Abstract


Tropical forests and peatlands provide important ecological, climate and socio‐economic benefits from the local to the global scale. However, these ecosystems and their associated benefits are threatened by anthropogenic activities, including agricultural conversion, timber harvesting, peatland drainage and associated fire. Here, we identify key challenges, and provide potential solutions and future directions to meet forest and peatland conservation and restoration goals in Indonesia, with a particular focus on Kalimantan.

Through a round‐table, dual‐language workshop discussion and literature evaluation, we recognized 59 political, economic, legal, social, logistical and research challenges, for which five key underlying factors were identified. These challenges relate to the 3Rs adopted by the Indonesian Peatland Restoration Agency (Rewetting, Revegetation and Revitalization), plus a fourth R that we suggest is essential to incorporate into (peatland) conservation planning: Reducing Fires.

Our analysis suggests that (a) all challenges have potential for impact on activities under all 4Rs, and many are inter‐dependent and mutually reinforcing, implying that narrowly focused solutions are likely to carry a higher risk of failure; (b) addressing challenges relating to Rewetting and Reducing Fire is critical for achieving goals in all 4Rs, as is considering the local socio‐political situation and acquiring local government and community support; and (c) the suite of challenges faced, and thus conservation interventions required to address these, will be unique to each project, depending on its goals and prevailing local environmental, social and political conditions.

With this in mind, we propose an eight‐step adaptive management framework, which could support projects in both Indonesia and other tropical areas to identify and overcome their specific conservation and restoration challenges.

A free Plain Language Summary can be found within the Supporting Information of this article.

Southeast Asia's tropical peat-swamp forests (TPSF) are globally important for carbon storage and biodiversity conservation, but are at risk from multiple threats and urgently require improved management. Ants are often used as ecological indicators in monitoring programmes to guide adaptive management, but data on TPSF ants are scarce. We conducted a twelve-month study on ants in the Sabangau TPSF in Indonesian Borneo using baited traps, to compare community composition across three disturbance categories (forest gaps, forest edge and relatively undisturbed interior forest) and between dry and wet season. The three disturbance categories supported distinct ant communities across seasons. Differences in canopy cover likely underlie these changes in ant community composition. Surveying was more effective in the dry season, because ant capture rates were higher and more indicator taxa were identified than in the wet season, but overall ant community composition did not differ significantly between seasons. These findings suggest a potentially useful role of ants as ecological indicators in TPSF. Further surveys should be conducted in Sabangau and other TPSFs to test the transferability of our findings.

Fire disturbance in many tropical forests, including peat swamps, has become more frequent and extensive in recent decades. These fires compromise a variety of ecosystem services, among which mitigating global climate change through carbon storage is particularly important for peat swamps. Indonesia holds the largest amount of tropical peat carbon globally, and mean annual CO2 emissions from decomposition of deforested and drained peatlands and associated fires in Southeast Asia have been estimated at ~2000 Mt y-1. A key component to understanding and therefore managing fire in the region is identifying the land use/land cover classes associated with fire ignitions. We assess the oft-asserted claim that escaped fires from oil palm concessions and smallholder farms near settlements are the primary sources of fire in a peat-swamp forest area in Central Kalimantan, Indonesia, equivalent to around a third of Kalimantan's total peat area. We use the MODIS Active Fire product from 2000 to 2010 to evaluate the fire origin and spread on the land use/land cover classes of legal, industrial oil palm concessions (the only type of legal concession in the study area), non-forest, and forest, as well as in relation to settlement proximity. We find that most fires (68-71%) originate in non-forest, compared to oil palm concessions (17%-19%), and relatively few (6-9%) are within 5 km of settlements. Moreover, most fires started within oil palm concessions and in close proximity to settlements stay within those boundaries (90% and 88%, respectively), and fires that do escape constitute only a small proportion of all fires on the landscape (2% and 1%, respectively). Similarly, a small proportion of fire detections in forest originate from oil palm concessions (2%) and within close proximity to settlements (2%). However, fire ignition density in oil palm (0.055 ignitions km-2) is comparable to that in non-forest (0.060 km-2 ignitions km-2), which is approximately ten times that in forest (0.006 ignitions km-2). Ignition density within 5 km of settlements is the highest at 0.125 ignitions km-2. Furthermore, increased anthropogenic activity in close proximity to oil palm concessions and settlements produces a detectable pattern of fire activity. The number of ignitions decreases exponentially with distance from concessions; the number of ignitions initially increases with distance from settlements, and, around from 7.2 km, then decreases with distance from settlements. These results refute the claim that most fires originate in oil palm concessions, and that fires escaping from oil palm concessions and settlements constitute a major proportion of fires in this study region. However, there is a potential for these land use types to contribute substantially to the fire landscape if their area expands. Effective fire management in this area should therefore target not just oil palm concessions, but also non-forested, degraded areas where ignitions and fires escaping into forest are most likely to occur.

Researchers have suggested that fallback foods (FBFs) shape primate food processing adaptations, whereas preferred foods drive harvesting adaptations, and that the dietary importance of FBFs is central in determining the expression of a variety of traits. We examine these hypotheses in extant apes. First, we compare the nature and dietary importance of FBFs used by each taxon. FBF importance appears greatest in gorillas, followed by chimpanzees and siamangs, and least in orangutans and gibbons (bonobos are difficult to place). Next, we compare 20 traits among taxa to assess whether the relative expression of traits expected for consumption of FBFs matches their observed dietary importance. Trait manifestation generally conforms to predictions based on dietary importance of FBFs. However, some departures from predictions exist, particularly for orang-utans, which express relatively more food harvesting and processing traits predicted for consuming large amounts of FBFs than expected based on observed dietary importance. This is probably due to the chemical, mechanical, and phenological properties of the apes' main FBFs, in particular high importance of figs for chimpanzees and hylobatids, compared to use of bark and leaves-plus figs in at least some Sumatran populations-by orang-utans. This may have permitted more specialized harvesting adaptations in chimpanzees and hylobatids, and required enhanced processing adaptations in orang-utans. Possible intercontinental differences in the availability and quality of preferred and FBFs may also be important. Our analysis supports previous hypotheses suggesting a critical influence of the dietary importance and quality of FBFs on ape ecology and, consequently, evolution. © 2011 the Author(s).

