Social network shrinking is explained by active and passive effects but not increasing selectivity with age in wild macaques.

Evidence of social disengagement, network narrowing and social selectivity with advancing age in several non-human animals challenges our understanding of the causes of social ageing. Natural animal populations are needed to test whether social ageing and selectivity occur under natural predation and extrinsic mortality pressures, and longitudinal studies are particularly valuable to disentangle the contribution of within-individual ageing from the demographic processes that shape social ageing at the population level. Data on wild Assamese macaques (Macaca assamensis) were collected between 2013 and 2020 at the Phu Khieo Wildlife Sanctuary, Thailand. We investigated the social behaviour of 61 adult females observed for 13 270 h to test several mechanistic hypotheses of social ageing and evaluated the consistency between patterns from mixed-longitudinal and within-individual analyses. With advancing age, females reduced the size of their social network, which could not be explained by an overall increase in the time spent alone, but by an age-related decline in mostly active, but also passive, behaviour, best demonstrated by within-individual analyses. A selective tendency to approach preferred partners was maintained into old age but did not increase. Our results contribute to our understanding of the driver of social ageing in natural animal populations and suggest that social disengagement and selectivity follow independent trajectories during ageing.

Selective responding to human ostensive communication is an early developing capacity of domestic dogs.

Ostensive communication, communication motivated not only by an informative intention, but also by an intention to make this informative intention overt, is thought to be restricted to human signalers and to rely on metacognitive skills. On the receiver side, human infants and dogs have been found to selectively respond to human ostensive communication, even if it remains unclear whether they are able to discern the underlying intentions. At present only animals with extensive experiences with humans have demonstrated a response to human attention calling, suggesting that this sensitivity may be an exaptation of communicative skills allowing for context-specific responding to intraspecific signals. We investigated whether dogs could respond to subtle cues of ostension without clear attention-calling and how this skill varies across age and with different human-given signals. Using a two-way object choice task, we evaluated whether dogs of different ages followed, with their gaze and with their choices, ostensive human pointing and gaze more than a formally similar but non-ostensive gesture and a directional gaze cue. Dogs followed pointing more than gaze, and older dogs followed the directional cues more accurately than younger dogs. Independent of cue type, we found that dogs of all ages responded to human ostensive signals more than similar directional cues motivated by no communicative intention. Given that dogs are sensitive to subtle cues of ostensive communication from an early age, and that this does not appear to change with age, we suggest that living close to humans may have selected for this skill. RESEARCH HIGHLIGHTS: Like human infants, dogs followed human ostensive cues more than non-ostensive cues both by their approach behavior and their differential looking behavior. Dogs differentiated between ostensive and non-ostensive human cues from an early age on, without their attention having been called explicitly in advance. Dogs followed human-given pointing cues more than human-given gaze cues, independent of age or intention (ostensive or non-ostensive). The early developing sensitivity of dogs to human ostension suggests that living close to humans may have selected for this skill.

Slow development of vocal sequences through ontogeny in wild chimpanzees (Pan troglodytes verus).

The development of the unique, hierarchical, and endless combinatorial capacity in a human language requires neural maturation and learning through childhood. Compared with most non-human primates, where combinatorial capacity seems limited, chimpanzees present a complex vocal system comprising hundreds of vocal sequences. We investigated how such a complex vocal system develops and the processes involved. We recorded 10,929 vocal utterances of 98 wild chimpanzees aged 0-55 years, from Taï National Park, Ivory Coast. We developed customized Generalized non-Linear Models to estimate the ontogenetic trajectory of four structural components of vocal complexity: utterance length, diversity, probability of panting (requiring phonation across inhalation and exhalation), and probability of producing two adjacent panted units. We found chimpanzees need 10 years to reach adult levels of vocal complexity. In three variables, the steepest increase coincided with the age of first non-kin social interactions (2-5 years), and plateaued in sub-adults (8-10 years), as individuals integrate into adult social life. Producing two adjacent panted units may require more neuromuscular coordination of the articulators, as its emergence and steepest increase appear later in development. These results suggest prolonged maturational processes beyond those hitherto thought likely in species that do not learn their vocal repertoire. Our results suggest that multifaceted ontogenetic processes drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation. As humans live in a complex social world, empirical support for the "social complexity hypothesis" may have relevance for theories of language evolution. RESEARCH HIGHLIGHTS: Chimpanzees need around 10 years to develop the vocal structural complexity present in the adult repertoire, way beyond the age of emergence of every single vocal unit. Multifaceted ontogenetic processes may drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation. Non-linear increases in vocal complexity coincide with social developmental milestones. Vocal sequences requiring rapid articulatory change emerge later than other vocal sequences, suggesting neuro-muscular maturational processes continue through the juvenile years.

