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Aim Climatic changes throughout the Pleistocene have strongly modified species distributions. We examine how these range shifts have affected the genetic diversity of a montane butterfly species and whether the genetic diversity in the extant populations is threatened by future climate change. Location Europe. Taxon Erebia epiphron Lepidoptera: Nymphalidae. Methods We analyzed mtDNA to map current genetic diversity and differentiation of E. epiphron across Europe to identify population refugia and postglacial range shifts. We used species distribution modeling (SDM) to hindcast distributions over the last 21,000 years to identify source locations of extant populations and to project distributions into the future (2070) to predict potential losses in genetic diversity. Results We found substantial genetic diversity unique to specific regions within Europe (total number of haplotypes = 31, number of unique haplotypes = 27, H d = 0.9). Genetic data and SDM hindcasting suggest long‐term separation and survival of discrete populations. Particularly, high rates of unique diversity in postglacially colonized sites in England (H d = 0.64) suggest this population was colonized from a now extinct cryptic refugium. Under future climate change, SDMs predict loss of climate suitability for E. epiphron, particularly at lower elevations (, The genetic diversification of cold‐adapted mountain species, such as E. epiphron, has been shaped by Pleistocene glaciations, resulting in unique genetic diversity in isolated populations. The unique genetic diversity in mountain and cold‐adapted species is at under future climate warming, and we predict E. epiphron will lose 38%–64% of its range in the future, resulting in the loss of genetic diversity, reducing its ability to adapt.

1. Human‐modified tropical landscapes are often invaded by exotic plant species, but relatively few species are able to colonise remnant areas of rainforest embedded within such landscapes. The functional traits of successful invaders of natural versus anthropogenic habitats are poorly known, especially in tropical regions, and identifying such traits provides insight into the mechanisms that drive invasion. Here, we examine the invasion of tropical rainforest remnants along a disturbance gradient, within a human‐modified agricultural landscape, and determine whether exotic species that invade these forests are selected according to particular traits. 2. We surveyed the occurrence of 18 exotic species along 100‐m transects in four habitats—oil palm road, forest‐oil palm edges and disturbed and intact forest within rainforest remnants—at 21 sites across Sabah, Malaysian Borneo. We collated data on four functional traits relevant to the barriers plants encounter when colonising new environments (e.g. dispersal and persistence) and tested whether trait filtering occurs during invasion of rainforest remnants. 3. Exotic species richness declined significantly from oil palm (mean 9.2 species per transect) to forest edge (7.8 species) to inside rainforest remnants (3.1 species in disturbed forest), and only one species, Clidemia hirta, invaded intact forest. Exotic communities within rainforest remnants had long‐distance (vertebrate) dispersal, were woodier and had taller maximum heights, compared to those found in oil palm. For each trait, the community‐weighted mean for the forest edge community was intermediate between oil palm and disturbed forest, suggesting trait filtering during the invasion of rainforest remnants. 4. Our study provides strong evidence that trait filtering occurs during invasion from human‐modified agricultural habitats into previously disturbed forests via the forest edge. Successful invasion of rainforest remnants requires relatively long‐distance dispersal, in particular by vertebrates, as well as traits that are more similar to those of native forest species (i.e. tall and woody), making these exotic species more able to compete and persist in that environment. Our results show that disturbed tropical rainforests with open canopies are susceptible to invasion and highlight the traits of exotic species which can invade rainforest habitats, and which may pose a threat to regenerating tropical rainforests.

Oligophagous insects can consume a wide range of different host plant species, but how these host plants vary in their metabolite compositions and the extent to which this variation affects the biochemistry of the insect herbivores is largely unknown. An understanding of how defensive metabolites from plants are processed by insects may help us develop more effective pesticides. We studied the interactions between the oligophagous insect herbivore Pieris rapae (Lepidoptera: Pieridae) and five species of its larval host plants (family Brassicaceae and Cleomaceae) by examining untargeted metabolic fingerprints of the plants and the larval herbivores feeding on them. Visualisation of the metabolic fingerprints of the different host plant species showed highly distinctive clusters in the PCA-X score plots. Larvae could also be distinguished based on the species of host plant they fed on but clusters overlapped to a greater extent. The fingerprints of larvae feeding on Cleome spinosa plants were most distinctive due to a large group of abundant metabolites also found in high abundance in C. spinosa, but not in the other host plants examined. We conclude that host plants influence the biochemistry of their larval herbivores, and that some metabolites are conserved from one trophic level to the next.

Populations undergoing rapid climate-driven range expansion experience distinct selection regimes dominated both by increased dispersal at the leading edges and steep environmental gradients. Characterisation of traits associated with such expansions provides insight into the selection pressures and evolutionary constraints that shape demographic and evolutionary responses. Here we investigate patterns in three components of wing morphology (size, shape, colour) often linked to dispersal ability and thermoregulation, along latitudinal gradients of range expansion in the Speckled Wood butterfly (Pararge aegeria) in Britain (two regions of expansion in England and Scotland). We measured 774 males from 54 sites spanning 799 km with a 10-year mean average temperature gradient of 4 °C. A geometric morphometric method was used to investigate variation in size and shape of forewings and hindwings; colour, pattern, and contrast of the wings were examined using a measure of lightness (inverse degree of melanism). Overall, wing size increased with latitude by ∼2% per 100 km, consistent with Bergmann’s rule. Forewings became more rounded and hindwings more elongated with history of colonisation, possibly reflecting selection for increased dispersal ability. Contrary to thermal melanism expectations, wing colour was lighter where larvae developed at cooler temperatures and unrelated to long-term temperature. Changes in wing spot pattern were also detected. High heterogeneity in variance among sites for all of the traits studied may reflect evolutionary time-lags and genetic drift due to colonisation of new habitats. Our study suggests that temperature-sensitive plastic responses for size and colour interact with selection for dispersal traits (wing size and shape). Whilst the plastic and evolutionary responses may in some cases act antagonistically, the rapid expansion ofP. aegeriaimplies an overall reinforcing effect between these two mechanisms.

