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Although experimentally simulating predator presence helps improve sample sizes in studies of free‐ranging animals, few studies have examined whether auditory playbacks and visual models produce similar results. Additionally, it is unclear if anti‐predator strategies are specific to predator hunting styles in understudied Neotropical pitheciid primates, limiting what we can generalize about this phenomenon across this taxonomic order. We conducted predator simulation experiments to assess whether wild Rylands' bald‐faced saki monkeys (Pithecia rylandsi) recognize predators based solely on acoustic cues, exhibit predator‐specific responses to different predator types, and vary responses to presentations in different sensory modes. In our playback experiments, sakis had weak responses to non‐predator control vocalizations compared to jaguar growls and harpy eagle shrieks. In most predator playbacks, subjects' first glance corresponded to the direction from which simulated predators would typically attack (above vs. below). However, although sakis exhibited appropriate movement responses to harpy playbacks (i.e., descending canopy), they exhibited no clear movement patterns when presented with jaguar playbacks. In contrast, jaguar model experiments consistently elicited fast approaches, mobbing‐style responses, and long alarm calling bouts. Thus, if we had relied on playbacks alone, we might have concluded that sakis have only generalized responses to terrestrial ambush predators. In fact, in all variables measured (e.g., latency, number of calls, and response duration), models of both predator species elicited stronger reactions than playbacks. Results indicate that bald‐faced sakis can identify predators based solely on vocalizations, but do not exhibit predator‐specific escape responses to terrestrial predators based on acoustic cues alone. The differential response to playbacks and models calls into question the reliability of using acoustic‐only stimuli to assess the specificity of anti‐predator behavior to predator hunting styles in some primate species.

Many animals use alarm calls in intraspecific communication to warn conspecifics of a predator's presence or to elicit coordinated group responses. However, alarm calls may also be aimed directly at the predator to discourage further pursuit. These 'pursuit-deterrent' signals are particularly important in the presence of ambush predators that rely on stealth to hunt prey. Here, we conducted playback experiments over a 16-month period on radiocollared ocelots, Leopardus pardalis, in Peru using audio stimuli of titi monkey (Callicebus toppini) and saki monkey (Pithecia rylandsi) alarm calls, with nonalarm loud calls as controls. We predicted that, if titi and saki alarm calls function as deterrent signals, then ocelots would move away from the sound source and leave the area following exposure to alarms but not following controls. We tracked ocelots via radiotelemetry for 30 min prior to and 30 min following experiments. At 15 min intervals we noted subject location, whether the cat was stationary or moving towards, away from or parallel to the playback area (calculated using a deflection angle) and distance travelled. Results showed a significantly different pattern in response movement between playback trials; ocelots moved away from the sound source in the majority of alarm trials but remained stationary/hidden or moved in a variety of directions following control trials. Ocelots also moved significantly farther following exposure to alarm trials than following exposure to controls. We conclude that ocelots can distinguish alarm calls from other loud calls and are deterred by alarm-calling monkeys. This is the first study to use playbacks on wild predators to test the pursuit-deterrent function of primate alarm calls.

Interspecific hybridization allows the introgression or movement of alleles from one genome to another. While some genomic regions freely exchange alleles during hybridization, loci associated with reproductive isolation do not intermix. In many model organisms, the X chromosome displays limited introgression compared to autosomes owing to the presence of multiple loci associated with hybrid sterility or inviability (the “large X-effect”). Similarly, if hybrids are produced, the heterogametic sex is usually inviable or sterile, a pattern known as Haldane’s rule. We analyzed the patterns of introgression of genetic markers located in the mitochondrial (control region) and nuclear (autosomal microsatellites and sex chromosome genes) genomes of two howler monkey species (Alouatta palliata and A. pigra) that form a natural hybrid zone in southern Mexico, to evaluate whether the large X-effect and Haldane’s rule affect the outcomes of hybridization between these sister species. To identify the level of admixture of each individual in the hybrid zone (N = 254) we analyzed individuals sampled outside the hybrid zone (109 A. pigra and 39 A. palliata) to determine allele frequencies of parental species and estimated a hybrid index based on nuclear markers. We then performed a cline analysis using individuals in the hybrid zone to determine patterns of introgression for each locus. Our analyses show that although the hybrid zone is bimodal (with no known F1 s and few recent generation hybrids) and quite narrow, there has been extensive introgression in both directions, and there is a large array of admixed individuals in the hybrid zone. Mitochondrial and most autosomal markers showed bidirectional introgression, but some had biased introgression toward one species or the other. All markers on the sex chromosomes and a few autosomal markers showed highly restricted introgression. This pattern is consistent with the hypothesis that the sex chromosomes make a disproportionate contribution to reproductive isolation, and our results broaden the taxonomic representation of these patterns across animal taxa.

