The acoustic structure of spider monkey (Ateles geoffroyi) calls is related both to caller goal and arousal.

Analyzing the factors related to the acoustic structure of primate calls is fundamental to identifying the potential information these signals convey, and therefore, their potential function. It is well-accepted that caller arousal is an important driver of acoustic variation in call structure. However, evidence suggests that arousal alone cannot explain the production of calls with different acoustic structures. The "caller goal" framework posits that a call type (e.g., bark and chitter) is a signal which evolved to elicit specific changes on receiver behavior, and a caller's goal, not its arousal state, determines the basic acoustic structure of animal calls. To get a better insight into the effects of caller goal and arousal on the acoustic structure of primate calls, we analyzed the acoustic variation of 382 calls produced by 27 free-ranging spider monkeys (Ateles geoffroyi) in six call contexts representing three caller goals (establishing contact, engage in aggression, and predator presence) across two arousal states (high and low). Discriminant function analyses and linear mixed models showed a strong effect of caller goal on calls' acoustic structure and supported the effect of arousal state on acoustic variation; misclassified calls mainly occurred between the same caller goal categories (e.g., alarm low arousal and alarm high arousal), and acoustic differences between arousal contexts showed the same pattern across the three caller goals (e.g., low center frequency [kHz] values in high arousal contexts). Our results supported the view that caller goal is the primary driver of acoustic difference and showed the effects of caller arousal on other structural differences.

Isolated catatonia-like executive dysfunction in mice with forebrain-specific loss of myelin integrity.

A key feature of advanced brain aging includes structural defects of intracortical myelin that are associated with secondary neuroinflammation. A similar pathology is seen in specific myelin mutant mice that model 'advanced brain aging' and exhibit a range of behavioral abnormalities. However, the cognitive assessment of these mutants is problematic because myelin-dependent motor-sensory functions are required for quantitative behavioral readouts. To better understand the role of cortical myelin integrity for higher brain functions, we generated mice lacking Plp1, encoding the major integral myelin membrane protein, selectively in ventricular zone stem cells of the mouse forebrain. In contrast to conventional Plp1 null mutants, subtle myelin defects were restricted to the cortex, hippocampus, and underlying callosal tracts. Moreover, forebrain-specific Plp1 mutants exhibited no defects of basic motor-sensory performance at any age tested. Surprisingly, several behavioral alterations reported for conventional Plp1 null mice (Gould et al., 2018) were absent and even social interactions appeared normal. However, with novel behavioral paradigms, we determined catatonia-like symptoms and isolated executive dysfunction in both genders. This suggests that loss of myelin integrity has an impact on cortical connectivity and underlies specific defects of executive function. These observations are likewise relevant for human neuropsychiatric conditions and other myelin-related diseases.

Introducing the Software CASE (Cluster and Analyze Sound Events) by Comparing Different Clustering Methods and Audio Transformation Techniques Using Animal Vocalizations.

Unsupervised clustering algorithms are widely used in ecology and conservation to classify animal sounds, but also offer several advantages in basic bioacoustics research. Consequently, it is important to overcome the existing challenges. A common practice is extracting the acoustic features of vocalizations one-dimensionally, only extracting an average value for a given feature for the entire vocalization. With frequency-modulated vocalizations, whose acoustic features can change over time, this can lead to insufficient characterization. Whether the necessary parameters have been set correctly and the obtained clustering result reliably classifies the vocalizations subsequently often remains unclear. The presented software, CASE, is intended to overcome these challenges. Established and new unsupervised clustering methods (community detection, affinity propagation, HDBSCAN, and fuzzy clustering) are tested in combination with various classifiers (k-nearest neighbor, dynamic time-warping, and cross-correlation) using differently transformed animal vocalizations. These methods are compared with predefined clusters to determine their strengths and weaknesses. In addition, a multidimensional data transformation procedure is presented that better represents the course of multiple acoustic features. The results suggest that, especially with frequency-modulated vocalizations, clustering is more applicable with multidimensional feature extraction compared with one-dimensional feature extraction. The characterization and clustering of vocalizations in multidimensional space offer great potential for future bioacoustic studies. The software CASE includes the developed method of multidimensional feature extraction, as well as all used clustering methods. It allows quickly applying several clustering algorithms to one data set to compare their results and to verify their reliability based on their consistency. Moreover, the software CASE determines the optimal values of most of the necessary parameters automatically. To take advantage of these benefits, the software CASE is provided for free download.

