Differential geographic patterns in song components of male Albert’s lyrebirds

Geographic variation in bird song has received much attention in evolutionary studies, yet few consider components within songs that may be subject to different constraints and follow different evolutionary trajectories. Here, we quantify patterns of geographic variation in the socially transmitted “whistle” song of Albert''s lyrebirds (Menura alberti), an oscine passerine renowned for its remarkable vocal abilities. Albert''s lyrebirds are confined to narrow stretches of suitable habitat in Australia, allowing us to map likely paths of cultural transmission using a species distribution model and least cost paths. We use quantitative methods to divide the songs into three components present in all study populations: the introductory elements, the song body, and the final element. We compare geographic separation between populations with variation in these components as well as the full song. All populations were distinguishable by song, and songs varied according to the geographic distance between populations. However, within songs, only the introductory elements and song body could be used to distinguish among populations. The song body and final element changed with distance, but the introductory elements varied independently of geographic separation. These differing geographic patterns of within‐song variation are unexpected, given that the whistle song components are always produced in the same sequence and may be perceived as a temporally discrete unit. Knowledge of such spatial patterns of within‐song variation enables further work to determine possible selective pressures and constraints acting on each song component and provides spatially explicit targets for preserving cultural diversity. As such, our study highlights the importance for science and conservation of investigating spatial patterns within seemingly discrete behavioral traits at multiple levels of organization.     

Song mimicry of Black-bellied Firefinch Lagonosticta rara and other finches by the brood-parasitic Cameroon Indigobird Vidua camerunensis in West Africa

Brood-parasitic finches Vidua spp. mimic songs of their foster species, with most Vidua species both mimicking songs and parasitizing nests of a single estrildid finch species. We describe a behavioural radiation in the Cameroon Indigobird Vidua camerunensis . Local populations are polymorphic in behaviour, each male mimicking songs of a single species, with certain males mimicking songs of one species and other males mimicking songs of another host species. The species most often mimicked in song are Black-bellied Firefinch Lagonosticta rara and African Firefinch L. rubricata ; other species mimicked in song are Brown Twinspot Clytospiza monteiri and Dybowski's Twinspot Euschistospiza dybowskii . Indigobirds in the different mimicry song populations do not differ morphologically in plumage colour or size. The lack of morphological differences between male indigobirds with different mimicry songs is consistent with a recent behavioural radiation through host shifts, perhaps facilitated by environmental change associated with prehistoric cultivation of grain. The mimicry song populations of indigobirds, behaviourally imprinted upon different host species, support the idea of a process of speciation driven by a shift to new host species.     

Randomization Analyses: Mimicry, Geographic Variation and Cultural Evolution of Song in Brood-Parasitic Straw-Tailed Whydahs, Vidua fischeri

Although bird song has been an important model for investigating questions of behavior development, cultural evolution and population differentiation, the quantitative methods of analysis have been problematic. Here we develop and apply quantitative randomization methods to test hypotheses about these processes in a natural population of birds. Songs of the African brood-parasitic straw-tailed whydahs ( Vidua fischeri ) and songs of their host species, the purple grenadier ( Granatina ianthinogaster ), were compared in audiospectrograms for similarity to test the following hypotheses: Whydahs mimic the songs of their host species, they have local song dialects, neighboring males match their song themes, local males match the songs of local hosts, remote populations have different songs according to their geographic distance, and songs undergo cultural evolution over time across generations. Randomization analyses were completed using (1) Mantel matrix statistics and (2) tree-based measures employing Sankoff optimization of Manhattan matrices and approximate randomizations. Our results provide evidence for song mimicry, local song dialects, matching song themes between neighboring males, song matching of local whydah mimics and grenadier song models, correspondence of song differences and geographic distance, and cultural continuity with change in song traditions within a local population. These randomization methods may be useful in other studies of animal communication, and they are sufficiently general for use both with distance matrices derived either from naturalistic impressions of song similarity as in our example or from acoustic measurements.     

