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Habitat degradation and climate change are thought to be altering the distributions and abundances of animals and plants throughout the world, but their combined impacts have not been assessed for any species assemblage 1 , 2 , 3 , 4 . Here we evaluated changes in the distribution sizes and abundances of 46 species of butterflies that approach their northern climatic range margins in Britain—where changes in climate and habitat are opposing forces. These insects might be expected to have responded positively to climate warming over the past 30 years, yet three-quarters of them declined: negative responses to habitat loss have outweighed positive responses to climate warming. Half of the species that were mobile and habitat generalists increased their distribution sites over this period (consistent with a climate explanation), whereas the other generalists and 89% of the habitat specialists declined in distribution size (consistent with habitat limitation). Changes in population abundances closely matched changes in distributions. The dual forces of habitat modification and climate change are likely to cause specialists to decline, leaving biological communities with reduced numbers of species and dominated by mobile and widespread habitat generalists.

We analyse distribution records for 51 British butterfly species to investigate altitudinal and latitudinal responses to twentieth century climate warming. Species with northern and/or montane distributions have disappeared from low elevation sites and colonized sites at higher elevations during the twentieth century, consistent with a climate explanation. We found no evidence for a systematic shift northwards across all species, even though 11 out of 46 southerly distributed species have expanded in the northern part of their distributions. For a subset of 35 species, we model the role of climate in limiting current European distributions and predict potential future distributions for the period 2070-2099. Most northerly distributed species will have little opportunity to expand northwards and will disappear from areas in the south, resulting in reduced range sizes. Southerly distributed species will have the potential to shift northwards, resulting in similar or increased range sizes. However, 30 out of 35 study species have failed to track recent climate changes because of lack of suitable habitat, so we revised our estimates accordingly for these species and predicted 65% and 24% declines in range sizes for northern and southern species, respectively. These revised estimates are likely to be more realistic predictions of future butterfly range sizes.

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.

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.

Despite growing concern, no consensus has emerged over the effects of habitat modification on species diversity in tropical forests. Even for comparatively well-studied taxa such as Lepidoptera, disturbance has been reported to increase and decrease diversity with approximately equal frequency. Species diversity within landscapes depends on the spatial scale at which communities are sampled, and the effects of disturbance in tropical forests have been studied at a wide range of spatial scales. Yet the question of how disturbance affects diversity at different spatial scales has not been addressed. We reanalyzed data from previous studies to examine the relationship between spatial scale and effects of disturbance on tropical-forest Lepidoptera. Disturbance had opposite effects on diversity at large and small scales: as scale decreased, the probability of a positive effect of disturbance on diversity increased. We also explicitly examined the relationship between spatial scale and the diversity of butterflies in selectively logged and unlogged forest in Maluku Province, Indonesia. Species richness increased with spatial scale in both logged and unlogged forest, but at a significantly faster rate in unlogged forest, whereas species evenness increased with scale in unlogged forest but did not increase with scale in logged forest. These data indicate that the effects of habitat modification on species diversity are heavily scale-dependent. As a result, recorded effects of disturbance were strongly influenced by the spatial scale at which species assemblages were sampled. Future studies need to account for this by explicitly examining the effects of disturbance at a number of different spatial scales. A further problem arises because the relationship between scale and diversity is likely to differ among taxa in relation to mobility. This may explain to some extent why the measured effects of disturbance have differed between relatively mobile and immobile taxa.

Mean global temperatures have risen this century, and further warming is predicted to continue for the next 50–100 years 1 , 2 , 3 . Some migratory species can respond rapidly to yearly climate variation by altering the timing or destination of migration 4 , but most wildlife is sedentary and so is incapable of such a rapid response. For these species, responses to the warming trend should be slower, reflected in poleward shifts of the range. Such changes in distribution would occur at the level of the population, stemming not from changes in the pattern of individuals' movements, but from changes in the ratios of extinctions to colonizations at the northern and southern boundaries of the range. A northward range shift therefore occurs when there is net extinction at the southern boundary or net colonization at the northern boundary. However, previous evidence has been limited to a single species 5 or to only a portion of the species' range 6 , 7 . Here we provide the first large-scale evidence of poleward shifts in entire species' ranges. In a sample of 35 non-migratory European butterflies, 63% have ranges that have shifted to the north by 35–240 km during this century, and only 3% have shifted to the south.

