Acoustic similarity of flight calls corresponds with the composition and structure of mixed-species flocks of migrating birds: evidence from a three-dimensional microphone array.

Animals that migrate in mixed-species groups may communicate with both conspecific and heterospecific individuals, providing a low-cost mechanism for navigation whenever individuals share similar migratory routes or destinations. Many migratory birds produce calls while flying, but the function of these calls, and the forces contributing to their evolution, are poorly known. We studied flight calls in mixed-species groups of wood warblers (Parulidae), a biodiverse group of migratory songbirds. We used a spatial approach to examine whether acoustic similarity of flight calls varies with group composition, recording flight calls of mixed-species flocks with a wireless microphone array and triangulating the positions of birds in three dimensions. We found that the acoustic similarity of flight calls was correlated with spatial proximity: birds with similar calls fly closer together during migration. We also found relationships between acoustic similarity, flock size and mixed-species flock diversity: birds with similar calls fly in smaller flocks and in flocks with lower species diversity. Our results support the idea that migrating birds use flight calls to maintain contact with acoustically similar individuals in mixed-species flocks, with communication transcending species boundaries. These results suggest that acoustically similar flight calls are used as cues of group assembly for migratory animals. This article is part of the theme issue 'Mixed-species groups and aggregations: shaping ecological and behavioural patterns and processes'.

Cumulative cultural evolution and mechanisms for cultural selection in wild bird songs.

Cumulative cultural evolution, the accumulation of sequential changes within a single socially learned behaviour that results in improved function, is prominent in humans and has been documented in experimental studies of captive animals and managed wild populations. Here, we provide evidence that cumulative cultural evolution has occurred in the learned songs of Savannah sparrows. In a first step, "click trains" replaced "high note clusters" over a period of three decades. We use mathematical modelling to show that this replacement is consistent with the action of selection, rather than drift or frequency-dependent bias. Generations later, young birds elaborated the "click train" song form by adding more clicks. We show that the new songs with more clicks elicit stronger behavioural responses from both males and females. Therefore, we suggest that a combination of social learning, innovation, and sexual selection favoring a specific discrete trait was followed by directional sexual selection that resulted in naturally occurring cumulative cultural evolution in the songs of this wild animal population.

Natal experience and pre-breeding environmental conditions affect lay date plasticity in Savannah Sparrows.

Phenotypic plasticity allows organisms to adjust the timing of life-history events in response to environmental and demographic conditions. Shifts by individuals in the timing of breeding with respect to variation in age and temperature are well documented in nature, and these changes are known to scale to affect population dynamics. However, relatively little is known about how organisms alter phenology in response to other demographic and environmental factors. We investigated how pre-breeding temperature, breeding population density, age, and rainfall in the first month of life influenced the timing and plasticity of lay date in a population of Savannah Sparrows (Passerculus sandwichensis) monitored over 33 yr (1987-2019). Females that experienced warmer pre-breeding temperatures tended to lay eggs earlier, as did older females, but breeding population density had no effect on lay date. Natal precipitation interacted with age to influence lay date plasticity, with females that experienced high precipitation levels as nestlings advancing lay dates more strongly over the course of their lives. We also found evidence for varied pace of life; females that experienced high natal precipitation had shorter lifespans and reduced fecundity, but more nesting attempts over their lifetimes. Rainfall during the nestling period increased through time, while population density and fecundity declined, suggesting that increased precipitation on the breeding grounds may be detrimental to breeding females and ultimately the viability of the population as a whole. Our results suggest that females adjust their laying date in response to pre-breeding temperature, and as they age, while presenting new evidence that environmental conditions during the natal period can affect phenological plasticity and generate downstream, population-level effects.

The evolution of wood warbler flight calls: Species with similar migrations produce acoustically similar calls.

