Oliver R. Wearn

Extinction Debt and Windows of Conservation Opportunity in the Brazilian Amazon

Growing Extinction Debt Predicting, and potentially preventing, extinction is a central goal of conservation biology. Wearn et al. (p. 228; see the Perspective by Rangel) describe a mathematical approach for predicting the time lags in extinction following habitat loss. The model was applied to the highly biodiverse Brazilian Amazon region, and used to reconstruct the spatial and temporal patterns of extinction and the accumulation of extinction debt from 1970 through to the present, and to extrapolate to 2050 under four deforestation scenarios. The Amazon basin sits at a critical point: Few species have been driven extinct to date, but an extinction debt is rapidly accumulating, which could lead to an increasing rate of extinction in the next four decades. Deforestation scenarios predict species extinction rates and identify targets of conservation efforts. Predicting when future species extinctions will occur is necessary for directing conservation investments but has proved difficult. We developed a new method for predicting extinctions over time, accounting for the timing and magnitude of habitat loss. We applied this to the Brazilian Amazon, predicting that local extinctions of forest-dependent vertebrate species have thus far been minimal (1% of species by 2008), with more than 80% of extinctions expected to be incurred from historical habitat loss still to come. Realistic deforestation scenarios suggest that local regions will lose an average of nine vertebrate species and have a further 16 committed to extinction by 2050. There is a window of opportunity to dilute the legacy of historical deforestation by concentrating conservation efforts in areas with greatest debt.

Logging cuts the functional importance of invertebrates in tropical rainforest

Invertebrates are dominant species in primary tropical rainforests, where their abundance and diversity contributes to the functioning and resilience of these globally important ecosystems. However, more than one-third of tropical forests have been logged, with dramatic impacts on rainforest biodiversity that may disrupt key ecosystem processes. We find that the contribution of invertebrates to three ecosystem processes operating at three trophic levels (litter decomposition, seed predation and removal, and invertebrate predation) is reduced by up to one-half following logging. These changes are associated with decreased abundance of key functional groups of termites, ants, beetles and earthworms, and an increase in the abundance of small mammals, amphibians and insectivorous birds in logged relative to primary forest. Our results suggest that ecosystem processes themselves have considerable resilience to logging, but the consistent decline of invertebrate functional importance is indicative of a human-induced shift in how these ecological processes operate in tropical rainforests.

Assessing the Status of Wild Felids in a Highly-Disturbed Commercial Forest Reserve in Borneo and the Implications for Camera Trap Survey Design

The proliferation of camera-trapping studies has led to a spate of extensions in the known distributions of many wild cat species, not least in Borneo. However, we still do not have a clear picture of the spatial patterns of felid abundance in Southeast Asia, particularly with respect to the large areas of highly-disturbed habitat. An important obstacle to increasing the usefulness of camera trap data is the widespread practice of setting cameras at non-random locations. Non-random deployment interacts with non-random space-use by animals, causing biases in our inferences about relative abundance from detection frequencies alone. This may be a particular problem if surveys do not adequately sample the full range of habitat features present in a study region. Using camera-trapping records and incidental sightings from the Kalabakan Forest Reserve, Sabah, Malaysian Borneo, we aimed to assess the relative abundance of felid species in highly-disturbed forest, as well as investigate felid space-use and the potential for biases resulting from non-random sampling. Although the area has been intensively logged over three decades, it was found to still retain the full complement of Bornean felids, including the bay cat Pardofelis badia, a poorly known Bornean endemic. Camera-trapping using strictly random locations detected four of the five Bornean felid species and revealed inter- and intra-specific differences in space-use. We compare our results with an extensive dataset of >1,200 felid records from previous camera-trapping studies and show that the relative abundance of the bay cat, in particular, may have previously been underestimated due to the use of non-random survey locations. Further surveys for this species using random locations will be crucial in determining its conservation status. We advocate the more wide-spread use of random survey locations in future camera-trapping surveys in order to increase the robustness and generality of inferences that can be made.

Making statistics biologically relevant in fragmented landscapes

The biological impacts of habitat fragmentation are routinely assessed using standard statistical modelling techniques that are used across many ecological disciplines. However, to assess the biological relevance of fragmentation impacts, we must consider an extra, spatial dimension to the standard statistical model: the biological importance of a significant and well supported model with large effect sizes crucially depends on the configuration of habitat within the study area. We argue that mapping the outputs from statistical models across a study area generates biologically meaningful estimates of fragmentation impacts. Integrating traditional statistical approaches with geographic information systems will facilitate rigorous comparisons of fragmentation impacts between taxa, studies and ecosystems.

