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Range shifting is vital for species persistence, but there is little consensus on why individual species vary so greatly in the rates at which their ranges have shifted in response to recent climate warming. Here, using 40 years of distribution data for 291 species from 13 invertebrate taxa in Britain, we show that interactions between habitat availability and exposure to climate change at the range margins explain up to half of the variation in rates of range shift. Habitat generalists expanded faster than more specialised species, but this intrinsic trait explains less of the variation in range shifts than habitat availability, which additionally depends on extrinsic factors that may be rare or widespread at the range margin. Similarly, while climate change likely underlies polewards expansions, we find that more of the between-species variation is explained by differences in habitat availability than by changes in climatic suitability. A model that includes both habitat and climate, and their statistical interaction, explains the most variation in range shifts. We conclude that climate-change vulnerability assessments should focus as much on future habitat availability as on climate sensitivity and exposure, with the expectation that habitat restoration and protection will substantially improve species' abilities to respond to uncertain future climates.

Extreme climatic events could be major drivers of biodiversity change, but it is unclear whether extreme biological changes are (i) individualistic (species- or group-specific), (ii) commonly associated with unusual climatic events and/or (iii) important determinants of long-term population trends. Using population time series for 238 widespread species (207 Lepidoptera and 31 birds) in England since 1968, we found that population 'crashes' (outliers in terms of species' year-to-year population changes) were 46% more frequent than population 'explosions'. (i) Every year, at least three species experienced extreme changes in population size, and in 41 of the 44 years considered, some species experienced population crashes while others simultaneously experienced population explosions. This suggests that, even within the same broad taxonomic groups, species are exhibiting individualistic dynamics, most probably driven by their responses to different, short-term events associated with climatic variability. (ii) Six out of 44 years showed a significant excess of species experiencing extreme population changes (5 years for Lepidoptera, 1 for birds). These 'consensus years' were associated with climatically extreme years, consistent with a link between extreme population responses and climatic variability, although not all climatically extreme years generated excess numbers of extreme population responses. (iii) Links between extreme population changes and long-term population trends were absent in Lepidoptera and modest (but significant) in birds. We conclude that extreme biological responses are individualistic, in the sense that the extreme population changes of most species are taking place in different years, and that long-term trends of widespread species have not, to date, been dominated by these extreme changes.This article is part of the themed issue 'Behavioural, ecological and evolutionary responses to extreme climatic events'., (© 2017 The Authors.)

Responses of species to climate change are extremely variable, perhaps because of climate-related changes to interactions among species. We show that temperature-related changes in the dependence of the butterfly Aricia agestis on different larval host plants have facilitated rapid range expansion. Historically, the butterfly was largely restricted to a single plant species, Helianthemum nummularium, but recent warmer conditions have enabled the butterfly to increasingly use the more widespread plant species Geranium molle. This has resulted in a substantial increase in available habitat and rapid range expansion by the butterfly (79 kilometers northward in Britain in 20 years). Interactions among species are often seen as constraints on species' responses to climate change, but we show that temperature-dependent changes to interspecific interactions can also facilitate change.

Thermal constraints on development are often invoked to predict insect distributions. These constraints tend to be characterized in species distribution models (SDMs) by calculating development time based on a constant lower development temperature (LDT). Here, we assessed whether species-specific estimates of LDT based on laboratory experiments can improve the ability of SDMs to predict the distribution shifts of six U.K. butterflies in response to recent climate warming. We find that species-specific and constant (5 degrees C) LDT degree-day models perform similarly at predicting distributions during the period of 1970-1982. However, when the models for the 1970-1982 period are projected to predict distributions in 1995-1999 and 2000-2004, species-specific LDT degree-day models modestly outperform constant LDT degree-day models. Our results suggest that, while including species-specific physiology in correlative models may enhance predictions of species' distribution responses to climate change, more detailed models may be needed to adequately account for interspecific physiological differences.

An appreciation of how some species are becoming more common despite unprecedented anthropogenic pressures could offer key insights for mitigating the global biodiversity crisis.  Research to date has largely focused on declining species, while species that are becoming more common have received relatively little attention. Macro-moths in Great Britain are well-studied and species-rich, making them an ideal group for addressing this knowledge gap. Here, we examine changes in 51 successful species between 1968 and 2016 using 4.5 million occurrence records and a systematic monitoring dataset. We employ 3D graphical analysis to visualise long-term multidimensional trends in prevalence (abundance and range) and use vector autoregression models to test whether past values of local abundance are useful for predicting changes in the extent of occurrence. The responses of Anthropocene winners are heterogeneous, suggesting multiple drivers are responsible. Changes in range and local abundance frequently occur intermittently through time, demonstrating the value of long-term, continuous monitoring. There is significant diversity among the winners themselves, which include widespread generalists, habitat specialists, and recent colonists. We offer brief discussion of possible causal factors and the wider ecosystem implications of these trends.

