Nonlinear acoustics in pant hoots of common chimpanzees (Pan troglodytes): frequency jumps, subharmonics, biphonation, and deterministic chaos

The pant hoot calls produced by common chimpanzees (Pan troglodytes) are multi-call vocalizations that have figured prominently in investigations of acoustic communication in this species. Although pant hoots are predominantly harmonically structured, they can exhibit an acoustic complexity that has recently been linked to nonlinearity in the vocal-fold dynamics underlying typical mammalian sound production. We examined the occurrence of these sorts of nonlinear phenomena in pant hoot vocalizations, contrasting quieter and lower-pitched "introduction" components with loud and high-pitched "climax" calls in the same bouts. Spectrographic evidence revealed four kinds of nonlinear phenomena, including discrete frequency jumps, subharmonics, biphonation, and deterministic chaos. While these events were virtually never observed during the introduction, they occurred in more than half of the climax calls. Biphonation was by far the most common phenomenon, followed by subharmonics, chaos, and frequency jumps. Individual callers varied in the degree to which their climax calls exhibited nonlinear phenomena, but were consistent in showing more biphonation than other forms. These outcomes show that nonlinear phenomena are routinely present in chimpanzee pant hoots, and help lay the foundation for investigating the function of such events.     

Acoustic variation of pant hoot calls by male chimpanzees: a playback experiment

Pant hoots, a type of long-distance calls of chimpanzees (Pan troglodytes), were played back to two male chimpanzees in a group of seven captive individuals to determine if chimpanzees would modify those vocalizations in response to strange males. Subjects emitted pant hoots with higher rates of delivery, shorter duration of buildup, and lower minimum fundamental frequency of climax when they were presented with pant hoots of strangers than when they produced the calls spontaneously. Considering the direction of acoustical change, we concluded that the rate of delivery, duration of buildup, and minimum frequency of climax might be associated with the underlying emotional states of the callers rather than call matching. Individual difference between two subject males was significant in minimum frequency and duration of climax and in average frequency of call, which appears to reflect differences of the caller's age and social status. These results suggest that different acoustic variables relate to within- and interindividual differences of these vocalizations. Received: March 7, 2000 / Accepted: May 8, 2000     

Phrase structure of wild chimpanzee pant hoots: Patterns of production and interpopulation variability

Recordings and behavioral observations of wild chimpanzees were made over a 2-year period in the Kibale National Park, Uganda (Kanyawara and Ngogo communities) to investigate patterns of acoustic variability in long-distance calls. The phrase structures of pant hoots, the species-typical loud call given predominantly by adult males, were analyzed. Analysis revealed that the build-up phrase was frequently absent from the pant hoots of Kibale chimpanzees. By contrast, analysis of published data on Gombe and Mahale chimpanzees (Tanzania) indicated that these animals typically included the build-up in their calls. These results were interpreted as evidence for phrase-level differences between populations in the acoustic morphology of this compound call. Data on age and sex differences in the context of production of pant hoots were analyzed, and their relevance to the possibility that aspects of pant hoot acoustic morphology are learned is discussed. Adult males initiated pant hoots more than subadult males, and adult males joined other pant hooting individuals with pant hoots more than subadult males did. It is suggested that younger males may pant hoot with specific adult males preferentially and that this may affect the development of their pant hoot acoustic morphologies. A peculiar pant hoot variant previously reported from Gombe, the whimper hoot, is described from Kibale. The production of this call by low-ranking individuals suggests that there are social constraints on pant hooting in the chimpanzee community. Ideas concerning the effect of social relationships on interpopulation variability in vocal signals are briefly discussed. © 1996 Wiley-Liss, Inc.     

Wild Chimpanzees Produce Group‐Specific Calls: a Case for Vocal Learning?

Vocal learning, where animals can modify the structure of their vocalizations as a result of experience, has been found in a range of birds and mammals. Although vocal learning is a fundamental aspect of developing spoken language, there is as yet little evidence that vocal learning occurs in primates. Here we examine whether vocal learning may occur in chimpanzees. We analysed whether wild male chimpanzees, Pan troglodytes verus, of four communities living in a similar habitat in the Taï Forest, Côte d'Ivoire, developed community specific pant hoots. If so, we expected males of three contiguous communities to have distinct pant hoots, while pant hoots of males from a fourth, distant community, located 70 km away, should only differ from those of the contiguous communities by chance. Our analysis confirmed these expectations. In addition, the acoustic distances between the pant hoots of pairs of individuals did not correlate with the genetic relatedness of those pairs, where genetic relatedness was determined using nuclear DNA analysis. Thus, neither habitat nor genetic differences accounted for the observation that there were acoustic differences in the pant hoot structure of males living in neighbouring communities, but not in those of males from a distant community. This suggests that chimpanzees may actively modify pant hoots to be different from their neighbours, providing support for the vocal learning hypothesis.     

