[1] Univ St Andrews, Ctr Biol Div, St Andrews, Fife, Scotland
[2] Univ Aveiro, Dept Biol, Aveiro, Portugal
[3] Univ Aveiro, CESAM, Aveiro, Portugal
[4] Univ Helsinki, Res Ctr Ecol Change, Organismal & Evolutionary Biol Res Programme, Helsinki, Finland