Species' distributions are moving polewards in response to climate change, and although range expansions of relatively warm‐adapted species are widely reported, reports of range retractions in cool‐adapted species are less common. Here, we analysed species' distribution shifts for 76 cool‐adapted moths in Great Britain using citizen science occurrence records from the National Moth Recording Scheme over a 40‐year period. Although we find evidence for trailing edge shifts to higher latitudes, shifts in species' range centroids are oriented towards the north‐west, and are more closely correlated with directional changes in total precipitation than average temperature. We also found that species' local extinction risk is higher in areas where temperature is high and precipitation is low, but this risk diminishes as precipitation increases. Adaptation efforts should therefore focus on maintaining or increasing water availability as the climate continues to change. [ABSTRACT FROM AUTHOR]

Advances in phenology (the annual timing of species' life-cycles) in response to climate change are generally viewed as bioindicators of climate change, but have not been considered as predictors of range expansions. Here, we show that phenology advances combine with the number of reproductive cycles per year (voltinism) to shape abundance and distribution trends in 130 species of British Lepidoptera, in response to ~0.5 °C spring-temperature warming between 1995 and 2014. Early adult emergence in warm years resulted in increased within- and between-year population growth for species with multiple reproductive cycles per year (n = 39 multivoltine species). By contrast, early emergence had neutral or negative consequences for species with a single annual reproductive cycle (n = 91 univoltine species), depending on habitat specialisation. We conclude that phenology advances facilitate polewards range expansions in species exhibiting plasticity for both phenology and voltinism, but may inhibit expansion by less flexible species. Many species' life cycles have moved earlier in the year because of climate change, but we do not know the consequences for range expansions. The authors show that these advances promote range expansions in species with multiple reproductive cycles per year, but not in species with only one. [ABSTRACT FROM AUTHOR]

A cornerstone of conservation is the designation and management of protected areas ( PAs): locations often under conservation management containing species of conservation concern, where some development and other detrimental influences are prevented or mitigated. However, the value of PAs for conserving biodiversity in the long term has been questioned given that species are changing their distributions in response to climatic change. There is a concern that PAs may become climatically unsuitable for those species that they were designated to protect, and may not be located appropriately to receive newly-colonizing species for which the climate is improving. In the present study, we analyze fine-scale distribution data from detailed resurveys of seven butterfly and 11 bird species in Great Britain aiming to examine any effect of PA designation in preventing extinctions and promoting colonizations. We found a positive effect of PA designation on species' persistence at trailing-edge warm range margins, although with a decreased magnitude at higher latitudes and altitudes. In addition, colonizations by range expanding species were more likely to occur on PAs even after altitude and latitude were taken into account. PAs will therefore remain an important strategy for conservation. The potential for PA management to mitigate the effects of climatic change for retracting species deserves further investigation. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●-●●. [ABSTRACT FROM AUTHOR]

Many species are extending their leading-edge (cool) range margins polewards in response to recent climate change. In the present study, we investigated range margin changes at the northern (cool) range margins of 1573 southerly-distributed species from 21 animal groups in Great Britain over the past four decades of climate change, updating previous work. Depending on data availability, range margin changes were examined over two time intervals during the past four decades. For four groups (birds, butterflies, macromoths, and dragonflies and damselflies), there were sufficient data available to examine range margin changes over both time intervals. We found that most taxa shifted their northern range margins polewards and this finding was not greatly influenced by changes in recorder effort. The mean northwards range margin change in the first time interval was 23 km per decade ( N = 13 taxonomic groups) and, in the second interval, was 18 km per decade ( N = 16 taxonomic groups) during periods when the British climate warmed by 0.21 and 0.28 °C per decade, respectively. For the four taxa examined over both intervals, there was evidence for higher rate of range margin change in the more recent time interval in the two Lepidoptera groups. Our analyses confirm a continued range margin shift polewards in a wide range of taxonomic groups. [ABSTRACT FROM AUTHOR]

