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The demand for timber products is facilitating the degradation and opening up of large areas of intact habitats rich in biodiversity. Logging creates an extensive network of access roads within the forest, yet these are commonly ignored or excluded when assessing impacts of logging on forest biodiversity. Here we determine the impact of these roads on the overall condition of selectively logged forests in Borneo, Southeast Asia. Focusing on dung beetles along > 40 km logging roads we determine: (i) the magnitude and extent of edge effects alongside logging roads; (ii) whether vegetation characteristics can explain patterns in dung beetle communities, and; (iii) how the inclusion of road edge forest impacts dung beetle assemblages within the overall logged landscape. We found that while vegetation structure was significantly affected up to 34 m from the road edge, impacts on dung beetle communities penetrated much further and were discernible up to 170 m into the forest interior. We found larger species and particularly tunnelling species responded more than other functional groups which were also influenced by micro-habitat variation. We provide important new insights into the long-term ecological impacts of tropical logging. We also support calls for improved logging road design both during and after timber extraction to conserve more effectively biodiversity in production forests, for instance, by considering the minimum volume of timber, per unit length of logging road needed to justify road construction. In particular, we suggest that governments and certification bodies need to highlight more clearly the biodiversity and environmental impacts of logging roads., Highlights • Distinct edge effects along roads penetrate > 100 m into the logged forest interior. • Vegetation changes however, differed within a maximum of 34 m of the road edge. • Large diurnal, and tunnelling dung beetles are affected more by proximity to roads. • Micro-habitat variations shown to influence large tunnelling species.

Freshwaters provide valuable habitat and important ecosystem services but are threatened worldwide by habitat loss and degradation. In Southeast Asia, rainforest streams are particularly threatened by logging and conversion to oil palm, but we lack information on the impacts of this on freshwater environmental conditions, and the relative importance of catchment versus riparian‐scale disturbance. We studied 16 streams in Sabah, Borneo, including old‐growth forest, logged forest, and oil palm sites. We assessed forest quality in riparian zones and across the whole catchment and compared it with stream environmental conditions including water quality, structural complexity, and organic inputs. We found that streams with the highest riparian forest quality were nearly 4 °C cooler, over 20 cm deeper, had over 40% less sand, greater canopy cover, more stored leaf litter, and wider channels than oil palm streams with the lowest riparian forest quality. Other variables were significantly related to catchment‐scale forest quality, with streams in the highest quality forest catchments having 40% more bedrock and 20 times more dead wood, along with higher phosphorus, and lower nitrate‐N levels compared to streams with the lowest catchment‐scale forest quality. Although riparian buffer strips went some way to protecting waterways, they did not maintain fully forest‐like stream conditions. In addition, logged forest streams still showed signs of disturbance 10–15 years after selective logging. Our results suggest that maintenance and restoration of buffer strips can help to protect healthy freshwater ecosystems but logging practices and catchment‐scale forest management also need to be considered.

South East Asia has the highest rate of lowland forest loss of any tropical region, with logging and deforestation for conversion to plantation agriculture being flagged as the most urgent threats. Detecting and mapping logging impacts on forest structure is a primary conservation concern, as these impacts feed through to changes in biodiversity and ecosystem functions. Here, we test whether high-spatial resolution satellite remote sensing can be used to map the responses of aboveground live tree biomass (AGB), canopy leaf area index (LAI) and fractional vegetation cover (FCover) to selective logging and deforestation in Malaysian Borneo. We measured these attributes in permanent vegetation plots in rainforest and oil palm plantations across the degradation landscape of the Stability of Altered Forest Ecosystems project. We found significant mathematical relationships between field-measured structure and satellite-derived spectral and texture information, explaining up to 62% of variation in biophysical structure across forest and oil palm plots. These relationships held at different aggregation levels from plots to forest disturbance types and oil palms allowing us to map aboveground biomass and canopy structure across the degradation landscape. The maps reveal considerable spatial variation in the impacts of previous logging, a pattern that was less clear when considering field data alone. Up-scaled maps revealed a pronounced decline in aboveground live tree biomass with increasing disturbance, impacts which are also clearly visible in the field data even a decade after logging. Field data demonstrate a rapid recovery in forest canopy structure with the canopy recovering to pre-disturbance levels a decade after logging. Yet, up-scaled maps show that both LAI and FCover are still reduced in logged compared to primary forest stands and markedly lower in oil palm stands. While uncertainties remain, these maps can now be utilised to identify conservation win–wins, especially when combining them with ongoing biodiversity surveys and measurements of carbon sequestration, hydrological cycles and microclimate.