Persistent anthrax as a major driver of wildlife mortality in a tropical rainforest.

Anthrax is a globally important animal disease and zoonosis. Despite this, our current knowledge of anthrax ecology is largely limited to arid ecosystems, where outbreaks are most commonly reported. Here we show that the dynamics of an anthrax-causing agent, Bacillus cereus biovar anthracis, in a tropical rainforest have severe consequences for local wildlife communities. Using data and samples collected over three decades, we show that rainforest anthrax is a persistent and widespread cause of death for a broad range of mammalian hosts. We predict that this pathogen will accelerate the decline and possibly result in the extirpation of local chimpanzee (Pan troglodytes verus) populations. We present the epidemiology of a cryptic pathogen and show that its presence has important implications for conservation.

Decay rates of arboreal and terrestrial nests of Eastern chimpanzees (Pan troglodytes schweinfurthii) in the Bugoma Central Forest Reserve, Uganda: Implications for population size estimates.

Chimpanzees were once thought to sleep primarily in the trees, but recent studies indicate that some populations also construct terrestrial night nests. This behavior has relevance not only to understanding the behavioral diversity of Pan troglodytes, but also to the conservation of the species, given that nest encounter rates are often used to estimate great ape population densities. A proper estimate of decay rates for ground nests is necessary for converting the encounter rate of nests to the density of weaned chimpanzees. Here we present the results of the first systematic comparative study between the decay rates of arboreal and terrestrial chimpanzee nests, from the Bugoma Central Forest Reserve in western Uganda. We followed the decay of 56 ground and 51 tree nests in eight nest groups between April 2020 and October 2021. For 15 of the ground and 19 of the tree nests, we collected detailed information on the condition of the nests every two weeks; we checked the remaining 73 nests only twice. On average, ground nests lasted 238 days versus 276 days for tree nests (p = 0.05). Of the 107 total nests surveyed, 51% of tree and 64% of ground nests had disappeared after six months. Based on our results, we propose a modification of the formula used to convert nest density into chimpanzee density. Our results highlight the importance of taking into account potential differences in decay rates between ground versus tree nests, which will likely influence our understanding of the distribution of ground nesting behavior in chimpanzee across tropical Africa, as well as our estimations of the densities of ground nesting populations.

Teaching varies with task complexity in wild chimpanzees.

Cumulative culture is a transformative force in human evolution, but the social underpinnings of this capacity are debated. Identifying social influences on how chimpanzees acquire tool tasks of differing complexity may help illuminate the evolutionary origins of technology in our own lineage. Humans routinely transfer tools to novices to scaffold their skill development. While tool transfers occur in wild chimpanzees and fulfill criteria for teaching, it is unknown whether this form of helping varies between populations and across tasks. Applying standardized methods, we compared tool transfers during termite gathering by chimpanzees in the Goualougo Triangle, Republic of Congo, and in Gombe, Tanzania. At Goualougo, chimpanzees use multiple, different tool types sequentially, choose specific raw materials, and perform modifications that improve tool efficiency, which could make it challenging for novices to manufacture suitable tools. Termite gathering at Gombe involves a single tool type, fishing probes, which can be manufactured from various materials. Multiple measures indicated population differences in tool-transfer behavior. The rate of transfers and probability of transfer upon request were significantly higher at Goualougo, while resistance to transfers was significantly higher at Gombe. Active transfers of tools in which possessors moved to facilitate possession change upon request occurred only at Goualougo, where they were the most common transfer type. At Gombe, tool requests were typically refused. We suggest that these population differences in tool-transfer behavior may relate to task complexity and that active helping plays an enhanced role in the cultural transmission of complex technology in wild apes.

Factors affecting nest height and ground nesting behaviour in Eastern chimpanzees (Pan troglodytes schweinfurthii) of the northern Democratic Republic of the Congo.

