Effectiveness of 20 years of conservation investments in protecting orangutans.

Conservation strategies are rarely systematically evaluated, which reduces transparency, hinders the cost-effective deployment of resources, and hides what works best in different contexts. Using data on the iconic and critically endangered orangutan (Pongo spp.), we developed a novel spatiotemporal framework for evaluating conservation investments. We show that around USD 1 billion was invested between 2000 and 2019 into orangutan conservation by governments, nongovernmental organizations, companies, and communities. Broken down by allocation to different conservation strategies, we find that habitat protection, patrolling, and public outreach had the greatest return on investment for maintaining orangutan populations. Given the variability in threats, land-use opportunity costs, and baseline remunerations in different regions, there were differential benefits per dollar invested across conservation activities and regions. We show that although challenging from a data and analysis perspective, it is possible to fully understand the relationships between conservation investments and outcomes and the external factors that influence these outcomes. Such analyses can provide improved guidance toward a more effective biodiversity conservation. Insights into the spatiotemporal interplays between the costs and benefits driving effectiveness can inform decisions about the most suitable orangutan conservation strategies for halting population declines. Although our study focuses on the three extant orangutan species of Sumatra and Borneo, our findings have broad application for evidence-based conservation science and practice worldwide.

Global Demand for Natural Resources Eliminated More Than 100,000 Bornean Orangutans.

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.

First integrative trend analysis for a great ape species in Borneo.

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.

Orangutan behavior in Kutai National Park after drought and fire damage: Adjustments to short- and long-term natural forest regeneration.

This study aimed to develop a long-term picture of orangutan (Pongo pygmaeus morio) behavioral adjustments to damaged masting forest around Mentoko, Kutai National Park, Indonesia. Mentoko is regenerating from two severe burnings and is one of few areas where orangutans were well-studied before and early after damage. We studied orangutans' feeding ecology, diet, and activity budgets 12-15 years after the second burning then compared our findings with earlier pre- and post-damage ones to assess the changes and factors involved. By our study, we predicted (1) improved feeding ecology compared to early in regeneration, (2) behavior diverging from the normal foraging strategy and (3) behavior shifting toward pre-damage patterns with improving feeding ecology. Data were behavioral observations on 42 orangutans (422 full day follows, 3,522 hr) and tree plot measures of feeding ecology. Findings were consistent with the first and third predictions but not the second: (1) feeding ecology had improved (plant food abundance was near per-damage levels, but species composition had changed); (2) foraging strategies showed no divergence from normal (fallback-preferred food switches in diet and activity budget adjustments were both normal, notably travel did not reduce), (3) diet and activity budgets had reverted to near pre-damage values by our study. Differently than post-damage studies on other orangutans but consistent with those on other primates, our comparisons showed behavioral adjustments were flexible, multiple vs. single, and influenced by multiple factors. Factors likely involved at Mentoko include type and spatial configuration of damage, duration of regeneration, and P. p. morio's recognized resilience. Findings have value for orangutan and nature conservation in showing that recovery to near-normal levels from severe habitat damage is possible within ca 15 years and in adding to understanding of the factors and processes that contribute to recovery.

Orangutan fish eating, primate aquatic fauna eating, and their implications for the origins of ancestral hominin fish eating.

This paper presents new evidence of fish eating in rehabilitant orangutans living on two Bornean islands and explores its contributions to understanding nonhuman primates' aquatic fauna eating and the origins of ancestral hominin fish eating. We assessed the prevalence of orangutans' fish eating, their techniques for obtaining fish, and possible contributors (ecology, individual differences, humans). We identified 61 events in which orangutans tried to obtain fish, including 19 in which they ate fish. All the orangutans were juvenile-adolescent; all the fish were disabled catfish; and most were obtained and eaten in drier seasons in or near shallow, slow-moving water. Orangutans used several techniques to obtain fish (inadvertent, opportunistic and deliberate hand-catch, scrounge, tool-assisted catch) and probably learned them in that order. Probable contributing factors were orangutan traits (age, pre-existing water or tool skills), island features (social density, water accessibility), and local human fishing. Our review of primates' aquatic fauna eating showed orangutans to be one of 20 species that eat aquatic fauna, one of nine confirmed to eat fish, and one of three that use tools to obtain fish. Primate fish eating is also site-specific within species, partly as a function of habitat (e.g., marine-freshwater, seasonality) and human influence (possibly fostered eating fish or other aquatic fauna at most sites, clearly induced it at some). At tropical freshwater sites, fish eating occurred most often in drier seasons around shallow water. Orangutan and primate findings are generally consistent with Stewart's (2010) reconstruction of the origins of ancestral hominin fish eating, but suggest that it, and tool-assisted fish catching, were possible much earlier.