Trends in insect abundance are well established in some datasets, but far less is known about how abundance measures translate into biomass trends. Moths (Lepidoptera) provide particularly good opportunities to study trends and drivers of biomass change at large spatial and temporal scales, given the existence of long‐term abundance datasets. However, data on the body masses of moths are required for these analyses, but such data do not currently exist.To address this data gap, we collected empirical data in 2018 on the forewing length and dry mass of field‐sampled moths, and used these to train and test a statistical model that predicts the body mass of moth species from their forewing lengths (with refined parameters for Crambidae, Erebidae, Geometridae and Noctuidae).Modeled biomass was positively correlated, with high explanatory power, with measured biomass of moth species (R 2 = 0.886 ± 0.0006, across 10,000 bootstrapped replicates) and of mixed‐species samples of moths (R 2 = 0.873 ± 0.0003), showing that it is possible to predict biomass to an informative level of accuracy, and prediction error was smaller with larger sample sizes.Our model allows biomass to be estimated for historical moth abundance datasets, and so our approach will create opportunities to investigate trends and drivers of insect biomass change over long timescales and broad geographic regions., Assessment of trends in insect biomass is hampered by a lack of data. To address this, we developed a method to estimate the biomass of historical samples of moths, where abundance and species identity have been recorded. Estimates made using this method explain >85% of total variation in sample biomass, allowing biomass to be predicted to an informative level of accuracy.

Many areas of tropical rainforest have been fragmented and the habitat quality of fragments is often poor. For example, on Borneo, many forest fragments are highly degraded by repeated logging of Dipterocarpaceae trees prior to fragmentation, and we examined the viability of enrichment planting as a potential management tool to enhance the conservation value of these forest fragments. We planted seedlings of three dipterocarp species with contrasting light demands and tolerances (Parashorea malaanonan (light demander), Dryobalanops lanceolata (intermediate), Hopea nervosa (shade tolerant)) in eight forest fragment sites (3–3529 ha), and compared seedling performance with four sites in continuous forest. Eighteen months after planting, survival rates of seedlings were equally high in fragment sites (mean survival = 63 %), and in continuous forest sites (mean survival = 68 %). By contrast, seedling growth and herbivory rates were considerably higher in fragments (by 60 % for growth and 45 % for herbivory) associated with higher light environments in degraded forest fragments compared with continuous forest sites. Among the three study species, H. nervosa seedlings had the highest survival rates overall, and P. malaanonan seedlings generally grew fastest and suffered highest herbivory rates. There were no interactions between species performance and the effects of fragment site area, forest structure or soil characteristics of sites suggesting that the three species responded similarly to fragmentation effects. High survival of planted seedlings implies that enrichment planting would be a successful forest management strategy to improve forest quality, and hence conservation value, of fragments.

Previously extensive tracts of primary rain forest have been degraded by human activities, and we examined how the effects of forest disturbance arising from habitat fragmentation and commercial selective logging affected ecosystem functioning in these habitats by studying leaf litter decomposition rates in litter bags placed on the forest floor. The rain forests of Borneo are dominated by trees from the family Dipterocarpaceae, and we compared leaf litter decomposition rates of three dipterocarp species at eight forest fragment sites (area 3-3529 ha) that had different histories of disturbance pre-fragmentation: four fragments had been selectively logged prior to fragmentation and four had been formed from previously undisturbed forest. We compared these logged and unlogged forest fragments with sites in continuous forest that had been selectively logged (two sites) and fully protected and undisturbed (two sites). After 120 d, undisturbed continuous forest sites had the fastest rates of decomposition (52% mass loss). Forest fragments formed from unlogged forest (32% mass loss) had faster decomposition rates than logged forest fragments (28% mass loss), but slower rates than continuous logged forest (39% mass loss). Leaves of a light-demanding species (Parashorea malaanonan) decomposed faster than those of a shade-tolerant species (Hopea nervosa), but decomposition of all three dipterocarp species that we studied responded similarly to logging and fragmentation effects. Reduced decomposition rates in logged and fragmented forest sites may affect nutrient cycling and thus have detrimental consequences for forest regeneration. Conservation management to improve forest quality should be a priority, particularly in logged forest fragments.

1. The ability of species' to undergo climate-driven range shifts across fragmented landscapes depends on their dispersal ability as well as the structure of the landscape. For species' range shifts to occur, individuals must first leave suitable habitat to seek new habitat; this is likely to depend on the rate of movement of individuals within habitat and the likelihood that a boundary is crossed, once it is encountered. For three species of butterfly with contrasting histories of recent range expansion, we examined the propensity of individuals to move within a habitat and their responses to habitat boundaries. 2. We quantified the extent to which Plebejus argus (Linnaeus) (a declining habitat specialist), Aricia agestis (Schiffermuller) (an expanding generalist) and Polymmatus icarus (Rottemburg) (a geographically ubiquitous generalist) crossed habitat boundaries into unsuitable habitat and moved within suitable habitat. The observed movement was then related to individual and environmental conditions. 3. Species differed in their activity levels in accordance within their recent distribution patterns (P. icarus > A. agestis > P. argus). Our results for P. argus suggest that movement may be motivated by nectar-seeking, and that males generally move more than females. All three species tended to avoid crossing habitat boundaries; however the proportion of individuals crossing habitat boundaries did not differ significantly among species. 4. We conclude that levels of activity within a habitat, which will affect the frequency with which individuals encounter habitat boundaries, rather than behavioural responses to the boundaries, may be important drivers of distribution change.