Diverse animal species engage in long-distance migrations. Many migrants travel in groups, and communication within these groups may be important to survival and successful migration. We examined migration and communication in 36 species of wood warblers (Parulidae), songbirds that breed in North America and migrate in mixed-species flocks to their wintering grounds. During migration, wood warblers produce short vocalizations called "flight calls." The function of flight calls and the patterns of acoustic similarity between species are poorly understood. We investigated whether acoustic similarity of flight calls of different species of warbler reflects the similarity in their migratory journeys or their phylogenetic relatedness. We found that phylogeny, similarity in breeding latitude, and overlap in the timing of migration predict acoustic flight call similarity across warbler species. Further, we found that phylogeny, similarity in migration distance, and overlap in wintering range predict acoustic flight call similarity in a subset of 12 species with highly similar calls, although this analysis has a small sample size. We conclude that migratory similarity may be an important force driving the evolution of acoustically similar calls in wood warblers, in addition to phylogenetic relatedness. Acoustic convergence in these species may facilitate communication between individuals with similar migrations.

Sympatry drives colour and song evolution in wood-warblers (Parulidae).

Closely related species often exhibit similarities in appearance and behaviour, yet when related species exist in sympatry, signals may diverge to enhance species recognition. Prior comparative studies provided mixed support for this hypothesis, but the relationship between sympatry and signal divergence is likely nonlinear. Constraints on signal diversity may limit signal divergence, especially when large numbers of species are sympatric. We tested the effect of sympatric overlap on plumage colour and song divergence in wood-warblers (Parulidae), a speciose group with diverse visual and vocal signals. We also tested how number of sympatric species influences signal divergence. Allopatric species pairs had overall greater plumage and song divergence compared to sympatric species pairs. However, among sympatric species pairs, plumage divergence positively related to the degree of sympatric overlap in males and females, while male song bandwidth and syllable rate divergence negatively related to sympatric overlap. In addition, as the number of species in sympatry increased, average signal divergence among sympatric species decreased, which is likely due to constraints on warbler perceptual space and signal diversity. Our findings reveal that sympatry influences signal evolution in warblers, though not always as predicted, and that number of sympatric species can limit sympatry's influence on signal evolution.

Social and physiological drivers of rapid colour change in a tropical toad.

Dynamic sexual dichromatism occurs when males and females differ in colouration for a limited time. Although this trait has been primarily studied in cephalopods, chameleons, and fishes, recent analyses suggest that dynamic dichromatism is prevalent among anurans and may be mediated through sexual selection and sex recognition. Yellow toads, Incilius luetkenii, exhibit dynamic dichromatism during explosive breeding events at the onset of the rainy season: males change from a cryptic brown to a bright yellow and back again during the brief mating event. We tested the hypothesis that dynamic dichromatism in yellow toads is influenced by conspecific interactions and mediated through sex hormones and stress hormones. We placed male toads into one of four social treatments (with three other males, one male, one female, or no other toads). Immediately before and after each one-hour treatment, we quantified male colour with a reflectance spectrometer and we collected a blood sample to assess plasma concentrations of both testosterone and corticosterone. We found that males held with conspecific animals showed the brightest yellow colour and showed little or no change in their corticosterone levels. Across treatments, toads with duller yellow colour had higher levels of corticosterone. Male colour showed no association with testosterone. Interestingly, males showed substantial temporal variation in colour and corticosterone: toads were duller yellow and exhibited greater levels of corticosterone post-treatment across subsequent days at the onset of the rainy season. Our findings reveal that both conspecific interactions and corticosterone are involved in the dynamic colour change of yellow toads.

Social group signatures in hummingbird displays provide evidence of co-occurrence of vocal and visual learning.