Aim To estimate whether species have shifted at equal rates at their leading edges (cool boundaries) and trailing edges (warm boundaries) in response to climate change. We provide the first such evidence for tropical insects, here examining elevation shifts for the upper and lower boundaries shifts of montane moths. Threats to species on tropical mountains are considered.Location Mount Kinabalu, Sabah, Malaysia.Methods We surveyed Lepidoptera (Geometridae) on Mount Kinabalu in 2007, 42 years after the previous surveys in 1965. Changes in species upper and lower boundaries, elevational extents and range areas were assessed. We randomly subsampled the data to ensure comparable datasets between years. Estimated shifts were compared for endemic versus more widespread species, and for species that reached their range limits at different elevations.Results Species that reached their upper limits at 2500-2700 m (n= 28 species, 20% of those considered) retreated at both their lower and upper boundaries, and hence showed substantial average range contractions (-300 m in elevational extent and -45 km2 in estimated range area). These declines may be associated with changes in cloud cover and the presence of ecological barriers (geological and vegetation transitions) which impede uphill movement. Other than this group, most species (n= 109, 80% of the species considered) expanded their upper boundaries upwards (by an average of 152 m) more than they retreated at their lower boundaries (77 m).Main conclusions Without constraints, leading margins shifted uphill faster than trailing margins retreated, such that many species increased their elevational extents. However, this did not result in increases in range area because the area of land available declines with increasing elevation. Species close to a major ecological/geological transition zone on the mountain flank declined in their range areas. Extinction risk may increase long before species reach the summit, even when undisturbed habitats are available.

Oil palm cultivation is expanding rapidly into many of the world’s most biodiverse tropical regions. One of the most functionally important and ecologically dominant animal groups in these environments is the ants. Here, we quantify the overall impacts of clear-felling lowland dipterocarp rainforest and conversion into oil palm plantation on ant diversity. At study sites in Sabah, Malaysia we collected ants from three microhabitats: 1 – the canopy, 2 – bird’s nest ferns (Asplenium nidus complex, a common epiphyte in forest and oil palm), and 3 – leaf litter. We also measured temperature, humidity and light at collection sites to assess their impacts on ant community composition. Total ant species richness decreased from 309 to 110 (−64%) between forest and oil palm plantation. However, this impact was not the same across all microhabitats, with bird’s nest ferns maintaining almost the same number of ant species in oil palm compared to forest (forest-oil palm, ferns: 36–35 (3% loss), canopy: 120–58 (52% loss), leaf litter: 216–56 (74% loss)). Relative abundance distributions remained the same for fern-dwelling ants, but became less even for oil palm ants in both the canopy and the leaf litter. These differences may be due in part to the ability of bird’s nest ferns to provide a stable microclimate in hot, dry plantations. We also found that non-native ant species were more abundant in oil palm than in forest, and few forest ant species survived in plantations in any of the microhabitats. Only 59 of the 309 forest species persisted in oil palm plantations, corresponding to an 81% loss of forest species resulting from habitat conversion. Although oil palm supports many more ant species than has been previously reported, converting forest into plantation still leads to a dramatic reduction in species richness. The maintenance of forested areas is therefore vital for the conservation of ant biodiversity.

