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Indirect wildlife population surveying largely depends upon counts of artifacts of behavior (e.g., nests or dung). Likelihood to encounter these artifacts is derived from both artifact production and decay, and variability in production behavior is considered to contribute minimally to inaccuracy in wildlife estimation. Here, we demonstrate how ignoring behavioral variability leads to significant population misestimation, using an example of an endangered ape, the bonobo (Pan paniscus). Until now, a single estimate of nest construction rate has been used to extrapolate bonobo densities, assumed to be broadly representative of bonobo sign production behavior. We estimated nest construction rates across seasons and social groups at the Kokolopori Bonobo Reserve, Democratic Republic of the Congo, and find nest construction rates in bonobos to be highly variable across populations as well as seasonal within populations. Failure to account for this variability led to degradation in the accuracy of bonobo population density estimates, accounting for a likely overestimation of bonobo numbers by 34%, and at worst as high as 80%. With this example, we demonstrate that failure to account for inter‐ and intrapopulation behavioral variation compromises the ability to estimate both relative and absolute wildlife abundances. We argue that variation in sign production is but one of the several potential ways that behavioral variability can affect conservation monitoring, should be measured across contexts whenever possible, and must be considered in population estimation confidence intervals. With increasing attention to behavioral variability as a potential tool for conservation, conservationists must also account for the impact that behavioral variability can have upon wildlife population estimation. Our results underline the importance of observational research to wildlife monitoring schemes as a critical component of conservation management. We discuss the avenues through which behavioral variability is likely to impact wildlife monitoring accuracy and precision and propose potential approaches for accounting for behavioral variability in wildlife monitoring. [ABSTRACT FROM AUTHOR]

The arbitrary relationship between signifier and signified is one of the features responsible for language's extreme lability, adaptability, and expressiveness. Understanding this arbitrariness and its emergence is essential in any account of the evolution of language. To shed light on the phylogeny of the phenomenon, comparative data examining the relationship between signal form and function in the communication systems of non-humans is central. Here we report the results of a study on the production and usage the whistle-high hoot call combination (W + HH) from two distant populations of wild bonobos (Pan paniscus): Lui Kotale, DRC, and Kokolopori, DRC. We find that the context in which bonobos produce the W + HHs varies systematically between populations. Our results suggest that variation in W + HH production may represent an example of signal-adjustment optionality, a key component of arbitrariness., (© 2024. The Author(s).)

Researchers investigating the evolution of human aggression look to our closest living relatives, bonobos (Pan paniscus) and chimpanzees (Pan troglodytes), as valuable sources of comparative data. 1 , 2 Males in the two species exhibit contrasting patterns: male chimpanzees sexually coerce females 3 , 4 , 5 , 6 , 7 , 8 and sometimes kill conspecifics, 9 , 10 , 11 , 12 whereas male bonobos exhibit less sexual coercion 13 , 14 and no reported killing. 13 Among the various attempts to explain these species differences, the self-domestication hypothesis proposes negative fitness consequences of male aggression in bonobos. 2 , 15 , 16 Nonetheless, the extent to which these species differ in overall rates of aggression remains unclear due to insufficiently comparable observation methods. 17 , 18 , 19 , 20 , 21 , 22 , 23 We used 14 community-years of focal follow data-the gold standard for observational studies 24 -to compare rates of male aggression in 3 bonobo communities at the Kokolopori Bonobo Reserve, Democratic Republic of Congo, and 2 chimpanzee communities at Gombe National Park, Tanzania. As expected, given that females commonly outrank males, we found that bonobos exhibited lower rates of male-female aggression and higher rates of female-male aggression than chimpanzees. Surprisingly, we found higher rates of male-male aggression among bonobos than chimpanzees even when limiting analyses to contact aggression. In both species, more aggressive males obtained higher mating success. Although our findings indicate that the frequency of male-male aggression does not parallel species difference in its intensity, they support the view that contrary to male chimpanzees, whose reproductive success depends on strong coalitions, male bonobos have more individualistic reproductive strategies. 25 ., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Reconstructing pathways to human peace can be hampered by superficial evaluations of similar processes in nonhuman species. A deeper understanding of bonobo social systems allows us to reevaluate the preconditions for peace to gain a greater insight on the evolutionary timescale of peace emergence.

Cooperation beyond familial and group boundaries is core to the functioning of human societies, yet its evolution remains unclear. To address this, we examined grooming, coalition, and food-sharing patterns in bonobos ( Pan paniscus ), one of our closest living relatives whose rare out-group tolerance facilitates interaction opportunities between groups. We show that, as in humans, positive assortment supports bonobo cooperation across borders. Bonobo cooperative attitudes toward in-group members informed their cooperative relationships with out-groups, in particular, forming connections with out-group individuals who also exhibited high cooperation tendencies. Our findings show that cooperation between unrelated individuals across groups without immediate payoff is not exclusive to humans and suggest that such cooperation can emerge in the absence of social norms or strong cultural dispositions.