High plasticity in marmoset monkey vocal development from infancy to adulthood.

The vocal behavior of human infants undergoes marked changes across their first year while becoming increasingly speech-like. Conversely, vocal development in nonhuman primates has been assumed to be largely predetermined and completed within the first postnatal months. Contradicting this assumption, we found a dichotomy between the development of call features and vocal sequences in marmoset monkeys, suggestive of a role for experience. While changes in call features were related to physical maturation, sequences of and transitions between calls remained flexible until adulthood. As in humans, marmoset vocal behavior developed in stages correlated with motor and social development stages. These findings are evidence for a prolonged phase of plasticity during marmoset vocal development, a crucial primate evolutionary preadaptation for the emergence of vocal learning and speech.

Individual Differences in the Vocal Communication of Malayan Tapirs (Tapirus indicus) Considering Familiarity and Relatedness.

Studies in animal communication have shown that many species have individual distinct calls. These individual distinct vocalizations can play an important role in animal communication because they can carry important information about the age, sex, personality, or social role of the signaler. Although we have good knowledge regarding the importance of individual vocalization in social living mammals, it is less clear to what extent solitary living mammals possess individual distinct vocalizations. We recorded and analyzed the vocalizations of 14 captive adult Malayan tapirs (Tapirus indicus) (six females and eight males) to answer this question. We investigated whether familiarity or relatedness had an influence on call similarity. In addition to sex-related differences, we found significant differences between all subjects, comparable to the individual differences found in highly social living species. Surprisingly, kinship appeared to have no influence on call similarity, whereas familiar subjects exhibited significantly higher similarity in their harmonic calls compared to unfamiliar or related subjects. The results support the view that solitary animals could have individual distinct calls, like highly social animals. Therefore, it is likely that non-social factors, like low visibility, could have an influence on call individuality. The increasing knowledge of their behavior will help to protect this endangered species.

Vocal convergence in a multi-level primate society: insights into the evolution of vocal learning.

The extent to which nonhuman primate vocalizations are amenable to modification through experience is relevant for understanding the substrate from which human speech evolved. We examined the vocal behaviour of Guinea baboons, Papio papio, ranging in the Niokolo Koba National Park in Senegal. Guinea baboons live in a multi-level society, with units nested within parties nested within gangs. We investigated whether the acoustic structure of grunts of 27 male baboons of two gangs varied with party/gang membership and genetic relatedness. Males in this species are philopatric, resulting in increased male relatedness within gangs and parties. Grunts of males that were members of the same social levels were more similar than those of males in different social levels (N = 351 dyads for comparison within and between gangs, and N = 169 dyads within and between parties), but the effect sizes were small. Yet, acoustic similarity did not correlate with genetic relatedness, suggesting that higher amounts of social interactions rather than genetic relatedness promote the observed vocal convergence. We consider this convergence a result of sensory-motor integration and suggest this to be an implicit form of vocal learning shared with humans, in contrast to the goal-directed and intentional explicit form of vocal learning unique to human speech acquisition.

Towards a new taxonomy of primate vocal production learning.

