Summary1. 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.

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.

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