A review of the behaviours of the Chondrichthyes: a multi-species ethogram for the chimaeras, sharks, and rays

In this review of the behavioural patterns of chondrichthyan fishes, we have strived to produce a comprehensive catalogue of events and states and develop standardized terminology. Hence, actions that are slightly different, will be pooled under inclusive titles. Those used by different investigators are included in quotations within the textual descriptions of the motor patterns. This standardized ethogram will ideally lead to an increase in inter-observer reliability, giving researchers more confidence when reading colleagues’ papers that report behaviours that appear similar to theirs despite being described for different species. The descriptions are presented under the following categories: (1) maintenance (2) courtship (3) filter feeding (4) scavenging (5) predation (6) sociality (7) aggression and (8) defence. The many actions are illustrated by line drawings and photographs in composite figures with an attempt to provide an example of each action for a chimaera, shark, and ray. The diversity of patterns is evident from this ethogram, consistent with observation that the brain-to-body mass ratios of cartilaginous fishes are greater than a third of the bird species and greater than those for some mammalian species. The major impetus for assembling this ethogram is to demonstrate the diversity of behaviours exhibited by members of the Chondrichthyes and to dispel the apocryphal belief that members of this taxon are ‘simple feeding machines’.