Objectives When closely related species overlap geographically, selection may favor species-specific mate recognition traits to avoid hybridization costs. Conversely, the need to recognize potential same-sex rivals may select for lower specificity, creating the possibility that selection in one domain constrains evolution in the other. Despite a wealth of data on mate recognition, studies addressing rival recognition between hybridizing species are limited to a few bird species. Using naive populations, we examine the extent to which failed rival recognition might have affected hybridization patterns when two species of howler monkeys (Alouatta pigra and A. palliata) first met after diverging in allopatry. Methods We simulated first contact between naive subjects using playback experiments in allopatric populations of the two purebred species. Using linear mixed models, we compared their look, move, and vocal responses to conspecific and heterospecific loud calls. Results Although not different in overall response strength to playbacks, the two species differed in reaction to heterospecific callers. Male A. pigra ignored calls from male A. palliata, but the reverse was not true. Discussion Despite striking differences in vocalizations, A. palliata respond equally to calls from both species whereas A. pigra respond only to conspecifics. This apparent failure of A. pigra males to recognize interspecific rivals might have biased hybridization (F1 hybrids = male A. palliata x female A. pigra), a pattern previously hypothesized based on genetic analysis of hybrids. Given that A. pigra males could be losing reproductive opportunities to heterospecific males, our findings add to growing evidence of potential costs for overly specific species recognition.

The relatively fixed vocal structure of nonhuman primates stands in stark contrast to humans. However, primate vocal–plasticity studies are particularly limited by ethical and logistic constraints. As an alternative approach, we take advantage of a confirmed howler hybrid zone (Alouatta palliata × A. pigra) to compare the effects of genetic ancestry and experience on vocal variation. Deviations from a tight phenotype–ancestry correlation can indicate potential plasticity. We also tested whether temporal features (e.g., syllable number, calling rate) show more plasticity than acoustic features (e.g., peak frequency, bandwidth) which might be morphologically constrained. Using 29 microsatellite markers, most hybrid male subjects fell at the extremes of the genetic ancestry distribution, consistent with the entire population’s distribution. We then analyzed 182 howling bouts and 231 loud call recordings from 33 male hybrid and purebred subjects from sympatric and allopatric populations. Acoustic features of hybrid calls clustered with calls from genetically similar purebred males, and calls from the only genetically intermediate hybrid in our sample fell between the two acoustic extremes, suggesting a strong effect of ancestry. However, temporal features tended to converge in sympatry (e.g., vocalizations from purebred A. palliata in the hybrid zone were more pigra-like than vocalizations from allopatric A. palliata, and indistinguishable from sympatric palliata-like and intermediate hybrid vocalizations). Our study shows that, at least for temporal features, primate vocal plasticity can extend beyond intrapopulation variation, consistent with a growing body of research suggesting that primate vocalizations may be more flexible than traditionally assumed.

Evolutionary research benefits form the integration of laboratory and field components to determine factors and processes that affect the evolutionary trajectories of species. Our shared interest in understanding hybridization with genetic admixture as a process that may impact social, behavioral, and ecological features of primates, brought us together in a collaborative project aimed at addressing how vocal variation in two species of howler monkeys in Mexico affects and is affected by hybridization. To achieve this goal, we joined our academic expertise in studying primate genetics, ecology, and behavior under different natural and experimental conditions. We took advantage of decades of experience studying and handing wild howler monkeys for translocation projects to safely sample and study wild populations for this project. Here, we describe the history of our collaboration highlighting how our different perspectives, academic realities, and individual strengths built the foundation for our successful collaboration. We also share our perspectives on how this collaboration opened up new academic venues, broadened our individual perspectives on the integration of different research approaches to address a complex topic, and allowed us to recognize the strength of international collaboration.

