Impacts of fire and prospects for recovery in a tropical peat forest ecosystem.

Uncontrolled fires place considerable burdens on forest ecosystems, compromising our ability to meet conservation and restoration goals. A poor understanding of the impacts of fire on ecosystems and their biodiversity exacerbates this challenge, particularly in tropical regions where few studies have applied consistent analytical techniques to examine a broad range of ecological impacts over multiyear time frames. We compiled 16 y of data on ecosystem properties (17 variables) and biodiversity (21 variables) from a tropical peatland in Indonesia to assess fire impacts and infer the potential for recovery. Burned forest experienced altered structural and microclimatic conditions, resulting in a proliferation of nonforest vegetation and erosion of forest ecosystem properties and biodiversity. Compared to unburned forest, habitat structure, tree density, and canopy cover deteriorated by 58 to 98%, while declines in species diversity and abundance were most pronounced for trees, damselflies, and butterflies, particularly for forest specialist species. Tracking ecosystem property and biodiversity datasets over time revealed most to be sensitive to recurrent high-intensity fires within the wider landscape. These megafires immediately compromised water quality and tree reproductive phenology, crashing commercially valuable fish populations within 3 mo and driving a gradual decline in threatened vertebrates over 9 mo. Burned forest remained structurally compromised long after a burn event, but vegetation showed some signs of recovery over a 12-y period. Our findings demonstrate that, if left uncontrolled, fire may be a pervasive threat to the ecological functioning of tropical forests, underscoring the importance of fire prevention and long-term restoration efforts, as exemplified in Indonesia.

Suspected Sunda clouded leopard (Neofelis diardi) predation attempts on two reintroduced Bornean orangutans (Pongo pygmaeus wurmbii) in Bukit Batikap Protection Forest, Central Kalimantan, Indonesia.

In February 2017 and August 2018, respectively, two Bornean orangutans (Pongo pygmaeus wurmbii) reintroduced into the Bukit Batikap Protection Forest in Central Kalimantan were found in weakened physical condition and with deep puncture wounds. The first individual was a sub-adult male, and the second an adult female whose 6- to 8-week-old infant was missing. Both individuals were rescued and transported back to the field base camp for treatment. Experienced veterinarians treating the injuries reported that the type of wounds appeared consistent with those expected from an attack by a large felid. The Sunda clouded leopard (Neofelis diardi) is the largest felid known to inhabit Bukit Batikap Protection Forest, and we suspect that these cases were unsuccessful predatory attacks by this species. Given the severity of his condition when found, the male orangutan would probably have died without medical intervention; however, both orangutans fully recovered following intensive treatment and were successfully returned to the forest. Predation attempts on orangutans are infrequently reported, thus our observations add to the body of knowledge about possible predation by clouded leopards on reintroduced, rehabilitant orangutans.

Implanted Radio Telemetry in Orangutan Reintroduction and Post-release Monitoring and its Application in Other Ape Species.

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.

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.

Nutritional Differences between Two Orangutan Habitats: Implications for Population Density.

Bottom-up regulatory factors have been proposed to exert a strong influence on mammalian population density. Studies relating habitat quality to population density have typically made comparisons among distant species or communities without considering variation in food quality among localities. We compared dietary nutritional quality of two Bornean orangutan populations with differing population densities in peatland habitats, Tuanan and Sabangau, separated by 63 km. We hypothesized that because Tuanan is alluvial, the plant species included in the orangutan diet would be of higher nutritional quality compared to Sabangau, resulting in higher daily caloric intake in Tuanan. We also predicted that forest productivity would be greater in Tuanan compared to Sabangau. In support of these hypotheses, the overall quality of the diet and the quality of matched dietary items were higher in Tuanan, resulting in higher daily caloric intake compared to Sabangau. These differences in dietary nutritional quality may provide insights into why orangutan population density is almost two times greater in Tuanan compared to Sabangau, in agreement with a potentially important influence of diet quality on primate population density.

Fine-scale habitat use by orang-utans in a disturbed peat swamp forest, central Kalimantan, and implications for conservation management.