Conversion of natural habitats to agriculture reduces species richness, particularly in highly diverse tropical regions, but its effects on species composition are less well-studied. The conversion of rain forest to oil palm is of particular conservation concern globally, and we examined how it affects the abundance of birds, beetles, and ants according to their local population size, body size, geographical range size, and feeding guild or trophic position. We re-analysed data from six published studies representing 487 species/genera to assess the relative importance of these traits in explaining changes in abundance following forest conversion. We found consistent patterns across all three taxa, with large-bodied, abundant forest species from higher trophic levels, declining most in abundance following conversion of forest to oil palm. Best-fitting models explained 39–66 % of the variation in abundance changes for the three taxa, and included all ecological traits that we considered. Across the three taxa, those few species found in oil palm tended to be small-bodied species, from lower trophic levels, that had low local abundances in forest. These species were often hyper-abundant in oil palm plantations. These results provide empirical evidence of consistent responses to land-use change among taxonomic groups in relation to ecological traits.

Conversion of natural forest to oil palm plantations is a major threat to biodiversity in Southeast Asia. The retention of natural forest habitats within plantations has been proposed as a method to reduce biodiversity losses in agricultural areas, and we examined whether forest areas resulted in spillover of species into adjacent oil palm plantations. We sampled ants and butterflies along two 2-km transects across an ecotone from plantation into adjacent forest in Sabah, Malaysian Borneo. Species richness of both taxa was reduced in plantations, but to a greater extent in butterflies (54% reduction) than in ants (25% reduction). Butterfly diversity increased in plantations with increasing proximity to forest primarily due to spillover of ‘vagrant’ forest species (whose larval host plants do not occur in plantations), although richness of species that could potentially breed in plantations also increased near to forest. By contrast, ants showed no spillover effects and were less sensitive to land-use changes, with much higher levels of similarity in species assemblages across habitats than for butterflies. Our results for butterflies suggest that despite the negative impacts of plantations on diversity, proximity to forest could improve diversity in adjacent plantations for some taxa. Spillover of forest species implies that retaining forest areas within plantations may be important for facilitating dispersal of some species through the landscape.

Many insects undertake long-distance migrations to exploit seasonally variable conditions at high latitudes, but the mechanisms used by migrants to select and maintain beneficial flight headings are poorly understood. Using computerized flight simulators, we performed controlled experiments to test the ability of an obligate migrant butterfly (Vanessa cardui) to orient in seasonally advantageous directions (i.e. northwards in spring and southwards in autumn). We also investigated the compass mechanism used to select and maintain these headings. Laboratory-reared autumn-generation butterflies flown in the U.K. displayed a highly significant mean orientation towards the south-southwest, consistent with return migration to winter breeding sites. However, seasonally adaptive flight headings were not observed in wild-caught adults flown at the same time. Spring-generation adults caught in Gibraltar (presumed to be migrating from winter breeding sites in North Africa into Europe) showed no evidence of northward flight headings, but produced a wide scatter of flight headings with a mean direction towards the west. Butterflies flown in the simulators when the sky was not visible produced a random scatter of flight headings and less-directed flight tracks, providing evidence that migrating V. cardui use a sun compass to select and maintain their flight headings. However, when butterflies were subjected to a 6 h clock shift, no change in orientation was observed relative to the control group. Field evidence for a return migration in autumn by V. cardui is surprisingly scarce in the literature, but we conclude that the species does attempt such southward movements and that individuals use a sun compass to select their migratory heading.

Recent climatic change in temperate regions has been rapid and there is mounting speculation that species are failing to keep track of suitable climate, perhaps necessitating assisted colonization for some species. An inability to spread into new areas may result in large reductions in species’ ranges in the future, and threaten the survival of some species. Here we use “speciesclimate” models to predict suitable sites for introductions beyond current range margins, using two U.K. butterfly species. We introduced Melanargia galathea (marbled white) and Thymelicus sylvestris (small skipper) into two sites in northern England, ∼65 and ∼35 km beyond their then-range margins, respectively, to sites that were predicted to be climatically suitable and that appeared to contain suitable habitat for the species. Both introduced populations grew and expanded their range over 6 years (2001–2006; still thriving in 2008), suggesting the existence of a colonization lag and providing evidence that well-planned assisted colonization can be successful. We suggest that assisted colonization may be a feasible and cost-effective means of enabling certain species to track climatic change.

Many areas of rain forest now exist as habitat fragments, and understanding the impacts of fragmentation is important for determining the viability of populations within forest remnants. We investigated impacts of forest fragmentation on genetic diversity in the butterfly Mycalesis orseis (Satyrinae) in Sabah (Malaysian Borneo). We investigated mtDNA diversity in 90 individuals from ten forest sites typical of the sizes of forest remnants that currently exist in the region. Nucleotide diversity declined with increasing isolation of remnants, but there was no effect of remnant size or population size, and haplotype diversity was similar among sites. Thus, approximately 50 y after forest fragmentation, few changes in genetic diversity were apparent and remnants apparently supported genetically viable populations of this butterfly. Many studies have shown that responses of species to habitat fragmentation usually follow a time delay, and so we developed a Monte Carlo simulation model to investigate changes in genetic diversity over time in small remnants. Model output indicated a substantial time delay (> 100 y) between fragmentation and genetic erosion, suggesting that, in the smallest study remnants, an increased risk of extinction from reduced genetic diversity is likely in the longer term.

