When presented with the option of either an immediate benefit or a larger, later reward, we may behave impatiently by choosing instant gratification. Nonetheless, when we can make the same decision ahead of time and plan for the future, we tend to make more patient choices. Here, we explored whether great apes share this core feature of human decision-making, often referred to as dynamic inconsistency. We found that orangutans, bonobos, and gorillas tended to act impatiently and with considerable variability between individuals when choosing between an immediate reward and a larger-later reward, which is a commonly employed testing method in the field. However, with the inclusion of a front-end delay for both alternatives, their decisions became more patient and homogeneous. These results show that great apes are dynamically inconsistent. They also suggest that, when choosing between future outcomes, they are more patient than previously reported. We advocate for the inclusion of diverse time ranges in comparative research, especially considering the intertwinement of intertemporal choices and future-oriented behavior.

This article applies object detection to CCTV video material to investigate the potential of using machine learning to automate behavior tracking. This study includes video tapings of two captive Bornean orangutans and their behavior. From a 2 min training video containing the selected behaviors, 334 images were extracted and labeled using Rectlabel. The labeled training material was used to construct an object detection model using Create ML. The use of object detection was shown to have potential for automating tracking, especially of locomotion, whilst filtering out false positives. Potential improvements regarding this tool are addressed, and future implementation should take these into consideration. These improvements include using adequately diverse training material and limiting iterations to avoid overfitting the model.

The rarity of Pongo fossils with precise absolute dating from the Middle Pleistocene hampers our understanding of the taxonomy and spatiotemporal distribution of Quaternary orangutans in southern China. Here, we report a newly discovered sample of 113 isolated teeth of fossil Pongo from Zhongshan Cave in the Bubing Basin, Guangxi, southern China. We describe the Pongo specimens from Zhongshan Cave and compare them metrically to other samples of fossil Pongo species (i.e., Pongo weidenreichi, Pongo devosi, Pongo duboisi, Pongo palaeosumatrensis, Pongo javensis, and Pongo sp.) and to extant orangutans (i.e., Pongo pygmaeus and Pongo abelii). The Zhongshan Pongo assemblage is dated using U-series and coupled electron spin resonance/U-series methods. Our results reasonably constrain the Zhongshan Pongo assemblage to 184 ± 16 ka, which is consistent with the biostratigraphic evidence. The Zhongshan Pongo teeth are only 6.5% larger on average than those of extant Pongo. The Zhongshan teeth are smaller overall than those of Pongo from all other cave sites in southern China, and they currently represent the smallest fossil orangutans in southern China. Based on their dental size, and the presence of a well-developed lingual pillar and lingual cingulum on the upper and lower incisors, an intermediate frequency of lingual cingulum remnants on the upper molars, and a higher frequency of moderate to heavy wrinkling on the upper and lower molars, we provisionally assign the Zhongshan fossils to P. devosi. Our results confirm earlier claims that P. weidenreichi is replaced by a smaller species in southern China, P. devosi, by the late Middle Pleistocene. The occurrence of P. devosi in Zhongshan Cave further extends its spatial and temporal distribution. The Pongo specimens from Zhongshan provide important new evidence to demonstrate that the dental morphological features of Pongo in southern China changed substantially during the late Middle Pleistocene.

