Vocal performance influences male receiver response in the banded wren

In a variety of songbirds the production of trilled song elements is constrained by a performance tradeoff between how fast a bird can repeat trill units (trill rate) and the range of frequencies each unit can span (frequency bandwidth). High-performance trills serve as an assessment signal for females, but little is known about the signal value of vocal performance for male receivers. We investigated the relationship between trill rate and frequency bandwidth in banded wren (Thryothorus pleurostictus) songs. Trilled song elements showed the same performance tradeoff found in other passerines and individuals differed in performance of some trill types. We tested the hypothesis that males of this species assess each other based on trill performance with a two-speaker experiment, in which territory owners were presented with alternating renditions of the same song type manipulated to differ in trill rate. Subjects were significantly more likely to approach the faster trill stimulus first. However, subjects that received trill types closer to the performance limit spent less time close to the fast speaker. Our results show that male banded wrens discriminate and respond differently to songs based on their vocal performance. Thus, performance of physically challenging songs may be important in intra- as well as inter-sexual assessment.     

Variation in Trill Characteristics in Tree Pipit Songs: Different Trills for Different Use?

Transmitting information about singer's quality is an important function of song in many bird species, and this information should be useful in territorial interactions. Fast trills, being physically demanding song structures, are particularly suitable candidates for signalling of quality or aggressive motivation. We have evaluated trill characteristics in songs within a population of the Tree Pipit, a common European songbird with no sexual dimorphism, in which song apparently plays a key role in territory defence as well as mate choice. Two types of relatively fast trills (each of them in multiple variants differing in complexity) were commonly observed in repertoires of Tree Pipit males. Trill rates significantly differed among individuals, suggesting that these song structures may carry information about male quality in this species. We tested by playback experiments whether both trill types are used in territorial encounters. Only one of the trill types was sung by males in response to playback, regardless on the trill type played to them. In an immediate response to playback, they increased the frequency of use of this trill, and also significantly increased the trill rate in comparison with spontaneous songs. This confirmed field observations, suggesting that this trill is important in male–male interactions. On the contrary, the use of the fastest, apparently more demanding, trill type actually decreased after the simulated territorial intrusion. We hypothesize that the latter one is more directed towards females, and that while performance of both trill types may reflect male quality, they are primarily used in different contexts.     

Vocal performance influences female response to male bird song: an experimental test

Female songbirds are thought to assess males based on aspectsof song, such as repertoire size or amount of singing, thatcould potentially provide information about male quality. Arelatively unexplored aspect of song that also might serve asan assessment signal is a male's ability to perform physicallychallenging songs. Trilled songs, such as those produced byswamp sparrows (Melospiza georgiana), present males with a performancechallenge because trills require rapid and precise coordinationof vocal tract movements, resulting in a trade-off between trillrate and frequency bandwidth. This trade-off defines a constrainton song production observed as a triangular distribution inacoustic space of trill rate by frequency bandwidth, with anupper boundary that represents a performance limit. Given thisbackground on song production constraints, we are able to identifya priori which songs are performed with a higher degree of proficiencyand, thus, which songs should be more attractive to females.We determined the performance limit for a population of swampsparrows and measured how well individual males performed songsrelative to this limit ("vocal performance"). We then comparedfemale solicitation responses to high-performance versus low-performanceversions of the same song type produced by different males.Females displayed significantly more to high-performance songsthan to low-performance songs, supporting the hypothesis thatfemales use vocal performance to assess males.     

Sound-pressure Levels and Song Preferences in Female Red-winged Blackbirds (Agelaius phoeniceus) (Aves,Emberizidae)

Female red-winged blackbirds (Agelaius phoeniceus) were tested for response to songs of male red-winged blackbirds differing in sound-pressure level (SPL) of playback. Subjects gave significantly more copulation-solicitation displays in response to playback of songs at 85 dB SPL than to playback of the same songs at 72 dB SPL. The strength of the preference, as judged by the ratio of response to high and low SPL playback (1.15:1), was lower than that of preferences for high SPL shown by insects and anurans. Female red-winged blackbirds responded preferentially to a conspecific song relative to a heterospecific imitation, even when the imitation was presented at an SPL 13 dB higher than that of the conspecific song. By contrast, female redwings did not maintain a preference for multiple conspecific song types over single types when the single song types were presented at the higher SPL. These results are compatible with Klump & Gerhardt's (1987) suggestion that the intensity independence of female preferences varies with the relative benefit females obtain from each preference.     

