[1] Wageningen Univ & Res, Resource Ecol Grp, Droevendaalsesteeg 3a, NL-6708 PB Wageningen, Netherlands
[2] Inst Marine Resources & Ecosyst Studies IMARES, NL-1976 CP Ijmuiden, Netherlands
[3] Zool Res Museum Alexander Koenig ZFMK, Adenauerallee 160, DE-53113 Bonn, Germany
[4] Univ Groningen, Dutch Montagus Harrier Fdn, Conservat Ecol Grp, Groningen Inst Evolutionary Life Sci GELIFES, POB 11103, NL-9700 CC Groningen, Netherlands