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.     

To trill or not to trill? Territorial response to a heterospecific vocal trait in male collared doves, Streptopelia decaocto

Responding of individuals outside the conspecific range hasbeen largely explained by biases in sensory or cognitive systemstoward particular traits or trait values. More recently, ithas been shown that such responses might occur if individualsstill respond to signal traits that have been lost over time.However, empirical evidence remains scarce. We report a casesupporting the latter mechanism. Phylogenetic analysis suggeststhat the collared dove, Streptopelia decaocto, had lost thetrilled vocalization present in most congeneric taxa. We testedwhether males retained the ability to respond to these trillsin the context of territory defense. We synthesized trilledsongs by inserting trills from the sister species S. roseogriseainto S. decaocto songs. We show that trilled songs yielded higherresponses than did natural conspecific songs, and that the intensityof the response depended on the number of trilled elements.We also show that trilled songs elicited as strong reactionsas frequency modulated coos, which are stronger releasers ofterritorial response than are nonmodulated coos, but are notproduced by every males. Additional tests suggest that the frequencypattern is the most important feature of the trill used by males.However, it is still unknown whether reactions to trilled andfrequency modulated coos have the same perceptual basis. Toour knowledge, this study is the first report of a stronglydeviating signal that is still effective in vocal intrasexualcommunication in birds.     

体征对灰眉岩鹀鸣唱特征的影响

声信号是鸟类交流的重要方式。鸟类的鸣声特征受到生境的塑造和性选择,也受到发声器官的影响,如种间的比较研究发现,体重大的鸟类鸣声频率低,喙短的鸟类鸣声频宽大。在种内是否也存在类似的规律,目前的结论还不一致。2012年6月,在北京小龙门地区(40°00'N,115°26'E)测量了17只灰眉岩鹀(Emberiza godlewskii)的形态参数以及鸣声的频率特征,未发现体重或喙长与鸣声的最高频率、最低频率、峰频或频宽存在显著的相关性。本研究的结果支持在小型雀形目鸟类中,体型对繁殖期鸣唱频率特征的影响不显著。     

Social Signals and Behaviors of Adult Alligators and Crocodiles

We compare and contrast the signalling systems and social behaviorsof Alligator mississippiensis, Crocodylus aculus, and Crocodylusmloticus. Our qualitative analysis focuses primarily on thebehaviors of adults during three phases of reproduction: I.Defense of Territory and Courtship, II. Nesting and Incubation,and III. Hatching and Post Hatching. Signals and signal elementsare very similar among the three species. For example, all havevocal, non-vocal acoustic, and visual signals, some transmittedthrough air or water and others through both media. In addition,each species' repertoire is composed of discrete, graded andcomplex signals. A few signals are unique to each species. However,their signalling systems differ in the temporal organizationof the behaviors, and in the relative frequency in which certainfunctional groups of signals occuror in which signals occurin a particular sensory mode. Apparently, the signalling systemsof C. acutus and C. niloticus are more similar to each otherthan either is to the signalling system of A. mississippiensis.The signalling systems of the crocodile species appear to beadapted to open water habitats in which visual signals are advantageousand to high density breeding groups and post-copulatory intersexualcontact. In contrast, the Alligatorsignalling system appearsadapted to a marsh habitat in which vocal signals are likelyfavored and to low density breeding groups.     

Physiological model analysis of involuntary human-voice tremor

Frequency variations in the human voice result from voluntary and involuntary changes in the parameters of the vocal system. The present work deals with involuntary frequency perturbations from two theoretical aspects: 1) the influence of pitch period variations on frequency changes in the band-limited signal which results from the resonant characteristics of the vocal tract; 2) the physiological parameters of the vocal system which are potentially able to govern involuntary frequency changes. It is shown that the modulation function of the vocal-cord wave can theoretically be derived from its harmonics using FM demodulation techniques, and that higher distortion may appear at higher harmonics. It is also shown that involuntary geometrical changes of the vocal tract and its terminal impedance as well as tension and initialarea changes of the vocal cord—changes well within the physiological range—can influence frequency changes in the human voice. The present results are correlated with our reported experimental findings on involuntary voice tremor, used in psychological stress evaluation. The role of the central nervous system, and possible mechanisms for these phenomena, are discussed.     

