Distribution and speciation in marine intertidal tardigrades: testing the roles of climatic and geographical isolation

Aim To analyse the importance of climatic and geographical isolation in determining the patterns of speciation and distribution of species within the tardigrade genus Echiniscoides. Location Marine intertidal zone, globally. Methods DNA was extracted from 465 individual tardigrades from 48 localities world-wide. The tardigrades were divided into clusters using several distance-based criteria. The phylogeny of these clusters was estimated with Bayesian analyses. The relationships between genetic distance and substrate, climate, and geographical distance were tested with a new improved Mantel test which incorporates phylogenetic uncertainties by analysing the raw tree data instead of the averaged tree. Results Approximately 40 clusters, each probably corresponding to species, were recovered from the genetic analysis; the number of clusters fluctuated depending on the criterion used for cluster delimitation. Each cluster had a limited temperature range and all clusters were confined to single oceans under all realistic criteria for cluster delimitation. Apart from a tropical cluster, each cluster occurred only in one hemisphere. Occurrence on different substrata was not correlated with genetic distance between clusters. Both geographical distance and climate were correlated with genetic distance; however, the correlation between geographical and genetic distance disappeared when the non-independence of climatic and geographical distance were controlled for. Main conclusions The distribution of individual species of Echiniscoides is limited by climate and geographical distance. Distance does not appear to be a major factor influencing phylogeny in this genus, but ecological speciation along a temperature gradient appears to be important.