Summary Animal acoustic communication is one of the most fruitful research areas in behavioural and evolutionary biology. Work in this area depends largely on quantifying the structure of acoustic signals, which has often depended upon closed‐source or graphical user interface (GUI)‐based software. Here, we describe the r package warbleR, a new package for the analysis of animal acoustic signal structure. The package offers functions for downloading avian vocalizations from the open‐access online repository Xeno‐Canto, displaying the geographic extent of the recordings, manipulating sound files, detecting acoustic signals or importing detected signals from other software, assessing performance of methods that measure acoustic similarity, conducting cross‐correlations, measuring acoustic parameters and analysing interactive vocal signals, among others. Functions working iteratively allow parallelization to improve computational efficiency. We present a case study showing how warbleR functions can be used in a workflow to evaluate the structure of acoustic signals. We analyse geographic variation in long‐billed hermit hummingbirds (Phaethornis longirostris) songs obtained from Xeno‐Canto. The code in warbleR can be executed by less experienced r users, but has also been thoroughly commented, which will facilitate further customization by advanced users. The combination of the tools described here with other acoustic analysis packages in r should significantly expand the range of analytical approaches available.
Despite longstanding interest in the evolutionary origins and maintenance of vocal learning, we know relatively little about how social dynamics influence vocal learning processes in natural populations. The “signaling group membership” hypothesis proposes that socially learned calls evolved and are maintained as signals of group membership. However, in fission–fusion societies, individuals can interact in social groups across various social scales. For learned calls to signal group membership over multiple social scales, they must contain information about group membership over each of these scales, a concept termed “hierarchical mapping.” Monk parakeets (Myiopsitta monachus), small parrots native to South America, exhibit vocal mimicry in captivity and fission–fusion social dynamics in the wild. We examined patterns of contact call acoustic similarity in Uruguay to test the hierarchical mapping assumption of the signaling group membership hypothesis. We also asked whether geographic variation patterns matched regional dialects or geographic clines that have been documented in other vocal learning species. We used visual inspection, spectrographic cross-correlation and random forests, a machine learning approach, to evaluate contact call similarity. We compared acoustic similarity across social scales and geographic distance using Mantel tests and spatial autocorrelation. We found high similarity within individuals, and low, albeit significant, similarity within groups at the pair, flock and site social scales. Patterns of acoustic similarity over geographic distance did not match mosaic or graded patterns expected in dialectal or clinal variation. Our findings suggest that monk parakeet social interactions rely more heavily upon individual recognition than group membership at higher social scales.
One way in which secondary sexual traits can influence differential reproductive success is by playing a key role in the outcome of direct physical contests for mates. Here we describe an undocumented trait in a species of hummingbird with a lek mating system, the Long-billed hermit (LBH, Phaethornis longirostris). The trait under consideration is a dagger-like structure at the bill tip, which we hypothesize is a secondary sexual trait that functions as a sexually dimorphic weapon. We tested our hypothesis by examining 5 leks during 4 consecutive years, and by employing morphological analyses, performance experiments, and behavioral observations. We found that 1) adult male bill tips were longer and pointier than their counterparts in females and juvenile males, 2) juvenile males acquired dagger-like tips during their transition to adulthood, 3) variation in bill tip morphology reflected puncture capability, and 4) males with larger and pointier bill tips were more successful in achieving lek territory tenure. Our study provides the first evidence of sexually dimorphic weapons in bird bills and stands as one of the few examples of male weaponry in birds. Our results suggest a role of sexual selection on the evolution of overall bill morphology, an alternative hypothesis to the prevailing "ecological causation" explanation for bill sexual dimorphism in hummingbirds.
Vocal learning in birds is typically restricted to a sensitive period early in life, with the few exceptions reported in songbirds and parrots. Here, we present evidence of open-ended vocal learning in a hummingbird, the third avian group with vocal learning. We studied vocalizations at four leks of the long-billed hermit Phaethornis longirostris during a four-year period. Individuals produce a single song repertoire, although several song-types can coexist at a single lek. We found that nine of 49 birds recorded on multiple days (18%) changed their song-type between consecutive recordings. Three of these birds replaced song-types twice. Moreover, the earliest estimated age when song replacement occurred ranged from 186 to 547 days (mean ¼ 307 days) and all nine birds who replaced songtypes produced a crystallized song before replacement. The findings indicate that song-type replacement is distinct from an initial early learning sensitive period. As half of lekking males do not survive past the first year of life in this species, song learning may well extend throughout the lifespan. This behaviour would be convergent to vocal learning programmes found in parrots and songbirds.