In order to achieve a better understanding of the factors that might have led our hominin ancestors to transition to a more terrestrial niche, including sleeping on the ground, we have conducted a study on the ground nesting behavior of chimpanzees (Pan troglodytes). Chimpanzees, like all other species of great apes, build nests in which to sleep each night, but little is known about regional differences in their nesting habits. Previously, nesting on the ground was considered typical of gorillas, but rare in most populations of chimpanzees. Using data acquired during our extensive chimpanzee nesting survey conducted between 2004 and 2013 across a > 50 000 km2 region in northern Democratic Republic of the Congo, we report a distinctive ground nesting behaviour of eastern chimpanzees (P. t. schweinfurthii). We have mapped the geographical distribution of ground nesting and compared its frequency at 20 survey areas on both sides of a large river, the Uele. We found that ground nests made up more than 1% of total nests at 15 of the 20 survey regions. For a subset of 16 of these regions, we utilized statistical models to investigate whether forest type and structure, as well as the abundance of carnivores and large herbivores, and the activities of humans impacted the frequency of ground nesting and nest height. We predicted that higher encounter rates of human and dangerous animal signs would be associated with lower rates of ground nesting as well as increased nest height. Overall, 10.4% of the Bili-Uéré chimpanzee nests were terrestrial, but the frequency of ground nesting varied extensively between the survey areas (0-29% of nests). The occurrence of ground nests was positively associated with denser forests (p = 0.004), herb patches (p < 0.001), and light gaps (p < 0.001). Light gaps (p < 0.001), herb patches (p = 0.044), and vine tangles (p = 0.016) also had a strong negative effect on nest height. Hunting by humans had a negative effect on the probability of the occurrence of ground nests (p = 0.001) and a positive one on nest height (p = 0.013), with a similar but likely marginal effect of large herbivores on nest height (p = 0.023). In addition, the chimpanzees nested at significantly lower heights with increasing distance from roads and settlements (p < 0.001). Carnivore encounter rates, however, had no significant impact on ground nest frequency or nest height. Our results indicate that ground nesting can no longer be considered a rare and patchily-occurring phenomenon in Pan troglodytes, but is instead a major component of the chimpanzee behavioural repertoire across a considerable fraction of the range of the Eastern subspecies. Our study highlights that neither the large body size of gorillas nor the taming of fire are necessary conditions for hominids to sleep overnight on the ground, even in areas inhabited by multiple species of large carnivore. Human hunting, however, appears to reduce the probability of ground nesting, or eliminate the behavior altogether.

Variation in aggression rates and urinary cortisol levels indicates intergroup competition in wild bonobos.

Intergroup competition is a widespread phenomenon across taxa and groups typically compete over access to limited resources, such as food and mates. Such competition may be quantified by changes in individuals' behavioral and physiological status in response to intergroup encounters (IGEs). Bonobos, one of our closest living relatives, are often regarded as xenophilic and exhibit high tolerance towards out-group individuals. This tolerance between groups may still be accompanied by intergroup competition over resources. We hereby compared variation in aggression rates and urinary cortisol levels of bonobos during and outside contexts of IGEs in the Kokolopori Bonobo Reserve and investigated whether food and mate availability influenced males' and females' aggression and cortisol levels, when controlling for dominance rank and the number of individuals present. We found that although females had higher aggression rates and urinary cortisol levels during than outside contexts of IGEs, these increases were not related to food availability or changes in between-group dynamics when maximally tumescent females were present, rather than absent. Furthermore, males showed higher aggression rates and urinary cortisol levels during than outside contexts of IGEs. However, males' responses during IGEs were not related to the presence of maximally tumescent females and food availability. Taken together, while competition intensified during seemingly tolerant IGEs in bonobos, such competition was unrelated to short-term changes in food and mate availability. Despite physical and physiological costs of aggression, bonobos associate with out-group individuals frequently and for extended periods. This suggests potential benefits of bonobo intergroup associations.

Kea (Nestor notabilis) show flexibility and individuality in within-session reversal learning tasks.