This study was conducted to see how orang-utans (Pongo pygmaeus wurmbii) were coping with fine-scale habitat disturbance in a selectively logged peat swamp forest in Central Kalimantan, Borneo. Seven habitat classes were defined, and orang-utans were found to use all of these, but were selective in their preference for certain classes over others. Overall, the tall forest classes (≥20 m) were preferred. They were preferred for feeding, irrespective of canopy connectivity, whereas classes with a connected canopy (canopy cover ≥75%), irrespective of canopy height, were preferred for resting and nesting, suggesting that tall trees are preferred for feeding and connected canopy for security and protection. The smaller forest classes (≤10 m high) were least preferred and were used mainly for travelling from patch to patch. Thus, selective logging is demonstrated here to be compatible with orang-utan survival as long as large food trees and patches of primary forest remain. Logged forest, therefore, should not automatically be designated as 'degraded'. These findings have important implications for forest management, forest classification and the designation of protected areas for orang-utan conservation.

Co-occurrence patterns of Bornean vertebrates suggest competitive exclusion is strongest among distantly related species.

Assessing the importance of deterministic processes in structuring ecological communities is a central focus of community ecology. Typically, community ecologists study a single taxonomic group, which precludes detection of potentially important biotic interactions between distantly related species, and inherently assumes competition is strongest between closely related species. We examined distribution patterns of vertebrate species across the island of Borneo in Southeast Asia to assess the extent to which inter-specific competition may have shaped ecological communities on the island and whether the intensity of inter-specific competition in present-day communities varies as a function of evolutionary relatedness. We investigated the relative extent of competition within and between species of primates, birds, bats and squirrels using species presence-absence and attribute data compiled for 21 forested sites across Borneo. We calculated for each species pair the checkerboard unit value (CU), a statistic that is often interpreted as indicating the importance of interspecific competition. The percentage of species pairs with significant CUs was lowest in within-taxon comparisons. Moreover, for invertebrate-eating species the percentage of significantly checkerboarded species pairs was highest in comparisons between primates and other taxa, particularly birds and squirrels. Our results are consistent with the interpretation that competitive interactions between distantly related species may have shaped the distribution of species and thus the composition of Bornean vertebrate communities. This research highlights the importance of taking into account the broad mammalian and avian communities in which species occur for understanding the factors that structure biodiversity.

Feeding ecology of red langurs in Sabangau tropical peat-swamp forest, Indonesian Borneo: extreme granivory in a non-masting forest.

Southeast Asia's lowland dipterocarp forests experience supra-annual "mast" fruiting and flowering events, in which the majority of trees reproduce simultaneously at irregular intervals, with extensive intervening periods of very low primate food availability. This scarcity of food results in a negative energy balance and a reliance on "fallback foods" in some primate species. By contrast, ombrogenous tropical peat-swamp forests are non-masting, and show lower variability of food availability. We sought to test the influence of fruit availability on primate diet and preference in peat-swamp habitats and assess whether it differs from masting forests. We collected behavioral-dependent feeding data on three adult females in a group of red langurs (Presbytis rubicunda: Colobinae) between January and December 2011 in Sabangau tropical peat-swamp forest, Central Kalimantan, Indonesia, as colobine monkeys are adaptated for folivory, and are therefore generally considered less reliant on temporally variable fruits than monogastric primates. We documented the highest level of granivory recorded to date in colobine monkeys: mean annual diet comprised 76.4% seeds and 7.3% other fruit parts; 7.7% young and 2.5% mature leaves; 2.8% flowers; 2.6% piths, and <1% on other minor food items. Fruit availability was fairly constant throughout the year and fruit parts were consumed at consistently higher levels than expected based on availability, confirming that fruit is preferred. Leaves and flowers were consumed consistently less than expected and thus are not preferred. There were no significant correlations between preferred food availability and consumption of potential fallback foods, suggesting that reliance on fallback foods did not occur in Sabangau during the study period. Furthermore, consumption of fruit was not significantly correlated with its availability. Our findings suggest that the relatively constant availability of fruit in this habitat affords P. rubicunda regular access to a nutritionally superior food, and that reliance on fallback foods is therefore not required.