A PERFORMANCE CONSTRAINT ON THE EVOLUTION OF TRILLED VOCALIZATIONS IN A SONGBIRD FAMILY (PASSERIFORMES: EMBERIZIDAE)

Behavioral evolution can be influenced by constraints, for example, of phylogeny and performance. In this paper I describe a pattern in the evolution of birdsongs that may reflect a constraint on vocal performance. Trilled vocalizations from 34 species of songbirds (Passeriformes: Emberizidae) were analyzed. Two acoustic variables, trill rate and frequency bandwidth, were measured for different trill types. In most species, maximal values of frequency bandwidth were found to decrease with increasing trill rates. Further, trills with low trill rates exhibited wide variance in frequency bandwidth, and trills with high trill rates exhibited only narrow frequency bandwidths. The bounded nature of this pattern suggests that performance constraints have limited the evolutionary diversification of trills. In particular, I explore the role of constraints associated with vocal tract modulations during song production and evolution. Identification of this constraint may enhance our ability to explain particular patterns of trill evolution.     

Imitation and Invention in Song Learning in Nightingales (Luscinia megarhynchos B., Turdidae)

Song learning in oscine birds is often defined solely as a process of song imitation; nonetheless, not all songs produced by laboratory‐tutored birds are imitations of the model songs. If song learning were strictly a process of imitation, these non‐imitated songs (inventions) would be expected to contain no learned attributes. To determine whether species‐typical song attributes can be learned in the absence of imitation, we compared the imitations and inventions of laboratory‐tutored nightingales (Luscinia megarhynchos B.) with the songs of wild nightingales and the songs of laboratory‐reared, untutored nightingales. The species‐typical song attribute measured was stereotypy. We quantified stereotypy by four variables: (1) percentage of notes shared between two renditions of the same song type (2) difference in repetition rates of the same trill in two renditions of the same song type (3) acoustic similarity of the same note in two renditions of the same song type, and (4) acoustic similarity of the same note repeated within a trill. Wild songs and imitated songs were significantly more stereotypical than the songs of untutored birds for all measures. For the percentage of notes shared (1), and the acoustic similarity of notes in two renditions of the same song type (3), invented songs did not differ from the songs of untutored birds, suggesting that imitation is necessary for the acquisition of these song characteristics. However, invented songs were significantly more stereotypical than the songs of untutored birds for measures of stereotypy related to trills (2 and 4), and neither imitated nor invented songs differed significantly from the songs of wild birds in terms of trill rate stereotypy (2). Thus, it appears that the process of learning to produce trills may differ from the process of learning non‐repetitive song components: increased stereotypy in trills occurs even when the trills themselves are not copied from song models. Strict imitation does not fully account for the acquisition of some learned song attributes.     

Rock wrens preferentially use song types that improve long distance signal transmission during natural singing bouts

When animals are capable of producing variable signals they may preferentially use some signal types over others. Among songbirds, individuals are known to alter song type form and usage patterns in contest and mating situations, but studies have not examined how song choice improves signal efficacy during broadcast song. For this study we investigated rock wren Salpinctes obsoletus song type use rates during natural singing bouts. We tested three hypotheses for adaptive song use during broadcast song: 1) birds improve signal content by increasing the use of high quality songs, 2) birds optimize for signal propagation by preferentially using songs that transmit well, and 3) birds maintain energy by reducing the use of costly songs. The study included 19 058 songs sung by 12 individuals, each of which had a measured song repertoire of between 52 and 117 song types which were produced at highly variable rates. Results indicated that rock wrens did not preferentially sing song types with shorter durations or fewer frequency switches, as would be expected if they selected song types to minimize delivery costs. They also did not favor songs with more rapid trills or more frequency switches, as would be expected if they adjusted song use primarily to indicate quality. Focal birds did preferentially sing significantly longer songs with lower bandwidths, lower frequencies, and slower trill rates. Results suggest that natural broadcast singing patterns are shaped more by the benefits of long distance transmission than by the benefits of advertising performance ability or the costs of song production.     