The extent to which vocal learning can be found in nonhuman primates is key to reconstructing the evolution of speech. Regarding the adjustment of vocal output in relation to auditory experience (vocal production learning in the narrow sense), effects on the ontogenetic trajectory of vocal development as well as adjustment to group-specific call features have been found. Yet, a comparison of the vocalizations of different primate genera revealed striking similarities in the structure of calls and repertoires in different species of the same genus, indicating that the structure of nonhuman primate vocalizations is highly conserved. Thus, modifications in relation to experience only appear to be possible within relatively tight species-specific constraints. By contrast, comprehension learning may be extremely rapid and open-ended. In conjunction, these findings corroborate the idea of an ancestral independence of vocal production and auditory comprehension learning. To overcome the futile debate about whether or not vocal production learning can be found in nonhuman primates, we suggest putting the focus on the different mechanisms that may mediate the adjustment of vocal output in response to experience; these mechanisms may include auditory facilitation and learning from success. This article is part of the theme issue 'What can animal communication teach us about human language?'

Insights into the evolution of social systems and species from baboon studies.

Baboons, members of the genus Papio, comprise six closely related species distributed throughout sub-Saharan Africa and southwest Arabia. The species exhibit more ecological flexibility and a wider range of social systems than many other primates. This article summarizes our current knowledge of the natural history of baboons and highlights directions for future research. We suggest that baboons can serve as a valuable model for complex evolutionary processes, such as speciation and hybridization. The evolution of baboons has been heavily shaped by climatic changes and population expansion and fragmentation in the African savanna environment, similar to the processes that acted during human evolution. With accumulating long-term data, and new data from previously understudied species, baboons are ideally suited for investigating the links between sociality, health, longevity and reproductive success. To achieve these aims, we propose a closer integration of studies at the proximate level, including functional genomics, with behavioral and ecological studies.

Conserved alarm calls but rapid auditory learning in monkey responses to novel flying objects.

To evaluate the flexibility in vocal production and comprehension in a non-human primate, we presented a drone to West African green monkeys, Chlorocebus sabaeus. Calls given in response to the drone were clearly distinct from those given to other predators, but highly similar to the aerial alarm calls of the East African vervet monkey, Chlorocebus pygerythrus, suggesting that call structure is conserved. To probe how rapidly the animals attached meaning to the sound of the drone, we played back the drone sound after one to three exposures. Subjects immediately scanned the sky and ran for cover. In contrast to vocal production, comprehension learning was rapid and open-ended.

Acoustic variation of spider monkey (Ateles geoffroyi) contact calls is related to caller isolation and affects listeners' responses.

Group living animals produce vocalizations denominated "contact calls" to maintain contact with out-of-sight group members. These calls have been shown to vary with caller identity and distance to potential listeners. However, it is not clear whether the acoustic variation of contact calls is related to caller social isolation (e.g., inside or outside a subgroup) and listeners' responses that can be helpful to maintain contact. Here, we addressed these questions in spider monkeys (Ateles geoffroyi), a Neotropical primate that exchanges contact calls denominated "whinnies", which show graded variation related to caller immediate behavior and distance between callers. Using 566 whinnies produced by 35 free-ranging adult spider monkeys recorded at ≤ 20 m from microphones, we first analyzed whether the acoustic variation of spontaneous whinnies (i.e., whinnies that are not responses to previous whinnies) is related to caller social isolation or whether acoustic variation is related to the likelihood of eliciting a response whinny from another individual. Secondly, we assessed whether listeners' responses (i.e., time to respond vocally, acoustic characteristics of response whinnies, orienting behaviors) were related to the acoustic variation of previous whinnies. Our study revealed that callers that were outside a subgroup produced whinnies with a lower fundamental frequency (F0), which travels longer distances, and increases the likelihood of producing a response whinny. Moreover, listeners (i.e., responders) responded faster to lower F0 whinnies. However, the acoustic variation (i.e., F0 variation) in response whinnies was better explained by the separation distance between callers, than by the acoustic variation of the previous whinny. Overall, our results suggest that whinny variation facilitates vocal contact to callers that are outside a subgroup, and that context and whinny variation affect listeners' responses.