Fecal glucocorticoid metabolites are increasingly used to investigate physiological stress. However, it is crucial for researchers to simultaneously investigate the effects of reproductive state because estradiol and placental hormones can affect circulating glucocorticoid concentrations. Reports on the relationships between glucocorticoids and reproductive state are inconsistent among females. Unlike several primate species that have heightened glucocorticoid activity during lactation, humans experience reduced glucocorticoid activity during lactation. Rather than a taxonomic difference, we hypothesize that this is a result of different environmental stressors, particularly the threat of infanticide. Here, we expand the number of wild primate species tested by validating a glucocorticoid assay for female Geoffroy's spider monkeys. We investigate the effects of reproductive state on their glucocorticoid concentrations. Utilizing a routine veterinary exam on a captive population, we determined that fecal glucocorticoid metabolites increase in response to a stressor (anesthesia), and this rise is detected approximately 24 hr later. Additionally, we found that extracted hormone patterns in a wild population reflected basic reproductive biology-estradiol concentrations were higher in cycling than lactating females, and in lactating females with older offspring who were presumably resuming their cycle. However, we found that estradiol and glucocorticoid concentrations were significantly correlated in lactating but not cycling females. Similarly, we found that reproductive state and estradiol concentration, but not stage of lactation, predicted glucocorticoid concentrations. Unlike patterns in several other primate species that face a relatively strong threat of infanticide, lactating spider monkeys experience reduced glucocorticoid activity, possibly due to attenuating effects of oxytocin and lower male-initiated aggression than directed at cycling females. More broadly, we conclude that future studies using fecal glucocorticoid metabolites to index stress should consider that reproductive state might confound glucocorticoid measurements. Am. J. Primatol. 77:925-935, 2015. © 2015 Wiley Periodicals, Inc.

Interspecific hybridization allows the introgression or movement of alleles from one genome to another. While some genomic regions freely exchange alleles during hybridization, loci associated with reproductive isolation do not intermix. In many model organisms, the X chromosome displays limited introgression compared to autosomes owing to the presence of multiple loci associated with hybrid sterility or inviability (the “large X-effect”). Similarly, if hybrids are produced, the heterogametic sex is usually inviable or sterile, a pattern known as Haldane’s rule. We analyzed the patterns of introgression of genetic markers located in the mitochondrial (control region) and nuclear (autosomal microsatellites and sex chromosome genes) genomes of two howler monkey species (Alouatta palliata and A. pigra) that form a natural hybrid zone in southern Mexico, to evaluate whether the large X-effect and Haldane’s rule affect the outcomes of hybridization between these sister species. To identify the level of admixture of each individual in the hybrid zone (N = 254) we analyzed individuals sampled outside the hybrid zone (109 A. pigra and 39 A. palliata) to determine allele frequencies of parental species and estimated a hybrid index based on nuclear markers. We then performed a cline analysis using individuals in the hybrid zone to determine patterns of introgression for each locus. Our analyses show that although the hybrid zone is bimodal (with no known F1 s and few recent generation hybrids) and quite narrow, there has been extensive introgression in both directions, and there is a large array of admixed individuals in the hybrid zone. Mitochondrial and most autosomal markers showed bidirectional introgression, but some had biased introgression toward one species or the other. All markers on the sex chromosomes and a few autosomal markers showed highly restricted introgression. This pattern is consistent with the hypothesis that the sex chromosomes make a disproportionate contribution to reproductive isolation, and our results broaden the taxonomic representation of these patterns across animal taxa. Electronic supplementary material The online version of this article (10.1007/s10764-018-0056-4) contains supplementary material, which is available to authorized users.

