasite community. An analysis of parasite community nestedness at the host population scale provided predictions about the order of infection at the individual scale, which were then tested using parasite community assembly data from individual hosts from the same populations. Nestedness analyses revealed parasite communities were significantly more structured than random. However, observed nestedness did not differ from null models in which parasite species abundance was kept constant. We did not find consistency between observed community structure at the host population scale and within‐host order of infection. Multi‐state Markov models of parasite community assembly showed that a host's likelihood of infection with one parasite did not consistently follow previous infection by a different parasite species, suggesting there is not a deterministic order of infection among the species we investigated in wild wood mice. Our results demonstrate that patterns at one scale (i.e., host population) do not reliably predict processes at another scale (i.e., individual host), and that neutral or stochastic processes may be driving the patterns of nestedness observed in these communities. We suggest that experimental approaches that manipulate parasite communities are needed to better link processes at multiple ecological scales.

    

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