Interspecific interactions between primates, birds, bats, and squirrels may affect community composition on Borneo.

For several decades, primatologists have been interested in understanding how sympatric primate species are able to coexist. Most of our understanding of primate community ecology derives from the assumption that these animals interact predominantly with other primates. In this study, we investigate to what extent multiple community assembly hypotheses consistent with this assumption are supported when tested with communities of primates in isolation versus with communities of primates, birds, bats, and squirrels together. We focus on vertebrate communities on the island of Borneo, where we examine the determinants of presence or absence of species, and how these communities are structured. We test for checkerboard distributions, guild proportionality, and Fox's assembly rule for favored states, and predict that statistical signals reflecting interactions between ecologically similar species will be stronger when nonprimate taxa are included in analyses. We found strong support for checkerboard distributions in several communities, particularly when taxonomic groups were combined, and after controlling for habitat effects. We found evidence of guild proportionality in some communities, but did not find significant support for Fox's assembly rule in any of the communities examined. These results demonstrate the presence of vertebrate community structure that is ecologically determined rather than randomly generated, which is a finding consistent with the interpretation that interactions within and between these taxonomic groups may have shaped species composition in these communities. This research highlights the importance of considering the broader vertebrate communities with which primates co-occur, and so we urge primatologists to explicitly consider nonprimate taxa in the study of primate ecology.

Understanding the impacts of land-use policies on a threatened species: is there a future for the Bornean orang-utan?

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.

Biodiversity Conservation in the REDD.

Deforestation and forest degradation in the tropics is a major source of global greenhouse gas (GHG) emissions. The tropics also harbour more than half the world's threatened species, raising the possibility that reducing GHG emissions by curtailing tropical deforestation could provide substantial co-benefits for biodiversity conservation. Here we explore the potential for such co-benefits in Indonesia, a leading source of GHG emissions from land cover and land use change, and among the most species-rich countries in the world. We show that focal ecosystems for interventions to reduce emissions from deforestation and forest degradation in Indonesia do not coincide with areas supporting the most species-rich communities or highest concentration of threatened species. We argue that inherent trade-offs among ecosystems in emission reduction potential, opportunity cost of foregone development and biodiversity values will require a regulatory framework to balance emission reduction interventions with biodiversity co-benefit targets. We discuss how such a regulatory framework might function, and caution that pursuing emission reduction strategies without such a framework may undermine, not enhance, long-term prospects for biodiversity conservation in the tropics.

Survey of Hylobates agilis albibarbis in a logged peat-swamp forest: Sabangau catchment, Central Kalimantan.

Few data are available on gibbon populations in peat-swamp forest. In order to assess the importance of this habitat for gibbon conservation, a population of Hylobates agilis albibarbis was surveyed in the Sabangau peat-swamp forest, Central Kalimantan, Indonesia. This is an area of about 5,500 km(2) of selectively logged peat-swamp forest, which was formally gazetted as a national park during 2005. The study was conducted during June and July 2004 using auditory sampling methods. Five sample areas were selected and each was surveyed for four consecutive days by three teams of researchers at designated listening posts. Researchers recorded compass bearings of, and estimated distances to, singing groups. Nineteen groups were located. Population density is estimated to be 2.16 (+/-0.46) groups/km(2). Sightings occurring either at the listening posts or that were obtained by tracking in on calling groups yielded a mean group size of 3.4 individuals, hence individual gibbon density is estimated to be 7.4 (+/-1.59) individuals/km(2). The density estimates fall at the mid-range of those calculated for other gibbon populations, thus suggesting that peat-swamp forest is an important habitat for gibbon conservation in Borneo. A tentative extrapolation of results suggests a potential gibbon population size of 19,000 individuals within the mixed-swamp forest habitat sub-type in the Sabangau. This represents one of the largest remaining continuous populations of Bornean agile gibbons. The designation of the Sabangau forest as a national park will hopefully address the problem of illegal logging and hunting in the region. Further studies should note any difference in gibbon density post protection.