Social differences between primate species may result from both flexible responses to current conditions or fixed differences across taxa, yet we know little about the relative importance of these factors. Here, we take advantage of a naturally occurring hybrid zone in Tabasco, Mexico to characterize the variation in social structure among two endangered howler monkey species, Alouatta pigra and A. palliata, and their hybrids. Work in pure populations has suggested that A. pigra females maintain closer proximity, exhibit higher rates of affiliation, and lower rates of agonism than A. palliata females, but we do not know what accounts for this difference. Using identical data collection and analysis methods across three populations, we first seek to confirm previously reported interspecific differences in social structure across all sexes. We next examine: (1) how female social relationships changed with ancestry (by comparing pure and hybrid individuals); (2) how female social relationships changed with group size (A. pigra have smaller groups than A. palliata); and (3) whether female social relationships differed between two taxonomic groups within a single forest fragment (thus controlling for ecological variation). We confirmed previously described species differences, including closer proximity among females than among males in all populations. We also found that smaller groups maintained closer proximity. However, even after accounting for variation in group size, A. pigra females had closer proximity and more affiliation than A. palliata females. Furthermore, differences between pigra-like and palliata-like hybrids paralleled differences between pure populations and persisted even after controlling for ecological variation. Together, our results suggest that flexibility cannot account for all of the social differences between A. pigra and A. palliata and indicate an important genetic component in primate social behavior.

Among male chacma baboons (Papio hamadryas ursinus), rank positions in the dominance hierarchy are fiercely contested. Physical fighting is costly but relatively rare in this species. Instead, disputes are frequently resolved using displays that include loud, repetitive “wahoo” (two-syllable bark) vocalizations. We previously found that males of all ranks adjust their contest behavior based on the relative fighting ability of opponents and that length of the second syllable (“hoo” duration), calling rate, and fundamental frequency reliably indicate fighting ability. To test whether males indeed attend to hoo duration when assessing opponents, we designed two sets of playback experiments in which call sequence pairs were identical except for this single modified feature. In experiment 1, we used calls recorded from high-ranking males unfamiliar to all subjects. In experiment 2, callers were familiar rivals that ranked one position below subjects in the dominance hierarchy. In paired analyses, subjects in both experiments responded more strongly to sequences with more intense signal features (most commonly associated with high-quality males) compared to sequences with relatively less exaggerated features (most often associated with low-quality males). Results suggest that males can use acoustic features to both indirectly evaluate strangers and to monitor the changing condition of those rivals that present the biggest intragroup threat to their position in the dominance hierarchy. Taken together with our previous research, baboons appear to follow a classic assessor strategy—signal features related to rank and condition are salient to males and directly affect their propensity to respond to rivals.

In chacma baboons (Papio hamadryas ursinus), adult males and lactating females form preferential associations, or ‘friendships’, that provide protection against potentially infanticidal attacks. Little is known about the mechanisms by which males and females form friendships, or the function of friendships for males. We examined the relationship between mating activity, paternity and friendships over a 3-year period. A male’s proportion of consort activity with a female within and outside of her period of most likely conception, along with his dominance rank, each contributed importantly in models of paternity outcomes and formation of friendships. In 87% of births, the male that monopolized the greatest proportion of a female’s consort days during her last conceptive cycle was the father of her offspring. In 82% of the births where the female’s primary consort partner was present at parturition, he became her friend. When the primary consort partner was absent at parturition, females formed friendships with males that had monopolized the second or third greatest proportion of their previous consort days. Similarly, 78% of males became friends with females with whom they had obtained their greatest consort success. Such a flexible strategy based on probabilistic assessment of paternity may be adaptive for females, given unpredictable fluctuations in male presence within the group. In a mating system with high dominance-based reproductive skew, males may benefit by investing preferentially in those infants that they are most likely to have fathered.