1. Species would be expected to shift northwards in response to current climate warming, but many are failing to do so because of fragmentation of breeding habitats. Dispersal is important for colonisation and an individual-based spatially explicit model was developed to investigate impacts of habitat availability on the evolution of dispersal in expanding populations. Model output was compared with field data from the speckled wood butterfly Pararge aegeria, which currently is expanding its range in Britain. 2. During range expansion, models simulated positive linear relationships between dispersal and distance from the seed location. This pattern was observed regardless of quantity (100% to 10% habitat availability) or distribution (random vs. gradient distribution) of habitat, although higher dispersal evolved at expanding range margins in landscapes with greater quantity of habitat and in gradient landscapes. Increased dispersal was no longer evident in any landscape once populations had reached equilibrium; dispersal values returned to those of seed populations. However, in landscapes with the least quantity of habitat, reduced dispersal (below that of seed populations) was observed at equilibrium. 3. Evolutionary changes in adult flight morphology were examined in six populations of P. aegeria along a transect from the distribution core to an expanding range margin in England (spanning a latitudinal distance of >200 km). Empirical data were in agreement with model output and showed increased dispersal ability (larger and broader thoraxes, smaller abdomens, higher wing aspect ratios) with increasing distance from the distribution core. Increased dispersal ability was evident in populations from areas colonised >30 years previously, although dispersal changes were generally evident only in females. 4. Evolutionary increases in dispersal ability in expanding populations may help species track future climate changes and counteract impacts of habitat fragmentation by promoting colonisation. However, at the highest levels of habitat loss, increased dispersal was less evident during expansion and reduced dispersal was observed at equilibrium indicating that, for many species, continued habitat fragmentation is likely to outweigh any benefits from dispersal.

1. Some species have expanded their ranges during recent climate warming and the availability of breeding habitat and species' dispersal ability are two important factors determining expansions. The exploitation of a wide range of larval host plants should increase an herbivorous insect species' ability to track climate by increasing habitat availability. Therefore we investigated whether the performance of a species on different host plants changed towards its range boundary, and under warmer temperatures. 2. We studied the polyphagous butterfly Polygonia c-album, which is currently expanding its range in Britain and apparently has altered its host plant preference from Humulus lupulus to include other hosts (particularly Ulmus glabra and Urtica dioica). We investigated insect performance (development time, larval growth rate, adult size, survival) and adult flight morphology on these host plants under four rearing temperatures (18-28.5 degrees C) in populations from core and range margin sites. 3. In general, differences between core and margin populations were small compared with effects of rearing temperature and host plant. In terms of insect performance, host plants were generally ranked U. glabra > or = U. dioica > H. lupulus at all temperatures. Adult P. c-album can either enter diapause or develop directly and higher temperatures resulted in more directly developing adults, but lower survival rates (particularly on the original host H. lupulus) and smaller adult size. 4. Adult flight morphology of wild-caught individuals from range margin populations appeared to be related to increased dispersal potential relative to core populations. However, there was no difference in laboratory reared individuals, and conflicting results were obtained for different measures of flight morphology in relation to larval host plant and temperature effects, making conclusions about dispersal potential difficult. 5. Current range expansion of P. c-album is associated with the exploitation of more widespread host plants on which performance is improved. This study demonstrates how polyphagy may enhance the ability of species to track climate change. Our findings suggest that observed differences in climate-driven range shifts of generalist vs. specialist species may increase in the future and are likely to lead to greatly altered community composition.

Highlight: Herbivore attacks on three species of Brassicales result in different metabolic fingerprints. Furthermore, there is no overlap among the three plants in the individual metabolites that are induced by herbivory., Studying the biochemical responses of different plant species to insect herbivory may help improve our understanding of the evolution of defensive metabolites found in host plants and their role in plant–herbivore interactions. Untargeted metabolic fingerprints measured as individual mass features were used to compare metabolite reactions in three Brassicales host-plant species (Cleome spinosa, Brassica oleracea, and Lunaria annua) to larval herbivore attack (Pieris rapae; Lepidoptera). Principal component analyses of metabolic fingerprints were able to distinguish among the three plant species and between uneaten control plants and plants that had been eaten. A large number of mass features (1186, 13% of mass features measured in control plants) were common to the three plant species. However, there were few similarities in the mass features that were induced (i.e. changed in abundance) following herbivory. Of the 87 and 68 induced mass features in B. oleracea and C. spinosa, respectively, there were only three that were induced in both plant species. By contrast, L. annua only had one mass feature induced by herbivory, and this was not induced in the other two plant species. The growth of the P. rapae larvae was poorer on the host plant L. annua than on B. oleracea and C. spinosa. The absence of common metabolites among the plants meant these induced responses could not be related to the performance of the herbivore. Thus, the response to herbivory by the same herbivore in these three host plants has evolved to be idiosyncratic in terms of the specific metabolites induced.

Polewards expansions of species' distributions have been attributed to climate warming, but evidence for climate-driven local extinctions at warm (low latitude/elevation) boundaries is equivocal. We surveyed the four species of butterflies that reach their southern limits in Britain. We visited 421 sites where the species had been recorded previously to determine whether recent extinctions were primarily due to climate or habitat changes. Coenonympha tullia had become extinct at 52% of study sites and all losses were associated with habitat degradation. Aricia artaxerxes was extinct from 50% of sites, with approximately one-third to half of extinctions associated with climate-related factors and the remainder with habitat loss. For Erebia aethiops (extinct from 24% of sites), approximately a quarter of the extinctions were associated with habitat and three-quarters with climate. For Erebia epiphron, extinctions (37% of sites) were attributed mainly to climate with almost no habitat effects. For the three species affected by climate, range boundaries retracted 70–100 km northwards (A. artaxerxes, E. aethiops) and 130–150 m uphill (E. epiphron) in the sample of sites analysed. These shifts are consistent with estimated latitudinal and elevational temperature shifts of 88 km northwards and 98 m uphill over the 19-year study period. These results suggest that the southern/warm range margins of some species are as sensitive to climate change as are northern/cool margins. Our data indicate that climate warming has been of comparable importance to habitat loss in driving local extinctions of northern species over the past few decades; future climate warming is likely to jeopardize the long-term survival of many northern and mountain species.

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.

