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Pangolins are the most trafficked mammals in the world and are severely threatened by poaching the loss, degradation, and fragmentation of habitat. In Malaysian Borneo, conservation initiatives for the Sunda pangolin (Manis javanica) are hindered by a paucity of data on their distribution and population size. Using MaxEnt niche modelling and consolidated species location data, we projected the distribution of Sunda pangolins in Sabah. Additionally, we assessed the accessibility of their forest habitats to humans to understand potential threats. Our model indicated that, as of 2015, approximately half of Sabah’s land area (39,530 km²) is suitable for pangolins, with 43% in protected forests, 38% in production forests, and 19% outside of these areas. Alarmingly, our data suggest that nearly all (91%) of these suitable habitats are relatively easily accessible to poachers. Our findings provide a state-level baseline understanding of Sunda pangolin distribution and assess potential threats in Sabah. These can inform short- and long-term conservation management plans for pangolin to safeguard this critically endangered species.

Various global-scale proposals exist to reduce the loss of biological diversity. These include the Half-Earth and Whole-Earth visions that respectively seek to set aside half the planet for wildlife conservation or to diversify conservation practices fundamentally and change the economic systems that determine environmental harm. Here we assess these visions in the specific context of Bornean orangutans Pongo pygmaeus and their conservation. Using an expert-led process we explored three scenarios over a 10-year time frame: continuation of Current Conditions, a Half-Earth approach and a Whole-Earth approach. In addition, we examined a 100-year population recovery scenario assuming 0% offtake of Bornean orangutans. Current Conditions were predicted to result in a population c. 73% of its current size by 2032. Half-Earth was judged comparatively easy to achieve and predicted to result in an orangutan population of c. 87% of its current size by 2032. Whole-Earth was anticipated to lead to greater forest loss and ape killing, resulting in a prediction of c. 44% of the current orangutan population for 2032. Finally, under the recovery scenario, populations could be c. 148% of their current size by 2122. Although we acknowledge uncertainties in all of these predictions, we conclude that the Half-Earth and Whole-Earth visions operate along different timelines, with the implementation of Whole-Earth requiring too much time to benefit orangutans. None of the theorized proposals provided a complete solution, so drawing elements from each will be required. We provide recommendations for equitable outcomes.

Abstract Conservation funding is currently limited; cost‐effective conservation solutions are essential. We suggest that the thousands of field stations worldwide can play key roles at the frontline of biodiversity conservation and have high intrinsic value. We assessed field stations’ conservation return on investment and explored the impact of COVID‐19. We surveyed leaders of field stations across tropical regions that host primate research; 157 field stations in 56 countries responded. Respondents reported improved habitat quality and reduced hunting rates at over 80% of field stations and lower operational costs per km2 than protected areas, yet half of those surveyed have less funding now than in 2019. Spatial analyses support field station presence as reducing deforestation. These “earth observatories” provide a high return on investment; we advocate for increased support of field station programs and for governments to support their vital conservation efforts by investing accordingly.

Abstract Despite decades of conservation management, many orangutan populations are on the brink of extinction. This is primarily due to habitat loss and direct killings. A study from 2008/2009 suggested that killing was impacting orangutan populations at a rate sufficient to cause local extinctions. As an illegal and taboo behavior that is difficult to measure, killing has been severely understudied since. We conducted 431 interviews in 79 villages across Kalimantan in 2020/2021. Ours is the first quantitative field study in more than 10 years to assess the state of killing of orangutans. We aimed to: (1) assess the current state of killing of orangutans in Kalimantan and compare this to the previous study; (2) determine whether conservation projects are affecting killing; and (3) explore drivers of killing. We examined killing of orangutans across villages with forest conservation projects, orangutan conservation projects, and no conservation projects. We assessed the existence of killing and used scenarios to examine perceived norms about illegal behavior relating to orangutans. We then used matching techniques to assess whether projects have any impact on these indicators. Overall, our findings suggest that killing has occurred in recent times, and our data does not indicate a clear attenuation of the behavior. As such, we argue that killing may still present a substantial threat to Bornean orangutan populations. We also found no statistically significant evidence that conservation projects are reducing killing. Conservation project managers could seek to understand the drivers of killing, and to invest in interventions that address these drivers. Research suggests that current allocation of conservation funding has been ineffective at abating orangutan population decline. We argue that a key part of improving Bornean orangutan conservation practice involves directly addressing killing of orangutans and the underlying drivers of killing.

Non-human great apes – chimpanzees, gorillas, bonobos, and orangutans – are threatened by agricultural expansion, particularly from rice, cacao, cassava, maize, and oil palm cultivation. Agriculture replaces and fragments great ape habitats, bringing them closer to humans and often resulting in conflict. Though the impact of agriculture on great apes is well-recognized, there is still a need for a more nuanced understanding of specific contexts and associated negative impacts on habitats and populations. Here we review these contexts and their implications for great apes. We estimate that within their African and South-East Asian ranges, there are about 100 people for each great ape. Given that most apes live outside strictly protected areas and the growing human population and increasing demand for resources in these landscapes, it will be challenging to balance the needs of both humans and great apes. Further habitat loss is expected, particularly in Africa, where compromises must be sought to re-direct agricultural expansion driven by subsistence farmers with small fields (generally