Recursive procedures that allow placing a vocal signal inside another of a similar kind provide a neuro-computational blueprint for syntax and phonology in spoken language and human song. There are, however, no known vocal sequences among nonhuman primates arranged in self-embedded patterns that evince vocal recursion or potential incipient or evolutionary transitional forms thereof, suggesting a neuro-cognitive transformation exclusive to humans. Here, we uncover that wild flanged male orangutan long calls feature rhythmically isochronous call sequences nested within isochronous call sequences, consistent with two hierarchical strata. Remarkably, three temporally and acoustically distinct call rhythms in the lower stratum were not related to the overarching rhythm at the higher stratum by any low multiples, which suggests that these recursive structures were neither the result of parallel non-hierarchical procedures nor anatomical artifacts of bodily constraints or resonances. Findings represent a case of temporally recursive hominid vocal combinatorics in the absence of syntax, semantics, phonology, or music. Second-order combinatorics, \'sequences within sequences\', involving hierarchically organized and cyclically structured vocal sounds in ancient hominids may have preluded the evolution of recursion in modern language-able humans.
Language is the most powerful communication tool known in nature. By combining a finite set of elements, it allows us to encode infinite messages. This enables communication about virtually anything, from alerting others to potential dangers, to recommending a favourite book. The prevailing theory of the last 70 years suggests that this ability rests on a computational process in the brain that is unique to humans, known as recursion. Recursion enables humans to produce and place a language element or pattern of elements inside another element or pattern of the same kind. In this way, a clause can be embedded inside another ‘carrier’ clause to extend a thought, argument, or scenario, for example, “the dog, which chased the cat, was barking”. While recursion offers a simple, yet potent, explanation for the endless possibilities of language, how and why recursion – and by extension language – emerged in humans but no other animals remains a mystery. Lameira et al. observed vocal patterns in wild orangutans that appeared to be composed of different elements. As orangutans and other great apes are our closest living relatives, they represent the most realistic model for studying the ability of human ancestors to use and comprehend language. Therefore, Lameira et al. set out to determine if this was a case of vocal patterning embedded within a similar vocal pattern, which could indicate that recursion underpins production of these calls. Analysing recordings of long calls made by wild male orangutans showed that they are organized as two layers, where calls with a regular beat (or tempo) are produced within another “carrier” call of a different tempo. Up to three different call types, each with their own signature tempo, can occur within the same carrier call. Further analysis confirmed these call types were unrelated to the carrier. The findings of Lameira et al. demonstrate that orangutans produce recursive vocal sequences that could represent a possible precursor to recursion in humans, offering a potential avenue for studying how recursion, and ultimately language, evolved in humans. In the future, better understanding of how language evolved may help to refine machine learning algorithms that aim to recognize, predict or generate text.

In human infants, exploratory object manipulation is a major vehicle for cognitive stimulation as well as an important way to learn about objects and basic physical concepts in general. The development of human infants\' exploratory object manipulation follows distinct developmental patterns. So far, the degree of evolutionary continuity of this developmental process remains unclear. We investigated the development of exploratory object manipulations in wild orangutans. Our data included 3200 exploration events collected on 13 immatures between the ages of 0.5 and 13 years, at the Suaq Balimbing monitoring station in Indonesia. Our results identify several parallels between the development of exploratory behaviour in humans and orangutans: on top of a highly similar overall age trajectory, we found an increase in variability of the actions used, an increase in the number of body parts involved in each event, and an overall decrease of mouthing of the objects. All in all, our results show that orangutans progress through a developmental sequence of different aspects of exploration behaviour. In combination with previous findings from captivity, our results also provide evidence that exploratory object manipulations reflect cognitive development and might function as a means of cognitive stimulation not just in humans but across the great apes.

For humans we implicitly assume that the way we do things is the product of social learning and thus cultural. For animals, this conclusion requires proof. Here, we first review the most commonly used procedure for documenting animal culture: the method of exclusion, which charts geographic behavioral variation between populations as evidence for culture. Using published data, we show that, whereas it is an adequate proof of principle, the method of exclusion has major deficiencies when capturing cultural diversity and complexity. Therefore, we propose a new method, namely the direct counting of socially learned skills, which we apply to previously collected data on wild orangutans. This method reveals a far greater cultural repertoire among orangutans, and a different distribution of cultural elements among behavioral domains than found by the method of exclusion, as well as clear ecological correlates for most cultural elements. The widespread occurrence of social learning ability throughout the animal kingdom suggests that these conclusions also apply to many other species. Culture is most likely more widespread and pervasive than commonly thought and an important avenue to local adaptation. The complex and normative dimensions of culture seem unique to our species, but were most likely built upon a very broad, pre-existing cultural capacity that we inherited from our ancestors.

The shape of the trapezium and first metacarpal (Mc1) markedly influence thumb mobility, strength, and the manual abilities of extant hominids. Previous research has typically focused solely on trapezium-Mc1 joint shape. Here we investigate how morphological integration and shape covariation between the entire trapezium (articular and non-articular surfaces) and the entire Mc1 reflect known differences in thumb use in extant hominids.
We analyzed shape covariation in associated trapezia and Mc1s across a large, diverse sample of Homo sapiens (n = 40 individuals) and other extant hominids (Pan troglodytes, n = 16; Pan paniscus, n = 13; Gorilla gorilla gorilla, n = 27; Gorilla beringei, n = 6; Pongo pygmaeus, n = 14; Pongo abelii, n = 9) using a 3D geometric morphometric approach. We tested for interspecific significant differences in degree of morphological integration and patterns of shape covariation between the entire trapezium and Mc1, as well as within the trapezium-Mc1 joint specifically.
Significant morphological integration was only found in the trapezium-Mc1 joint of H. sapiens and G. g. gorilla. Each genus showed a specific pattern of shape covariation between the entire trapezium and Mc1 that was consistent with different intercarpal and carpometacarpal joint postures.
Our results are consistent with known differences in habitual thumb use, including a more abducted thumb during forceful precision grips in H. sapiens and a more adducted thumb in other hominids used for diverse grips. These results will help to infer thumb use in fossil hominins.

