Adult Chimpanzees Research Articles

Social play fosters cooperation in wild adult chimpanzees.

Adult social play is a universal human trait, promoting the tolerance, bonding, cooperation, and collective action that sustain our large and complex societies.1,2 Play serves as a conduit for transmitting positive emotions, thereby stimulating psychological resilience to stressors and facilitating the positive intent and trust3,4,5,6 essential for cooperation emergence. In contrast, non-human adult social play is considered rare, and its role in cooperation remains unknown. We address this gap by studying the play behavior of 57 adult chimpanzees (Pan troglodytes) in Taï National Park, Côte d'Ivoire, where adult social play and collective action regularly occur. We show that adult female and male chimpanzees play more during times of increased mate competition (with males mainly playing with immatures) and with adult partners they had recent disputes with, highlighting the role of play in regulating social tension that can undermine cooperation. Chimpanzees also preferred playing with adult partners with whom they share strong affiliative bonds, aligning with the idea that play is associated with social familiarity and trust. Finally, adult chimpanzees were more likely to play before collectively defending their territory against outsiders and hunting monkeys. Those who played together were subsequently more likely to collaborate, reinforcing the notion that the positive feedback signaled via play can facilitate cooperation.5 Our findings demonstrate the sustained significance of adult social play throughout the chimpanzee lifespan, providing valuable insights into the evolution of adult social play and its societal functions, from diffusing tension to supporting social bonds and collective action.

Sex differences in positional behavior of chimpanzees (Pan troglodytes schweinfurthii) living in the dry and open habitat of Issa Valley, Tanzania.

Many early fossil hominins are associated with savanna-mosaic paleohabitats, and high sexual dimorphism that may reflect differences in positional behavior between sexes. However, reconstructions of hominin behavior and the selective pressures they faced in an open habitat are limited by a lack of studies of extant apes living in contemporary, analogous habitats. Here, we describe adult chimpanzee positional behavior in the savanna-mosaic habitat of the Issa Valley, Tanzania, to test whether Issa chimpanzees show larger sex-differences in positional behavior than their forest-dwelling counterparts. We quantified and compared adult locomotor and postural behavior across sexes (6 females, 7 males) in the riparian forest (closed) and miombo woodland (open) vegetation types at Issa Valley (13,743 focal observations). We then compared our results to published data of chimpanzee communities living in more forested habitats. Issa females and males both spent less time arboreally in open vegetation and showed similar locomotor and postural behavior on the same substrates, notably using a high level of suspensory locomotion when arboreal. Females were, however, more arboreal than males during locomotor behavior, as well as compared with females from other communities. Issa males behaved similarly to males from other communities. Results suggest that open habitats do not elicit less arboreal behaviors in either sex, and may even select for suspensory locomotion to effectively navigate an open canopy. An open habitat may, however, increase sex differences in positional behavior by driving female arboreality. We suggest this is because of higher energetic demands and predator pressures associated with open vegetation, which are likely exaggerated for reproducing females. These results have implications for the interpretation of how sexual dimorphism may influence reconstructions of hominin positional behavior.

Surgical Management of Left-Hand Traumatic Injury in an Adult Female Captive Chimpanzee (Pan troglodytes): A Case Report

Captive chimpanzees are reported to present a higher frequency of injuries compared to those obtained in the wild due to their intraspecific aggression. Wounds are inescapable events in life following physical, chemical, thermal, microbial, or immunological insult to a body tissue, resulting in disruptions, breakage and loss of cellular and anatomical or functional continuity of living tissues or organs. The attention of the Veterinary Teaching Hospital, University of Jos was drawn to an injured adult female chimpanzee at the Jos Zoo. History revealed that the wound was the result of aggression from a counterpart chimpanzee. History reveals also that the wound was managed for 1 month before presentation with no significant improvement. Close examination revealed degloved left index and ring digits with pussy discharges. Assessment of the wound was undertaken after chemical immobilization and the resulting defects were reconstructed following proper aseptic surgical preparation utilizing chlorhexidine gluconate (Purit®) and amputation of the non-viable segments of the affected digits. Postoperatively, daily cleaning and administration of oxytetracycline spray (topical), doxycycline capsules 5 mg/kg ϰ1/52 PO and Dolometa B® ϰ5/7 PO were instituted. This case demonstrated the important role of prompt and thorough clinical evaluation in the adoption of management options, especially in wildlife management.