Assessing where wildlife populations are at risk from future habitat loss is particularly important for land-use planning and avoiding biodiversity declines. Combining projections of future deforestation with species density information provides an improved way to anticipate such declines. Using the critically endangered Bornean orangutan (Pongo pygmaeus) as a case study we applied a spatio-temporally explicit deforestation model to forest loss data from 2001 to 2017 and projected future impacts on orangutans to the 2030s. Our projections point to continued deforestation across the island, amounting to a potential loss of forest habitat for 26,200 orangutans. Populations currently persisting in forests gazetted for industrial timber and oil palm concessions, or unprotected forests outside of concessions, were projected to experience the worst losses within the next 15 years, amounting to 15,400 individuals. Our analysis indicates the importance of protecting orangutan habitat in plantation landscapes, maintaining protected areas and efforts to prevent the conversion of logged forests for the survival of highly vulnerable wildlife. The modeling framework could be expanded to other species with available density or occurrence data. Our findings highlight that species conservation should not only act on the current information, but also anticipate future changes to be effective.

Critically Endangered orangutans are translocated in several situations: reintroduced into historic range where no wild populations exist, released to reinforce existing wild populations, and wild-to-wild translocated to remove individuals from potentially risky situations. Translocated orangutans exposed to human diseases, including Coronavirus Disease 2019 (COVID-19), pose risks to wild and previously released conspecifics. Wildlife disease risk experts recommended halting great ape translocations during the COVID-19 pandemic to minimize risk of disease transmission to wild populations. We collected data on orangutan releases and associated disease risk management in Indonesia during the COVID-19 pandemic, and developed a problem description for orangutan disease and conservation risks. We identified that at least 15 rehabilitated ex-captive and 27 wild captured orangutans were released during the study period. Identified disease risks included several wild-to-wild translocated orangutans in direct contact or proximity to humans without protective equipment, and formerly captive rehabilitated orangutans that have had long periods of contact and potential exposure to human diseases. While translocation practitioners typically employ mitigation measures to decrease disease transmission likelihood, these measures cannot eliminate all risk, and are not consistently applied. COVID-19 and other diseases of human origin can be transmitted to orangutans, which could have catastrophic impacts on wild orangutans, other susceptible fauna, and humans should disease transmission occur. We recommend stakeholders conduct a Disease Risk Analysis for orangutan translocation, and improve pathogen surveillance and mitigation measures to decrease the likelihood of potential outbreaks. We also suggest refocusing conservation efforts on alternatives to wild-to-wild translocation including mitigating human-orangutan interactions, enforcing laws and protecting orangutan habitats to conserve orangutans in situ.

Historically, orangutans (Pongo spp.) lived in large contiguous areas of intact rainforest. Today, they are also found in highly modified and fragmented landscapes dominated by oil palm or industrial timber plantations; a situation that calls for new conservation approaches. Here we report signs of orangutan presence in more than 120 small forest fragments of

Abstract 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.

In conservation, Collaboration is thought to improve returns from investment and is frequently encouraged, however not all collaborations are equal and may therefore lack characteristics important for addressing collective action problems. Furthermore, partnerships that are advantageous for a collective may not necessarily be advantageous for an individual. This study investigated collaboration within the Bornean orangutan (Pongo pygmaeus) conservation sector – a system with reported inefficiencies and for which there has been a renewed call for collaborative partnerships. Collaborative partnerships were conceptualised as a social network and analysed using exponential random graph modelling. The prevalence of structural attributes associated with social processes considered to be important for solving collective action problems such as trust and innovation were investigated. Qualitative surveying techniques were used to measure the perceptions of collaboration held by individual actors within the network and the impact of organizational attributes on network formation and perceptions was assessed. Collaboration was found to be occurring within the conservation network and was positively perceived at the individual organisational level. At the collective level, the current collaborative network contains some structural characteristics important for addressing the collective-action problem of orangutan conservation, particularly through innovation and knowledge sharing. However efforts to develop trust between organisations may be needed. To improve returns on investment, future collaborative partnerships must be strategically implemented with individual roles and desired overall outcomes explicitly articulated. Increased operational transparency and improved performance evaluation will be critical for achieving improved collaborative efficiency.

The rise of palm oil as the world's most consumed vegetable oil has coincided with exponential growth in palm oil research activity. Bibliometric analysis of research outputs reveals a distinct imbalance in the type of research being undertaken, notably a disproportionate focus on biofuel and engineering topics. Recognizing the expansion of oil palm agriculture across the tropics and the increasing awareness of environmental, social, and economic impacts, we seek to reorientate the existing research agenda toward one that addresses the most fundamental and urgent questions defined by the palm oil stakeholder community. Following consultation with 659 stakeholders from 38 countries, including palm oil growers, government agencies, non-governmental organizations, and researchers, the highest priority research questions were identified within 13 themes. The resulting 279 questions, including 26 ranked as top priority, reveal a diversity of environmental and social research challenges facing the industry, ranging from the ecological and ecosystem impacts of production, to the livelihoods of plantation workers and smallholder communities. Analysis of the knowledge type produced from these questions underscores a clear need for fundamental science programmes, and studies that involve the consultation of non-academic stakeholders to develop “transformative” solutions to the oil palm sector. Stakeholders were most aligned in their choice of priority questions across the themes of policy and certification related themes, and differed the most in environmental feedback, technology and smallholder related themes. Our recommendations include improved regional academic leadership and coordination, greater engagement with private and public stakeholders in Africa, and Central and South America, and enhanced collaborative efforts with researchers in the major consuming countries of India and China.

