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Animals move in ‘modes’ where movement patterns relate to specific behaviours. Despite much work on the movement of butterflies, their behavioural modes are relatively unexplored. Here we analysed the behaviour of the model butterfly species the meadow brown, Maniola jurtina. We identified modes in both sexes and across habitats varying in resource density. We found that, in nectar-rich habitats, males had more diverse behaviour than females, engaging in a unique ‘high-flight’ mode associated with mate search, whereas females were primarily nectaring or inactive. In nectar-poor habitats, both sexes were similar, switching between flight and inactivity. We also identified the movement parameters of the modes, finding that, for both sexes, movements associated with nectaring were slower and more tortuous and, for males, the mode associated with mate searching was straighter and faster. Using an individual-based random-walk model, we investigated the effects of behaviour on movement predictions by comparing a mode-switching model with a version including intraspecific variation and another assuming homogeneity between individuals. For both sexes, including modes affected the mean and shape of the displacement rate compared to models assuming homogeneity, although for females modes increased displacement 1.5 times while for males they decreased it by a third. Both models also differed substantially from models assuming intraspecific variation. Finally, using a new model of search behaviour we investigated the general conditions under which individuals should engage in an exclusive search for host plants or receptive females. Parameterized for M. jurtina, the model predicted males should engage exclusively in mate search, but females only when searching is very efficient. The model provides a framework for analysing the searching behaviour of other butterfly species.

Dispersal ability is key to species persistence in times of environmental change. Assessing a species' vulnerability and response to anthropogenic changes is often performed using one of two methods: correlative approaches that infer dispersal potential based on traits, such as wingspan or an index of mobility derived from expert opinion, or a mechanistic modeling approach that extrapolates displacement rates from empirical data on short‐term movements.Here, we compare and evaluate the success of the correlative and mechanistic approaches using a mechanistic random‐walk model of butterfly movement that incorporates relationships between wingspan and sex‐specific movement behaviors.The model was parameterized with new data collected on four species of butterfly in the south of England, and we observe how wingspan relates to flight speeds, turning angles, flight durations, and displacement rates.We show that flight speeds and turning angles correlate with wingspan but that to achieve good prediction of displacement even over 10 min the model must also include details of sex‐ and species‐specific movement behaviors.We discuss what factors are likely to differentially motivate the sexes and how these could be included in mechanistic models of dispersal to improve their use in ecological forecasting., Dispersal is often predicted using correlations with species traits. Here, we present new data that evaluate the success of the trait approach for predicting the movement behavior of four species of butterfly. We show that wingspan correlates well with flight speeds and turning angles but that to achieve good prediction of displacement over 10 min the model must also include the proportion of time spent flying.

The importance of wild bees for crop pollination is well established, but less is known about which species contribute to service delivery to inform agricultural management, monitoring and conservation. Using sites in Great Britain as a case study, we use a novel qualitative approach combining ecological information and field survey data to establish a national list of crop pollinating bees for four economically important crops (apple, field bean, oilseed rape and strawberry). A traits data base was used to establish potential pollinators, and combined with field data to identify both dominant crop flower visiting bee species and other species that could be important crop pollinators, but which are not presently sampled in large numbers on crops flowers. Whilst we found evidence that a small number of common, generalist species make a disproportionate contribution to flower visits, many more species were identified as potential pollinators, including rare and specialist species. Furthermore, we found evidence of substantial variation in the bee communities of different crops. Establishing a national list of crop pollinators is important for practitioners and policy makers, allowing targeted management approaches for improved ecosystem services, conservation and species monitoring. Data can be used to make recommendations about how pollinator diversity could be promoted in agricultural landscapes. Our results suggest agri-environment schemes need to support a higher diversity of species than at present, notably of solitary bees. Management would also benefit from targeting specific species to enhance crop pollination services to particular crops. Whilst our study is focused upon Great Britain, our methodology can easily be applied to other countries, crops and groups of pollinating insects., SCOPUS: ar.j, info:eu-repo/semantics/published

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.

Carbonic anhydrase (CA) mimics are often studied with a focus on the hydration of CO2 for atmospheric carbon capture. Consequently, the reverse reaction (dehydration of HCO3–) has received minimal attention, so much so that the rate-limiting step of the dehydration reaction in CA mimics is currently unknown. The rate-limiting step of the hydration reaction is reported to be the bicarbonate-bound intermediate step, and thus is susceptible to product inhibition. It is not, however, clear if this inhibition is a consequence of an increase in the rate of the competing dehydration reaction or resulting from the strong affinity of bicarbonate to the mimic. To address this, insight into the dehydration reaction kinetics is needed. We therefore report the most comprehensive study of a CA mimic to date. The dehydration profile of the fastest small-molecule CA mimic, ZnL1S, was characterized, and consequently evidence for the rate-limiting step for the dehydration reaction was seen to be the bicarbonate-bound intermediate step, much like the hydration reaction. This experimental validation of the rate-limiting step was achieved through a variety of methods including NMR experiments and the effect of inhibitors, substrate concentration, and metal center on activity. With this understanding, an improvement in the favorability of the rate-limiting step was achieved, resulting in decreased bicarbonate inhibition. Thus, an increase in the mimic’s kcat for both reactions was observed, resulting in the largest rate constants of any small-molecule CA mimic reported to date (28 093 and 579 M–1 s–1 for hydration and dehydration, respectively). Enzyme-like kcat/km values were obtained for ZnL1S (5.9 × 105 M–1 s–1 for CO2 hydration), and notably there is only a difference of 2.5 orders of magnitude from the enzyme, the closest of any CA mimic reported in the literature. The results from this work can be applied to the development and improvement of future and existing mimics toward attaining increased activities.