Socioecological correlates of Chimpanzee (Pan troglodytes) hydration status at Gombe National Park, Tanzania

Abstract Maintaining water balance is imperative for organismal survival. Various socioecological conditions such as sex, reproductive state, and dominance rank can impact water intake and hydration status. Eastern Chimpanzees (Pan troglodytes schweinfurthii) present an interesting biological model to investigate the socioecological correlates of hydration status, given adult sex differences in behavior. Males are more gregarious than females, which may afford them less behavioral flexibility to reduce dehydration risk. However, females invest heavily in their offspring, producing a water heavy milk for up to 5 years, and adult females experience disparities in access to water in foods dependent on dominance rank, which may cause variation in hydration status. In the present study, we investigated correlates of adult chimpanzee hydration status at Gombe National Park, Tanzania, using a measure of urine concentration—urine specific gravity (USG)—highly concentrated urine results in a higher USG, indicating dehydration. We found that both males and females had significantly higher USG in the dry season, but males had lower USG than females during the dry season. Within females, reproductive state did not significantly impact hydration status; however, females in middle lactation tended to have higher USG than nonpregnant, nonlactating females. Additionally, high-ranking females had lower USG than mid- and low-ranking females. While our study showed consistency with USG values that have been demonstrated at other Chimpanzee research sites, we found evidence that this population may be chronically dehydrated by human standards, but more research is needed across populations to determine the health and fitness effects of chronic dehydration in Chimpanzees.

The influence of multiple variables on bipedal context in wild chimpanzees: implications for the evolution of bipedality in hominins

Investigations into the role of selection in the origin of human bipedalism using ape models have relied heavily on behavioral frequency data. However, analysis of video of wild apes has the advantage of capturing the details of the entirety of each rare, brief bipedal bout witnessed, not just the moment detected in observational studies. We used video to explore the behavioral context and effects of several variables on bipedalism across all ages in wild forest-dwelling chimpanzees from Ngogo, Uganda. We found, as in earlier studies, that adult chimpanzees used bipedalism in the context of foraging; however, unlike earlier studies, we found that while foraging was the predominant behavioral context during arboreal bipedalism, terrestrial bipedalism was more varied in contextual composition. We also found that these different behavioral contexts of bipedalism were associated with different variables. Specifically, foraging was associated with arboreality, hand assistance, and adulthood; antagonism was associated with adulthood, locomotion, and males; play was associated with terrestriality and subadulthood; and travel was associated with locomotion and females. Given that several variables influence bipedalism across multiple behavioral contexts in chimpanzees, it is likely that the early evolution of human bipedalism occurred under the influence of numerous factors. This exploratory study thus suggests that more comprehensive models should be used when reconstructing the transition to bipedalism from the Last Common Ancestor of humans and chimpanzees.

Ecological variation in adult social play reveals a hidden cost of motherhood for wild chimpanzees

Though common among humans, social play by adults is an uncommon occurrence in most animals, even between parents and offspring.1,2,3 The most common explanation for why adult play is so rare is that its function and benefits are largely limited to development, so that social play has little value later in life.3,4,5,6 Here, we draw from 10 years of behavioral data collected by the Kibale Chimpanzee Project to consider an alternative hypothesis: that despite its benefits, adult play in non-humans is ecologically constrained by energy shortage or time limitations. We further hypothesized that, since they may be the only available partners for their young offspring, mother chimpanzees pay greater costs of play than other adults. Our analysis of nearly 4,000 adult play bouts revealed that adult chimpanzees played both among themselves and with immature partners. Social play was infrequent when diet quality was low but increased with the proportion of high-quality fruits in the diet. This suggests that adults engage in play facultatively when they have more energy and/or time to do so. However, when diet quality was low and most adult play fell to near zero, play persisted between mothers and offspring. Increased use of play by adult chimpanzees during periods of resource abundance suggests that play retains value as a social currency beyond development but that its costs constrain its use. At the same time, when ecological conditions constrain opportunities for young to play, play by mothers fills a critical role to promote healthy offspring development.

Sign Language Studies with Chimpanzees in Sanctuary.

