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.Turnover in community composition across space, or “beta diversity,” reflects eco-evolutionary processes that determine range limits of species (
1–
3). These processes include adaptive specialization on particular habitats, barriers to dispersal, and interactions among species (
4–
6). Antagonistic interactions between hosts and parasites may have an underappreciated effect on turnover (
7), as evidenced by the sensitivity of host populations to novel parasites. For example, introductions of avian malaria (
Plasmodium relictum) and avian pox (
Avipoxvirus) led to extinctions or range contractions for dozens of endemic Hawaiian honeycreeper species (
8). Introduced parasites have also driven shifts in community composition when competing hosts differ in susceptibility to infection (
9). While these cases highlight extreme impacts of parasites on host communities, it remains unclear whether host–parasite interactions generally drive turnover in continental faunas, whether such effects are reciprocal or unidirectional, and whether these interactions also impact diversity patterns at regional scales or over evolutionary time.A persistent challenge in studying the factors that underlie community assembly is that turnover is dynamic and exhibits nonlinear variation over space and time (
10). As a result, different processes may underlie turnover, depending on the scale at which the community is defined (
11–
13). For instance, numerous studies have asserted that adaptive specialization on abiotic conditions and barriers to dispersal drive regional turnover patterns while biotic interactions filter communities locally (
2,
14). Still, the spatial scales of these various processes are uncertain (
11,
15,
16), and empirical tests are complicated by the fact that potential drivers of turnover tend to be spatially autocorrelated (
17).To determine the drivers and scale of community turnover in complex systems, we need appropriate, nonlinear analytical tools. Generalized dissimilarity models (GDMs) are an extension of matrix regression that provides two notable innovations: 1) GDMs can incorporate various biotic and abiotic predictors into a single model, and 2) GDMs explicitly model the curvilinear relationship between community dissimilarity and ecological or geographic distance (
4,
10,
18). This modeling framework is better suited than linear matrix regression to identifying key factors underlying turnover in complex environments (
19–
21). In addition, by incorporating phylogenetic measures of community diversity and similarity, we can use GDMs to test how drivers of turnover have varied over evolutionary time (
22). Comparing “phylogenetic turnover” to species turnover allows us to distinguish deep-time processes that may restrict the ranges of clades from contemporary processes that may constrain the range limits of individual species (
2). For example, evolutionary conservation of traits may exclude entire clades from certain habitats, leading to strong phylogenetic turnover over ecological gradients (
3). Alternatively, if traits that underpin environmental associations are evolutionarily labile, species turnover will be higher than phylogenetic turnover and better predicted by ecological variation.The tropical Andes provide an ideal natural laboratory for investigating community turnover in response to biotic and abiotic changes in the environment. Habitable elevational gradients spanning more than 5,000 vertical meters encompass rapid changes in vegetation structure, temperature, atmospheric pressure, ultraviolet (UV) exposure, and precipitation (
23,
24). The Andean cordillera generates broad orographic precipitation, but its complex topography also creates a patchwork of rain shadows. Rain-shadowed slopes and valleys fragment the ranges of humid and dry-adapted species, particularly those occurring at higher elevations (
25–
29). Environmental change across elevational gradients of the Andes is exceptionally rapid compared to change along axes parallel to the cordillera. As a result, spatial distance and environmental difference are decoupled. Pairs of communities separated by the same geographic distance may have similar or contrasting environments. In this way, this landscape provides the opportunity to pinpoint environmental effects on community turnover and distinguish them from the effects of dispersal limitation.The Andes are a global hot spot for species richness and turnover, evolutionary distinctness, and small-ranged species (
30–
33). Species interactions are thought to be particularly important in shaping Andean community turnover: For example, Andean birds are often highly specialized on particular habitats and resources (
13,
34), and competitive exclusion is thought to further limit and reinforce range boundaries (
7,
35–
37). However, parasitism has received less attention as a driver of turnover compared to competition (
35,
37) and bird–plant mutualisms (
36,
38,
39). One important group that could affect bird turnover is the hemosporidians (Apicomplexa: Haemosporida), a diverse clade of vector-borne parasites in the genera
Haemoproteus, Parahaemoproteus, Plasmodium, and
Leucocytozoon (
40,
41). These parasites can reduce the fitness of their hosts, even in low-level chronic infections (
42), and are thought to have the potential to shape avian biogeographic patterns (
40,
43). Hemosporidian communities in turn are thought to be influenced to varying degrees by host community, climate, and barriers to dispersal (
44–
51), but improved modeling frameworks with new data are needed to reciprocally test the causes of host and parasite turnover across biodiverse, tropical landscapes.In this study, we identified and compared the drivers of diversity in interacting bird and hemosporidian communities of the Peruvian Andes. First, we tested whether similar or different drivers affect host and parasite turnover; second, we tested how drivers of turnover vary with spatial scale; and third, we tested how drivers of turnover have changed over evolutionary time. Then, we used a complementary modeling approach to identify sources of variation in species richness among host and parasite communities, respectively. We used these models to map host and parasite turnover and richness to identify hot spots for faunal overlap and transition, critical zones for biodiversity study and protection.
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