1. Pollination is a key ecosystem service for global agriculture but evidence of pollinator populationdeclines is growing. Reliable spatial modelling of pollinator abundance is essential if we are toidentify areas at risk of pollination service deficit and effectively target resources to support pollinatorpopulations. Many models exist which predict pollinator abundance but few have been calibratedagainst observational data from multiple habitats to ensure their predictions are accurate. 2. We selected the most advanced process-based pollinator abundance model available and calibrated it for bumblebees and solitary bees using survey data collected at 239 sites across GreatBritain. We compared three versions of the model: one parameterised using estimates based on expert opinion, one where the parameters are calibrated using a purely data-driven approach and onewhere we allow the expert opinion estimates to inform the calibration process. 3. All three model versions showed significant agreement with the survey data, demonstrating thismodel’s potential to reliably map pollinator abundance. However, there were significant differencesbetween the nesting/floral attractiveness scores obtained by the two calibration methods and fromthe original expert opinion scores. 4. Our results highlight a key universal challenge of calibrating spatially-explicit, process-basedecological models. Notably, the desire to reliably represent complex ecological processes in finelymapped landscapes necessarily generates a large number of parameters, which are challenging to calibrate with ecological and geographical data that is often noisy, biased, asynchronous and sometimesinaccurate. Purely data-driven calibration can therefore result in unrealistic parameter values, despite appearing to improve model-data agreement over initial expert opinion estimates. We thereforeadvocate a combined approach where data-driven calibration and expert opinion are integrated intoan iterative Delphi-like process, which simultaneously combines model calibration and credibility assessment. This may provide the best opportunity to obtain realistic parameter estimates and reliablemodel predictions for ecological systems with expert knowledge gaps and patchy ecological data.

1. Sustainable agriculture aims to produce sufficient food while minimizing environmental damage. To achieve this, we need to understand the role of agricultural landscapes in providing diverse ecosystem services and how these affect crop production and resilience, that is, maintaining yields despite environmental perturbation. 2. We used 10 years of English wheat yield data to derive three metrics of resilience (relative yield across the time series, yield stability around a moving average and resistance to an extreme weather event) at 10 km × 10 km resolution. We used remotely sensed maps to calculate measures of landscape structure, including composition (proportions of different land cover types) and configuration (metrics of connectivity and proximity), known to affect ecosystem service delivery (e.g. control of pests by beneficial invertebrates). We then used an information‐theoretic approach to identify the best‐fitting combination of landscape structure predictors for each resilience metric, using a potential yield model to account for the effects of climate and soils. 3. Relative yield showed a strongly positive relationship with the area of arable land. For yield stability, this relationship was evident but alongside other landscape structure variables in the best‐fitting model. No relationship with arable land was evident for resistance. 4. Yield stability showed a strongly positive effect of proximity to semi‐natural habitats. For resistance, the best‐fitting model included positive relationships with the cover of semi‐natural habitats and proximity to semi‐natural grasslands. 5. Synthesis and applications. Landscapes with the highest relative wheat yields did not show the highest yield stability or resistance to extreme events. As resilience metrics were derived from shorter portions of the time series, the importance of semi‐natural habitats compared to arable land increased. This is probably driven by the complex interplay between landscape structure, agricultural management and ecosystem services. These results demonstrate that measuring relative yield over time may be insufficient to capture the full effect that non‐arable components of the landscape, and the ecosystem services they deliver, have on other aspects of resilience, and that there are clear trade‐offs in managing agricultural landscapes to maximize different aspects of crop yield resilience.