New Digs Many studies predict range alterations among species in response to climate change. Species, however, cannot be thought of as truly independent entities, because they all exist as a part of an interacting community. Most discussions of such interactions have focused on the potential restrictions they might place on range expansion; however, Pateman et al. (p. 1028 ) show that climate change has the potential to increase the number of species that can interact and thus facilitate expansion. British data collected by the general public on sightings of the Brown Argus butterfly and its host plants revealed an increased affiliation between the butterflies and a previously little-used group of plants. As summers warmed, populations associated with the new host were more productive than those associated with the more “traditional” host, which facilitated expansion of the butterflies into novel regions.

1. Salix lapponum host plants at an upper altitudinal site differed significantly in size, structural density, phenology, growth performance, and spatial isolation from those growing at a lower site. 2. Plant differences were paralleled by significant differences in psyllid population density and phenology parameters, with psyllid population density, percentage of catkins occupied, and phenological development relatively lower or retarded at the upper site. Population densities at the upper site, nevertheless, remained high. 3. Plant measurements were good predictors of insect density, often explaining up to 73% of the variance in abundance among plants at a given site. 4. Sets of four plant characters identified by best subsets regression were better predictors of psyllid density and development than single factors, although differences were often not great and the combinations of characters selected by multiple regression sometimes differed from the best single predictors. 5. Best single predictors of psyllid density on catkins were measurements of plant size, particularly height, length, and basal stem diameter. Shoot density and catkin phenology were occasionally important but plant isolation and prior growth performance were less important. 6. By contrast with density, age structure of the psyllid population was predicted best from plant phenological measurements, notably catkin phenology.

Since the 1940s, the distributions of several butterfly species have been expanding in northern Europe, probably in response to climate warming. We focus on the speckled wood butterfly Pararge aegeria in order to determine impacts of habitat availability on expansion rates. We analyse observed expansion rates since 1940 and also use a spatially explicit mechanistic model (MIGRATE) to simulate range expansion in two areas of the UK which differ in their distribution of breeding habitat (woodland). Observed and simulated expansion rates were in very close agreement but were 42%–45% slower in an area that had 24% less woodland. Unlike P. aegeria, the majority of butterfly species are not currently expanding, almost certainly because of lack of suitable habitat. Incorporating the spatial distribution of habitat into investigations of range changes is likely to be important in determining those species that can and cannot expand, and for predicting potential future range changes.

The single young of shearwaters grows slowly and is fed at long, often highly variable, intervals. We monitored the growth patterns of nestling Short-tailed Shearwaters Puffinus tenuirostris exchanged between nests in order to assess whether the patterns of parental visits or the demands of their young determined the rate of food provisioning in this species. These cross-fostering trials indicate a strong parental influence upon the rate of food delivery, but did not exclude the possibility of a minor influence from chicks.

Individuals colonizing unoccupied habitats typically possess characters associated with increased dispersal and, in insects, colonization success has been related to flight morphology. The speckled wood butterfly, Pararge aegeria, has undergone recent major expansions in its distribution: in the north of its range, P. aegeria has colonized many areas in north and east England, and in the south, it was first recorded on Madeira in 1976. We examined morphological traits associated with flight and reproduction in the northern subspecies tircis, and in the southern subspecies aegeria, from sites colonized about 20 years ago in northern England and on Madeira, respectively. Investment in flight was measured as relative wing area and thorax mass, and investment in reproduction as relative abdomen mass. All measurements were from individuals reared in a common environment and there were significant family effects in most of the variables measured. Compared with individuals from sites continuously occupied in recent history, colonizing individuals were larger (adult live mass). In the subspecies tircis, colonizing individuals also had relatively larger thoraxes and lower wing aspect ratios indicating that evolutionary changes in flight morphology may be related to colonization. However, sex by site interactions in analyses of thorax mass and abdomen mass suggest different selection pressures on flight morphology between the sexes in relation to colonization. Overall, the subspecies aegeria was smaller (adult live mass) and had a relatively larger thorax and wings, and smaller abdomen than subspecies tircis. Evolutionary changes in flight morphology and dispersal rate may be important determinants of range expansion, and may affect responses to future climate change.

Mean global temperatures have risen this century, and further warming is predicted to continue for the next 50-100 years(1-3) Some migratory species can respond rapidly to yearly climate variation ...

In modern landscapes, many populations of rare species are restricted to fragments of formerly extensive habitat. However, the potential for evolutionary changes in dispersal ability to occur within these fragmented populations has received little attention. We examined morphological traits associated with flight and reproduction in fragmented populations of the silver-spotted skipper butterfly, Hesperia comma . Investment in flight was measured as relative wing area and thorax mass, and investment in reproduction as relative abdomen mass. All measurements were made on individuals reared in a common environment. In the UK, Hesperia comma was once fairly widely distributed in southern and eastern England, but retracted to its smallest UK distribution in the 1970s and 1980s. It then partially re-expanded in the North and South Downs in S.E. England. We first compared traits from colonised and refuge sites 2.9–4.5 km apart, and found no differences in relative investment in flight or reproduction. There were, however, significant differences in both relative thorax and abdomen mass between regions >40 km apart. Populations with the highest relative investment in the thorax occurred in a region where population expansion has been most rapid. These results, in combination with other studies linking butterfly morphology to patch area and/or isolation, suggest that evolutionary responses to habitat fragmentation could be widespread. ©

exploitation along an altitudinal gradient: the willow psyllids (Cacopsylla spp.) on Salix lapponum. - Ecography 21: 289-296. Three congeneric species of Salix-feeding psyllid Cacopsylla spp. occur along an altitudinal gradient (988-1222 m a.s.l.) in southern Norway. The 27-km transect covered the local altitudinal range of the three species. Each species showed a different characteristic altitudinal distribution: C. propinqua occurred over the entire altitudinal range of the transect; C. palmeni was restricted to higher altitudes and C. brunneipennis occurred mainly at lower altitudes. This pattern of local altitudinal distribution mirrored the broader geographical distributions of these psyllids along latitudinal gradients. All three species developed on willow catkins but also made use of leaves. Species differed in their use of leaves with