Adult chimpanzees Tatu and Loulis lived at the Fauna Foundation sanctuary. They had acquired signs of American Sign Language (ASL) while young and continued to use them as adults. Caregivers with proficiency in ASL maintained daily sign language records during interactions and passive observation. Sign checklists were records of daily vocabulary use. Sign logs were records of signed interactions with caregivers and other chimpanzees. This study reports sign use from eight years of these records. Tatu and Loulis used a majority of their base vocabularies consistently over the study period. They used signs that they had acquired decades earlier and new signs. Their utterances served a variety of communicative functions, including responses, conversational devices, requests, and descriptions. They signed to caregivers, other chimpanzees, including those who did not use signs, and to themselves privately. This indicates the importance of a stimulating and interactive environment to understand the scope of ape communication and, in particular, their use of sign language.

Comparative transcriptomics reveals human-specific cortical features.

The cognitive abilities of humans are distinctive among primates, but their molecular and cellular substrates are poorly understood. We used comparative single-nucleus transcriptomics to analyze samples of the middle temporal gyrus (MTG) from adult humans, chimpanzees, gorillas, rhesus macaques, and common marmosets to understand human-specific features of the neocortex. Human, chimpanzee, and gorilla MTG showed highly similar cell-type composition and laminar organization as well as a large shift in proportions of deep-layer intratelencephalic-projecting neurons compared with macaque and marmoset MTG. Microglia, astrocytes, and oligodendrocytes had more-divergent expression across species compared with neurons or oligodendrocyte precursor cells, and neuronal expression diverged more rapidly on the human lineage. Only a few hundred genes showed human-specific patterning, suggesting that relatively few cellular and molecular changes distinctively define adult human cortical structure.

PROSPECTIVE POSTMORTEM ASSESSMENT OF THE INCIDENCE OF MAMMARY NEOPLASIA IN CHIMPANZEES (PAN TROGLODYTES).

When evaluating the last 25 yr of morbidity and mortality from adult chimpanzees managed within the Association of Zoo and Aquarium Chimpanzee Species Survival Plan® for North American zoos, only two female chimpanzees were diagnosed with mammary neoplasia: one incidentally antemortem and one with a terminal metastatic neoplasia. When comparing this observation of prevalence of mammary neoplasia to humans, a substantial disparity is apparent. Mammary neoplasia is the second most common cancer in adult female humans, with a lifetime risk of 1:8 in the United States. The reason for the disparity between humans and chimpanzees, as closely related species, is unknown. The true prevalence in chimpanzees may be higher than currently noted, because routine examination of mammary tissue in chimpanzees is generally less complete than for other tissues postmortem, and antemortem assessment is generally limited to mammary palpation. This study was performed on intact, bilateral mammary glands harvested at postmortem examination of adult female chimpanzees (n = 7) from six institutions. With mammography, complete histopathologic sectioning, and genetic evaluation, the risk of mammary neoplasia was evaluated more thoroughly than during a typical postmortem exam in zoo populations during 2017-2019. No chimpanzees in the study were diagnosed with mammary neoplasia. Overall, this study supports the previous impression that chimpanzees do not develop mammary neoplasia at a similar rate as humans, even when comparable diagnostic modalities for evaluation are used.

Comparative Histoanatomy of the Epiglottis and Pre-epiglottic Space of the Chimpanzee Larynx

Understanding of the evolution of the larynx clarifies the physiology and functional histoanatomy of the present-day human organ. Comparative histoanatomy of the epiglottis and pre-epiglottic space of the chimpanzee larynx (a hominid, phyletic closest relative of humans) was investigated. Two normal adult chimpanzee larynges obtained from natural deaths were investigated. The whole organ serial section technique was employed. The histoanatomical structures of the chimpanzees' epiglottis and pre-epiglottic space were considerably similar to those of human adults. The chimpanzees' epiglottic cartilage was relatively thin and composed of elastic cartilage. These histologic findings of epiglottis indicate that the chimpanzee's epiglottis is flexible and plays the role of retroflection. The chimpanzees' larynges had a pre-epiglottic space composed of adipose tissue and loose connective tissue. Epiglottic cartilage was connected to the thyroid cartilage anteroinferiorly with an intervening thyroepiglottic ligament and to the hyoid bone anteriorly with an intervening hyoepiglottic ligament. These histoanatomical structures of the epiglottis and pre-epiglottic space reflect the fact that chimpanzees have a descended larynx and acquire the pharyngeal space of the vocal tract. The results of this study are consistent with the hypothesis that, in the process of evolution, the histoanatomical structures of the epiglottis and pre-epiglottic space change and allow the larynx to descend and lengthen the pharyngeal space of the vocal tract which facilitates speech production in humans. Moreover, the distribution of the human pre-epiglottic space likely allows the epiglottis to more effectively play the role of retroflection during swallowing in order to prevent aspiration.