The European Union's Natura 2000 (N2000) is among the largest international networks of protected areas. One of its aims is to secure the status of a predetermined set of (targeted) bird and butterfly species. However, nontarget species may also benefit from N2000. We evaluated how the terrestrial component of this network affects the abundance of nontargeted, more common bird and butterfly species based on data from long-term volunteer-based monitoring programs in 9602 sites for birds and 2001 sites for butterflies. In almost half of the 155 bird species assessed, and particularly among woodland specialists, abundance increased (slope estimates ranged from 0.101 [SD 0.042] to 3.51 [SD 1.30]) as the proportion of landscape covered by N2000 sites increased. This positive relationship existed for 27 of the 104 butterfly species (estimates ranged from 0.382 [SD 0.163] to 4.28 [SD 0.768]), although most butterflies were generalists. For most species, when land-cover covariates were accounted for these positive relationships were not evident, meaning land cover may be a determinant of positive effects of the N2000 network. The increase in abundance as N2000 coverage increased correlated with the specialization index for birds, but not for butterflies. Although the N2000 network supports high abundance of a large spectrum of species, the low number of specialist butterflies with a positive association with the N2000 network shows the need to improve the habitat quality of N2000 sites that could harbor open-land butterfly specialists. For a better understanding of the processes involved, we advocate for standardized collection of data at N2000 sites.Efectos de Natura 2000 sobre las Especies No Focales de Aves y Mariposas con Base en Datos de Ciencia Ciudadana Resumen La red Natura 2000 (N2000) de la Unión Europea está entre las redes internacionales más grandes de áreas protegidas. Uno de sus objetivos es asegurar el estado de un conjunto predeterminado de especies de aves y mariposas (focales). Sin embargo, las especies no focales también pueden beneficiarse con la N2000. Evaluamos cómo el componente terrestre de esta red afecta la abundancia de las especies de aves y mariposas no focales más comunes con base en los datos de programas de monitoreo voluntario a largo plazo en 9,602 sitios para aves y en 2,001 sitios para mariposas. En casi la mitad de las 155 especies de aves evaluadas, particularmente entre aquellas especies especialistas en zonas boscosas, la abundancia incrementó (los estimaciones de la pendiente variaron desde 0.101 [DS 0.042] hasta 3.51 [DS 1.30]) conforme incrementó la proporción del paisaje cubierto por sitios de la N2000. Esta relación positiva existió en 27 de las 104 especies de mariposas (con una variación de estimaciones desde 0.382 [DS 0.163] hasta 4.28 [DS 0.768]), aunque la mayoría de las especies de mariposas fueron generalistas. Cuando se consideraron las covarianzas de cobertura de suelo estas relaciones positivas no fueron evidentes para la mayoría de las especies, lo que significa que la cobertura de suelo puede ser una determinante de los efectos positivos de la red N2000. El incremento en la abundancia conforme aumentó la cobertura de la N2000 estuvo correlacionado con el índice de especialización de las aves, pero no el de las mariposas. Aunque la red N2000 sostiene la abundancia alta de un espectro amplio de especies, el bajo número de mariposas especialistas con una asociación positiva a la red N2000 demuestra la necesidad de mejorar la calidad del hábitat de los sitios N2000 que podrían albergar a mariposas especialistas de campo abierto. Para un mejor entendimiento de los procesos involucrados, promovemos una recolección estandarizada de datos en los sitios de la red N2000.欧盟的 Natura 2000(N2000) 是全球最大的保护地网络之一, 它的目标之一是保护一批目标鸟类和蝴蝶物种的现状。不过, 非目标物种也可能从 N2000 中获益。本研究评估了该网络陆地保护区如何影响更为常见的非目标鸟类和蝴蝶的丰度, 所用数据来自长期的志愿者监测项目, 包括了 9602 个鸟类监测点和 2001 个蝴蝶监测点。在评估的 155 种鸟类中, 几乎一半的鸟类, 特别是林地专性种, 在 N2000 位点覆盖的景观比例上升时数量增加 (斜率估计值在 0.101 [SD 0.042] 到3.51 [SD 1.30] 之间) 。虽然大多数蝴蝶都是广幅种, 但 104 种蝴蝶中也有 27 种存在这种正相关关系 (斜率估计值在 0.382 [SD 0.163] 到 4.28 [SD 0.768]) 。对于大多数物种来说, 当考虑土地覆盖协变量时, 这样的正相关关系并不明显, 这意味着土地覆盖可能是 N2000 网络能否对物种产生积极影响的关键因素。物种的丰度随着 N2000 覆盖度增加而增加的现象与鸟类的生境特化指数相关, 而在蝴蝶中则无关。虽然 N2000 网络使许多物种得以维持较高丰度, 但获得其积极影响的蝴蝶专性种数量仍较少, 这说明需要在 N2000 网络中提高专性生活在开阔地的蝴蝶的生境质量。为了更好地理解其中的过程, 我们建议收集更多标准化的 N2000 位点数据。 【翻译: 胡怡思; 审校: 聂永刚】.

Dispersal is a key process affecting population persistence and major factors affecting dispersal rates are the amounts, connectedness and properties of habitats in landscapes. We present new data on the butterfly Maniola jurtina in flower-rich and flower-poor habitats that demonstrates how movement and behaviour differ between sexes and habitat types, and how this effects consequent dispersal rates. Females had higher flight speeds than males, but their total time in flight was four times less. The effect of habitat type was strong for both sexes, flight speeds were ~ 2.5 × and ~ 1.7 × faster on resource-poor habitats for males and females, respectively, and flights were approximately 50% longer. With few exceptions females oviposited in the mown grass habitat, likely because growing grass offers better food for emerging caterpillars, but they foraged in the resource-rich habitat. It seems that females faced a trade-off between ovipositing without foraging in the mown grass or foraging without ovipositing where flowers were abundant. We show that taking account of habitat-dependent differences in activity, here categorised as flight or non-flight, is crucial to obtaining good fits of an individual-based model to observed movement. An important implication of this finding is that incorporating habitat-specific activity budgets is likely necessary for predicting longer-term dispersal in heterogeneous habitats, as habitat-specific behaviour substantially influences the mean (> 30% difference) and kurtosis (1.4 × difference) of dispersal kernels. The presented IBMs provide a simple method to explicitly incorporate known activity and movement rates when predicting dispersal in changing and heterogeneous landscapes. Electronic supplementary material The online version of this article (10.1007/s00442-020-04638-4) contains supplementary material, which is available to authorized users.

