To explain social learning without invoking the cognitively complex concept of imitation, many learning mechanisms have been proposed. Borrowing an idea used routinely in cognitive psychology, we argue that most of these alternatives can be subsumed under a single process, priming, in which input increases the activation of stored internal representations. Imitation itself has generally been seen as a “special faculty.” This has diverted much research towards the all-or-none question of whether an animal can imitate, with disappointingly inconclusive results. In the great apes, however, voluntary, learned behaviour is organized hierarchically. This means that imitation can occur at various levels, of which we single out two clearly distinct ones: the “action level,” a rather detailed and linear specification of sequential acts, and the “program level,” a broader description of subroutine structure and the hierarchical layout of a behavioural “program.” Program level imitation is a high-level, constructive mechanism, adapted for the efficient learning of complex skills and thus not evident in the simple manipulations used to test for imitation in the laboratory. As examples, we describe the food-preparation techniques of wild mountain gorillas and the imitative behaviour of orangutans undergoing “rehabilitation” to the wild. Representing and manipulating relations between objects seems to be one basic building block in their hierarchical programs. There is evidence that great apes suffer from a stricter capacity limit than humans in the hierarchical depth of planning. We re-interpret some chimpanzee behaviour previously described as “emulation” and suggest that all great apes may be able to imitate at the program level. Action level imitation is seldom observed in great ape skill learning, and may have a largely social role, even in humans.

We made an observational study of spontaneous imitation in orangutans (Pongo pygmaeus). Previous studies may have underestimated great apes' imitative capacities by studying subjects under inhibiting conditions. We used subjects living in enriched environments, namely, rehabilitation. We collected a sample of spontaneous imitations and analyzed the most complex incidents for the likelihood that true imitation, learning new actions by observing rather than by doing, was involved in their acquisition. From 395 hr of observation and other reports on 26 orangutans, we identified 354 incidents of imitation. Of these, 54 complex incidents were difficult to explain by forms of imitation based on associative processes grounded in experimental learning alone; they were, however, congruent with acquisition processes that include true imitation. These findings suggest that orangutans may be capable of true imitation and point to critical eliciting factors.

Unsustainable exploitation of natural resources is increasingly affecting the highly biodiverse tropics [1, 2]. Although rapid developments in remote sensing technology have permitted more precise estimates of land-cover change over large spatial scales [3-5], our knowledge about the effects of these changes on wildlife is much more sparse [6, 7]. Here we use field survey data, predictive density distribution modeling, and remote sensing to investigate the impact of resource use and land-use changes on the density distribution of Bornean orangutans (Pongo pygmaeus). Our models indicate that between 1999 and 2015, half of the orangutan population was affected by logging, deforestation, or industrialized plantations. Although land clearance caused the most dramatic rates of decline, it accounted for only a small proportion of the total loss. A much larger number of orangutans were lost in selectively logged and primary forests, where rates of decline were less precipitous, but where far more orangutans are found. This suggests that further drivers, independent of land-use change, contribute to orangutan loss. This finding is consistent with studies reporting hunting as a major cause in orangutan decline [8-10]. Our predictions of orangutan abundance loss across Borneo suggest that the population decreased by more than 100,000 individuals, corroborating recent estimates of decline [11]. Practical solutions to prevent future orangutan decline can only be realized by addressing its complex causes in a holistic manner across political and societal sectors, such as in land-use planning, resource exploitation, infrastructure development, and education, and by increasing long-term sustainability [12]. VIDEO ABSTRACT.

of feeding time to different food types such as fruit, vegetable matter, bark, insects, and other items (e.g. Fox et al. 2004; Wich et al. 2006a; Chapter 8 this volume). Understanding a species' diet, however, also requires a comprehensive list of the food items eaten (Tutin et al. 1994; Rodman 2002). 9.1 Introduction This chapter compares orangutan diets, both across orangutan sites and relative to other great apes, based on current food lists. Orangutan diets are typically represented in terms of the allocation

For many threatened species the rate and drivers of population decline are difficult to assess accurately: species’ surveys are typically restricted to small geographic areas, are conducted over short time periods, and employ a wide range of survey protocols. We addressed methodological challenges for assessing change in the abundance of an endangered species. We applied novel methods for integrating field and interview survey data for the critically endangered Bornean orangutan (Pongo pygmaeus), allowing a deeper understanding of the species’ persistence through time. Our analysis revealed that Bornean orangutan populations have declined at a rate of 25% over the last 10 years. Survival rates of the species are lowest in areas with intermediate rainfall, where complex interrelations between soil fertility, agricultural productivity, and human settlement patterns influence persistence. These areas also have highest threats from human-wildlife conflict. Survival rates are further positively associated with forest extent, but are lower in areas where surrounding forest has been recently converted to industrial agriculture. Our study highlights the urgency of determining specific management interventions needed in different locations to counter the trend of decline and its associated drivers.

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We present an exploratory study of forest-living orangutan pantomiming, i.e. gesturing in which they act out their meaning, focusing on its occurrence, communicative functions, and complexities. Studies show that captive great apes may elaborate messages if communication fails, and isolated reports suggest that great apes occasionally pantomime. We predicted forest-living orangutans would pantomime spontaneously to communicate, especially to elaborate after communication failures. Mining existing databases on free-ranging rehabilitant orangutans' behaviour identified 18 salient pantomimes. These pantomimes most often functioned as elaborations of failed requests, but also as deceptions and declaratives. Complexities identified include multimodality, re-enactments of past events and several features of language (productivity, compositionality, systematicity). These findings confirm that freeranging rehabilitant orangutans pantomime and use pantomime to elaborate on their messages. Further, they use pantomime for multiple functions and create complex pantomimes that can express propositionally structured content. Thus, orangutan pantomime serves as a medium for communication, not a particular function. Mining cases of complex great ape communication originally reported in functional terms may then yield more evidence of pantomime.

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