Acoustic analysis of wild chimpanzee pant hoots: Do Kibale Forest chimpanzees have an acoustically distinct food arrival pant hoot?

Recordings were made of 193 pant hoots given by 6 adult males during ad lib sampling over a 16-month period in 1988 and 1989. The presence or absence of a let-down phase, and acoustic measures of the let-down and climax phases of the calls, were compared for different call contexts to determine if an acoustically distinct pant hoot was given uniquely upon arrival at fruiting trees. The greatest proportion of pant hoots with a let-down (LD pant hoots) occurred immediately upon arrival at fruiting trees. However, LD pant hoots also occurred at other times during feeding bouts. The frequency and duration of the first exhaled element of 19 letdown phases were measured, and four measures were made on the highest pitched element of 49 climax phases: duration, maximum and minimum frequencies, and average frequency. No differences were found in these acoustic measures that distinguished calls given immediately upon arrival at a food tree from calls given later during feeding bouts. Thus no evidence was found that an acoustically distinct pant hoot was given uniquely upon arrival at fruiting trees. However, the analyses did suggest that identifiable pant hoot variants are given in different social contexts. The proportion of LD pant hoots decreased in more interactive social contexts, and other acoustic features may be available to distinguish this pant hoot variant at long distances. We suggest that different pant hoot variants might broadcast information specific to social, as opposed to ecological, context. © 1993 Wiley-Liss, Inc.     

Buttress drumming by wild chimpanzees: Temporal patterning, phrase integration into loud calls, and preliminary evidence for individual distinctiveness

Wild chimpanzees (Pan troglodytes) generate low-frequency sounds that are audible to humans from a distance of at least 1 km away by hitting the buttresses of trees with their hands and feet. This buttress drumming occurs in discrete bouts of rapidly delivered beats that usually accompany “pant hoots,” the species-specific long-distance vocalization. Individual differences in male chimpanzee (P.t. verus) drumming were investigated during a 6-month field study in the Taï National Park, Ivory Coast. Analysis of drumming bouts recorded from six adult males revealed significant differences between individuals in three acoustic features: (1) mean duration of inter-beat interval; (2) mean number of beats per bout; and (3) mean bout duration. Preliminary analysis indicated that individuals differ in their tendency to deliver drum beats in temporally close pairs separated by longer interbeat intervals. Qualitative examination also suggested that individuals may differ in the temporal integration of drumming into the pant hoot vocalization. These results suggest that there may be acoustic cues available for chimpanzees to recognize unseen males by their drumming performances alone. Drumming by Taï chimpanzees was also compared to drumming by chimpanzees (P.t. schweinfurthii) from the Kanyawara study group in Kibale National Park. Uganda. The Kanyawara chimpanzees appeared to drum more often without vocalizing than did the Taï chimpanzees. When they did drum and vocalize together, the Kanyawara chimpanzees appeared to integrate their drumming later into the associated pant hoots than did the Taï chimpanzees. These results suggest the possibility that interpopulation variation exists in chimpanzee buttress drumming.     

Responses of wild chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) toward seismic aftershocks in the Mahale Mountains National Park,Tanzania

This is the first report documenting the responses of wild chimpanzees (Pan troglodytes schweinfurthii) to seismic activities. During our long-term fieldwork in Mahale Mountains National Park, Tanzania, a high-intensity earthquake with a Richter magnitude of 6.8 occurred at 15:19 hours local time on 5 December 2005. During the main tremor, the chimpanzees displayed the “wraa” call, “scream,” and “pant bark” or “bark” vocalizations. Many mild aftershocks followed the main tremor, and the wild chimpanzees displayed a variety of responses to these. In several cases, they climbed trees or stopped activities such as grooming, moving, and feeding. These responses are similar to those previously reported in nonhuman primates. During the observations, a unique behavior, one never reported before was exhibited by a female chimpanzee. She placed her right palm on the ground giving the impression she was inspecting the trembling of the ground.     