Species interactions have a spatio‐temporal component driven by environmental cues, which if altered by climate change can drive shifts in community dynamics. There is insufficient understanding of the precise time‐windows during which inter‐annual variation in weather drives phenological shifts and the consequences for mismatches between interacting species and resultant population dynamics – particularly for insects. We use a 20‐year study on a tri‐trophic system: sycamore Acer pseudoplatanus, two associated aphid species Drepanosiphum platanoidis and Periphyllus testudinaceus, and their hymenopteran parasitoids. Using a sliding window approach, we assess climatic drivers of phenology in all three trophic levels. We quantify the magnitude of resultant trophic mismatches between aphids and their plant hosts and parasitoids, and then model the impacts of these mismatches, direct weather effects and density dependence on local‐scale aphid population dynamics. Warmer temperatures in mid‐March to late‐April were associated with advanced sycamore budburst, parasitoid attack and (marginally) D. platanoidis emergence. The precise time‐window during which spring weather advances phenology varies considerably across each species. Crucially, warmer temperatures in late winter delayed the emergence of both aphid species. Seasonal variation in warming rates thus generate marked shifts in the relative timing of spring events across trophic levels and mismatches in the phenology of interacting species. Despite this, we found no evidence that aphid population growth rates were adversely impacted by the magnitude of mismatch with their host plants or parasitoids, or direct impacts of temperature and precipitation. Strong density dependence effects occurred in both aphid species and probably buffered populations, through density dependent compensation, from adverse impacts of the marked inter‐annual climatic variation that occurred during the study period. These findings explain the resilience of aphid populations to climate change and uncover a key mechanism, warmer winter temperatures delaying insect phenology, by which climate change drives asynchronous shifts between interacting species.

Oxygenic photosynthesis starts with the oxidation of water to O2, a light-driven reaction catalysed by photosystem II. Cyanobacteria are the only prokaryotes capable of water oxidation and therefore, it is assumed that the origin of oxygenic photosynthesis is a late innovation relative to the origin of life and bioenergetics. However, when exactly water oxidation originated remains an unanswered question. Here we use phylogenetic analysis to study a gene duplication event that is unique to photosystem II: the duplication that led to the evolution of the core antenna subunits CP43 and CP47. We compare the changes in the rates of evolution of this duplication with those of some of the oldest well-described events in the history of life: namely, the duplication leading to the Alpha and Beta subunits of the catalytic head of ATP synthase, and the divergence of archaeal and bacterial RNA polymerases and ribosomes. We also compare it with more recent events such as the duplication of Cyanobacteria-specific FtsH metalloprotease subunits and the radiation leading to Margulisbacteria, Sericytochromatia, Vampirovibrionia, and other clades containing anoxygenic phototrophs. We demonstrate that the ancestral core duplication of photosystem II exhibits patterns in the rates of protein evolution through geological time that are nearly identical to those of the ATP synthase, RNA polymerase, or the ribosome. Furthermore, we use ancestral sequence reconstruction in combination with comparative structural biology of photosystem subunits, to provide additional evidence supporting the premise that water oxidation had originated before the ancestral core duplications. Our work suggests that photosynthetic water oxidation originated closer to the origin of life and bioenergetics than can be documented based on phylogenetic or phylogenomic species trees alone., Highlights • The origin of oxygenic photosynthesis cannot be located on a species tree. • Vampirovibrionia, Sericytochromatia and Margulisbacteria lost photosynthesis. • Photosystem II can be as old as the oldest enzymes. • Earliest type II reaction centres were structurally like water-splitting Photosystem II. • Anoxygenic photosynthesis is not more primitive than oxygenic photosynthesis.

Non-technical summary Resilience is a cross-disciplinary concept that is relevant for understanding the sustainability of the social and environmental conditions in which we live. Most research normatively focuses on building or strengthening resilience, despite growing recognition of the importance of breaking the resilience of, and thus transforming, unsustainable social-ecological systems. Undesirable resilience (cf. lock-ins, social-ecological traps), however, is not only less explored in the academic literature, but its understanding is also more fragmented across different disciplines. This disparity can inhibit collaboration among researchers exploring interdependent challenges in sustainability sciences. In this article, we propose that the term lock-in may contribute to a common understanding of undesirable resilience across scientific fields. Technical summary Resilience is an extendable concept that bridges the social and life sciences. Studies increasingly interpret resilience normatively as a desirable property of social-ecological systems, despite growing awareness of resilient properties leading to social and ecological degradation, vulnerability or barriers that hinder sustainability transformations (i.e., 'undesirable' resilience). This is the first study to qualify, quantify and compare the conceptualization of 'desirable' and 'undesirable' resilience across academic disciplines. Our literature analysis found that various synonyms are used to denote undesirable resilience (e.g., path dependency, social-ecological traps, institutional inertia). Compared to resilience as a desirable property, research on undesirable resilience is substantially less frequent and scattered across distinct scientific fields. Amongst synonyms for undesirable resilience, the term lock-in is more frequently and evenly used across academic disciplines. We propose that lock-in therefore has the potential to reconcile diverse interpretations of the mechanisms that constrain system transformation - explicitly and coherently addressing characteristics of reversibility and plausibility - and thus enabling integrative understanding of social-ecological system dynamics. Social media summary 'Lock-in' as a bridging concept for interdisciplinary understanding of barriers to desirable sustainability transitions.

