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1

Plumptre, A. J., and V. Reynolds. "Censusing chimpanzees in the Budongo Forest, Uganda." International Journal of Primatology 17, no. 1 (February 1996): 85–99. http://dx.doi.org/10.1007/bf02696160.

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2

Tweheyo, Mnason, and Kare A. Lye. "Patterns of frugivory of the Budongo Forest chimpanzees, Uganda." African Journal of Ecology 43, no. 4 (December 2005): 282–90. http://dx.doi.org/10.1111/j.1365-2028.2005.00566.x.

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3

Newton-Fisher, Nicholas E. "The diet of chimpanzees in the Budongo Forest Reserve, Uganda." African Journal of Ecology 37, no. 3 (September 1999): 344–54. http://dx.doi.org/10.1046/j.1365-2028.1999.00186.x.

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4

Arnold, Kate, and Andrew Whiten. "POST-CONFLICT BEHAVIOUR OF WILD CHIMPANZEES (PAN TROGLODYTES SCHWEINFURTHII) IN THE BUDONGO FOREST, UGANDA." Behaviour 138, no. 5 (2001): 649–90. http://dx.doi.org/10.1163/156853901316924520.

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AbstractSince de Waal & van Roosmalen (1979) first documented the occurrence of reconciliation between former opponents in captive chimpanzees, the study of the post-conflict behaviour of primates has provided valuable information about some of the details of primate social organisation. The vast majority of these studies have been carried out on captive subjects and it has been assumed that these findings are representative of wild primates. We set out to investigate whether this was true for the Sonso community of wild chimpanzees (Pan troglodytes schweinfurthii) of the Budongo Forest, Uganda, using controlled procedures comparable with those used in captive studies. We found that these chimpanzees were much less likely to reconcile than their captive counterparts. Only one dimension of relationship quality had an effect on the likelihood of reconciliation. Individuals which were highly compatible, in terms of time spent affiliating, reconciled conflicts more often than those with weak relationships. Captive chimpanzees have also been shown to 'console' one another (de Waal & van Roosmalen, 1979; de Waal & Aureli, 1996), where uninvolved bystanders initiate affiliative contacts with victims of aggression. This study did not confirm that consolatory behaviour was characteristic of wild chimpanzee post-conflict behaviour. Nor did these chimpanzees use explicit gestures during post-conflict interactions as they have been shown to do in two out of three captive studies. We conclude that the post-conflict behaviour of chimpanzees is more variable than has previously been thought and is likely to be dependent on the prevailing social environment.
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5

McLennan, Matthew R., David Hyeroba, Caroline Asiimwe, Vernon Reynolds, and Janette Wallis. "Chimpanzees in mantraps: lethal crop protection and conservation in Uganda." Oryx 46, no. 4 (October 2012): 598–603. http://dx.doi.org/10.1017/s0030605312000592.

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AbstractA main concern of farmers worldwide is how to reduce crop losses to wildlife. Some potentially lethal crop protection methods are non-selective. It is important to understand the impact of such methods on species of conservation concern. Uganda has important populations of Endangered eastern chimpanzees Pan troglodytes schweinfurthii. Farmers sometimes use large metal mantraps to guard their fields against crop-raiding wildlife, particularly baboons Papio anubis and wild pigs Potamochoerus sp.. Chimpanzees that range onto farmland also step in these illegal devices and without rapid veterinary invention face severe injury or eventual death. Unlike inadvertent snaring of great apes in African forests, the problem of mantraps in forest–farm ecotones has received little attention. We report 10 cases of entrapped chimpanzees in the cultivated landscape surrounding Uganda's Budongo Forest during 2007–2011, undoubtedly only a portion of the actual number of cases. Mantraps currently present a substantial threat to ape populations in this important conservation landscape. Our data underscore the need for conservation programmes to consider the techniques used by rural farmers to protect their livelihoods from wild animals.
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6

Newton-Fisher, Nicholas E. "Termite eating and food sharing by male chimpanzees in the Budongo Forest, Uganda." African Journal of Ecology 37, no. 3 (September 1999): 369–71. http://dx.doi.org/10.1046/j.1365-2028.1999.00187.x.

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7

Emery Thompson, M., N. E. Newton-Fisher, and V. Reynolds. "Probable Community Transfer of Parous Adult Female Chimpanzees in the Budongo Forest, Uganda." International Journal of Primatology 27, no. 6 (December 5, 2006): 1601–17. http://dx.doi.org/10.1007/s10764-006-9098-0.

