Phylogeography of North Atlantic intertidal tardigrades: refugia, cryptic speciation and the history of the Mid-Atlantic Islands

Abstract

Aim To analyse the phylogeographical history of intertidal tardigrades in the North Atlantic in order to improve our understanding of geographical differentiation in microscopic organisms, and to understand the potential importance of the Mid-Atlantic Islands as stepping stones between the American and European coasts of the Atlantic Ocean. Location Twenty-four localities from the Mid-Atlantic Islands (Greenland, Iceland and the Faroe Islands) and both sides of the North Atlantic Ocean. Methods A mitochondrial marker (cytochrome c oxidase subunit I) was sequenced from individual tardigrades belonging to the genus Echiniscoides. The existence of cryptic species was detected using generalized mixed Yule coalescence analysis; lineage ages were estimated with relaxed clock methods; and the degree of geographical differentiation was analysed with samova analyses, haplotype networks and Mantel tests. Results Echiniscoides hoepneri, previously known only from Greenland, was recovered throughout the Mid-Atlantic Islands. The Faroe Islands population was isolated from Greenland and Iceland, but overall genetic variation was low. The morphospecies Echiniscoides sigismundi had high genetic variation and consisted of at least two cryptic species. A northern and a southern species were both recovered on both sides of the Atlantic, but only the northern species was found on the Mid-Atlantic Islands. The northern species showed signs of long-term isolation between the Western and Eastern Atlantic, despite the potential of the Mid-Atlantic islands to act as stepping-stones. There was no sign of long-term isolation in the southern species. The Mid-Atlantic individuals of the northern species were of Eastern Atlantic origin, but Greenland and Iceland showed signs of long-term isolation. The genetic pattern found in the southern species is not clearly geographical, and can probably be best explained by secondary contact between former isolated populations. Main conclusions North Atlantic intertidal tardigrades from the genus Echiniscoides showed strong geographical differentiation, and the Mid-Atlantic Islands seemed unimportant as stepping stones across the Atlantic. The geographical variation of the northern species of E. sigismundi suggests post-glacial recolonization from several refugia.