Vocal learning, in which animals modify their vocalizations based on social experience, has evolved in several lineages of mammals and birds, including humans. Despite much attention, the question of how this key cognitive trait has evolved remains unanswered. The motor theory for the origin of vocal learning posits that neural centres specialized for vocal learning arose from adjacent areas in the brain devoted to general motor learning. One prediction of this hypothesis is that visual displays that rely on complex motor patterns may also be learned in taxa with vocal learning. While learning of both spoken and gestural languages is well documented in humans, the occurrence of learned visual displays has rarely been examined in non-human animals. We tested for geographical variation consistent with learning of visual displays in long-billed hermits ( Phaethornis longirostris), a lek-mating hummingbird that, like humans, has both learned vocalizations and elaborate visual displays. We found lek-level signatures in both vocal parameters and visual display features, including display element proportions, sequence syntax and fine-scale parameters of elements. This variation was not associated with genetic differentiation between leks. In the absence of genetic differences, geographical variation in vocal signals at small scales is most parsimoniously attributed to learning, suggesting a significant role of social learning in visual display ontogeny. The co-occurrence of learning in vocal and visual displays would be consistent with a parallel evolution of these two signal modalities in this species.

Wild Birds Learn Songs from Experimental Vocal Tutors.

In eight groups of animals, including humans and songbirds, juveniles are understood to learn vocalizations by listening to adults [1-4]. Experimental studies of laboratory-reared animals support this hypothesis for vocal learning [5-7], yet we lack experimental evidence of vocal learning in wild animals. We developed an innovative playback technology involving automated loudspeakers that broadcast songs with distinctive acoustic signatures. We used this technology to simulate vocal tutors in the wild and conducted year-long tutoring sessions to five cohorts of free-living migratory Savannah Sparrows in eastern Canada. We confirm that wild birds learn songs by listening to adult conspecific animals, and we show that they pass these songs on to subsequent generations. Further, we provide the first experimental evidence in the wild that the timing of exposure to tutor song influences vocal learning: wild Savannah Sparrows preferentially learn songs heard during both their natal summer and at the outset of their first breeding season. This research provides direct experimental evidence of song learning by wild animals and shows that wild birds learn songs during two critical stages of development early in life. VIDEO ABSTRACT.

Hot temperatures during the dry season reduce survival of a resident tropical bird.

Understanding how climate change will shape species distributions in the future requires a functional understanding of the demographic responses of animals to their environment. For birds, most of our knowledge of how climate influences population vital rates stems from research in temperate environments, even though most of Earth's avian diversity is concentrated in the tropics. We evaluated effects of Southern Oscillation Index (SOI) and local temperature and rainfall at multiple temporal scales on sex-specific survival of a resident tropical bird, the rufous-and-white wren Thryophilus rufalbus, studied over 15 years in the dry forests of northwestern Costa Rica. We found that annual apparent survival of males was 8% higher than females, more variable over time, and responded more strongly to environmental variation than female survival, which did not vary strongly with SOI or local weather. For males, mean and maximum local temperatures were better predictors of survival than either rainfall or SOI, with high temperatures during the dry season and early wet season negatively influencing survival. These results suggest that, even for species adapted to hot environments, further temperature increases may threaten the persistence of local populations in the absence of distributional shifts.

Parallel evolutionary forces influence the evolution of male and female songs in a tropical songbird.

Given the important role that animal vocalizations play in mate attraction and resource defence, acoustic signals are expected to play a significant role in speciation. Most studies, however, have focused on the acoustic traits of male animals living in the temperate zone. In contrast to temperate environments, in the tropics, it is commonplace for both sexes to produce complex acoustic signals. Therefore, tropical birds offer the opportunity to compare the sexes and provide a more comprehensive understanding of the evolution of animal signals. In this study, we quantified patterns of acoustic variation in Rufous-and-white Wrens (Thryophilus rufalbus) from five populations in Central America. We quantified similarities and differences between male and female songs by comparing the role that acoustic adaptation, cultural isolation and neutral genetic divergence have played in shaping acoustic divergence. We found that males and females showed considerable acoustic variation across populations, although females exhibited greater population divergence than males. Redundancy analysis and partial-redundancy analysis revealed significant relationships between acoustic variation and ecological variables, genetic distance, and geographic distance. Both ambient background noise and geographic distance explained a high proportion of variance for both males and females, suggesting that both acoustic adaptation and cultural isolation influence song. Overall, our results indicate that parallel evolutionary forces act on male and female acoustic signals and highlight the important role that cultural drift and selection play in the evolution of both male and female songs.

