[1] Univ Aberdeen, Inst Biol & Environm Sci, Aberdeen AB24 2TZ, Scotland
[2] Univ Leeds, Inst Integrat & Comparat Biol, Leeds LS2 9JT, W Yorkshire, England
[3] Univ Potsdam, D-14469 Potsdam, Germany
[4] Univ York, Dept Biol, York YO10 5YW, N Yorkshire, England