A wide variety of tools aim to support decision making by modelling, mapping and quantifying ecosystem services. If decisions are to be properly informed, the accuracy and potential limitations of these tools must be well understood. However, dedicated studies evaluating ecosystem service models against empirical data are rare, especially over large areas. In this paper, we report on the national-scale assessment of a new ecosystem service model for nutrient delivery and retention, the InVEST Nutrient Delivery Ratio model. For 36 river catchments across the UK, we modelled total catchment export of phosphorus (P) and/or nitrogen (N) and compared model outputs to measurements derived from empirical water chemistry data.\ud The model performed well in terms of relative magnitude of nutrient export among catchments (best Spearman’s rank correlation for N and P, respectively: 0.81 and 0.88). However, there was wide variation among catchments in the accuracy of the model, and absolute values of nutrient exports frequently showed high percentage differences between modelled and empirically-derived exports (best median absolute percentage difference for N and P, respectively: ± 64%, ± 44%). The model also showed a high degree of sensitivity to nutrient loads and hydrologic routing input parameters and these sensitivities varied among catchments. \ud These results suggest that the InVEST model can provide valuable information on nutrient fluxes to decision makers, especially in terms of relative differences among catchments. However, caution is needed if using the absolute modelled values for decision-making. Our study also suggests particular attention should be paid to researching input nutrient loadings and retentions, and the selection of appropriate input data resolutions and threshold flow accumulation values. Our results also highlight how availability of empirical data can improve model calibration and performance assessment and reinforce the need to include such data in ecosystem service modelling studies.

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.

1. Abundance data are the foundation for many ecological and conservation projects, but are only available for a few taxonomic groups. In contrast, distribution records (georeferenced presence records) are more widely available. Here we examine whether year-to-year changes in numbers of distribution records, collated over a large spatial scale, can provide a measure of species' population variability, and hence act as a metric of abundance changes. 2. We used 33 British butterfly species to test this possibility, using distribution and abundance data (transect counts) from 1976 to 2012. 3. Comparing across species, we found a strong correlation between mean year-to-year changes in total number of distribution records and mean year-to-year changes in abundance (N = 33 species; r2 = 0.66). This suggests that annual distribution data can be used to identify species with low versus high population variability. 4. For individual species, there was considerable variation in the strength of relationships between year-to-year changes in total number of distribution records and abundance. Between-year changes in abundance can be identified from distribution records most accurately for species whose populations are most variable (i.e. have high annual variation in numbers of records). 5. We conclude that year-to-year changes in distribution records can indicate overall population variability within a taxon, and are a reasonable proxy for year-to-year changes in abundance for some types of species. This finding opens up more opportunities to inform ecological and conservation studies about population variability, based on the wealth of citizen science distribution records that are available for other taxa.

1. Focusing on food production, in this paper we define resilience in the food security context as maintaining production of sufficient and nutritious food in the face of chronic and acute environmental perturbations. In agri-food systems, resilience is manifest over multiple spatial scales: field, farm, regional and global. Metrics comprise production and nutritional diversity as well as socio-economic stability of food supply. 2. Approaches to enhancing resilience show a progression from more ecologically-based methods at small scales to more socially-based interventions at larger scales. At the field scale, approaches include the use of mixtures of crop varieties, livestock breeds and of forage species, polycultures, and boosting ecosystem functions. Stress-tolerant crops, or with greater plasticity, provide technological solutions. 3. At the farm scale, resilience may be conferred by diversifying crops and livestock and by farmers implementing adaptive approaches in response to perturbations. Biodiverse landscapes may enhance resilience, but the evidence is weak. At regional to global scales, resilient food systems will be achieved by coordination and implementation of resilience approaches among farms, advice to farmers and targeted research. 4. Synthesis. Threats to food production are predicted to increase under climate change and land degradation. Holistic responses are needed that integrate across spatial scales. Ecological knowledge is critical, but should be implemented alongside agronomic solutions and socio-economic transformations.

1. Measuring functional connectivity, the ability of species to move between resource patches, is essential for conservation in fragmented landscapes. However, current methods are highly time consuming and expensive. 2. Population synchrony- the correlation in time series of counts between two long-term population monitoring sites, has been suggested as a possible proxy measure of functional connectivity. To date, population synchrony has been shown to correlate with proxies for movement frequency such as the coverage of suitable habitat types in intervening landscapes, and also least cost distances around hostile land cover types. 3. This provides tentative evidence that synchrony is directly driven by movements of the focal species, but an alternative explanation is that these land cover types affect the movement of interacting species (e.g. natural enemies of the focal species) which can also drive synchronous population dynamics. Therefore, what is needed is a test directly relating population synchrony to movement frequencies of a focal species. 4. Here we use data from a 21 year mark-release-recapture study and show that population synchrony does indeed predict movements of a focal butterfly species Boloria eunomia (Esper). 5. There is growing evidence that population synchrony can be a useful conservation tool to measure functional connectivity.

Background Understanding the factors influencing movement is essential to forecasting species persistence in a changing environment. Movement is often studied using mechanistic models, extrapolating short-term observations of individuals to longer-term predictions, but the role of weather variables such as air temperature and solar radiation, key determinants of ectotherm activity, are generally neglected. We aim to show how the effects of weather can be incorporated into individual-based models of butterfly movement thus allowing analysis of their effects. Methods We constructed a mechanistic movement model and calibrated it with high precision movement data on a widely studied species of butterfly, the meadow brown (Maniola jurtina), collected over a 21-week period at four sites in southern England. Day time temperatures during the study ranged from 14.5 to 31.5 °C and solar radiation from heavy cloud to bright sunshine. The effects of weather are integrated into the individual-based model through weather-dependent scaling of parametric distributions representing key behaviours: the durations of flight and periods of inactivity. Results Flight speed was unaffected by weather, time between successive flights increased as solar radiation decreased, and flight duration showed a unimodal response to air temperature that peaked between approximately 23 °C and 26 °C. After validation, the model demonstrated that weather alone can produce a more than two-fold difference in predicted weekly displacement. Conclusions Individual Based models provide a useful framework for integrating the effect of weather into movement models. By including weather effects we are able to explain a two-fold difference in movement rate of M. jurtina consistent with inter-annual variation in dispersal measured in population studies. Climate change for the studied populations is expected to decrease activity and dispersal rates since these butterflies already operate close to their thermal optimum. Electronic supplementary material The online version of this article (10.1186/s40462-019-0171-7) contains supplementary material, which is available to authorized users.

