Behavior Of Apes Research Articles

Dental findings in wild great apes from macerated skull analysis.

Oral health is a crucial aspect of overall well-being in both humans and nonhuman primates. Understanding the oral pathologies and dental conditions in apes can provide valuable insights into their evolutionary history, dietary habits, and overall health. The present study evaluates dental findings in wild great apes from museum specimens to gain insights into the influence of natural nutrition on dental health. Complete macerated skulls of wild, adult great apes from the collection of the Museum of Natural History, Berlin, Germany, were examined. We analyzed skulls of 53 gorillas (Gorilla gorilla), 63 chimpanzees (Pan troglodytes), and 41 orangutans (Pongo spp.). For each skull, we recorded wear of dental hard tissues (Lussi and Ganss index), carious lesions, and periodontal bone loss. Incisal and occlusal dental hard tissue defects were found in all skulls, as well as considerable external staining. In all species, incisors and canines showed the greatest loss of tissue, followed by molars. The wear of molars decreased from the first to the third molars, premolars showed the least pronounced defects. Some individuals had apical osteolytic defects along with severe dental hard tissue loss with pulp involvement or after dental trauma, respectively (n = 5). Our study did not observe any carious lesions among the examined great ape skulls. However, we did find evidence for localized or generalized periodontal bone loss in a subset of the specimens (n = 3 chimpanzees, n = 7 orangutans). The natural diet and foraging behavior of great apes induces abrasion and attrition of dental hard tissue but does not yield carious lesions. The occurrence of periodontitis in individual apes indicates that the natural circumstances can induce periodontal bone loss even in the wild, despite physiological nutrition.

Evolution of the human tongue and emergence of speech biomechanics.

The tongue is one of the organs most central to human speech. Here, the evolution and species-unique properties of the human tongue is traced, via reference to the apparent articulatory behavior of extant non-human great apes, and fossil findings from early hominids - from a point of view of articulatory phonetics, the science of human speech production. Increased lingual flexibility provided the possibility of mapping of articulatory targets, possibly via exaptation of manual-gestural mapping capacities evident in extant great apes. The emergence of the human-specific tongue, its properties, and morphology were crucial to the evolution of human articulate speech.

Friends in high places: Interspecific grooming between chimpanzees and primate prey species in Budongo Forest

While cases of interspecies grooming have been reported in primates, no comprehensive cross-site review has been published about this behavior in great apes. Only a few recorded observations of interspecies grooming events between chimpanzees and other primate species have been reported in the wild, all of which have thus far been in Uganda. Here, we review all interspecies grooming events recorded for the Sonso community chimpanzees in Budongo Forest Reserve, Uganda, adding five new observations to the single, previously reported event from this community. A new case of interspecies play involving three juvenile male chimpanzees and a red-tailed monkey is also detailed. All events took place between 1993 and 2021. In all of the six interspecific grooming events from Budongo, the ‘groomer’ was a female chimpanzee between the ages of 4–6 years, and the ‘recipient’ was a member of the genus Cercopithecus. In five of these events, chimpanzee groomers played with the tail of their interspecific grooming partners, and except for one case, initiated the interaction. In three cases, chimpanzee groomers smelled their fingers after touching distinct parts of the receiver’s body. While a single function of chimpanzee interspecies grooming remains difficult to determine from these results, our review outlines and assesses some hypotheses for the general function of this behavior, as well as some of the costs and benefits for both the chimpanzee groomers and their sympatric interspecific receivers. As allogrooming is a universal behavior in chimpanzees, investigating the ultimate and proximate drivers of chimpanzee interspecies grooming may reveal further functions of allogrooming in our closest living relatives, and help us to better understand how chimpanzees distinguish between affiliative and agonistic species and contexts.