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8

Whiten, Andrew, and Kate Arnold. "Grooming Interactions Among the Chimpanzees of the Budongo Forest, Uganda: Tests of Five Explanatory Models." Behaviour 140, no. 4 (2003): 519–52. http://dx.doi.org/10.1163/156853903322127968.

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AbstractPatterns of allogrooming among the Sonso community of chimpanzees of the Budongo Forest, Uganda, were examined and found to closely resemble those at other study sites. Strong affiliative bonds among males were reflected in high levels of grooming compared with other sex combinations. Adult males groomed, and received grooming most often from, other adult males and also adolescent females which were the only females with regular oestrous cycles during the study. Males had a wider diversity of grooming partners than females and groomed more equitably. However, males concentrated the majority of their effort on a very small number of partners compared with other sites. Grooming reciprocity was found among all age/sex combinations with the exception of adult male-female dyads once immediate reciprocation in the form of synchronous mutual grooming was removed from the analysis. Since grooming among males is thought to play a major role in servicing relationships and agonistic coalitions that can improve dominance status, competition for high-ranking grooming partners was predicted to influence the distribution of grooming among males. Grooming was indeed directed up the male hierarchy and closely ranked males groomed each other more often than those that were distantly ranked. However, when only adult males were considered, rank had little effect on grooming distributions. High rank appeared to influence access to females, but did not attract more female grooming partners. Grooming distributions in this average-sized community did not fit a number of alternative priority of access models which assume competition for high-ranking grooming partners that Watts (2000b) found to have some explanatory value in one very large community of chimpanzees, but not in a smaller, more representative one. Although rank is highly likely to influence coalition partner choice, whether such relationships depend upon strategic grooming partner choices in wild chimpanzees is presently unclear.
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9

Newton-Fisher, Nicholas E. "The home range of the Sonso community of chimpanzees from the Budongo Forest, Uganda." African Journal of Ecology 41, no. 2 (June 2003): 150–56. http://dx.doi.org/10.1046/j.1365-2028.2003.00408.x.

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10

Newton-Fisher, Nicholas E., Melissa Emery Thompson, Vernon Reynolds, Christophe Boesch, and Linda Vigilant. "Paternity and social rank in wild chimpanzees (Pan troglodytes) from the Budongo Forest, Uganda." American Journal of Physical Anthropology 142, no. 3 (December 23, 2009): 417–28. http://dx.doi.org/10.1002/ajpa.21241.

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11

Bakuneeta, C., K. Johnson, R. Plumptre, and V. Reynolds. "Human uses of tree species whose seeds are dispersed by chimpanzees in the Budongo Forest, Uganda." African Journal of Ecology 33, no. 3 (September 1995): 276–78. http://dx.doi.org/10.1111/j.1365-2028.1995.tb00807.x.

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12

Cameron, Elissa, Kerry Slater, Johan du Toit, and Trudy Turner. "The influence of oestrous swellings on the grooming behaviour of chimpanzees of the Budongo Forest, Uganda." Behaviour 145, no. 9 (2008): 1235–46. http://dx.doi.org/10.1163/156853908785387629.

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13

McGrew, William C. "Chimpanzees of the Budongo Forest, Uganda Vernon Reynolds . Review ofThe Chimpanzees of the Budongo Forest: Ecology, Behaviour, and Conservation. Oxford University Press. Oxford. 297ISBN: 0-19-851546-4. Paperback: US$ 69.50. 2005." Primate Conservation 21 (August 2006): 179–80. http://dx.doi.org/10.1896/0898-6207.21.1.179.

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14

Dezecache, Guillaume, Klaus Zuberbühler, Marina Davila-Ross, and Christoph D. Dahl. "Skin temperature changes in wild chimpanzees upon hearing vocalizations of conspecifics." Royal Society Open Science 4, no. 1 (January 2017): 160816. http://dx.doi.org/10.1098/rsos.160816.

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A growing trend of research using infrared thermography (IRT) has shown that changes in skin temperature, associated with activity of the autonomic nervous system, can be reliably detected in human and non-human animals. A contact-free method, IRT provides the opportunity to uncover emotional states in free-ranging animals during social interactions. Here, we measured nose and ear temperatures of wild chimpanzees of Budongo Forest, Uganda, when exposed to naturally occurring vocalizations of conspecifics. We found a significant temperature decrease over the nose after exposure to conspecifics' vocalizations, whereas we found a corresponding increase for ear temperature. Our study suggests that IRT can be used in wild animals to quantify changes in emotional states in response to the diversity of vocalizations, their functional significance and acoustical characteristics. We hope that it will contribute to more research on physiological changes associated with social interactions in wild animals.
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15

Reynolds, V., A. J. Plumptre, J. Greenham, and J. Harborne. "Condensed tannins and sugars in the diet of chimpanzees ( Pan troglodytes schweinfurthii ) in the Budongo Forest, Uganda." Oecologia 115, no. 3 (July 3, 1998): 331–36. http://dx.doi.org/10.1007/s004420050524.