1. Procellariiform seabirds such as Manx Shearwaters, Puffinus puffinus, accumulate large quantities of lipid during the nestling period, the purpose of which remains unclear. This paper examines temporal and age-specific variation in the pattern of food delivery to nestling Manx Shearwaters, to determine whether food delivery was likely to be determined primarily by the availability of food to adults at sea or by a chick’s short-term nutritional requirements. The variability in daily food delivery to chicks in the context of nestling obesity is then discussed. 2. For all but the youngest and oldest age classes, individual meals delivered by adults averaged 49 g, which was 12% of mean adult mass over the same period. More than 80% of chicks were fed each night and the maximum interval between feeding events for any chick was four nights. Feeding conditions did exert some influence over the ability of adults to feed their offspring, but deviations from expected values for both feeding frequency and meal size were restricted to a small number of nights, included values both higher and lower than expected and did not persist for more than one night in succession. These results were not compatible with the notion that large lipid reserves accumulated by chicks during their development provide insurance against prolonged interruptions to feeding. 3. Although adults did not appear to modulate food delivery over the nestling period, there was some evidence of short-term adjustment of feeding frequency (but not meal size) according to a chick’s immediate nutritional requirements. This suggests that variation in the provisioning rates of chicks did not result solely from stochastic variation in parental foraging success. However, the effect was slight and despite such minor adjustments, the coefficient of variation in daily provisioning rate was not substantially lower than in other species where the chick has no influence on parental food delivery. This suggests that even among those species that feed their chicks at comparatively frequent intervals, lipid accumulation among Procellariiformes may result from chronic overfeeding of chicks in order to reduce the impact of variable food provisioning by individual parents.

1. Three independent methods were used to investigate population structure in the butterfly Plebejus argus. First, migration and dispersal ability were measured by mark–release–recapture in seven adjacent habitat patches, and by release of butterflies in unoccupied habitat. Secondly, colonization of newly created habitat was observed over 7 years. Finally, genetic differentiation of local populations within a metapopulation was investigated. Sampled local populations included parts of the mark–release–recapture study area. 2. Plebejus argus is relatively sedentary: the maximum movement detected was 395 m, and only 2% of individuals moved further than 100 m between recaptures on different days. None the less, adjacent local populations in the mark–release–recapture study area were linked by occasional migration, with ≈ 1.4% of individuals moving between patches separated by 13–200 m. 3. Despite low mobility, observed colonizations occurred rapidly over distances of 1 km. Because P. argus occurs at high population densities, 1.4% migration can generate enough migrants to colonize newly suitable habitat quickly at this spatial scale. 4. Mark–release–recapture data were used to predict that there would be limited genetic differentiation through drift between local populations at this spatial scale. The prediction was supported by allele frequency data for the same local populations. 5. Genetic differentiation often indicates higher levels of migration than are revealed by the movements of marked individuals. This study shows that when experimental releases and extensive marking are undertaken in areas that are large relative to most movements, indirect measures of gene flow and direct measures of dispersal can concur. 6. Evidence from the three different approaches was complementary, indicating that P. argus occurs as metapopulations within the study area.

Some species are expanding their ranges polewards during current climate warming. However, anthropogenic fragmentation of suitable habitat is affecting expansion rates and here we investigate interactions between range expansion, habitat fragmentation and genetic diversity. We examined three closely related Satyrinae butterflies, which differ in their habitat associations, from six sites along a transect in England from distribution core to expanding range margin. There was a significant decline in allozyme variation towards an expanding range margin in Pararge aegeria , which has the most restricted habitat availability, but not in Pyronia tithonus whose habitat is more widely available, or in a non-expanding ‘control species’ ( Maniola jurtina ). Moreover, data from another transect in Scotland indicated that declines in genetic diversity in P. aegeria were evident only on the transect in England, which had greater habitat fragmentation. Our results indicate that fragmentation of breeding habitats leads to more severe founder events during colonization, resulting in reduced diversity in marginal populations in more specialist species. The continued widespread loss of suitable habitats in the future may increase the likelihood of loss of genetic diversity in expanding species, which may affect whether or not species can adapt to future environmental change.

Long-range, seasonal migration is a widespread phenomenon among insects, allowing them to track and exploit abundant but ephemeral resources over vast geographical areas. However, the basic patterns of how species shift across multiple locations and seasons are unknown in most cases, even though migrant species comprise an important component of the temperate-zone biota. The painted lady butterfly Vanessa cardui is such an example; a cosmopolitan continuously-brooded species which migrates each year between Africa and Europe, sometimes in enormous numbers. The migration of 2009 was one of the most impressive recorded, and thousands of observations were collected through citizen science programmes and systematic entomological surveys, such as high altitude insect-monitoring radar and ground-based butterfly monitoring schemes. Here we use V. cardui as a model species to better understand insect migration in the Western Palaearctic, and we capitalise on the complementary data sources available for this iconic butterfly. The migratory cycle in this species involves six generations, encompassing a latitudinal shift of thousands of kilometres (up to 60 degrees of latitude). The cycle comprises an annual poleward advance of the populations in spring followed by an equatorward return movement in autumn, with returning individuals potentially flying thousands of kilometres. We show that many long-distance migrants take advantage of favourable winds, moving downwind at high elevation (from some tens of metres from the ground to altitudes over 1000 m), pointing at strong similarities in the flight strategies used by V. cardui and other migrant Lepidoptera. Our results reveal the highly successful strategy that has evolved in these insects, and provide a useful framework for a better understanding of long-distance seasonal migration in the temperate regions worldwide.