Song discrimination suggests premating isolation among sympatric indigobird species and host races

Indigobirds (Vidua spp.) are host-specific brood parasites thathave diversified in a recent radiation apparently driven byhost colonization. Behavioral imprinting of both male and femaleindigobirds on host song is thought to promote rapid speciationbecause it results in assortative mating between indigobirdsassociated with a particular host. We conducted a song playbackexperiment to test whether male indigobirds discriminate amongpotential competitors based on song. Of particular interestwas the behavior of two sympatric host races of the Cameroonindigobird Vidua camerunensis that differ only in host songmimicry and other components of their vocal repertoires. Territorialmales of the two V. camerunensis host races and Vidua raricola,a morphologically distinct indigobird species, were tested withplaybacks of each other's songs. Males of all three groups respondedmost aggressively to songs of their own species and/or hostrace, as evidenced by strong and highly significant differencesin a variety of response variables. This differential territorialdefense suggests that an intruding male with different songsdoes not represent a competitive threat and is expected if femalesmate assortatively with respect to song. Thus, our results provideevidence of premating reproductive isolation among recentlyevolved indigobird species and host races.     

Song Frequency Does Not Reflect Differences in Body Size among Males in Two Oscine Species

Large animals, having large vocal organs, produce low sound frequencies more efficiently. Accordingly, the frequency of vocalizations is often negatively related to body size across species, and also among individuals of many species, including several non‐oscine birds (non‐songbirds). Little is known about whether song frequency reveals information about body size within oscine species, which are characterized by song learning and large repertoires. We asked whether song frequency is related to body size in two oscines that differ in repertoire size: the dark‐eyed junco (Junco hyemalis) and the serin (Serinus serinus). We also asked whether the extent to which receivers sample repertoires might influence the reliability of their assessment of body size. We found that none of the frequency traits of song that we investigated was related to male body size, nor did more extensive sampling of repertoires lead to any relationship between frequency and body size. Possible reasons for these results are the small range of variation in size within species, or the elaborate vocal physiology of oscines that gives them great control over a wide frequency range. We discuss these results as they relate to female preferences for high‐frequency song that have been previously reported for oscine species.     

Effects of territorial intrusions on eavesdropping neighbors: communication networks in nightingales

Animal communication often occurs in communication networksin which multiple signalers and receivers are within signalingrange of each other. In such networks, individuals can obtaininformation on the quality and motivation of territorial neighborsby eavesdropping on their signaling interactions. In songbirds,extracting information from interactions involving neighborsis thought to be an important factor in the evolution of strategiesof territory defense. In a playback experiment with radio-taggednightingales Luscinia megarhynchos we here demonstrate thatterritorial males use their familiar neighbors' performancein a vocal interaction with an unfamiliar intruder as a standardfor their own response. Males were attracted by a vocal interactionbetween their neighbor and a simulated stranger and intrudedinto the neighbor's territory. The more intensely the neighborhad interacted with playback, the earlier the intrusions weremade, indicating that males eavesdropped on the vocal contestinvolving a neighbor. However, males never intruded when wehad simulated by a second playback that the intruder had retreatedand sang outside the neighbor's territory. These results suggestthat territorial males use their neighbors' singing behavioras an early warning system when territorial integrity is threatened.Simultaneous responses by neighboring males towards unfamiliarrivals are likely to be beneficial to the individuals in maintainingterritorial integrity.     

Mate choice in lekking sage grouse revisited: the roles of vocal display, female site fidelity, and copying

In lekking sage grouse (Centrocercus urophasianus), femalesexhibit relatively unanimous mate choice for particular males,but a satisfactory explanation for this unanimity has been elusive.We present analyses of mating distributions from two leks over4 years that provide evidence for female choice based on differencesin vocal display performance of males, the locations at whichhens mated in the previous year, and the choices of other females(copying). The unanimity of female choice varied markedly amongleks and years in correlation with changes in the mean numbersof hens that mated at the same time and hence the opportunityto copy. The results confirm that hens assess phenotypic traitsof males directly but also indicate that the secondary tacticsof site fidelity and copying are often important componentsof female choice. The occurrence of these secondary tacticshas three implications: the variance in mating success amonglek males will be a poor predictor of the intensity of sexualselection on specific traits; female preferences may generatemore clustered dispersions of displaying males than predictedby hotspot settlement models; and direct assessment of malesby females may be difficult or costly, a conclusion that supportsadaptive models of sexual selection over a nonadaptive Fisherianprocess. [Behav Ecol 1991;2:165–180]     

Variation among vocalizations of Taraba major (Aves: Thamnophilidae) subspecies

Geographical variation of bird vocalizations may be related to factors influencing sound production and sound propagation. If birds, e.g. the Great Antshrike (Taraba major), produce vocalizations that develop normally in the absence of learning, these variations may reflect evolutionary divergence within species. In this case, vocal variation could be influenced by habitat structure, since abiotic features and vegetal cover affect sound propagation through environment. Selective pressures may be acting on populations in different ways, which could culminate in a process of speciation. Thus, we searched for structural variation of Great Antshrike vocalizations between subspecies and sought for relationships between these vocal variation and environmental structure. We found variations in frequency and time features of vocalizations among subspecies, which are correlated to latitude, elevation and climate. We also observed an increase in vocal differences along with an increase in distances between individuals, which could reflect isolation of subspecies and the vocal adaptation to different environments.     