Designed as a new method to facilitate the reintroduction and post-release monitoring of orangutans and other apes, implanted radio-telemetry (IRT) was developed and first deployed in 2009. Since that time, it has been necessary to collate and review information on its uptake and general efficacy to inform its ongoing development and that of other emerging tracking technologies. We present here technical specifications and the surgical procedure used to implant miniaturized radio transmitters, as well as a formal testing procedure for measuring detectable transmission distances of implanted devices. Feedback from IRT practitioners (veterinarians and field managers) was gathered through questionnaires and is also presented. To date, IRT has been used in at least 250 individual animals (mainly orangutans) from four species of ape in both Asia and Africa. Median surgical and wound healing times were 30 min and 15 days, respectively, with implants needing to be removed on at least 36 separate occasions. Confirmed failures within the first year of operation were 18.1%, while longer distances were reported from positions of higher elevation relative to the focal animal. IRT has been a transformational technology in facilitating the relocation of apes after their release, resulting in much larger amounts of post-release data collection than ever before. It is crucial however, that implant casings are strengthened to prevent the requirement for recapture and removal surgeries, especially for gradually adapting apes. As with all emerging technological solutions, IRT carries with it inherent risk, especially so due to the requirement for subcutaneous implantation. These risks must, however, be balanced with the realities of releasing an animal with no means of relocation, as has historically been, and is still, the case with orangutans and gorillas.

Abstract The PREDICTS project—Projecting Responses of Ecological Diversity In Changing Terrestrial Systems (https://www.nhm.ac.uk/our-science/our-work/biodiversity/predicts.html)—has collated from published studies a large, reasonably representative database of comparable samples of biodiversity from multiple sites that differ in the nature or intensity of human impacts relating to land use. We have used this evidence base to develop global and regional statistical models of how local biodiversity responds to these measures. We describe and make freely available this 2016 release of the database, containing more than 3.2 million records sampled at over 26,000 locations and representing over 47,000 species. We outline how the database can help in answering a range of questions in ecology and conservation biology. To our knowledge, this is the largest and most geographically and taxonomically representative database of spatial comparisons of biodiversity that has been collated to date; it will be useful to researchers and international efforts wishing to model and understand the global status of biodiversity.

The Bornean orangutan is critically endangered and monitoring its population is needed to inform effective conservation management. In this paper, we present results of 2014-17 aerial nest surveys of the major orangutan populations in Sabah and compare them with baseline data produced during surveys conducted in 2002-03 using similar methods. Our results show three important points: a) by increasing the survey effort (estimated at 15-25% cover), sparsely scattered orangutan sub-populations not recorded in the previous aerial surveys were located and the accuracy of the nest count estimates is expected to improve; b) large populations in the interior forests of Sabah, occupying sustainably managed logged and unlogged forests, have been stable over 15 years and are of vital importance for the species' conservation; c) fragmented populations located in eastern Sabah, that are surrounded by extensive oil palm plantations, have declined at varying rates.

Abstract Biodiversity continues to decline in the face of increasing anthropogenic pressures such as habitat destruction, exploitation, pollution and introduction of alien species. Existing global databases of species’ threat status or population time series are dominated by charismatic species. The collation of datasets with broad taxonomic and biogeographic extents, and that support computation of a range of biodiversity indicators, is necessary to enable better understanding of historical declines and to project – and avert – future declines. We describe and assess a new database of more than 1.6 million samples from 78 countries representing over 28,000 species, collated from existing spatial comparisons of local‐scale biodiversity exposed to different intensities and types of anthropogenic pressures, from terrestrial sites around the world. The database contains measurements taken in 208 (of 814) ecoregions, 13 (of 14) biomes, 25 (of 35) biodiversity hotspots and 16 (of 17) megadiverse countries. The database contains more than 1% of the total number of all species described, and more than 1% of the described species within many taxonomic groups – including flowering plants, gymnosperms, birds, mammals, reptiles, amphibians, beetles, lepidopterans and hymenopterans. The dataset, which is still being added to, is therefore already considerably larger and more representative than those used by previous quantitative models of biodiversity trends and responses. The database is being assembled as part of the PREDICTS project (Projecting Responses of Ecological Diversity In Changing Terrestrial Systems – https://www.nhm.ac.uk/our-science/our-work/biodiversity/predicts.html). We make site‐level summary data available alongside this article. The full database will be publicly available in 2015.

Reducing Emissions from Deforestation and forest Degradation (REDD+) aims to avoid forest conversion to alternative land-uses through financial incentives. Oil-palm has high opportunity costs, which according to current literature questions the financial competitiveness of REDD+ in tropical lowlands. To understand this more, we undertook regional fine-scale and coarse-scale analyses (through carbon mapping and economic modelling) to assess the financial viability of REDD+ in safeguarding unprotected forest (30,173 ha) in the Lower Kinabatangan floodplain in Malaysian Borneo. Results estimate 4.7 million metric tons of carbon (MgC) in unprotected forest, with 64% allocated for oil-palm cultivations. Through fine-scale mapping and carbon accounting, we demonstrated that REDD+ can outcompete oil-palm in regions with low suitability, with low carbon prices and low carbon stock. In areas with medium oil-palm suitability, REDD+ could outcompete oil palm in areas with: very high carbon and lower carbon price; medium carbon price and average carbon stock; or, low carbon stock and high carbon price. Areas with high oil palm suitability, REDD+ could only outcompete with higher carbon price and higher carbon stock. In the coarse-scale model, oil-palm outcompeted REDD+ in all cases. For the fine-scale models at the landscape level, low carbon offset prices (US $3 MgCO2e) would enable REDD+ to outcompete oil-palm in 55% of the unprotected forests requiring US $27 million to secure these areas for 25 years. Higher carbon offset price (US $30 MgCO2e) would increase the competitiveness of REDD+ within the landscape but would still only capture between 69%-74% of the unprotected forest, requiring US $380-416 million in carbon financing. REDD+ has been identified as a strategy to mitigate climate change by many countries (including Malaysia). Although REDD+ in certain scenarios cannot outcompete oil palm, this research contributes to the global REDD+ debate by: highlighting REDD+ competitiveness in tropical floodplain landscapes; and, providing a robust approach for identifying and targeting limited REDD+ funds.