Social,contextual, and individual factors affecting the occurrence and acoustic structure of drumming bouts in wild chimpanzees (Pan troglodytes)

The production of structured and repetitive sounds by striking objects is a behavior found not only in humans, but also in a variety of animal species, including chimpanzees (Pan troglodytes). In this study we examined individual and social factors that may influence the frequency with which individuals engage in drumming behavior when producing long distance pant hoot vocalizations, and analyzed the temporal structure of those drumming bouts. Male chimpanzees from Budongo Forest, Uganda, drummed significantly more frequently during travel than feeding or resting and older individuals were significantly more likely to produce drumming bouts than younger ones. In contrast, we found no evidence that the presence of estrus females, high ranking males and preferred social partners in the caller's vicinty had an effect on the frequency with which an individual accompanied their pant hoot vocalization with drumming. Through acoustic analyses, we demonstrated that drumming sequences produced with pant hoots may have contained information on individual identity and that qualitatively, there was individual variation in the complexity of the temporal patterns produced. We conclude that drumming patterns may act as individually distinctive long‐distance signals that, together with pant hoot vocalizations, function to coordinate the movement and spacing of dispersed individuals within a community, rather than as signals to group members in the immediate audience. Am J Phys Anthropol 156:125–134, 2015 © 2014 Wiley Periodicals, Inc.     

Selection for acoustic individuality within the vocal repertoire of wild chimpanzees

Individual primates typically produce acoustically distinct calls. To investigate the factors that facilitate the evolution of individual vocal signatures, we examined two components of the call repertoire of chimpanzees: the pant hoot and pant grunt. Pant hoots are long-distance signals whose recipients can be several hundred meters away, while pant grunts are short-range calls given to conspecifics within close visual range. Given their markedly different contexts of emission, we predicted that natural selection would favor the elaboration of individually distinctive acoustic features in pant hoots compared with pant grunts. Analyses of nine acoustic features revealed that pant hoots are more stereotyped within-individuals and variable between-individuals than pant grunts. These data are consistent with the hypothesis that selection may act to encode varying degrees of individuality in different components of the vocal repertoire of a single species.     

Geographic variation in the calls of wild chimpanzees: A reassessment

Male chimpanzees produce a species‐typical call, the pant hoot, to communicate to conspecifics over long‐distances. Calls given by males from the well‐known Gombe and Mahale populations typically consist of four different phases: an introduction, build‐up, climax, and let‐down. Recent observations suggest that chimpanzees living in the Kibale National Park, Uganda, consistently give calls that lack a build‐up and are thus qualitatively distinguishable acoustically from those made by other East African conspecifics. We analyzed additional recordings from Mahale and Kibale to re‐examine geographic variation in chimpanzee calls. Results indicate that males from both sites produce pant hoots containing all four parts of the call. Calls made by chimpanzees from the two populations, however, differ in quantitative acoustic measures. Specifically, males at Kibale initiate their calls with significantly longer elements and build‐up over briefer periods at slower rates than individuals from Mahale. Kibale males also deliver acoustically less variable calls than chimpanzees at Mahale. Although climax elements do not differ between populations in any single acoustic feature, discriminant function analysis reveals that acoustic variables can be used in combination to assign calls to the correct population at rates higher than that expected by chance. Ecological factors related to differences in habitat acoustics, the sound environment of the local biota, and body size are likely to account for these observed macrogeographic variations in chimpanzee calls. Am. J. Primatol. 47:133–151, 1999. © 1999 Wiley‐Liss, Inc.     

Dialects in wild chimpanzees?

Chimpanzees emit a loud, species-typical long distance call known as the pant hoot. Geographic variation between the pant hoots of chimpanzees living in two neighboring populations, the Mahale Mountains and Gombe Stream National Parks, Tanzania, was examined. Analysis of six acoustic features revealed subtle differences in the way chimpanzees from the two populations called. Individuals from the Mahale study site uttered one section of their pant hoots at a faster rate and with shorter elements than animals from Gombe Stream. In addition, individuals at Mahale delivered broader-band, higher pitched “climax” elements than males from Gombe. While genetic factors, anatomical differences, variations in the use of calls at the two sites, and changes in calling over time may account for the variability between populations, we suggest the additional possibility that differences in pant hooting may be due to learning. © 1992 Wiley-Liss, Inc.     

Communication of male quality in owl hoots

The evolution of communication through intrasexual selection is expected to lead signalers to transmit honest information on their fighting ability. Here we studied the information encoded in the acoustic structure of the territorial calls of a nocturnal raptor. During territorial contests, male scops owls give hoots composed of a downward frequency shift followed by a stable plateau. We found that the frequency of the hoot was negatively correlated with the body weight of the vocalizer. We shifted the frequency contour of natural hoots in order to create resynthesized calls corresponding to individuals of varying body weight and used these stimuli in playback experiments simulating an intrusion into the territory of established breeders. Territory owners responded less intensely when they heard hoots simulating heavier intruders, and males with heavier apparent weight tended to give hoots with a lower plateau in response to playbacks simulating heavier intruders.     