Predicting Achievable Fundamental Frequency Ranges in Vocalization Across Species

Vocal folds are used as sound sources in various species, but it is unknown how vocal fold morphologies are optimized for different acoustic objectives. Here we identify two main variables affecting range of vocal fold vibration frequency, namely vocal fold elongation and tissue fiber stress. A simple vibrating string model is used to predict fundamental frequency ranges across species of different vocal fold sizes. While average fundamental frequency is predominantly determined by vocal fold length (larynx size), range of fundamental frequency is facilitated by (1) laryngeal muscles that control elongation and by (2) nonlinearity in tissue fiber tension. One adaptation that would increase fundamental frequency range is greater freedom in joint rotation or gliding of two cartilages (thyroid and cricoid), so that vocal fold length change is maximized. Alternatively, tissue layers can develop to bear a disproportionate fiber tension (i.e., a ligament with high density collagen fibers), increasing the fundamental frequency range and thereby vocal versatility. The range of fundamental frequency across species is thus not simply one-dimensional, but can be conceptualized as the dependent variable in a multi-dimensional morphospace. In humans, this could allow for variations that could be clinically important for voice therapy and vocal fold repair. Alternative solutions could also have importance in vocal training for singing and other highly-skilled vocalizations.     

Signature information and individual recognition in the isolation calls of Amazonian manatees,Trichechus inunguis (Mammalia: Sirenia)

Acoustic signals are assumed to form the basis of manatee communication. Empirical evidence of individual vocal recognition has been reported. If manatees can recognize one another by acoustical means, it should be possible to identify individual vocal patterns. We recorded vocalizations of 14 individually housed Amazonian manatees and then digitized selected vocalizations, allowing seven variables to be measured and subjected to multivariate statistical treatment. Discriminant function analysis indicated that individuals can be separated on the basis of variables related to the fundamental frequency and signal duration. We observed significant differences in the vocal patterns between sexes and age classes. Females tended to have greater fundamental frequency and shorter note duration than males. Calves had shorter note durations and greater values for the fundamental frequency range than subadults and adults. An inverse relationship between total body length and fundamental frequency range suggests that the fundamental frequency becomes more defined as the animal ages. The similar individual patterns in the vocalizations of a mother and calf pair are discussed. Individual recognition by Amazonian manatees according to their vocal patterns is suggested through a preliminary playback experiment.     

The success of alternative reproductive tactics in monogyne populations of the ant Solenopsis invicta: significance for transitions in social organization

Newly produced queens from monogyne (single-queen) coloniesof the ant Solenopsis invicta usually initiate reproductionindependently, that is, without worker assistance. However,some recently mated queens attempt to bypass this risky phaseof new colony foundation by entering established nests to reproduce,although it is unclear how often these queens are successfulin natural populations. We surveyed a mature monogyne populationof S. invicta in both 1995 and 1996 for colonies headed by queensincapable of independent colony founding (diploid-male-producingqueens) in order to estimate the frequency of colonies thatare headed by queens that initiated reproduction within establishednests (adopted queens). Using the frequency of diploid-male-producingqueens among the recently mated queens in this population, weestimated that the overall rate of queen replacement by adoptedqueens is about 0.7% per colony per year. Although theory suggeststhat a change to a novel queen reproductive tactic could beassociated with a fundamental change in social organization(queen number), this does not appear to be the case in monogyneS. invicta. However, the evolution of nest-infiltrating reproductivetactics by queens in a monogyne population and the evolutionof multiple-queen societies may result from similar ecologicalpressures facing newly mated queens. We therefore incorporatethis strategy into an existing theoretical framework that wasdeveloped to explain the evolution of alternative social organizationsin ants, providing testable predictions regarding the distributionand frequency of queen adoption in other single-queen ant societies.     