Reproductive inequality, or reproductive skew, drives natural selection, but has been difficult to assess, particularly for males in species with promiscuous mating and slow life histories, such as bonobos ( Pan paniscus ) and chimpanzees ( Pan troglodytes ). Although bonobos are often portrayed as more egalitarian than chimpanzees, genetic studies have found high male reproductive skew in bonobos. Here, we discuss mechanisms likely to affect male reproductive skew in Pan , then re-examine skew patterns using paternity data from published work and new data from the Kokolopori Bonobo Reserve, Democratic Republic of Congo and Gombe National Park, Tanzania. Using the multinomial index ( M ), we found considerable overlap in skew between the species, but the highest skew occurred among bonobos. Additionally, for two of three bonobo communities, but no chimpanzee communities, the highest ranking male had greater siring success than predicted by priority-of-access. Thus, an expanded dataset covering a broader demographic range confirms that bonobos have high male reproductive skew. Detailed comparison of data from Pan highlights that reproductive skew models should consider male-male dynamics including the effect of between-group competition on incentives for reproductive concessions, but also female grouping patterns and factors related to male-female dynamics including the expression of female choice. This article is part of the theme issue 'Evolutionary ecology of inequality'.

Characteristics of food availability and distribution are key components of a species' ecology. Objective ecological surveying used in animal behavior research does not consider aspects of selection by the consumer and therefore may produce imprecise measures of availability. We propose a method to integrate ecological sampling of an animal's environment into existing behavioral data collection systems by using the consumer as the surveyor. Here, we evaluate the consumer-centric method (CCM) of assessing resource availability for its ability to measure food resource abundance, distribution, and dispersion. This method catalogs feeding locations observed during behavioral observation and uses aggregated data to characterize these ecological metrics. We evaluated the CCM relative to traditional vegetation plot surveying using accumulated feeding locations across 3 years visited by a tropical frugivore, the bonobo ( Pan paniscus ), and compared it with data derived from over 200 vegetation plots across their 50 km 2 + home ranges. We demonstrate that food species abundance estimates derived from the CCM are comparable to those derived from traditional vegetation plot sampling in less than 2 years of data collection, and agreement improved when accounting for aspects of consumer selectivity in objective vegetation plot sampling (e.g., tree size minima). Density correlated between CCM and plot-derived estimates and was relatively insensitive to home range inclusion and other species characteristics, however, it was sensitive to sampling frequency. Agreement between the methods in relative distribution of resources performed better across species than expected by chance, although measures of dispersion correlated poorly. Once tested in other systems, the CCM may provide a robust measure of food availability for use in relative food availability indices and can be incorporated into existing observational data collection. The CCM has an advantage over traditional sampling methods as it incorporates sampling biases relevant to the consumer, thereby serving as a promising method for animal behavioral research., Competing Interests: The authors have no conflict of interest to declare., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

In animal societies, control over resources and reproduction is often biased towards one sex. Yet, the ecological and evolutionary underpinnings of male-female power asymmetries remain poorly understood. We outline a comprehensive framework to quantify and predict the dynamics of male-female power relationships within and across mammalian species. We show that male-female power relationships are more nuanced and flexible than previously acknowledged. We then propose that enhanced reproductive control over when and with whom to mate predicts social empowerment across ecological and evolutionary contexts. The framework explains distinct pathways to sex-biased power: coercion and male-biased dimorphism constitute a co-evolutionary highway to male power, whereas female power emerges through multiple physiological, morphological, behavioural, and socioecological pathways., Competing Interests: Declaration of interests No interests are declared., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Sociality is widespread among group-living primates and is beneficial in many ways. Sociality amongst female bonobos (Pan paniscus) has been proposed to have evolved as a female counterstrategy to male infanticide and sexual coercion. In male-philopatric bonobo societies, females mostly form relationships with unrelated females. Among these social relationships, it has been proposed that females with infants (also referred to as mothers) tend to have strong relationships with each other (mother-bonding hypothesis). In this paper, we use the case of an infant death in a group of wild bonobos in the Kokolopori Bonobo Reserve, Democratic Republic of Congo, to test this hypothesis. By using dyadic sociality indices for grooming, proximity, and aggression, we investigated whether the infant death influenced dyadic relationships the mother had with other group members. Before the infant death, grooming index (GI) and proximity index (PI) scores were the highest between the focal mother and another mother. After the death, the relationship of this mother dyad weakened, as indicated by lower GI and PI scores, whereas the relationship of another mother dyad became stronger. Aggression index scores among the mothers were comparable before and after the death, suggesting that changes in mother affiliative relationships were not a by-product of changes in overall interaction frequencies. Also, PI scores increased between the focal mother and three non-mothers after the death. Collectively, the shift in social dynamics between the focal mother and other group members after the infant death partially supported the mother-bonding hypothesis., (© 2022. The Author(s).)

Emergency Ultrasound Training for and with Medical Students Abstract. Practical basic skills in sonography are a mandatory part of Swiss medical schools since 2018. The universities of Basel and Bern teach students the content of the POCUS component "Basic Emergency Sonography" of the SGUM and have developed the e-learning tool "POCUS Emergency Sonography" for this purpose in cooperation. By using this innovative blend- ed learning concept, students acquire basic skills in sonography and can build upon this know-how in their further education.

