Great Ape Species Research Articles

Comparative isotope ecology of African great apes

The isotope ecology of great apes is a useful reference for palaeodietary reconstructions in fossil hominins. As extant apes live in C3-dominated habitats, variation in isotope signatures is assumed to be low compared to hominoids exploiting C4-plant resources. However, isotopic differences between sites and between and within individuals are poorly understood due to the lack of vegetation baseline data. In this comparative study, we included all species of free-ranging African great apes (Pan troglodytes, Pan paniscus, Gorilla sp.). First, we explore differences in isotope baselines across different habitats and whether isotopic signatures in apes can be related to feeding niches (faunivory and folivory). Secondly, we illustrate how stable isotopic variations within African ape populations compare to other extant and extinct primates and discuss possible implications for dietary flexibility. Using 701 carbon and nitrogen isotope data points resulting from 148 sectioned hair samples and an additional collection of 189 fruit samples, we compare six different great ape sites. We investigate the relationship between vegetation baselines and climatic variables, and subsequently correct great ape isotope data to a standardized plant baseline from the respective sites. We obtained temporal isotopic profiles of individual animals by sectioning hair along its growth trajectory. Isotopic signatures of great apes differed between sites, mainly as vegetation isotope baselines were correlated with site-specific climatic conditions. We show that controlling for plant isotopic characteristics at a given site is essential for faunal data interpretation. While accounting for plant baseline effects, we found distinct isotopic profiles for each great ape population. Based on evidence from habituated groups and sympatric great ape species, these differences could possibly be related to faunivory and folivory. Dietary flexibility in apes varied, but temporal variation was overall lower than in fossil hominins and extant baboons, shifting from C3 to C4-resources, providing new perspectives on comparisons between extinct and extant primates.

Watering holes: The use of arboreal sources of drinking water by Old World monkeys and apes

Water is one of the most important components of an animal’s diet, as it is essential for life. Primates, as do most animals, procure water directly from standing or free-flowing sources such as pools, ponds and rivers, or indirectly by the ingestion of certain plant parts. The latter is frequently described as the main source of water for predominantly arboreal species. However, in addition to these, many species are known to drink water accumulated in tree-holes. This has been commonly observed in several arboreal New World primate species, but rarely reported systematically from Old World primates. Here, we report observations of this behaviour from eight great ape and Old World monkey species, namely chimpanzee, orangutan, siamang, western hoolock gibbon, northern pig-tailed macaque, bonnet macaque, rhesus macaque and the central Himalayan langur. We hypothesise three possible reasons why these primates drink water from tree-holes: (1) coping with seasonal or habitat-specific water shortages, (2) predator/human conflict avoidance, and (3) potential medicinal benefits. We also suggest some alternative hypotheses that should be tested in future studies. This behaviour is likely to be more prevalent than currently thought, and may have significant, previously unknown, influences on primate survival and health, warranting further detailed studies.

Quantifying ocular morphologies in extant primates for reliable interspecific comparisons

Cross-examining evidence from comparative morphology has become a way to support specific hypotheses in social cognition. In particular, ocular morphology has been compared across extant primate species in order to argue that humans present a unique morphology that enables also an uniquely human array of socio-cognitive functions—including a crucial role in the acquisition and practice of linguistic abilities. Even though some of these comparisons have relied on quantifiable dimensions of ocular morphology, other aspects of the comparison have been established qualitatively, based on subjective ratings. In this article, I present a new method that intends to restrain the focus of attention to one specific perceptual aspect of ocular morphology—namely, the contrast between scleral and iridal colors, as it is thought to enable (or ease) gaze following from other individuals. The method also allows for more reliable comparisons across extant primate species, as it relies on quantifiable measurements. I exemplify the potential of this method with a three-fold comparison of three great ape species ( Pongo pygmaeus , Pongo abelii , and Homo sapiens ).