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16

Waller, John C., and Vernon Reynolds. "Limb injuries resulting from snares and traps in chimpanzees (Pan troglodytes schweinfurthii) of the Budongo Forest, Uganda." Primates 42, no. 2 (April 2001): 135–39. http://dx.doi.org/10.1007/bf02558140.

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17

Brownlow, A. R., A. J. Plumptre, V. Reynolds, and R. Ward. "Sources of variation in the nesting behavior of chimpanzees (Pan troglodytes schweinfurthii) in the Budongo forest, Uganda." American Journal of Primatology 55, no. 1 (2001): 49–55. http://dx.doi.org/10.1002/ajp.1038.

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18

Kosheleff, Valerie P., and Christian N. K. Anderson. "Temperature's influence on the activity budget, terrestriality, and sun exposure of chimpanzees in the Budongo Forest, Uganda." American Journal of Physical Anthropology 139, no. 2 (June 2009): 172–81. http://dx.doi.org/10.1002/ajpa.20970.

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19

Reynolds, Vernon, Andrew W. Lloyd, Fred Babweteera, and Christopher J. English. "Decaying Raphia farinifera Palm Trees Provide a Source of Sodium for Wild Chimpanzees in the Budongo Forest, Uganda." PLoS ONE 4, no. 7 (July 10, 2009): e6194. http://dx.doi.org/10.1371/journal.pone.0006194.

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20

Dezecache, Guillaume, Claudia Wilke, Nathalie Richi, Christof Neumann, and Klaus Zuberbühler. "Skin temperature and reproductive condition in wild female chimpanzees." PeerJ 5 (December 5, 2017): e4116. http://dx.doi.org/10.7717/peerj.4116.

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Infrared thermal imaging has emerged as a valuable tool in veterinary medicine, in particular for evaluating reproductive processes. Here, we explored differences in skin temperature of twenty female chimpanzees in Budongo Forest, Uganda, four of which were pregnant during data collection. Based on previous literature in other mammals, we predicted increased skin temperature of maximally swollen reproductive organs of non-pregnant females when approaching peak fertility. For pregnant females, we made the same prediction because it has been argued that female chimpanzees have evolved mechanisms to conceal pregnancy, including swellings of the reproductive organs, conspicuous copulation calling, and solicitation of male mating behaviour, to decrease the infanticidal tendencies of resident males by confusing paternity. For non-pregnant females, we found slight temperature increases towards the end of the swelling cycles but no significant change between the fertile and non-fertile phases. Despite their different reproductive state, pregnant females had very similar skin temperature patterns compared to non-pregnant females, suggesting little potential for males to use skin temperature to recognise pregnancies, especially during maximal swelling, when ovulation is most likely to occur in non-pregnant females. We discuss this pattern in light of the concealment hypothesis, i.e., that female chimpanzees have evolved physiological means to conceal their reproductive state during pregnancy.
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21

Krief, Sabrina, Camille Daujeard, Marie-Hélène Moncel, Noemie Lamon, and Vernon Reynolds. "Flavouring food: the contribution of chimpanzee behaviour to the understanding of Neanderthal calculus composition and plant use in Neanderthal diets." Antiquity 89, no. 344 (April 2015): 464–71. http://dx.doi.org/10.15184/aqy.2014.7.

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In a recent study, Hardy et al. (2012) examined ten samples of dental calculus from five Neanderthal individuals from El Sidrón in northern Spain (occupation dates between 47300 and 50600 BP). In calculus from a young adult, they discovered the presence of compounds (dihydroazulene, chamazulene and methylherniarin) that occur in yarrow (Achillea millefolium) and camomile (Matriarca chamomilla). In preference to other hypotheses, the authors proposed that these two plants were used for self-medication. In this paper, we do not reject the self-medication hypothesis, but our observations of wild chimpanzees in Uganda, at Sonso in the Budongo Forest Reserve and at Kanyawara and Sebitoli in Kibale National Park (separated by about 150km), as well as ethnological and palaeontological evidence, lead us to propose three other explanations for the presence of these compounds. In addition, data on Neanderthal behaviour suggest that their subsistence and technological strategies were complex.
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22

Newton-Fisher, Nicholas. "ASSOCIATION BY MALE CHIMPANZEES: A SOCIAL TACTIC?" Behaviour 136, no. 6 (1999): 705–30. http://dx.doi.org/10.1163/156853999501531.