Whip it good! Geographic consistency in male songs and variability in female songs of the duetting eastern whipbird Psophodes olivaceus

Geographic variation in male bird songs has been studied extensively, but there have been few investigations of geographic variation in female songs or sex differences in patterns of geographic variation. We compared patterns of variation in male and female songs of eastern whipbirds Psophodes olivaceus by analyzing recordings from 16 populations across the species’ geographic range in eastern Australia. We found remarkably different patterns of geographic variation between the sexes. Female eastern whipbird songs are easily categorized into discrete song types. Song types are shared between nearby females, but female songs show pronounced differences at a continental scale. In contrast, male eastern whipbird songs show high consistency throughout the species’ geographic range. All recorded males share the ability to transpose the frequency of the introductory whistle and most recorded males share the ability to vary the direction of the slope of the terminal whip crack. For eight of nine measured variables, female songs show significantly higher levels of variation than male songs. We discuss whether sex differences in dispersal, song learning strategies, and song function may explain these sex differences in patterns of song variation. We suggest that eastern whipbirds have experienced a decoupling of male and female song learning strategies and that the songs of each sex have responded to different selective pressures in the context of countersinging interactions. We speculate that consistency in male songs throughout the geographic range of eastern whipbirds may arise through female preference for males that perform large bandwidth whip cracks.     

Song learning and deceptive mimicry

Payne, 1978, Payne, 1981a, Payne, 1981b has suggested that young males of some bird species copy the songs of the previous occupant or the songs of local established males when setting up a territory and thereby increase their likelihood of success. According to this view, song copying is a form of deceptive mimicry. We show that first-year great tits do not share more songs than expected with the previous occupants of their territories, nor are shared songs used more often than other songs in their repertoires. Songs are shared with neighbours but not selectively with established neighbours and there is no difference in reproductive success between great tits sharing with established neighbours and with first-year neighbours. We discuss the general applicability of the deceptive mimicry hypothesis and suggest that song sharing between neighbours may be advantageous for reasons other than deceptive mimicry.     

Mimicry as a novel pathway linking biodiversity functions and individual behavioural performances

The feedback of biodiversity on individual trait variation is a poorly explored mechanistic pathway in ecological research. We analysed the relationship between biodiversity and individual performance by focusing on vocal mimicry, a widespread interaction that may serve in intra- and interspecific communication. We studied the songs of two lark species (genus Galerida) that increase the complexity of their song displays by mimicking other birds, and analysed the influence of bird species richness on individual song performance. The diversity of mimicked species and the prevalence of mimicry increased in areas characterized by great α and γ diversity (i.e. where larks experience more diverse encounters with community members, many of them being highly vocal owing to breeding). Conversely, the variability in species-specific song components peaked where larks were abundant, probably matching the complexity of conspecific social milieu. Some trade-offs existed between homo- and heterospecific complexity, suggesting that larks could change from population- to community-driven song variation by tracking the composition of the auditory environment. Mimicry, which serves communication with conspecifics or predators, may mediate interactions, ultimately cascading to aspects of ecological diversity other than those promoting its complexity.     

Non‐Salient Geographic Variation in Birdsong in a Species That Learns by Improvisation

Geographic variation in birdsong and differential responses of territorial males to local and non‐local song variants have been documented in a number of songbird species in which males learn their songs through imitation. Here, we investigated geographic song variation and responses to local and non‐local song in the grasshopper sparrow (Ammodramus savannarum), a species in which males develop song by improvisation rather than imitation, as a first step toward understanding how the extent and salience of geographic song variation is related to the mode of song development. To describe the geographic variation in song, we compared songs from populations in eastern Maryland and central Ohio, USA, using multiple acoustic analysis techniques. We then conducted a playback experiment in Maryland using local and non‐local (Ohio) songs to test how territorial males responded to this geographic variation. We found acoustic differences between songs from the two sites. However, males responded similarly to playback of these songs, suggesting that this geographic variation is not behaviorally salient in a territorial context. Together with previous studies, our results suggest that across species, geographic song variation and the extent to which this variation functions in communication may be correlated with the accuracy with which song models are imitated during song development.     