A suite of idealised ensemble experiments are used to investigate the sensitivity of the Southern Hemisphere atmospheric circulation to the location of SST anomalies over the tropical Pacific Ocean. Of particular interest is the response of South American rainfall during austral summer. The experiments reveal an approximately opposite response in rainfall over South America when the tropical SST forcing is applied over the western and eastern Pacific Ocean, respectively. The contrasting tropical rainfall conditions are due primarily to the displaced Walker circulation anomalies. Further south, the atmospheric circulation over the South American subtropics is modulated by teleconnection patterns that appear as a wave train. The resulting circulation manifests as an anomalous cyclone over central‐eastern South America which in turn leads to a northward displacement of the westerly moisture transport when the SST forcing is located further west. The opposite pattern occurs when the SST forcing is applied to the eastern equatorial Pacific.

The impacts of inter‐El Niño events on South American circulation during austral summer are investigated using observations and an atmospheric general circulation model (AGCM). The AGCM was forced with sea surface temperature (SST) anomalies in the tropical Pacific for the two El Niño events of 1997/1998 (EN97) and 2002/2003 (EN02). The strong eastern Pacific SST anomaly of EN97 resulted in a typical displacement of the Walker circulation, causing a decrease in precipitation across the north of South America. A strengthened low‐level jet (LLJ) east of the Andes during EN97 enhanced the moisture transport from low latitudes to the subtropics, leading to intensified precipitation over southeastern South America. The simulated circulation in EN02 reveals a weakened LLJ and anomalous convergence of moisture over eastern South America, which can be attributed to a displacement of the Pacific‐South American (PSA) mode in response to the different location of the heat sources along the tropical Pacific Ocean.

1. There is an urgent need to understand the impacts of anthropogenic habitat disturbance on biodiversity in tropical forests, but no consensus has yet emerged. We reviewed the literature for the most frequently studied taxon (birds, 37 studies) and found that increased and decreased diversity in response to disturbance (selective logging and shifting agriculture) were reported with approximately equal frequency. 2. The spatial scale at which studies were carried out significantly affected the reported response to disturbance: studies where disturbed and undisturbed habitats were sampled at large spatial scales were more likely to report increased diversity following disturbance, whereas studies that sampled habitats at small spatial scales were more likely to report decreased diversity. These results were not a consequence of sampling method: we divided the studies into those using capture methods and those using observation methods and the same result was obtained when the analysis was restricted to only those studies using observation methods. 3. Previously, we have shown that reported impacts of disturbance on Lepidoptera are also affected by the spatial scale of study. We reviewed the Lepidoptera literature published since then and showed that all 12 new studies conformed to the predicted pattern. 4. While sampling scale significantly affected the reported responses of both birds and Lepidoptera, there were opposite effects of scale in the two taxa: large-scale bird studies and small-scale Lepidoptera studies were more likely to report increased diversity following disturbance. Bird studies were generally carried out at larger spatial scales than those of Lepidoptera and these opposite impacts of scale were probably due to a nonlinear effect of habitat disturbance on habitat heterogeneity at different spatial scales. 5. Synthesis and applications. The rapid loss and degradation of tropical forests means that an understanding of the general patterns of responses of species to habitat disturbance is urgently needed. However, there has been little discussion of the most appropriate methods to ensure comparability of results between studies. Data presented here indicate that the spatial scale of sampling chosen in studies has a marked effect on the results obtained, and future studies need to account for this by examining explicitly the effects of disturbance at different spatial scales. The effect of spatial scale differs between taxa, and this may explain why the search for indicator taxa of disturbance effects has so far proved elusive.

Tropical forest gaps are ephemeral and patchily distributed within forest areas and have very different light environments compared with closed-canopy forest. We used fruit-baited traps to investigate if gaps are exploited by more opportunistic butterfly species compared with closed-canopy forest. Gaps supported a higher diversity of butterflies in terms of species evenness but closed-canopy sites contained species with more restricted geographical distributions. There was little similarity between the assemblages of butterflies trapped in the canopy and those in either gap or closed-canopy sites, but the greater similarity was with gaps, and increased diversity in gaps was partly due to canopy species turning up in gaps. Dispersal rates (as measured by recapture rates) were higher in gaps and there was evidence that butterflies in gaps had relatively larger and broader thoraxes, indicating a flight morphology adapted for faster flight. These results support the notion of a distinctive gap fauna comprising more widespread, mobile species. Habitat modification that opens up the canopy is likely to result in an increase in these widespread species and a decline in understorey species with restricted distributions.