Free hand hitting of stone-like objects in wild gorillas

The earliest stone tool types, sharp flakes knapped from stone cores, are assumed to have played a crucial role in human cognitive evolution. Flaked stone tools have been observed to be accidentally produced when wild monkeys use handheld stones as tools. Holding a stone core in hand and hitting it with another in the absence of flaking, free hand hitting, has been considered a requirement for producing sharp stone flakes by hitting stone on stone, free hand percussion. We report on five observations of free hand hitting behavior in two wild western gorillas, using stone-like objects (pieces of termite mound). Gorillas are therefore the second non-human lineage primate showing free-hand hitting behavior in the wild, and ours is the first report for free hand hitting behavior in wild apes. This study helps to shed light on the morphofunctional and cognitive requirements for the emergence of stone tool production as it shows that a prerequisite for free hand percussion (namely, free hand hitting) is part of the spontaneous behavioral repertoire of one of humans’ closest relatives (gorillas). However, the ability to combine free hand hitting with the force, precision, and accuracy needed to facilitate conchoidal fracture in free hand percussion may still have been a critical watershed for hominin evolution.

Comparison of the arm‐lowering performance between Gorilla and Homo through musculoskeletal modeling

AbstractObjectivesContrary to earlier hypotheses, a previous biomechanical analysis indicated that long‐documented morphological differences between the shoulders of humans and apes do not enhance the arm‐raising mechanism. Here, we investigate a different interpretation: the oblique shoulder morphology that is shared by all hominoids but humans enhances the arm‐lowering mechanism.Materials and methodsMusculoskeletal models allow us to predict performance capability to quantify the impact of muscle soft‐tissue properties and musculoskeletal morphology. In this study, we extend the previously published gorilla shoulder model by adding glenohumeral arm‐lowering muscles, then comparing the arm‐lowering performance to that of an existing human model. We further use the models to disentangle which morphological aspects of the shoulder affect arm‐lowering capacity and result in interspecific functional differences.ResultsOur results highlight that arm‐lowering capacity is greater in Gorilla than in Homo. The enhancement results from greater maximum isometric force capacities and moment arms of two important arm‐lowering muscles, teres major, and pectoralis major. More distal muscle insertions along the humerus together with a more oblique shoulder configuration cause these greater moment arms.DiscussionThe co‐occurrence of improved arm‐lowering capacity and high‐muscle activity at elevation angles used during vertical climbing highlight the importance of a strong arm‐lowering mechanism for arboreal locomotor behavior in nonhuman apes. Therefore, our findings reveal certain skeletal shoulder features that are advantageous in an arboreal context. These results advance our understanding of adaptation in living apes and can improve functional interpretations of the hominin fossil record.

Talar trochlear morphology may not be a good skeletal indicator of locomotor behavior in humans and great apes

To reconstruct locomotor behaviors of fossil hominins and understand the evolution of bipedal locomotion in the human lineage, it is important to clarify the functional morphology of the talar trochlea in humans and extant great apes. Therefore, the present study aimed to investigate the interspecific-differences of the talar trochlear morphology among humans, chimpanzees, gorillas, and orangutans by means of cone frustum approximation to calculate an apical angle and geometric morphometrics for detailed variability in the shape of the talar trochlea. The apical angles in gorillas and orangutans were significantly greater than those in humans and chimpanzees, but no statistical difference was observed between humans and chimpanzees, indicating that the apical angle did not necessarily correspond with the degree of arboreality in hominoids. The geometric morphometrics revealed clear interspecific differences in the trochlear morphology, but no clear association between the morphological characteristics of the trochlea and locomotor behavior was observed. The morphology of the trochlea may not be a distinct skeletal correlate of locomotor behavior, possibly because the morphology is determined not only by locomotor behavior, but also by other factors such as phylogeny and body size.

Morphological differences in the calcaneus among extant great apes investigated by three-dimensional geometric morphometrics

Investigating the morphological differences of the calcaneus in humans and great apes is crucial for reconstructing locomotor repertories of fossil hominins. However, morphological variations in the calcaneus of the great apes (chimpanzees, gorillas, and orangutans) have not been sufficiently studied. This study aims to clarify variations in calcaneal morphology among great apes based on three-dimensional geometric morphometrics. A total of 556 landmarks and semilandmarks were placed on the calcaneal surface to calculate the principal components of shape variations among specimens. Clear interspecific differences in calcaneal morphology were extracted, corresponding to the degree of arboreality of the three species. The most arboreal orangutans possessed comparatively more slender calcaneal tuberosity and deeper pivot region of the cuboid articular surface than chimpanzees and gorillas. However, the most terrestrial gorillas exhibited longer lever arm of the triceps surae muscle, larger peroneal trochlea, more concave plantar surface, more inverted calcaneal tuberosity, more everted cuboid articular surface, and more prominent plantar process than the orangutans and chimpanzees. These interspecific differences possibly reflect the functional adaptation of the calcaneus to locomotor behavior in great apes. Such information might be useful for inferring foot functions and reconstructing the locomotion of fossil hominoids and hominids.