Plant recovery rates after herbivory are thought to be a key factor driving recursion by herbivores to sites and plants to optimise resource-use but have not been investigated as an explanation for recursion in large herbivores. We investigated the relationship between plant recovery and recursion by elephants (Elephas maximus borneensis) in the Lower Kinabatangan Wildlife Sanctuary, Sabah. We identified 182 recently eaten food plants, from 30 species, along 14 × 50 m transects and measured their recovery growth each month over nine months or until they were re-browsed by elephants. The monthly growth in leaf and branch or shoot length for each plant was used to calculate the time required (months) for each species to recover to its pre-eaten length. Elephant returned to all but two transects with 10 eaten plants, a further 26 plants died leaving 146 plants that could be re-eaten. Recursion occurred to 58% of all plants and 12 of the 30 species. Seventy-seven percent of the re-eaten plants were grasses. Recovery times to all plants varied from two to twenty months depending on the species. Recursion to all grasses coincided with plant recovery whereas recursion to most browsed plants occurred four to twelve months before they had recovered to their previous length. The small sample size of many browsed plants that received recursion and uneven plant species distribution across transects limits our ability to generalise for most browsed species but a prominent pattern in plant-scale recursion did emerge. Plant recovery time was a good predictor of time to recursion but varied as a function of growth form (grass, ginger, palm, liana and woody) and differences between sites. Time to plant recursion coincided with plant recovery time for the elephant’s preferred food, grasses, and perhaps also gingers, but not the other browsed species. Elephants are bulk feeders so it is likely that they time their returns to bulk feed on these grass species when quantities have recovered sufficiently to meet their intake requirements. The implications for habitat and elephant management are discussed.

Industrial oil palm plantations in South East Asia have caused significant biodiversity losses and perverse social outcomes. To address concerns over plantation practices and in an attempt to improve sustainability through market mechanisms, civil society organisations and industry representatives developed the Roundtable on Sustainable Palm Oil (RSPO) in 2004. The effectiveness of RSPO in improving the sustainability of the palm oil industry is frequently debated and to date, few quantitative analyses have been undertaken to assess how successful RSPO has been in delivering the social, economic and environmental sustainability outcomes it aims to address. With the palm oil industry continuing to expand in South East Asia and significant estates being planted in Africa and South America, this paper evaluates the effectiveness of RSPO plantations compared to non-certified plantations by assessing the relative performance of several key sustainability metrics compared to business as usual practices. Using Indonesian Borneo (Kalimantan) as a case study, a novel dataset of RSPO concessions was developed and causal analysis methodologies employed to evaluate the environmental, social and economic sustainability of the industry. No significant difference was found between certified and non-certified plantations for any of the sustainability metrics investigated, however positive economic trends including greater fresh fruit bunch yields were revealed. To achieve intended outcomes, RSPO principles and criteria are in need of substantial improvement and rigorous enforcement.

The Sumatran orangutan is currently listed by the IUCN as critically endangered and the Bornean species as endangered. Unless effective conservation measures are enacted quickly, most orangutan populations without adequate protection face a dire future. Two main strategies are being pursued to conserve orangutans: (i) rehabilitation and reintroduction of ex-captive or displaced individuals; and (ii) protection of their forest habitat to abate threats like deforestation and hunting. These strategies are often mirrored in similar programs to save other valued and endangered mega-fauna. Through GIS analysis, collating data from across the literature, and combining this information within a modelling and decision analysis framework, we analysed which strategy or combination of strategies is the most cost-effective at maintaining wild orangutan populations, and under what conditions. We discovered that neither strategy was optimal under all circumstances but was dependent on the relative cost per orangutan, the timescale of management concern, and the rate of deforestation. Reintroduction, which costs twelve times as much per animal as compared to protection of forest, was only a cost-effective strategy at very short timescales. For time scales longer than 10-20 years, forest protection is the more cost-efficient strategy for maintaining wild orangutan populations. Our analyses showed that a third, rarely utilised strategy is intermediate: introducing sustainable logging practices and protection from hunting in timber production forest. Maximum long-term cost-efficiency is achieved by working in conservation forest. However, habitat protection involves addressing complex conservation issues and conflicting needs at the landscape level. We find a potential resolution in that well-managed production forests could achieve intermediate conservation outcomes. This has broad implications for sustaining biodiversity more generally within an economically productive landscape. Insights from this analysis should provide a better framework to prioritize financial investments, and facilitate improved integration between the organizations that implement these strategies.