Data on energy intake and the effects of fluctuations in fruit availability on energy intake for African apes, and orangutans in mast-fruiting habitats, indicate that orangutans may face greater energetic challenges than do their African counterparts. Comparable data on orangutans in nonmasting forests, which experience lower fluctuations in fruit availability, have been lacking, however, complicating interpretations. We conducted a 46-mo study of orangutan energetics in the nonmasting Sabangau peat-swamp forest, Indonesian Borneo. Sabangau orangutans experienced periods of negative energy balance apparently even longer than in mast-fruiting habitats, as indicated by comparisons of observed energy intake with theoretical requirements and analysis of urinary ketones. Daily energy intake was positively related to fruit availability in flanged males, but not in adult females or unflanged males. This may represent different foraging strategies between age-sex classes and suggests that fruit availability is not always an accurate indicator of ape energy intake/balance. Urinary ketone levels were not generally related to fruit availability, daily energy intake, day range, or party size. This is probably due to low energy intake, and consequently high ketone production, throughout much of the study period. Comparisons with published results on African apes support the hypothesis that orangutans are unique among hominoids in regularly experiencing prolonged periods of negative energy balance. This has important effects on orangutan behavior and socioecology, and has likely been a key factor driving the evolutionary divergence of orangutans and African apes. © 2010 Springer Science+Business Media, LLC.

Reforestation is promoted to address the dual global climate and biodiversity crises. This is particularly relevant for carbon-rich, biodiverse tropical peatlands, for which active reforestation typically involves two post-germination stages: nursery rearing of seedlings, then outplanting. Yet, linkages between these stages and cumulative seedling performance are rarely quantified during tropical peatland reforestation. By monitoring tree seedling survival and growth, we investigate factors influencing seedling performance (species identity, seedling source, treatments, and climate), whether nursery performance predicts outplanting performance, and calculate cumulative survival (nursery plus outplanting) in Sebangau National Park, Indonesian Borneo. Standardized survival at 2 years was higher in the nursery (mean 67% across 40 species) than outplanting (44% across 24 species). For nursery and outplanting, species identity was the main source of variation in survival and height growth. Seedling source, treatments, site condition, and precipitation had no significant impact on survival but did influence growth in some cases. Nursery survival did not predict outplanting survival, but nursery height did predict outplanting height. Across species, around a quarter of seedlings survived from nursery to outplanting over 4 years. Cumulative survival represents a more realistic basis for assessing the genetic and other resource costs of tropical peatland reforestation. Our two-phase approach identified outplanting as the greater bottleneck to cumulative seedling survivability. We argue that the nursery stage may be used to harden seedlings for degraded peatland conditions by selecting more relevant treatments (e.g. flooding) and screening for resilience to common disturbances (e.g. fire) to enhance outplanted, and thus cumulative, seedling survival.

Assessing a taxon’s response to change in environmental variables is fundamental knowledge to understanding trends in species diversity, abundance, and distribution patterns. This is particularly needed on Borneo, where knowledge on Odonata populations in different habitats is poor. To address this gap, we present the first study investigating the relationship between morphology and species distribution of Odonata communities in a heath (kerangas)-dominated mixed-mosaic-lowland forest in southern Borneo. We sampled 250-m line transects in three habitat types: mixed peatcswamp, kerangas, and low-pole peatcswamp, with weekly surveys from December 2019 to February 2020. A total of 309 individuals were detected from 25 species. Anisoptera and Zygoptera diversity was the highest in mixed peatcswamp and lowest in low pole, while abundance was the highest in low pole and lowest in kerangas; with kerangas notably harboring a very small sample size. Odonata community assemblages differed most between mixed peat swamp and low pole. Morphological data were compared between suborders and habitats. Anisoptera showed significantly larger thoraces, hindwings, and hindwing-to-body ratio than Zygoptera. Anisoptera in low pole were significantly smaller in body, thorax, and hindwing compared to both kerangas and mixed peat swamp. Anisoptera showed a strong association with pools and Zygoptera with flowing water. Heterogeneity, habitat characteristics, presence of specialists, body size, and the interaction between species’ morphological traits and habitat characteristics likely explained the trends observed.

Peatlands are highly dynamic systems, able to accumulate carbon over millennia under natural conditions, but susceptible to rapid subsidence and carbon loss when drained. Short-term, seasonal and long-term peat surface elevation changes are closely linked to key peatland attributes such as water table depth (WTD) and carbon balance, and may be measured remotely using satellite radar and LiDAR methods. However, field measurements of peat elevation change are spatially and temporally sparse, reliant on low-resolution manual subsidence pole measurements, or expensive sensor systems. Here we describe a novel, simple and low-cost image-based method for measuring peat surface motion and WTD using commercially available time-lapse cameras and image processing methods. Based on almost two years’ deployment of peat cameras across contrasting forested, burned, agricultural and oil palm plantation sites in Central Kalimantan, Indonesia, we show that the method can capture extremely high resolution (sub-mm) and high-frequency (sub-daily) changes in peat surface elevation over extended periods and under challenging environmental conditions. WTD measurements were of similar quality to commercially available pressure transducers. Results reveal dynamic peat elevation response to individual rain events, consistent with variations in WTD. Over the course of the relatively severe 2019 dry season, cameras in deep-drained peatlands recorded maximum peat shrinkage of over 8 cm, followed by partial rebound, leading to net annual subsidence of up to 5 cm. Sites with higher water tables, and where borehole irrigation was used to maintain soil moisture, had lower subsidence, suggesting potential to reduce subsidence through altered land-management. Given the established link between subsidence and CO2 emissions, these results have direct implications for the management of peatlands to reduce high current greenhouse gas (GHG) emissions. Camera-based sensors provide a simple, low-cost alternative to commercial elevation, WTD and GHG flux monitoring systems, suitable for deployment at scale, and in areas where existing approaches are impractical or unaffordable. If ground-based observations of peat motion can be linked to measured GHG fluxes and with satellite-based monitoring tools, this approach offers the potential for a large-scale peatland monitoring tool, suitable for identifying areas of active carbon loss, targeting climate change mitigation interventions, and evaluating intervention outcomes.