A key challenge in the study of mutualism is to understand the mechanisms that prevent cheating. In some systems, host retaliation against cheaters prevents the breakdown of cooperation. Here, we focus on the converse of this demonstration, and ask whether hosts that fail to retaliate are commonly inhabited by cheaters. We do so using the classic antplant interaction, in which plants provide ant-housing (domatia) in return for protection from herbivores. Our model system is the rattan ant-palm Korthalsia furtadoana, which grows swollen leaf sheaths as domatia and associates with two species of obligate host-ants, Camponotus sp90 and C. sp93, and with facultative Crematogaster and 'tramp' ant species. One ant-tree species is known to retaliate by tying the growth of domatia to the successful protection of new leaves, and non-protecting cheaters are rare. In contrast, K. furtadoana grows the domatium before the new leaf develops, suggesting that sanctioning may not be possible. We experimentally simulated herbivory by cutting leaves from shoots and found no difference in the mortality and growth of domatia on such 'cheated' shoots than on controls, confirming that K. furtadoana cannot sanction non-protectors. We then investigated the intensity of protection that Camponotus and Crematogaster ant- symbionts provide K. furtadoana. We demonstrate that C. sp90, which only inhabit half of colonised plants, vigorously protects leaves, that C. sp93 rarely protects, and that Crematogaster never protects. We then show that plants inhabited by C. sp90 have a higher growth rate than those inhabited by C. sp93. We conclude that C. sp90 is a protection mutualist, while C. sp93 and Crematogaster are parasites, the first such demonstrations for an antpalm interaction. The presence of commonly occurring parasites, as well as rare tramp ants, provides the first clear correlative evidence that an inability to punish results in abundant cheating. A key challenge in the study of inter-specific cooperation, or mutualisms, is to understand the mechanisms that prevent cheating. Several overlapping theories, with different de- grees of generality, have been developed to explain the maintenance of cooperation in the face of (a) invasion by parasites that purloin the benefits one mutualist provides another, and (b) selection for cheating behaviour among the

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., Invertebrates are key components in the ecological functioning of tropical forests. Here, Ewers et al. show that, compared to primary forest, logging halves the contribution of invertebrate species to several key ecosystem processes, including litter decomposition.

Summary 1 Widespread and rapid losses of tropical rain forests have made understanding the responses of species to rain forest fragmentation an area of major concern. In this study we examined the impacts of habitat fragmentation on the species richness and faunal composition of butterflies in tropical rain forests in Sabah, Borneo. We analysed patterns of both α- and β-diversity to assess the relative importance of differences in patch size, isolation and vegetation structure on the diversity and similarity of species assemblages. We used additive partitioning to assess the relative contributions of intact forest and forest remnants to overall species richness at a landscape scale and we examined which traits of species best predicted their responses to fragmentation. 2 Species richness and diversity in rain forest remnants was significantly positively related to remnant size and significantly negatively related to isolation, in keeping with theories of island biogeography. Species assemblages at different sites were significantly nested, with those species most adversely affected by forest fragmentation having a narrow range of larval host-plants and, to a lesser extent, being large-bodied. No species endemic to Borneo was recorded in forest remnants smaller than 4000 ha, but even the smallest remnant (120 ha) supported species with geographical distributions confined within Sundaland (West Malaysia and the islands of the Sunda Shelf). 3 Although assemblages were significantly nested, they departed substantially from perfect nestedness, with some species recorded only or predominantly in small, relatively depauperate remnants. As a result there was substantial β-diversity among sites, which was related to variation in both fragment size and vegetation structure. At the landscape scale, diversity within sites was less than that between sites, and the majority of the diversity between sites was related to variation in fragment size. 4 Synthesis and applications. Substantial diversity was added to the assemblage of butterflies in Bornean rain forests by virtue of species differences among fragments, which were related mainly to differences in patch size and vegetation structure. The data reported indicate that, despite having lower species richness, relatively small and isolated remnants of rain forest make a substantial contribution to regional diversity. Small isolated forest remnants are generally accorded low conservation status and given little protection, with the result that they often disappear over time because of continued anthropogenic disturbance. The results of this study indicate that the conservation value of small remnants of forest, in particular their contribution to environmental heterogeneity, should not be overlooked.