The Pan social brain: An evolutionary history of neurochemical receptor genes and their potential impact on sociocognitive differences

Humans have unique cognitive capacities that, compared with apes, are not only simply expressed as a higher level of general intelligence, but also as a quantitative difference in sociocognitive skills. Humans’ closest living relatives, bonobos (Pan paniscus), and chimpanzees (Pan troglodytes), show key between-species differences in social cognition despite their close phylogenetic relatedness, with bonobos arguably showing greater similarities to humans. To better understand the evolution of these traits, we investigate the neurochemical mechanisms underlying sociocognitive skills by focusing on variation in genes encoding proteins with well-documented roles in mammalian social cognition: the receptors for vasopressin (AVPR1A), oxytocin (OXTR), serotonin (HTR1A), and dopamine (DRD2). Although these genes have been well studied in humans, little is known about variation in these genes that may underlie differences in social behavior and cognition in apes. We comparatively analyzed sequence data for 33 bonobos and 57 chimpanzees, together with orthologous sequence data for other apes. In all four genes, we describe genetic variants that alter the amino acid sequence of the respective receptors, raising the possibility that ligand binding or signal transduction may be impacted. Overall, bonobos show 57% more fixed substitutions than chimpanzees compared with the ancestral Pan lineage. Chimpanzees, show 31% more polymorphic coding variation, in line with their larger historical effective population size estimates and current wider distribution. An extensive literature review comparing allelic changes in Pan with known human behavioral variants revealed evidence of homologous evolution in bonobos and humans (OXTR rs4686301(T) and rs237897(A)), while humans and chimpanzees shared OXTR rs2228485(A), DRD2 rs6277(A), and DRD2 rs11214613(A) to the exclusion of bonobos. Our results offer the first in-depth comparison of neurochemical receptor gene variation in Pan and put forward new variants for future behavior–genotype association studies in apes, which can increase our understanding of the evolution of social cognition in modern humans.

Human–Ape Interactions in a Zoo Setting: Gorillas and Orangutans Modify Their Behavior Depending upon Human Familiarity

ABSTRACTUnderstanding how human presence influences animal behavior, specifically the behavior of nonhuman primates, has important implications for methodology in cognitive and behavioral studies, for our understanding of human–animal relationships and for animal welfare in captive settings. Conducted in the San Diego Zoo, this study partially replicates an existing study on human–animal relationships, which focused on familiarity effects on the human-directed behavior of great apes residing in the Toronto Zoo. Such a replication in a new location and with different individuals is necessary to expand sample size and to better understand how ontogeny and living environment influence human-directed ape behavior. All-occurrence sampling of human-directed behavior was used to gather data on two gorilla groups and one orangutan group, all housed at the San Diego Zoo. The aim was to compare ape-initiated behaviors toward zoo staff with apeinitiated visitor-directed behavior, in accordance with predictions of the Human–Animal Relationship (HAR) model. Overall, human-directed behaviors were more frequent toward visitors than toward staff, mainly caused by increased visitor-directed visual behavior, specifically brief glances. Given that visual monitoring has been seen as fearful behavior, potentially related to predator monitoring, this would confirm the expectation that lack of familiarity will lead to increased visual vigilance. However, the visitor-directed visual behaviors may not be antagonistic or fearful. Considering the specific behaviors and exhibit designs, curiosity may be an equally likely explanation in this context. Aiming to also understand whether these groups exhibited species differences in human-directed behavior, it was found that orangutans engaged in more affiliative behavior overall than gorillas. Comparing our results with the previous study, we conclude that while effects of familiarity are present, those effects are modulated by both individual ontogeny and exhibit design. While the HAR model is partially supported, additional factors may need to be included to fully interpret the observed behavior. Additional research is needed to further understand the effects of species, sex, and ontogeny on the predictions of the HAR model.