Rufous‐capped Warblers Basileuterus rufifrons show seasonal,temporal and annual variation in song use

In the majority of songbird species, males have repertoires of multiple song types used for mate attraction and territory defence. The wood‐warblers (family Parulidae) are a diverse family of songbirds in which males of many migratory species use different song types or patterns of song delivery (known as ‘singing modes’) depending on context. The vocal behaviour of most tropical resident warblers remains undescribed, although these species differ ecologically and behaviourally from migratory species, and may therefore differ in their vocal behaviour. We test whether male Rufous‐capped Warblers Basileuterus rufifrons use distinct singing modes by examining song structure and context‐dependent variation in their songs. We recorded multiple song bouts from 50 male Warblers in a Costa Rican population over 3 years to describe seasonal, diel and annual variation in song structure and vocal behaviour. We found that Rufous‐capped Warbler songs are complex, with many syllable types shared both within and between males’ repertoires. Males varied their song output depending on context: they sang long songs at a high rate at dawn and during the breeding season, and shortened songs in the presence of a vocalizing female mate. Unlike many migratory species, Rufous‐capped Warblers do not appear to have different singing modes; they did not change the song variants used or the pattern of song delivery according to time of day, season or female vocal activity. Our research provides the first detailed vocal analysis of any Basileuterus warbler species, and enhances our understanding of the evolution of repertoire specialization in tropical resident songbirds.     

Responses of a sub-oscine bird during playback: effects of different song variants and breeding period

Vermilion flycatchers (Pyrocephalus rubinus) vary their song rate and song length across the breeding season. Males sing more and longer songs after nest construction than before. Here we explored the possibility that this variation is meaningful to territorial males. Using a playback approach, we tested several males with different variations in song output (i.e. variations in song length and song rate) in different periods of the breeding season (i.e. before and after the onset of nest construction). We found that males call more in response to playbacks of long and short songs before the onset of nest construction. However, after nest construction began they responded flying more when exposed to long songs than to short songs. These results show that vermilion flycatcher discriminates between different variants of song length, and suggest that males react to long songs as if they were more threatening signals than short songs, especially after the onset of nest construction. We did not find evidence of males discriminating between a high and a low song rate. We discuss some possible implications for song function in this sub-oscine species, and compare these findings with other results in oscine species.     

Male territorial aggression does not drive conformity to local vocal culture in a passerine bird

In many songbird species, young individuals learn songs from neighbors and then settle nearby, thus creating neighborhoods of conformity to local vocal culture. In some species, individuals appear to postpone song learning until after dispersal, possibly to facilitate conformity to local dialects. Despite decades of study, we still lack a consensus regarding the selective pressures driving this delayed song learning. Two common hypothetical benefits to conformity, and thus delayed song learning, are rooted in territorial interactions; individuals preferentially produce local song either to avoid detection as new arrivals (deceptive mimicry) or to be more effectively recognized as conspecific territory holders. The dickcissel (Spiza americana) is an ideal species in which to study these hypotheses. Males of this species appear to delay song learning until they arrive at their first adult territory, each individual sings a single song type, and conformity to the local song culture is high. Using playback, we contradicted both of the territorial hypotheses described above; male dickcissels did not respond differentially to local vs foreign song playback treatment. We are confident in this lack of difference because dickcissels clearly responded less strongly to a third treatment, neighbor song, than to the other two treatments, demonstrating sufficient power in our experimental design (and providing the first evidence of the dear‐enemy effect in dickcissels). Our results raise the question of why dickcissels respond equally aggressively to both local and foreign songs when other bird species often show reduced aggression toward foreign song. If reduced aggression to foreign song is not ubiquitous in species that achieve conformity through delayed learning, then selection from aggressive territorial interaction seems unlikely to be a general explanation for such delayed learning. Reduced aggression in response to foreign songs in other species may be due to reduced exposure to the stimulus of foreign song or to different cost‐benefit trade‐offs when responding to songs that deviate from the local average.     