Comparative studies on tool technologies in extant primates, especially in our closest living relatives, offer a window into the evolutionary foundations of tool use in hominins. Whereas chimpanzee tool technology is well studied across populations, the scarcity of described tool technology in wild populations of our other closest living relative, the bonobo, is a mystery. Here we provide a first report of the tool use repertoire of the Kokolopori bonobos and describe in detail the use of leaf-umbrellas during rainfall, with the aim to improve our knowledge of bonobo tool use capacity in the wild. The tool use repertoire of the Kokolopori bonobos was most similar to that of the nearby population of Wamba and comprised eight behaviors, none in a foraging context. Further, over a 6-month period we documented 44 instances of leaf-umbrella use by 22 individuals from three communities, suggesting that this behavior is habitual. Most leaf-umbrella tool users were adult females, and we observed a nonadult using a leaf-umbrella on only a single occasion. While the study and theory of tool technologies is often based on the use of tools in foraging tasks, tool use in bonobos typically occurs in nonforaging contexts across populations. Therefore, incorporating both foraging and nonforaging contexts into our theoretical framework is essential if we wish to advance our understanding of the evolutionary trajectories of tool technology in humans., (© 2021 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Here we show that sexual signaling affects patterns of female spatial association differently in chimpanzees and bonobos, indicating its relevance in shaping the respective social systems. Generally, spatial association between females often mirrors patterns and strength of social relationships and cooperation within groups. While testing for proposed differences in female-female associations underlying female coalition formation in the species of the genus Pan, we find only limited evidence for a higher female-female gregariousness in bonobos. While bonobo females exhibited a slightly higher average number of females in their parties, there is neither a species difference in the time females spent alone, nor in the number of female party members in the absence of sexually attractive females. We find that the more frequent presence of maximally tumescent females in bonobos is associated with a significantly stronger increase in the number of female party members, independent of variation in a behavioural proxy for food abundance. This indicates the need to look beyond ecology when explaining species differences in female sociality as it refutes the idea that the higher gregariousness among bonobo females is driven by ecological factors alone and highlights that the temporal distribution of female sexual receptivity is an important factor to consider when studying mammalian sociality., (© 2021. The Author(s).)

Nonstaple food is a food resource which sole consumption does not allow the maintenance of regular physiological functions, thus constituting a minor portion of an individual's diet. Many primates consume nonstaple food such as meat, insects, and fungi. Hypotheses on the dietary importance of nonstaple food include its role as fallback food and as source of specific nutrients. We tested these two hypotheses by investigating mycophagy (i.e., the consumption of fungi) in a population of wild bonobos in the Kokolopori Bonobo Reserve, DRC. Specifically, we examined the relationship between fungus consumption and various factors relevant to bonobo feeding ecology (i.e., fruit abundance and the consumption of other food types). Additionally, we measured the deviation from linear travel when bonobos searched for fungi to evaluate the nature of fungus consumption (e.g., opportunistic or targeted). Lastly, we examined the nutritional content of the major fungus species consumed (Hysterangium bonobo) to test whether this food item was potentially consumed as source of specific nutrients. We found that bonobos spent a higher proportion of their time feeding on fungi when fruit abundance was higher, indicating that fungi were not consumed as a fallback food. Moreover, bonobos deviated from linear travel when visiting fungus patches more than observed when visiting fruit patches, suggesting that they actively sought out fungi. Lastly, initial analyses suggest that H. bonobo samples contained high concentration of sodium. Collectively, these results suggest that subterranean fungi appear to be attractive food source to Kokolopori bonobos, and that mycophagy may serve to supplement nutrients, like sodium, in bonobo diet., (© 2021 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Humans harbor diverse communities of microorganisms, the majority of which are bacteria in the gastrointestinal tract. These gut bacterial communities in turn host diverse bacteriophage (hereafter phage) communities that have a major impact on their structure, function, and, ultimately, human health. However, the evolutionary and ecological origins of these human-associated phage communities are poorly understood. To address this question, we examined fecal phageomes of 23 wild nonhuman primate taxa, including multiple representatives of all the major primate radiations. We find relatives of the majority of human-associated phages in wild primates. Primate taxa have distinct phageome compositions that exhibit a clear phylosymbiotic signal, and phage-superhost codivergence is often detected for individual phages. Within species, neighboring social groups harbor compositionally and evolutionarily distinct phageomes, which are structured by superhost social behavior. Captive nonhuman primate phageome composition is intermediate between that of their wild counterparts and humans. Phage phylogenies reveal replacement of wild great ape-associated phages with human-associated ones in captivity and, surprisingly, show no signal for the persistence of wild-associated phages in captivity. Together, our results suggest that potentially labile primate-phage associations have persisted across millions of years of evolution. Across primates, these phylosymbiotic and sometimes codiverging phage communities are shaped by transmission between groupmates through grooming and are dramatically modified when primates are moved into captivity., Competing Interests: The authors declare no competing interest.