Adapted to roar: functional morphology of tiger and lion vocal folds

Vocal production requires active control of the respiratory system, larynx and vocal tract. Vocal sounds in mammals are produced by flow-induced vocal fold oscillation, which requires vocal fold tissue that can sustain the mechanical stress during phonation. Our understanding of the relationship between morphology and vocal function of vocal folds is very limited. Here we tested the hypothesis that vocal fold morphology and viscoelastic properties allow a prediction of fundamental frequency range of sounds that can be produced, and minimal lung pressure necessary to initiate phonation. We tested the hypothesis in lions and tigers who are well-known for producing low frequency and very loud roaring sounds that expose vocal folds to large stresses. In histological sections, we found that the Panthera vocal fold lamina propria consists of a lateral region with adipocytes embedded in a network of collagen and elastin fibers and hyaluronan. There is also a medial region that contains only fibrous proteins and hyaluronan but no fat cells. Young's moduli range between 10 and 2000 kPa for strains up to 60%. Shear moduli ranged between 0.1 and 2 kPa and differed between layers. Biomechanical and morphological data were used to make predictions of fundamental frequency and subglottal pressure ranges. Such predictions agreed well with measurements from natural phonation and phonation of excised larynges, respectively. We assume that fat shapes Panthera vocal folds into an advantageous geometry for phonation and it protects vocal folds. Its primary function is probably not to increase vocal fold mass as suggested previously. The large square-shaped Panthera vocal fold eases phonation onset and thereby extends the dynamic range of the voice.     

Convergence and divergence in Diana monkey vocalizations

Individually distinct vocalizations are widespread among social animals, presumably caused by variation in vocal tract anatomy. A less-explored source of individual variation is due to learned movement patterns of the vocal tract, which can lead to vocal convergence or divergence in social groups. We studied patterns of acoustic similarity in a social call produced by 14 female Diana monkeys (Cercopithecus diana) in two free-ranging groups. Calls showed variability in fundamental frequency contours owing to individual identity and external context. Vocal divergence increased significantly between females during poor visibility and tended to increase in the presence of neighbours. In contrast, vocal convergence increased significantly between females during vocal interactions, because females matched the frequency contour of their own call with another female's preceding call. Our findings demonstrate that these primates have some control over the acoustic fine structure of their most important social vocalization. Vocal convergence and divergence are two opposing processes that enable callers to ensure spatial proximity and social cohesion with other group members.     

Songbird frequency selectivity and temporal resolution vary with sex and season

Many species of songbirds exhibit dramatic seasonal variation in song output. Recent evidence suggests that seasonal changes in auditory processing are coincident with seasonal variation in vocal output. Here, we show, for the first time, that frequency selectivity and temporal resolution of the songbird auditory periphery change seasonally and in a sex-specific manner. Male and female house sparrows (Passer domesticus) did not differ in their frequency sensitivity during the non-breeding season, nor did they differ in their temporal resolution. By contrast, female house sparrows showed enhanced frequency selectivity during the breeding season, which was matched by a concomitant reduction of temporal resolution. However, males failed to show seasonal plasticity in either of these auditory properties. We discuss potential mechanisms generating these seasonal patterns and the implications of sex-specific seasonal changes in auditory processing for vocal communication.     

Long-distance communication of acoustic cues to social identity in African elephants

Research on long-distance vocal communication in mammals has tended to focus on the maximum distances over which a vocal signal might be physically detectable. For example, because elephants and some whales communicate using infrasonic calls, and low frequencies are particularly resilient to attenuation, it has often been assumed that these species can communicate over very long distances. However, a wide range of acoustic characteristics typically carry information on individual identity in mammalian calls, and frequency components crucial for social recognition could be distorted or lost as distance from the source increases. We used long-distance playback experiments to show that female African elephants, Loxodonta africana, can recognize a contact call as belonging to a family or bond group member over distances of 2.5 km, but that recognition is more usually achieved over distances of 1-1.5 km. We analysed female contact calls to distinguish source- and filter-related vocal characteristics that have the potential to code individual identity, and rerecorded contact calls 0.5-3.0 km from the loudspeaker to determine how different frequencies persist with distance. Our analyses suggest that the most important frequency components for long-distance communication of social identity may be well above the infrasonic range. When frequency components around 115 Hz become immersed in background noise, once propagation distances exceed 1 km, abilities for long-distance social recognition become limited. Our results indicate that the possession of an unusually long vocal filter, which appears to incorporate the trunk, may be a more important attribute for long-distance signalling in female African elephants than the ability to produce infrasound. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Learned Vocal Variation Is Associated with Abrupt Cryptic Genetic Change in a Parrot Species Complex