Measuring Hair Cortisol Concentrations to Assess the Effect of Anthropogenic Impacts on Wild Chimpanzees (Pan troglodytes)

Non-human primates face major environmental changes due to increased human impacts all over the world. Although some species are able to survive in certain landscapes with anthropogenic impact, their long-term viability and fitness may be decreased due to chronic stress. Here we assessed long-term stress levels through cortisol analysis in chimpanzee hair obtained from sleeping nests in northwestern Uganda, in order to estimate welfare in the context of ecotourism, forest fragmentation with human-wildlife conflicts, and illegal logging with hunting activity (albeit not of primates), compared with a control without human contact or conflict. Concerning methodological issues, season [F(2,129) = 37.4, p < 0.0001, r2 = 0.18] and the age of nests [F(2,178) = 20.3, p < 0.0001, r2 = 0.11] significantly predicted hair cortisol concentrations (HCC). With regard to effects of anthropogenic impacts, our results neither showed elevation of HCC due to ecotourism, nor due to illegal logging compared to their control groups. We did, however, find significantly increased HCC in the fragment group compared to chimpanzees living in a nearby intact forest [F(1,88) = 5.0, p = 0.03, r2 = 0.20]. In conclusion, our results suggest that hair cortisol analysis is a powerful tool that can help understanding the impact of anthropogenic disturbances on chimpanzee well-being and could be applied to other great ape species.

Differences in the Visual Perception of Symmetric Patterns in Orangutans (Pongo pygmaeus abelii) and Two Human Cultural Groups: A Comparative Eye-Tracking Study.

Symmetric structures are of importance in relation to aesthetic preference. To investigate whether the preference for symmetric patterns is unique to humans, independent of their cultural background, we compared two human populations with distinct cultural backgrounds (Namibian hunter-gatherers and German town dwellers) with one species of non-human great apes (Orangutans) in their viewing behavior regarding symmetric and asymmetric patterns in two levels of complexity. In addition, the human participants were asked to give their aesthetic evaluation of a subset of the presented patterns. The results showed that humans of both cultural groups fixated on symmetric patterns for a longer period of time, regardless of the pattern’s complexity. On the contrary, Orangutans did not clearly differentiate between symmetric and asymmetric patterns, but were much faster in processing the presented stimuli and scanned the complete screen, while both human groups rested on the symmetric pattern after a short scanning time. The aesthetic evaluation test revealed that the fixation preference for symmetric patterns did not match with the aesthetic evaluation in the Hai//om group, whereas in the German group aesthetic evaluation was in accordance with the fixation preference in 60 percent of the cases. It can be concluded that humans prefer well-ordered structures in visual processing tasks, most likely because of a positive processing bias for symmetry, which Orangutans did not show in this task, and that, in humans, an aesthetic preference does not necessarily accompany the fixation preference.

Enduring voice recognition in bonobos.

Long-term social recognition is vital for species with complex social networks, where familiar individuals can encounter one another after long periods of separation. For non-human primates who live in dense forest environments, visual access to one another is often limited, and recognition of social partners over distances largely depends on vocal communication. Vocal recognition after years of separation has never been reported in any great ape species, despite their complex societies and advanced social intelligence. Here we show that bonobos, Pan paniscus, demonstrate reliable vocal recognition of social partners, even if they have been separated for five years. We experimentally tested bonobos’ responses to the calls of previous group members that had been transferred between captive groups. Despite long separations, subjects responded more intensely to familiar voices than to calls from unknown individuals - the first experimental evidence that bonobos can identify individuals utilising vocalisations even years after their last encounter. Our study also suggests that bonobos may cease to discriminate between familiar and unfamiliar individuals after a period of eight years, indicating that voice representations or interest could be limited in time in this species.

Young children spontaneously invent wild great apes' tool-use behaviours.

The variety and complexity of human-made tools are unique in the animal kingdom. Research investigating why human tool use is special has focused on the role of social learning: while non-human great apes acquire tool-use behaviours mostly by individual (re-)inventions, modern humans use imitation and teaching to accumulate innovations over time. However, little is known about tool-use behaviours that humans can invent individually, i.e. without cultural knowledge. We presented 2- to 3.5-year-old children with 12 problem-solving tasks based on tool-use behaviours shown by great apes. Spontaneous tool use was observed in 11 tasks. Additionally, tasks which occurred more frequently in wild great apes were also solved more frequently by human children. Our results demonstrate great similarity in the spontaneous tool-use abilities of human children and great apes, indicating that the physical cognition underlying tool use shows large overlaps across the great ape species. This suggests that humans are neither born with special physical cognition skills, nor that these skills have degraded due to our species’ long reliance of social learning in the tool-use domain.

Great ape Y Chromosome and mitochondrial DNA phylogenies reflect subspecies structure and patterns of mating and dispersal.