Behavioral flexibility is considered by some to be one of the hallmarks of advanced cognitive ability. One measure of behavioral flexibility is how subjects respond to novel objects. Despite growing interest in comparative cognition, no comparative research on neophilia in wild primates has been conducted. Here, we compare responses to novel objects in wild chacma baboons (Papio ursinus) and geladas (Theropithecus gelada). Baboons and geladas are closely related taxa, yet they differ in their ecology and degree of social tolerance: (1) baboons are habitat and dietary generalists, whereas geladas have one of the most specialized primate diets (90% grass); (2) baboons exhibit an aversion toward extra-group individuals, whereas geladas typically exhibit an attraction toward them. Using subjects of all age and sex classes, we examined responses to three different objects: a plastic doll, a rubber ball, and a metal can. Overall, baboon subjects exhibited stronger responses to the objects (greater neophilia and exploration) than gelada subjects, yet we found no evidence that the geladas were afraid of the objects. Furthermore, baboons interacted with the objects in the same way they might interact with a potential food item. Responses were unrelated to sex, but immatures showed more object exploration than adults. Results corroborate novel object research conducted in captive populations and suggest that baboons and geladas have differences in behavioral flexibility (at least in this cognitive domain) that have been shaped by ecological (rather than social) differences between the two species.

Abstract Group members do not always act cohesively when facing extra-group rivals. When benefits such as group-defence are not monopolizable, it poses an economics problem: who should contribute to public goods and who should freeload? A collective action framework compliments existing theoretical explanations for cooperation, and provides testable hypotheses about group-level behaviour based on individual costs and benefits. Using this approach, we review research on intergroup encounters in non-human primates published over the last 20 years, focusing on participation by different classes of individuals. While food- and mate-defence explain much between-sex variation in participation, rank and reproductive access frequently explain within-sex variation. In some species, individuals may use intergroup interactions to survey potential transfer locations and mating options, which might coincidently intimidate rivals. Experimental evidence suggests that when intergroup dominance is based on relative number of fighters, individual participation still varies with sex, rank, companion behaviour and dependent offspring presence. Relatively few studies have examined how factors such as relationships within and between groups or individual temperament mediate aggression. Long-term studies of multiple habituated groups and methodological advances (e.g., playback experiments) will continue to improve our understanding of how complex group-level patterns are predictable when viewed from an individual perspective.

Fecal samples were collected from black howler monkeys (Alouatta pigra) in north central Belize and analyzed for evidence of endoparasite life stages. At least six types of endoparasites were found in Alouatta pigra fecal samples collected in the Lamanai Archaeological Reserve and the Community Baboon Sanctuary in 1999. These include a digenean trematode, an oxyurid nematode, a strongyle-type nematode, an ascarid presumed to be Ascaris sp., Entamoeba coli and Iodamoeba butschlii. Higher trematode prevalence was found in adult Alouatta pigra compared to juveniles and higher prevalence of nematode larvae in all animals was found in the wet season compared to the dry season.

During group defense, the contribution of female black howler monkeys (Alouatta pigra) may help deter male intruders; however, their involvement during natural intergroup encounters is facultative. Experimental playback trials simulating potentially infanticidal males were used to examine whether a female's reproductive investment and/or her group's relative fighting ability would influence her participation in loud call displays. Female howlers never responded to recordings without alpha male accompaniment, but their response patterns did not simply mimic his. For example, unlike alpha males, females with small, vulnerable offspring were no more likely than females without infants to participate in howling displays during playback trials. Further, although females without any small infants in their group got closer to speakers than females living in groups with small infants, males did not respond in the same way. To artificially simulate different "numeric odds" scenarios, recordings of one or three howling males (simulated intruders) were broadcast to females living in groups with 1-3 resident males. Responses were consistent with the hypothesis that females assessed intergroup fighting ability. As in alpha males, the weakest female responses occurred when the numeric odds were against their group. However, whereas alpha males participated most when numeric odds were in their favor, females had the most intense approach responses when the number of defending and intruding males was equal. Females appeared to use a cost-effective strategy, reserving their assistance for when their participation could have the greatest impact.

