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A two-year longitudinal study of deferred imitation of object manipulation in a juvenile chimpanzee (Pan troglodytes) and orangutan (Pongo pygmaeus).

Changes in deferred imitation of novel actions on objects were assessed over a 2-year period in two enculturated, juvenile great apes (one chimpanzee, Pan troglodytes, and one orangutan, Pongo pygmaeus). Both apes displayed deferred imitation, and both displayed improve ments in deferred imitation over the 2-year period, although the magnitude of improvement was greater for the chimpanzee. This is, to our knowledge, the first experimental demonstration of longitudinal improvements of deferred imitation in great apes. The results were interpreted as reflecting maturationally paced cognitive differences consistent with other cognitive accomplishments in these species, and as demonstrating the influence that a species-atypical rearing environment can have on cognitive abilities in juvenile great apes.

Aging↗

The ontogeny of sexual dimorphism in the cranium of Bornean orang-utans (Pongo pygmaeus pygmaeus): I. Univariate analyses.

Based on a homogeneous sample of 212 individuals spanning all postnatal age periods, we examine the ontogeny of cranial sexual dimorphism in Bornean orang-utans (Pongo pygmaeus pygmaeus) by means of univariate statistics. A distinct pattern emerges at the early juvenile stage and continues at all subsequent stages with males tending to exceed females in all cranial dimensions. In conjunction, starting at mid-juvenile stage, there is a strong tendency for an increase in number and strength of significant cranial sex differences, all of them in favor of males. Significant sex differences in the viscerocranium, reflecting stronger prognathism in males, emerge prior to those in the neurocranium. The total ontogenetic pattern of cranial sexual dimorphism in orang-utans is remarkably similar to that of gorillas, except that there is no evidence of a sex difference in timing of the adolescent growth spurt in the orang-utan. As for other catarrhine species (Wood 1976), male variance of cranial dimensions tends to be greater than that of females, thus lending support to Leutenegger & Cheverud's (1982, 1985) model on the evolution of character dimorphism by means of variance dimorphism.

Analysis of Variance↗

The ontogeny of sexual dimorphism in the cranium of Bornean orang-utans (Pongo pygmaeus pygmaeus): II. Allometry and heterochrony.

Based on a homogeneous sample of 212 individuals spanning all postnatal periods, we examine the ontogeny of cranial sexual dimorphism in Bornean orang-utans (Pongo pygmaeus pygmaeus) by means of allometric analysis and in terms of heterochrony. The bivariate growth allometries of 20 cranial dimensions against basicranial length yield two major patterns. Confirming the null hypothesis, strong ontogenetic scaling, where growth regressions of both sexes fall along a single ontogenetic continuum, and where shape differences between adult males and females result from the extension of relative growth in the smaller females to larger size in males, is found in 10 cases. Ontogenetic scaling is particularly strong in proportions of (1) the neurocranium directly associated with brain size, (2) the orbital region, and (3) the dental arcade. In terms of heterochrony such a pattern most likely is the result of a process termed "time hypermorphosis", i.e. an extension of the growth period in time in males. The second major pattern seen in the remaining 10 cases shows a departure from ontogenetic scaling, with males exhibiting a significantly steeper slope than females. Departures from ontogenetic scaling, where size and shape are dissociated with adult males being disproportionately larger than adult females, are found in proportions of cranial regions directly associated with secondary sexual character development: prognathism, canine size, and cheek pad area. In terms of heterochrony such a pattern most likely is the result of a process termed "acceleration", i.e. the rate of shape change is increased in males.

Animals↗

Tool-use and tool-making by captive, group-living orangutans (Pongo pygmaeus abelii) at an artificial termite mound.

The present study examined the use and making of tools to obtain foodstuffs in artificial-mound holes by five captive, group-living Sumatran orangutans (Pongo pygmaeus abelii). Three adult orangutans frequently stripped leaves and twigs from a branch provided (tool-making), and then inserted the tool into a hole to obtain foodstuffs (tool-using). A 5-year-old female juvenile usually used the tools that adult orangutans had previously used, but rarely made tools herself. A 2-year-old male infant did not use any tools. The adult orangutans tend to leave one to several leaves at the top of the branch than to leave many leaves on the branch or to strip all leaves. It seemed likely that tools with appropriate leaves are easier to insert into holes and obtain more foodstuffs, compared with branches with many leaves or sticks without any leaves. When the orangutans were unable to insert a tool into a hole, they usually modified the tool and/or changed their tool-using technique, such as changing how they grasped the tool. These findings are discussed from the perspectives of the orangutan's behavioral flexibility regarding tool-use skills and hierarchical organization in food-processing techniques.

