![]() ![]() Woodpeckers themselves are part of the family Picidae, which together with puffbirds (family: Bucconidae), Jacamars (family: Galbulidae), and a variety of toucan and barbets (infraorder: Ramphastides), form the order Piciformes ( Jarvis et al., 2014). They are contained within the clade Coraciimorphae, which includes a large assemblage of cavity nesting species such as trogons (order: Trogoniformes), hornbills and hoopoes (order: Bucerotiformes), and rollers and kingfishers (order: Coraciiformes) ( Jarvis et al., 2014). Woodpeckers are an intriguing group of birds. Woodpeckers: A Family-Wide Model for Studies of Integrative Evolution Our aim is to begin to merge our understanding of the ecological factors associated with the diversification of “beak behavior” and the physiological and mechanical factors that shape this behavior. We do this by reviewing the relatively limited literature on the topic and then developing a model for how drumming may be controlled. We then move the discussion to the physiological and biomechanical basis of woodpecker drilling and drumming. A deeper exploration of the evolution of drumming displays (a territorial signal that is produced by rapidly hitting their bill against a resonant substrate) allows us to assess how selection promotes behavioral diversity, particularly in the face of morphological constraint. We then review how these birds use their bill for important naturally selected and sexually selected behavioral traits. We start by discussing the woodpecker clade and its diversity. ![]() We do this by focusing on the evolution of woodpecker “beak behavior,” or the actions of drilling for foraging and nest creation as well as drumming for social signaling. Here, we explore the evolutionary interplay among behavioral evolution and organismal ecology and physiology. Our understanding of behavioral evolution from this mechanistic standpoint is murkier than it is from the ecological standpoint-yet, both perspectives are necessary to fully uncover the complex processes by which behavioral changes can (or cannot) occur. In these cases, complex behaviors occur through concomitant changes to the nervous and/or musculoskeletal systems that determine how individuals express behavior ( Bauwens et al., 1995 Clifton et al., 2015 Ding et al., 2016 Fuxjager et al., 2016 Barkan et al., 2018). ![]() Yet, at the same time, we must remember that behavior itself is often a manifestation of complex neurobiological and physiological processes. Such work has resulted in an extremely rich knowledge of environmental factors that create selection pressures, which in turn modify the way that individuals interact with their social and physical surroundings to better survive and reproduce. Indeed, most research within the field of behavioral ecology that addresses this objective explores the ecological factors that influence changes to a species’ behavioral program over time ( Westneat and Fox, 2010). Understanding how behavioral traits evolve is a longstanding goal of organismal biology. Our aim is to introduce the woodpecker as an ideal study system to study the physiological basis of behavioral evolution and how it relates to selection born through different ecological factors. We synthesize research related to principles of avian muscle physiology and ecology to guide inferences about the biomechanical basis of woodpecker drumming. We describe how different components of the display-namely, speed (bill strikes/beats sec –1), length (total number of beats), and rhythm-differentially evolve likely in response to sexual selection by male-male competition, whereas other components of the display appear more evolutionarily static, possibly due to morphological or physiological constraints. Individuals produce this behavior by rapidly hammering their bill on trees in their habitat, and it serves as an aggressive signal during territorial encounters. Our paper explores the interplay between ecological, physiological, and mechanical factors that shape the evolution of an elaborate display in woodpeckers called the drum. Yet organismal physiology and biomechanics also play a role in this process by defining the types of behavioral traits that are more or less likely to arise. Historically, this topic is examined from an ecological perspective, where behavioral evolution is thought to occur in response to selection pressures that arise through different social and environmental factors. Understanding how and why behavioral traits diversify during the course of evolution is a longstanding goal of organismal biologists. 2Department of Ecology, Evolution, and Organismal Biology, Brown University, Providence, RI, United States. ![]()
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