Abstract


Interactions between conservation and the social sciences are frequently characterized by either critique (of conservation by social scientists) or co‐optation (of social scientific methods and insights by conservationists).

This article seeks to push beyond these two dominant positions by exploring how conservationists and social scientists can engage in mutually transformative dialogue. Jointly authored by conservation scientists and social scientists, it uses the global nexus of orangutan conservation as a lens onto current challenges and possibilities facing the conservation–social science relationship.

We begin with a cross‐disciplinary overview of recent developments in orangutan conservation—particularly those concerned with its social, political and other human dimensions.

The article then undertakes a synthetic analysis of key challenges in orangutan conservation—working across difference, juggling scales and contexts and dealing with politics and political economy—and links them to analogous concerns in the conservation–social science relationship.

Finally, we identify some ways by which orangutan conservation specifically, and the conservation–social science relationship more generally, can move forward: through careful use of proxies as bridging devices, through the creation of new, shared spaces, and through a willingness to destabilize and overhaul status quos. This demands an open‐ended, unavoidably political commitment to critical reflexivity and self‐transformation on the part of both conservationists and social scientists.

A free Plain Language Summary can be found within the Supporting Information of this article.

Conservation projects are likely to fail if plans to preserve important wildlife habitats and species are not co-developed between conservation organisations and local communities to reflect the needs and diverse values of the latter. Tropical peatland conservation represents a case in point: local community livelihoods have only recently come into focus, particularly within academic literature. Instead, many previous studies emphasise the need to conserve intact peat swamp forests for their carbon storage, as a habitat for flagship species such as the orangutan, and to provide fire-free landscapes. Here, we explore the socio-environmental issues being faced in the peatland landscapes of Central Kalimantan, Indonesia. This includes the loss of peat-swamp forest, decreases in peatland fish populations and related socio-cultural challenges such as potential loss of fishing livelihoods along with historic and continued experiences of marginalisation of indigenous communities. To find solutions to these complex and interrelated problems, an interdisciplinary approach which focuses on interdependencies and includes multiple worldviews is required. We propose an approach which deploys both Ethan Miller's use of livelihoods (incl. Miller, 2019) and biocultural approaches to conservation to analyse human-nonhuman relationships, with a focus on fish and fishing livelihoods. We draw on data from in-depth social and ecological research in two village communities in Central Kalimantan, and in so doing illustrate how fish conservation has the potential to support important biocultural and livelihood relationships between human and nonhuman communities in peatland areas. Our findings lend support to previous calls for biocultural approaches to conservation in other socio-ecological contexts, and lead us to conclude that tropical peatland conservation initiatives that integrate such approaches will result in improved outcomes for peatlands, forests, biodiversity and people. These findings will be relevant to other tropical peatland areas with high dependence on fishing as a source of livelihood, such as the peatlands of the Amazon and Congo basins.

The COVID-19 pandemic has caused global disruption, with the emergence of this and other pandemics having been linked to habitat encroachment and/or wildlife exploitation. High impacts of COVID-19 are apparent in some countries with large tropical peatland areas, some of which are relatively poorly resourced to tackle disease pandemics. Despite this, no previous investigation has considered tropical peatlands in the context of emerging infectious diseases (EIDs). Here, we review: (i) the potential for future EIDs arising from tropical peatlands; (ii) potential threats to tropical peatland conservation and local communities from COVID-19; and (iii) potential steps to help mitigate these risks. We find that high biodiversity in tropical peat-swamp forests, including presence of many potential vertebrate and invertebrate vectors, combined, in places, with high levels of habitat disruption and wildlife harvesting represent suitable conditions for potential zoonotic EID (re-)emergence. Although impossible to predict precisely, we identify numerous potential threats to tropical peatland conservation and local communities from the COVID-19 pandemic. This includes impacts on public health, with the potential for haze pollution from peatland fires to increase COVID-19 susceptibility a noted concern; and on local economies, livelihoods and food security, where impacts will likely be greater in remote communities with limited/no medical facilities that depend heavily on external trade. Research, training, education, conservation and restoration activities are also being affected, particularly those involving physical groupings and international travel, some of which may result in increased habitat encroachment, wildlife harvesting or fire, and may therefore precipitate longer-term negative impacts, including those relating to disease pandemics. We conclude that sustainable management of tropical peatlands and their wildlife is important for mitigating impacts of the COVID-19 pandemic, and reducing the potential for future zoonotic EID emergence and severity, thus strengthening arguments for their conservation and restoration. To support this, we list seven specific recommendations relating to sustainable management of tropical peatlands in the context of COVID-19/disease pandemics, plus mitigating the current impacts of COVID-19 and reducing potential future zoonotic EID risk in these localities. Our discussion and many of the issues raised should also be relevant for non-tropical peatland areas and in relation to other (pandemic-related) sudden socio-economic shocks that may occur in future.

Underground peat fires are a major hazard to health and livelihoods in Indonesia, and are a major contributor to carbon emissions globally. Being subterranean, these fires can be difficult to detect and track, especially during periods of thick haze and in areas with limited accessibility. Thermal infrared detectors mounted on drones present a potential solution to detecting and managing underground fires, as they allow large areas to be surveyed quickly from above and can detect the heat transferred to the surface above a fire. We present a pilot study in which we show that underground peat fires can indeed be detected in this way. We also show that a simple temperature thresholding algorithm can be used to automatically detect them. We investigate how different thermal cameras and drone flying strategies may be used to reliably detect underground fires and survey fire-prone areas. We conclude that thermal equipped drones are potentially a very powerful tool for surveying for fires and firefighting. However, more investigation is still needed into their use in real-life fire detection and firefighting scenarios.