Animals↗

Maintenance of self-imposed delay of gratification by four chimpanzees (Pan troglodytes) and an orangutan (Pongo pygmaeus).

Delay maintenance, which is the continuance over time of the choice to forgo an immediate, less preferred reward for a future, more preferred reward, was examined in 4 chimpanzees (Pan troglodytes) and 1 orangutan (Pongo pygmaeus). In the 1st experiment, the apes were presented with 20 chocolate pieces that were placed, one at a time, into a bowl that was within their reach. The apes could consume the available chocolate pieces at any time during a trial, but no additional pieces would be given. The total length of time taken to place the 20 items into the bowl ranged from 60 s to 180 s. All 5 apes delayed gratification on a majority of trials until all 20 chocolate pieces were presented. Unlike in most experiments with human children using this test situation, attention by the apes to the reward was not detrimental to delay maintenance. In a 2nd experiment with the chimpanzees only, 4 foods of differing incentive value were presented in different trials in the same manner as in Experiment 1. The chimpanzees were highly successful in obtaining all food pieces, and there was no difference in performance as a function of food type.

Animals↗

Airsacculitis in fourteen juvenile southern Bornean orangutans (Pongo pygmaeus wurmbii).

BACKGROUND: Airsacculitis is a clinical condition which has been reported in a range of primates species, including orangutans. METHODS: This report describes the occurence and management of airsacculitis in fourteen juvenile Southern Bornean orangutans (Pongo pygmaeus wurmbii) that presented beween January 1st 1999 and January 31st 2001 at the Orangutan Care Center and Quarantine (OCC&Q), Kalimantan Tengah, Indonesia (S 2 degrees 43' 49.2"; E 111 degrees 38' 54.2"). Details of the signalment, clinical history, presenting clinical signs, clinicopathological findings and bacterial isolates in each case were reviewed. RESULTS: Cough, halitosis and nasal discharge were the most frequently observed clinical signs. A range of Gram-negative bacteria were isolated from infected air sacs, including Pseudomonas sp., Enterobacter sp. and Klebsiella pneumoniae. A simple drainage and lavage technique was used in cases where surgical intervention was indicated, in combination with local and systemic antibiotic therapy. CONCLUSIONS: The importance of early diagnosis, prompt management and antibiotic selection, based on bacterial culture and sensitivity profiles, is outlined.

Air Sacs↗

Referential understanding of videos in chimpanzees (Pan troglodytes), orangutans (Pongo pygmaeus), and children (Homo sapiens).

Performance on identical search tasks based on cues directly perceived or indirectly perceived through video were compared among a group of 4 adult chimpanzees (Pan troglodytes), a group of 2 adult orangutans (Pongo pygmaeus), and a group of 36 children (between 2 and 3 years of age). Children comprehended directly perceived cues but had difficulty with video cues. In contrast, chimpanzees and 1 orangutan were successful in using video to guide their search for a hidden object. Two follow-up studies with 3-year-old children demonstrated the importance of more distinct perceptual and verbal cues in aiding children's understanding of video as referring to real-world events.

Animals↗

Social learning by orangutans (Pongo abelii and Pongo pygmaeus) in a simulated food-processing task.

Increasing evidence for behavioral differences between populations of primates has created a resurgence of interest in examining mechanisms of information transfer between individuals. The authors examined the social transmission of information in 15 captive orangutans (Pongo abelii and Pongo pygmaeus) using a simulated food-processing task. Experimental subjects were shown 1 of 2 methods for removing a suite of defenses on an "artificial fruit." Control subjects were given no prior exposure before interacting with the fruit. Observing a model provided a functional advantage in the task, as significantly more experimental than control subjects opened the fruit. Within the experimental groups, the authors found a trend toward differences in the actual behaviors used to remove 1 of the defenses. Results support observations from the wild implying horizontal transfer of information in orangutans and show that a number of social learning processes are likely to be involved in the transfer of knowledge in this species.