We examined aggressive displays among male chacma baboons (Papio ursinus) over a 23-mo period in the Okavango Delta of Botswana. High-ranking males were more likely than middle- or low-ranking males to participate in displays. Regardless of rank, all males were more likely to participate in chases or physical fights if their opponent’s rank was similar to their own. Most chases and fights, including those that led to injuries, were also between similarly-ranked males. The rate of both aggressive displays and approachretreat interactions increased in the weeks before rank reversals, suggesting that rank challenges were preceded by a period when males assessed each others’ competitive ability and/or motivation. Aggressive displays between disparately-ranked opponents occurred most frequently in contests involving resources of high fitness value: the defense of meat, the defense of estrous females, and the protection of infants against infanticidal attacks. Silent displays were more likely to occur in these three contexts than were displays that occurred as part of more slowly escalating interactions, in which opponents first exchanged calls. Results suggest that competitive encounters among male baboons follow patterns predicted by evolutionary game theory.

Free-ranging adult male baboons give loud two-syllable ‘wahoo’ calls during dawn choruses, interactions between groups, when chasing females, and in aggressive interactions with other males. Previous research has shown that the rate and duration of these contest wahoos are correlated with a male’s competitive ability: high-ranking males call more often, call at faster rates, and call for longer bouts than do low-ranking males. Here we report that acoustic features of wahoos also reveal information about male competitive ability. High-ranking males give wahoos with higher fundamental frequencies (F0) and longer ‘hoo’ syllables. Within-subject analyses revealed that, as males fall in rank, the hoo syllables tend to shorten within a period of months. As males age and continue to fall in rank, F0 declines, hoo syllables shorten, and formant dispersion decreases. Independent of age and rank, within bouts of calling F0 declines and hoo syllables become shorter. Because wahoos are often given while males are running or leaping through trees, variation in these acoustic features may function as an indicator of a male’s stamina. The acoustic features of contest wahoos thus potentially allow listeners to assess a male’s competitive ability.

We present results of a 10-year study of free-ranging gray-footed chacma baboons (Papio ursinus griseipes) in the Okavango Delta of Botswana. The majority of deaths among adult females and juveniles were due to predation, while infants were more likely to die of infanticide. There were strong seasonal effects on birth and mortality, with the majority of conceptions occurring during the period of highest rainfall. Mortality due to predation and infanticide was highest during the 3-mo period when flooding was at its peak, when the group was more scattered and constrained to move along predictable routes. The reproductive parameters most likely to be associated with superior competitive ability—interbirth interval and infant growth rates—conferred a slight fitness advantage on high-ranking females. However, it was counterbalanced by the effects of infanticide and predation. Infanticide affected highand low-ranking females more than middle-ranking females, while predation affected females of all ranks relatively equally. As a result, there were few rank-related differences in estimated female lifetime reproductive success.

When assessing a group's overall fighting ability, functional group size (i. e. the number of cooperative members willing to confront opponents) may be more important than actual group size. Despite obvious benefits, group members do not always act in a collective manner. For example, participation by subordinate male black howler monkeys (Alouatta pigra) in natural group-level contests is highly variable. Using playback experiments, we examined whether alpha-subordinate relationships, number of intruders, or age affected subordinate contest participation. Long-term census data were used to estimate the nature and duration of intra-group male associations. Some subordinate males had long-term relationships with the alpha that existed prior to residence in the current group or they were the alpha male's grown sons. Other subordinate males were in more recent associations with the alpha that formed under seemingly antagonistic circumstances (e. g. the deposed alpha or his grown sons). We found that subordinate males in long-term alpha-subordinate associations had stronger howling and approach responses than males in short-term associations. Younger long-term associates had the strongest reactions to simulated intruders, similarly aged shortterm associates rarely responded, and older males in both association categories responded at intermediate levels. We discuss the variable strategies of subordinate male black howlers and suggest that males who were more likely to participate in group-defence might be gaining direct or indirect fitness benefits by group living.

Inter-group encounters among baboons range from peaceful to aggressive. During 23 months we observed 110 inter-group interactions involving four groups of chacma baboons in the Okavango Delta, Botswana. Results supported the hypothesis that male behavior functions to prevent extra-group males from gaining access to sexually receptive females. Males were more likely to chase females in their own group when estrous females were present, and their chases targeted estrous females more often than expected. Males also chased members of the opposing group more when estrous females were present. When estrous females were absent, male displays were shorter in duration, involved fewer participants, were less intense, and were more likely to result in peaceful mingling between groups. The alpha male was the individual most actively involved in inter-group chases and displays, but males of all ranks participated, especially when they were in consort with a female. However, males did not cooperate in group defense. While behavior during encounters was affected by the presence of estrous females, the outcome of encounters was affected by location and rival group identity. Groups were more likely to approach and displace opponents in the core of their range and more likely to retreat in the periphery. Correcting for location, we also found some evidence for an inter-group dominance hierarchy based on the relative number of males.