The distribution of genetic diversity in great ape species is likely to have been affected by patterns of dispersal and mating. This has previously been investigated by sequencing autosomal and mitochondrial DNA (mtDNA), but large-scale sequence analysis of the male-specific region of the Y Chromosome (MSY) has not yet been undertaken. Here, we use the human MSY reference sequence as a basis for sequence capture and read mapping in 19 great ape males, combining the data with sequences extracted from the published whole genomes of 24 additional males to yield a total sample of 19 chimpanzees, four bonobos, 14 gorillas, and six orangutans, in which interpretable MSY sequence ranges from 2.61 to 3.80 Mb. This analysis reveals thousands of novel MSY variants and defines unbiased phylogenies. We compare these with mtDNA-based trees in the same individuals, estimating time-to-most-recent common ancestor (TMRCA) for key nodes in both cases. The two loci show high topological concordance and are consistent with accepted (sub)species definitions, but time depths differ enormously between loci and (sub)species, likely reflecting different dispersal and mating patterns. Gorillas and chimpanzees/bonobos present generally low and high MSY diversity, respectively, reflecting polygyny versus multimale–multifemale mating. However, particularly marked differences exist among chimpanzee subspecies: The western chimpanzee MSY phylogeny has a TMRCA of only 13.2 (10.8–15.8) thousand years, but that for central chimpanzees exceeds 1 million years. Cross-species comparison within a single MSY phylogeny emphasizes the low human diversity, and reveals species-specific branch length variation that may reflect differences in long-term generation times.

Complex MHC Class I Gene Transcription Profiles and Their Functional Impact in Orangutans.

MHC haplotypes of humans and the African great ape species have one copy of the MHC-A, -B, and -C genes. In contrast, MHC haplotypes of orangutans, the Asian great ape species, exhibit variation in the number of gene copies. An in-depth analysis of the MHC class I gene repertoire in the two orangutan species, Pongo abelii and Pongo pygmaeus, is presented in this article. This analysis involved Sanger and next-generation sequencing methodologies, revealing diverse and complicated transcription profiles for orangutan MHC-A, -B, and -C. Thirty-five previously unreported MHC class I alleles are described. The data demonstrate that each orangutan MHC haplotype has one copy of the MHC-A gene, and that the MHC-B region has been subject to duplication, giving rise to at least three MHC-B genes. The MHC-B*03 and -B*08 lineages of alleles each account for a separate MHC-B gene. All MHC-B*08 allotypes have the C1-epitope motif recognized by killer cell Ig-like receptor. At least one other MHC-B gene is present, pointing to MHC-B alleles that are not B*03 or B*08. The MHC-C gene is present only on some haplotypes, and each MHC-C allotype has the C1-epitope. The transcription profiles demonstrate that MHC-A alleles are highly transcribed, whereas MHC-C alleles, when present, are transcribed at very low levels. The MHC-B alleles are transcribed to a variable extent and over a wide range. For those orangutan MHC class I allotypes that are detected by human monoclonal anti-HLA class I Abs, the level of cell-surface expression of proteins correlates with the level of transcription of the allele.

The Time Scale of Recombination Rate Evolution in Great Apes.

We present three linkage-disequilibrium (LD)-based recombination maps generated using whole-genome sequence data from 10 Nigerian chimpanzees, 13 bonobos, and 15 western gorillas, collected as part of the Great Ape Genome Project (Prado-Martinez J, et al. 2013. Great ape genetic diversity and population history. Nature 499:471-475). We also identified species-specific recombination hotspots in each group using a modified LDhot framework, which greatly improves statistical power to detect hotspots at varying strengths. We show that fewer hotspots are shared among chimpanzee subspecies than within human populations, further narrowing the time scale of complete hotspot turnover. Further, using species-specific PRDM9 sequences to predict potential binding sites (PBS), we show higher predicted PRDM9 binding in recombination hotspots as compared to matched cold spot regions in multiple great ape species, including at least one chimpanzee subspecies. We found that correlations between broad-scale recombination rates decline more rapidly than nucleotide divergence between species. We also compared the skew of recombination rates at centromeres and telomeres between species and show a skew from chromosome means extending as far as 10-15 Mb from chromosome ends. Further, we examined broad-scale recombination rate changes near a translocation in gorillas and found minimal differences as compared to other great ape species perhaps because the coordinates relative to the chromosome ends were unaffected. Finally, on the basis of multiple linear regression analysis, we found that various correlates of recombination rate persist throughout the African great apes including repeats, diversity, and divergence. Our study is the first to analyze within- and between-species genome-wide recombination rate variation in several close relatives.