Animal Communication↗

Hand preferences in captive orangutans (Pongo pygmaeus).

The strength of the evidence for population-level handedness in the great apes is a topic of considerable debate, yet there have been few studies of handedness in orangutans. We conducted a study of manual lateralization in a captive group of eight orangutans (Pongo pygmaeus) ranking the degrees of manual preference according to a defined framework. We analyzed five behavioral patterns: eat (one- and two-handed), make/modify tool, oral tool-use, and manual tool-use. Although some individuals showed significant manual preferences for one or more tasks, at the group-level both one-handed and two-handed eating, oral tool-use, and make/modify tool were ranked at level 1 (unlateralized). Manual tool-use was ranked at level 2, with four subjects demonstrating significant hand preferences, but no group-level bias to the right or left. Four subjects also showed hand specialization to the right or left across several tasks. These results are consistent with most previous studies of manual preference in orangutans. The emergence of manual lateralization in orangutans may relate to more complex manipulative tasks. We hypothesize that more challenging manual tasks elicit stronger hand preferences.

Animals↗

Gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) encode relevant problem features in a tool-using task.

Two important elements in problem solving are the abilities to encode relevant task features and to combine multiple actions to achieve the goal. The authors investigated these 2 elements in a task in which gorillas (Gorilla gorilla) and orangutans (Pongo pygmaeus) had to use a tool to retrieve an out-of-reach reward. Subjects were able to select tools of an appropriate length to reach the reward even when the position of the reward and tools were not simultaneously visible. When presented with tools that were too short to retrieve the reward, subjects were more likely to refuse to use them than when tools were the appropriate length. Subjects were proficient at using tools in sequence to retrieve the reward.

Animals↗

Use of limestone karst forests by Bornean orangutans (Pongo pygmaeus morio) in the Sangkulirang peninsula, east Kalimantan, Indonesia.

The Indonesian province of East Kalimantan is home to some of the largest remaining contiguous tracts of lowland Dipterocarp forest on the island of Borneo. Nest surveys recently conducted in these forests indicated the presence of a substantial population of Eastern Bornean orangutans (Pongo pygmaeus morio) in the Berau and East Kutai regencies in the northern half of the province. The Sangkulirang Peninsula contains extensive limestone karst forests in close proximity to the lowland Dipterocarp forests inhabited by orangutans in these regencies. Orangutans have been sighted in these limestone karst forests, but the importance of this forest type for orangutans has been unclear. Therefore, we conducted 49 km of nest surveys in limestone karst forest to obtain the first quantitative estimates of orangutan densities in this habitat, and walked 28 km of surveys in nearby lowland Dipterocarp forests for comparison. We also gathered basic ecological data along our transects in an attempt to identify correlates of orangutan abundance across these habitat types. Undisturbed limestone karst forests showed the lowest orangutan densities (147 nests/km(2), 0.82 indiv/km(2)), disturbed limestone forests had intermediate densities (301 nests/km(2), 1.40 indiv/km(2)), and undisturbed lowland Dipterocarp forests contained the highest density (987 nests/km(2), 5.25 indiv/km(2)), significantly more than the undisturbed limestone karst forests. This difference was not correlated with variation in liana abundance, fig stem density, or stump density (an index of forest disturbance). Therefore, other factors, such as the relatively low tree species diversity of limestone karst forests, may explain why orangutans appear to avoid these areas. We conclude that limestone karst forests are of low relevance for safeguarding the future of orangutans in East Kalimantan.

Animals↗

The Alu I-induced bands in metaphase chromosomes of orangutan (Pongo pygmaeus). Implications for the distribution pattern of highly repetitive DNA sequences.

Restriction endonucleases have been recently proved to be active on fixed chromosomes, thus they are useful in chromatin structure studies. Within this class of enzymes, Alu I is able to detect the presence and localization of highly repetitive DNA sequences in human and in other mammalian and dipteran species. In this paper the pattern obtained on fixed metaphase chromosomes of orangutan (Pongo pygmaeus) by Alu I digestion and Giemsa staining is shown. The results are discussed in the light of the distribution, in this species, of the I-IV human satellite DNAs. It is also suggested that in Pongo some highly repetitive sequences, different from the major human satellites, are present.

