Environmental, anthropogenic, and spatial factors affecting species composition and species associations in helminth communities of water frogs (Pelophylax esculentus complex) in Latvia

We investigated factors affecting species composition and patterns of species associations in parasite communities of water frogs (Pelophylax esculentus complex), applying the distance-based redundancy analysis for component communities (assemblages harboured by host populations) and Markov random fields modelling for infracommunities (assemblages harboured by individual hosts), respectively. We asked (a) What are the relative effects of variation in environmental, land use (i.e., anthropogenic), and spatial factors on the variation in the species composition of component communities (i.e., in a locality)? and (b) What is the dominant pattern of species associations in infracommunities (in a host individual), and how do these associations vary along environmental and/or anthropogenic gradients? In component communities, the greatest portion of variation in helminth species composition was explained by the combined effects of space, anthropogenic pressure, and NDVI, with the pure effect of the spatial predictor being much stronger than the effects of the remaining predictors. In infracommunities, the probability of occurrence of some, but not all, helminth species depended on the occurrence of another species, with the numbers of negative and positive co-occurrences being equal. The strength and/or sign of associations of some species pairs were spatially stable, whereas interactions between other species pairs varied along the gradient of the amount of green vegetation, from negative to positive and vice versa. We conclude that the processes in parasite infracommunities and component communities in frogs are intertwined, with both bottom-up and top-down effects acting at different hierarchical scales.