Fatal Chimpanzee Attack in Loango National Park,Gabon

In some populations, chimpanzees engage in lethal aggression within and between social units. We report a fatal attack on an adult male chimpanzee at a new research site in Loango National Park, Gabon. We found a fresh corpse of an adult male chimpanzee only a few hundred meters from the research camp, after noting numerous vocalizations and chimpanzee movements the previous evening. Previous contacts with chimpanzees and fresh tracks in the area around the corpse suggest that 2 communities of chimpanzees range where the attack occurred and that members of the neighboring community killed the chimpanzee. To support the conclusion, we conducted genetic analysis for 13 Y-chromosome loci and 9 microsatellite loci of fecal samples from the dead individual, 5 possible attackers, and 2 members of the other community Though we cannot exclude the possibility of an intracommunity killing, the combined observational and genetic evidence suggest an intercommunity attack. The case study adds to the growing evidence that intercommunity killings are a rare but widespread phenomenon among chimpanzees and not an artifact of human provisioning or habituation.     

Chimpanzee vocal signaling points to a multimodal origin of human language

The evolutionary origin of human language and its neurobiological foundations has long been the object of intense scientific debate. Although a number of theories have been proposed, one particularly contentious model suggests that human language evolved from a manual gestural communication system in a common ape-human ancestor. Consistent with a gestural origins theory are data indicating that chimpanzees intentionally and referentially communicate via manual gestures, and the production of manual gestures, in conjunction with vocalizations, activates the chimpanzee Broca's area homologue--a region in the human brain that is critical for the planning and execution of language. However, it is not known if this activity observed in the chimpanzee Broca's area is the result of the chimpanzees producing manual communicative gestures, communicative sounds, or both. This information is critical for evaluating the theory that human language evolved from a strictly manual gestural system. To this end, we used positron emission tomography (PET) to examine the neural metabolic activity in the chimpanzee brain. We collected PET data in 4 subjects, all of whom produced manual communicative gestures. However, 2 of these subjects also produced so-called attention-getting vocalizations directed towards a human experimenter. Interestingly, only the two subjects that produced these attention-getting sounds showed greater mean metabolic activity in the Broca's area homologue as compared to a baseline scan. The two subjects that did not produce attention-getting sounds did not. These data contradict an exclusive "gestural origins" theory for they suggest that it is vocal signaling that selectively activates the Broca's area homologue in chimpanzees. In other words, the activity observed in the Broca's area homologue reflects the production of vocal signals by the chimpanzees, suggesting that this critical human language region was involved in vocal signaling in the common ancestor of both modern humans and chimpanzees.     

Association partners of young chimpanzees in the Mahale Mountains National Park,Tanzania

Association partners of young chimpanzees at the Mahale Mountains National Park were analyzed. Juvenile and adolescent chimpanzees associated frequently with their mothers, although mother-offspring association decreased as the offspring grew up. Males tended to leave their mothers and associate with adult males, while females remained frequently associating with their mothers in early adolescence. In late adolescence and young adulthood, males usually associated with adult males and cycling adult females. Females may transfer into neighboring unit-groups in this stage. Although an immigrant female tended to be alone when her estrous cycle stopped, she associated with many individuals, in particular with adult males, when she resumed cycling. Some orphans were observed to associate frequently with particular adults. The findings were discussed in relation to the unique characteristics of chimpanzee social system.     

Case study of a chimpanzee recovered from poachers and temporarily released with wild conspecifics

This case study describes the behavior ofBahati (BA), a captive, wild-born, 4 – 6 yr old, female chimpanzee (Pan troglodytes schweinfurthii), before and after her temporary release into Kibale National Park, Uganda. Post-release interactions with habituated, wild chimpanzees were recorded.BA was not attacked by the wild chimpanzees at the time of introduction and observations during her period of travel with conspecifics indicate that she received significantly more affiliative contact than aggressive.BA foraged further from humans, glanced less at observers and climbed ever higher in the canopy as the project proceeded.BA associated closely with two wild chimpanzees: a fully adult male and a young juvenile female. In the latter stages of the project, she spent progressively less time in proximity to conspecifics. After six weeks,BA no longer followed wild chimpanzees and repeatedly returned to human settlements. The attempt was terminated after two months andBA returned to captivity. On the basis of this case study, we suggest possible adjustments to re-introduction criteria, and discuss the outcomes of pre-release training, public involvement, and health screening under field conditions.     

