Plumage pattern function and evolution: a phylogenetic and comparative approach

Abstract

Visual patterns, such as bars and spots, are common in the animal kingdom. In no other group are patterns so exquisite in their arrangement and coloration than in birds. Although bird plumage patterns appear to be visually diverse there are only four types of patterns, which can be broadly categorized into irregular and regular patterning. That these types of irregular and regular patterning are recursive is intriguing and speaks of an underlying shared mechanism on which selection can act. The prevailing assumption is that patterns predominantly function in camouflage, however evidence suggests that they also function in communication in a small number of birds. In particular it has been suggested that barred plumage patterns could be a signal of individual quality.

In visual ecology, communication and camouflage seem to be in conflict with one another – visual signals are often conspicuous whereas camouflage has evolved to provide concealment. These ideas of pattern function need not be incongruous if patterns evolved a) for camouflage first and were subsequently co-opted by sexual selection for communication, and/or b) some patterns, specifically barred plumage, evolved for both camouflage and communication to overcome this functional compromise. To test these alternative ideas of pattern evolution I test whether a) patterns were co-opted for signaling in the model group waterfowl and gamebirds, and b) if the evolution of sexual dimorphism in barred plumage indicate camouflage and ⁄ or signaling functions across the class Aves. Additionally, I investigated whether development poses a constraint on pattern evolution in waterfowl and gamebirds.

Tracing the most probable evolutionary pathway of plumage pattern evolution revealed that the ancestral state of plumage was uniform coloration. From uniform coloration, patterns initially evolved to be predominantly monomorphic, and subsequently evolved to be sexually dimorphic. In sexually dimorphic patterns, barred plumage frequently evolved in females and males, suggesting a role for both camouflage and communication. However, dimorphic spotted plumage only evolved in males suggesting it predominantly evolved for communication. Overall, it is likely Plumage pattern function and evolution: a phylogenetic and comparative approach ii that patterns originally evolved for camouflage and were subsequently co-opted for signaling.

Focusing on the evolution of barred patterns by comparing their prevalence between the sexes I found a higher frequency of female- rather than male-biased sexual dimorphism, indicating that camouflage is its most common function. But I also found that, compared with other pigmentation patterns, barred plumage is more frequently biased towards males and its expression more frequently restricted to adulthood, suggesting that barred plumage often evolves or is maintained as a sexual communication signal. This illustrates how visual traits can accommodate the apparently incompatible functions of camouflage and communication.

Lastly, I studied the recurrence of irregular and regular plumage patterns to explore why there are different kinds of patterns that are broadly recursive. By modeling pattern evolution I derived directionality and show that where species exhibit a single pattern, selection need not be constrained by development. However, instances of irregular and regular patterns in the same species are a result of selection on existing patterns. Together this demonstrates that the evolution of patterns is not difficult and that states of multiple pattern types are a result of selection.