The midsession reversal paradigm confronts an animal with a two-choice discrimination task where the reward contingencies are reversed at the midpoint of the session. Species react to the reversal with either win-stay/lose-shift, using local information of reinforcement, or reversal estimation, using global information, e.g. time, to estimate the point of reversal. Besides pigeons, only mammalian species were tested in this paradigm so far and analyses were conducted on pooled data, not considering possible individually different responses. We tested twelve kea parrots with a 40-trial midsession reversal test and additional shifted reversal tests with a variable point of reversal. Birds were tested in two groups on a touchscreen, with the discrimination task having either only visual or additional spatial information. We used Generalized Linear Mixed Models to control for individual differences when analysing the data. Our results demonstrate that kea can use win-stay/lose-shift independently of local information. The predictors group, session, and trial number as well as their interactions had a significant influence on the response. Furthermore, we discovered notable individual differences not only between birds but also between sessions of individual birds, including the ability to quite accurately estimate the reversal position in alternation to win-stay/lose-shift. Our findings of the kea's quick and flexible responses contribute to the knowledge of diversity in avian cognitive abilities and emphasize the need to consider individuality as well as the limitation of pooling the data when analysing midsession reversal data.

Range-wide indicators of African great ape density distribution.

Species distributions are influenced by processes occurring at multiple spatial scales. It is therefore insufficient to model species distribution at a single geographic scale, as this does not provide the necessary understanding of determining factors. Instead, multiple approaches are needed, each differing in spatial extent, grain, and research objective. Here, we present the first attempt to model continent-wide great ape density distribution. We used site-level estimates of African great ape abundance to (1) identify socioeconomic and environmental factors that drive densities at the continental scale, and (2) predict range-wide great ape density. We collated great ape abundance estimates from 156 sites and defined 134 pseudo-absence sites to represent additional absence locations. The latter were based on locations of unsuitable environmental conditions for great apes, and on existing literature. We compiled seven socioeconomic and environmental covariate layers and fitted a generalized linear model to investigate their influence on great ape abundance. We used an Akaike-weighted average of full and subset models to predict the range-wide density distribution of African great apes for the year 2015. Great ape densities were lowest where there were high Human Footprint and Gross Domestic Product values; the highest predicted densities were in Central Africa, and the lowest in West Africa. Only 10.7% of the total predicted population was found in the International Union for Conservation of Nature Category I and II protected areas. For 16 out of 20 countries, our estimated abundances were largely in line with those from previous studies. For four countries, Central African Republic, Democratic Republic of the Congo, Liberia, and South Sudan, the estimated populations were excessively high. We propose further improvements to the model to overcome survey and predictor data limitations, which would enable a temporally dynamic approach for monitoring great apes across their range based on key indicators.

Human local adaptation of the TRPM8 cold receptor along a latitudinal cline.

Ambient temperature is a critical environmental factor for all living organisms. It was likely an important selective force as modern humans recently colonized temperate and cold Eurasian environments. Nevertheless, as of yet we have limited evidence of local adaptation to ambient temperature in populations from those environments. To shed light on this question, we exploit the fact that humans are a cosmopolitan species that inhabit territories under a wide range of temperatures. Focusing on cold perception-which is central to thermoregulation and survival in cold environments-we show evidence of recent local adaptation on TRPM8. This gene encodes for a cation channel that is, to date, the only temperature receptor known to mediate an endogenous response to moderate cold. The upstream variant rs10166942 shows extreme population differentiation, with frequencies that range from 5% in Nigeria to 88% in Finland (placing this SNP in the 0.02% tail of the FST empirical distribution). When all populations are jointly analyzed, allele frequencies correlate with latitude and temperature beyond what can be explained by shared ancestry and population substructure. Using a Bayesian approach, we infer that the allele originated and evolved neutrally in Africa, while positive selection raised its frequency to different degrees in Eurasian populations, resulting in allele frequencies that follow a latitudinal cline. We infer strong positive selection, in agreement with ancient DNA showing high frequency of the allele in Europe 3,000 to 8,000 years ago. rs10166942 is important phenotypically because its ancestral allele is protective of migraine. This debilitating disorder varies in prevalence across human populations, with highest prevalence in individuals of European descent-precisely the population with the highest frequency of rs10166942 derived allele. We thus hypothesize that local adaptation on previously neutral standing variation may have contributed to the genetic differences that exist in the prevalence of migraine among human populations today.