Study Objective: Delirium is frequently observed in the postoperative and intensive care unit (ICU) population. Due to the multifactorial origin of delirium and according to international guidelines (e.g., American Geriatrics Society; Prevention and Management of Pain, Agitation/Sedation, Delirium, Immobility, and Sleep Disruption (PADIS) guideline), there are several but no incontestable options for prevention and symptomatic treatment. The purpose of the Baden PRIDe (Prevention and Reduction of Incidence of postoperative Delirium) trial was to determine whether postoperative cognitive dysfunction and delirium could be prevented by the combination of possible preventive agents such as haloperidol and ketamine. In addition, pre- and postoperative levels of the biomarkers cortisol, neuron specific enolase (NSE) and S100β were measured to investigate their dynamics in delirious and non-delirious patients after surgery., Design: The Baden PRIDe Trial was an investigator-initiated, phase IV, two-centre, randomised, placebo-controlled, double-blind clinical trial., Setting: Perioperative care., Patients: 182 adult patients that underwent elective or emergency surgery under general or combined (i.e., general and regional) anaesthesia., Interventions: Pre-anaesthetic, pharmacologic prevention of postoperative brain dysfunction with haloperidol, ketamine, and the combination of both vs. placebo., Measurements: Assessment of cognitive performance pre- and postoperatively with the MMSE, the DOS, the Nursing Delirium Screening Scale (Nu-DESC) or the Intensive Care Delirium Screening Checklist (ICDSC) during ICU stay., Main Results: None of the three study arms - haloperidol, ketamine, or both drugs combined - was significantly superior to placebo for prevention of postoperative brain dysfunction and delirium (P = 0.39). Measured levels of postoperative cortisol were significantly higher in delirious patients. S-100β levels were significantly higher in all postoperative outcome groups (cognitive impairment, delirium, no cognitive decline), whereas postoperative NSE levels declined in all groups., Conclusions: The study results offer no possibility for a novel recommendation for prevention of postoperative cognitive decline including delirium. Perioperative S-100β trajectories in patients with cognitive deterioration suggest affection of glial cells in particular., Trial Registration: ClinicalTrials.govNCT02433041; registered on April 7, 2015., (Copyright © 2020 The Authors. Published by Elsevier Inc. All rights reserved.)

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., (Copyright © 2021 The Authors. Published by Elsevier Inc. All rights reserved.)

Competing Interests: Conflict of interest The author declares that he has no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

The importance of cultural processes to behavioural diversity in our closest living relatives is central to revealing the evolutionary origins of human culture. However, the bonobo is often overlooked as a candidate model. Further, a prominent critique to many examples of proposed animal cultures is premature exclusion of environmental confounds known to shape behavioural phenotypes. We addressed these gaps by investigating variation in prey preference between neighbouring bonobo groups that associate and overlap space use. We find group preference for duiker or anomalure hunting otherwise unexplained by variation in spatial usage, seasonality, or hunting party size, composition, and cohesion. Our findings demonstrate that group-specific behaviours emerge independently of the local ecology, indicating that hunting techniques in bonobos may be culturally transmitted. The tolerant intergroup relations of bonobos offer an ideal context to explore drivers of behavioural phenotypes, the essential investigations for phylogenetic constructs of the evolutionary origins of culture., Competing Interests: LS, FW, MS No competing interests declared, (© 2020, Samuni et al.)

Several theories have been generated to understand the socio-cognitive mechanisms underlying the unique cooperative abilities of humans. The 'interdependence hypothesis' posits first, that the cognitive dimension of human cooperation evolved in contexts when several individuals needed to act together to achieve a common goal, like when hunting large prey. Second, the more interdependent individuals are, the more likely they are to provide services to conspecifics in other contexts. Alternatively, the 'social tolerance hypothesis' proposes that higher social tolerance allows conspecifics to cooperate more efficiently and with a wider range of partners. We conducted the first field experimental evaluation of both hypotheses in our closest living relatives by contrasting chimpanzees to the less interdependent but more tolerant bonobos. We compared each species' performance during a cooperative task: informing conspecifics about a danger. We presented Gaboon viper models to 82 individuals from five wild communities. Chimpanzees arriving late at the snake were significantly more likely to have heard a call and less likely to startle, indicating that chimpanzees were better informed about the presence of the threat than bonobos. This stems from clear species differences in how individuals adjusted their calling decisions to the level of information already available. Chimpanzees were more likely to call and produced more alarm calls when they had not yet heard a call, whereas bonobos did so when they already heard a call. Our results confirm the link between interdependence and cooperation performance. These species differences were most likely driven by differences in motivation rather than in cognitive capacities because both species tended to consider audience knowledge in their decision to call. Our results inform theories on the evolution of human cooperation by linking inter-group competition pressure and in-group cooperative motivation and/or capability.