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AbstractWhether and with whom to associate are possibly some of the most fundamental of choices faced by a chimpanzee, choices constrained by and dependent upon similar decisions made by potential association partners. Although much work has focused on the impact of ecological factors on the size of the temporary associations, or parties, it seems that chimpanzees, especially the males, form parties to gain social benefits, and that these benefits are dependent the particular composition of these parties. This paper examines the social determinates of the association patterns of male chimpanzees, members of the Sonso community in the Budongo Forest Reserve, western Uganda. Male chimpanzees showed clear preferences for association partners, and these partners had a strong tendency to associate in small parties. Strength of association within dyads accounted for a significant fraction of the variance in party size [all parties containing males: r2 = 0.30; male parties only: r2 = 0.24 (all dyads), r2 = 0 . 50 (dyads with positive associations only)]. These results supported the hypothesis that association was the result of tactical decisions, and refuted predictions of alternate hypotheses: random and passive (independent attraction to the same locations, such as fruiting trees) association. Further support for the tactical association hypothesis was provided by observations that association between males was flexible and that association patterns resolved into alternative association strategies, with some males shifting between strategies over time. The presence of cycling females influenced party size, which increased with the number present. The number of adult males in a party was also greater when cycling females were present, although remained similar whether one, two, three or four such females were in the party. Samples sizes for parties containing more than two cycling females were low, however. It is suggested that, for a Machiavellian primate in fission-fusion social system, the need to make decisions concerning association partners is cognitively demanding and may be an important, frequently used, function of the intelligence demonstrated by captive chimpanzees.
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23

Crockford, Catherine, Thibaud Gruber, and Klaus Zuberbühler. "Chimpanzee quiet hoo variants differ according to context." Royal Society Open Science 5, no. 5 (May 2018): 172066. http://dx.doi.org/10.1098/rsos.172066.

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In comparative studies of evolution of communication, the function and use of animal quiet calls have typically been understudied, despite that these signals are presumably under selection like other vocalizations, such as alarm calls. Here, we examine vocalization diversification of chimpanzee quiet ‘hoos’ produced in three contexts—travel, rest and alert—and potential pressures promoting diversification. Previous playback and observational studies have suggested that the overarching function of chimpanzee hoos is to stay in contact with others, particularly bond partners. We conducted an acoustic analysis of hoos using audio recordings from wild chimpanzees ( Pan troglodytes schweinfurthii ) of Budongo Forest, Uganda. We identified three acoustically distinguishable, context-specific hoo variants. Each call variant requires specific responses from receivers to avoid breaking up the social unit. We propose that callers may achieve coordination by using acoustically distinguishable calls, advertising their own behavioural intentions. We conclude that natural selection has acted towards acoustically diversifying an inconspicuous, quiet vocalization, the chimpanzee hoo. This evolutionary process may have been favoured by the fact that signallers and recipients share the same goal, to maintain social cohesion, particularly among those who regularly cooperate, suggesting that call diversification has been favoured by the demands of cooperative activities.
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24

Langergraber, Kevin E., Carolyn Rowney, Catherine Crockford, Roman Wittig, Klaus Zuberbühler, and Linda Vigilant. "Genetic analyses suggest no immigration of adult females and their offspring into the Sonso community of chimpanzees in the Budongo Forest Reserve, Uganda." American Journal of Primatology 76, no. 7 (January 16, 2014): 640–48. http://dx.doi.org/10.1002/ajp.22258.

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25

Tweheyo, Mnason, and Joseph Obua. "Feeding habits of chimpanzees (Pan troglodytes ), red-tail monkeys (Cercopithecus ascanius schmidti ) and blue monkeys (Cercopithecus mitis stuhlmanii ) on figs in Budongo Forest Reserve, Uganda." African Journal of Ecology 39, no. 2 (June 2001): 133–39. http://dx.doi.org/10.1046/j.1365-2028.2001.00290.x.

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26

Bouchard, Alice, and Klaus Zuberbühler. "An intentional cohesion call in male chimpanzees of Budongo Forest." Animal Cognition, January 19, 2022. http://dx.doi.org/10.1007/s10071-022-01597-6.