Nymphal development of Cacopsylla moscovita (And.) (Homoptera : Psylloidea) takes place only on female catkins of Salix repens L. and close phenological synchrony is crucial because development times of catkins and nymphs are similar. Eggs are laid on catkins as soon as they develop and close synchrony between oviposition and budburst maximizes time available for nymphal development. Sampling adult C. moscovita in the field revealed little synchrony between egg development and budburst, with over 60% of females containing mature eggs four weeks before catkins first appeared. In the laboratory, egg development was influenced by both temperature and photoperiod. At 10 °C, egg development occurred approximately one month earlier than at 5 °C and two weeks earlier than in the field. Adult survival in the laboratory was substantially reduced at increased temperature, with only 20% of adults surviving longer than two weeks at 10 °C, compared with over 95% at 5 °C. Body condition (weight corrected for size) of males and females decreased significantly in the field over winter. However, body condition of females levelled off before budburst, coincident with egg development implying that females had resumed feeding. We discuss our results in relation to regulation of phenological synchrony between oviposition and catkin appearance.

Summary. 1 The geographical distributions of three species of jumping plant lice (psyllids) along an altitudinal transect (988–1300 m a.s.l.) in southern Norway were restricted within the range of their host plant Salix lapponum. One species, Cacopsylla propinqua, occurred at all sampling locations between 988 and 1222 m, whereas C.palmeni was confined to higher altitudes (1153–1222 m) and C.brunneipennis was more abundant at lower altitudes (988–1101 m). 2 C.brunneipennis and C.palmeni developed only on female catkins. Development times of catkins and psyllids were similar (approximately 50 days) and successful psyllid development depended on close phenological synchrony with catkins. 3 Thermal requirements for development of female catkins were greater at low altitude (988 m) compared with higher altitude (1222 m), showing local adaptation of S.lapponum to altitude. In general, thermal requirements of psyllids were less than those of catkins at the same location. C.brunneipennis had higher thermal requirements than C.palmeni. 4 Field experiments, using polythene enclosures to elevate temperatures at two sites at different altitudes (by 0.6–1.4 deg. C), showed that insects had an enhanced relative rate of development under elevated temperatures compared with their host plants. 5 Indices of phenological synchrony were calculated from thermal requirements of psyllids and catkins. Under elevated temperatures, phenological synchrony decreased at both sites. This resulted in the subsequent development of smaller adult insects at low altitude, although at higher altitude, insects developing under elevated temperatures were larger and had a higher survival rate compared with controls. 6 Effects of temperature on phenological synchrony may explain the limits to the geographical range of psyllids. The consequences of climate change on psyllid populations will depend on the effects of decreased phenological synchrony on insect development and this may differ within the insect's geographical range.

1 Adults of the noctuid moth Autographa gamma undertake seasonal migrations into areas where they are unable to breed continuously. Individuals migrate into Britain each spring, and offspring of these migrants probably return in autumn to over-wintering areas in North Africa and the Middle East. However, the existence of these return migrations has been questioned. 2 Insects usually migrate during the adults' pre-reproductive period (PRP). The length of this period is therefore an index of migratory potential since individuals with longer PRPs have more time to express their potential for flight and to travel further before becoming sexually mature. 3 Offspring of insects from Morocco, Britain and Germany showed that, under both long and short photoperiods, PRP increased with latitude over the range 34–53°N. There were significant genotype—environment interactions in the effect of photoperiod on length of PRP. 4 However, PRPs of offspring from insects collected in Sweden were no different under a long photoperiod from those of offspring collected further south, and were significantly shorter under a short photoperiod than those of insects from Morocco. 5 With the exception of Swedish moths, offspring from insects collected at higher latitudes had longer PRPs and, therefore, the potential to travel further during migrations. The ability of some individuals to respond to environmental cues signalling habitat deterioration will further increase their potential to return to over-wintering areas further south.

Genetic control of the pre-reproductive period in Autographa gamma (L.) (Silver Y moth) (Lepidoptera: Noctuidae)

Adults ofAutographa gamma(Linnaeus) make regular, seasonal migrations into areas where they are unable to breed continuously. Individuals migrate into Britain each spring, and after one, two or three generations, offspring of the spring migrants return to over-wintering areas in North Africa and the Middle East. The larvae are highly polyphagous and have been recorded damaging many crops, especiallyBrassicaspp. Insects usually migrate during the adults' pre-reproductive period (PRP). The length of this period is therefore an index of migratory potential since individuals with longer PRPs have more time to express their capacity for flight and to travel further. Individuals reared outside in Britain in autumn have longer PRPs than individuals reared outside in summer. Laboratory experiments show that PRPs are extended at low temperatures and under short photoperiods and there is some evidence that PRPs are also extended under gradually decreasing photoperiods. By extending PRPs in response to environmental conditions typical of Britain in autumn, individuals are able to increase their potential for pre-reproductive migratory flight and therefore their potential for reaching suitable overwintering sites further south.

During recent climate warming, some species have expanded their ranges northwards to keep track of climate changes. Evolutionary changes in dispersal have been demonstrated in these expanding populations and here we show that increased dispersal is associated with reduced investment in reproduction in populations of the speckled wood butterfly, Pararge aegeria. Evolutionary changes in flight versus reproduction will affect the pattern and rate of expansion at range boundaries in the future, and understanding these responses will be crucial for predicting the distribution of species in the future as climates continue to warm.

