Song Traits Research Articles

Song divergence indicates an unclear relationship between the Neotropical and Nearctic Horned Larks

BackgroundSome studies have indicated that the Horned Larks (Eremophila alpestris) should be considered as a species complex. Recently it split into four species or clades and seven subclades based on genetic, morphological, and biogeographic data. However, other aspects like song divergence have not been studied and several subspecies have not been evaluated, leading to important information gaps in this group of birds. In this work, we aimed to assess the differences in song traits and playback response between the Nearctic subclade and the Neotropical or Colombian subspecies E. a. peregrina.MethodsWe compared six song traits between these groups and performed field playback experiments, to test the response of the Neotropical larks to both songs. We tested the difference in the variables for separate as well as by principal component analysis (PCA).ResultsWe found significant differences (p < 0.05) in the individual song traits and the PCA analysis between the two groups. Further, the PCA analysis showed a clearer divergence of the Neotropical songs in comparison to the Nearctic songs of different locations within North America. Similarly, the playback analysis showed a significantly lower response of E. a. peregrina to the songs of the Nearctic larks.ConclusionsBesides this song divergence, there are important ecological and biogeographic differences between the Neotropical and Nearctic Horned Larks, that indicate an unclear relationship between these two groups. Thus, further morphological and genetic studies are required to clarify the taxonomy of the Neotropical Horned Lark and define if they share the same evolutionary history as the other subspecies of the Nearctic subclade.

The effect of social environment on bird song: listener-specific expression of a sexual signal.

Animal signals should consistently differ among individuals to convey distinguishable information about the signalers. However, behavioral display signals, such as bird song are also loaded with considerable within-individual variance with mostly unknown function. We hypothesized that the immediate social environment may play a role in mediating such variance component, and investigated in the collared flycatcher (Ficedula albicollis) if the identity and quality of listeners could affect song production in signalers. After presenting territorial males with either a female or male social stimulus, we found in the subsequent song recordings that the among-stimulus effects corresponded to non-zero variance components in several acoustic traits indicating that singing males are able to plastically adjust their songs according to stimulus identity. Male and female stimuli elicited different responses as the identity of the female stimuli affected song complexity only, while the identity of male stimuli altered also song length, maximum frequency, and song rate. The stimulus-specific effect on song in some cases decreased with time, being particularly detectable right after the removal of the stimulus and ceasing later, but this pattern varied across the sex of the stimulus and the song traits. We were able to identify factors that can explain the among-stimulus effects (e.g., size and quality of the stimuli) with roles that also varied among song traits. Our results confirm that the variable social environment can raise considerable variation in song performance, highlighting that within-individual plasticity of bird song can play important roles in sexual signaling.

Senescence of song revealed by a long-term study of the Seychelles warbler (Acrocephalus sechellensis)

Senescence is widespread in nature, often resulting in diminishing survival or reproduction with age, but its role in age-dependent variation in sexual traits is often poorly understood. One reason is that few studies of sexual traits consider non-linear relationships with age, or only consider a narrow range of years relative to the life span of the species. Birdsong has evolved to allow assessment of conspecific quality in numerous bird species. Whilst theory and empirical work suggests that song may become more elaborate with age, there are a paucity of long-term studies testing whether song is associated with age or longevity. In particular, the occurrence of song senescence has rarely been demonstrated. Using an exceptional long-term dataset for the Seychelles warbler (Acrocephalus sechellensis), we analysed relationships between male song, age, survival, and longevity. This species is a long-lived songbird with early life increases, followed by senescent declines, in survival and reproduction. The study population (Cousin Island, Seychelles) is a closed population, with no depredation of adults, providing an excellent opportunity to study senescence in free-living animals. We tested whether song traits were related to age at recording, future survival, longevity, and territory quality. We found age-dependent changes in five song traits (duration, maximum frequency, peak frequency of songs, and duration and frequency bandwidth of trills). Relationships with age were quadratic, indicating reversal in the expression of song coinciding with the onset of senescence in reproduction and survival in this species. One song trait (trill bandwidth) had a quadratic relationship with future survival, but no song traits were related to longevity, suggesting age-related patterns were not the result of selective disappearance. Our study provides one of the first examples of functional senescence in song, offering new insights into avian senescence. Late-life declines in avian song, and possibly other sexual traits, may be more common than currently known, and may play a fundamental role in age-dependent changes in reproductive success.