Dispersal influences genetic and acoustic spatial structure for both males and females in a tropical songbird.

Animals exhibit diverse dispersal strategies, including sex-biased dispersal, a phenomenon common in vertebrates. Dispersal influences the genetic structure of populations as well as geographic variation in phenotypic traits. Patterns of spatial genetic structure and geographic variation may vary between the sexes whenever males and females exhibit different dispersal behaviors. Here, we examine dispersal, spatial genetic structure, and spatial acoustic structure in Rufous-and-white Wrens, a year-round resident tropical bird. Both sexes sing in this species, allowing us to compare acoustic variation between males and females and examine the relationship between dispersal and song sharing for both sexes. Using a long-term dataset collected over an 11-year period, we used banding data and molecular genetic analyses to quantify natal and breeding dispersal distance in Rufous-and-white Wrens. We quantified song sharing and examined whether sharing varied with dispersal distance, for both males and females. Observational data and molecular genetic analyses indicate that dispersal is female-biased. Females dispersed farther from natal territories than males, and more often between breeding territories than males. Furthermore, females showed no significant spatial genetic structure, consistent with expectations, whereas males showed significant spatial genetic structure. Overall, natal dispersal appears to have more influence than breeding dispersal on spatial genetic structure and spatial acoustic structure, given that the majority of breeding dispersal events resulted in individuals moving only short distances. Song sharing between pairs of same-sex animals decreases with the distance between their territories for both males and females, although males exhibited significantly greater song sharing than females. Lastly, we measured the relationship between natal dispersal distance and song sharing. We found that sons shared fewer songs with their fathers the farther they dispersed from their natal territories, but that song sharing between daughters and mothers was not significantly correlated with natal dispersal distance. Our results reveal cultural differences between the sexes, suggesting a relationship between culture and sex-biased dispersal.

Operant discrimination of relative frequency ratios in black-capped chickadee song.

The two-note fee bee song of the black-capped chickadee (Poecile atricapillus) is sung at many different absolute frequencies, but the relative frequencies, or "pitch ratios", between the start and end of the fee note (glissando) and the fee and the bee notes (inter-note interval) are preserved with each pitch-shift. Ability to perceive these ratios and their relative salience varies with sex of the bird and setting: while both sexes appear to perceive changes in the inter-note interval, males appear to attend to the glissando in the field, and females appear to attend to both ratios. In this study, we compared directly whether male and female chickadees could discriminate between normal fee bee songs and songs that had one or both of the pitch ratios altered, and whether birds attended to one type of alteration over another. Both sexes learned to discriminate normal from altered songs; songs lacking an inter-note interval were more easily discriminated than songs with only the glissando removed. Females performed slightly better than males, including in the most difficult task with the stimuli lacking the glissando. Our study illustrates the value of using perceptual tasks to directly compare performance between the sexes and to demonstrate the difference between perception of and attention to acoustic features of vocal communication.

Male song sparrows have elevated testosterone in response to neighbors versus strangers.