Tropical rain forests are well known as centres of insect diversity and much effort has focused on the role of larval host plant specificity in generating and maintaining this diversity, but fewer studies have examined the exploitation of different food resources by adults in this context. Tropical butterflies feed as adults on a wide range of resources and we examined the diversity and ecology of species feeding on rotting fruit and carrion in a tropical lowland rain forest in Sabah, Borneo. We found that species richness and diversity were significantly higher on carrion than on fruit, and that this pattern was repeated at genus and family level. There was little similarity in species assemblages on the two substrates and β-diversity between carrion and fruit comprised 33% of the total diversity of butterflies feeding on decaying matter. β-diversity between canopy gap and shade microhabitats comprised 21% of total species diversity on carrion but only 7% of the total on fruit, indicating greater functional diversity on carrion in terms of light preferences. Captures were strongly male-biased on carrion but not on fruit, and recapture rates were much lower on carrion than on fruit. Species from two subfamilies (Nymphalinae and Charaxinae) exploited both substrates and for Charaxinae, there was evidence from adult flight morphology that species on carrion were capable of faster more-powerful flight. These results support the notion of a distinctive carrion-feeding fauna comprising more mobile species, which may use carrion to meet additional nitrogen requirements resulting from greater musculature. However there was no relationship between flight morphology and substrate choice in the Nymphalinae, and carrion-feeding may not have a unitary explanation.

We used traps baited with fruit to examine how the temporal variation of butterflies within primary forest in Sabah, Borneo differed between species. In addition, we compared patterns of temporal variation in primary and selectively logged forest, and we tested the hypothesis that selective logging has different recorded impacts on species diversity of adults during the wet monsoon period and the drier remaining half of the year. Species of Satyrinae and Morphinae had significantly less-restricted flight periods than did species of Nymphalinae and Charaxinae, which were sampled mainly during the drier season, especially in primary forest. Species diversity of adults was significantly higher during the drier season in primary forest, but did not differ between seasons in logged forest. As a consequence, logging had opposite recorded impacts on diversity during wetter and drier seasons: primary forest had significantly higher diversity than logged forest during the drier season but significantly lower diversity than logged forest during the wetter monsoon season. The results of this study have important implications for the assessment of biodiversity in tropical rain forests, particularly in relation to habitat disturbance: short-term assessments that do not take account of seasonal variation in abundance are likely to produce misleading results, even in regions where the seasonal variation in rainfall is not that great.

1. The impacts of habitat disturbance on biodiversity within tropical forests are an area of current concern but are poorly understood and difficult to predict. This is due in part to a poor understanding of how species respond to natural variation in environmental conditions within primary forest and how these conditions alter following anthropogenic disturbance. Within this context, the main aim of this study was to test the hypothesis that the gap and shade preferences of fruit-feeding butterflies in primary forest in northern Borneo can be used to predict species' responses to selective logging and thus explain changes in diversity and geographical distinctness in relation to habitat disturbance. 2. Overall, there was little difference in butterfly diversity between primary forest and forest that had been selectively logged 10-12 years previously. In contrast, there were marked differences in the composition of the butterfly assemblages in the two habitats, which were strongly associated with species' gap preferences and geographical distributions. In Satyrinae and Morphinae, those species with higher shade preferences and narrower geographical distributions were most adversely affected by logging, whereas cosmopolitan species with high light preferences benefited from logging. In Nymphalinae and Charaxinae the opposite was observed: those species with wider geographical distributions were adversely affected and those species with relatively restricted distributions were more common in logged forest. 3. These changes in butterfly assemblages were associated with changes in vegetation structure following selective logging, which resulted in much lower habitat heterogeneity with less dense shade and fewer open gaps in logged forest. Areas of dense shade, which were more common in unlogged forest, supported species of Satyrinae and Morphinae with restricted geographical distributions, whereas open gaps, which were also more common in unlogged forest, attracted widespread species of Nymphalinae and Charaxinae. These butterfly-habitat associations in primary forest explain the opposite responses of the two groups of butterflies to selective logging. 4. Synthesis and applications. This study highlights the need to sample at a sufficiently large spatial scale to account for impacts of disturbance on heterogeneity in forest environments. It also demonstrates how understanding the responses of species to natural variation in environmental conditions within undisturbed forest is crucial to interpreting responses of species to anthropogenic habitat modification. The results further indicate that selectively logged forests can make an important contribution to the conservation of tropical biodiversity, provided that they are managed in a way that maintains environmental heterogeneity.