Is it for real? Structural differences between play and real fighting in adult chimpanzees (Pan troglodytes).

In primates, as well as in other mammals, play fighting (PF) is a complex form of playful activity that is structurally similar to real fighting (RF) and may also be used in a competitive way. Here, we verify the structural key differences that can distinguish PF from RF in adult chimpanzees (Pan troglodytes). We collected 962 h of video recording on 30 adult individuals belonging to four chimpanzee groups (Mona Chimpanzee Sanctuary, Spain; La Vallée des Singes and ZooParc de Beauval, France). We applied different indices-two of which were borrowed from the ecological measures of biodiversity-to test for structural differences between PF (345 sessions) and RF (461 sessions) in the levels of behavior repetition (Repeatability of Same Behavior Index, RSBI), distribution uniformity (Pielou Index, J), variability (Shannon Index, H') and, symmetry (i.e., reciprocal exchange of offensive/defensive behaviors; Asymmetry Index, AI). Moreover, we compared the session duration between PF and RF. We found that duration and RSBI were higher in PF than RF while AI was higher in RF than PF. No difference was found between J and H'. Interestingly, both females and males maintained similar ranking positions (determined via Normalized David's scores) in RF and PF. Our study indicates that session duration, behavior repetition, and symmetry can be distinctive structural key features of PF whereas dominance role-reversal, behavior variability, and distribution uniformity were not. PF in adult chimpanzees may have elements of serious contexts (e.g., absence of role-reversal as in RF) which is in line with the view that play is a blended, multifunctional behavior deriving from the re-combination of different behavioral systems. Our findings highlight the need to investigate play structure and manifestation in a nuanced way to better understand the actual motivation that underlies what appears to be play.

Lean muscle mass, not aggression, mediates a link between dominance rank and testosterone in wild male chimpanzees

Testosterone promotes mating effort, which involves intraspecific aggression for males of many species. Therefore, males with higher testosterone levels are often thought to be more aggressive. For mammals living in multimale groups, aggression is hypothesized to link male social status (i.e. dominance rank) and testosterone levels, given that high status predicts mating success and is acquired partly through aggressive intragroup competition. In male chimpanzees, Pan troglodytes, dominance rank has been repeatedly linked to interindividual variation in testosterone levels, but evidence directly linking interindividual variation in testosterone and aggression is lacking. In the present study, we test both aggression levels and lean muscle mass, as measured by urinary creatinine, as links between dominance rank and testosterone levels in a large sample of wild male chimpanzees. Multivariate analyses indicated that dominance rank was positively associated with total rates of intragroup aggression, average urinary testosterone levels and average urinary creatinine levels. Testosterone was positively associated with creatinine levels but negatively associated with total aggression rates. Furthermore, mediation analyses showed that testosterone levels facilitated an association between dominance rank and creatinine levels. Our results indicate that (1) adult male chimpanzees with higher average testosterone levels are often higher ranking but not more aggressive than males with lower testosterone and (2) lean muscle mass links dominance rank and testosterone levels in Ngogo males. We assert that aggression rates are insufficient to explain links between dominance rank and testosterone levels in male chimpanzees and that other social variables (e.g. male-male relationship quality) may regulate testosterone's links to aggression.

The less-is-better effect: a developmental perspective.

The less-is-better effect emerges when an option of lesser quantitative value is preferred or overvalued relative to a quantitively greater alternative (e.g., 24-piece dinnerware set > 24-piece dinnerware set with 16 additional broken dishes; Hsee, 1998, Journal of Behavioral Decision Making, 11, 107-121). This decisional bias emerges when the option of lesser quantitative value is perceived as qualitatively better (e.g., smaller set of intact dishes > larger set of partially broken dishes). Interestingly, this effect emerges for adult humans when options are evaluated separately but dissipates when options are considered simultaneously. The less-is-better bias has been attributed to the evaluability hypothesis: individuals judge objects on the basis of easy-to-evaluate attributes when judged in isolation, such as the brokenness of items within a set, yet shift to quantitative information when evaluated jointly, such as the overall number of dishes. This bias emerges for adult humans and chimpanzees in a variety of experimental settings but has not yet been evaluated among children. In the current study, we presented a joint evaluation task (larger yet qualitatively inferior option vs. smaller yet qualitatively superior option) to children aged 3 to 9 years old to better understand the developmental trajectory of the less-is-better effect. Children demonstrated the bias across all choice trials, preferring an objectively smaller set relative to a larger yet qualitatively poorer alternative. These developmental findings suggest that young children rely upon salient features of a set to guide decision-making under joint evaluation versus more objective attributes such as quantity/value.