Upon hearing a conspecific signal, animals must assess their relationship with the signaller and respond appropriately. Territorial animals usually respond more aggressively to strangers than neighbors in a phenomenon known as the "dear enemy effect". This phenomenon likely evolved because strangers represent a threat to an animal's territory tenure and parentage, whereas neighbors only represent a threat to an animal's parentage because they already possess a territory (providing territory boundaries are established and stable). Although the dear enemy effect has been widely documented using behavioral response variables, little research has been conducted on the physiological responses of animals to neighbors versus strangers. We sought to investigate whether the dear enemy effect is observed physiologically by exposing territorial male song sparrows (Melospiza melodia) to playback simulating a neighbor or a stranger, and then collecting blood samples to measure plasma testosterone levels. We predicted that song sparrows would exhibit increased testosterone levels after exposure to stranger playback compared to neighbor playback, due to the role testosterone plays in regulating aggression. Contrary to our prediction, we found that song sparrows had higher testosterone levels after exposure to neighbor playback compared to stranger playback. We discuss several explanations for our result, notably that corticosterone may regulate the dear enemy effect in male song sparrows and this may inhibit plasma testosterone. Future studies will benefit from examining corticosterone in addition to testosterone, to better understand the hormonal underpinnings of the dear enemy effect.

The biogeographic and evolutionary history of an endemic clade of Middle American sparrows: Melozone and Aimophila (Aves: Passerellidae).

The large number of endemic species in Middle America is frequently attributed to the interplay of geographical barriers and historical climatic changes in the region. This process promotes genetic divergence between populations, and given enough time, may yield new species. Animals that inhabit mid-elevation or highland habitats may be disproportionately affected in this way. Genetic analyses of animals in this region allow us to better understand how historical patterns of isolation have influenced the generation of new species in this biodiversity hotspot. We studied the biogeography and systematics of two closely related genera of sparrows (Passerellidae): Melozone and Aimophila. Collectively, this group is distributed from the southwestern United States and southward as far as central Costa Rica. We sampled 81 individuals of 8 Melozone and 2 Aimophila species, from 19 localities distributed throughout their ranges. We reconstructed phylogenetic relationships and time-calibrated species trees using multilocus sequence data comprised of one mitochondrial gene and five nuclear genes. We conducted an ancestral area reconstruction analysis to determine the probability of ancestral range at each divergent event. Despite analyzing six loci, we were unable to obtain a fully resolved phylogenetic tree. We recovered four main lineages: lineage 1 includes four Melozone species distributed north of Isthmus of Tehuantepec (M. albicollis, M. crissalis, M. aberti, M. fusca); lineage 2 includes three Melozone species distributed south of the Isthmus of Tehuantepec (M. biarcuata, M. cabanisi, M. leucotis); lineage 3 lineage consists of a single species endemic to the Pacific coast of Mexico (M. kieneri); and lineage 4 includes the more widely distributed sparrows in the genus Aimophila. Our analyses suggest that these genera probably originated during the late Miocene in the Madrean Highlands of southern Mexico. We identified dispersal as the prevalent cause of speciation in this clade with most lineages dispersing to their current distributions from southern Mexico either to the north following a developing and expanding Madro-Tertiary flora, or to the south across the Isthmus of Tehuantepec. A similar pattern of dispersal from this biogeographic region has been reported in other taxa including fishes, reptiles, and birds. Our results reveal that the four lineages identified represent geographically coherent and ecologically similar assemblages of taxa. Finally, when our genetic results are considered, along with apparent differences in morphology and song, the allopatric forms M. b. cabanisi and M. l. occipitalis warrant recognition as biological species.

Vocal production and playback of altered song do not affect ZENK expression in black-capped chickadees (Poecile atricapillus).

The two-note fee bee song of the black-capped chickadee (Poecile atricapillus) is sung at many different absolute frequencies, but the relative frequencies between the start and end of the fee note (the glissando) and between the fee and the bee notes (the inter-note ratio) are preserved regardless of absolute frequency. If these relative frequencies are experimentally manipulated, birds exhibit reduced behavioural responses to playback of altered songs both in field studies and laboratory studies. Interestingly, males appear to be sensitive to alterations in the glissando, while females appear to be sensitive to alterations in both the glissando and the inter-note ratio. In this study, we sought to determine whether the behaviour of male and female chickadees corresponds to differences in zenk protein immunoreactivity (ZENK-ir) in auditory perceptual regions following playback of fee bee songs with typical and altered pitch ratios. Overall, there was a small but significant sex difference in ZENK-ir (females>males), but altering relative frequencies did not reduce ZENK-ir compared to typical song. Birds did vocalize less in response to playback of songs that lacked an inter-note interval, but amount of singing fee bee song, chick-a-dee calls, or gargles was not correlated with ZENK-ir in perceptual regions (caudomedial nidopallium, NCM and caudomedial mesopallium, CMM) or in HVC, which is part of the song system. Our results confirm that ZENK-ir in NCM and CMM is not involved in fine-grain perceptual discrimination, however it did not support the idea that increased vocalizing increases ZENK-ir in HVC.