Factors affecting reproduction in rehabilitant female orangutans: young age at first birth and short inter-birth interval.

This study investigated the reproductive parameters of free-ranging rehabilitant female orangutans. We aimed to assess the factors that influence these parameters and provide information that could assist with the management of orangutan reintroduction programs. We analyzed the birth records of free-ranging female rehabilitants at Bukit Lawang, Bukit Tigapuluh, Sepilok, Camp Leakey, Kaja Island, Sungai Wain, and Meratus and compared them with reproductive parameters reported in wild and zoo populations. Females' ages at first birth were 10.6-14.7 years, significantly earlier than those of wild and zoo orangutans. Computed inter-birth intervals (IBIs) calculated by the Kaplan-Meier method were 65.1-90.1 months; the values for Camp Leakey and Bukit Lawang rehabilitants were significantly shorter than those reported for wild Sumatran orangutans. Infant mortality rates were 18-61%; the values for Bukit Lawang and Sepilok were significantly higher than those reported for wild Sumatran and zoo orangutans. In rehabilitants, young ages at first birth and shorter IBIs may result from the high energy intake enabled by provisioning, although the possibility exists that they reflect underestimations of age on arrival at rehabilitation centers. The observed high infant mortality rate may reflect poor mothering skills due to human rearing and/or increased disease transmission. This study demonstrates that accelerated reproductive rates (younger age at first birth and shorter IBI) are common in provisioned rehabilitant females on both Sumatra and Borneo.

Pantomime in great apes: Evidence and implications.

We recently demonstrated, by mining observational data, that forest-living orangutans can communicate using gestures that qualify as pantomime.1 Pantomimes, like other iconic gestures, physically resemble their referents.2,3 More elaborately, pantomimes involve enacting their referents.4 Holding thumb and finger together at the lips and blowing between them to mean "balloon" is one example.5 Here we sketch evidence of pantomime in other great apes, methodological concerns and sophisticated cognitive capabilities that great ape pantomimes suggest.

Orangutan pantomime: elaborating the message.

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 free-ranging 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.

Life history: the energy-efficient orangutan.

A study of orangutans' daily energy expenditure confirmed exceptionally slow metabolism. It suggests they evolved a lifestyle designed to minimize energy use. If so, shifting to a higher energy-use strategy may help explain how humans evolved.

How orangutans (Pongo pygmaeus) innovate for water.

We report an observational field study that aimed to identify innovative processes in rehabilitant orangutans' (Pongo pygmaeus) water innovations on Kaja Island, Central Kalimantan, Indonesia. We tested for the basic model of innovating (make small changes to old behavior), 4 contributors (apply old behavior to new ends, accidents, independent working out, social cross-fertilization), development, and social rank. Focal observations of Kaja rehabilitants' behavior over 20 months yielded 18 probable innovations from among 44 water variants. We identified variants by function and behavioral grain, innovations by prevalence, and innovative processes by relations between innovations, other behaviors, and social encounters. Findings indicate innovating by small changes and some involvement of all 4 contributors; midrank orangutans were the most innovative; and rehabilitants' adolescent age profile, orphaning, and intense sociality probably enhanced innovativeness. Important complexities include: orangutan innovating may favor certain behavioral levels and narrowly defined similarities, and it may constitute a phase-like process involving a succession of changes and contributors. Discussion focuses on links with great ape cognition and parallels with innovating in humans and other nonhuman species.

Orangutan leaf-carrying for nest-building: toward unraveling cultural processes.

We report an empirical study on leaf-carrying, a newly discovered nest-building technique that involves collecting nest materials before reaching the nest site. We assessed whether leaf-carrying by rehabilitant orangutans on Kaja Island, Central Kalimantan, owes to cultural influences. Findings derive from ca 600 h observational data on nesting skills and nesting associations in Kaja's 42 resident rehabilitants, which yielded 355 nests and 125 leaf-carrying cases by 34 rehabilitants. Regional contrasts with 14 other communities (7 rehabilitant, 7 wild) indicated cultural influences on leaf-carrying on Kaja. Association data showed exceptional social learning opportunities for leaf-carrying on Kaja, with residents taking differential advantage of these opportunities as a function of development, experience, and social position. Juvenile males with basic nesting skills were most influenced by social input. Most (27) leaf-carriers had probably learned leaf-carrying when caged and 7 probably learned it on Kaja. Social priming was probably the main impetus to leaf-carrying on Kaja, by simply prompting observers to copy when leaf-carrying associates collected nesting materials, what they collected, and where they used their collected materials. Implications concern acquisition processes and ontogenetic schedules that orchestrate sets of features-needs or interests, cognitive abilities, social preferences-which enable cultural transmission.