Microclimate has been known to drive variation in the distribution and abundance of insects for some time. Until recently however, quantification of microclimatic effects has been limited by computing constraints and the availability of fine-scale biological data. Here, we tested fine-scale patterns of persistence/extinction in butterflies and moths against two computed indices of microclimate derived from Digital Elevation Models: a summer solar index, representing fine-scale variation in temperature, and a topographic wetness index, representing fine-scale variation in moisture availability. We found evidence of microclimate effects on persistence in each of four 20 × 20 km British landscapes selected for study (the Brecks, the Broads, Dartmoor, and Exmoor). Broadly, local extinctions occurred more frequently in areas with higher minimum or maximum solar radiation input, while responses to wetness varied with landscape context. This negative response to solar radiation is consistent with a response to climatic warming, wherein grid squares with particularly high minimum or maximum insolation values provided an increasingly adverse microclimate as the climate warmed. The variable response to wetness in different landscapes may have reflected spatially variable trends in precipitation. We suggest that locations in the landscape featuring cooler minimum and/or maximum temperatures could act as refugia from climatic warming, and may therefore have a valuable role in adapting conservation to climatic change. [ABSTRACT FROM AUTHOR]

It is uncertain whether Protected Areas (PAs) will conserve high abundances of species as their distributions and abundances shift in response to climate change. We analyzed large datasets for 57 butterfly and 42 odonate species (including four that have recently colonized Britain). We found that 73 of 94 species with sufficient data for analysis were more abundant inside than outside PAs in the historical parts of their British distributions, showing that PAs have retained high conservation value. A significant majority (61 of 99 species) was also more abundant inside PAs in regions they have colonized during the last 30-40 years of climate warming. Species with relatively high abundances inside PAs in long-established parts of their distributions were also disproportionately associated with PAs in recently colonized regions, revealing a set of relatively PA-reliant species. Pas, therefore, play a vital role in the conservation of biodiversity as species' ranges become more dynamic. [ABSTRACT FROM AUTHOR]

There is little consensus as to why there is so much variation in the rates at which different species' geographic ranges expand in response to climate warming. Here we show that the relative importance of species' abundance trends and habitat availability for British butterfly species vary over time. Species with high habitat availability expanded more rapidly from the 1970s to mid-1990s, when abundances were generally stable, whereas habitat availability effects were confined to the subset of species with stable abundances from the mid-1990s to 2009, when abundance trends were generally declining. This suggests that stable (or positive) abundance trends are a prerequisite for range expansion. Given that species' abundance trends vary over time for non-climatic as well as climatic reasons, assessment of abundance trends will help improve predictions of species' responses to climate change, and help us to understand the likely success of different conservation strategies for facilitating their expansions. [ABSTRACT FROM AUTHOR]

The phenology of many species has been shown to shift under climate change. However, because species respond at different rates, ecological communities may be disrupted leading to species extinctions and loss of ecosystem services. Hence, there is a need to monitor and understand phenological change., Population data, gathered by standardised monitoring schemes, can be used to this end. However, such schemes require significant organisation and financial resources. Distribution data (georeferenced biological records with dates) are easier and cheaper to collect and may be an unexploited resource for phenology analyses. This would allow analysis of more taxa from more regions of the world. However, distribution data are potentially biased due to the unstandardised behaviour of biological recorders., Here, the ability of distribution data record dates to accurately predict phenology is investigated by using the British butterfly fauna as a model system. We used the total number of distribution records per unit time across Great Britain as a proxy for butterfly abundance. Phenology metrics of mean flight date and flight period length were then calculated from the resulting abundance-time relationships for each year in a 15-year time series. These estimates were validated against those generated from a standardised-effort population monitoring scheme., We analysed 1 078 328 records from 30 British butterflies and found that distribution data accurately predicted the mean flight date for 22 of the 30 species tested. Flight period length was only predicted accurately for seven of 30 species., We found a nonlinear but consistent positive relationship between the accuracy of mean flight date estimates and sample size (number of records) at both inter- and intraspecific scales. Our results suggest that a threshold sample size of c. 6500 distribution records (430 per year) is a pragmatic compromise between accuracy and recording effort, leading to little loss of accuracy in phenology predictions (an average decrease in accuracy of 2·9 days was observed)., The results suggest that distribution data are a potentially useful resource for phenology research. This may allow practitioners to monitor particular regions and previously unstudied species relatively cheaply using existing mapping schemes. [ABSTRACT FROM AUTHOR]