Female choice for song characteristics in the house finch

Song has been proposed to function in mate choice, and such a role has been demonstrated experimentally in a number of species. We presented captive female house finches, Carpodacus mexicanus, with a choice of two songs, each of which was identical except for a single parameter that had been either accentuated or minimized. To ensure that our playback tapes offered choices within the normal range of variation in house finch song, we recorded wild male finches, then created playback tapes using values for the song parameters under consideration that were above or below the mean for the local population, but within the natural range of variation. We presented females that had not previously mated with choices between songs that (1) were long or short, (2) were given at fast or slow rates, and (3) included a large or small repertoire of unique song elements. Females showed significant preferences for long songs and for songs presented at a faster rate, but there was no significant preference for large or small repertoires of unique song elements. Song length and rate each seem likely to indicate a male's energy reserves, and thus could be important sources of information for females choosing mates.     

Song learning by prairie warblers: When,where, and from whom

Songbirds of many species acquire their songs by imitating the songs of conspecific singers. Conclusive evidence of such imitation comes from controlled laboratory studies, but such studies do not reveal when and where songbirds learn their songs under natural conditions. To determine the timing and location of song learning in a population of prairie warblers, we compared the songs of yearling prairie warblers of known hatching location to the songs of other birds in the yearlings' natal and first breeding areas. The comparisons yielded a likely model song (and model singer) for each of the song types used by the focal yearlings. We supplemented our findings from the song comparisons with inferences drawn from an analysis of local geographic variation in songs. This analysis revealed that shared song types showed no tendency to be geographically clustered within the study area. Taken together, our data suggest that prairie warblers learn their songs during the hatch year, at locations somewhat distant (mean distance 1,437 m) from their natal site, most likely as birds wander about during the post-fledging period.     

Whip it good! Geographic consistency in male songs and variability in female songs of the duetting eastern whipbird Psophodes olivaceus

Geographic variation in male bird songs has been studied extensively, but there have been few investigations of geographic variation in female songs or sex differences in patterns of geographic variation. We compared patterns of variation in male and female songs of eastern whipbirds Psophodes olivaceus by analyzing recordings from 16 populations across the species' geographic range in eastern Australia. We found remarkably different patterns of geographic variation between the sexes. Female eastern whipbird songs are easily categorized into discrete song types. Song types are shared between nearby females, but female songs show pronounced differences at a continental scale. In contrast, male eastern whipbird songs show high consistency throughout the species' geographic range. All recorded males share the ability to transpose the frequency of the introductory whistle and most recorded males share the ability to vary the direction of the slope of the terminal whip crack. For eight of nine measured variables, female songs show significantly higher levels of variation than male songs. We discuss whether sex differences in dispersal, song learning strategies, and song function may explain these sex differences in patterns of song variation. We suggest that eastern whipbirds have experienced a decoupling of male and female song learning strategies and that the songs of each sex have responded to different selective pressures in the context of countersinging interactions. We speculate that consistency in male songs throughout the geographic range of eastern whipbirds may arise through female preference for males that perform large bandwidth whip cracks.     

Acoustic but no genetic divergence in migratory and sedentary populations of blackcaps,Sylvia atricapilla

In songbirds, territorial songs are key regulators of sexual selection and are learned from conspecifics. The cultural transmission of songs leads to divergence in song characteristics within populations, which can ultimately lead to speciation. Many songbirds also migrate, and individual differences in migratory behaviours can influence population genetic structure and local song differentiation. Blackcaps, Sylvia atricapilla, exhibit versatile territorial songs and show diversity in migration behaviours. They therefore comprise a good model for investigating the relationships between migratory patterns, song variation, and genetic diversity. We studied a migratory population (two groups near Paris) and a sedentary population (three groups in Corsica). All of the birds were ringed and blood sampled to investigate genetic relatedness using 17 microsatellite loci. A detailed song analysis showed that this species has a complex repertoire (> 100 syllables), which required the development of a semi‐supervised method to classify different categories of syllables and compare sequences of syllables. Our analysis showed no genetic structuring among populations: individuals belonging to the same group were not genetically closer than those from different groups. However, we found a strong wingsize difference between sedentary and migratory populations. We also showed that geographical variations in songs rely at least on both syllable and sequence content. Unexpectedly, despite a higher turnover of individuals, migratory groups share as many syllables and sequences as sedentary groups, which raises interesting issues on song learning and the maintenance of dialects in migratory birds.     