The postnatal development of ultrasonic vocalization-associated breathing is altered in glycine transporter 2-deficient mice.

KEY POINTS: Newborn mice produce ultrasonic vocalization to communicate with their mother. The neuronal glycine transporter (GlyT2) is required for efficient loading of synaptic vesicles in glycinergic neurons. Mice lacking GlyT2 develop a phenotype that resembles human hyperekplexia and the mice die in the second postnatal week. In the present study, we show that GlyT2-knockout mice do not acquire adult ultrasonic vocalization-associated breathing patterns. Despite the strong impairment of glycinergic inhibition, they can produce sufficient expiratory airflow to produce ultrasonic vocalization. Because mouse ultrasonic vocalization is a valuable read-out in translational research, these data are highly relevant for a broad range of research fields. ABSTRACT: Mouse models are instrumental with respect to determining the genetic basis and neural foundations of breathing regulation. To test the hypothesis that glycinergic synaptic inhibition is required for normal breathing and proper post-inspiratory activity, we analysed breathing and ultrasonic vocalization (USV) patterns in neonatal mice lacking the neuronal glycine transporter (GlyT2). GlyT2-knockout (KO) mice have a profound reduction of glycinergic synaptic currents already at birth, develop a severe motor phenotype and survive only until the second postnatal week. At this stage, GlyT2-KO mice are smaller, have a reduced respiratory rate and still display a neonatal breathing pattern with active expiration for the production of USV. By contrast, wild-type mice acquire different USV-associated breathing patterns that depend on post-inspiratory control of air flow. Nonetheless, USVs per se remain largely indistinguishable between both genotypes. We conclude that GlyT2-KO mice, despite the strong impairment of glycinergic inhibition, can produce sufficient expiratory airflow to produce ultrasonic vocalization.

Baboon vocal repertoires and the evolution of primate vocal diversity.

A remarkable and derived trait of humans is the faculty for language, and considerable research effort has been devoted to understanding the evolution of speech. In contrast to spoken language, which constitutes a (learned) symbolic communication system, the acoustic structure of nonhuman primate vocalizations is largely genetically fixed. Yet, appreciable differences between different genera and species may exist. Environmental conditions, sexual selection, and characteristics of the social system have been invoked to explain these differences. Here, we studied the acoustic variation of call types and vocal repertoires in the genus Papio. Because the genus comprises both stable groups as well as multi-level societies, and reveals striking variation in the degree of aggressiveness from south to north, it constitutes a promising model to assess the link between social system characteristics and vocal communication. We found that, the vocal repertoires of the different species were composed of the same general call types. A quantitative analysis of the acoustic features of the grunts and loud calls of chacma (Papio ursinus), olive (P. anubis), and Guinea (P. papio) baboons showed subtle acoustic differences within call types, however. Social system characteristics did not map onto acoustic variation. We found no correlation between the structure of grunts and geographic distance; the same was true for female loud calls. Only for male loud calls from three populations, call structure varied with geographic distance. Our findings corroborate the view that the structure of nonhuman primate vocalizations is highly conserved, despite the differences in social systems. Apparently, variation in rate and intensity of occurrence of signals, probably due to different behavioral dispositions in species, are sufficient to allow for plasticity at the level of the social relationships, mating patterns, and social organization.

Do sexual calls in female black capuchin monkeys (Sapajus nigritus) vary with fertility? An acoustic analysis.