Lethal aggression in Pan is better explained by adaptive strategies than human impacts.

Observations of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) provide valuable comparative data for understanding the significance of conspecific killing. Two kinds of hypothesis have been proposed. Lethal violence is sometimes concluded to be the result of adaptive strategies, such that killers ultimately gain fitness benefits by increasing their access to resources such as food or mates. Alternatively, it could be a non-adaptive result of human impacts, such as habitat change or food provisioning. To discriminate between these hypotheses we compiled information from 18 chimpanzee communities and 4 bonobo communities studied over five decades. Our data include 152 killings (n = 58 observed, 41 inferred, and 53 suspected killings) by chimpanzees in 15 communities and one suspected killing by bonobos. We found that males were the most frequent attackers (92% of participants) and victims (73%); most killings (66%) involved intercommunity attacks; and attackers greatly outnumbered their victims (median 8:1 ratio). Variation in killing rates was unrelated to measures of human impacts. Our results are compatible with previously proposed adaptive explanations for killing by chimpanzees, whereas the human impact hypothesis is not supported.

Selective overimitation in dogs.

Dogs have not only shown different kinds of social learning, from either conspecifics or humans, including do-as-I-do imitation, deferred imitation, and selective imitation, but in two previous studies they have also shown an eagerness to copy causally irrelevant actions. This so-called overimitation is prevalent in humans but is totally absent in great apes. Whereas in one of two previous studies dogs copied actions from an experimenter (Johnston, Holden, & Santos in Developmental Science, 20, e12460, 2017), in the other a reasonable number of the dogs copied the irrelevant actions from their human caregiver (Huber, Popovová, Riener, Salobir, & Cimarelli in Learning & Behavior, 46, 387-397, 2018). Dogs have not only been domesticated to live and work with us, but many companion dogs develop strong affiliative relationships with their caregiver, which are akin to the attachment bonds between human children and their mother. We therefore assumed that overimitation in dogs might be strongly motivated by social factors, such as affiliation or conformity. To test this hypothesis, we confronted dogs with the same demonstration of causally relevant and irrelevant actions as in the previous study (Huber et al. in Learning & Behavior, 46, 387-397, 2018), but this time with an unfamiliar experimenter instead of the caregiver as the demonstrator. The results strongly supported our hypothesis: Whereas half of the subjects in the previous study replicated the causally irrelevant action demonstrated by their caregiver, only very few did so when the actions were demonstrated by the experimenter. We conclude that the eagerness of dogs to learn from humans and to copy even unnecessary actions is strongly facilitated by their relationship with the particular human.

Oxytocin reactivity during intergroup conflict in wild chimpanzees.

Intergroup conflict is evident throughout the history of our species, ubiquitous across human societies, and considered crucial for the evolution of humans' large-scale cooperative nature. Like humans, chimpanzee societies exhibit intragroup coordination and coalitionary support during violent intergroup conflicts. In both species, cooperation among group members is essential for individuals to gain access to benefits from engaging in intergroup conflict. Studies suggest that a contributive mechanism regulating in-group cooperation during intergroup conflicts in humans involves the neuropeptide hormone oxytocin, known to influence trust, coordination, and social cognition, although evidence from natural settings is lacking. Here, applying a noninvasive method, we investigate oxytocinergic system involvement during natural intergroup conflicts in wild chimpanzees. We found that chimpanzees of both sexes had significantly higher urinary oxytocin levels immediately before and during intergroup conflict compared with controls. Also, elevated hormone levels were linked with greater cohesion during intergroup conflicts, rather than with the level of potential threat posed by rival groups, intragroup affiliative social interactions, or coordinated behavior alone. Thus, the oxytocinergic system, potentially engendering cohesion and cooperation when facing an out-group threat, may not be uniquely human but rather a mechanism with evolutionary roots shared by our last common ancestor with chimpanzees, likely expediting fitness gains during intergroup conflict.

Joint attention skills in wild Arabian babblers ( Turdoides squamiceps): a consequence of cooperative breeding?