The native forests of Borneo have been impacted by selective logging, fire, and conversion to plantations at unprecedented scales since industrial-scale extractive industries began in the early 1970s. There is no island-wide documentation of forest clearance or logging since the 1970s. This creates an information gap for conservation planning, especially with regard to selectively logged forests that maintain high conservation potential. Analysing LANDSAT images, we estimate that 75.7% (558,060 km2) of Borneo's area (737,188 km2) was forested around 1973. Based upon a forest cover map for 2010 derived using ALOS-PALSAR and visually reviewing LANDSAT images, we estimate that the 1973 forest area had declined by 168,493 km2 (30.2%) in 2010. The highest losses were recorded in Sabah and Kalimantan with 39.5% and 30.7% of their total forest area in 1973 becoming non-forest in 2010, and the lowest in Brunei and Sarawak (8.4%, and 23.1%). We estimate that the combined area planted in industrial oil palm and timber plantations in 2010 was 75,480 km2, representing 10% of Borneo. We mapped 271,819 km of primary logging roads that were created between 1973 and 2010. The greatest density of logging roads was found in Sarawak, at 0.89 km km-2, and the lowest density in Brunei, at 0.18 km km-2. Analyzing MODIS-based tree cover maps, we estimate that logging operated within 700 m of primary logging roads. Using this distance, we estimate that 266,257 km2 of 1973 forest cover has been logged. With 389,566 km2 (52.8%) of the island remaining forested, of which 209,649 km2 remains intact. There is still hope for biodiversity conservation in Borneo. Protecting logged forests from fire and conversion to plantations is an urgent priority for reducing rates of deforestation in Borneo.

Lowland tropical forests are increasingly threatened with conversion to oil palm as global demand and high profit drives crop expansion throughout the world's tropical regions. Yet, landscapes are not homogeneous and regional constraints dictate land suitability for this crop. We conducted a regional study to investigate spatial and economic components of forest conversion to oil palm within a tropical floodplain in the Lower Kinabatangan, Sabah, Malaysian Borneo. The Kinabatangan ecosystem harbours significant biodiversity with globally threatened species but has suffered forest loss and fragmentation. We mapped the oil palm and forested landscapes (using object-based-image analysis, classification and regression tree analysis and on-screen digitising of high-resolution imagery) and undertook economic modelling. Within the study region (520,269 ha), 250,617 ha is cultivated with oil palm with 77% having high Net-Present-Value (NPV) estimates ($413/ha-yr-$637/ha-yr); but 20.5% is under-producing. In fact 6.3% (15,810 ha) of oil palm is commercially redundant (with negative NPV of $-299/ha-yr-$-65/ha-yr) due to palm mortality from flood inundation. These areas would have been important riparian or flooded forest types. Moreover, 30,173 ha of unprotected forest remain and despite its value for connectivity and biodiversity 64% is allocated for future oil palm. However, we estimate that at minimum 54% of these forests are unsuitable for this crop due to inundation events. If conversion to oil palm occurs, we predict a further 16,207 ha will become commercially redundant. This means that over 32,000 ha of forest within the floodplain would have been converted for little or no financial gain yet with significant cost to the ecosystem. Our findings have globally relevant implications for similar floodplain landscapes undergoing forest transformation to agriculture such as oil palm. Understanding landscape level constraints to this crop, and transferring these into policy and practice, may provide conservation and economic opportunities within these seemingly high opportunity cost landscapes.

We investigated why orangutans are being killed in Kalimantan, Indonesia, and the role of conflict in these killings. Based on an analysis of interview data from over 5,000 respondents in over 450 villages, we also assessed the socio-ecological factors associated with conflict and non-conflict killings. Most respondents never kill orangutans. Those who reported having personally killed an orangutan primarily did so for non-conflict reasons; for example, 56% of these respondents said that the reason they had killed an orangutan was to eat it. Of the conflict-related reasons for killing, the most common reasons orangutans were killed was fear of orangutans or in self-defence. A similar pattern was evident among reports of orangutan killing by other people in the villages. Regression analyses indicated that religion and the percentage of intact forest around villages were the strongest socio-ecological predictors of whether orangutans were killed for conflict or non-conflict related reasons. Our data indicate that between 44,170 and 66,570 orangutans were killed in Kalimantan within the respondents' active hunting lifetimes: between 12,690 and 29,024 for conflict reasons (95%CI) and between 26,361 and 41,688 for non-conflict reasons (95% CI). These findings confirm that habitat protection alone will not ensure the survival of orangutans in Indonesian Borneo, and that effective reduction of orangutan killings is urgently needed.

We ascertained villagers' perceptions about the importance of forests for their livelihoods and health through 1,837 reliably answered interviews of mostly male respondents from 185 villages in Indonesian and Malaysian Borneo. Variation in these perceptions related to several environmental and social variables, as shown in classification and regression analyses. Overall patterns indicated that forest use and cultural values are highest among people on Borneo who live close to remaining forest, and especially among older Christian residents. Support for forest clearing depended strongly on the scale at which deforestation occurs. Deforestation for small-scale agriculture was generally considered to be positive because it directly benefits people's welfare. Large-scale deforestation (e.g., for industrial oil palm or acacia plantations), on the other hand, appeared to be more context-dependent, with most respondents considering it to have overall negative impacts on them, but with people in some areas considering the benefits to outweigh the costs. The interviews indicated high awareness of negative environmental impacts of deforestation, with high levels of concern over higher temperatures, air pollution and loss of clean water sources. Our study is unique in its geographic and trans-national scale. Our findings enable the development of maps of forest use and perceptions that could inform land use planning at a range of scales. Incorporating perspectives such as these could significantly reduce conflict over forest resources and ultimately result in more equitable development processes.