Despite their ubiquity, in most cases little is known about the impact of eukaryotic parasites on their mammalian hosts. Comparative approaches provide a powerful method to investigate the impact of parasites on host ecology and evolution, though two issues are critical for such efforts: controlling for variation in methods of identifying parasites and incorporating heterogeneity in sampling effort across host species. To address these issues, there is a need for standardized methods to catalogue eukaryotic parasite diversity across broad phylogenetic host ranges. We demonstrate the feasibility of a metabarcoding approach for describing parasite communities by analysing faecal samples from 11 nonhuman primate species representing divergent lineages of the primate phylogeny and the full range of sampling effort (i.e. from no parasites reported in the literature to the best-studied primates). We detected a number of parasite families and regardless of prior sampling effort, metabarcoding of only ten faecal samples identified parasite families previously undescribed in each host (x̅ = 8.5 new families per species). We found more overlap between parasite families detected with metabarcoding and published literature when more research effort-measured as the number of publications-had been conducted on the host species' parasites. More closely related primates and those from the same continent had more similar parasite communities, highlighting the biological relevance of sampling even a small number of hosts. Collectively, results demonstrate that metabarcoding methods are sensitive and powerful enough to standardize studies of eukaryotic parasite communities across host species, providing essential new tools for macroecological studies of parasitism., (© 2019 The Authors. Molecular Ecology Resources published by John Wiley & Sons Ltd.)

In some species habitual same-sex sexual behavior co-occurs with high levels of intra-sexual alliance formation, suggesting that these behaviors may be linked. We tested for such a link by comparing behavioral and physiological outcomes of sex with unrelated same- and opposite-sex partners in female bonobos (Pan paniscus). We analyzed behavioral outcomes following 971 sexual events involving n = 19 female and n = 8 male adult and sub-adult members of a wild, habituated bonobo community. We additionally collected n = 143 urine samples before and after sexual interactions to non-invasively measure oxytocin (OT), which modulates female sexual behavior and facilitates cooperation in other species. The majority of sexual events (65%) consisted of female same-sex genito-genital rubbing (or GG-rubbing). Female dyads engaged in significantly more sexual interactions than did inter-sexual dyads, and females were more likely to remain within close proximity to their partners following GG-rubbing. Females also exhibited greater increases in urinary OT following GG-rubbing compared with copulations, indicating a physiological basis for increased motivation to cooperate among females. The frequency of coalitionary support among non-kin was positively predicted by the frequency of sexual interactions for female as well opposite-sex dyads, although coalitionary support tended to be more frequent among females. The emergence of habitual same-sex sexual behavior may have been an important step in the evolution of cooperation outside of kinship and pair-bonds in one of our closest phylogenetic relatives., (Copyright © 2019 The Authors. Published by Elsevier Inc. All rights reserved.)

Primate individuals use a variety of strategies in intergroup encounters, from aggression to tolerance; however, recent focus on the evolution of either warfare or peace has come at the cost of characterizing this variability. We identify evolutionary advantages that may incentivize tolerance toward extra-group individuals in humans and nonhuman primates, including enhanced benefits in the domains of transfer, mating, and food acquisition. We highlight the role these factors play in the flexibility of gorilla, chimpanzee, bonobo, and human behavior. Given humans have an especially broad range of intergroup behavior, we explore how the human foraging ecology, especially large spatial and temporal fluctuations in resource availability, may have selected for a greater reliance on tolerant between-community relationships-relationships reinforced by status acquisition and cultural institutions. We conclude by urging careful, theoretically motivated study of behavioral flexibility in intergroup encounters in humans and the nonhuman great apes., (© 2019 The Authors. Evolutionary Anthropology: Issues, News, and Reviews published by Wiley Periodicals, Inc.)

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., (© 2019 Wiley Periodicals, Inc.)

Temperament and personality research in humans and nonhuman animals measures behavioral variation in individual, population, or species-specific traits with implications for survival and fitness, such as social status, foraging, and mating success [1-5]. Curiosity and risk-taking tendencies have been studied extensively across taxa by measuring boldness and exploration responses to experimental novelty exposure [3, 4, 6-15]. Here, we conduct a natural field experiment using wildlife monitoring technology to test variation in the reaction of wild great apes (43 groups of naive chimpanzees, bonobos, and western gorillas across 14 field sites in Africa) to a novel object, the camera trap. Bonobo and gorilla groups demonstrated a stronger looking impulse toward the camera trap device compared to chimpanzees, suggesting higher visual attention and curiosity. Bonobos were also more likely to show alarm and other fearful behaviors, although such neophobic (and conversely, neophilic) responses were generally rare. Among all three species, individuals looked at cameras longer when they were young, were associating with fewer individuals, and did not live near a long-term research site. Overall, these findings partially validate results from great ape novelty paradigms in captivity [7, 8]. We further suggest that species-typical leadership styles [16] and social and environmental effects, including familiarity with humans, best explain novelty responses of wild great apes. In sum, this study illustrates the feasibility of large-scale field experiments and the importance of both intrinsic and extrinsic factors in shaping animal curiosity. VIDEO ABSTRACT., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Coalitions among males during within group conflicts have a strong influence on the competitive and social environment within social groups. To evaluate possible variation in the occurrence of such coalitions in our closest relatives, chimpanzees and bonobos, we compared male aggression and coalitionary behavior in two natural communities, one of each species, with a similar size and composition. Furthermore we compared affiliative behavior that might be related to coalition formation among males. We found higher frequencies of aggression and a greater likelihood to form coalitions during within-group conflicts among wild male chimpanzees at Taï compared to wild male bonobos at LuiKotale. The species differed in the predominant sex of the male coalition partners, with male bonobos forming coalitions more often with females, while male chimpanzees formed coalitions more often with other males. Compared to male bonobos, male chimpanzees showed higher rates of grooming and tended to reconcile more conflicts with other males. Overall our results showed lower frequencies of reconciliation among bonobos than those described in captivity and at artificial feeding sites. These findings add to the evidence that male cooperation and conflict resolution are potentially very different in bonobos and chimpanzees, despite the fact that these two species are closely related, live in multi-male, multi-female communities with a high degree of fission-fusion dynamics and have female-biased migration patterns. Given the correlation between aggressive, cooperative and some affiliative patterns within the species in our study, we hypothesize that the fitness benefits of male relationships are greater in chimpanzees compared to bonobos., (© 2017 Wiley Periodicals, Inc.)

