[1] Univ Oxford, Dept Zool, Oxford OX1 3PS, England
[2] Univ Roma La Sapienza, Ist Pasteur, Fondaziane Cenci Bolognetti, Dipartimento Genet & Biol Mol, I-00185 Rome, Italy
[3] Univ Roma La Sapienza, Consiglio Nazl Ric, Ist Biol & Patol Mol, I-00185 Rome, Italy
[4] Univ Oxford, Life Sci Interface Doctoral Training Ctr, Oxford OX1 3QD, England
[5] Univ Oxford, Dept Stat, Oxford OX1 3SY, England