. 1. Population declines among insects are inadequately quantified, yet of vital importance to national and global biodiversity assessments and have significant implications for ecosystem services. 2. Substantial declines in abundance and distribution have been reported recently within a species-rich insect taxon, macro-moths, in Great Britain and other European countries. These declines are of concern because moths are important primary consumers and prey items for a wide range of other taxa, as well as contributing to ecosystem services such as pollination. 3. I summarise these declines and review potential drivers of change. Direct evidence for causes of moth declines is extremely limited, but correlative studies and extrapolation from closely related taxa suggest that habitat degradation (particularly because of agricultural intensification and changing silviculture) and climate change are likely to be major drivers. There is currently little evidence of negative population-level effects on moths caused by chemical or light pollution, non-native species or direct exploitation. 4. I make suggestions for future research with a focus on quantifying impacts of land management practices, light pollution and climate change on moth population dynamics and developing evidence-based measures that can be incorporated into agri-environment schemes and other policy initiatives to help reverse the widespread decline of moths in Great Britain and beyond. [ABSTRACT FROM AUTHOR]

Many species are expanding at their leading-edge range boundaries in response to climate warming. Species are known to respond individualistically to climate change, but there has been little consideration of whether responses are consistent over time. We compared responses of 37 southerly distributed British butterflies over two study periods, first between 1970-1982 and 1995-1999 and then between 1995-1999 and 2005-2009, when mean annual temperature increased regionally by 0.03 °C yr−1 (a significant rate of increase) and 0.01 °C yr−1(a nonsignificant increase) respectively. Our study species might be expected to benefit from climate warming. We measured three responses to climate to investigate this; changes in range margin, distribution area and abundance. In general, the responses of species were inconsistent over time. Species that increased their distribution areas during the first period tended to do so again during the second period, but the relationship was weak. Changes in range margins and abundance were not consistent. In addition, only 5/37 species showed qualitatively similar responses in all three response variables over time (three species increased and two species declined in all variables in both periods). Overall rates of range expansion and distribution area change were significantly greater in the second study period, despite the lower rate of warming, perhaps due to species exploiting climate-distribution lags remaining from the earlier, warmer period. However, there was a significantly greater decline in abundance during the second study period, so range expansions northwards were not necessarily accompanied by increases in distribution area and/or abundance. Hence, species ranges have been thinning as they have expanded northwards. The idiosyncratic responses of these species likely reflect the balance of climatic and habitat drivers of species distribution and abundance changes. [ABSTRACT FROM AUTHOR]

Conservation planners often wish to predict how species distributions will change in response to environmental changes. Species distribution models (SDMs) are the primary tool for making such predictions. Many methods are widely used; however, they all make simplifying assumptions, and predictions can therefore be subject to high uncertainty. With global change well underway, field records of observed range shifts are increasingly being used for testing SDM transferability. We used an unprecedented distribution dataset documenting recent range changes of British vascular plants, birds, and butterflies to test whether correlative SDMs based on climate change provide useful approximations of potential distribution shifts. We modelled past species distributions from climate using nine single techniques and a consensus approach, and projected the geographical extent of these models to a more recent time period based on climate change; we then compared model predictions with recent observed distributions in order to estimate the temporal transferability and prediction accuracy of our models. We also evaluated the relative effect of methodological and taxonomic variation on the performance of SDMs. Models showed good transferability in time when assessed using widespread metrics of accuracy. However, models had low accuracy to predict where occupancy status changed between time periods, especially for declining species. Model performance varied greatly among species within major taxa, but there was also considerable variation among modelling frameworks. Past climatic associations of British species distributions retain a high explanatory power when transferred to recent time - due to their accuracy to predict large areas retained by species - but fail to capture relevant predictors of change. We strongly emphasize the need for caution when using SDMs to predict shifts in species distributions: high explanatory power on temporally-independent records - as assessed using widespread metrics - need not indicate a model's ability to predict the future. [ABSTRACT FROM AUTHOR]

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 “species-climate” 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. [ABSTRACT FROM AUTHOR]