Analysis of female song provides insight into the evolution of sex differences in a widely studied songbird

Understanding the patterns and processes related to sexual dimorphism and sex differences in diverse animal taxa is a foundational research topic in ecology and evolution. Within the realm of animal communication, studies have traditionally focused on male signals, assuming that female choice and male–male competition have promoted sex differences via elaboration of male traits, but selection on females also has the potential to drive divergence. Here, we describe female song in barn swallows, Hirundo rustica erythrogaster , for the first time, report rates of female song production and couple song data with plumage data to explore the relative degree to which sex differences in phenotypic traits are consistent with contemporary selection on males versus females. During previous intensive study of male song over 2 years, we opportunistically recorded songs for 15 females, with matched phenotypic and fitness data. We randomly selected 15 high-quality samples from our larger male data set to test whether sex differences in song and plumage are more strongly associated with fledgling success for females or genetic paternity for males. Analyses included 35 potential sexual signals including 22 song parameters and 13 plumage traits. Outcomes indicate that female songs were used in multiple contexts, restricted primarily to the beginning of the breeding season; song traits showed greater sexual differentiation than visual plumage traits; and trait correlations with reproductive success in females, rather than males, predicted sex-based differences in song and plumage. These results are consistent with phylogenetic studies showing that sex-based phenotypic differences are driven by changes in females, highlighting the potential role of female trait evolution in explaining patterns of sexual differentiation. To achieve a better understanding of sex differences and dimorphism, we require comprehensive studies that measure the same traits in males and females and their fitness consequences. • We describe and quantify female song in barn swallows for the first time. • Females sang about 10 times less than males, and only early in the breeding season. • Song traits showed greater sex differences than visual plumage traits. • Sexual differentiation was correlated with female, but not male, reproductive success.

Social context and noise affect within and between male song adjustments in a common passerine

AbstractAcross populations, animals that inhabit areas with high anthropogenic noise produce vocalizations that differ from those inhabiting less noisy environments. Such patterns may be due to individuals rapidly adjusting their songs in response to changing noise, but individual variation is seldom explored. Therefore, we tested the hypothesis that male house wrens (Troglodytes aedon) immediately adjust their songs according to changing noise and that social context further modifies responses. We recorded songs, quantified noise, and defined social context within pairs as female fertile status and between males as number of conspecific neighbors. We used a reaction-norm approach to compare song trait intercepts (between-male effects) and slopes (within-male effects) as a function of noise. Individuals immediately adjusted song duration in response to changing noise. How they achieved adjustments varied: some sang shorter and others longer songs with greater noise, and individuals varied in the extent to which they adjusted song duration. Variation in song duration could be affected by competition as between-male noise levels interacted with number of neighbors to affect syllable duration. Neither within- nor between-male noise effects were detected for frequency traits. Rather, males with fertile mates sang lower-frequency songs and increased peak frequency with more neighbors. Among males, social context but not noise affected song frequency, whereas temporal structure varied between and within individuals depending on noise and social factors. Not all males adjusted signals the same way in response to noise, and selection could favor different levels of variation according to noise.

Vocal recognition suggests premating isolation between lineages of a lekking hummingbird

AbstractSpecies with genetically differentiated allopatric populations commonly differ in phenotypic traits due to drift and/or selection, which can be important drivers of reproductive isolation. Wedge-tailed sabrewing (Campylopterus curvipennis) is a species complex composed of three genetically and acoustically differentiated allopatric lineages that correspond to currently recognized subspecies in Mexico: C. c. curvipennis (Sierra Madre Oriental), C. c. pampa (Yucatán Peninsula), and C. c. excellens (Los Tuxtlas). Although excellens is taxonomically recognized as a distinct species, there is genetic evidence that lineages excellens and curvipennis have diverged from each other later than pampa. In this study, we experimentally tested C. c. curvipennis song recognition as a major factor in premating reproductive isolation for lineage recognition. To this end, we conducted a song playback experiment to test whether territorial males of one C. c. curvipennis lek discriminate among potential competitors based on male songs from the three lineages. Males of curvipennis responded more aggressively to songs of their own lineage and excellens, than to songs of the most divergent lineage pampa, as evidenced by significant differences in a variety of intensity and latency response variables. This indicate that the pampa male song does not represent a competitive threat as curvipennis and excellens songs, in which divergence and song recognition represent premating reproductive isolation between these isolated lineages. However, the acoustic limits between curvipennis and excellens might be attenuated by gene flow in case of secondary contact between them, despite the strong and relatively rapid divergence of their sexually selected song traits.