Human cooperation strongly relies on the ability of interlocutors to coordinate each other's attentional state: joint attention. One predominant hypothesis postulates that this hallmark of the unique cognitive system of humans evolved due to the combination of an ape-like cognitive system and the prosocial motives that facilitate cooperative breeding. Here, we tested this hypothesis by investigating communicative interactions of a cooperatively breeding bird species, the Arabian babbler ( Turdoides squamiceps). The behaviour of 12 wild social groups was observed focusing on two distinct communicative behaviours: object presentation and babbler walk. The results showed that both behaviours fulfilled the criteria for first-order intentional communication and involved co-orientation of recipients' attention. In turn, recipients responded with cooperative and communicative acts that resulted in coordinated joint travel between interlocutors. These findings provide the first evidence that another animal species shows several key criteria traditionally used to infer joint attention in prelinguistic human infants. Furthermore, they emphasize the influence of cooperative breeding on sophisticated socio-cognitive performances, while questioning the necessity of an ape-like cognitive system underlying joint attentional behaviour.

Chimpanzees flexibly update working memory contents and show susceptibility to distraction in the self-ordered search task.

Working memory (WM) is a core executive function that allows individuals to hold, process and manipulate information. WM capacity has been repeatedly nominated as a key factor in human cognitive evolution; nevertheless, little is known about the WM abilities of our closest primate relatives. In this study, we examined signatures of WM ability in chimpanzees (Pan troglodytes). Standard WM tasks for humans (Homo sapiens) often require participants to continuously update their WM. In Experiment 1, we implemented this updating requirement in a foraging situation: zoo-housed chimpanzees (n = 13) searched for food in an array of containers. To avoid redundant searches, they needed to continuously update which containers they had already visited (similar to WM paradigms for human children) with 15 s retention intervals in between each choice. We examined chimpanzees' WM capacity and to what extent they used spatial cues and object features to memorize their previous choices. In Experiment 2, we investigated how susceptible their WM was to attentional interference, an important signature, setting WM in humans apart from long-term memory. We found large individual differences with some individuals remembering at least their last four choices. Chimpanzees used a combination of spatial cues and object features to remember which boxes they had chosen already. Moreover, their performance decreased specifically when competing memory information was introduced. Finally, we found that individual differences in task performance were highly reliable over time. Together, these findings show remarkable similarities between human and chimpanzee WM abilities despite evolutionary and life-history differences.

Sun, age and test location affect spatial orientation in human foragers in rainforests.

The ability to know the direction of food sources is important for the foraging success of hunter-gatherers, especially in rainforests where dense vegetation limits visual detection distances. Besides sex and age, prior experience with the environment and the use of environmental cues are known to influence orientation abilities of humans. Among environmental cues, the position of the sun in the sky is important for orientation of diurnal animal species. However, whether or to what extent humans use the sun is largely unknown. Here, we investigated orientation abilities of the Mbendjele BaYaka people in the Republic of Congo, by conducting pointing tests (Nparticipants = 54, age: 6-76 years) in different locations in the rainforest. The Mbendjele were overall highly accurate at pointing to out-of-sight targets (median error: 6°). Pointing accuracy increased with age, but sex did not affect accuracy. Crucially, sun visibility increased pointing accuracy in young participants, especially when they were far from the camp. However, this effect became less apparent in older participants who exhibited high pointing accuracy, also when the sun was not visible. This study extends our understandings of orientation abilities of human foragers and provides the first behavioural evidence for sun compass use in humans.

Towards systematic and evidence-based conservation planning for western chimpanzees.

As animal populations continue to decline, frequently driven by large-scale land-use change, there is a critical need for improved environmental planning. While data-driven spatial planning is widely applied in conservation, as of yet it is rarely used for primates. The western chimpanzee (Pan troglodytes verus) declined by 80% within 24 years and was uplisted to Critically Endangered by the IUCN Red List of Threatened Species in 2016. To support conservation planning for western chimpanzees, we systematically identified geographic areas important for this taxon. We based our analysis on a previously published data set of modeled density distribution and on several scenarios that accounted for different spatial scales and conservation targets. Across all scenarios, typically less than one-third of areas we identified as important are currently designated as high-level protected areas (i.e., national park or IUCN category I or II). For example, in the scenario for protecting 50% of all chimpanzees remaining in West Africa (i.e., approximately 26,500 chimpanzees), an area of approximately 60,000 km2 was selected (i.e., approximately 12% of the geographic range), only 24% of which is currently designated as protected areas. The derived maps can be used to inform the geographic prioritization of conservation interventions, including protected area expansion, "no-go-zones" for industry and infrastructure, and conservation sites outside the protected area network. Environmental guidelines by major institutions funding infrastructure and resource extraction projects explicitly require corporations to minimize the negative impact on great apes. Therefore, our results can inform avoidance and mitigation measures during the planning phases of such projects. This study was designed to inform future stakeholder consultation processes that could ultimately integrate the conservation of western chimpanzees with national land-use priorities. Our approach may help in promoting similar work for other primate taxa to inform systematic conservation planning in times of growing threats.