Combining protected areas with natural forest timber concessions may sustain larger forest landscapes than is possible via protected areas alone. However, the role of timber concessions in maintaining natural forest remains poorly characterized. An estimated 57% (303,525 km²) of Kalimantan's land area (532,100 km²) was covered by natural forest in 2000. About 14,212 km² (4.7%) had been cleared by 2010. Forests in oil palm concessions had been reduced by 5,600 km² (14.1%), while the figures for timber concessions are 1,336 km² (1.5%), and for protected forests are 1,122 km² (1.2%). These deforestation rates explain little about the relative performance of the different land use categories under equivalent conversion risks due to the confounding effects of location. An estimated 25% of lands allocated for timber harvesting in 2000 had their status changed to industrial plantation concessions in 2010. Based on a sample of 3,391 forest plots (1×1 km; 100 ha), and matching statistical analyses, 2000-2010 deforestation was on average 17.6 ha lower (95% C.I.: -22.3 ha- -12.9 ha) in timber concession plots than in oil palm concession plots. When location effects were accounted for, deforestation rates in timber concessions and protected areas were not significantly different (Mean difference: 0.35 ha; 95% C.I.: -0.002 ha-0.7 ha). Natural forest timber concessions in Kalimantan had similar ability as protected areas to maintain forest cover during 2000-2010, provided the former were not reclassified to industrial plantation concessions. Our study indicates the desirability of the Government of Indonesia designating its natural forest timber concessions as protected areas under the IUCN Protected Area Category VI to protect them from reclassification.

Nature Communications 6: Article number:6819 (2015); Published 14 April 2015; Updated 14 June 2016 The original version of this Article contained an error in the spelling of the author Hugh P. Possingham, which was incorrectly given as Hugh P. Posssingham. This has now been corrected in both the PDFand HTML versions of the Article.

The approximately 300 (298, 95% CI: 152-581) elephants in the Lower Kinabatangan Managed Elephant Range in Sabah, Malaysian Borneo are a priority sub-population for Borneo's total elephant population (2,040, 95% CI: 1,184-3,652). Habitat loss and human-elephant conflict are recognized as the major threats to Bornean elephant survival. In the Kinabatangan region, human settlements and agricultural development for oil palm drive an intense fragmentation process. Electric fences guard against elephant crop raiding but also remove access to suitable habitat patches. We conducted expert opinion-based least-cost analyses, to model the quantity and configuration of available suitable elephant habitat in the Lower Kinabatangan, and called this the Elephant Habitat Linkage. At 184 km(2), our estimate of available habitat is 54% smaller than the estimate used in the State's Elephant Action Plan for the Lower Kinabatangan Managed Elephant Range (400 km(2)). During high flood levels, available habitat is reduced to only 61 km(2). As a consequence, short-term elephant densities are likely to surge during floods to 4.83 km(-2) (95% CI: 2.46-9.41), among the highest estimated for forest-dwelling elephants in Asia or Africa. During severe floods, the configuration of remaining elephant habitat and the surge in elephant density may put two villages at elevated risk of human-elephant conflict. Lower Kinabatangan elephants are vulnerable to the natural disturbance regime of the river due to their limited dispersal options. Twenty bottlenecks less than one km wide throughout the Elephant Habitat Linkage, have the potential to further reduce access to suitable habitat. Rebuilding landscape connectivity to isolated habitat patches and to the North Kinabatangan Managed Elephant Range (less than 35 km inland) are conservation priorities that would increase the quantity of available habitat, and may work as a mechanism to allow population release, lower elephant density, reduce human-elephant conflict, and enable genetic mixing.

Nature-based tourism can generate important revenue to support conservation of biodiversity. However, constant exposure to tourists and subsequent chronic activation of stress responses can produce pathological effects, including impaired cognition, growth, reproduction, and immunity in the same animals we are interested in protecting. Utilizing fecal samples (N = 53) from 2 wild habituated orangutans (Pongo pygmaeus morio) (in addition to 26 fecal samples from 4 wild unhabituated orangutans) in the Lower Kinabatangan Wildlife Sanctuary of Sabah, Malaysian Borneo, we predicted that i) fecal glucocorticoid metabolite concentrations would be elevated on the day after tourist visitation (indicative of normal stress response to exposure to tourists on the previous day) compared to samples taken before or during tourist visitation in wild, habituated orangutans, and ii) that samples collected from habituated animals would have lower fecal glucocorticoid metabolites than unhabituated animals not used for tourism. Among the habituated animals used for tourism, fecal glucocorticoid metabolite levels were significantly elevated in samples collected the day after tourist visitation (indicative of elevated cortisol production on the previous day during tourist visitation). Fecal glucocorticoid metabolite levels were also lower in the habituated animals compared to their age-matched unhabituated counterparts. We conclude that the habituated animals used for this singular ecotourism project are not chronically stressed, unlike other species/populations with documented permanent alterations in stress responses. Animal temperament, species, the presence of coping/escape mechanisms, social confounders, and variation in amount of tourism may explain differences among previous experiments. Acute alterations in glucocorticoid measures in wildlife exposed to tourism must be interpreted conservatively. While permanently altered stress responses can be detrimental, preliminary results in these wild habituated orangutans suggest that low levels of predictable disturbance can likely result in low physiological impact on these animals.