Contact zones between subspecies or closely related species offer valuable insights into speciation processes. A typical feature of such zones is the presence of clinal variation in multiple traits. The nature of these traits and the concordance among clines are expected to influence whether and how quickly speciation will proceed. Learned signals, such as vocalizations in species having vocal learning (e.g. humans, many birds, bats and cetaceans), can exhibit rapid change and may accelerate reproductive isolation between populations. Therefore, particularly strong concordance among clines in learned signals and population genetic structure may be expected, even among continuous populations in the early stages of speciation. However, empirical evidence for this pattern is often limited because differences in vocalisations between populations are driven by habitat differences or have evolved in allopatry. We tested for this pattern in a unique system where we may be able to separate effects of habitat and evolutionary history. We studied geographic variation in the vocalizations of the crimson rosella (Platycercus elegans) parrot species complex. Parrots are well known for their life-long vocal learning and cognitive abilities. We analysed contact calls across a ca 1300 km transect encompassing populations that differed in neutral genetic markers and plumage colour. We found steep clinal changes in two acoustic variables (fundamental frequency and peak frequency position). The positions of the two clines in vocal traits were concordant with a steep cline in microsatellite-based genetic variation, but were discordant with the steep clines in mtDNA, plumage and habitat. Our study provides new evidence that vocal variation, in a species with vocal learning, can coincide with areas of restricted gene flow across geographically continuous populations. Our results suggest that traits that evolve culturally can be strongly associated with reduced gene flow between populations, and therefore may promote speciation, even in the absence of other barriers.     

Sound Production in the Salientia: Mechanism and Evolution of the Emitter

Frog sounds involve expulsion of air through the larynx. Inmating, release, rain, and territorial calls, the air vibratesvocal cords and/or arytenoid cartilages. Sound is amplifiedand radiated by the distended buccal cavity and vocal sacs.Distress calls are emitted with open mouth, with minimum laryngealmodulation. The trunk is filled by inflation cycles, but airis driven out by synchronized contractions of the body wallmusculature. The pressure levels are more than five times thoseduring ventilation. In the release call of Bufo valliceps the dilatators and constrictorsof the larynx fire simultaneously keeping the larynx closed.As the pulmonary pressure reaches a peak they cease firing.The arytenoids then separate and vibrate, as do the vocal cords.The dilatators terminate the sound pulse by pulling vibratorsout of the air stream, hence the very sharp termination. Prolongedrelease call sequences include interpulse Teinflations thatreturn air from buccal cavity to lung. Frogs apparently evolved from amphibians too small to use aspirationbreathing. Vocalization represented a critical factor in theirsocial organization and its importance locked these animalsinto reliance upon pulse-pumping rather than the more efficientaspiration breathing.     

Shift down,look up: A test of the non‐linearity and fear hypothesis in a non‐vocal skink

The non‐linearity and fear hypothesis predicts that certain non‐linear sounds are one way to evoke antipredator responses in both birds and mammals. This hypothesis, however, has not been studied in non‐vocal species or in reptiles. Such a study would be important because if non‐linear sounds are evocative even in a species that does not produce sounds, then there may be generally salient cues of risk in these sounds. We asked whether non‐vocal lizards, white‐bellied copper‐striped skinks (Emoia cyanura), respond to experimentally broadcast non‐linearities. This species is ideal to ask the question in because prior research has shown that they respond to predator sounds and alarm calls of other species even though they are not vocal. We conducted playback experiments with three computer‐generated simulated non‐linearities to assess whether or not skinks increased antipredator behavior after hearing them. We controlled for novelty by broadcasting a 3‐kHz, 500‐ms pure tone and tropical kingbird (Tyrannus melancholicus) song. Our treatments consisted of a 3‐kHz, 400‐ms pure tone followed by a frequency shift up to 5‐kHz for 100‐ms, a 3‐kHz, 400‐ms pure tone to frequency shift down to 1‐kHz for 100‐ms, and a pure tone followed by 100‐ms of white noise. Following a total of 222 playbacks, we categorized responses into looking, locomotion, and high locomotion, focusing on how skinks changed their rates of time allocation from baseline. We examined 95% confidence intervals to identify whether skinks responded to playbacks and fitted general linear models followed by pairwise comparisons to ask whether skinks discriminated between broadcast stimuli. We found that skinks were especially responsive to frequency downshifts: They significantly increased looking and locomotion, consistent with our predictions based on the non‐linearity and fear hypothesis. Surprisingly, they decreased rates of looking behavior after hearing frequency upshifts, possibly suggesting an increase in relaxed behavior. While skinks responded to noise by increasing their rate of locomotion, this response was not significantly different from controls. We conclude that skinks increase antipredator behavior after hearing downshifts more than any other type of non‐linearity. This provides some support for the non‐linearity and fear hypothesis; even non‐vocal species may respond fearfully to specific types of non‐linear sounds.