Aggression by male bonobos against immature individuals does not fit with predictions of infanticide.

The selective advantage of male infanticide is enhancement of reproductive success of the aggressor. This implies that aggression is directed at individuals sired by others, infant loss shortens the mother's inter-birth interval, and the aggressor has a greater likelihood of siring the next offspring of the victims' mother. As these conditions are not always met, the occurrence of male infanticide is expected to vary, and hominoid primates offer an interesting example of variation in male infanticide. Infanticide has been reported in gorillas and chimpanzees but appears to be absent in orangutans and bonobos. One argument for the absence of infanticide in bonobos is reduction of male aggression. However, given that male aggression against immature individuals occurs and that females engage in behavior that is considered to be counterstrategy against male infanticide, the risk of male infanticide may pose a potential threat. Here, we explored whether aggression by male bonobos fits predictions of male infanticide. Male aggression toward immature individuals was rare and did not have lethal consequences, but the majority of observed cases exposed targets to risks of injury. Males did not target their own offspring less frequently than unrelated immatures, and the risk of being the target of male aggression increased with the targets' age. Overall, these results do not match the predictions of the adaptive male infanticide hypothesis. Instead, aggression by males may promote the emigration of the targets and older males may reinforce their superior status toward individuals that will soon compete for the same resources.

Bili-Uéré: A Chimpanzee Behavioural Realm in Northern Democratic Republic of Congo.

To improve our understanding of the evolutionary origins of culture and technology in humans, it is vital that we document the full extent of behavioural diversity in our great ape relatives. About half of the world's remaining chimpanzees (Pan spp.) live in the Democratic Republic of Congo (DRC), yet until now we have known almost nothing about their behaviour. Here we describe the insect-related tool technology of Bili-Uéré chimpanzees (Pan troglodytes schweinfurthii) inhabiting an area of at least a 50,000-km2 area in northern DRC, as well as their percussive technology associated with food processing. Over a 12-year period, we documented chimpanzee tools and artefacts at 20 survey areas and gathered data on dung, feeding remains and sleeping nests. We describe a new chimpanzee tool kit: long probes used to harvest epigaeic driver ants (Dorylus spp.), short probes used to extract ponerine ants and the arboreal nests of stingless bees, wands to dip for D. kohli and stout digging sticks used to access underground meliponine bee nests. Epigaeic Dorylus tools were significantly longer than the other tool types, and D. kohli tools were significantly thinner. Tools classified as terrestrial honey-digging sticks were a significant predictor for brushed and blunted tool ends, consistent with their presumed use. We describe two potential new tool types, an "ant scoop" and a "fruit hammer." We document an extensive percussive technology used to process termite mounds of Cubitermes sp. and Thoracotermes macrothorax and hard-shelled fruits such as Strychnos, along with evidence of the pounding open of African giant snails and tortoises. We encountered some geographic variation in behaviour: we found honey-digging tools, long driver ant probes and fruit-pounding sites only to the north of the Uele River; there were more epigaeic Dorylus tools to the north and more ponerine ant tools to the south. We found no evidence of termite-fishing, despite the availability of Macrotermes muelleri mounds throughout the region. This lack of evidence is consistent with the results of dung washes, which revealed a substantial proportion of driver ants, but no evidence of Macrotermes or other termites. Our results allow us to describe a new chimpanzee behavioural complex, characterised by a general similarity of multiple behaviours across a large, ecologically diverse region but with subtle differences in prey choice and techniques. We propose that this widespread and related suite of behaviours be referred to as the Bili-Uéré Chimpanzee Behavioural Realm. Possible explanations for this pattern are a recent chimpanzee expansion across the region and the interconnectedness of this population across at least the entirety of northern DRC.