Females across a range of animal taxa produce vocalizations and signals uniquely associated with periods of mating. While such signals may ultimately function to increase female attractivity to males, conflicting findings challenge the extent to which these signals co-vary in accordance with the probability of conception. Female black capuchin monkeys (Sapajus nigritus) display an elaborate repertoire of both vocal and visual components as part of their socio-sexual behavior, and previous analyses have shown that the rates of production of visual, but not vocal, components provide graded information on female ovulation. It remains possible, however, that the acoustic parameters of these sexual calls, rather than their rate of productions, co-vary with female fertility. To test this, we analyzed structural and temporal call parameters from estrous calls and post-copulatory calls recorded over five consecutive mating seasons in 12 sexually mature females at Iguazú National Park, Argentina. Calls given during the fertile phase of the female ovarian cycle were compared with those given during the non-fertile phase, as determined by profiles of female reproductive hormones. Similarly, within the fertile phase, we tested whether temporal or spectral acoustic parameters of calls gradually change with the approach of ovulation. We did not find any significant relationship between call parameters and the two measures of female fertility in either female estrous calls or post-copulatory calls. However, some differences between pre- and post-copulatory calls were apparent. Overall, our results indicate that sexual calls in black capuchin females do not provide precise information about the timing of ovulation, but may allow listeners to make probabilistic inferences about whether copulations have taken place. This, combined with previous findings, suggests that females in our study may use signals in different modalities to convey information about their fertility and sexual behavior with varying degrees of precision.

Acoustic variation of spider monkeys' contact calls (whinnies) is related to distance between vocalizing individuals and immediate caller behavior.

Due to several factors such as ecological conditions, group size, and social organization, primates frequently spend time out of visual contact with individuals of their own group. Through the use of long-distance vocalizations, often termed "contact calls," primates are able to maintain contact with out-of-sight individuals. Contact calls have been shown to be individually distinct, and reverberation and attenuation provide information about caller distance. It is less clear, however, whether callers actively change the structure of contact calls depending on the distance to the presumed listeners. We studied this question in spider monkeys (Ateles geoffroyi), a species with complex spatial dynamics (fission-fusion society) that produces highly frequency modulated contact calls, denominated "whinnies." We determined the acoustic characteristics of 566 whinnies recorded from 35 free-ranging spider monkeys that belong to a community located in Mexico, and used cluster analyses, discriminant function analyses, and generalized linear mixed models to assess if they varied in relation to the presumed distance to the listener. Whinnies could be grouped into five subtypes. Since the lowest frequency subtype was mainly produced by spider monkeys that exchanged whinnies at longer distances, and lower frequency calls propagate across longer distances, our results suggest that whinnies vary in order to enhance vocal contact between individuals separated by different distances. Our results also revealed that whinnies convey potential information about caller immediate behaviors and corroborated that these calls are individually distinct. Overall, our results suggest that whinny acoustic variation facilitates the maintenance of vocal contact between individuals living in a society with complex spatial dynamics.

Hypersocial behavior and biological redundancy in mice with reduced expression of PSD95 or PSD93.

The postsynaptic density proteins 95 (PSD95) and 93 (PSD93) belong to a family of scaffolding proteins, the membrane-associated guanylate kinases (MAGUKs), which are highly enriched in synapses and responsible for organizing the numerous protein complexes required for synaptic development and plasticity. Genetic studies have associated MAGUKs with diseases like autism and schizophrenia, but knockout mice show severe, complex defects with difficult-to-interpret behavioral abnormalities due to major motor dysfunction which is atypical for psychiatric phenotypes. Therefore, rather than studying loss-of-function mutants, we comprehensively investigated the behavioral consequences of reduced PSD95 expression, using heterozygous PSD95 knockout mice (PSD95+/-). Specifically, we asked whether heterozygous PSD95 deficient mice would exhibit alterations in the processing of social stimuli and social behavior. Additionally, we investigated whether PSD95 and PSD93 would reveal any indication of functional or biological redundancy. Therefore, homozygous and heterozygous PSD93 deficient mice were examined in a similar behavioral battery as PSD95 mutants. We found robust hypersocial behavior in the dyadic interaction test in both PSD95+/- males and females. Additionally, male PSD95+/- mice exhibited higher levels of aggression and territoriality, while female PSD95+/- mice showed increased vocalization upon exposure to an anesthetized female mouse. Both male and female PSD95+/- mice revealed mild hypoactivity in the open field but no obvious motor deficit. Regarding PSD93 mutants, homozygous (but not heterozygous) knockout mice displayed prominent hypersocial behavior comparable to that observed in PSD95+/- mice, despite a more severe motor phenotype, which precluded several behavioral tests or their interpretation. Considering that PSD95 and PSD93 reduction provoke strikingly similar behavioral consequences, we explored a potential substitution effect and found increased PSD93 protein expression in hippocampal synaptic enrichment preparations of PSD95+/- mice. These data suggest that both PSD95 and PSD93 are involved in processing of social stimuli and control of social behavior. This important role may be partly assured by functional/behavioral and biological/biochemical redundancy.