Abstract


Tropical forests and peatlands provide important ecological, climate and socio‐economic benefits from the local to the global scale. However, these ecosystems and their associated benefits are threatened by anthropogenic activities, including agricultural conversion, timber harvesting, peatland drainage and associated fire. Here, we identify key challenges, and provide potential solutions and future directions to meet forest and peatland conservation and restoration goals in Indonesia, with a particular focus on Kalimantan.

Through a round‐table, dual‐language workshop discussion and literature evaluation, we recognized 59 political, economic, legal, social, logistical and research challenges, for which five key underlying factors were identified. These challenges relate to the 3Rs adopted by the Indonesian Peatland Restoration Agency (Rewetting, Revegetation and Revitalization), plus a fourth R that we suggest is essential to incorporate into (peatland) conservation planning: Reducing Fires.

Our analysis suggests that (a) all challenges have potential for impact on activities under all 4Rs, and many are inter‐dependent and mutually reinforcing, implying that narrowly focused solutions are likely to carry a higher risk of failure; (b) addressing challenges relating to Rewetting and Reducing Fire is critical for achieving goals in all 4Rs, as is considering the local socio‐political situation and acquiring local government and community support; and (c) the suite of challenges faced, and thus conservation interventions required to address these, will be unique to each project, depending on its goals and prevailing local environmental, social and political conditions.

With this in mind, we propose an eight‐step adaptive management framework, which could support projects in both Indonesia and other tropical areas to identify and overcome their specific conservation and restoration challenges.

A free Plain Language Summary can be found within the Supporting Information of this article.

The authors apologise for the fact that in the original published version of this article the name of the fourth author was misspelled. The online article has been corrected.

Southeast Asia's tropical peat-swamp forests (TPSF) are globally important for carbon storage and biodiversity conservation, but are at risk from multiple threats and urgently require improved management. Ants are often used as ecological indicators in monitoring programmes to guide adaptive management, but data on TPSF ants are scarce. We conducted a twelve-month study on ants in the Sabangau TPSF in Indonesian Borneo using baited traps, to compare community composition across three disturbance categories (forest gaps, forest edge and relatively undisturbed interior forest) and between dry and wet season. The three disturbance categories supported distinct ant communities across seasons. Differences in canopy cover likely underlie these changes in ant community composition. Surveying was more effective in the dry season, because ant capture rates were higher and more indicator taxa were identified than in the wet season, but overall ant community composition did not differ significantly between seasons. These findings suggest a potentially useful role of ants as ecological indicators in TPSF. Further surveys should be conducted in Sabangau and other TPSFs to test the transferability of our findings.

Tropical tree species distributions are determined by a wide range of biotic and abiotic factors, including topography and hydrology. Tropical peat swamp forests (TPSFs) are characterized in part by small-scale variations in topography (‘hummocks’ and ‘hollows’) that create distinct microhabitats and thus may contribute to niche diversification among TPSF tree species. Using tree elevations calibrated to daily peat water levels collected using a data logger and a permutation test, we evaluated topographical microhabitat preferences for 21 tree species in a relatively undisturbed TPSF in Central Kalimantan, Indonesia, to determine whether these species show preferential association with hummocks or hollows and to quantify the prevalence of microhabitat specialization among them. Only one species, Tetractomia tetrandrum, emerged as a hollow specialist, with no hummock specialists among the species tested. The remaining 20 species, including Psydrax dicoccos, which had the lowest mean observed elevation, and Maasia hypoleuca, which had the highest mean observed elevation, showed no clear microtopographic preference. This suggests that many TPSF species may be resilient to the natural hydrologic variations that occur in relatively intact peat swamp forests. Such studies of microtopographic preferences of tree species in TPSF and other wetland forest ecosystems can inform selection of tree species for reforestation projects, and potentially also provide information on how future climate change may impact these habitats and their resident tree species.

Tropical peat swamp forests provide important ecosystem services, ranging from carbon storage and fire prevention to fish provision. In the Sebangau catchment of Central Kalimantan, Indonesia, we completed the first detailed spatial and temporal assessments of local fish biodiversity in peat swamp forest and blackwater river habitats. Monthly environmental and fish data were collected over a 15-month period in both riverine and forest habitats. This resulted in a species list of 55 species from 16 different families. Species richness in the river was almost 1.5 times higher than in the forest, probably due to the sampling methods and trap selectivity. Average monthly river fish catches were negatively correlated with average monthly river depth. River fish surveys were conducted pre- and post- fire in 2015, with results showing increased river acidity and reduced fish catches post-fire. The fish and environmental data presented form a baseline for future monitoring projects and highlight a previously overlooked potential impact of fire on local biodiversity in Indonesia, namely that fire is likely to have negative impacts on the sizes of fish populations and catches. There are direct implications for human communities that depend on fishing for their livelihoods. Because peatlands and their rivers face continued human disturbance and degradation, assessments of fish biodiversity and water quality are of high priority.