Dynamic sexual dichromatism produces a sex signal in an explosively breeding Neotropical toad: A model presentation experiment.

Many animals breed in large mating aggregations, where males must rapidly discriminate between prospective mates and rivals. Selection may favour features that facilitate rapid discrimination in these aggregations. The explosively breeding Neotropical Yellow Toad, Incilius luetkenii, exhibits a rapid and dramatic colour change; males change from a cryptic brown to a conspicuous lemon yellow for their brief breeding period. Females, in contrast, remain cryptic brown throughout the year. The function of this temporary, sex-specific colour change is unknown. We tested the hypothesis that yellow colouration displayed by male I. luetkenii facilitates sex recognition during both daytime and nighttime mating aggregations. We created yellow and brown model toads and presented them to males during a breeding event. Male I. luetkenii responded significantly more intensely to brown models compared to yellow models, approaching them and making more amplexus attempts on the brown versus yellow models. This strong pattern held true regardless of ambient light intensity, making this the first study to expose a dynamic colour signal that operates during both day and night. Our results indicate that male I. luetkenii use colouration to quickly discriminate between males and females during their brief, explosive mating aggregations. Our findings suggest that the rapid, dramatic colour change of male I. luetkenii facilitates sex recognition, which could provide a significant fitness advantage to males in the form of reduced energy expenditure and reduced risk of injury by other males. Dynamic dichromatism may provide similar fitness benefits in any organisms that mate in large, competitive aggregations.

Playback responses of socially monogamous black-fronted titi monkeys to simulated solitary and paired intruders.

Many birds and primates use loud vocalizations to mediate agonistic interactions with conspecifics, either as solos by males or females, or as coordinated duets. The extensive variation in duet complexity, the contribution of each sex, and the context in which duets are produced suggest that duets may serve several functions, including territory and mate defense. Titi monkeys (Callicebus spp.) are believed to defend their home range via solo loud calls or coordinated duets. Yet there are remarkably few experimental studies assessing the function of these calls. Observations of interactions between wild established groups and solitary individuals are rare and, therefore, controlled experiments are required to simulate such situations and evaluate the mate and joint territorial defense hypotheses. We conducted playback experiments with three free-ranging groups of habituated black-fronted titi monkeys (Callicebus nigrifrons) to test these hypotheses. We found that titi monkeys responded to the three conspecific playback treatments (duets, female solos, and male solos) and did not respond to the heterospecific control treatment. The monkeys did not show sex-specific responses to solos (N = 12 trials). Partners started to duet together in 79% of their responses to playback-simulated rivals (N = 14 calls in response to playback). Males started to approach the loudspeaker before females regardless of the type of stimulus. The strength of the response of mated pairs to all three conspecific treatments was similar. Overall, our results are consistent with the idea that black-fronted titi monkeys use their loud calls in intergroup communication as a mechanism of joint territorial defense.

Analysis of plumage, morphology, and voice reveals species-level differences between two subspecies of Prevost's Ground-sparrow Melozone biarcuata (Prévost and Des Murs) (Aves: Emberizidae).