Social interactions and activity patterns of old Barbary macaques: Further insights into the foundations of social selectivity.

Human aging is accompanied by a decrease in social activity and a narrowing in social networks. Studies in nonhuman primates may provide valuable comparative insights in which way aging impacts social life, in the absence of cultural conventions and an awareness of a limited lifetime. For female Barbary macaques at "La Forêt des Singes" in Rocamadour, France, we previously reported an age-associated decrease in active grooming time and network size. Here, we aimed to extend these findings by investigating in which way physical decline, spatial proximity, and aggression vary with age in female Barbary macaques. We analyzed >1,200 hr of focal observations for 46 females aged 5-29 years. As expected, older females engaged less frequently in challenging locomotor activity, such as climbing or running, than younger ones. The previously reported decrease in grooming time was not due to shorter grooming bout duration. Instead, active grooming bouts lasted even longer, which discounts the idea that manual fatigue explains the shift in grooming pattern. We found that older females tended to be spatially reclusive and that they were less frequently the targets of aggression. Although older females showed aggressive behaviors at similar rates as younger females, the proportion of low-level aggression (i.e., threats) increased with age. We suggest that these threats are not simply a signal of dominance, but also function to deter approaches by others. Overall, these findings are in line with the idea that older females aim to avoid potentially negative interactions, specifically if these are costly. In sum, these findings support the idea that shifts in female Barbary macaques' grooming activity, do not simply result from physical deterioration, but are instead due to a higher selectivity in the choice of social partners.

Vocal repertoire of long-finned pilot whales (Globicephala melas) in northern Norway.

The knowledge of the vocal repertoire of pilot whales is very limited. In this paper, the vocal repertoire of long-finned pilot whales recorded during different encounters in the Vestfjord in northern Norway between November 2006 and August 2010 are described. Sounds were analysed using two different methods: (1) an observer-based audio-visual inspection of FFT-derived spectrograms, with which, besides a general variety of clicks, buzzes, nonharmonic sounds, and whistles, 129 different distinct call types and 25 subtypes were distinguished. These call types included pulsed calls and discrete structured whistles varying from simple to highly complex structures composed of several segments and elements. In addition, ultrasonic whistles previously not described for pilot whales were found. In addition to the diversity of single calls, call sequences consisting of repetitions and combinations of specific call types were recorded and (2) a parametric approach that permitted the confirmation of the high variability in pilot whale call structures was developed. It is concluded that the pilot whale vocal repertoire is among the most complex for the mammalian species and the high structural variability, along with call repetitions and combinations, require a closer investigation to judge their importance for vocal communication.

Charting the neglected West: The social system of Guinea baboons.