Fire disturbance in many tropical forests, including peat swamps, has become more frequent and extensive in recent decades. These fires compromise a variety of ecosystem services, among which mitigating global climate change through carbon storage is particularly important for peat swamps. Indonesia holds the largest amount of tropical peat carbon globally, and mean annual CO2 emissions from decomposition of deforested and drained peatlands and associated fires in Southeast Asia have been estimated at ~2000 Mt y-1. A key component to understanding and therefore managing fire in the region is identifying the land use/land cover classes associated with fire ignitions. We assess the oft-asserted claim that escaped fires from oil palm concessions and smallholder farms near settlements are the primary sources of fire in a peat-swamp forest area in Central Kalimantan, Indonesia, equivalent to around a third of Kalimantan's total peat area. We use the MODIS Active Fire product from 2000 to 2010 to evaluate the fire origin and spread on the land use/land cover classes of legal, industrial oil palm concessions (the only type of legal concession in the study area), non-forest, and forest, as well as in relation to settlement proximity. We find that most fires (68-71%) originate in non-forest, compared to oil palm concessions (17%-19%), and relatively few (6-9%) are within 5 km of settlements. Moreover, most fires started within oil palm concessions and in close proximity to settlements stay within those boundaries (90% and 88%, respectively), and fires that do escape constitute only a small proportion of all fires on the landscape (2% and 1%, respectively). Similarly, a small proportion of fire detections in forest originate from oil palm concessions (2%) and within close proximity to settlements (2%). However, fire ignition density in oil palm (0.055 ignitions km-2) is comparable to that in non-forest (0.060 km-2 ignitions km-2), which is approximately ten times that in forest (0.006 ignitions km-2). Ignition density within 5 km of settlements is the highest at 0.125 ignitions km-2. Furthermore, increased anthropogenic activity in close proximity to oil palm concessions and settlements produces a detectable pattern of fire activity. The number of ignitions decreases exponentially with distance from concessions; the number of ignitions initially increases with distance from settlements, and, around from 7.2 km, then decreases with distance from settlements. These results refute the claim that most fires originate in oil palm concessions, and that fires escaping from oil palm concessions and settlements constitute a major proportion of fires in this study region. However, there is a potential for these land use types to contribute substantially to the fire landscape if their area expands. Effective fire management in this area should therefore target not just oil palm concessions, but also non-forested, degraded areas where ignitions and fires escaping into forest are most likely to occur.

1. Tropical fruit-feeding Nymphalidae butterflies are widely used in research, monitoring, and conservation projects, but to date a key aspect of their behaviour-dispersal-remains poorly understood. They have anecdotally been described as 'relatively sedentary' based on movement vectors from mark-recapture studies, but this may be inaccurate because plot-based studies in small sampling areas often underrepresent long-distance movements. 2. Based on data from a peat-swamp forest in Borneo, it was found that these butterflies may be much more mobile than previously thought, as they frequently moved distances of 1-2km between sampling plots. Median daily movements were approximately 200-250m, and over lifespans of one or more months these movements may sum to total life-time dispersals of several kilometres. 3. Recapture rates for long-distance movements between sampling plots were between 28.2% and 41.6% of the re-encounter rates that would be predicted by a random-walk approximation/Brownian motion (without accounting for survival rates), supporting the hypothesis that it is a suitable model of dispersal in this group, although further research is needed to confirm this. 4. There was no evidence that butterflies occupied permanent home ranges, and it is suggested that a permanent home range or territorial behaviour would be maladaptive in this group. Pseudo-replication caused by 'trap-happy' behaviour was not found to be widespread, and some recommendations are provided regarding the treatment of recapture data in trap-based studies. 5. These findings substantially increase the spatial parameters for future research and conservation projects in this group and are also applicable to theoretical modelling studies. An abstract for this article in Bahasa Indonesia is included in the online supporting information File S1.

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.

Responses of biodiversity to changes in both land cover and climate are recognized [1] but still poorly understood [2]. This poses significant challenges for spatial planning as species could shift, contract, expand, or maintain their range inside or outside protected areas [2-4]. We examine this problem in Borneo, a global biodiversity hotspot [5], using spatial prioritization analyses that maximize species conservation under multiple environmental-change forecasts. Climate projections indicate that 11%-36% of Bornean mammal species will lose ≥ 30% of their habitat by 2080, and suitable ecological conditions will shift upslope for 23%-46%. Deforestation exacerbates this process, increasing the proportion of species facing comparable habitat loss to 30%-49%, a 2-fold increase on historical trends. Accommodating these distributional changes will require conserving land outside existing protected areas, but this may be less than anticipated from models incorporating deforestation alone because some species will colonize high-elevation reserves. Our results demonstrate the increasing importance of upland reserves and that relatively small additions (16,000-28,000 km(2)) to the current conservation estate could provide substantial benefits to biodiversity facing changes to land cover and climate. On Borneo, much of this land is under forestry jurisdiction, warranting targeted conservation partnerships to safeguard biodiversity in an era of global change.

The orangutan is the world's largest arboreal mammal, and images of the red ape moving through the tropical forest canopy symbolise its typical arboreal behaviour. Records of terrestrial behaviour are scarce and often associated with habitat disturbance. We conducted a large-scale species-level analysis of ground-based camera-trapping data to evaluate the extent to which Bornean orangutans Pongo pygmaeus come down from the trees to travel terrestrially, and whether they are indeed forced to the ground primarily by anthropogenic forest disturbances. Although the degree of forest disturbance and canopy gap size influenced terrestriality, orangutans were recorded on the ground as frequently in heavily degraded habitats as in primary forests. Furthermore, all age-sex classes were recorded on the ground (flanged males more often). This suggests that terrestrial locomotion is part of the Bornean orangutan's natural behavioural repertoire to a much greater extent than previously thought, and is only modified by habitat disturbance. The capacity of orangutans to come down from the trees may increase their ability to cope with at least smaller-scale forest fragmentation, and to cross moderately open spaces in mosaic landscapes, although the extent of this versatility remains to be investigated.