Gaze Duration Biases for Colours in Combination with Dissonant and Consonant Sounds: A Comparative Eye-Tracking Study with Orangutans.

Research on colour preferences in humans and non-human primates suggests similar patterns of biases for and avoidance of specific colours, indicating that these colours are connected to a psychological reaction. Similarly, in the acoustic domain, approach reactions to consonant sounds (considered as positive) and avoidance reactions to dissonant sounds (considered as negative) have been found in human adults and children, and it has been demonstrated that non-human primates are able to discriminate between consonant and dissonant sounds. Yet it remains unclear whether the visual and acoustic approach–avoidance patterns remain consistent when both types of stimuli are combined, how they relate to and influence each other, and whether these are similar for humans and other primates. Therefore, to investigate whether gaze duration biases for colours are similar across primates and whether reactions to consonant and dissonant sounds cumulate with reactions to specific colours, we conducted an eye-tracking study in which we compared humans with one species of great apes, the orangutans. We presented four different colours either in isolation or in combination with consonant and dissonant sounds. We hypothesised that the viewing time for specific colours should be influenced by dissonant sounds and that previously existing avoidance behaviours with regard to colours should be intensified, reflecting their association with negative acoustic information. The results showed that the humans had constant gaze durations which were independent of the auditory stimulus, with a clear avoidance of yellow. In contrast, the orangutans did not show any clear gaze duration bias or avoidance of colours, and they were also not influenced by the auditory stimuli. In conclusion, our findings only partially support the previously identified pattern of biases for and avoidance of specific colours in humans and do not confirm such a pattern for orangutans.

In vitro strain in human metacarpal bones during striking: testing the pugilism hypothesis of hominin hand evolution.

The hands of hominins (i.e. bipedal apes) are distinguished by skeletal proportions that are known to enhance manual dexterity but also allow the formation of a clenched fist. Because male-male physical competition is important in the mating systems of most species of great apes, including humans, we tested the hypothesis that a clenched fist protects the metacarpal bones from injury by reducing the level of strain during striking. We used cadaver arms to measure in vitro strain in metacarpals during forward strikes with buttressed and unbuttressed fist postures and during side slaps with an open palm. If the protective buttressing hypothesis is correct, the clenched fist posture should substantially reduce strain in the metacarpal bones during striking and therefore reduce the risk of fracture. Recorded strains were significantly higher in strikes in which the hand was secured in unbuttressed and slapping postures than in the fully buttressed posture. Our results suggest that humans can safely strike with 55% more force with a fully buttressed fist than with an unbuttressed fist and with twofold more force with a buttressed fist than with an open-hand slap. Thus, the evolutionary significance of the proportions of the hominin hand may be that these are the proportions that improved manual dexterity while at the same time making it possible for the hand to be used as a club during fighting.

High-throughput RNA sequencing reveals structural differences of orthologous brain-expressed genes between western lowland gorillas and humans.

The human brain and human cognitive abilities are strikingly different from those of other great apes despite relatively modest genome sequence divergence. However, little is presently known about the interspecies divergence in gene structure and transcription that might contribute to these phenotypic differences. To date, most comparative studies of gene structure in the brain have examined humans, chimpanzees, and macaque monkeys. To add to this body of knowledge, we analyze here the brain transcriptome of the western lowland gorilla (Gorilla gorilla gorilla), an African great ape species that is phylogenetically closely related to humans, but with a brain that is approximately one-third the size. Manual transcriptome curation from a sample of the planum temporale region of the neocortex revealed 12 protein-coding genes and one noncoding-RNA gene with exons in the gorilla unmatched by public transcriptome data from the orthologous human loci. These interspecies gene structure differences accounted for a total of 134 amino acids in proteins found in the gorilla that were absent from protein products of the orthologous human genes. Proteins varying in structure between human and gorilla were involved in immunity and energy metabolism, suggesting their relevance to phenotypic differences. This gorilla neocortical transcriptome comprises an empirical, not homology- or prediction-driven, resource for orthologous gene comparisons between human and gorilla. These findings provide a unique repository of the sequences and structures of thousands of genes transcribed in the gorilla brain, pointing to candidate genes that may contribute to the traits distinguishing humans from other closely related great apes.

