A global analysis of song frequency in passerines provides no support for the acoustic adaptation hypothesis but suggests a role for sexual selection.

Animals use acoustic signals for communication, implying that the properties of these signals can be under strong selection. The acoustic adaptation hypothesis predicts that species in dense habitats emit lower-frequency sounds than those in open areas because low-frequency sounds propagate further in dense vegetation than high-frequency sounds. Signal frequency may also be under sexual selection because it correlates with body size and lower-frequency sounds are perceived as more intimidating. Here, we evaluate these hypotheses by analysing variation in peak song frequency across 5,085 passerine species (Passeriformes). A phylogenetically informed analysis revealed that song frequency decreases with increasing body mass and with male-biased sexual size dimorphism. However, we found no support for the predicted relationship between frequency and habitat. Our results suggest that the global variation in passerine song frequency is mostly driven by natural and sexual selection causing evolutionary shifts in body size rather than by habitat-related selection on sound propagation.

Automatic acoustic identification of individuals in multiple species: improving identification across recording conditions.

Many animals emit vocal sounds which, independently from the sounds' function, contain some individually distinctive signature. Thus the automatic recognition of individuals by sound is a potentially powerful tool for zoology and ecology research and practical monitoring. Here, we present a general automatic identification method that can work across multiple animal species with various levels of complexity in their communication systems. We further introduce new analysis techniques based on dataset manipulations that can evaluate the robustness and generality of a classifier. By using these techniques, we confirmed the presence of experimental confounds in situations resembling those from past studies. We introduce data manipulations that can reduce the impact of these confounds, compatible with any classifier. We suggest that assessment of confounds should become a standard part of future studies to ensure they do not report over-optimistic results. We provide annotated recordings used for analyses along with this study and we call for dataset sharing to be a common practice to enhance the development of methods and comparisons of results.

Song complexity-no correlation between standard deviation of frequency and traditionally used song complexity metrics in passerines: A comment on Pearse et al. (2018).

Recently, Pearse et al. explored the macroecology of passerine song using a large citizen science database of bird songs and machine learning techniques. They used standard deviation of frequency (SDF) as a surrogate for song complexity, finding only weak support for correlation between SDF and life-history traits such as monogamy and sexual dimorphism. Their finding that song complexity increases toward more productive environments and warmer areas seemingly contradicts several previous multitaxonomic studies. By comparing SDF scores with traditionally used song complexity metrics (syllable repertoire size and the number of syllable types per song), we found no evidence of any correlation. This may help to explain the discrepancy between their findings and findings of previous studies. While we agree that simple metrics that can be quantified and compared between multiple, highly variable species are crucial for progress in large-scale analysis of birdsong complexity, the biological relevance of SDF remains unclear and more research is needed to clarify its relevance for further studies of birdsong complexity.

Being angry, singing fast? Signalling of aggressive motivation by syllable rate in a songbird with slow song.

Syllable rate has been shown to play a role in male-male aggressive interactions and has been proposed to serve as a male quality indicator in several bird species. In those with fast syllable rates, males often increase rates when singing in aggressive context, and respond differently to test stimuli of varying rates. We asked whether the syllable rate fulfils a similar signalling function in the chiffchaff (Phylloscopus collybita), a songbird species with a slow syllable rate. We confronted 36 chiffchaff males with one of three playback types differing in syllable rate: control (non-manipulated rate), fast, or slow (artificially increased and decreased syllable rate, respectively). We recorded tested males' songs and behaviour before and during the experiment. Our results indicate that syllable rate might be an aggressive signal in chiffchaff. Males that physically attacked the loudspeaker during experiments sang faster songs spontaneously, and those that continued singing during the playback responded to fast and non-manipulated stimuli with substantial increase of syllable rate. Indirect evidence further suggests that syllable rate in chiffchaff is unlikely constrained by respiratory demands; thus, we propose that syllable rate in this species functions as a conventional signal of male aggressiveness rather than an index of quality.

The causes and evolutionary consequences of mixed singing in two hybridizing songbird species (Luscinia spp.).

Bird song plays an important role in the establishment and maintenance of prezygotic reproductive barriers. When two closely related species come into secondary contact, song convergence caused by acquisition of heterospecific songs into the birds' repertoires is often observed. The proximate mechanisms responsible for such mixed singing, and its effect on the speciation process, are poorly understood. We used a combination of genetic and bioacoustic analyses to test whether mixed singing observed in the secondary contact zone of two passerine birds, the Thrush Nightingale (Luscinia luscinia) and the Common Nightingale (L. megarhynchos), is caused by introgressive hybridization. We analysed song recordings of both species from allopatric and sympatric populations together with genotype data from one mitochondrial and seven nuclear loci. Semi-automated comparisons of our recordings with an extensive catalogue of Common Nightingale song types confirmed that most of the analysed sympatric Thrush Nightingale males were 'mixed singers' that use heterospecific song types in their repertoires. None of these 'mixed singers' possessed any alleles introgressed from the Common Nightingale, suggesting that they were not backcross hybrids. We also analysed songs of five individuals with intermediate phenotype, which were identified as F1 hybrids between the Thrush Nightingale female and the Common Nightingale male by genetic analysis. Songs of three of these hybrids corresponded to the paternal species (Common Nightingale) but the remaining two sung a mixed song. Our results suggest that although hybridization might increase the tendency for learning songs from both parental species, interspecific cultural transmission is the major proximate mechanism explaining the occurrence of mixed singers among the sympatric Thrush Nightingales. We also provide evidence that mixed singing does not substantially increase the rate of interspecific hybridization and discuss the possible adaptive value of this phenomenon in nightingales.

Territorial meadow pipit males (Anthus pratensis; Passeriformes) become more aggressive in female presence.

Although mate guarding as prevention of extra-pair copulation is common among birds, evidence for aggressive behaviour involving physical contact related to mate guarding in passerines is scarce and cases of the presence of one partner directly influencing the aggressiveness of the other are lacking. We investigated the intra-specific territorial behaviour of male meadow pipits (Anthus pratensis; Passeriformes: Motacillidae) at the beginning of the breeding season by placing a pipit model accompanied by an intra-specific song playback in the territory of socially paired males and compared the responses of males whose mates were physically present during trials with those whose females were out of sight. The level of aggression of males was significantly higher in the presence of the female; half of the males in this group physically attacked the model (the most intense and risky aggressive behaviour). Physical attacks did not occur among males whose female was absent during the trial; response to the playback by most of these males was only weak. This pattern may be related to the prevention of extra-pair copulation; if the risks involved in the conflict are outweighed by potential loss of paternity, such aggressive mate guarding may pay off. The apparently overlooked effect on the territorial behaviour of a partner's passive physical presence during conflict should be further evaluated because it may be important for the design and interpretation of results of behavioural experiments.