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

Forest degradation is leading to widespread negative impacts on biodiversity in South-east Asia. Tropical peat-swamp forests are one South-east Asian habitat in which insect communities, and the impacts of forest degradation on them, are poorly understood. To address this information deficit, we investigated the impacts of forest gaps on fruit-feeding butterflies in the Sabangau peat-swamp forest, Central Kalimantan, Indonesia. Fruit-baited traps were used to monitor butterflies for 3months during the 2009 dry season. A network of 34 traps (ngap=17, nshade=17) was assembled in a grid covering a 35ha area. A total of 445 capture events were recorded, comprising 384 individuals from 8 species and 2 additional species complexes classified to genera. On an inter-site scale, canopy traps captured higher species richness than understory traps; however, understory traps captured higher diversity within each site. Species richness was positively correlated with percent canopy cover and comparisons of diversity indices support these findings. Coupled with results demonstrating morphological differences in thorax volume and forewing length between species caught in closed-canopy traps vs. those in gaps, this indicates that forest degradation has a profound effect on butterfly communities in this habitat, with more generalist species being favored in disturbed conditions. Further studies are necessary to better understand the influences of macro-habitat quality and seasonal variations on butterfly diversity and community composition in South-east Asian peat-swamp forests. © 2012 Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society.

Increased resource investment in conservation is generating greater appreciation of the importance of ecological monitoring programmes to assess the effectiveness of conservation interventions in achieving their stated goals. A key component for developing such a programme is an appropriate conceptual model of ecosystem function and the effects of human activities on this. Tropical peatswamp forests are a particularly important ecosystem for reducing carbon emissions and biodiversity conservation. This is leading to increased investment in their protection and, consequently, an increased need for effective ecological monitoring programmes and conceptual models of ecosystem function on which to base these. Here, I adapt previous conceptual models of ecosystem function developed for terrestrial forests and other wetland ecosystems, to create a habitat-specific model for tropical peat-swamp forests. This provides a tool to guide thinking in developing ecological monitoring studies in this habitat and understanding (i) ecosystem processes and function, (ii) the impact of human activities on these, and (iii) subsequent impacts on ecosystem services. This is relevant for monitoring the effectiveness of conservation interventions with varying goals in tropical peat-swamp forest, including emission reductions; and highlights considerations relevant for conservation management and ecological monitoring in other wetland habitats. © US Government 2013.

Development of a standard monitoring protocol for assessing the impacts of REDD+ (Reduced Emissions from Deforestation and Degradation) projects on biodiversity is desirable. Drawing on the conservation literature and our personal experience, we review whether such a one-size-fits-all approach is justifiable on scientific and practical grounds. We conclude that achieving a model biodiversity monitoring protocol suitable for use in all potential REDD+ sites is probably an unrealistic objective, owing to the huge differences among the world's forests in terms of structure, species composition, ecological interactions and ecosystem services provided. Moreover, to provide useful feedback for conservation managers, ecological monitoring programmes must be designed around a project's conservation goals, which will vary from project to project, owing to these differences in forest ecology and human threats faced. Thus, site-specific biodiversity monitoring programmes are needed. No single monitoring method is likely to be optimal, or even suitable for use, in all REDD+ forests. Instead, we suggest that a standard approach be adopted, in which ecological monitoring research is (i) designed to reflect a project's biodiversity conservation goals; (ii) based upon scientifically-tractable, policy-relevant questions regarding the impacts of management interventions on the ecosystem; (iii) founded on detailed knowledge of the habitat type in question; (iv) includes monitoring of a number of indicators, as appropriate to the project; and (v) defines appropriate reference/baseline conditions against which progress can be assessed. © Mark E. Harrison, Arjan Boonman, Susan M. Cheyne, Simon J. Husson, Nicholas C. Marchant and Matthew J. Struebig.

Here, we show how the mechanical properties of a thick-shelled tropical seed are adapted to permit them to germinate while preventing their predation. The seed has evolved a complex heterogeneous microstructure resulting in hardness, stiffness and fracture toughness values that place the structure at the intersection of these competing selective constraints. Analyses of different damage mechanisms inflicted by beetles, squirrels and orangutans illustrate that cellular shapes and orientations ensure damage resistance to predation forces imposed across a broad range of length scales. This resistance is shown to be around the upper limit that allows cracking the shell via internal turgor pressure (i.e. germination). Thus, the seed appears to strike an exquisitely delicate adaptive balance between multiple selection pressures. This journal is © 2011 the Royal Society.

The influence of habitat structure and support availability on support use is an important aspect of understanding locomotor behavior in arboreal primates. We compared habitat structure and support availability in three orangutan study sites-two on Sumatra (Pongo abelii) in the dry-lowland forest of Ketambe and peat swamp forest of Suaq Balimbing, and one on Borneo (Pongo pygmaeus wurmbii) in the disturbed peat swamp forest of Sabangau-to better understand orangutan habitat use. Our analysis revealed vast differences in tree and liana density between the three sites. Sabangau had a much higher overall tree density, although both Sumatran sites had a higher density of larger trees. The two peat swamp forests were more similar to each other than to Ketambe, particularly with regard to support availability. Ketambe had a wider variety of supports of different sizes and types, and a higher density of larger lianas than the two peat swamps. Orangutans in all three sites did not differ substantially in terms of their preferred supports, although Sumatran orangutans had a strong tendency to use lianas, not observed in Sabangau. Differences in observed frequencies of locomotor behavior suggest the homogeneous structure of Sabangau limits the locomotor repertoire of orangutans, with high frequencies of fewer behaviors, whereas the wider range of supports in Ketambe appears to have facilitated a more varied locomotor repertoire. There were no differences among age-sex classes in the use of arboreal pathways in Suaq Balimbing, where orangutans selected larger trees than were typically available. This was less apparent in Sabangau, where orangutans generally used trees in relation to their environmental abundance, reflecting the homogeneous nature of disturbed peat swamp forest. These results demonstrate that forest architecture has an important influence on orangutan locomotion, which may become increasingly important as the structure of orangutan habitat continues to be altered through human disturbance. © 2012 Wiley Periodicals, Inc.