Adult male baboons (Papio cynocephalus) give loud two-syllable 'wahoo' calls during dawn choruses, interactions between groups, when chasing females, and in aggressive interactions with other males. These 'contest' wahoos are acoustically different from 'alarm' wahoos given to predators. In a study of free-ranging baboons in the Okavango Delta, Botswana, we found no significant correlations between the acoustic features of wahoos and adult male size; however, acoustic features were correlated with male dominance rank, age, and calling bout length. Here we show that other measures of calling behavior also appear to function as honest indicators of stamina and competitive ability. High-ranking males were more likely than middle- or low-ranking males to participate in wahoo bouts. They called at significantly higher rates, and their bouts were longer and contained more calls. All males were significantly more likely to participate in wahoo bouts with another male if their opponent's rank was similar to their own. Bouts involving males of similar ranks were longer, contained more wahoos, and involved calling at higher rates, than other bouts. In contests between males of similar ranks, the subordinate and dominant were equally likely to end the bout, whereas in contests involving males of disparate ranks, subordinates were significantly more likely to end the bout. Bouts involving males of similar rank were significantly more likely than others to escalate and result in physical fighting.

One of the most striking features of howler monkeys’ natural history is their loud call, which gives the genus Alouatta its common name in English. However, the disproportionate focus on functional aspects of those calls has driven attention away from other relevant issues related to their vocal behavior. In this chapter, we review the studies of acoustic structure conducted so far on these peculiar calls, highlighting the variation among and within the species of this genus. The variation we uncover runs against the notion of uniformity among howler monkeys, but we do find that the relationship between loud call structure and phylogeny compliments genetic work in this genus. We also show how the anatomy of howler monkey’s vocal organs can explain the unusual features of their loud calls and possibly the variation found between species, while also pointing to the various gaps that exist in our knowledge regarding the role of the several components of their highly specialized vocal apparatus. Additionally, we review some basic concepts about sound propagation and geographic variation in long-distance communication. Unlike loud calls, we know relatively little about the low-amplitude calls of howler monkeys. Such calls have received a great deal of attention in the literature, particularly in Old World monkeys, because they can offer insights into the social lives of these animals. Because few comparable studies have been conducted on howler monkeys, we propose some lines of future research that we deemed potentially interesting. We conclude with some methodological approaches to recording howler monkey calls in the field and for sharing vocalizations with other researchers.

Beyond the unique sound of howler monkey vocalizations, their vigorous loud calling displays are perplexing given the otherwise sedentary lifestyle of these primates. Here we provide potential explanations for this energetic investment by reviewing all available functional studies conducted to date. We highlight the variation among and even within species when we explore whether male loud calls are used in group cohesion, predator avoidance, attraction of females, or competition with other males or other groups over resources. In the competition scenario, we examine strategies of avoidance versus direct competition and whether contests are focused on defense of space, food, mates, or infants. We suggest that much of the debate surrounding the function of loud calls stems from methodological differences among researchers and from the varied levels of analyses used, although we also demonstrate that studies of form and function can be intertwined. We emphasize the need to examine different call types separately and discuss the role of howling in intragroup male relationships. Finally, we address the understudied role of female loud calling and the potential use of hybrid populations to examine the evolution of species-typical loud calls. We conclude with some practical hints for designing field tests to uncover functional significance.

