Phylogeography of the calanoid copepods Calanus helgolandicus and C-euxinus suggests Pleistocene divergences between Atlantic, Mediterranean, and Black Sea populations

Molecular systematic analyses of marine taxa are crucial for recording ocean biodiversity, so too are elucidation of the history of population divergence and the dynamics of speciation. In this paper we present the joined phylogeography of the calanoid copepod Calanus helgolandicus (Claus 1863) from the North East (NE) Atlantic and the Adriatic Sea and the closely related C. euxinus (Hulsemann 1991) from the Black Sea based on sequences of a mitochondrial Cytochrome Oxidase subunit I (COI) fragment. Coalescent-based Bayesian methods and minimum spanning networks are used to reconstruct the history of population divergence. Our results reveal that copepod populations from all three basins share a great number of haplotypes and demonstrate a close genetic affinity of C. euxinus with C. helgolandicus. The data do not support significant genetic structuring among samples within seas. Coalescent analyses suggest divergences between NE Atlantic, Mediterranean, and Black Sea populations dating back to the middle Pleistocene, with the NE Atlantic-Mediterranean divergence being the earliest and the Mediterranean-Black Sea divergence the most recent. These middle Pleistocene dates are much older than the estimated dates of colonisation of the Mediterranean and Black Seas based on paleoclimatic scenarios. Our results do not rule out that the assumed colonisations took place but they indicate that the populations colonising the Mediterranean and the Black Sea were already, and have since remained, diverged. The chaetognath Sagitta setosa, which has a comparable distribution pattern and feeds upon the copepods, provides a unique opportunity to compare phylogeographic patterns and distinguish among alternative hypotheses. The dates produced in this paper are in agreement with those estimated elsewhere for S. setosa. We propose that a great deal of the genetic make-up of marine planktonic populations comprises divergences that date back to long before the last glacial maximum. We consider questions on the taxonomic status of C. euxinus to remain open. However, its high genetic affinity to the C. helgolandicus calls for further investigation.