Species turnover between age groups of horses and positive network of co-occurrences define the structure of horse strongylid communities: a meta-analysis

Abstract

Grazing horses are infected by a wide range of strongylid species mostly located in the large intestine. Despite their impact on equine health and the emergence of drug resistant isolates, the phenology of these nematodes has been poorly characterized and the rules structuring their assembly as a community are not understood. Here, we compiled data on 46 equine strongylid species collected worldwide at the regional or horse scales (upon deworming or after necropsy) to analyse their richness, diversity and associated factors of variation. Worldwide, twelve species from the Cylicocyclus (n = 4), Cylicostephanus (n = 3), Coronocyclus (n = 2) and Cyathostomum (n = 2) genera were found in at least 75% of sites. Geoclimatic conditions had a limited effect on strongylid communities, but reduced species richness was found under the temperate European area. The recovery method did not affect species richness and differences on the temporal and sampling effort scales between studies applying either methods underpinned heterogeneous variances in community diversity. At the horse level, rarefaction curves correlated poorly to parasite egg excretion, suggesting little contribution of community diversity to this trait. Using a diversity partitioning approach, we found that within-host diversity represented half of overall diversity underscoring the importance of host density and environmental contamination to the diversity of strongylid communities. While this is expected to erase diversity across communities, species turnover between age classes was the second most important contributor to overall diversity (23.9%). This was associated with a network of positive co-occurrences between the four most prevalent genera that we resolved at the anatomical niche level. Altogether this pattern of {beta}-diversity maintenance across age classes combined with positive co-occurrences may be grounded by priority effects between major species. Our findings set the first assembly rules of equine strongylid communities.