Evidence of anthropogenic global climate change is accumulating, but its potential consequences for insect distributions have received little attention. We use a 'climate response surface' model to investigate distribution changes at the northern margin of the speckled wood butterfly, Pararge aegeria. We relate its current European distribution to a combination of three bioclimatic variables. We document that P. aegeria has expanded its northern margin substantially since 1940, that changes in this species' distribution over the past 100 years are likely to have been due to climate change, and that P. aegeria will have the potential to shift its range margin substantially northwards under predicted future climate change. At current rates of expansion, this species could potentially colonize all newly available climatically suitable habitat in the UK over the next 50 years or more. However, fragmentation of habitats can affect colonization, and we show that availability of habitat may be constraining range expansion of this species at its northern margin in the UK. These lag effects may be even more pronounced in less-mobile species inhabiting more fragmented landscapes, and highlight how habitat distribution will be crucial in predicting species' responses to future climate change.

Procellariiform seabirds such as the Manx shearwater Puffinus puffinus, rear only one chick at a time but may breed many times in their lives; parents should thus limit food delivery to the chick in keeping with the balance between current and future reproductive output. Yet procellariiform chicks accumulate large quantities of lipid, which may provide a buffer against pronounced and unpredictable variation in food provisioning, resulting in part from an inability of parents to regulate food supply to the nest. We switched chicks between nests to examine the roles of parents and offspring in controlling food delivery. The serial autocorrelation in age-specific body masses for unmanipulated chicks decreased from 0.61 (P< 0.01) to 0.35 (NS) over a period of 15 days and remained nonsignificant thereafter. By contrast, the serial autocorrelation for switched chicks increased from 0.64 (P< 0.01) to 0.83 (P< 0.001) and the serial cross-correlation rose from 0.23 (NS) to 0.50 (P< 0.05). These results supported both chick determination and parental determination models of food provisioning, indicating that chicks conveyed information about their nutritional status, which parents acted upon by adjusting their rate of food delivery. We discuss these results in relation to the optimization of nestling lipid reserves and parental foraging effort. We suggest that information conveyed by the chick's begging intensity serves to reduce the provisioning rate to well-fed chicks, but parents cannot or do not increase food provisioning to poorly fed chicks. Such adjustment of food provisioning does not refute the hypothesis that nestling obesity provides a buffer against highly variable food delivery. Copyright 1999 The Association for the Study of Animal Behaviour.

Cory's Shearwaters, in common with other species of Procellariiformes, accumulate large quantities of body fat after hatching. This pattern of development has been related both to temporal variation in feeding conditions around the colony and to stochastic variation in foraging success and food provisioning by individual parents. Chicks need not necessarily always accept food provided by their parents, yet few studies have examined parental attendance independently of the masses of food fed to chicks overnight. This paper examines variability in provisioning and growth of nestling Cory's Shearwaters over three years between 1991 and 1994, and uses observations of individually marked adults coupled with radio-tracking to examine the nest attendance patterns of individual parents. Chicks were fed on 77% of nights on average, with little temporal variation in food delivery at the level of the colony. Individual chicks were nonetheless fed highly variably, and this to some extent supports the notion that nestling obesity results from chronic overfeeding by the parents to reduce the impact of variable food provisioning at an individual level. Chicks were attended by 1.5 parents overnight on average, and chicks were attended but not fed on 16% of occasions. Parents may have returned on these occasions without any food for the chick, or the chick may have refused to accept food from the parents. A minority of nests accounted for most of the occasions when a chick was attended but not fed, and there was evidence to suggest that these chicks were consistently heavy for their size. A refusal by these chicks to accept food on some occasions is consistent with the notion that lipid stores of chicks should be optimized rather than maximized, and this requires further investigation.

1. Metapopulation dynamics of the silver-spotted skipper butterfly Hesperia comma were studied between 1982 and 1991 along a 25-km stretch of chalk hills on the North Downs, Surrey, UK. Sixty-nine patches of suitable habitat were identified, of which 48 were occupied in 1982. Over the 9-year period, 12 patches were colonized, seven went extinct and nine patches remained vacant. Patches were more likely to be colonized if they were relatively large and close to other large, occupied patches. Local populations in small, isolated patches were more likely to go extinct. 2. Within a 70-ha section of a metapopulation, mark-release-recapture techniques were used in 1994 to investigate the effects of local patch area and isolation on movement of individuals among local patches. 988 butterflies were marked of which 133 moved between patches. 67% of between-patch movements were less than 50 m, although the longest recorded distance moved was 1070 m. Butterflies were most likely to move between large patches that were close together. 3. Metapopulation models have assumed that the distribution of distances moved by migrants follows a negative-exponential function. In our study, this distribution fitted an inverse-power function better than a negative-exponential. The under-estimation of long-distance migration by negative exponentials, compared with inverse power functions, may explain why theoretical models have under-estimated the number of occupied patches in metapopulations of H. comma following natural colonization. 4. Per capita emigration and immigration rates were significantly higher in small patches (area < 0.07 ha) compared with medium-sized (0.33-0.78 ha) or large patches (5.66 ha). However, in absolute numbers, more emigrants came from the largest patch where the source population was the largest. 5. The population system studied here shares attributes of several theoretical types of spatially structured population (patchy population, metapopulation, mainland-island system, non-equilibrium system) depending on the temporal and spatial scale examined. The distribution and dynamics of metapopulations should be regarded as being affected by a variety of behavioural and population processes, and real metapopulations can rarely be characterized as a single theoretical type.

We examined the pattern of chick feeding in Cory's shearwater Calonectris diomedea on Selvagem Grande by weighing 50 chicks at 4-h intervals throughout nine successive nights, then daily for a further 11 days (20 days in all). Meal sizes were estimated from the sum of positive mass increments recorded over 4-h intervals (SUM). Mass increments over 24-h (NET) were linearly related to SUMs by the equation NET = 0.83 SUM −40.1 (r 2 = 0.82). Individual meals fed to chicks averaged 74 g (SD = 26.4 g, range = 20 - 100 g). A mean of 79% of chicks were fed each night and the mean interval between feeds was 1.3 nights, with a maximum of four nights. There was little day-to-day variation in feeding rate, and no relationship with ambient light conditions over a half lunar cycle