In several group-living species, individuals' social preferences are thought to be influenced by cooperation. For some societies with fission-fusion dynamics, sex-specific association patterns reflect sex differences in cooperation in within- and between-group contexts. In our study, we investigated this hypothesis further by comparing sex-specific association patterns in two closely related species, chimpanzees and bonobos, which differ in the level of between-group competition and in the degree to which sex and kinship influence dyadic cooperation. Here, we used long-term party composition data collected on five chimpanzee and two bonobo communities and assessed, for each individual of 10 years and older, the sex of its top associate and of all conspecifics with whom it associated more frequently than expected by chance. We found clear species differences in association patterns. While in all chimpanzee communities males and females associated more with same-sex partners, in bonobos males and females tended to associate preferentially with females, but the female association preference for other females is lower than in chimpanzees. Our results also show that, for bonobos (but not for chimpanzees), association patterns were predominantly driven by mother-offspring relationships. These species differences in association patterns reflect the high levels of male-male cooperation in chimpanzees and of mother-son cooperation in bonobos. Finally, female chimpanzees showed intense association with a few other females, and male chimpanzees showed more uniform association across males. In bonobos, the most differentiated associations were from males towards females. Chimpanzee male association patterns mirror fundamental human male social traits and, as in humans, may have evolved as a response to strong between-group competition. The lack of such a pattern in a closely related species with a lower degree of between-group competition further supports this notion., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Objectives: Female bonobos (Pan paniscus) are characterized as highly affiliative and cooperative, but few studies have quantified the strength and stability of female intra-sexual relationships or explored how variation in social relationships influences cooperation. We measure female social preferences, identify causes of variation in preferences, and test whether variation in social preferences predicts food sharing or coalitionary support., Methods: Data were collected over 3 years from females in the Bompusa community at LuiKotale, DRC. We measured genetic relatedness and constructed social preference indices for party association, proximity, grooming, GG-rubbing and aggression. We identified preferred social partners based on permutation tests and measured stability using Mantel tests. We used factor analysis to identify inter-relationships between preference indices and used LMMs to test whether variation in social preferences was explained by relatedness, rank differences, having dependent young or co-residency time. We used GLMMs to test whether variation in social preferences predicted food sharing or coalitionary support., Results: All females had preferred non-kin partners for proximity, grooming or GG-rubbing, but only grooming preferences were stable across years. Association indices were higher among lactating females, and aggression was lower among females with longer co-residency times. The factor analysis identified one factor, representing proximity and GG-rubbing preferences, labeled behavioral coordination. Dyads with higher levels of behavioral coordination were more likely to share food., Conclusions: Female bonobos exhibit stable, differentiated grooming relationships outside of kinship and philopatry. Females also exhibit flexible proximity and GG-rubbing preferences that may facilitate cooperation with a wider range of social partners., (© 2017 Wiley Periodicals, Inc.)

Dietary ecology of extant great apes is known to respond to environmental conditions such as climate and food availability, but also to vary depending on social status and life history characteristics. Bonobos (Pan paniscus) live under comparatively steady ecological conditions in the evergreen rainforests of the Congo Basin. Bonobos are an ideal species for investigating influences of sociodemographic and physiological factors, such as female reproductive status, on diet. We investigate the long term dietary pattern in wild but fully habituated bonobos by stable isotope analysis in hair and integrating a variety of long-term sociodemographic information obtained through observations. We analyzed carbon and nitrogen stable isotopes in 432 hair sections obtained from 101 non-invasively collected hair samples. These samples represented the dietary behavior of 23 adult bonobos from 2008 through 2010. By including isotope and crude protein data from plants we could establish an isotope baseline and interpret the results of several general linear mixed models using the predictors climate, sex, social rank, reproductive state of females, adult age and age of infants. We found that low canopy foliage is a useful isotopic tracer for tropical rainforest settings, and consumption of terrestrial herbs best explains the temporal isotope patterns we found in carbon isotope values of bonobo hair. Only the diet of male bonobos was affected by social rank, with lower nitrogen isotope values in low-ranking young males. Female isotope values mainly differed between different stages of reproduction (cycling, pregnancy, lactation). These isotopic differences appear to be related to changes in dietary preference during pregnancy (high protein diet) and lactation (high energy diet), which allow to compensate for different nutritional needs during maternal investment., Competing Interests: The authors have declared that no competing interests exist.