The geographic distribution of Bornean orang-utans and its overlap with existing land-use categories (protected areas, logging and plantation concessions) is a necessary foundation to prioritize conservation planning. Based on an extensive orang-utan survey dataset and a number of environmental variables, we modelled an orang-utan distribution map. The modelled orang-utan distribution map covers 155,106 km(2) (21% of Borneo's landmass) and reveals four distinct distribution areas. The most important environmental predictors are annual rainfall and land cover. The overlap of the orang-utan distribution with land-use categories reveals that only 22% of the distribution lies in protected areas, but that 29% lies in natural forest concessions. A further 19% and 6% occurs in largely undeveloped oil palm and tree plantation concessions, respectively. The remaining 24% of the orang-utan distribution range occurs outside of protected areas and outside of concessions. An estimated 49% of the orang-utan distribution will be lost if all forest outside of protected areas and logging concessions is lost. To avoid this potential decline plantation development in orang-utan habitats must be halted because it infringes on national laws of species protection. Further growth of the plantation sector should be achieved through increasing yields in existing plantations and expansion of new plantations into areas that have already been deforested. To reach this goal a large scale island-wide land-use masterplan is needed that clarifies which possible land uses and managements are allowed in the landscape and provides new standardized strategic conservation policies. Such a process should make much better use of non-market values of ecosystem services of forests such as water provision, flood control, carbon sequestration, and sources of livelihood for rural communities. Presently land use planning is more driven by vested interests and direct and immediate economic gains, rather than by approaches that take into consideration social equity and environmental sustainability.

BACKGROUND: Southeast Asian deforestation rates are among the world's highest and threaten to drive many forest-dependent species to extinction. Climate change is expected to interact with deforestation to amplify this risk. Here we examine whether regional incentives for sustainable forest management will be effective in improving threatened mammal conservation, in isolation and when combined with global climate change mitigation. METHODOLOGY/PRINCIPAL FINDINGS: Using a long time-series of orangutan nest counts for Sabah (2000-10), Malaysian Borneo, we evaluated the effect of sustainable forest management and climate change scenarios, and their interaction, on orangutan spatial abundance patterns. By linking dynamic land-cover and downscaled global climate model projections, we determine the relative influence of these factors on orangutan spatial abundance and use the resulting statistical models to identify habitat crucial for their long-term conservation. We show that land-cover change the degradation of primary forest had the greatest influence on orangutan population size. Anticipated climate change was predicted to cause reductions in abundance in currently occupied populations due to decreased habitat suitability, but also to promote population growth in western Sabah by increasing the suitability of presently unoccupied regions. CONCLUSIONS/SIGNIFICANCE: We find strong quantitative support for the Sabah government's proposal to implement sustainable forest management in all its forest reserves during the current decade; failure to do so could result in a 40 to 80 per cent regional decline in orangutan abundance by 2100. The Sabah orangutan is just one (albeit iconic) example of a forest-dependent species that stands to benefit from sustainable forest management, which promotes conservation of existing forests.

Human-orangutan conflict and hunting are thought to pose a serious threat to orangutan existence in Kalimantan, the Indonesian part of Borneo. No data existed prior to the present study to substantiate these threats. We investigated the rates, spatial distribution and causes of conflict and hunting through an interview-based survey in the orangutan's range in Kalimantan, Indonesia. Between April 2008 and September 2009, we interviewed 6983 respondents in 687 villages to obtain socio-economic information, assess knowledge of local wildlife in general and orangutan encounters specifically, and to query respondents about their knowledge on orangutan conflicts and killing, and relevant laws. This survey revealed estimated killing rates of between 750 and 1800 animals killed in the last year, and between 1950 and 3100 animals killed per year on average within the lifetime of the survey respondents. These killing rates are higher than previously thought and are high enough to pose a serious threat to the continued existence of orangutans in Kalimantan. Importantly, the study contributes to our understanding of the spatial variation in threats, and the underlying causes of those threats, which can be used to facilitate the development of targeted conservation management.

Species conservation is difficult. Threats to species are typically high and immediate. Effective solutions for counteracting these threats, however, require synthesis of high quality evidence, appropriately targeted activities, typically costly implementation, and rapid re-evaluation and adaptation. Conservation management can be ineffective if there is insufficient understanding of the complex ecological, political, socio-cultural, and economic factors that underlie conservation threats. When information about these factors is incomplete, conservation managers may be unaware of the most urgent threats or unable to envision all consequences of potential management strategies. Conservation research aims to address the gap between what is known and what knowledge is needed for effective conservation. Such research, however, generally addresses a subset of the factors that underlie conservation threats, producing a limited, simplistic, and often biased view of complex, real world situations. A combination of approaches is required to provide the complete picture necessary to engage in effective conservation. Orangutan conservation (Pongo spp.) offers an example: standard conservation assessments employ survey methods that focus on ecological variables, but do not usually address the socio-cultural factors that underlie threats. Here, we evaluate a complementary survey method based on interviews of nearly 7,000 people in 687 villages in Kalimantan, Indonesia. We address areas of potential methodological weakness in such surveys, including sampling and questionnaire design, respondent biases, statistical analyses, and sensitivity of resultant inferences. We show that interview-based surveys can provide cost-effective and statistically robust methods to better understand poorly known populations of species that are relatively easily identified by local people. Such surveys provide reasonably reliable estimates of relative presence and relative encounter rates of such species, as well as quantifying the main factors that threaten them. We recommend more extensive use of carefully designed and implemented interview surveys, in conjunction with more traditional field methods.

