[1] Univ Rennes 1, Biol Stn, CNRS, Lab Ecobio Ecosyst Biodivers & Evolut,UMR 6553, F-35380 Paimpont, France
[2] Univ Rennes 1, Biol Stn, CNRS, Lab EVE Ethol Evolut & Ecol,UMR 6552, F-35380 Paimpont, France
[3] Zoolog Forsch Museum Alexander Koenig, D-53113 Bonn, Germany
[4] CNRS, USM 601, Lab Mammiferes & Oiseaux, Dept Syst & Evolut,Museum Natl Hist Nat,UMR 5202, F-75005 Paris, France
[5] Inst Mar IMAR, Dept Oceanog & Pescas, P-9901862 Horta, Portugal