The Environmental Legacy of Modern Tropical Deforestation

Summary Tropical deforestation has caused a significant share of carbon emissions and species losses, but historical patterns have rarely been explicitly considered when estimating these impacts [1]. A deforestation event today leads to a time-delayed future release of carbon, from the eventual decay either of forest products or of slash left at the site [2]. Similarly, deforestation often does not result in the immediate loss of species, and communities may exhibit a process of “relaxation” to their new equilibrium over time [3]. We used a spatially explicit land cover change model [4] to reconstruct the annual rates and spatial patterns of tropical deforestation that occurred between 1950 and 2009 in the Amazon, in the Congo Basin, and across Southeast Asia. Using these patterns, we estimated the resulting gross vegetation carbon emissions [2, 5] and species losses over time [6]. Importantly, we accounted for the time lags inherent in both the release of carbon and the extinction of species. We show that even if deforestation had completely halted in 2010, time lags ensured there would still be a carbon emissions debt of at least 8.6 petagrams, equivalent to 5–10 years of global deforestation, and an extinction debt of more than 140 bird, mammal, and amphibian forest-specific species, which if paid, would increase the number of 20th-century extinctions in these groups by 120%. Given the magnitude of these debts, commitments to reduce emissions and biodiversity loss are unlikely to be realized without specific actions that directly address this damaging environmental legacy.

BIOFRAG - a new database for analyzing BIOdiversity responses to forest FRAGmentation

Habitat fragmentation studies have produced complex results that are challenging to synthesize. Inconsistencies among studies may result from variation in the choice of landscape metrics and response variables, which is often compounded by a lack of key statistical or methodological information. Collating primary datasets on biodiversity responses to fragmentation in a consistent and flexible database permits simple data retrieval for subsequent analyses. We present a relational database that links such field data to taxonomic nomenclature, spatial and temporal plot attributes, and environmental characteristics. Field assessments include measurements of the response(s) (e.g., presence, abundance, ground cover) of one or more species linked to plots in fragments within a partially forested landscape. The database currently holds 9830 unique species recorded in plots of 58 unique landscapes in six of eight realms: mammals 315, birds 1286, herptiles 460, insects 4521, spiders 204, other arthropods 85, gastropods 70, annelids 8, platyhelminthes 4, Onychophora 2, vascular plants 2112, nonvascular plants and lichens 320, and fungi 449. Three landscapes were sampled as long-term time series (>10 years). Seven hundred and eleven species are found in two or more landscapes. Consolidating the substantial amount of primary data available on biodiversity responses to fragmentation in the context of land-use change and natural disturbances is an essential part of understanding the effects of increasing anthropogenic pressures on land. The consistent format of this database facilitates testing of generalizations concerning biologic responses to fragmentation across diverse systems and taxa. It also allows the re-examination of existing datasets with alternative landscape metrics and robust statistical methods, for example, helping to address pseudo-replication problems. The database can thus help researchers in producing broad syntheses of the effects of land use. The database is dynamic and inclusive, and contributions from individual and large-scale data-collection efforts are welcome.

Grain‐dependent responses of mammalian diversity to land use and the implications for conservation set‐aside

Diversity responses to land-use change are poorly understood at local scales, hindering our ability to make forecasts and management recommendations at scales which are of practical relevance. A key barrier in this has been the underappreciation of grain-dependent diversity responses and the role that β-diversity (variation in community composition across space) plays in this. Decisions about the most effective spatial arrangement of conservation set-aside, for example high conservation value areas, have also neglected β-diversity, despite its role in determining the complementarity of sites. We examined local-scale mammalian species richness and β-diversity across old-growth forest, logged forest, and oil palm plantations in Borneo, using intensive camera- and live-trapping. For the first time, we were able to investigate diversity responses, as well as β-diversity, at multiple spatial grains, and across the whole terrestrial mammal community (large and small mammals); β-diversity was quantified by comparing observed β-diversity with that obtained under a null model, in order to control for sampling effects, and we refer to this as the β-diversity signal. Community responses to land use were grain dependent, with large mammals showing reduced richness in logged forest compared to old-growth forest at the grain of individual sampling points, but no change at the overall land-use level. Responses varied with species group, however, with small mammals increasing in richness at all grains in logged forest compared to old-growth forest. Both species groups were significantly depauperate in oil palm. Large mammal communities in old-growth forest became more heterogeneous at coarser spatial grains and small mammal communities became more homogeneous, while this pattern was reversed in logged forest. Both groups, however, showed a significant β-diversity signal at the finest grain in logged forest, likely due to logging-induced environmental heterogeneity. The β-diversity signal in oil palm was weak, but heterogeneity at the coarsest spatial grain was still evident, likely due to variation in landscape forest cover. Our findings suggest that the most effective spatial arrangement of set-aside will involve trade-offs between conserving large and small mammals. Greater consideration in the conservation and management of tropical landscapes needs to be given to β-diversity at a range of spatial grains.

