From molecules to macroevolution: Venom as a model system for evolutionary biology across levels of life.

Biological systems are inherently hierarchical. Consequently, any field which aims to understand an aspect of biology holistically requires investigations at each level of the hierarchy of life, and venom research is no exception. This article aims to illustrate the structure of the field in light of a 'levels of life' perspective. In doing so, I highlight how traditional fields and approaches fit into this structure as focussing on describing levels or investigating links between levels, and emphasise where implicit assumptions are made due to lack of direct information. Taking a 'levels of life' perspective to venom research enables us to understand the complementarity of different research programmes and identify avenues for future research. Moreover, it provides a broader view that, in itself, shows how new questions can be addressed. For instance, understanding how adaptations develop and function from molecular to organismal scales, and what the consequences are of those adaptations at scales from molecular to macroevolutionary, is a general question relevant to a great deal of biology. As a trait which is molecular in nature and has clearer and more direct links between genotype and phenotype than many other traits, venom provides a relatively simple system to address such questions. Furthermore, because venom is also diverse at each level of life, the complexity within the hierarchical structure provides variation that enables powerful analytical approaches to answering questions. As a result, venom provides an excellent model system for understanding big questions in evolutionary biology.