Chimpanzee Cooperation Is Fast and Independent From Self-Control

Large-scale cooperation is a hallmark of our species and appears to be unique among primates. Yet the evolutionary mechanisms that drove the emergence of humanlike patterns of cooperation remain unclear. Studying the cognitive processes underlying cooperative behavior in apes, our closest living relatives, can help identify these mechanisms. Accordingly, we employed a novel test battery to assess the willingness of 40 chimpanzees to donate resources, instrumentally help others, and punish a culpable thief. We found that chimpanzees were faster to make prosocial than selfish choices and that more prosocial individuals made the fastest responses. Further, two measures of self-control did not predict variation in prosocial responding, and individual performance across cooperative tasks did not covary. These results show that chimpanzees and humans share key cognitive processes for cooperation, despite differences in the scope of their cooperative behaviors.

The comparative neuroprimatology 2018 (CNP-2018) road map for research on How the Brain Got Language

Abstract We present a new road map for research on “How the Brain Got Language” that adopts an EvoDevoSocio perspective and highlights comparative neuroprimatology – the comparative study of brain, behavior and communication in extant monkeys and great apes – as providing a key grounding for hypotheses on the last common ancestor of humans and monkeys (LCA-m) and chimpanzees (LCA-c) and the processes which guided the evolution LCA-m → LCA-c → protohumans → H. sapiens. Such research constrains and is constrained by analysis of the subsequent, primarily cultural, evolution of H. sapiens which yielded cultures involving the rich use of language.

Introducing a special issue

Abstract The paper introduces a Special Issue of Interaction Studies which includes 21 papers based on presentations and discussion at a workshop entitled “How the Brain Got Language: Towards a New Road Map.” Unifying themes include the comparative study of brain, behavior and communication in monkeys, apes and humans, and an EvoDevoSocio framework for approaching biological and cultural evolution within a shared perspective. The final article of the special issue builds on the previous papers to present “The Comparative Neuroprimatology 2018 (CNP-2018) Road Map for Research on How the Brain Got Language.”

Parallel overinterpretation of behavior of apes and corvids.

The report by Kabadayi and Osvath (Science, 357(6347), 202-204, 2017) does not demonstrate planning in ravens. The behavior of corvids and apes is fascinating and will be best appreciated through well-designed experiments that explicitly test alternative explanations and that are interpreted without unjustified anthropomorphic embellishment.

Tie one on: 'nest tying' by wild chimpanzees at Bulindi-a variant of a universal great ape behavior?

With data accumulating from a growing pool of chimpanzee field studies, new behaviors as well as novel variants on common behaviors continue to be described. Nest construction is a universal behavior in wild great apes. Among chimpanzee populations, reported variation in nest building behavior mostly reflects environmental constraints. Despite the ubiquity of nest making by chimpanzees, only ground nesting has been recognized as a behavioral variant, potentially determined by both environmental and social factors. In a study of nests made by chimpanzees (Pan troglodytes schweinfurthii) in Bulindi, Uganda, I identified a hitherto undescribed nest construction technique, termed 'nest tying'. Five observed nests lacked strong weight-bearing structures beneath them, such as large branches or a supporting trunk. Instead, the nests appeared 'tied' (or 'tethered') to an adjacent trunk by looping leafy stems or palm fronds around it and interweaving these into the nest mattress, securing the nest against the trunk; thus, nest tying presumably functions to provide addedstability and support. This preliminary report presents a description of the observed nests. Irrespective of whether nest tying constitutes true knot making-commonly considered absent in wild great apes-this nest construction technique would seem to require advanced dexterity and a sophisticated understanding of the mechanical properties of the plants used. Forest fragments in Bulindi are highly degraded. Thus, nest tying-and construction of integrated nests (i.e., utilizing multiple plants, often small trees and shrubs) generally-may be promoted by a relative paucity of suitable nesting trees at this site. Still, insofar as nest building is learned in chimpanzees, different construction techniques including nest tying are potentially acquired through social learning. Further investigation is required to ascertain the prevalence and acquisition of this nest construction technique at Bulindi, and to verify its presence or absence in other habitats.