Song amplitude and population density in two sympatric warblers, Phylloscopus schwarzi and P. fuscatus

ABSTRACT In passerine birds, song is widely used broadcasting signal. One song trait that so far has received little attention is amplitude, i.e. the sound pressure level of a song. It has been shown that there is a considerable variation in song amplitude between species, but the causes and consequences of this variation are poorly studied. Here, we measured song amplitude in two similar-sized sympatric Phylloscopus, Radde’s Warbler P. schwarzi and Dusky Warbler P. fuscatus. We found that Radde’s Warbler males sing much more loudly than males of Dusky Warbler. The sound pressure level values of songs of Radde’s and Dusky Warblers were 78.6 ± 2.4 dB(A) (n = 16 males) and 70.9 ± 1.5 dB(A) (n = 16) at a distance of 1 m, respectively. The analysis of video-recordings revealed the larger extension of the neck region (‘throat sac’) in Radde’s Warbler than in Dusky Warbler. We hypothesised that the larger maximum ‘throat sac’ volume might be responsible for the louder song of Radde’s Warbler. We also found that Dusky Warbler males settled twice as close to each other than Radde’s Warblers. We hypothesised that the louder song of Radde’s Warbler could be useful for males of this species given their need to communicate through a larger distance.

Principles of interpreting P. Tychyna’s poetry in the vocal cycle “Enharmonic” by L. Dychko

Principles of interpreting P. Tychyna’s poetry in the vocal cycle “Enharmonic” by L. Dychko

Genetic mating system, population genetics and effective size of Saffron Finches breeding in southern South America.

Thraupidae (Tanagers and allies) show a remarkable array of behaviors, ecologies, morphologies and plumage colors, offering a great opportunity to investigate the evolution of avian mating strategies. We characterize the population genetics and mating system of Sicalis flaveola pelzelni, a socially monogamous Neotropical songbird with biparental care. We found moderate to high levels of neutral genetic variation, similar across three breeding seasons, consistent with large and stable populations, and no temporal genetic structure. Parentage analyses of 114 adults and 198 nestlings (54 nests) revealed 31.8% of extra-pair offspring (EPO) and 51.8% of broods with at least one extra-pair chick. Extra-pair paternity (EPP) rates varied across seasons and were significantly higher in 2014/2015. Neither breeding synchrony nor pair genetic relatedness was significantly associated to EPP rates. Males paired with females in nests with and without EPO were equally heterozygous, and EPO were significantly less heterozygous than within-pair offspring, not supporting the 'indirect genetic benefits' hypotheses. Females were more related to their extra-pair mates than to their social mates, not supporting the 'inbreeding avoidance' hypothesis. The non-monogamous genetic mating system uncovered here seems not to lower the effective size of the population, which was higher than the sample size of adult breeders. We report and discuss possible cases of quasi-parasitism, as indicated by maternity exclusion patterns. We contribute novel information to expand the knowledge about the largely unexplored genetic mating systems of Thraupidae. Our findings also set the stage for further studies examining if plumage coloration or song traits predict paternity gain or loss in Saffron Finches.