OBJECTIVES: Primate social systems are remarkably diverse, and thus play a central role in understanding social evolution, including the biological origin of human societies. Although baboons have been prominently featured in this context, historically little was known about the westernmost member of the genus, the Guinea baboon (Papio papio). MATERIAL AND METHODS: Here, we summarize the findings from the first years of observations at the field site CRP Simenti in the Niokolo Koba National Park in Senegal. RESULTS: Guinea baboons reveal a nested multi-level social organization, with reproductive units comprising one "primary" male, one to several females, young, and occasionally "secondary" males at the base of the society. Three to five units form "parties," which team up with other parties to form a "gang." Different gangs have largely overlapping home ranges and agonistic interactions between different parties or gangs are rare. Some but not all strongly socially bonded males are highly related, and population genetic and behavioral evidence indicate female-biased dispersal. Females play an important role in intersexual bond formation and maintenance, and female tenure length varies between a few weeks to several years. DISCUSSION: While the social organization resembles that of hamadryas baboons (P. hamadryas), the social structure differs considerably, specifically in terms of low male aggressiveness and female freedom. Despite substantial differences in social organization and social structure, the acoustic structure of Guinea baboon vocalizations does not differ substantially from that of other baboon taxa. With its multi-level organization, stable bonds between males and females, as well as a high-degree of male-male cooperation and tolerance, Guinea baboons constitute an intriguing model for reconstructing human social evolution.

Does the Structure of Female Rhesus Macaque Coo Calls Reflect Relatedness and/or Familiarity?

In social animals, kin relations strongly shape the social structure of a group. In female-bonded species, maternal relatedness is likely to be mediated via familiarity, but evidence is accumulating that non-human primates are able to recognize kin that they are not familiar with and adjust their behavior accordingly. In playback experiments, female rhesus macaques showed increased interest in 'coo' calls produced by unfamiliar paternal half-sisters compared to 'coo' calls produced by unfamiliar unrelated females, suggesting that these calls should have some common structural characteristics that facilitate the discrimination of kin from non-kin. Here we analyzed 'coo' calls of 67 adult female rhesus macaques from four groups and seven matrilines living on the island of Cayo Santiago (Puerto Rico). We tested whether the call structure of closely maternal and/or paternal related females, as determined from extensive pedigree data, differed from the call structure of unrelated females, while controlling for familiarity (i.e., group-matrilineal membership and age difference) of subjects. In contrast to our expectation, kinship did not predict similarities in 'coo' call structure, whereas 'coo' structure was more similar when produced by females of similar age as well as by females with higher familiarity, suggesting that experience is more decisive than genetic background. The high number of individuals in the analysis and the high accuracy of the assignment of calls to individuals render a lack of power as an unlikely explanation. Thus, based on the results of this study, kin recognition in rhesus monkeys does neither appear to be based on the assessment of self-similarity, nor on the comparison among related subjects (i.e., acoustic phenotype matching), but appears to be mediated by different or multiple cues. Furthermore, the results support the notion that frequent social interactions result in increasing acoustic similarity within largely innate call types ('vocal accommodation').

Motivational Shifts in Aging Monkeys and the Origins of Social Selectivity.

As humans age, they become more selective regarding their personal goals [1] and social partners [2]. Whereas the selectivity in goals has been attributed to losses in resources (e.g., physical strength) [3], the increasing focus on emotionally meaningful partners is, according to socioemotional selectivity theory, driven by the awareness of one's decreasing future lifetime [2]. Similar to humans, aging monkeys show physical losses [4] and reductions in social activity [2, 5-7]. To disentangle a general resource loss and the awareness of decreasing time, we combined field experiments with behavioral observations in a large age-heterogeneous population of Barbary macaques (Macaca sylvanus) at La Forêt des Singes. Novel object tests revealed a loss of interest in the nonsocial environment in early adulthood, which was modulated by the availability of a food reward. Experiments using vocal and visual representations of social partners indicated that monkeys maintained an interest in social stimuli and a preferential interest in friends and socially important individuals into old age. Old females engaged in fewer social interactions, although other group members continued to invest in relationships with them. Consequently, reductions in sociality were not due to a decrease in social interest. In conclusion, some of the motivational shifts observed in aging humans, particularly the increasing focus on social over nonsocial stimuli, may occur in the absence of a limited time perspective and are most likely deeply rooted in primate evolution. Our findings highlight the value of nonhuman primates as valuable models for understanding human aging [8, 9].