Unexpected terrestrial hand posture diversity in wild mountain gorillas.

Gorillas, along with chimpanzees and bonobos, are ubiquitously described as 'knuckle-walkers.' Consequently, knuckle-walking (KW) has been featured pre-eminently in hypotheses of the pre-bipedal locomotor behavior of hominins and in the evolution of locomotor behavior in apes. However, anecdotal and behavioral accounts suggest that mountain gorillas may utilize a more complex repertoire of hand postures, which could alter current interpretations of African ape locomotion and its role in the emergence of human bipedalism. Here we documented hand postures during terrestrial locomotion in wild mountain gorillas to investigate the frequency with which KW and other hand postures are utilized in the wild. Multiple high-speed cameras were used to record bouts of terrestrial locomotion of 77 habituated mountain gorillas at Bwindi Impenetrable National Park (Uganda) and Volcanoes National Park (Rwanda). We captured high-speed video of hand contacts in 8% of the world's population of mountain gorillas. Our results reveal that nearly 40% of these gorillas used "non-KW" hand postures, and these hand postures constituted 15% of all hand contacts. Some of these "non-KW" hand postures have never been documented in gorillas, yet match hand postures previously identified in orangutans. These results highlight a previously unrecognized level of hand postural diversity in gorillas, and perhaps great apes generally. Although present at lower frequencies than KW, we suggest that the possession of multiple, versatile hand postures present in wild mountain gorillas may represent a shared feature of the African ape and human clade (or even great ape clade) rather than KW per se.

Factors Influencing the Survival of Sympatric Gorilla (Gorilla gorilla gorilla) and Chimpanzee (Pan troglodytes troglodytes) Nests

Accurate and precise surveys of primate abundance provide the basis for understanding species ecology and essential information for conservation assessments. Owing to the elusive nature of wild apes and the vast region of dense forest they inhabit, population estimates of central chimpanzees (Pan troglodytes troglodytes) and western lowland gorillas (Gorilla gorilla gorilla) have largely relied on surveys of their nests. Specific information about the nesting behavior of apes permits the estimation of the number of nests built (nest creation rate). Similarly, information on nest characteristics and environmental factors can be used to estimate the time it takes nests to decay (nest decay rate). Nest creation and decay rates are then used to convert nest density estimates to absolute ape densities. Population estimates that use site-specific estimates of nest creation and decay rates are more accurate and precise. However, it is common practice to generalize these conversion factors across sites because of the additional cost of studies required to gather the information to estimate them. Over a 9-mo study period, we detected and monitored the time to decay of gorilla nests (N = 514) and chimpanzee nests (N = 521) in northern Republic of Congo. We investigated the influence of nest characteristics and environmental factors on nest survivorship and estimated the mean time to nest decay (or equivalently survival) using MARK. Key factors influencing nest decay rate included ape species, forest type, nest height, mean rainfall, nest structure, nest type, and primary aspects of nest construction. Our findings highlight the synergistic effect of behavior and environment on great ape nest degradation, as well as providing practical insights for improving measures to monitor remaining populations of these endangered species.

Through the looking glass, and what gibbons found there – New observations on the behaviour of lesser apes when confronted with their own reflection and a novel modification of the mirror-test

Through the looking glass, and what gibbons found there – New observations on the behaviour of lesser apes when confronted with their own reflection and a novel modification of the mirror-test

Toward the Language-Ready Brain: Biological Evolution and Primate Comparisons.