We investigated the effects of rainfall on the distribution and abundance of the satyrine butterfly Ragadia makuta in selectively logged and unlogged forest on Borneo. In 1997-98, there was a severe El Niño-Southern Oscillation (ENSO) drought, and annual surveys over a 4-y period showed that abundance of R. makuta was greatly reduced during the drought, but that populations quickly recovered after it. Monthly surveys over a 12-mo period of typical rainfall showed that high rainfall in the month preceding surveys significantly reduced butterfly abundance. Butterfly abundance and distribution did not differ between selectively logged and unlogged areas in either monthly or annual surveys and there was no difference between selectively logged and unlogged areas in the pattern of post-drought recovery. These results indicate that the abundance of R. makuta was significantly reduced both after high rainfall and during severe drought, but that these impacts were short-lived and were not affected by habitat disturbance. ENSO droughts on Borneo naturally often lead to widespread forest fires and thus impacts of ENSO events for butterflies are more likely to be due to indirect effects of habitat loss, rather than direct effects of drought on butterfly population dynamics.

The diversity of macromoths in the various fast-growing exotic tree plantations and natural secondary forest in Brumas, Sabah, Malaysia was assessed by means of an annual cycle of light-trap samples. The moth diversity in the forest plantations, viz. Acacia mangium, Gmelina arborea, Paraserianthes (=Albizia) falcataria, Pinus caribaea, and in particular Eucalyptus deglupta, was unexpectedly high. Eucalyptus deglupta showed moth diversity as high as that in the natural secondary forest, a finding attributed to the fact that the E. deglupta plantation had a very diverse understorey both in terms of plant species (secondary regrowth species) and architecture, and thus supported a more diverse moth fauna. Subsidiary samples showed that primary natural forest in the neighbouring Danum Valley does not show higher moth diversity compared to the disturbed forest habitats in Brumas, though moth diversity at Danum is lower than that recorded in other Bornean primary forests. The value of plantation forests for conservation of invertebrate diversity is discussed, together with the implications for plantation management strategy.

Physiological research suggests that tropical insects are particularly sensitive to temperature, but information on their responses to climate change has been lacking—even though the majority of all terrestrial species are insects and their diversity is concentrated in the tropics. Here, we provide evidence that tropical insect species have already undertaken altitude increases, confirming the global reach of climate change impacts on biodiversity. In 2007, we repeated a historical altitudinal transect, originally carried out in 1965 on Mount Kinabalu in Borneo, sampling 6 moth assemblages between 1,885 and 3,675 m elevation. We estimate that the average altitudes of individuals of 102 montane moth species, in the family Geometridae, increased by a mean of 67 m over the 42 years. Our findings indicate that tropical species are likely to be as sensitive as temperate species to climate warming, and we urge ecologists to seek other historic tropical samples to carry out similar repeat surveys. These observed changes, in combination with the high diversity and thermal sensitivity of insects, suggest that large numbers of tropical insect species could be affected by climate warming. As the highest mountain in one of the most biodiverse regions of the world, Mount Kinabalu is a globally important refuge for terrestrial species that become restricted to high altitudes by climate warming.

