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Funder: Woolf Fisher Trust, Understanding how ecological communities are assembled remains a key goal of ecosystem ecology. Because communities are hierarchical, factors acting at multiple scales can contribute to patterns of community structure. Parasites provide a natural system to explore this idea, as they exist as discrete communities within host individuals, which are themselves part of a community and metacommunity. We aimed to understand the relative contribution of multi-scale drivers in parasite community assembly and assess how patterns at one level may mask those occurring at another. Specifically, we wanted to disentangle patterns caused by passive sampling from those determined by ecological drivers, and how these vary with scale. We applied a Markov Random Fields model and assessed measures of β-diversity and nestedness for 420 replicate parasite infracommunities (parasite assemblages in host individuals) across two freshwater mussel host species, three sites and two time periods, comparing our results to simulations from four different ecologically relevant null models. We showed that β-diversity between sites (explaining 25% of variation in parasite distribution) and host species (41%) is greater than expected, and β-diversity between individual hosts is smaller than expected, even after accounting for parasite prevalence and characteristics of host individuals. Furthermore, parasite communities were significantly less nested than expected once parasite prevalence and host characteristics were both accounted for, but more nested than expected otherwise, suggesting a degree of modularity at the within-host level that is masked if underlying host and parasite characteristics are not taken into account. The Markov Random Fields model provided evidence for possible competitive within-host parasite interactions, providing a mechanism for the observed infracommunity modularity. An integrative approach that examines factors at multiple scales is necessary to understand the composition of ecological communities. Furthermore, patterns at one level can alter the interpretation of ecologically important drivers at another if variation at higher scales is not accounted for.

Funder: Woolf Fisher Trust, Parasite conservation is important for the maintenance of ecosystem diversity and function. Conserving parasites relies first on understanding parasite biodiversity and second on estimating the extinction risk to that biodiversity. Although steps have been taken independently in both these areas, previous studies have overwhelmingly focused on helminths in vertebrate hosts over broad scales, providing low resolution and excluding a large proportion of possible host and parasite diversity. We estimated both total obligate parasite richness and parasite extinction risk in freshwater mussels (Unionidae and Margaritiferidae) from Europe and the United States to provide a case study for considering parasite conservation in a severely understudied system. We used currently reported host-parasite relationships to extrapolate parasite diversity to all possible mussel hosts and then used the threat levels of those hosts to estimate the extinction risk for both described and undescribed parasites. An estimated 67% of parasite richness in freshwater mussels is undescribed and over 80% of the most host-specific groups (digenean trematodes and ciliates) are undescribed. We estimated that 21% of this total parasite fauna is at immediate risk of extinction, corresponding to 60 unique species, many of which will likely go extinct before being described. Given the important roles parasites play in community structure and function and the strong ecosystem engineering capacities of freshwater mussels, such extinctions are likely to severely affect freshwater ecosystems. Our detailed study of mussel parasites provides compelling evidence for the hidden conservation threat to parasites through extinction cascades and shows parasites are deserving of immediate attention.