1. Department of Biology, University of York , York, North Yorkshire, YO10 5DD, UK
2. Oxford University Museum of Natural History , Oxford, Oxfordshire, OX1 3PW, UK
3. Department of Biology and Biochemistry, University of Bath , Bath and North East Somerset, BA2 7AX, UK
4. Leverhulme Centre for Anthropocene Biodiversity, University of York , York, North Yorkshire, YO10 5DD, UK