Physically Challenging Song Traits,Male Quality,and Reproductive Success in House Wrens

Physically challenging signals are likely to honestly indicate signaler quality. In trilled bird song two physically challenging parameters are vocal deviation (the speed of sound frequency modulation) and trill consistency (how precisely syllables are repeated). As predicted, in several species, they correlate with male quality, are preferred by females, and/or function in male-male signaling. Species may experience different selective pressures on their songs, however; for instance, there may be opposing selection between song complexity and song performance difficulty, such that in species where song complexity is strongly selected, there may not be strong selection on performance-based traits. I tested whether vocal deviation and trill consistency are signals of male quality in house wrens (Troglodytes aedon), a species with complex song structure. Males’ singing ability did not correlate with male quality, except that older males sang with higher trill consistency, and males with more consistent trills responded more aggressively to playback (although a previous study found no effect of stimulus trill consistency on males’ responses to playback). Males singing more challenging songs did not gain in polygyny, extra-pair paternity, or annual reproductive success. Moreover, none of the standard male quality measures I investigated correlated with mating or reproductive success. I conclude that vocal deviation and trill consistency do not signal male quality in this species.     

A Tradeoff Between Performance and Accuracy in Bird Song Learning

Some of nature’s most complex behaviors, such as human speech and oscine bird song, are acquired through imitative learning. Accurate imitative learning tends to preserve patterns of behavior across generations, thus limiting the scope of cultural evolution. Less well studied are the routes by which cultural novelties arise during development, beyond simple copy error. In this study we assess, in a species of songbird, the relationship in song learning between two potentially conflicting learning goals: accuracy in copying and maximization of vocal performance. In our study species, the swamp sparrow (Melospiza georgiana), vocal performance can be defined for a given song type and frequency range by the rate of note repetition (‘trill rate’), with faster trills being more difficult to sing. We trained young swamp sparrows with song models with experimentally modified trill rates and characterized both the accuracy and performance levels of copies. Our main finding is that birds elevated the trill rates of low‐performance models, but at the expense of imitative accuracy. By contrast, birds reproduced normal and high‐performance models with typically high accuracy in structure and timing. Developmental mechanisms that enable songbirds to balance imitative accuracy and vocal performance are likely favored by sexual selection and may help explain some current patterns of variation in birdsong. Such mechanisms may also explain how behaviors that are learned by imitation can nevertheless respond to selection for high‐performance levels in their expression.     

Morphological adaptation influences the evolution of a mating signal

Theory predicts that forces of natural selection can reduce the intensity of sexually selected traits. In this study, I investigate how morphological adaptation to feeding ecology influences a mating signal. In birds, changes in feeding ecology can cause rapid divergence in bill morphology. Because bills are also important for song production, feeding ecology may influence song divergence. During song, birds can rapidly change vocal tract resonance using bill movement, yet are constrained in rate and magnitude of bill movements resulting in a trade-off between trill rate and frequency bandwidth. Male swamp sparrows vary in their ability to produce rapid, broad-band trills and females prefer more physically demanding songs. Populations of swamp sparrows adapted to the feeding ecology of tidal marshes have larger bills than inland populations. Larger bills should increase the constraints of producing rapid, broad-band trills allowing for a test of how changes in feeding ecology affect a feature of song used in mate choice. I found significant differences in acoustic features of song consistent with the hypothesis that coastal males are less able to meet the physical demands of song production because of the constraints of having larger bills. As possible compensation for decreases in song performance, coastal populations exhibit an increase in song complexity. These changes support the current model of how motor constraints influence song production and suggest a mechanism by which feeding ecology can influence signal evolution.     

Song is a reliable signal of bill morphology in Darwin's small tree finch Camarhynchus parvulus, and vocal performance predicts male pairing success

The shape and movement of the vocal tract are known to influence bird song. Current theory predicts that large bill and body size are correlated with low frequency song and slow trill rate. It is also widely accepted that song characteristics are important for mate choice by females. We investigated the relationship between bill morphology, song characteristics, and pairing success in Darwin's small tree finch Camarhynchus parvulus , on the Galapagos Islands. Contrary to predictions from a previous cross-species study on Darwin's finches, we found that individuals with larger bill size produced songs with slow trill rate, high dominant frequency, and broad frequency bandwidth, indicating that song is a reliable signal of bill morphology. Vocal performance as indicated by the deviation from an upper performance limit was higher in paired than unpaired males. Pairing was not skewed in favour of a particular bill size, and both small and large billed males that sang high performance song had high pairing success. The reliable signalling function of song has implications for female choice and territorial defence, given that both females and conspecific competitors can assess the relative size of males' bills through song, while females may use vocal performance as a signal of male quality.     