Considerable research has been conducted on the effects of inter-institutional transfers, but far less consideration has been given to intra-institutional transfers and extended housing in off-habitat holding. On 15 May 2018, The Oregon Zoo\'s orangutans (n = 3) were moved from the Red Ape Reserve (RAR) to the Veterinary Medical Center (VMC) indoor holding areas and remained there until 22 December 2020, resulting in over two years of housing in a facility not specifically designed for orangutans. This study aimed to quantify potential changes in fecal glucocorticoid metabolites (fGM) typically associated with increased adrenal activity as a result of transfers, as well as potential differences in fGM concentrations associated with housing in the two different types of locations. We collected fecal samples from all orangutans during three time periods: the initial housing at RAR (RAR1), the time spent at VMC holding (VMC), and the return to RAR (RAR2). Samples were analyzed using enzyme-immunoassay (EIA) analyses and compared using two-way ANOVA tests with Games-Howell post-hoc evaluations. The results of our analyses showed the following: (1) significant differences in fGM concentrations based on location in two orangutans, with the highest fGM concentration occurring in fecal samples collected at the VMC; and (2) a lack of significant fGM peaks following multiple intra-institutional transfers for all three orangutans. Though requiring further corroboration through future studies, we speculated that pre-transfer behavior training and intensive, continued care by familiar animal care staff may have helped to mitigate the stress responses commonly associated with transfers and major changes in housing. Furthermore, this study highlights the individualistic nature of the stress response, as illustrated by the substantial variation in fGM concentrations across different housing regimens in the three orangutans.

Pongo fossils with precise absolute age brackets are rare, limiting our understanding of their taxonomy and spatiotemporal distribution in southern China during the Late Pleistocene. Twenty-four isolated teeth of fossil orangutans were recently discovered during excavations at Yicun Cave in Guangxi Zhuang Autonomous Region, southern China. Here, we dated the fossil-bearing layer using Uranium-series dating of the associated flowstone and soda straw stalactites. Our results date the Yicun orangutan fossils to between 66 ± 0.32 ka and 57 ± 0.26 ka; thus, these fossils currently represent the last appearance datum of Pongo in southern China. We further conducted a detailed morphological comparison of the Yicun fossil teeth with large samples of fossil (n = 2454) and extant (n = 441) orangutans from mainland and island Southeast Asia to determine their taxonomic position. Compared to other fossil and extant orangutan samples, the Yicun Pongo assemblage has larger teeth and displays greater variation in occlusal structure. Based on the high frequency of cingular remnants and light to moderate enamel wrinkling of the molars, we assigned the Yicun fossils to Pongo weidenreichi, a species that was widespread in southern China throughout the Pleistocene. Lastly, we used published stable carbon isotope data of Early to Late Pleistocene mammalian fossil teeth from mainland Southeast Asia to reconstruct changes in the paleoenvironment and to interpret dental size variation of Pongo assemblages in a broader temporal and environmental context. The carbon isotope data show that dental size reduction in Pongo is associated with environmental changes. These morphological changes in Pongo appear to coincide with the expansion of savannah biomes and the contraction of forest habitats from the Middle Pleistocene onward. The variation in dental size of forest-dwelling Pongo in mainland Southeast Asia may have resulted from habitat differentiation during the Pleistocene.

Gestures play an essential role in primate communication. However, little is known about how complexity of gestural use (in terms of repertoire size, intentional use, flexibility and use of gestural sequences) relates to individual and dyadic measures of sociality and whether more complex gestural use is more effective in eliciting a response. We observed 19 captive chimpanzees (Pan troglodytes), 16 Sumatran orangutans (Pongo abelii) and 18 siamangs (Symphalangus syndactylus) to assess the complexity and effectiveness of their gestural use. We found that, beyond interspecies variation, the number of gesture types used in a dyad was higher when individuals had stronger social bonds; the probability of accounting for others\' attention increased with age, especially for visual gestures; and sequences were more likely used by younger or socially less integrated individuals. In terms of effectiveness, older individuals and those using fewer sequences were more likely to be responded to, while across dyads, the probability of obtaining a response was higher when both individuals accounted for the other\'s attention and when they used fewer sequences. Overall, this confirms the link between sociality and complex gestural use and suggests that more complex forms of communication, at least in terms of intentional use, may be more effective at achieving communicative goals. This article is part of the theme issue \'Cognition, communication and social bonds in primates\'.