Animals↗

How the great apes (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) perform on the reversed contingency task: the effects of food quantity and food visibility.

S. T. Boysen and G. G. Berntson (1995) found that chimpanzees performed poorly on a reversed contingency task in which they had to point to the smaller of 2 food quantities to acquire the larger quantity. The authors compared the performance of 4 great ape species (Pan troglodytes, Pongo pygmaeus, Pan paniscus, and Gorilla gorilla) on the reversed contingency task while manipulating food quantity (0-4 or 1-4) and food visibility (visible pairs or covered pairs). Results showed no systematic species differences but large individual differences. Some individuals of each species were able to solve the reversed contingency task. Both quantity and visibility of the food items had a significant effect on performance. Subjects performed better when the disparity between quantities was smaller and the quantities were not directly visible.

Age Factors↗

Gait parameters in vertical climbing of captive, rehabilitant and wild Sumatran orang-utans (Pongo pygmaeus abelii).

Vertical climbing is central to the locomotor and foraging strategies of the great apes and, indeed, to theories about the evolution of locomotor specialisations of hominoid primates. Nevertheless, its kinematics have yet to be fully evaluated. Here, we present spatio-temporal parameters of 80 climbing sequences containing 560 limb cycles obtained from video recordings of captive, rehabilitant and wild Sumatran orang-utans (Pongo pygmaeus abelii). Gait parameters such as cycle duration, duty factor, laterality of footfalls, relative stride length and normalised speed were analysed to identify the influence of body size (through an intraspecific comparison of age/sex classes), the influence of the environment (through an intraspecific comparison of animals living in different habitats) and the influence of morphology (through an interspecific comparison with the climbing characteristics of other primates) on climbing ability in orang-utans. Results show that there are only few differences between the climbing of wild, rehabilitant and captive individuals. Cycle duration is longer and speed is lower for the wild individuals than for other groups due to the complexity of their environment and lack of familiarity with each substrate, which results in a more cautious gait. Sex/age-related differences in climbing were found to be small, although juveniles generally exhibited a shorter cycle duration and lower duty factor than other groups. The spatio-temporal gait parameters of adult females carrying infants were not found to differ significantly from those for females without an infant, which suggests that climbing kinematics are not affected by the presence of a clinging infant. Extended-elbow vertical climbing is primarily characterised by a higher duty factor than flexed-elbow climbing, indicating that the former is an energetically more demanding form of locomotion. In comparison with other primates, orang-utans exhibit a longer cycle duration, longer strides but lower climbing speed, reflecting a compromise between the demands of a large body mass and extreme joint mobility.

Age Factors↗

Developmental changes in the facial morphology of the Borneo orangutan (Pongo pygmaeus): possible signals in visual communication.

Orangutans display remarkable developmental changes and sexual differences in facial morphology, such as the flanges or cheek-pads that develop only on the face of dominant adult males. These changes suggest that facial morphology is an important factor in visual communication. However, developmental changes in facial morphology have not been examined in detail. We studied developmental changes in the facial morphology of the Borneo orangutan (Pongo pygmaeus) by observing 79 individuals of various ages living in the Sepilok Orangutan Rehabilitation Centre (SORC) in Malaysia and in Japanese zoos. We also analyzed photographs of one captive male that were taken over a period of more than 16 years. There were clear morphological changes that occurred with growth, and we identified previously unreported sexual and developmental differences in facial morphology. Light-colored skin around the eyes and mouth is most prominent in animals younger than 3 years, and rapidly decreases in area through the age of approximately 7 years. At the same time, the scattered, erect hairs on the head (infant hair) become thick, dense hairs lying on the head (adult hair) in both sexes. The results suggest that these features are infant signals, and that adult signals may include darkened face color, adult hair, whiskers, and a beard, which begin to develop after the age of approximately 7 years in both sexes. In females, the eyelids remain white even after 10 years, and turn black at around the age of 20; in males, the eyelids turn black before the age of 10. The whiskers and beards of adults are thicker in males than in females, and are fully developed before the age of 10 in males, while they begin to develop in females only after approximately 20 years. White eyelids and undeveloped whiskers and beards may be visual signals that are indicative of young adult females. Our results also show that the facial morphology of the unflanged male is similar to that of the adult female, although it has also been pointed out that unflanged males resemble younger individuals.