Comparative ontogeny of functional aspects of human cervical vertebrae.

Differences between adult humans and great apes in cervical vertebral morphology are well documented, but the ontogeny of this variation is still largely unexplored. This study examines patterns of growth in functionally relevant features of C1, C2, C4, and C6 in extant humans and apes to understand the development of their disparate morphologies. Linear and angular measurements were taken from 530 cervical vertebrae representing 146 individual humans, chimpanzees, gorillas, and orangutans. Specimens were divided into three age-categories based on dental eruption: juvenile, adolescent, and adult. Inter- and intraspecific comparisons were evaluated using resampling methods. Of the eighteen variables examined here, seven distinguish humans from apes at the adult stage. Human-ape differences in features related to atlantoaxial joint function tend to be established by the juvenile stage, whereas differences in features related to the nuchal musculature and movement of the subaxial elements do not fully emerge until adolescence or later. The orientation of the odontoid process-often cited as a feature that distinguishes humans from apes-is similar in adult humans and adult chimpanzees, but the developmental patterns are distinct, with human adultlike morphology being achieved much earlier. The biomechanical consequences of the variation observed here is poorly understood. Whether the differences in growth patterns represent functional links to cranial development or postural changes, or both, requires additional investigation. Determining when humanlike ontogenetic patterns evolved in hominins may provide insight into the functional basis driving the morphological divergence between extant humans and apes.

Similarity and structured representation in human and nonhuman apes

How we judge the similarity between objects in the world is connected ultimately to how we represent those objects. It has been argued extensively that object representations in humans are ‘structured’ in nature, meaning that both individual features and the relations between them can influence similarity. In contrast, popular models within comparative psychology assume that nonhuman species appreciate only surface-level, featural similarities. By applying psychological models of structural and featural similarity (from conjunctive feature models to Tversky's Contrast Model) to visual similarity judgements from adult humans, chimpanzees, and gorillas, we demonstrate a cross-species sensitivity to complex structural information, particularly for stimuli that combine colour and shape. These results shed new light on the representational complexity of nonhuman apes, and the fundamental limits of featural coding in explaining object representation and similarity, which emerge strikingly across both human and nonhuman species.

Integrating Reference Intervals into Chimpanzee Welfare Research

Animal welfare researchers are committed to developing novel approaches to enhance the quality of life of chimpanzees living in professional care. To systematically monitor physical, mental, and emotional states, welfare scientists highlight the importance of integrating non-invasive, animal-based welfare indicators. This study aimed to create species-specific reference intervals for behavioral measures and physiological biomarkers. Specifically, we analyzed data from 40 adult chimpanzees (22 females, 18 males) residing at 16 zoological facilities to generate reference intervals for behavioral states and events, behavioral diversity, fecal glucocorticoid metabolites (GCMs), and fecal immunoglobulin-A (IgA). Comparisons of sex and age using linear regression models revealed significant differences for several behaviors. The proportion of time spent engaged in mutual/multiple social grooming significantly decreased as individuals aged. Furthermore, males spent a higher proportion of time performing aggressive contact behaviors and displaying to other chimpanzees when compared to females. Males also performed sexual examination behaviors at a higher rate than females. Behavioral diversity, fecal GCM, and fecal IgA did not vary by sex or age. In the future, values for individual chimpanzees can be compared to the ranges reported here for particular age/sex classes. Ultimately, animal care professionals can utilize reference intervals to make evidence-based decisions regarding management practices and environmental conditions.