Song characterization in the spectacled warbler (Sylvia conspicillata): a circum-Mediterranean species with a complex song structure

The spectacled warbler (Sylvia conspicillata) is a small passerine with a patchy distribution throughout the circum-Mediterranean region, including the North Atlantic archipelagos of Madeira, Canary Islands and Cape Verde. Here we characterize the species song structure on the island of Fuerteventura, quantifying repertoire size, inter- and intra-individual spectrographic variation, to determine whether acoustic variation occurred within an island population. Male song display was organized in song bouts of a variable number of song phrases, which in turn were made up of 4–69 syllables. We classified syllable types to derive a measure of repertoire size (number of different syllables) per song bout, and then used rarefaction methods to calculate the estimated repertoire size for our population of males. Three categories of song bout length were considered in analyses: short song bouts of 10 phrases, average bouts of 19 phrases and long bouts of ≥ 29 phrases. The observed and estimated repertoire size per male (between 43 and 126 syllables per male) increased with song bout duration, although the relationship was not significant for the estimated values. To test whether songs could be individually specific, we measured 11 spectrotemporal parameters of the song. A discriminant analysis using these variables performed poorly in classifying songs to the individuals that uttered them, but we found less variation in the individual than in the population for three out of the 11 variables. These individually specific variables, involving the first or the most common syllable of the song, the trill, were the duration of the first syllable of the phrase, the duration and the dominant frequency of the trill syllable. Our study emphasizes the complexity of spectacled warbler songs, in which males continuously add novel syllables over the entire song bout. This complexity appears to be determined by individual innovation capabilities rather than by the behaviour of copying neighbour repertoires, since songs of close birds were not more similar than songs from far-away territories.     

A new species of firefinch Lagonosticta from northern Nigeria and its association with the Jos Plateau Indigobird Vidua maryae

A firefinch of the Jos Plateau, northern Nigeria, is described as a new species, Rock Fire-finch Lagonosticta sanguinodorsalis. This firefinch has distinctive songs and a distinctive habitat—bushy and grassy rocky outcrops on the plateau and inselbergs to the north and east. It has a blue-grey bill, red back in the male and reddish brown back in the female and juvenile and broad primaries in both the adult and juvenile. It appears most closely related to the Mali Firefinch Lagonosticta virata and the Chad Firefinch Lagonosticta umbrinodorsalis. Its songs are mimicked by the brood-parasitic Jos Plateau Indigobird Vidua maryae , which occurs within the range of the firefinch and whose songs led to the discovery of the firefinch. The songs of the firefinch, the song mimicry of the indigobird and the apparent continuity of song mimicry across 27 years indicate a unique association of this distinct pair of species.     

Cultural isolation is greater than genetic isolation across an avian hybrid zone

Elucidating the relationship between genetic and cultural evolution is important in understanding speciation, as learned premating barriers might be involved in maintaining species differences. Here, we test this relationship by examining a widely recognized premating barrier, bird song, in a hybrid zone between black‐throated green (Setophaga virens) and Townsend's warblers (S. townsendi). We use song analysis, genomic techniques and playback experiments to characterize the cultural and genetic backgrounds of individuals in this region, expecting that if song is an important reproductive barrier between these species, there should be a strong relationship between song and genotype. We show that songs in the hybrid zone correspond to the distinctly different songs found in allopatry but that song and genotype are not tightly coupled in sympatry. Allopatric individuals responded only to local songs, indicating that individuals may have learned to respond to songs they commonly hear. We observed discordance between song and genotype clines; a narrower cline suggests that cultural selection on song is stronger than natural selection on genotype. These findings indicate that song is unlikely to play a role in reproductive isolation between these species, and we suggest that spatial variation in song may nonetheless be maintained by frequency‐dependent cultural selection. This decoupling of genes and culture may contribute to hybridization in this region.     