Many countries have conservation plans for threatened species, but such plans have generally been developed without taking into account the potential impacts of climate change. Here, we apply a decision framework, specifically developed to identify and prioritise climate change adaptation actions and demonstrate its use for 30 species threatened in the UK. Our aim is to assess whether government conservation recommendations remain appropriate under a changing climate. The species, associated with three different habitats (lowland heath, broadleaved woodland and calcareous grassland), were selected from a range of taxonomic groups (primarily moths and vascular plants, but also including bees, bryophytes, carabid beetles and spiders). We compare the actions identified for these threatened species by the decision framework with those included in existing conservation plans, as developed by the UK Government’s statutory adviser on nature conservation. We find that many existing conservation recommendations are also identified by the decision framework. However, there are large differences in the spatial prioritisation of actions when explicitly considering projected climate change impacts. This includes recommendations for actions to be carried out in areas where species do not currently occur, in order to allow them to track movement of suitable conditions for their survival. Uncertainties in climate change projections are not a reason to ignore them. Our results suggest that existing conservation plans, which do not take into account potential changes in suitable climatic conditions for species, may fail to maximise species persistence. Comparisons across species also suggest a more habitat-focused approach could be adopted to enable climate change adaptation for multiple species.

Understanding spatial variation in the structure and stability of plant-pollinator networks, and their relationship with anthropogenic drivers, is key to maintaining pollination services and mitigating declines. Constructing sufficient networks to examine patterns over large spatial scales remains challenging. Using biological records (citizen science), we constructed potential plant-pollinator networks at 10km resolution across Great Britain, comprising all potential interactions inferred from recorded floral visitation and species co-occurrence. We calculated network metrics (species richness, connectance, pollinator and plant generality) and adapted existing methods to assess robustness to sequences of simulated plant extinctions across multiple networks. We found positive relationships between agricultural land cover and both pollinator generality and robustness to extinctions under several extinction scenarios. Increased robustness was attributable to changes in plant community composition (fewer extinction-prone species) and network structure (increased pollinator generality). Thus, traits enabling persistence in highly agricultural landscapes can confer robustness to potential future perturbations on plant-pollinator networks.

1.Planning for nature conservation has increasingly emphasised the concepts of resilience and spatial networks. Although the importance of habitat networks for individual species is clear, their significance for long‐term ecological resilience and multi‐species conservation strategies is less established. 2.Referencing spatial network theory, we describe the conceptual basis for defining and assessing a network of wildlife areas that supports species’ resilience to multiple forms of perturbations and pressures. We explore actions that could enhance network resilience at a range of scales, based on ecological principles, with reference to four well‐established strategies for intervention in a spatial network (“Better, Bigger, More and Joined”) from the influential Making Space for Nature report by Lawton et al. (2010). 3.Building existing theory into useable and scalable approaches applicable to large numbers of species is challenging but tractable. We illustrate the policy context, describe the elements of a long‐term adaptive management plan and provide example actions, metrics and targets for early implementation using England as a case study, where there is an opportunity to include large‐scale ecological planning in a newly launched 25‐year environment plan. 4.Policy implications. The concept of resilient ecological networks has attracted scientific and political support, but there is no consensus on what a resilient network would look like, or how to assess it. Therefore, it is unclear whether existing targets for action will be sufficient to achieve network resilience. We show that the scientific principles to place resilience and network theory at the heart of large‐scale and long‐term environmental planning are established and ready to implement in practice. Delivering a resilient network to support nature recovery is achievable and can be integrated with ongoing conservation actions and targets, by assessing their effectiveness on properties of the entire network. England's 25 Year Environment Plan promises to deliver a natural environment that is protected and enhanced for the future and so provides the ideal testbed.

The importance of Cultural Ecosystem Services (CES) to human wellbeing is widely recognised. However, quantifying these non-material benefits is challenging and consequently they are often not assessed. Mapping approaches are increasingly being used to understand the spatial distribution of different CES and how this relates to landscape characteristics. This study uses an online Public Participation Geographic Information System (PPGIS) to elicit information on outdoor locations important to respondents in Wiltshire, a dynamic lowland landscape in southern England. We analysed these locations in a GIS with spatial datasets representing potential influential factors, including protected areas, land use, landform, and accessibility. We assess these characteristics at different spatial and visual scales for different types of cultural engagement. We find that areas that are accessible, near to urban centres, with larger views, and a high diversity of protected habitats, are important for the delivery of CES. Other characteristics including a larger area of woodland and the presence of sites of historic interest in the surrounding landscape were also influential. These findings have implications for land-use planning and the management of ecosystems, by demonstrating the benefits of high quality ecological sites near to towns. The importance of maintaining and restoring landscape features, such as woodlands, to enhance the delivery of CES were also highlighted. [Abstract copyright: Crown Copyright © 2017. Published by Elsevier Ltd. All rights reserved.]

