Drivers of symbiont diversity in freshwater snails: a comparative analysis of resource availability, community heterogeneity, and colonization opportunities.

Decades of community ecology research have highlighted the importance of resource availability, habitat heterogeneity, and colonization opportunities in driving biodiversity. Less clear, however, is whether a similar suite of factors explains the diversity of symbionts. Here, we used a hierarchical dataset involving 12,712 freshwater snail hosts representing five species to test the relative importance of potential factors in driving symbiont richness. Specifically, we used model selection to assess the explanatory power of variables related to host species identity, resource availability (average body size, host density), ecological heterogeneity (richness of hosts and other taxa), and colonization opportunities (wetland size and amount of neighboring wetland area) on symbiont richness in 146 snail host populations in California, USA. We encountered a total of 23 taxa of symbionts, including both obligatory parasites such as digenetic trematodes as well as more commensal, mutualistic, or opportunistic groups such as aquatic insect larvae, annelids, and leeches. After validating richness estimates per host population using species accumulative curves, we detected positive effects on symbiont richness from host body size, total richness of the aquatic community, and colonization opportunities. Neither snail density nor the richness of snail species accounted for significant variation in symbiont diversity. Host species identity also affected symbiont richness, with higher gamma and average alpha diversity among more common host species with higher local abundances. These findings highlight the importance of multiple, concurrent factors in driving symbiont richness that extend beyond epidemiological measures of host abundance or host diversity alone.