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AbstractMany social animals travel in cohesive groups but some species, including chimpanzees, form flexible fission–fusion systems where individuals have some control over group cohesion and proximity to others. Here, we explored how male chimpanzees of the Sonso community of Budongo Forest, Uganda, use communication signals during resting, a context where the likelihood of group fission is high due to forthcoming travel. We focused on a context-specific vocalisation, the ‘rest hoo’, to investigate its function and determine whether it is produced intentionally. We found that this call was typically given towards the end of typical silent resting bouts, i.e., the period when individuals need to decide whether to continue travelling after a brief stop-over or to start a prolonged resting bout. Subjects rested longer after producing ‘rest hoos’ and their resting time increased with the number of calls produced. They also rested longer if their calls were answered. Furthermore, focal subjects’ resting time was prolonged after hearing others’ ‘rest hoos’. Subjects called more when with top proximity partners and in small parties and rested longer if a top proximity partner called. We also found an interaction effect between rank and grooming activity, with high-ranking males with a high grooming index calling less frequently than other males, suggesting that vocal communication may serve as a cohesion strategy alternative to tactile-based bonding. We discuss these different patterns and conclude that chimpanzee ‘rest hoos’ meet key criteria for intentional signalling. We suggest that ‘rest hoos’ are produced to prolong resting bouts with desired partners, which may function to increase social cohesion.
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27

Badihi, Gal, Kelsey Bodden, Klaus Zuberbühler, Liran Samuni, and Catherine Hobaiter. "Flexibility in the social structure of male chimpanzees ( Pan troglodytes schweinfurthii ) in the Budongo Forest, Uganda." Royal Society Open Science 9, no. 9 (September 2022). http://dx.doi.org/10.1098/rsos.220904.

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Individuals of social species experience competitive costs and social benefits of group living. Substantial flexibility in humans' social structure and the combination of different types of social structure with fission–fusion dynamics allow us to live in extremely large groups—overcoming some of the costs of group living while capitalizing on the benefits. Non-human species also show a range of social strategies to deal with this trade-off. Chimpanzees are an archetypical fission–fusion species, using dynamic changes in day-to-day association to moderate the costs of within-group competition. Using 4 years of association data from two neighbouring communities of East African chimpanzees ( Pan troglodytes schweinfurthii ), we describe an unexplored level of flexibility in chimpanzee social structure. We show that males from the larger Waibira community ( N = 24–31) exhibited additional structural levels of semi-stable core–periphery society, while males from the smaller Sonso community ( N = 10–13) did not. This novel core–periphery pattern adds to previous results describing alternative modular social structure in other large communities of chimpanzees. Our data support the hypothesis that chimpanzees can incorporate a range of strategies in addition to fission–fusion to overcome costs of social living, and that their social structures may be closer to that of modern humans than previously described.
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28

McLennan, Matthew R., Georgia A. Lorenti, Simon Mugenyi, Jonan Muganzi, and Jacqueline Rohen. "Digging for clues: Stick tools used for honey digging in a second community of ‘forest fragment chimpanzees’ outside the Budongo and Bugoma Forests, Uganda." Revue de primatologie, no. 10 (December 1, 2019). http://dx.doi.org/10.4000/primatologie.6718.

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29

Soldati, Adrian, Geresomu Muhumuza, Guillaume Dezecache, Pawel Fedurek, Derry Taylor, Josep Call, and Klaus Zuberbühler. "The Ontogeny of Vocal Sequences: Insights from a Newborn Wild Chimpanzee (Pan troglodytes schweinfurthii)." International Journal of Primatology, August 16, 2022. http://dx.doi.org/10.1007/s10764-022-00321-y.

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AbstractObservations of early vocal behaviours in non-human primates (hereafter primates) are important for direct comparisons between human and primate vocal development. However, direct observations of births and perinatal behaviour in wild primates are rare, and the initial stages of behavioural ontogeny usually remain undocumented. Here, we report direct observations of the birth of a wild chimpanzee (Pan troglodytes schweinfurthii) in Budongo Forest, Uganda, including the behaviour of the mother and other group members. We monitored the newborn’s vocal behaviour for approximately 2 hours and recorded 70 calls. We categorised the vocalisations both qualitatively, using conventional call descriptions, and quantitatively, using cluster and discriminant acoustic analyses. We found evidence for acoustically distinct vocal units, produced both in isolation and in combination, including sequences akin to adult pant hoots, a vocal utterance regarded as the most complex vocal signal produced by this species. We concluded that chimpanzees possess the capacity to produce vocal sequences composed of different call types from birth, albeit in rudimentary forms. Our observations are in line with the idea that primate vocal repertoires are largely present from birth, with fine acoustic structures undergoing ontogenetic processes. Our study provides rare and valuable empirical data on perinatal behaviours in wild primates.
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