One major concern regarding the biodiversity impacts of Reduced Emissions from Deforestation and Degradation (REDD+) is "leakage" of threats from REDD+ to non-REDD+ forests, particularly if those forests storing the highest amounts of carbon - and thus prioritized under REDD+ - do not coincide with those most important for biodiversity conservation. This concern applies globally, and has been previously discussed in Indonesia, where the highest-carbon forests on peat are known to support lower species diversity and concentrations of threatened species than lowland mineral-soil forests. To help refine management of this risk, we discuss previously overlooked considerations regarding biodiversity threat leakage, suggest three strategies for managing leakage risk, and outline important questions to address with respect to these. We emphasize (1) the need to recognize intrinsic differences in threat displacement vulnerability among forests not currently protected/proposed to be protected under REDD+; and (2) that not pursuing REDD+ in high-carbon forests in an attempt to avoid leakage will not necessarily reduce this risk in low-carbon, non-REDD+ forests, due to the often high intrinsic vulnerability of these forests. Further to previous recommendations, suggested strategies for reducing risk of threat displacement include (1) focusing "traditional" conservation resources on the most vulnerable high-biodiversity forests not scheduled for protection under REDD+; (2) reducing costs, simplifying procedures and encouraging community-based approaches for pursuing REDD+ in low-carbon, high-biodiversity forests; and (3) developing more creative measures, especially fiscal and financial incentives, for protecting vulnerable low-carbon forests. Inter-disciplinary research is urgently needed to evaluate the feasibility and effectiveness of these strategies to successfully manage biodiversity leakage risk from pursuing REDD+ in high-carbon forests and, thus, for ensuring REDD+ achieves its potential for generating biodiversity conservation gains. © Mark E. Harrison and Gary D. Paoli.

Widespread hunting of flying foxes has generated concern regarding population declines and the spread of emerging infectious diseases. To investigate the potential impacts of this trade, we conducted questionnaires in 45 settlements across 12 population centres within Central Kalimantan, Indonesia, a region previously identified as a hunting hotspot. By combining results from 63 hunter and 88 vendor interviews, we highlight two population centres (Palangka Raya and Buntok/Tamiang Layang) with higher hunting rates than other areas, which act as flying fox trading hubs. Flying fox populations were perceived to be declining province-wide: declines in captures and sales were reported by 81% of hunters and 60% of market vendors, who also reported availability as the key factor underlying temporal variations in trade. There was substantial risk of zoonotic disease transmission between bats, hunters and traders: the vast majority of respondents were unaware that flying foxes carry potentially fatal viruses, and so few people protected themselves from physical contact. Moreover, both hunters and vendors were frequently bitten and the majority of bites drew blood. Most hunters (58%) also reported unintentional by-catches that included keystone bird species and slow lorises. The scale of hunting over Central Kalimantan represents a serious threat to the long-term viability of flying fox populations (and potentially those of other species), and could have serious public health implications. Reducing or eliminating hunting and trade would mitigate the risk of disease transmission, while maintaining the economic and ecosystem benefits that flying foxes provide, in terms of pollination and seed dispersal. © 2011 Elsevier Ltd.

Researchers have suggested that fallback foods (FBFs) shape primate food processing adaptations, whereas preferred foods drive harvesting adaptations, and that the dietary importance of FBFs is central in determining the expression of a variety of traits. We examine these hypotheses in extant apes. First, we compare the nature and dietary importance of FBFs used by each taxon. FBF importance appears greatest in gorillas, followed by chimpanzees and siamangs, and least in orangutans and gibbons (bonobos are difficult to place). Next, we compare 20 traits among taxa to assess whether the relative expression of traits expected for consumption of FBFs matches their observed dietary importance. Trait manifestation generally conforms to predictions based on dietary importance of FBFs. However, some departures from predictions exist, particularly for orang-utans, which express relatively more food harvesting and processing traits predicted for consuming large amounts of FBFs than expected based on observed dietary importance. This is probably due to the chemical, mechanical, and phenological properties of the apes' main FBFs, in particular high importance of figs for chimpanzees and hylobatids, compared to use of bark and leaves-plus figs in at least some Sumatran populations-by orang-utans. This may have permitted more specialized harvesting adaptations in chimpanzees and hylobatids, and required enhanced processing adaptations in orang-utans. Possible intercontinental differences in the availability and quality of preferred and FBFs may also be important. Our analysis supports previous hypotheses suggesting a critical influence of the dietary importance and quality of FBFs on ape ecology and, consequently, evolution. © 2011 the Author(s).

Orang-utans (Pongo spp.) are primarily frugivorous (Morrogh-Bernard et al. 2009) and are often regarded as important seed dispersers (Corlett 1998). In Tanjung Puting, Borneo, Galdikas (1982) found intact seeds in 94% of faecal samples, with a median 111 seeds per defecation; and in Ketambe, Sumatra, Rijksen (1978) found seeds in 44% of faecal samples. Furthermore, orang-utans have large day ranges (e.g. mean = 968 m, range = 280-2834 m across adults in Sabangau; Harrison 2009) and slow passage rates of digesta through the gut (Caton et al. 1999), and, hence, may disperse seeds far from parent trees. Many seeds are also spat out or discarded at distances up to 75 m from parent trees (Galdikas 1982). © 2011 Cambridge University Press.

Faecal samples from 163 captive and semi-captive individuals, 61 samples from wild individuals and 38 samples from captive groups of Bornean orangutans (Pongo pygmaeus) in Kalimantan, Indonesia, were collected during one rainy season (November 2005May 2006) and screened for intestinal parasites using sodium acetate-acetic acid-formalin-concentration (SAFC), sedimentation, flotation, McMaster- and Baermann techniques. We aimed to identify factors influencing infection risk for specific intestinal parasites in wild orangutans and individuals living in captivity. Various genera of Protozoa (including Entamoeba, Endolimax, Iodamoeba, Balantidium, Giardia and Blastocystis), nematodes (such as Strongyloides, Trichuris, Ascaris, Enterobius, Trichostrongylus and hookworms) and one trematode (a dicrocoeliid) were identified. For the first time, the cestode Hymenolepis was detected in orangutans. Highest prevalences were found for Strongyloides (individuals 37%; groups 58%), hookworms (41%; 58%), Balantidium (40%; 61%), Entamoeba coli (29%; 53%) and a trichostrongylid (13%; 32%). In re-introduction centres, infants were at higher risk of infection with Strongyloides than adults. Infection risk for hookworms was significantly higher in wild males compared with females. In groups, the centres themselves had a significant influence on the infection risk for Balantidium. Ranging patterns of wild orangutans, overcrowding in captivity and a shift of age composition in favour of immatures seemed to be the most likely factors leading to these results. © 2009 Cambridge University Press.