Despite a large increase in the area of selectively logged tropical forest worldwide, the carbon stored in deadwood across a tropical forest degradation gradient at the landscape scale remains poorly documented. Many carbon stock studies have either focused exclusively on live standing biomass or have been carried out in primary forests that are unaffected by logging, despite the fact that coarse woody debris (deadwood with ≥10 cm diameter) can contain significant portions of a forest’s carbon stock. We used a field-based assessment to quantify how the relative contribution of deadwood to total above-ground carbon stock changes across a disturbance gradient, from unlogged old-growth forest to severely degraded twice-logged forest, to oil palm plantation. We measured in 193 vegetation plots (25 × 25 m), equating to a survey area of >12 ha of tropical humid forest located within the Stability of Altered Forest Ecosystems Project area, in Sabah, Malaysia. Our results indicate that significant amounts of carbon are stored in deadwood across forest stands. Live tree carbon storage decreased exponentially with increasing forest degradation 7–10 years after logging while deadwood accounted for >50% of above-ground carbon stocks in salvage-logged forest stands, more than twice the proportion commonly assumed in the literature. This carbon will be released as decomposition proceeds. Given the high rates of deforestation and degradation presently occurring in Southeast Asia, our findings have important implications for the calculation of current carbon stocks and sources as a result of human-modification of tropical forests. Assuming similar patterns are prevalent throughout the tropics, our data may indicate a significant global challenge to calculating global carbon fluxes, as selectively-logged forests now represent more than one third of all standing tropical humid forests worldwide.

We used microscopic pollen analysis to investigate the diversity and similarity of pollen diets of six colonies of stingless bees (Apidae; Meliponini) located within one monospecific (three colonies of Trigona collina) and one mixed nesting aggregation (one colony of T. collina, and one colony of each of the close relatives T. melina and T. melanocephala) in lowland tropical rain forest in Sabah, Malaysia. Samples of 20 corbicular loads, collected six times over a period of three months from each colony, contained a total of 74 different morphotypes of pollen grains with an average between 4.7 to 8.5 per sample for the different colonies. In an analysis on total diet composition intraspecific similarity was much greater than interspecific similarity. The focal colony of Trigona collina from the mixed aggregation distinctly clustered according to species rather than nest location, suggesting that some interspecific resource partitioning occurs. The sampling period was accompanied by a drastic increase in flowering activity as evidenced by data from a flower phenology transect. At times of limited flowering similarity of pollen diets was generally low, both within and between species. It is hypothesized that this is so because bees are forced to forage from scattered subsets of flower patches spread out over a large foraging range. In times of increased flowering pollen diet similarity significantly increased between colonies of the monospecific aggregation, presumably because colonies concentrated on more profitable sources in closer proximity. In contrast, similarity remained low within the mixed aggregation, suggesting that innate differences in foraging preferences precluded any effect of diet convergence.

The diversity of beetle assemblages in different habitat types (primary forest, logged forest, acacia plantation and oil palm plantation) in Sabah, Malaysia was investigated using three different methods based on habitat levels (Winkler sampling, flight-interception-trapping and mist-blowing). The overall diversity was extremely high, with 1711 species recorded from only 8028 individuals and 81 families (115 family and subfamily groups). Different degrees of environmental changes had varying effects on the beetle species richness and abundance, with oil palm plantation assemblage being most severely affected, followed by acacia plantation and then logged forest. A few species became numerically dominant in the oil palm plantation. In terms of beetle species composition, the acacia fauna showed much similarity with the logged forest fauna, and the oil palm fauna was very different from the rest. The effects of environmental variables (number of plant species, sapling and tree densities, amount of leaf litter, ground cover, canopy cover, soil pH and compaction) on the beetle assemblage were also investigated. Leaf litter correlated with species richness, abundance and composition of subterranean beetles. Plant species richness, tree and sapling densities correlated with species richness, abundance and composition of understorey beetles while ground cover correlated only with the species richness and abundance of these beetles. Canopy cover correlated only with arboreal beetles. In trophic structure, predators represented more than 40% of the species and individuals. Environmental changes affected the trophic structure with proportionally more herbivores (abundance) but fewer predators (species richness and abundance) in the oil palm plantation. Biodiversity, conservation and practical aspects of pest management were also highlighted in this study.