[7] The five genera are:[1] Leiuperinae display several anti-predator mechanisms, including eyespots, hidden bright colours, macroglands in a inguinal/lumbar position, defensive behaviours, and/or chemical defence.
[8] A 2021 study showed the most recent ancestor of Edalorhina, Engystomops, Physalaemus, and Pleurodema evolved a particular type of serous gland (the main component of macroglands) in the lumbar skin along with the absence of the Eberth-Katschenko layer.
[8][9] Leiuperines with macroglands exhibit a defensive behaviour which includes four displays ('crouching down', rear elevation, body inflation, and eye protection) all of which are present in this same common ancestor.
[8] Several species independently evolved the two elements associated with aposematism (hidden bright colours and eyespots).
[8][10] Overall, the study provided phylogenetic evidence for the startle-first hypothesis,[11] i.e. behavioural displays arise as sudden movements in camouflaged individuals to avoid predatory attacks, prior to the origin of bright coloration.