Repeatability of signalling traits in the avian dawn chorus

BackgroundBirdsong, a key model in animal communication studies, has been the focus of intensive research. Song traits are commonly considered to reflect differences in individual or territory quality. Yet, few studies have quantified the variability of song traits between versus within individuals (i.e. repeatability), and thus whether certain song traits indeed provide reliable individual-specific information. Here, we studied the dawn chorus of male great tits (Parus major) to determine if key song traits are repeatable over multiple days and during different breeding stages. Additionally, we examined whether repeatability was associated with exploration behaviour, a relevant personality trait. Finally, we tested if variation in song traits could be explained by breeding stage, lowest night temperature, and exploration behaviour.ResultsWe show that the start time of an individual’s dawn song was indeed repeatable within and across breeding stages, and was more repeatable before, than during, their mate’s egg laying stage. Males started singing later when the preceding night was colder. Daily repertoire size was repeatable, though to a lesser extent than song start time, and no differences were observed between breeding stages. We did not find evidence for an association between exploration behaviour and variation in dawn song traits. Repertoire composition, and specifically the start song type, varied across days, but tended to differ less than expected by chance.ConclusionsOur findings that individuals consistently differ in key song traits provides a better understanding of the information receivers can obtain when sampling songs of different males. Surprisingly, start time, despite being influenced by a highly variable environmental factor, appeared to be a more reliable signal of individual differences than repertoire size. Against expectation, singers were more repeatable before than during their mate’s egg laying stage, possibly because before egg laying, females are less constrained to move around unguarded and thus may then already sample (and compare) different singers. Combining repeated dawn song recordings with spatial tracking could reveal if the sampling strategies of receivers are indeed important drivers of repeatability of song traits. Such a complementary approach will further advance our insights into the dynamics and evolution of animal signalling systems.

The variability of song variability in zebra finch (Taeniopygia guttata) populations.

Birdsong is a classic example of a learned social behaviour. Song behaviour is also influenced by genetic factors, and understanding the relative contributions of genetic and environmental influences remains a major goal. In this study, we take advantage of captive zebra finch populations to examine variation in a population-level song trait: song variability. Song variability is of particular interest in the context of individual recognition and in terms of the neuro-developmental mechanisms that generate song novelty. We find that the Australian zebra finch Taeniopygia guttata castanotis (TGC) maintains higher song diversity than the Timor zebra finch T. g. guttata (TGG) even after experimentally controlling for early life song exposure, suggesting a genetic basis to this trait. Although wild-derived TGC were intermediate in song variability between domesticated TGC populations and TGG, the difference between domesticated and wild TGC was not statistically significant. The observed variation in song behaviour among zebra finch populations represents a largely untapped opportunity for exploring the mechanisms of social behaviour.

Physical linkage and mate preference generate linkage disequilibrium for behavioral isolation in two parapatric crickets.

Behavioral isolation is a potent barrier to gene flow and a source of striking diversity in the animal kingdom. However, it remains unclear if the linkage disequilibrium (LD) between sex‐specific traits required for behavioral isolation results mostly from physical linkage between signal and preference loci or from directional mate preferences. Here, we test this in the field crickets Gryllus rubens and G. texensis. These closely related species diverged with gene flow and have strongly differentiated songs and preference functions for the mate calling song rhythm. We map quantitative trait loci for signal and preference traits (pQTL) as well as for gene expression associated with these traits (eQTL). We find strong, positive genetic covariance between song traits and between song and preference. Our results show that this is in part explained by incomplete physical linkage: although both linked pQTL and eQTL couple male and female traits, major effect loci for different traits were never on the same chromosome. We suggest that the finely tuned, highly divergent preference functions are likely an additional source of LD between male and female traits in this system. Furthermore, pleiotropy of gene expression presents an underappreciated mechanism to link sexually dimorphic phenotypes.

Dialects of Metrioptera bicolor (Orthoptera: Tettigonioidea): Intraspecific Acoustic Signal Divergence in a Small-Scale Geographical Distribution

We present an analysis of the calling songs of species Metrioptera bicolor isolated from different locations within Northeast China illustrating the intraspecies calling song divergence. Recording and analysis of the male calling songs of some acoustic characters were performed and five morphological traits of M. bicolor were examined under stereoscope microscope and scanning electron microscope. A cluster analysis was made based on the song properties and the morphology of stridulatory organs. All the song traits, except the pulse number per pulse group, differ distinctly within species collected at different localities, while the pulse number of each pulse group and all the five morphological characters were consistent across locations. The cluster analysis showed that M. bicolor collected from eight sampling sites grouped into two branches and were consistent with geographical distribution.The dialect phenomenon exists in insects and the divergence of acoustic signals occurs in a small-scale geographical region and may be an early stage of speciation. We inferred that the divergence of intraspecific acoustic signals on a small-scale may result from different habitats. This study provides evidence for insect acoustic signal divergence, speciation, and insect acoustic signal evolution.