Inferring performance in the songs of dark-eyed juncos (Junco hyemalis)

Within bird species, songs differ in their attractiveness tofemales or effectiveness in male–male interactions. Somesongs are more difficult to sing than others, and receiversmay use a singer's performance of difficult songs as a meansfor evaluating the quality of the singer. The concept of songperformance aims at quantifying how physiologically demandingare different songs. Using variation between song types of dark-eyedjuncos, Junco hyemalis, we show that some song traits tradeoff with costly aspects of song output—short intervalsbetween syllables or loud sound amplitude—suggesting thatthose traits are difficult to sing. First, after controllingfor other traits, long syllables require longer intervals forrecovery. This supports the idea that a measure of "respiratoryperformance" could be based on the relative lengths of syllablesand intervals. Second, some syllable traits trade off stronglywith sound amplitude, suggesting that these traits may be difficultto sing at high amplitudes. The ratio of frequency bandwidthand trill rate has been used to infer performance in other birdspecies, but we found no evidence that frequency bandwidth tradesoff with any aspect of song output in the junco. The negativeassociation of bandwidth with trill rate may instead be a passiveconsequence of syllable length, with longer syllables randomlyaccumulating frequency modulation. We conclude that bird receiversmay best evaluate how well a song is performed if they integratemultiple cues and discuss how researchers may similarly devisemeasures of song performance.     

Birdsong performance and the evolution of simple (rather than elaborate) sexual signals

Sexual signals are often elaborate as a result of sexual selection for signals of individual quality. Contrary to expectation, however, the elaboration of signals such as birdsong is not related to the strength of sexual selection across species. With a comparative study across wood warblers (family Parulidae), we show a compromise between advertising the performance of trills (syllable repetitions) and song complexity, which can result in the evolution of simple, rather than elaborate, song. Species with higher trill performance evolved simple songs with more extensive trilled syntax. This advertises trill performance but reduces syllable diversity in songs. These two traits are commonly sexually selected in songbirds, but indexes of sexual selection were not related to either in wood warblers. This is consistent with sexual selection targeting different traits in different species, sometimes resulting in simple signals. We conclude that the evolution of sexual signals can be unpredictable when their physiology affords multiple or, as here, opposing ways of advertising individual quality.     

Are good ornaments bad armaments? Male chaffinch perception of songs with varying flourish length

Birdsong functions as both a territorial and a mate-attracting signal, and is therefore subject to inter- and intrasexual selection through male-male competition and female mate choice. Consequently, differences in song perception between males and females could potentially lead to conflicting selection pressures on song structure. Chaffinch,Fringilla coelebs , songs have two distinctive parts: a series of rapidly repeated notes (the trill) followed by a sequence of more complex and often nonrepeated elements (the flourish). Males often spontaneously sing incomplete songs consisting mainly of the trill part, but their communicative function is unclear. We tested the reaction of males to songs with and without the terminal flourish in a dual speaker playback design by measuring their approach to the speakers. Like female chaffinches tested with the same stimuli in a previous study, males responded more to songs with both trill and flourish than to those consisting of a trill only. However, although females had preferred songs with a relatively longer flourish, males showed the strongest reaction (closest approach to the speaker) to songs with longer trills and shorter flourishes. The results suggest that males and females may perceive aspects of chaffinch song structure differently, in which case it could be under conflicting pressure of intra- and intersexual selection. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Habituation,Recovery and the Similarity of Song Types within Repertoires in Red-winged Blackbirds (Agelaius phoeniceus) (Aves,Emberizidae)

Male red-winged blackbirds (Agelaius phoeniceus) respond to playback of conspecific song on their territories with the song spread, a graded aggressive display in which males extend their wings to expose their red epaulets while singing. We show that the intensity of song spread display declines with repeated presentation of one song type, and recovers when song types are switched. Recovery is greater for switches between song types that are acoustically dissimilar than for switches between song types that are acoustically similar. Recovery is no different for switches between two song types taken from the repertoires of different males than for switches between song types recorded from the same male. Analysis of acoustic features also indicates that song types recorded from different males are not more dissimilar than are song types from the same male. Our results do not support the idea that repertoires of red-winged blackbirds are composed of similar song types in order to facilitate individual recognition. Rather, repertoires may be constructed of dissimilar song types, so as to help maintain the response of listeners despite habituation.     