The approach to language evolution suggested here focuses on three questions: How did the human brain evolve so that humans can develop, use, and acquire languages? How can the evolutionary quest be informed by studying brain, behavior, and social interaction in monkeys, apes, and humans? How can computational modeling advance these studies? I hypothesize that the brain is language ready in that the earliest humans had protolanguages but not languages (i.e., communication systems endowed with rich and open-ended lexicons and grammars supporting a compositional semantics), and that it took cultural evolution to yield societies (a cultural constructed niche) in which language-ready brains could become language-using brains. The mirror system hypothesis is a well-developed example of this approach, but I offer it here not as a closed theory but as an evolving framework for the development and analysis of conflicting subhypotheses in the hope of their eventual integration. I also stress that computational modeling helps us understand the evolving role of mirror neurons, not in and of themselves, but only in their interaction with systems "beyond the mirror." Because a theory of evolution needs a clear characterization of what it is that evolved, I also outline ideas for research in neurolinguistics to complement studies of the evolution of the language-ready brain. A clear challenge is to go beyond models of speech comprehension to include sign language and models of production, and to link language to visuomotor interaction with the physical and social world.

Great Apes Do Not Learn Novel Tool Use Easily: Conservatism, Functional Fixedness, or Cultural Influence?

Animal culture has been of interest for decades but the concept remains controversial. Many researchers feel that animal behavior should not be granted the label “culture” because the latter includes more than “simple” behavioral variation. In recent years, the study of animal culture has been concerned mainly with social learning mechanisms, often claimed to be human specific, which led to the uniqueness of human culture. In addition, failure to innovate novel cultural behavior as often as humans is usually explained by psychological mechanisms presented as more parsimonious, such as conservatism or functional fixedness. However, it is unclear how cognitively complex these mechanisms are in the first place. Here, I analyze recent data obtained with wild chimpanzees (Pan troglodytes schweinfurthii) and wild-born orangutans (Pongo abelii) using the honey-trap experiment, in which individuals must devise a solution to reach inaccessible honey. I compare behavior of apes that developed a tool-based solution in the task with those that did not, and test whether conservatism and functional fixedness can explain individual variation in ape behavior. I find evidence of conservatism, with apes relying on their existing knowledge, and evidence of functional fixedness, with this knowledge potentially preventing them from innovating. However, the apes showed large intraspecific variability. I discuss the two mechanisms through a representational perspective to tackle their cognitive complexity. Understanding how conservatism and functional fixedness interact with ape cultural knowledge to limit innovation of novel tool use appears necessary to understand the full extent of ape cultures and how they compare to human cultures.

Effects of crowd size on exhibit use by and behavior of chimpanzees (Pan troglodytes) and Western lowland gorillas (Gorilla gorilla) at a zoo

Several factors are known to affect the overall well-being of captive apes, including facility design and the presence of and interaction with both familiar and unfamiliar humans. Several studies have examined the effects crowds of varying sizes have on the behavior of a wide taxonomic range of zoo-housed animals, including apes. However, previous investigations of visitor effects on ape behavior vary in scope and methodology, leading to inconsistent results across groups and institutions. Using data gathered through two long-term observation protocols, we aimed to determine how 6 chimpanzees (Pan troglodytes) and 5 Western lowland gorillas (Gorilla gorilla) housed at Lincoln Park Zoo responded to visitor crowds of various sizes. The dataset, gathered over 12 consecutive months, included both behavioral data collected via focal animal follows, and exhibit use data collected via whole group scans. As with previous studies, we investigated if and how the behavior of the apes varied as the number of visitors near their exhibits increased across several levels of visitor density. In addition, we looked at whether the apes in this study use the indoor areas of their exhibits that are nearest the public (within one meter) at a rate proportional to the size of that area, and whether their use of this space changed as crowd size varied. As crowd size increased, neither chimpanzees (P=0.97) nor gorillas (P=0.18) were observed to alter their use of the areas of their exhibit closest to zoo visitors. In addition, both chimpanzees and gorillas were observed in this area at a rate equal to or greater than expected by random movements at all three levels of crowd size analyzed. Crowd size had no effect on the frequency of most behaviors analyzed including abnormal, agonism, and grooming (P>0.05). We observed a slight effect of crowd size on self-directed behaviors in both species though it was not the case that the largest crowds caused the highest rates of self-directed behavior. Although these data were collected at a single institution, our findings support the argument that well-designed zoo exhibits can be effective in removing the potentially negative effects of large crowds.