Duetting correlates with territory quality and reproductive success in a suboscine bird with low extra-pair paternity

AbstractDuetting has been intensively studied, but we still have little consensus about its fitness consequences. Some studies suggest that duetting functions in acoustic mate guarding to prevent cuckoldry (acoustic paternity guarding hypothesis), whereas other studies argue that duetting is a cooperative behavior to defend common resources (territory defense hypothesis). We tested these 2 hypotheses by investigating the relationships among song traits, extra-pair paternity, territory quality and reproductive success in the Rufous Hornero (Furnarius rufus), a Neotropical, socially monogamous bird. We found a low rate of extra-pair paternity (3.33% of 120 offspring and 6.52% of 46 broods), which suggests that acoustic paternity guarding is probably not a primary function of duetting behavior in this species. Female song output was positively correlated with territory quality, measured as the availability of territory foraging patches. The number of young that survived post-fledging was positively correlated with duet duration, but not with territory features. Our results support the territory defense hypothesis, suggesting that female song in duets might be used in the defense of food resources within territories. Our study provides the first evidence that song traits correlate with both territory features and reproductive success in a duetting species.

Behavioural mechanisms of sexual isolation involving multiple modalities and their inheritance.

Speciation research dissects the genetics and evolution of reproductive barriers between parental species. Hybrids are the "gatekeepers" of gene flow, so it is also important to understand the behavioural mechanisms and genetics of any potential isolation from their parental species. We tested the role of multiple behavioural barriers in reproductive isolation among closely related field crickets and their hybrids (Teleogryllus oceanicus and Teleogryllus commodus). These species hybridize in the laboratory, but the behaviour of hybrids is unusual and there is little evidence for gene flow in the wild. We found that heterospecific pairs exhibited reduced rates of courtship behaviour due to discrimination by both sexes, and that this behavioural isolation was symmetrical. However, hybrids were not sexually selected against and exhibited high rates of courtship behaviour even though hybrid females are sterile. Using reciprocal hybrid crosses, we characterized patterns of interspecific divergence and inheritance in key sexual traits that might underlie the mating patterns we found: calling song, courtship song and cuticular hydrocarbons (CHCs). Song traits exhibited both sex linkage and transgressive segregation, whereas CHCs exhibited only the latter. Calculations of the strength of isolation exerted by these sexual traits suggest that close-range signals are as important as long-distance signals in contributing to interspecific sexual isolation. The surprisingly weak mating barriers observed between hybrids and parental species highlight the need to examine reproductive isolating mechanisms and their genetic bases across different potential stages of introgressive hybridization.

Social effects on fruit fly courtship song.

Courtship behavior in Drosophila has often been described as a classic innate behavioral repertoire, but more recently extensive plasticity has been described. In particular, prior exposure to acoustic signals of con‐ or heterspecific males can change courtship traits in both sexes that are liable to be important in reproductive isolation. However, it is unknown whether male courtship song itself is socially plastic. We examined courtship song plasticity of two species in the Drosophila melanogaster subgroup. Sexual isolation between the species is influenced by two male song traits, the interpulse interval (IPI) and sinesong frequency (SSF). Neither of these showed plasticity when males had prior experience of con‐ and heterospecific social partners. However, males of both species produced longer bursts of song during courtship when they were exposed to social partners (either con‐ or heterospecific) than when they were reared in isolation. D. melanogaster carrying mutations affecting short‐ or medium‐term memory showed a similar response to the social environment, not supporting a role for learning. Our results demonstrate that the amount of song a male produces during courtship is plastic depending on the social environment, which might reflect the advantage of being able to respond to variation in intrasexual competition, but that song structure itself is relatively inflexible, perhaps due to strong selection against hybridization.

The effect of early life conditions on song traits in male dippers (Cinclus cinclus).

Song complexity and singing frequency in male birds are shaped by female choice; they signal male quality because song is costly to develop and produce. The timing of song learning and the development of the brain structures involved occur during a period when chicks are exposed to a number of potential stressors. The quality and quantity of song produced by adults may therefore reflect the level of stress experienced during early life, a theory known as the ‘developmental stress hypothesis’. We tested this hypothesis using song recordings and life-history data from an individually marked, long-term study population of wild dippers (Cinclus cinclus). The extent to which early life conditions predict adult song traits was investigated using natal brood size as a measure of sibling competition; the rate of provisioning by parents as a proxy for nutritional stress; and residuals of the linear regression between body mass and tarsus length as a measure of nestling condition. The syllable diversity in the songs of adult males was positively correlated with their body condition as nestlings, but there was no significant correlation with either provisioning rate or brood size. Provisioning rate did, however, predict song rate; males in relatively poor condition as nestlings or those raised in smaller broods which were fed more frequently by their parents sang at a higher rate in adulthood. These results support the developmental stress hypothesis and provide some of the first evidence from a wild bird of how the conditions experienced during early life impact adult song. Song traits may therefore provide females with information regarding both the current condition and developmental history of males.