The intensification of agricultural practices throughout the twentieth century has had large detrimental effects on biodiversity and these are likely to increase as the human population rises, with consequent pressure on land. To offset these negative impacts, agri-environment schemes have been widely implemented, offering financial incentives for land-owners to create or maintain favourable habitats that enhance or maintain biodiversity. While some evidence is available on the resulting species richness and abundance for groups such as natural predators, pollinating insects including butterflies and moths, this is costly to obtain and it is difficult to predict the effects of specific habitat designs. To alleviate this problem we here develop an individual-based model (IBM), modelling the detailed movement behaviour, foraging, and energy budget of a grassland butterfly Maniola jurtina Linn. in patches of varying dimensions and quality. The IBM is successfully validated against data on M. jurtina densities, movement behaviour, resource use, fecundity and lifespan in habitats of varying quality. We use the IBM to quantify the benefits for life-history outcomes of M. jurtina of increasing the quantity and the quality of field margins within agricultural landscapes. We find that increasing the quantity of field margin habitat from 1 to 3 ha per 100 ha, as recommended in agri-environment schemes, increases the average number of eggs laid across a two-week period by 60% and adds an extra day to the average lifespan. Similar effects are reported for variation in the quality of field margins. We discuss the implications of the result for modelling butterfly responses to management scenarios.

A variety of tools have emerged with the goal of mapping the current delivery of ecosystem services and quantifying the impact of environmental changes. An important and often overlooked question is how accurate the outputs of these models are in relation to empirical observations. In this paper we validate a hydrological ecosystem service model (InVEST Water Yield Model) using widely available data. We modelled annual water yield in 22 UK catchments with widely varying land cover, population and geology, and compared model outputs with gauged river flow data from the UK National River Flow Archive. Values for input parameters were selected from existing literature to reflect conditions in the UK and were subjected to sensitivity analyses. We also compared model performance between precipitation and potential evapotranspiration data sourced from global- and UK-scale datasets. We then tested the transferability of the results within the UK by additional validation in a further 20 catchments.Whilst the model performed only moderately with global-scale data (linear regression of modelled total water yield against empirical data; slope = 0.763, intercept = 54.45, R2 = 0.963) with wide variation in performance between catchments, the model performed much better when using UK-scale input data, with closer fit to the observed data (slope = 1.07, intercept = 3.07, R2 = 0.990). With UK data the majority of catchments showed less than 10% difference between measured and modelled water yield but there was a minor but consistent overestimate per hectare (86 m3/ha/year). Additional validation on a further 20 UK catchments was similarly robust, indicating that these results are transferable within the UK. These results suggest that relatively simple models can give accurate measures of ecosystem services. However, the choice of input data is critical and there is a need for further validation in other parts of the world.

Climate change is increasingly altering the composition of ecological communities, in combination with other environmental pressures such as high‐intensity land use. Pressures are expected to interact in their effects, but the extent to which intensive human land use constrains community responses to climate change is currently unclear. A generic indicator of climate change impact, the community temperature index (CTI), has previously been used to suggest that both bird and butterflies are successfully ‘tracking’ climate change. Here, we assessed community changes at over 600 English bird or butterfly monitoring sites over three decades and tested how the surrounding land has influenced these changes. We partitioned community changes into warm‐ and cold‐associated assemblages and found that English bird communities have not reorganized successfully in response to climate change. CTI increases for birds are primarily attributable to the loss of cold‐associated species, whilst for butterflies, warm‐associated species have tended to increase. Importantly, the area of intensively managed land use around monitoring sites appears to influence these community changes, with large extents of intensively managed land limiting ‘adaptive’ community reorganization in response to climate change. Specifically, high‐intensity land use appears to exacerbate declines in cold‐adapted bird and butterfly species, and prevent increases in warm‐associated birds. This has broad implications for managing landscapes to promote climate change adaptation., This study investigates how bird and butterfly communities have changed during three decades of climate warming. We present the first evidence to show that community changes appear to be constrained by high‐intensity land use. This has broad implications for managing landscapes to promote climate change adaptation.

How much of something do we need to keep people safe and well? This question is frequently asked by those working in risk management. Across diverse sectors from flood protection to health care, practitioners assess risk as the product of the impact of a given event and the probability of its occurrence. Although these estimates are often uncertain, policy-makers must ultimately make spending decisions aimed at averting these risks, because the costs of inaction to society can be substantial. Biodiversity loss is a similarly critical, yet uncertain, issue. On page 288 of this issue, Newbold et al. ( 1 ) quantify global biodiversity losses, providing much-needed information on the encroachment of proposed “safe limits.”

Drought events are projected to increase in frequency and magnitude, which may alter the composition of ecological communities. Using a functional community metric that describes abundance, life history traits and conservation status, based upon Grime's CSR (Competitive – Stress tolerant – Ruderal) scheme, we investigated how British butterfly communities changed during an extreme drought in 1995. Throughout Britain, the total abundance of these insects had a significant tendency to increase, accompanied by substantial changes in community composition, particularly in more northerly, wetter sites. Communities tended to shift away from specialist, vulnerable species, and towards generalist, widespread species and, in the year following, communities had yet to return to equilibrium. Importantly, heterogeneity in surrounding landscapes mediated community responses to the drought event. Contrary to expectation, however, community shifts were more extreme in areas of greater topographic diversity, whilst land-cover diversity buffered community changes and limited declines in vulnerable specialist butterflies.