Fruit Finding by Mangabeys (Lophocebus albigena): Are Monitoring of Fig Trees and Use of Sympatric Frugivore Calls Possible Strategies?

Frugivorous forest primates face a continual challenge to locate ripe fruit due to the poor visibility characterizing a heavily vegetated habitat and the spatial and temporal unpredictability of their fruit sources. We present two hypotheses regarding fruit finding in gray-cheeked mangabeys (Lophocebus albigena). The first hypothesis is that mangabeys monitor nonfruiting fig trees by visiting and checking them for fruit at a higher rate than control trees that do not produce preferred fruit. We test this hypothesis by comparing rates of visitation to focal fig trees and control trees. The second hypothesis is that mangabeys use sympatric frugivore loud calls to locate fruit sources. We test this hypothesis (1) observationally, by comparing the rates at which mangabeys visit calling sites of sympatric frugivores and matched control areas; and (2) experimentally, by following mangabey responses to playbacks of tape-recorded calls: the black-and-white-casqued hornbill (Bycanistes subcylindricus) long call, the great blue turaco (Corythaeola cristata) rattling kok, the adult male mangabey whoopgobble, and the chimpanzee (Pan troglodytes) pant hoot. We tested the hypotheses via data from a single group of mangabeys in the Kibale National Park, Uganda. There is no evidence that mangabeys monitor fig trees for the presence of fruit, but they may use the calls of hornbills to locate fruit. Statistical evidence that mangabeys use conspecific whoopgobbles and chimpanzee pant hoots in fruit finding is lacking, though anecdotal observations suggest this possibility. There is no evidence for use of turaco calls in fruit finding.     

An observed birth in a free-living chimpanzee (Pan troglodytes schweinfurthii) in Gombe National Park,Tanzania

During 19 years of study of chimpanzees in the Gombe National Park only one birth has been observed; this is probably the first such observation for any chimpanzee in the natural habitat. The birth took place in a nest in a tall tree during the morning. Details were recorded by Tanzanian field staff from a neighbouring tree. Labour and parturition are described as well as the mother’s care of the infant immediately after birth. The mother consumed the placenta as she lay in another nest. Throughout the birth process the mother’s juvenile son remained close by and watched with apparent interest. Another mother and her offspring were present during the birth and an adult male approached the mother while she was feeding on the placenta. Their behaviour is described.     

Visual Search for Human Gaze Direction by a Chimpanzee (Pan troglodytes)

Background

Humans detect faces with direct gazes among those with averted gazes more efficiently than they detect faces with averted gazes among those with direct gazes. We examined whether this “stare-in-the-crowd” effect occurs in chimpanzees (Pan troglodytes), whose eye morphology differs from that of humans (i.e., low-contrast eyes, dark sclera).

Methodology/Principal Findings

An adult female chimpanzee was trained to search for an odd-item target (front view of a human face) among distractors that differed from the target only with respect to the direction of the eye gaze. During visual-search testing, she performed more efficiently when the target was a direct-gaze face than when it was an averted-gaze face. This direct-gaze superiority was maintained when the faces were inverted and when parts of the face were scrambled. Subsequent tests revealed that gaze perception in the chimpanzee was controlled by the contrast between iris and sclera, as in humans, but that the chimpanzee attended only to the position of the iris in the eye, irrespective of head direction.

Conclusion/Significance

These results suggest that the chimpanzee can discriminate among human gaze directions and are more sensitive to direct gazes. However, limitations in the perception of human gaze by the chimpanzee are suggested by her inability to completely transfer her performance to faces showing a three-quarter view.     

Rosalià Abreu and the Apes of Havana

Rosalià Abreu (1862–1930) was a wealthy amateur collector and keeper of primates, including apes. She was the first person to keep orangutans and chimpanzees alive in captivity for their natural lifespans, and the first to breed chimpanzees. Although not a scientist, she made her animals available for scientific study, and it was on her animals that the first published observations of chimpanzee mating, birth, and development were made. The system of husbandry that she designed for apes, with an emphasis on spacious cages, vegetarian diet, cleanliness, and social contact (preferably conspecific, but heterospecific where conspecifics were not available), were groundbreaking in their time. Through her extensive correspondence, her methods of husbandry spread and formed the basis of captive ape practices around the world. Though much of Abreu’s work was forward-looking, other aspects, such as her belief that chimpanzees were monogamous and had psychic abilities, strike modern readers as eccentric. Nonetheless primatologists today may be helped to see the cultural assumptions that underlie today’s research by noting those that guided research in the past.