These studies serve as a good example where very different techniques and lines of evidence support a hypothesis regarding the behaviour of an extinct organism. We strongly advocate that such holistic and integrative approaches should be brought to bear on the interpretation of behaviour from the fossil record wherever possible (see discussion below). Even with these techniques and sources of data, work on the behaviour of animals from the fossil record has understandably lagged that of traditional ethology. It is clearly difficult to attempt to reconstruct the behaviour of

an extinct animal (Benton, 2010). Many details cannot possibly be reconstructed as no trace of them could be preserved or determined with any degree of accuracy. For example the exact nature of a courtship ritual, whether or not monoagamous Daporinad mouse species indulged in extra pair Trametinib datasheet copulations (which even today can only be determined unambiguously through genetic analysis of parentage), or whether or not a species was territorial will likely never be determined for any truly ancient species. However, many fundamental issues can potentially be inferred such as whether or not a species was social or asocial, or was herbivorous or carnivorous, though

not always without controversy. New and innovative research has produced new insights into the behaviour of fossils animals (e.g. Finite-element analysis, Rayfield et al., 2007 or computed tomography work to reveal the brain structure of birds, Walsh et al., 2009), and this has been supported by more traditional functional analyses (e.g. Snively & Russell, 2001) and the discovery of exceptional fossils that show evidence of behaviour [e.g. dinosaurs that show unequivocal evidence for MCE公司 brooding

on a nest (Norell et al., 1995) or scavenging (Hone & Watabe, 2010)]. Unfortunately, despite this new wealth of data, many hypotheses have been proposed to explain the behaviour of fossil animals (from individual specimens through to entire clades) that are based on little evidence, equivocal data or that rely on huge extrapolation, or misunderstandings of, behaviour. The incredible adaptability of animals to their environments and the complexities of interactions between and within species (Faulkes & Bennett, 2013; Kappeler et al., 2013), combined with the limited evidence from the fossil record perhaps makes this inevitable (Fig. 1). There are not even that many good correlations between many behaviours or types of behaviour and the kinds of anatomical features that readily preserve in the fossil record (or ichnological traces that can be correctly tied to a given species). Here we document some of the pitfalls that have beset previous statements about the behaviour of extinct animals. We attempt to layout a framework for the future establishment of viable hypotheses and how evidence could be accumulated for these.