Species Richness and Distribution of Primates in Disturbed and Converted Forest Landscapes in Northern Borneo

Knowledge of fundamental aspects of ecology such as species richness and distribution, and the factors affecting them, is increasingly used to identify priority areas for conservation and to effectively manage threatened species. We investigated the species richness and distribution pattern of nonhuman primate communities inhabiting 10 sampling sites in four different habitat classes corresponding to increasing habitat disturbance level, that is, old growth forest, twice logged forest, repeatedly logged forest, and oil palm plantation, in and around Kalabakan Forest Reserve, in central Sabah, Malaysian Borneo. By using direct and indirect survey methods, we confirmed the existence of 9 out of the total 10 primate species, found in Sabah, within the surveyed areas. Based on the monthly number of primate species detected, our results indicated a general trend of decreasing primate species richness with increasing habitat disturbance level. However, the response within the disturbed forest sites showed some variations with some sites in repeatedly logged forest displaying comparable primate species number to that of the undisturbed forest sites. We also found that within the forest habitats, tree density is a good predictor of the richness of the primate community with a positive effect. Hence, tree density may be a key indicator for evaluating primate communities in forest habitats. Overall, the results of our study suggest that although not equivalent to areas of undisturbed forest, degraded forests—including those that have been repeatedly logged—are still valuable for primate conservation. In contrast, oil palm plantations have mainly negative effects on the primate community.

Terrestrial Mammal Species Richness and Composition in Three Small Forest Patches within an Oil Palm Landscape in Sabah, Malaysian Borneo

Abstract. Small and highly degraded forest patches are usually scattered across oil palm plantation landscapes and often exist as permanent features. By using a combination of camera-trapping and line-transect methods, we evaluated the usefulness of three such forest patches (< 30 ha) for terrestrial mammal species conservation in a mature oil palm plantation located near (< 1.7 km) a large continuous tract of logged forest in eastern Sabah, Malaysian Borneo. Of the 29 terrestrial mammal species recorded in this study, 28 were found in the continuous logged forest habitat including six species that are either large-bodied, wide ranging, locally rare or are of high conservation concern. In comparison, 18 species were recorded across the three forest patches collectively; consisting mostly of species that are widespread, well-adapted to living in highly modified habitats and of low conservation concern. The presence of small forest patches within the oil palm habitat matrix seemed to be useful to some extent for some mammal species. However, many of the species were likely only transient in this habitat. The maintenance of large continuous tracts of natural forest is critical to the continued survival of many terrestrial mammal species on Borneo, particularly for species that are of high conservation value.

Influence of microhabitat structure and disturbance on detection of native and non-native murids in logged and unlogged forests of northern Borneo

Understanding the habitat preferences of native and non-native species may offer valuable insights into the mechanisms favouring invasion of disturbed habitats. This study investigated the determinants of trap-site detection probability of three native (Maxomys surifer, Maxomys whiteheadi and Leopoldamys sabanus) and one invasive (Rattus rattus) species of terrestrial murid (Muridae) in logged and unlogged forests of northern Borneo. We established four andtwotrappinggridsinrepeatedlyloggedandunloggedforest,respectively,foratotalof500sampledtrapsites.From these, we obtained 504 detections of the four species over 3420 trap nights. For each species, probability of detection wasmodelledasafunctionofboththestructuralcomponentsanddisturbanceleveloftheforestpatchmeasuredaround each trap site. Each of the four species showed contrasting microhabitat preferences: M. surifer favoured increased canopy closure and intermediate ground and understorey vegetation cover; M. whiteheadi preferred increased ground vegetation cover and canopy height;L. sabanusfavoured sites with larger amounts of coarse woody debris and less leaf litter;andR.rattuswasassociatedwithincreasedgroundvegetationcover.Withinloggedforest,detectionprobabilities of the three native species did not vary significantly with level of patch disturbance, whereas that of the invasive R. rattus increased markedly in more degraded sites. This latter finding will have increasingly important implications when considering the rapid degradation of forests in the region, and the resulting expansion of suitable habitat for this competitive species.