Age Factors↗

Nonenculturated orangutans' (Pongo pygmaeus) use of experimenter-given manual and facial cues in an object-choice task.

Several experiments have been performed, to examine whether nonhuman primates are able to make use of experimenter-given manual and facial (visual) cues to direct their attention to a baited object. Contrary to the performance of prosimians and monkeys, great apes repeatedly have shown task efficiency in experiments such as these. However, many great ape subjects used have been "enculturated" individuals. In the present study, 3 nonenculturated orangutans (Pongo pygmaeus) were tested for their ability to use experimenter-given pointing, gazing, and glancing cues in an object-choice task. All subjects readily made use of the pointing gesture. However, when subjects were left with only gazing or glancing cues, their performance deteriorated markedly, and they were not able to complete the task.

Animals↗

Inverted and satellited Y chromosome in the orangutan (Pongo pygmaeus).

An inverted and satellited Y chromosome of almost acrocentric appearance was detected in seven of 14 male orangutans. In the remaining seven animals a submetacentric Y chromosome without NORs occurred. The high frequency with which the satellited Y chromosomes were associated with acrocentric autosomes and the positive AgNO3-staining of their satellite stalks clearly indicate the active state of the NOR on the Y chromosomes. DNA fingerprinting in two orangutan families showed that the inverted and satellited Y chromosomes in carrier orangutan males do not interfere with normal fertility. Within our sample of male orangutans studied, the inverted and satellited Y chromosome is restricted to Sumatran animals; all Bornean specimens possessed the submetacentric Y chromosome. The question arises whether these two kinds of Y chromosome differ constitutively between the Pongo pygmaeus subpopulations.

Animals↗

Hypothesis for the causes and periodicity of repetitive linear enamel hypoplasia in large, wild African (Pan troglodytes and Gorilla gorilla) and Asian (Pongo pygmaeus) apes.

Repetitive linear enamel hypoplasia (rLEH) is often observed in recent large-bodied apes from Africa and Asia as well as Mid- to Late Miocene sites from Spain to China. The ubiquity and periodicity of rLEH are not understood. Its potential as an ontogenetic marker of developmental stress in threatened species (as well as their ancient relatives) makes rLEH an important if enigmatic problem. We report research designed to show the periodicity of rLEH among West African Pan troglodytes (12 male, 32 female), Gorilla gorilla (10 male, 10 female), and Bornean and Sumatran Pongo pygmaeus (11 male, 9 female, 9 unknown) from collections in Europe. Two methods were employed. In the common chimpanzees and gorillas, the space between adjacent, macroscopically visible LEH grooves on teeth with two or more episodes was expressed as an absolute measure and as a ratio of complete unworn crown height. In the orangutans, the number of perikymata between episode onsets, as well as duration of rLEH, was determined from scanning electron micrographs of casts of incisors and canines. We conclude that stress in the form of LEH commences as early as 2.5 years of age in all taxa and lasts for several years, and even longer in orangutans; the stress is not chronic but episodic; the stressor has a strong tendency to occur in pulses of two occurrences each; and large apes from both land masses exhibit rLEH with an average periodicity of 6 months (or multiples thereof; Sumatran orangutans seem to show only annual stress), but this needs further research. This is supported by evidence of spacing between rLEH as well as perikymata counts. Duration of stress in orangutans averages about 6 weeks. Finally, the semiannual stressor transcends geographic and temporal boundaries, and is attributed to regular moisture cycles associated with the intertropical convergence zone modified by the monsoon. While seasonal cycles can influence both disease and nutritional stress, it is likely the combination of seasonal variation in fruiting cycles with specific stressors (malaria and/or intestinal parasites, especially hookworm) that results in this widespread phenomenon. This seasonal stress is sufficiently common and of long duration (6 weeks on average in orangutans) that we think rLEH may reflect significant stress in recent and, inferentially, fossil apes. Increasing seasonality may have impinged negatively on later Miocene apes, especially if they lacked a clear birth peak or seasonality in their reproductive cycles.

Animals↗