Extreme selective sweeps independently targeted the X chromosomes of the great apes

The unique inheritance pattern of the X chromosome exposes it to natural selection in a way that is different from that of the autosomes, potentially resulting in accelerated evolution. We perform a comparative analysis of X chromosome polymorphism in 10 great ape species, including humans. In most species, we identify striking megabase-wide regions, where nucleotide diversity is less than 20% of the chromosomal average. Such regions are found exclusively on the X chromosome. The regions overlap partially among species, suggesting that the underlying targets are partly shared among species. The regions have higher proportions of singleton SNPs, higher levels of population differentiation, and a higher nonsynonymous-to-synonymous substitution ratio than the rest of the X chromosome. We show that the extent to which diversity is reduced is incompatible with direct selection or the action of background selection and soft selective sweeps alone, and therefore, we suggest that very strong selective sweeps have independently targeted these specific regions in several species. The only genomic feature that we can identify as strongly associated with loss of diversity is the location of testis-expressed ampliconic genes, which also have reduced diversity around them. We hypothesize that these genes may be responsible for selective sweeps in the form of meiotic drive caused by an intragenomic conflict in male meiosis.

Looking ahead? Computerized maze task performance by chimpanzees (Pan troglodytes), rhesus monkeys (Macaca mulatta), capuchin monkeys (Cebus apella), and human children (Homo sapiens).

Human and nonhuman primates are not mentally constrained to the present. They can remember the past and-at least to an extent-anticipate the future. Anticipation of the future ranges from long-term prospection such as planning for retirement to more short-term future-oriented cognition such as planning a route through a maze. Here we tested a great ape species (chimpanzees), an Old World monkey species (rhesus macaques), a New World monkey species (capuchin monkeys), and human children on a computerized maze task. All subjects had to move a cursor through a maze to reach a goal at the bottom of the screen. For best performance on the task, subjects had to "plan ahead" to the end of the maze to move the cursor in the correct direction, avoid traps, and reverse directions if necessary. Mazes varied in difficulty. Chimpanzees were better than both monkey species, and monkeys showed a particular deficit when moving away from the goal or changing directions was required. Children showed a similar pattern to monkeys regarding the effects of reversals and moves away from the goal, but their overall performance in terms of correct maze completion was similar to the chimpanzees. The results highlight similarities as well as differences in planning across species and the role that inhibitory control may play in future-oriented cognition in primates.

Playing hide and seek with primates: A comparative study of Theory of Mind

The ability to anticipate how another individual will behave is a critical skill in the animal kingdom. Humans typically deal with this problem using their Theory of Mind (ToM), i.e. the ability to decipher beliefs and intentions of others. Although non-human primates (apes in particular) exhibit elaborate social behavior, there is no established consensus on whether this mirrors some form of sophisticated ToM. We suggest that ToM sophistication can be operationally defined in terms of the depth of recursive beliefs, as in think that you think that I think.... Importantly, mutual social interactions may call for highly recursive beliefs: not only do I think that you think, but I also think that you think that I think... This study aims at comparing how primate species (including humans) differ with respect to this form of ToM sophistication. The main novelty of our experimental approach is to estimate each primate’s ToM sophistication from its performance in repeated interactive games that are calibrated using information theoretic models of (artificial) recursive ToM. We first validated our model in human subjects and then tested 6 species of captive non-human primates (3 species of great apes: orangutans, gorillas, chimpanzees, and 3 species of old-world monkeys: lion-tailed macaques, rhesus macaques and collared mangabeys) in an adaptation of the task used in the human study. Our paradigm consisted in a “hide and seek” game in which individuals played against a human opponent (a familiar zoo keeper). In each trial, the individual had to infer where food had been hidden by the experimenter opponent (either in his right or left hand). In fact, the location of the food was chosen by learning algorithms, some of which endowed with artificial ToM. Critically, we varied the sophistication of these algorithms, yielding three conditions, ranging from a control (a simple random biased sequence) to a mildly sophisticated ToM agent (1-ToM). The profile of primates’ performances against the different algorithms was then used to infer their ToM sophistication. Preliminary results show that the pattern of performance of nonhuman primates differ from those of humans. Although most nonhuman primates tended to win in the control condition, they lost against the 1-ToM agent, in contrast to the human group (that won). Surprisingly, comparing the performances between the non-human primates' species did not reveal any significant inter-species difference. Therefore, great apes did not perform better than monkeys in our task. Interestingly, variability in the performances actually seems more to rely on inter-individual differences rather than on inter-species differences. Moreover, in every species, at least one individual exhibited mild (but statistically significant) ToM sophistication. Taken together, these data indicate that the distribution of ToM sophistication within each of these non-human primate species is quite heterogeneous. Further analyses are in progress to identify predictive factors (e.g. familiarity with humans, age, motivation and frustration during the game) that could explain away inter-individual differences, thus eventually revealing (or not) inter‑species differences [Authors Aurore San‑Galli and Marie Devaine contributed equally to this work].

