“…In a separate study, we were interested in how a component of this signal, rhythm, that is in part used for species recognition is also influenced by sexual selection. We find that greater SSD values positively predict whether complex cadences and acceleration patterns are present in a particular species (Miles et al, 2020). Woodpecker species with larger males are therefore more likely to produce a drum that either increases or decreases in speed (or both) at a linear (e.g., consistent slowdown in interbeat interval) or non-linear cadence (e.g., exponential increase in inter-beat interval speed).…”
Section: Evolution Of Drumming Behaviormentioning
confidence: 73%
“…Individuals, for example, might modify the drum's speed (number of beats produced over time) or length (total number of beats) (Figure 2). Likewise, individuals can adjust elements of the signal's rhythm by altering elements of cadence (description of how speed changes over time) or acceleration (description of the direction of speed change over time) (see Figure 2; Miles et al, 2018Miles et al, , 2020. Of course, individuals may also modify amplitude (volume) or dominant frequencies within the broadband spectra that define beat acoustics.…”
Section: Drum Displays: Speed Length and Rhythmmentioning
confidence: 99%
“…Green boxes illustrate how drums vary across the woodpecker phylogeny. Woodpecker drums differ in terms of speed (beats/sec), length (total number of beats), and rhythm (Miles et al, 2018(Miles et al, , 2020. The Japanese pygmy woodpecker (Yungipicus kizuki) exhibits the fastest drums.…”
Section: Drum Displays: Speed Length and Rhythmmentioning
Understanding how and why behavioral traits diversify during the course of evolution is a longstanding goal of organismal biologists. 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. 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. Our paper explores the interplay between ecological, physiological, and mechanical factors that shape the evolution of an elaborate display in woodpeckers called the drum. Individuals produce this behavior by rapidly hammering their bill on trees in their habitat, and it serves as an aggressive signal during territorial encounters. 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. We synthesize research related to principles of avian muscle physiology and ecology to guide inferences about the biomechanical basis of woodpecker drumming. 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.
“…In a separate study, we were interested in how a component of this signal, rhythm, that is in part used for species recognition is also influenced by sexual selection. We find that greater SSD values positively predict whether complex cadences and acceleration patterns are present in a particular species (Miles et al, 2020). Woodpecker species with larger males are therefore more likely to produce a drum that either increases or decreases in speed (or both) at a linear (e.g., consistent slowdown in interbeat interval) or non-linear cadence (e.g., exponential increase in inter-beat interval speed).…”
Section: Evolution Of Drumming Behaviormentioning
confidence: 73%
“…Individuals, for example, might modify the drum's speed (number of beats produced over time) or length (total number of beats) (Figure 2). Likewise, individuals can adjust elements of the signal's rhythm by altering elements of cadence (description of how speed changes over time) or acceleration (description of the direction of speed change over time) (see Figure 2; Miles et al, 2018Miles et al, , 2020. Of course, individuals may also modify amplitude (volume) or dominant frequencies within the broadband spectra that define beat acoustics.…”
Section: Drum Displays: Speed Length and Rhythmmentioning
confidence: 99%
“…Green boxes illustrate how drums vary across the woodpecker phylogeny. Woodpecker drums differ in terms of speed (beats/sec), length (total number of beats), and rhythm (Miles et al, 2018(Miles et al, , 2020. The Japanese pygmy woodpecker (Yungipicus kizuki) exhibits the fastest drums.…”
Section: Drum Displays: Speed Length and Rhythmmentioning
Understanding how and why behavioral traits diversify during the course of evolution is a longstanding goal of organismal biologists. 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. 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. Our paper explores the interplay between ecological, physiological, and mechanical factors that shape the evolution of an elaborate display in woodpeckers called the drum. Individuals produce this behavior by rapidly hammering their bill on trees in their habitat, and it serves as an aggressive signal during territorial encounters. 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. We synthesize research related to principles of avian muscle physiology and ecology to guide inferences about the biomechanical basis of woodpecker drumming. 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.
“…Yet, given their widespread presence within this clade 41 (Fig. 1b ) and some evidence of sexual selection on drumming duration and cadence 43 , 52 , 54 , 55 , we predicted (1) that signal structure has evolved to at least maintain, and potentially increase, species-specific information during the clade radiation, and (2) that drumming signals provide sufficient information to allow receivers to discriminate conspecifics from sympatric woodpecker species. Fig.…”
Section: Introductionmentioning
confidence: 97%
“…1b ). Drumming is an innate behaviour 51 , whose divergence has been relatively limited during woodpecker radiation 52 , potentially given the strong constraints inherent to its production mechanism 53 . Yet, given their widespread presence within this clade 41 (Fig.…”
Communicating species identity is a key component of many animal signals. However, whether selection for species recognition systematically increases signal diversity during clade radiation remains debated. Here we show that in woodpecker drumming, a rhythmic signal used during mating and territorial defense, the amount of species identity information encoded remained stable during woodpeckers’ radiation. Acoustic analyses and evolutionary reconstructions show interchange among six main drumming types despite strong phylogenetic contingencies, suggesting evolutionary tinkering of drumming structure within a constrained acoustic space. Playback experiments and quantification of species discriminability demonstrate sufficient signal differentiation to support species recognition in local communities. Finally, we only find character displacement in the rare cases where sympatric species are also closely related. Overall, our results illustrate how historical contingencies and ecological interactions can promote conservatism in signals during a clade radiation without impairing the effectiveness of information transfer relevant to inter-specific discrimination.
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