Pair collaborative behavior may play an important role in avian reproduction. However, evidence for this mainly comes from certain ecological groups (e.g. passerines). We studied the coordination of parents in foraging and its effect on food provisioning rate and chick growth in a small seabird, the Dovekie (Little auk, Alle alle). The species exhibits a dual foraging strategy, where provisioning adults make foraging trips of short (mean~2 h; to provide food for the chick) and long duration (mean~13 h; mainly for adults self-maintenance, although the food is also brought to the chick). We expected that offspring would benefit if parents coordinate their foraging patterns: one making short trips in the time when the other performing the long one. We examined this hypothesis using Monte Carlo randomization tests on field data collected during observations of individually marked birds. We found that parents did indeed adjust provisioning, making their long and short trips in an alternating pattern with respect to each other. Furthermore, we found that a higher level of coordination is associated with a lower variability in the duration of inter-feeding intervals, although this does not affect chick growth. Nevertheless, our results provide compelling evidence on the coordinated behavior of breeding partners.
Animals produce a wide array of sounds with highly variable acoustic structures. It is possible to understand the causes and consequences of this variation across taxa with phylogenetic comparative analyses. Acoustic and evolutionary analyses are rapidly increasing in sophistication such that choosing appropriate acoustic and evolutionary approaches is increasingly difficult. However, the correct choice of analysis can have profound effects on output and evolutionary inferences. Here, we identify and address some of the challenges for this growing field by providing a roadmap for quantifying and comparing sound in a phylogenetic context for researchers with a broad range of scientific backgrounds. Sound, as a continuous, multidimensional trait can be particularly challenging to measure because it can be hard to identify variables that can be compared across taxa and it is also no small feat to process and analyse the resulting high‐dimensional acoustic data using approaches that are appropriate for subsequent evolutionary analysis. Additionally, terminological inconsistencies and the role of learning in the development of acoustic traits need to be considered. Phylogenetic comparative analyses also have their own sets of caveats to consider. We provide a set of recommendations for delimiting acoustic signals into discrete, comparable acoustic units. We also present a three‐stage workflow for extracting relevant acoustic data, including options for multivariate analyses and dimensionality reduction that is compatible with phylogenetic comparative analysis. We then summarize available phylogenetic comparative approaches and how they have been used in comparative bioacoustics, and address the limitations of comparative analyses with behavioural data. Lastly, we recommend how to apply these methods to acoustic data across a range of study systems. In this way, we provide an integrated framework to aid in quantitative analysis of cross‐taxa variation in animal sounds for comparative phylogenetic analysis. In addition, we advocate the standardization of acoustic terminology across disciplines and taxa, adoption of automated methods for acoustic feature extraction, and establishment of strong data archival practices for acoustic recordings and data analyses. Combining such practices with our proposed workflow will greatly advance the reproducibility, biological interpretation, and longevity of comparative bioacoustic studies.
Although song development in songbirds has been much studied as an analogue of language development in humans, the development of vocal interaction rules has been relatively neglected in both groups. Duetting avian species provide an ideal model to address the acquisition of interaction rules as duet structure involves time and pattern-specific relationships among the vocalizations from different individuals. In this study, we address the development of the most striking properties of duets: the specific answering rules that individuals use to link their own phrase types to those of their partners (duet codes) and precise temporal coordination. By performing two removal experiments in canebrake wrens (Cantorchilus zeledoni), we show that individuals use a fixed phrase repertoire to create new phrase pairings when they acquire a new partner. Furthermore, immediately after pairing, individuals perform duets with poor coordination and poor duet code adherence, but both aspects improve with time. These results indicate that individuals need a learning period to be able to perform well-coordinated duets that follow a consistent duet code. We conclude that both duet coordination and duet code adherence are honest indicators of pair-bond duration.
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