Data on energy intake and the effects of fluctuations in fruit availability on energy intake for African apes, and orangutans in mast-fruiting habitats, indicate that orangutans may face greater energetic challenges than do their African counterparts. Comparable data on orangutans in nonmasting forests, which experience lower fluctuations in fruit availability, have been lacking, however, complicating interpretations. We conducted a 46-mo study of orangutan energetics in the nonmasting Sabangau peat-swamp forest, Indonesian Borneo. Sabangau orangutans experienced periods of negative energy balance apparently even longer than in mast-fruiting habitats, as indicated by comparisons of observed energy intake with theoretical requirements and analysis of urinary ketones. Daily energy intake was positively related to fruit availability in flanged males, but not in adult females or unflanged males. This may represent different foraging strategies between age-sex classes and suggests that fruit availability is not always an accurate indicator of ape energy intake/balance. Urinary ketone levels were not generally related to fruit availability, daily energy intake, day range, or party size. This is probably due to low energy intake, and consequently high ketone production, throughout much of the study period. Comparisons with published results on African apes support the hypothesis that orangutans are unique among hominoids in regularly experiencing prolonged periods of negative energy balance. This has important effects on orangutan behavior and socioecology, and has likely been a key factor driving the evolutionary divergence of orangutans and African apes. © 2010 Springer Science+Business Media, LLC.

The results of comparisons of behavioral data between individuals, age-sex classes, seasons, sites and possibly even species may depend on sample size and the computational method used. To establish whether these are valid concerns, we compared results for percentage time spent feeding on major food types (fruit, leaves, flowers, invertebrates, bark, pith and other) for two orangutan populations in Sabangau (24 months) and Tuanan (29 months), Indonesian Borneo. Both the minimum follow limit included in analyses and the computational method used produced small, but statistically significant, differences in the results obtained, and the differences were more common for food types eaten less frequently. In addition, using different computational methods produced more significant differences than did including different minimum follow lengths in analyses. The computational method used also influenced the results of tests for differences in diet composition between age-sex classes. Thus, the method used can influence the results obtained and, hence, it is important to state explicitly the minimum follow limit included and computational method used to compile averages, and to ensure standardization in methods when comparing data between age-sex classes, time periods, field sites or species. Am. J. Primatol. 71:353-358, 2009. © 2008 Wiley-Liss, Inc.

The hippopotamus population in Liwonde National Park, Malawi was studied from December 2002 to June 2003. Motorboat censuses along the River Shire counted 640 animals, but because of the large number of hippos in temporary water sources at the time, the true number is probably closer to 950. Marked shifts in hippo distribution from the Shire into temporary water sources occurred as the wet season advanced. Because of the Shire's year-round water supply, the hippo population is not regulated by the availability of aquatic refuges, but by food availability. By following feeding tracks, dry season grazing range was estimated to extend 5 km east and 1 km west of the river. Grazing intensity transects and visual estimates indicated only a small portion of this area is suitable for hippo grazing, leading to over-grazing in suitable areas. Coupled with low primary productivity levels in Liwonde, this means that dry season food competition between hippos and other herbivores is probably high. Recent proposals to raise the Shire's dry season water level should be considered very carefully, as this will flood late dry season grazing grounds, thereby intensifying grazing competition and increasing grazing pressure in remaining grazing areas, having potentially serious impacts on the animal community. © 2007 the Authors.

Pteropus vampyrus natunae, the Bornean subspecies of the large flying fox, has important roles in pollination but unsustainable hunting has been reported in Malaysian states. We provide the first description of hunting techniques and intensity in Indonesian Borneo. In forests around Palangka Raya this species is captured in canopy-level nets to support trade in the provincial capital. We estimate that in 2003 4,500 individuals were extracted from a single location in 30 days, which, together with trends reported in interviews with hunters and traders, suggests that hunting in this region is intensive and probably causing severe population declines. Further surveys are needed throughout Kalimantan to determine if this trend is occurring around other cities and whether intervention is needed to safeguard viable populations. © 2007 FFI.

The orang-utan is unique among apes in having an unusually long male developmental period and two distinct adult male morphs (flanged and unflanged), which generally, but not exclusively, employ different reproductive strategies (call-and-wait vs. sneak-and-rape). Both morphs have recently been shown to have roughly similar levels of reproductive success in the one site where such a study has been conducted. This is in stark contrast to the unimale polygynous gorilla, in which dominant males sire almost all infants. Despite this, evidence on sexual dimorphism, life history, diet, and socioecology of extant and extinct apes, as well as the ontogeny, reproductive morphology, and physiology of extant apes, all indicate that the orang-utan's present-day mating system most likely evolved from a gorilla-like base, with one dominant male guarding a harem of females. The available evidence indicates that, due chiefly to the likely onset of the El Niño Southern Oscillation (generally regarded as the trigger for mast fruiting in dipterocarps) approximately 3-5 Ma, southeast Asian forests would have begun to experience longer and more severe periods of low food availability. This change in food availability would have meant that full-time gregariousness was no longer energetically tolerable and, as a result, females dispersed more widely in search of food and adult/flanged males were no longer able to effectively guard a harem of females. A niche for a quiet, quick, opportunistic "sexual predator" (i.e. the unflanged male) then became available. This finding implies that, despite being anatomically quite chimpanzee-like, the ancestral hominoid probably had a social and mating system more similar to the gorilla than any other living ape. © 2006 Elsevier Ltd. All rights reserved.