Aerosols play an important role in global climate change by their interactions with incoming solar radiation and outgoing longwave radiation from the planetary surface. The climate effects of aerosols depend on their scattering and absorption properties. This article describes the development of an instrument (ASTER: Aerosol Scattering To Extinction Ratio) that simultaneously measures the scattering and extinction of single aerosol particles. ASTER uses a high-Q cavity to amplify the extinction signal and innovative optics to collect the scattered light. It can distinguish many partially absorbing particles from a few black ones even if the bulk absorption is the same. Optical sizing and single-scattering albedo measurements were made for laboratory-generated particles with diameters from about 300 nanometers to above one micrometer. Using this prototype instrument, changes in albedo for single particles of 20% or greater were detected by measurement of the scattering and extinction. Optical sizing of the individual particles to within ∼ 50 nm was accomplished using the ratio of the forward scattered light to the total scattering. Initial measurements of laboratory air showed a mode of highly absorbing particles. This article reports design and early laboratory tests on ASTER. [ABSTRACT FROM AUTHOR]

Many factors, including climate, resource availability, and habitat diversity, have been proposed as determinants of global diversity, but the links among them have rarely been studied. Using structural equation modeling (SEM), we investigated direct and indirect effects of climate variables, host-plant richness, and habitat diversity on butterfly species richness across Britain, at 20-km grid resolution. These factors were all important determinants of butterfly diversity, but their relative contributions differed between habitat generalists and specialists, and whether the effects were direct or indirect. Climate variables had strong effects on habitat generalists, whereas host-plant richness and habitat diversity contributed relatively more for habitat specialists. Considering total effects (direct and indirect together), climate variables had the strongest link to butterfly species richness for all groups of species. The results suggest that different mechanistic hypotheses to explain species richness may be more appropriate for habitat generalists and specialists, with generalists hypothesized to show direct physiological limitations and specialists additionally being constrained by trophic interactions (climate affecting host-plant richness). [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Evidence is accumulating of shifts in species' distributions during recent climate warming. However, most of this information comes predominantly from studies of a relatively small selection of taxa (i.e., plants, birds and butterflies), which may not be representative of biodiversity as a whole. Using data from less well-studied groups, we show that a wide variety of vertebrate and invertebrate species have moved northwards and uphill in Britain over approximately 25 years, mirroring, and in some cases exceeding, the responses of better-known groups. [ABSTRACT FROM AUTHOR]

The Rothamsted Insect Survey has operated a Great Britain-wide network of light-traps since 1968. From these data we estimated the first ever national abundance indices and 35-year population trends for 338 species of common macro-moths. Although the number of trap sites which run each year is not constant, there is a representative, well-distributed core of traps that have run for = 15 years. The proportion of operating sites catching a species and the annual geometric mean catch of successful traps were used to provide estimates of species range and absolute abundance.T, an index of long-term population trends, was used to compare trends among species.Twas not biased by trap site turnover. The percentage of species displaying significant decreases (54%) was more than double that displaying increases (22%). Species found throughout Great Britain are decreasing most rapidly in the south and especially the southeast but species with a southerly distribution are increasing. Results of a preliminary overview suggest habitat and climate change may both play a role in changing species dynamics. The existence of estimates of abundances and trends for such a large species pool opens the way for much further research, linking trends with land-use changes, climate change and inter-specific dynamics. [ABSTRACT FROM AUTHOR]

It is important for conservationists to be able to assess the risks that climate change poses to species, in order to inform decision making. Using standardised and repeatable methods, we present a national-scale assessment of the risks of range loss and opportunities for range expansion that climate change could pose for over 3000 plants and animals. Species were selected by their occurrence in England, the primary focus of the study, but climate change impacts were assessed across Great Britain, widening their geographical relevance. A basic risk assessment that compared projected future changes in potential range with recently observed changes classified 21% of species as being at high risk and 6% at medium risk of range loss under a B1 climate change scenario. A greater number of species were classified as having a medium (16%) or high (38%) opportunity to potentially expand their distribution. A more comprehensive assessment, incorporating additional ecological information, including potentially confounding and exacerbating factors (e.g. dispersal, habitat availability and other constraints), was applied to 402 species, of which 35% were at risk of range loss and 42% may expand their range extent. This study covers a temperate region with a significant proportion of species at their poleward range limit; the balance of risks and opportunities from climate change may be different elsewhere. The outcome of both risk assessments varied between taxonomic groups, with bryophytes and vascular plants containing the greatest proportion of species at risk from climate change. Upland habitats contained more species at risk than other habitats. Whilst the overall pattern was clear, confidence was generally low for individual assessments, with the exception of well-studied taxa such as birds. In response to climate change, nature conservation needs to plan for changing species distributions and an uncertain future. [ABSTRACT FROM AUTHOR]