Conservation of multivariate female preference functions and preference mechanisms in three species of trilling field crickets

Divergence in mate recognition systems among closely related species is an important contributor to assortative mating and reproductive isolation. Here, we examine divergence in male song traits and female preference functions in three cricket species with songs consisting of long trills. The shape of female preference functions appears to be mostly conserved across species and follows the predictions from a recent model for song recognition. Multivariate preference profiles, combining the pulse and trill parameters, demonstrate selectivity for conspecific pulse rates and high trill duty cycles. The rules for integration across pulse and trill timescales were identical for all three species. Generally, we find greater divergence in male song traits than in associated female preferences. For pulse rate, we find a strong match between divergent male traits and female peak preferences. Preference functions for trill parameters and carrier frequency are similar between species and show less congruence between signal and preference. Differences among traits in the degree of trait–preference (mis)match may reflect the strength of preferences and the potential for linkage disequilibrium, selective constraints and alternative selective pressures, but appear unrelated to selection for mate recognition per se.     

Temporal stability and change in a microgeographical pattern of song variation in the rufous-collared sparrow

We studied the pattern of song variation in the rufous-collared sparrow, Zonotrichia capensis, in the 1987 and 2000 breeding seasons in a 7-km² area covering open and closed habitats. We measured 14 quantitative song variables over a total of 390 individuals and found consistent differences between habitats in both years. In particular, songs of individuals from closed habitat sites had trills with longer trill intervals and lower frequencies than those of individuals from open habitats. This pattern of variation is interpreted as a song cline that correlates with the environmental gradient. Although this cline was stable in location and shape, it also showed several differences between years. The songs recorded during 2000 had trills with longer trill intervals and lower minimum frequencies than those recorded during 1987. Thus, this study is the first to directly document temporal changes in song variables in a Z. capensis population. This change would be expected if all the habitats studied were more closed in 2000 than in 1987, but we did not find obvious differences in habitat features between seasons. However, this modification of song structure is compatible with alternative explanations, such as subtle differences in habitat characteristics, changes in climatic variables, a delayed effect of a past modification of the environment and neutrality of the change.     

Female Lincoln's sparrows modulate their behavior in response to variation in male song quality

Sexually reproducing organisms should mate with the highest quality individuals that they can. When female songbirds choose a mate, they are thought to use several aspects of male song that reflect his quality. Under resource-limited environmental conditions, male Lincoln's sparrows (Melospiza lincolnii) vary among one another in several aspects of song quality, including song length, song complexity, and trill performance. In a 2-pronged approach, we tested whether variation in song quality of male Lincoln's sparrows influences the behavior of females that are in a reproductive-like state. Over two trials, we exposed females to songs from the high and low ends of the distribution of naturally occurring song quality variation and found a higher level of behavioral activity in females exposed to high-quality songs, especially when they had first been exposed to low-quality songs. We also examined female phonotaxis toward antiphonally played songs with experimentally elevated and reduced trill performance and found that females moved preferentially toward the songs with elevated trill performance. Contrary to most studies investigating the behavioral responses of wild, female songbirds to variation in male song, we obtained our results without administering exogenous estradiol, which can artificially perturb the female's physiology. Our results demonstrate that the behavior of female Lincoln's sparrows is modulated by the quality of male songs to which they are exposed and that trill performance plays a significant role in this behavioral modulation. Furthermore, as the order of song quality presentation matters, it appears that recent song experience also influences female behavior.     

The predictive value of trill performance in a large repertoire songbird,the nightingale Luscinia megarhynchos

In animal communication, elaborate signals have been shown to be under sexual selection and often to reliably indicate a signaler's quality, condition, or motivation. For instance, the performance of physically challenging signals such as trills – i.e. rapidly repeated elements of broad frequency bandwidth – is considered to reflect signaler quality. Nightingales Luscinia megarhynchos are renowned for their outstanding song repertoire sizes, and most songs include a variety of complex trills. In the present study, we examined whether performance of trills can reliably reflect male quality. We show that vocal performance of trills predicts the age of a male. Older males sang trills that were closer to the performance limit than did younger males. Moreover, males with narrower beaks sang more consistent trills than did males with wider beaks. Vocal performance of trills, however, did not significantly predict other measures of biometric quality such as body size or body condition of the males. The findings suggest that receivers could benefit from the predictive value of physically demanding song traits in assessing age as an important quality component of potential mates or rivals. Particularly in species with high singing versatility, signaler assessment based on readily assessable structures may be adaptive, as this will allow receivers to quickly gather relevant information about the singer without attending to the full song repertoire.