Multi-modal signal evolution in birds: re-examining a standard proxy for sexual selection.

Sexual selection is proposed to be an important driver of speciation and phenotypic diversification in animal systems. However, previous phylogenetic tests have produced conflicting results, perhaps because they have focused on a single signalling modality (visual ornaments), whereas sexual selection may act on alternative signalling modalities (e.g. acoustic ornaments). Here, we compile phenotypic data from 259 avian sister species pairs to assess the relationship between visible plumage dichromatism-a standard index of sexual selection in birds-and macroevolutionary divergence in the other major avian signalling modality: song. We find evidence for a strong negative relationship between the degree of plumage dichromatism and divergence in song traits, which remains significant even when accounting for other key factors, including habitat type, ecological divergence and interspecific interactions. This negative relationship is opposite to the pattern expected by a straightforward interpretation of the sexual selection-diversification hypothesis, whereby higher levels of dichromatism indicating strong sexual selection should be related to greater levels of mating signal divergence regardless of signalling modality. Our findings imply a 'trade-off' between the elaboration of visual ornaments and the diversification of acoustic mating signals, and suggest that the effects of sexual selection on diversification can only be determined by considering multiple alternative signalling modalities.

Male competition drives song divergence along an ecological gradient in an avian ring species.

Sexual selection operates via female choice and male competition, which can act independently, in concert, or in opposition. Female choice is typically considered the stronger selective force, but how these two processes interact to shape phenotypic divergence is poorly understood. I tested the hypothesis that variation in sexual selection in different habitats drives song divergence in the greenish warbler ring species. I evaluated the strength, direction, and targets of female choice and male competition in three populations spanning 2400km of latitude. Average song length increased with latitude, concomitant with a decline in population density. Within populations, males sang longer songs when females were fertile and shorter songs during territory establishment. Females consistently preferred males with longer songs and larger song repertoires. By contrast, playback experiments showed that males used short songs in territory defense. Songs were shortest at high densities, and in the highest density population only, song traits preferred by females correlated with male territoriality. Stronger male competition at high population densities likely constrains maximum song length, whereas weaker competition at low densities allows expression of female choice for long songs. Interactions between male competition and ecology may be a crucial but oft-overlooked component of phenotypic divergence and speciation.

When to change your tune? Unpaired and paired male house wrens respond differently to anthropogenic noise

In response to anthropogenic noise, many bird species adjust their song frequency, presumably to optimize song transmission and overcome noise masking. But the costs of song adjustments may outweigh the benefits during different stages of breeding, depending on the locations of potential receivers. Selection might favor unpaired males to alter their songs because they sing to attract females that may be widely dispersed, whereas paired males might not if mates and neighbors are primary receivers of their song. We hypothesized male house wrens (Troglodytes aedon) respond differently to noise depending on their pairing status. To test our hypothesis we synthesized pink noise, which mimics anthropogenic noise, and played it at three intensities in territories of paired and unpaired focal males. We recorded their songs and analyzed whether song structure varied with pairing status and noise treatment. To validate our study design, we tested whether noise playback affected measurement of spectral song traits and changed noise levels within territories of focal males. Consistent with our predictions, unpaired males sang differently than paired males, giving longer songs at higher rates. Contrary to predictions, paired males changed their songs by increasing peak frequency during high intensity noise playback, whereas unpaired males did not. If adjusting song frequency in noise is beneficial for long-distance communication we would have expected unpaired males to change their songs in response to noise. By adjusting song frequency, paired males reduce masking and produce a song that is easier to hear. However, if females prefer low frequency song, then unpaired males may be constrained by female preference. Alternatively, if noise adjustments are learned and vary with experience or quality, unpaired males in our study population may be younger, less experienced, or lower quality males.