Within- and between-species variation in male mating strategies has been attributed to a multitude of factors including male competitive ability and the distribution of fertile females across space and time. Differences in energy balance across and within males allow for the identification of some of the trade-offs associated with certain social and mating strategies. Bonobos live in groups with a high degree of fission-fusion dynamics, there is co-dominance between the sexes and a linear dominance hierarchy among males. Males compete over access to females, breeding is aseasonal, and females exhibit sexual swellings over extended time periods. In this study we use urinary C-peptide (UCP) levels in male bonobos (Pan paniscus) obtained from 260 urine samples from a wild bonobo community, to quantify male energy balance during mate competition and levels of gregariousness in the species. Although high ranking males are more aggressive, spend more time in proximity to maximally tumescent females, and have higher mating frequencies, we found no indication that mate guarding or mate competition affected male energy balance. Our results showed a positive correlation between monthly mean UCP levels and mean party size. When traveling in large parties, high ranking males had higher UCP levels than those of the low ranking males. These results support the hypothesis that patterns of fission-fusion dynamics in bonobos are either linked to energy availability in the environment or to the energetic costs of foraging. The finding of a rank-bias in UCP levels in larger parties could also reflect an increase in contest competition among males over access to food., (Copyright © 2015. Published by Elsevier Inc.)

The emotional mediation hypothesis proposes a mediating role of social bonds in the exchange of services. This model predicts that the form of short-term exchange of services depends on the relationship between the individuals involved. Here, we test this prediction in the exchange of grooming among males in a wild bonobo community for which close relatedness could be excluded. As bonobo males hardly engage in food sharing or agonistic support, grooming is mainly exchanged for grooming. While overall grooming, both given and received, correlates across dyads and within sessions, the form of grooming exchange within a given session differs according to dyadic association preferences. Individuals with a higher tendency to associate, ergo more familiar individuals, exhibit larger time differences and reduced reciprocation in consecutive grooming bouts than less familiar individuals. These results support the idea that emotional components are involved in the exchange of services between unrelated individuals.

In long-lived social mammals such as primates, individuals can benefit from social bonds with close kin, including their mothers. In the patrilocal chimpanzee (Pan troglodytes spp.) and bonobo (Pan paniscus), sexually mature males reside and reproduce in their natal groups and can retain post-dependency bonds with their mothers, while immatures of both sexes might also have their paternal grandmothers available. However, quantitative information on the proportion of males and immatures that co-reside with both types of these close female relatives is limited for both species. Combining genetic parentage determination and group composition data from five communities of wild chimpanzees and three communities of wild bonobos, we estimated the frequency of co-residence between (1) mature males and their mothers, and (2) immature males and females and their paternal grandmothers. We found that adult males resided twice as frequently with their mothers in bonobos than in chimpanzees, and that immature bonobos were three times more likely to possess a living paternal grandmother than were immature chimpanzees. Patterns of female and male survivorship from studbook records of captive individuals of both species suggest that mature bonobo females survive longer than their chimpanzee counterparts, possibly contributing to the differences observed in mother-son and grandmother-immature co-residency levels. Taking into account reports of bonobo mothers supporting their sons' mating efforts and females sharing food with immatures other than their own offspring, our findings suggest that life history traits may facilitate maternal and grandmaternal support more in bonobos than in chimpanzees.

A positive interaction is any interaction between individuals of the same or different species (mutualism) that provides a benefit to both partners such as increased fitness. Here we focus on seed dispersal mutualism between an animal (bonobo, Pan paniscus) and a plant (velvet tamarind trees, Dialium spp.). In the LuiKotale rainforest southwest of Salonga National Park, Democratic Republic of Congo, seven species of the genus Dialium account for 29.3% of all trees. Dialium is thus the dominant genus in this forest. Dialium fruits make up a large proportion of the diet of a habituated bonobo community in this forest. During the 6 months of the fruiting season, more than half of the bonobos' feeding time is devoted to Dialium fruits. Furthermore, Dialium fruits contribute a considerable proportion of sugar and protein to bonobos' dietary intake, being among the richest fruits for these nutrients. Bonobos in turn ingest fruits with seeds that are disseminated in their feces (endozoochory) at considerable distances (average: 1.25 km after 24 hr of average transit time). Endozoochory through the gut causes loss of the cuticle protection and tegumentary dormancy, as well as an increase in size by water uptake. Thus, after gut passage, seeds are better able to germinate. We consider other primate species as a potential seed disperser and conclude that Dialium germination is dependent on passage through bonobo guts. This plant-animal interaction highlights positive effects between two major organisms of the Congo basin rainforest, and establishes the role of the bonobo as an efficient disperser of Dialium seeds. Periodicals, Inc., (© 2013 Wiley Periodicals, Inc.)