Classification of group-specific variations in songs within House Wren species using machine learning models

Songbirds have shown variation in vocalizations across different populations and different geographical ranges. Such variations can over time lead to divergence in song characteristics, sometimes referred to as dialects. House Wren (Troglodytes aedon) is one such widely distributed bird species that has shown variation in its song characteristics within different populations. Traditionally, such studies have been conducted using manual approaches for classification. In this work we explore the use of machine learning models that can assist in performing classification of bird songs at a conspecific level. Two machine learning techniques, the random forest and a shallow feed forward neural network, are fed with pre-computed sound features to classify vocal variation in House Wren species across different reported population groups and latitudinal areas. A randomized approach is employed to create balanced subsets of sounds from different locations for repeated classification runs in order to provide a reliable estimate of performance. It is observed that such an automated approach is able to classify variations in songs within House Wren with high accuracy. We were also able to confirm the latitudinal variation of House Wren songs reported in previous studies. Given these results, we believe, such a purely data-driven way of analyzing bird songs in general can provide useful hints to biologists on where to look for interesting patterns in order to understand the evolutionary divergence in song characteristics.     

What exactly is ‘local song’ in a population of ortolan buntings with a common dialect?

Previous studies have shown that ortolan buntings (Emberiza hortulana) exhibit apparent dialect variation. Neighbouring males typically share the same final phrase of a simple two-part song. Consequently, the final phrase was considered to be the dialect cue important for discriminating between males from a local population and strangers. Recently, it was shown that in an isolated and fragmented population of the ortolan buntings in Norway there was no local dialect in the above-mentioned sense. Norwegian males often had song types with different final phrases in their repertoire, and had larger repertoires and a lower level of song type sharing than in other populations. It was experimentally revealed that only local songs (L) evoked a strong response in Norwegian males, while these birds did not respond strongly to foreign (F) or hybrid songs composed of local and foreign initial and final phrases in any composition (i.e. both FL and LF songs). These results suggest that, in the Norwegian population, the final phrase of the song is not a sufficient cue for local song dialect recognition. This paper is a further study in which we tested the response of the ortolan bunting males to L, F, FL and LF songs in a typical continuous population (in Poland) in which males share the same single final phrase, i.e. have a common dialect. We found that ortolan bunting males in Poland responded with similar strength to L, FL and LF songs. The majority of males did not respond as strongly to playback of only F songs. Our results suggest that a common final phrase for a population should not be treated as the only signal of ‘locality’. Our results show clear asymmetry in response to hybrid songs with non-local phrases in different populations.     

Parallel plasticity of mating songs and preferences in the field cricket Gryllus rubens

In animal communication systems, matching mating signals and preferences enable species identification and successful reproduction. In some species, the environment introduces substantial variation in signals and/or preferences. Only a few studies have tested how the match between signals and preferences is maintained despite phenotypic variation. Signal–preference coupling in the context of phenotypic plasticity is the focus of this study. The bivoltine cricket Gryllus rubens displays seasonal differences in the pulse rate of its mating songs. The seasonal effect on other fine‐temporal characters of the songs besides pulse rate, such as pulse and interval duration, duty cycle, as well as the dominant frequency, is not known and is described in the first part of the study for a Kentucky population. In the second part of the study, we tested preferences of spring and fall females to determine whether they match the seasonal plasticity of male songs using single‐speaker phonotaxis experiments. We found that fall songs had a faster pulse rate, shorter pulse and interval durations, and a higher dominant frequency than spring songs. Female preferences shifted in parallel with male song plasticity, that is, spring females preferred the spring song and fall females the fall song. In addition, female responsiveness to male song was plastic as well, that is, fall females were significantly more responsive than spring females. The parallel plasticity of male songs and female preferences facilitates successful communication despite the environmentally induced variation. The potential origin and function of behavioral plasticity in G. rubens are discussed.