Fecal glucocorticoid metabolites are increasingly used to investigate physiological stress. However, it is crucial for researchers to simultaneously investigate the effects of reproductive state because estradiol and placental hormones can affect circulating glucocorticoid concentrations. Reports on the relationships between glucocorticoids and reproductive state are inconsistent among females. Unlike several primate species that have heightened glucocorticoid activity during lactation, humans experience reduced glucocorticoid activity during lactation. Rather than a taxonomic difference, we hypothesize that this is a result of different environmental stressors, particularly the threat of infanticide. Here, we expand the number of wild primate species tested by validating a glucocorticoid assay for female Geoffroy's spider monkeys. We investigate the effects of reproductive state on their glucocorticoid concentrations. Utilizing a routine veterinary exam on a captive population, we determined that fecal glucocorticoid metabolites increase in response to a stressor (anesthesia), and this rise is detected approximately 24 hr later. Additionally, we found that extracted hormone patterns in a wild population reflected basic reproductive biology-estradiol concentrations were higher in cycling than lactating females, and in lactating females with older offspring who were presumably resuming their cycle. However, we found that estradiol and glucocorticoid concentrations were significantly correlated in lactating but not cycling females. Similarly, we found that reproductive state and estradiol concentration, but not stage of lactation, predicted glucocorticoid concentrations. Unlike patterns in several other primate species that face a relatively strong threat of infanticide, lactating spider monkeys experience reduced glucocorticoid activity, possibly due to attenuating effects of oxytocin and lower male-initiated aggression than directed at cycling females. More broadly, we conclude that future studies using fecal glucocorticoid metabolites to index stress should consider that reproductive state might confound glucocorticoid measurements. Am. J. Primatol. 77:925-935, 2015. © 2015 Wiley Periodicals, Inc.

Social differences between primate species may result from both flexible responses to current conditions or fixed differences across taxa, yet we know little about the relative importance of these factors. Here, we take advantage of a naturally occurring hybrid zone in Tabasco, Mexico to characterize the variation in social structure among two endangered howler monkey species, Alouatta pigra and A. palliata, and their hybrids. Work in pure populations has suggested that A. pigra females maintain closer proximity, exhibit higher rates of affiliation, and lower rates of agonism than A. palliata females, but we do not know what accounts for this difference. Using identical data collection and analysis methods across three populations, we first seek to confirm previously reported interspecific differences in social structure across all sexes. We next examine: (1) how female social relationships changed with ancestry (by comparing pure and hybrid individuals); (2) how female social relationships changed with group size (A. pigra have smaller groups than A. palliata); and (3) whether female social relationships differed between two taxonomic groups within a single forest fragment (thus controlling for ecological variation). We confirmed previously described species differences, including closer proximity among females than among males in all populations. We also found that smaller groups maintained closer proximity. However, even after accounting for variation in group size, A. pigra females had closer proximity and more affiliation than A. palliata females. Furthermore, differences between pigra-like and palliata-like hybrids paralleled differences between pure populations and persisted even after controlling for ecological variation. Together, our results suggest that flexibility cannot account for all of the social differences between A. pigra and A. palliata and indicate an important genetic component in primate social behavior.

Predation, food competition, and infanticide all negatively impact female reproductive success. Female dominance rank may mitigate these effects, if competitive exclusion allows high-ranking females to gain priority of access to critical food resources. It may also exacerbate them, if low-ranking females are forced to feed or rest in marginal habitats where they are at increased risk. In this chapter, we present data on reproduction, mortality, and female reproductive success from a 10-year study of free-ranging chacma baboons (Papio hamadryas ursinus) in the Okavango Delta of Botswana and examine the influence of predation, infanticide, and dominance rank on female reproductive success. Predation appeared to be the cause of most deaths among adult females and juveniles, whereas infanticide was the most likely cause of deaths among infants. Seasonality strongly affected both births and mortality: The majority of conceptions occurred during the period of highest rainfall. Mortality due to predation and infanticide was highest during the 3-month period when flooding was at its peak, most likely because the group was more constrained to move along predictable routes during this time. Those reproductive parameters most likely to be associated with superior competitive ability-interbirth interval and infant growth rates-conferred a slight fitness advantage on high-ranking females. This fitness advantage was counterbalanced, however, by the effects of infanticide and predation. Infanticide affected high- and low-ranking females more than middle-ranking females, while predation affected females of all ranks relatively equally. As a result, there were few rank-related differences in estimated female lifetime reproductive success.