The rapid disruption of tropical forests probably imperils global biodiversity more than any other contemporary phenomenon. With deforestation advancing quickly, protected areas are increasingly becoming final refuges for threatened species and natural ecosystem processes. However, many protected areas in the tropics are themselves vulnerable to human encroachment and other environmental stresses. As pressures mount, it is vital to know whether existing reserves can sustain their biodiversity. A critical constraint in addressing this question has been that data describing a broad array of biodiversity groups have been unavailable for a sufficiently large and representative sample of reserves. Here we present a uniquely comprehensive data set on changes over the past 20 to 30 years in 31 functional groups of species and 21 potential drivers of environmental change, for 60 protected areas stratified across the world’s major tropical regions. Our analysis reveals great variation in reserve ‘health’: about half of all reserves have been effective or performed passably, but the rest are experiencing an erosion of biodiversity that is often alarmingly widespread taxonomically and functionally. Habitat disruption, hunting and forest-product exploitation were the strongest predictors of declining reserve health. Crucially, environmental changes immediately outside reserves seemed nearly as important as those inside in determining their ecological fate, with changes inside reserves strongly mirroring those occurring around them. These findings suggest that tropical protected areas are often intimately linked ecologically to their surrounding habitats, and that a failure to stem broad-scale loss and degradation of such habitats could sharply increase the likelihood of serious biodiversity declines.

Cortisol excretion in males of group living species is often associated with social rank and competition for oestrous females. Rank-related patterns of cortisol levels can be used to study mechanisms of rank maintenance and costs associated with mate competition. Bonobos (Pan paniscus) are interesting because males form a linear dominance hierarchy but are not dominant over females and therefore aggressive male-male competition over access to females alone is not considered to be a successful reproductive strategy. In this study on social correlates of urinary cortisol in wild male bonobos, we investigated the relationship between cortisol levels and several aspects of mate competition, including male rank, aggression rates, and association time with oestrous females. We found that cortisol levels correlated positively with dominance rank when oestrous females were present, but not when they were absent. This result is consistent with the idea that aggressive behaviour plays a minor role in maintenance of high rank. While aggression received from males and females explained within-individual variation in cortisol levels, it was the time spent in association with oestrous females that best explained between-individual variation in male cortisol levels. The observed increase in male cortisol may be associated with spatial proximity to oestrous females and could result from anticipated aggression from other group members, reduced feeding time in the males, or a combination of both., (Copyright © 2012 Elsevier Inc. All rights reserved.)

In primates, age, sex, and social status can strongly influence access to food resources. In Pan, these criteria are assumed to influence access to vertebrate meat. However, the significance of meat in terms of its role in the nutrition of Pan is still debated. Here we present a study using stable carbon and nitrogen isotope ratios in hair samples from habituated, wild bonobos (Pan paniscus) to explore these issues. Over a period of 5 mo hair samples were collected from fresh bonobo nests at LuiKotale, Democratic Republic of Congo. Hair samples were assigned to known individuals and were of sufficient length to allow the evaluation of isotopic variation over several months. Samples of plant foods and sympatric fauna were also analyzed. The δ(13)C and δ(15)N results of the bonobo hair were remarkably homogeneous over time and for the group as a whole. There are no differences in diet between the sexes. Within the group of males, however, there was a positive correlation between dominance status and δ(15)N. The isotopic data indicate that the contribution of fauna to bonobo diet is marginal and that plant food is the dietary protein source. In only some cases did elevated δ(15)N hair values correlate with observed faunivory and not correspond to the δ(15)N measured in the dominant plant foods. Given the large variation in hunting and meat eating of Pan across the African continent, the detection of seasonal changes in faunivory by elevated δ(15)N values in sectioned ape hair is a promising approach.

Variation in male mating success is often related to rank differences. Males who are unable to monopolize oestrous females alone may engage in coalitions, thus enhancing their mating success. While studies on chimpanzees and dolphins suggest that coalitions are independent of kinship, information from female philopatric species shows the importance of kin support, especially from mothers, on the reproductive success of females. Therefore, one might expect a similar effect on sons in male philopatric species. We evaluate mating success determinants in male bonobos using data from nine male individuals from a wild population. Results reveal a steep, linear male dominance hierarchy and a positive correlation between dominance status and mating success. In addition to rank, the presence of mothers enhances the mating success of sons and reduces the proportion of matings by the highest ranking male. Mothers and sons have high association rates and mothers provide agonistic aid to sons in conflicts with other males. As bonobos are male-philopatric and adult females occupy high dominance status, maternal support extends into adulthood and females have the leverage to intervene in male conflicts. The absence of female support to unrelated males suggests that mothers gain indirect fitness benefits by supporting their sons.

We present evidence for the consumption of a diurnal, arboreal, group living primate by bonobos. The digit of an immature black mangabey (Lophocebus aterrimus) was found in the fresh feces of a bonobo (Pan paniscus) at the Lui Kotale study site, Democratic Republic of Congo. In close proximity to the fecal sample containing the remains of the digit, we also found a large part of the pelt of a black mangabey. Evidence suggests that the Lui Kotale bonobos consume more meat than other bonobo populations and have greater variation in the mammalian species exploited than previously thought [Hohmann & Fruth, Folia primatologica 79:103-110]. The current finding supports Stanford's argument [Current Anthropology 39:399-420] that some differences in the diet and behavior between chimpanzees (P. troglodytes) and bonobos are an artefact of the limited number of bonobo study populations. If bonobos did obtain the monkey by active hunting, this would challenge current evolutionary models relating the intra-specific aggression and violence seen in chimpanzees and humans to hunting and meat consumption [Wrangham, Yearbook of Physical Anthropology 42:1-30]., ((c) 2008 Wiley-Liss, Inc.)