Innovative multi-material tool use in the pant-hoot display of a chimpanzee

‘Pant-hoot displays’ are a species-typical, multi-modal communicative behaviour in chimpanzees in which pant-hoot vocalisations are combined with varied behavioural displays. In both captivity and the wild, individuals commonly incorporate striking or throwing elements of their environment into these displays. In this case study, we present five videos of an unenculturated, captive, adult male chimpanzee combining a large rubber feeding tub with excelsior (wood wool) in a multi-step process, which was then integrated into the subject’s pant-hoot displays as a percussive tool or ‘instrument’. During the construction process, the subject demonstrated an understanding of the relevant properties of these materials, ‘repairing’ the tub to be a more functional drum when necessary. We supplement these videos with a survey of care staff from the study site for additional detail and context. Although care must be taken in generalising data from a single individual, the behaviour reported here hints at three intriguing features of chimpanzee communicative cognition: (1) it suggests a degree of voluntary control over vocal production, (2) it is a so-far unique example of compound tool innovation and use in communicative behaviour and (3) it may represent an example of forward planning in communicative behaviour. Each of these would represent hitherto undocumented dimensions of flexibility in chimpanzee communication, mapping fertile ground for future research.

Age, adrenal steroids, and cognitive functioning in captive chimpanzees (Pan troglodytes).

Dehydroepiandrosterone-sulfate is the most abundant circulating androgen in humans and other catarrhines. It is involved in several biological functions, such as testosterone production, glucocorticoid antagonist actions, neurogenesis and neuroplasticty. Although the role of dehydroepiandrosterone-sulfate (DHEAS) in cognition remains elusive, the DHEAS/cortisol ratio has been positively associated with a slower cognitive age-decline and improved mood in humans. Whether this relationship is found in nonhuman primates remains unknown. We measured DHEAS and cortisol levels in serum of 107 adult chimpanzees to investigate the relationship between DHEAS levels and age. A subset of 21 chimpanzees was used to test the potential associations between DHEAS, cortisol, and DHEAS/cortisol ratio in cognitive function, taking into account age, sex, and their interactions. We tested for cognitive function using the primate cognitive test battery (PCTB) and principal component analyses to categorize cognition into three components: spatial relationship tasks, tool use and social communication tasks, and auditory-visual sensory perception tasks. DHEAS levels, but not the DHEAS/cortisol ratio, declined with age in chimpanzees. Our analyses for spatial relationships tasks revealed a significant, positive correlation with the DHEAS/cortisol ratio. Tool use and social communication had a negative relationship with age. Our data show that the DHEAS/cortisol ratio, but not DHEAS individually, is a promising predictor of spatial cognition in chimpanzees.

Molecular and cellular evolution of the primate dorsolateral prefrontal cortex.

The granular dorsolateral prefrontal cortex (dlPFC) is an evolutionary specialization of primates that is centrally involved in cognition. We assessed more than 600,000 single-nucleus transcriptomes from adult human, chimpanzee, macaque, and marmoset dlPFC. Although most cell subtypes defined transcriptomically are conserved, we detected several that exist only in a subset of species as well as substantial species-specific molecular differences across homologous neuronal, glial, and non-neural subtypes. The latter are exemplified by human-specific switching between expression of the neuropeptide somatostatin and tyrosine hydroxylase, the rate-limiting enzyme in dopamine production in certain interneurons. The above molecular differences are also illustrated by expression of the neuropsychiatric risk gene FOXP2, which is human-specific in microglia and primate-specific in layer 4 granular neurons. We generated a comprehensive survey of the dlPFC cellular repertoire and its shared and divergent features in anthropoid primates.

Hierarchical object combination and tool use in the great apes and human children

Object manipulation can be used as a comparative scale of cognitive development among primates, including humans. Combinatory object manipulation is a precursor of tool-using behavior that indicates material intelligence in primates. However, developmental data on it regarding the great apes other than chimpanzees is insufficient. We conducted a longitudinal investigation of humans and chimpanzees as well as a cross-sectional examination of other great-ape infants (two bonobos, three gorillas, and four orangutans) in captive settings by using two kinds of tasks that required either inserting or stacking combinatory action. The four species of great apes and humans demonstrated both types of combinatory object manipulation during infancy. However, the order of development in different types of combinatory object manipulations varied among the great apes. Furthermore, we applied a nesting-cup task to examine the hierarchical complexity in the combinatory strategies of human children and adult chimpanzees. Both of them exhibited highly hierarchical combinations in the nesting-cup task and employed the subassembly strategy, indicating that an action merge may exist not only in human children but also in adult chimpanzees. The results were discussed with reviews of the tool-use literature from the wild great apes. The early acquisition of an inserting action in the chimpanzees may explain the tool utilization commonality reported in wild chimpanzees. The combinatory object manipulation may have worked as an external enhancer to achieve an additional hierarchical complexity in cognition and behavior, eventually leading humans to develop a language system.