Melozone biarcuata (Prevost's Ground-sparrow) has traditionally been divided into two allopatric groups based on differences in vocalizations and plumage characteristics: M. b. cabanisi in Costa Rica and M. b. biarcuata/M. b. hartwegi in northern Central America. However, the relationship between these subspecies has not been studied using a modern taxonomic approach. In this study, our objective was to provide the first detailed taxonomic comparison between these three subspecies using an integrative multi-trait analysis. We analyzed morphometric features, qualitative plumage patterns, and quantitative plumage measurements using spectral reflectance from all three subspecies, and we analyzed vocalizations for subspecies M. b. biarcuata and M. b. cabanisi. Our results show that M. b. cabanisi can be readily distinguished from the two other subspecies on the basis of morphometrics (M. b. cabanisi are smaller), plumage patterns (M. b. cabanisi have different facial markings and plumage patches), color differences (M. b. cabanisi have plumage patches that differ in color and brightness), and vocalizations (M. b. cabanisi have songs and calls that are acoustically distinct from those of M. b. biarcuata). By contrast, the two northern subspecies M. b. biarcuata and M. b. hartwegi were very similar for most traits, supporting previous suggestions that the two northern subspecies should be considered a single subspecies. Our data reveal that the differentiation in phenotypic characteristics between M. b. cabanisi versus M. b. biarcuata and M. b. hartwegi is similar to that reported for other complexes of subspecies where species status has been recognized. We argue that M. b. cabanisi should be treated as a species separate from M. biarcuata and propose that it be called Melozone cabanisi, White-faced Ground-sparrow. Our findings will contribute to the conservation efforts of the White-faced Ground-sparrow, which is endemic to Costa Rica's Central Valley and Turrialba Valley, by bringing focus to conservation policies that preserve ground-sparrow habitat (thickets, shade coffee plantations, and young secondary forest).

Vocal behavior of black-fronted titi monkeys (Callicebus nigrifrons): acoustic properties and behavioral contexts of loud calls.

Loud calls can be heard over long distances due to their high amplitude and low frequency. These calls are commonly used for both within- and between-group communication in many bird and primate species. In the Neotropics, mated pairs of socially monogamous titi monkeys (genus Callicebus) emit conspicuous, coordinated loud calls. These vocalizations appear to play a role in territorial defense, a hypothesis derived from studies of only three of the 31 recognized Callicebus species. Here, we describe the acoustic properties and organization of the loud calls of black-fronted titi monkeys (Callicebus nigrifrons). We compare the behavioral and ecological contexts associated with these vocalizations to investigate their role in within- and between-group communication, resource defense, and mate defense. Black-fronted titi monkeys create loud calls by combining a finite number of syllables to form more complex phrases, which are assembled to compose long sequences of loud calls. Bioacoustic features distinguish the loud calls used in different contexts, involving communication within- and between-groups. We found support for the hypothesis that vocalizations used for between-group communication are cooperative displays used by the mated pair and other group members to regulate access to important food resources, such as fruits. On the other hand, we only found weak support for the mate defense hypothesis.

Uneven sampling and the analysis of vocal performance constraints.

Studies of trilled vocalizations provide a premiere illustration of how performance constraints shape the evolution of mating displays. In trill production, vocal tract mechanics impose a trade-off between syllable repetition rate and frequency bandwidth, with the trade-off most pronounced at higher values of both parameters. Available evidence suggests that trills that simultaneously maximize both traits are more threatening to males or more attractive to females, consistent with a history of sexual selection favoring high-performance trills. Here, we identify a sampling limitation that confounds the detection and description of performance trade-offs. We reassess 70 data sets (from 26 published studies) and show that sampling limitations afflict 63 of these to some degree. Traditional upper-bound regression, which does not control for sampling limitations, detects performance trade-offs in 33 data sets; yet when sampling limitations are controlled, performance trade-offs are detected in only 15. Sampling limitations therefore confound more than half of all performance trade-offs reported using the traditional method. An alternative method that circumvents this sampling limitation, which we explore here, is quantile regression. Our goal is not to question the presence of mechanical trade-offs on trill production but rather to reconsider how these trade-offs can be detected and characterized from acoustic data.