Bornean orangutan geophagy: analysis of ingested and control soils.

Geophagy among orangutans is the most poorly documented in contrast to the knowledge of soil-eating practices of other great ape species. Observations of soil consumption by orangutans in the Sungai Wain Forest Preserve (Wanariset) of Borneo are presented, along with physico-mineral-chemical analyses of the ingested soil in an effort to understand what might stimulate the activity. The consumed soils are: light colored, not excessively weathered by normal standards, higher in the clay size fraction relative to controls, and are comprised of a mix of clay minerals without any specificity of 1:1, 2:1 and/or 2:1:1 (Si:Al) species. The geophagic soils contain chlorides below detection limits, effectively eliminating salt as a stimulus. Soil chemical and geochemical analyses confirm that orangutans prefer soils with pH levels near or above 4.0, while controls are consistently lower (pH=3.5-4.0), a considerable difference in acidity for at least four out of six soils consumed. Geochemical analysis shows Al, Fe and K are high in the consumed vs control samples; higher Al follows from higher clay percentages in the consumed earth. Iron and K may play physiological roles, but Fe is mostly in the ferrous form (Fe(+2)) and may not be readily taken up by the animals. The preferential choice of consumed samples, with pH above 4.0 and higher clay contents, may promote a more beneficial intestinal environment.

Children Conform to the Behavior of Peers; Other Great Apes Stick With What They Know

All primates learn things from conspecifics socially, but it is not clear whether they conform to the behavior of these conspecifics—if conformity is defined as overriding individually acquired behavioral tendencies in order to copy peers’ behavior. In the current study, chimpanzees, orangutans, and 2-year-old human children individually acquired a problem-solving strategy. They then watched several conspecific peers demonstrate an alternative strategy. The children switched to this new, socially demonstrated strategy in roughly half of all instances, whereas the other two great-ape species almost never adjusted their behavior to the majority’s. In a follow-up study, children switched much more when the peer demonstrators were still present than when they were absent, which suggests that their conformity arose at least in part from social motivations. These results demonstrate an important difference between the social learning of humans and great apes, a difference that might help to account for differences in human and nonhuman cultures.

Lineage sorting in apes.

Recombination allows different parts of the genome to have different genealogical histories. When a species splits in two, allelic lineages sort into the two descendant species, and this lineage sorting varies along the genome. If speciation events are close in time, the lineage sorting process may be incomplete at the second speciation event and lead to gene genealogies that do not match the species phylogeny. We review different recent approaches to model lineage sorting along the genome and show how it is possible to learn about population sizes, natural selection, and recombination rates in ancestral species from application of these models to genome alignments of great ape species.

Food Neophobia and Social Learning Opportunities in Great Apes

Finding food resources and maintaining a balanced diet are major concerns for all animals. A compromise between neophobia and neophilia is hypothesised to enable animals to enlarge their diet while limiting the risk of poisoning. However, little is known about how primates respond to novel food items and whether their use is socially transmitted. By comparing how four different species of great apes respond to novel food items, we investigated how differences in physiology (digestive tract size and microbial content), habitats (predictability of food availability), and social systems (group size and composition) affect their response toward novelty. We presented two familiar foods, one novel fruit, four novel aromatic plants from herbal medicine, and kaolin to captive chimpanzees (Pan troglodytes), western gorillas (Gorilla gorilla), Bornean orangutans (Pongo pygmaeus) and Sumatran orangutans (Pongo abelii). We recorded smelling, approach-taste delays, ingestion, interindividual observations, and food transfers with continuous sampling. We found that behaviors differed between the apes: chimpanzees were the most cautious species and observed their conspecifics handling the items more frequently than the other apes. Close observations and food transfers were extremely rare in gorillas in comparison to orangutans and chimpanzees. We suggest that a low neophobia level reflects an adaptive response to digestive physiological features in gorillas and to unpredictable food availability in orangutans. Social interactions appeared to be predominant in chimpanzees and in both orangutan species to overcome food neophobia. They reflect higher social tolerance and more opportunities for social learning and cultural transmission in a feeding context.