The rapid expansion of systematic monitoring schemes necessitates robust methods to reliably assess species' status and trends. Insect monitoring poses a challenge where there are strong seasonal patterns, requiring repeated counts to reliably assess abundance. Butterfly monitoring schemes (BMSs) operate in an increasing number of countries with broadly the same methodology, yet they differ in their observation frequency and in the methods used to compute annual abundance indices. Using simulated and observed data, we performed an extensive comparison of two approaches used to derive abundance indices from count data collected via BMS, under a range of sampling frequencies. Linear interpolation is most commonly used to estimate abundance indices from seasonal count series. A second method, hereafter the regional generalized additive model (GAM), fits a GAM to repeated counts within sites across a climatic region. For the two methods, we estimated bias in abundance indices and the statistical power for detecting trends, given different proportions of missing counts. We also compared the accuracy of trend estimates using systematically degraded observed counts of the Gatekeeper Pyronia tithonus (Linnaeus 1767). The regional GAM method generally outperforms the linear interpolation method. When the proportion of missing counts increased beyond 50%, indices derived via the linear interpolation method showed substantially higher estimation error as well as clear biases, in comparison to the regional GAM method. The regional GAM method also showed higher power to detect trends when the proportion of missing counts was substantial.Synthesis and applications. Monitoring offers invaluable data to support conservation policy and management, but requires robust analysis approaches and guidance for new and expanding schemes. Based on our findings, we recommend the regional generalized additive model approach when conducting integrative analyses across schemes, or when analysing scheme data with reduced sampling efforts. This method enables existing schemes to be expanded or new schemes to be developed with reduced within-year sampling frequency, as well as affording options to adapt protocols to more efficiently assess species status and trends across large geographical scales.Monitoring offers invaluable data to support conservation policy and management, but requires robust analysis approaches and guidance for new and expanding schemes. Based on our findings, we recommend the regional generalized additive model approach when conducting integrative analyses across schemes, or when analysing scheme data with reduced sampling efforts. This method enables existing schemes to be expanded or new schemes to be developed with reduced within-year sampling frequency, as well as affording options to adapt protocols to more efficiently assess species status and trends across large geographical scales.

In our recent Research Article “Individualistic sensitivities and exposure to climate change explain variation in species’ distribution and abundance changes” (1), we presented an analysis relating species-specific measures of sensitivity and exposure to climate, to species’ recent population changes. While our measure and interpretation of species’ climate sensitivity remain correct, we now recognize our interpretation of the exposure measure was inaccurate: Our climate models included an intercept, representing a nonzero average population growth rate; thus, the exposure measure incorporated not only climate effects but other nonclimatic—and potentially unmeasured climatic—effects as well. While our results still demonstrate that a significant proportion of variation in population trends can be explained by exposure and sensitivity, the correct interpretation of the exposure measure means that the explained variation is not solely due to climate. As such, our conclusion that a large proportion of variation in population changes can be explained by individualistic responses to climate is misleading. Given this, and to avoid confusion, we are wholly retracting the Research Article, and we apologize that this was not picked up sooner.

A recent paper by Mori [1] states the need for a unification of studies of ‘engineering’ and ‘ecological’ frameworks of resilience. Engineering resilience focuses on the capacity of a system to recover to equilibrium following some kind of perturbation, while ecological resilience (ER) explicitly recognizes multiple stable states and the capacity for systems to resist ‘regime shifts’ between alternative states. We find Mori's argument somewhat surprising given the number of recent biodiversity–ecosystem functioning (B-EF) studies that incorporate aspects of both resistance and recovery (e.g., see references in [2,3]).

The composition of species communities is changing rapidly through drivers such as habitat loss and climate change, with potentially serious consequences for the resilience of ecosystem functions on which humans depend. To assess such changes in resilience, we analyse trends in the frequency of species in Great Britain that provide key ecosystem functions—specifically decomposition, carbon sequestration, pollination, pest control and cultural values. For 4,424 species over four decades, there have been significant net declines among animal species that provide pollination, pest control and cultural values. Groups providing decomposition and carbon sequestration remain relatively stable, as fewer species are in decline and these are offset by large numbers of new arrivals into Great Britain. While there is general concern about degradation of a wide range of ecosystem functions, our results suggest actions should focus on particular functions for which there is evidence of substantial erosion of their resilience., Global change may affect the resilience of ecosystem functions by altering community composition. Here, Oliver et al. show that in Great Britain since the 1970s there have been significant net declines among animal species that provide key ecosystem functions such as pollination and pest control.

Analysis of butterfly and moth species shows that responses to climate change may be more predictable than previously recognized., The responses of animals and plants to recent climate change vary greatly from species to species, but attempts to understand this variation have met with limited success. This has led to concerns that predictions of responses are inherently uncertain because of the complexity of interacting drivers and biotic interactions. However, we show for an exemplar group of 155 Lepidoptera species that about 60% of the variation among species in their abundance trends over the past four decades can be explained by species-specific exposure and sensitivity to climate change. Distribution changes were less well predicted, but nonetheless, up to 53% of the variation was explained. We found that species vary in their overall sensitivity to climate and respond to different components of the climate despite ostensibly experiencing the same climate changes. Hence, species have undergone different levels of population “forcing” (exposure), driving variation among species in their national-scale abundance and distribution trends. We conclude that variation in species’ responses to recent climate change may be more predictable than previously recognized.