Ecological studies of orangutans have almost exclusively focused on populations living in primary or selectively logged rainforest. The response of orangutans to severe habitat degradation remains therefore poorly understood. Most experts assume that viable populations cannot survive outside undisturbed or slightly disturbed forests. This is a concern because nearly 75% of all orangutans live outside protected areas, where degradation of natural forests is likely to occur, or where these are replaced by planted forests. To improve our understanding of orangutan survival in highly altered forest habitats, we conducted population density surveys in two pulp and paper plantation concessions in East Kalimantan, Indonesia. These plantations consist of areas planted with fast-growing exotics intermixed with stands of highly degraded forests and scrublands. Our rapid surveys indicate unexpectedly high orangutan densities in plantation landscapes dominated by Acacia spp., although it remains unclear whether such landscapes can maintain long-term viable populations. These findings indicate the need to better understand how plantation-dominated landscapes can potentially be incorporated into orangutan conservation planning. Although we emphasize that plantations have less value for overall biodiversity conservation than natural forests, they could potentially boost the chances of orangutan survival. Our findings are based on a relatively short study and various methodological issues need to be addressed, but they suggest that orangutans may be more ecologically flexible than previously thought.

BackgroundBornean orangutans (Pongo pygmaeus) currently occur at low densities and seeing a wild one is a rare event. Compared to present low encounter rates of orangutans, it is striking how many orangutan each day historic collectors like Alfred Russel Wallace were able to shoot continuously over weeks or even months. Does that indicate that some 150 years ago encounter rates with orangutans, or their densities, were higher than now?Methodology/principal findingsWe test this hypothesis by quantifying encounter rates obtained from hunting accounts, museum collections, and recent field studies, and analysing whether there is a declining trend over time. Logistic regression analyses of our data support such a decline on Borneo between the mid-19th century and the present. Even when controlled for variation in the size of survey and hunting teams and the durations of expeditions, mean daily encounter rates appear to have declined about 6-fold in areas with little or no forest disturbance.Conclusions/significanceThis finding has potential consequences for our understanding of orangutans, because it suggests that Bornean orangutans once occurred at higher densities. We explore potential explanations-habitat loss and degradation, hunting, and disease-and conclude that hunting fits the observed patterns best. This suggests that hunting has been underestimated as a key causal factor of orangutan density and distribution, and that species population declines have been more severe than previously estimated based on habitat loss only. Our findings may require us to rethink the biology of orangutans, with much of our ecological understanding possibly being based on field studies of animals living at lower densities than they did historically. Our approach of quantifying species encounter rates from historic data demonstrates that this method can yield valuable information about the ecology and population density of species in the past, providing new insight into species' conservation needs.

BackgroundToday the majority of wild great ape populations are found outside of the network of protected areas in both Africa and Asia, therefore determining if these populations are able to survive in forests that are exploited for timber or other extractive uses and how this is managed, is paramount for their conservation.Methodology/principal findingsIn 2007, the "Kinabatangan Orang-utan Conservation Project" (KOCP) conducted aerial and ground surveys of orang-utan (Pongo pygmaeus morio) nests in the commercial forest reserves of Ulu Segama Malua (USM) in eastern Sabah, Malaysian Borneo. Compared with previous estimates obtained in 2002, our recent data clearly shows that orang-utan populations can be maintained in forests that have been lightly and sustainably logged. However, forests that are heavily logged or subjected to fast, successive coupes that follow conventional extraction methods, exhibit a decline in orang-utan numbers which will eventually result in localized extinction (the rapid extraction of more than 100 m(3) ha(-1) of timber led to the crash of one of the surveyed sub-populations). Nest distribution in the forests of USM indicates that orang-utans leave areas undergoing active disturbance and take momentarily refuge in surrounding forests that are free of human activity, even if these forests are located above 500 m asl. Displaced individuals will then recolonize the old-logged areas after a period of time, depending on availability of food sources in the regenerating areas.Conclusion/significanceThese results indicate that diligent planning prior to timber extraction and the implementation of reduced-impact logging practices can potentially be compatible with great ape conservation.

Great ape populations are undergoing a dramatic decline, which is predicted to result in their extinction in the wild from entire regions in the near future. Recent findings have particularly focused on African apes, and have implicated multiple factors contributing to this decline, such as deforestation, hunting, and disease. Less well-publicised, but equally dramatic, has been the decline in orang-utans, whose distribution is limited to parts of Sumatra and Borneo. Using the largest-ever genetic sample from wild orang-utan populations, we show strong evidence for a recent demographic collapse in North Eastern Borneo and demonstrate that this signature is independent of the mutation and demographic models used. This is the first demonstration that genetic data can detect and quantify the effect of recent, human-induced deforestation and habitat fragmentation on an endangered species. Because current demographic collapses are usually confounded by ancient events, this suggests a much more dramatic decline than demographic data alone and emphasises the need for major conservation efforts.

Great apes are threatened with extinction, but precise information about the distribution and size of most populations is currently lacking. We conducted orangutan nest counts in the Malaysian state of Sabah (North Borneo), using a combination of ground and helicopter surveys, and provided a way to estimate the current distribution and size of the populations living throughout the entire state. We show that the number of nests detected during aerial surveys is directly related to the estimated true animal density and that a helicopter is an efficient tool to provide robust estimates of orangutan numbers. Our results reveal that with a total estimated population size of about 11,000 individuals, Sabah is one of the main strongholds for orangutans in North Borneo. More than 60% of orangutans living in the state occur outside protected areas, in production forests that have been through several rounds of logging extraction and are still exploited for timber. The role of exploited forests clearly merits further investigation for orangutan conservation in Sabah.