Pollination by hoverflies in the Anthropocene.

Pollinator declines, changes in land use and climate-induced shifts in phenology have the potential to seriously affect ecosystem function and food security by disrupting pollination services provided by insects. Much of the current research focuses on bees, or groups other insects together as 'non-bee pollinators', obscuring the relative contribution of this diverse group of organisms. Prominent among the 'non-bee pollinators' are the hoverflies, known to visit at least 72% of global food crops, which we estimate to be worth around US$300 billion per year, together with over 70% of animal pollinated wildflowers. In addition, hoverflies provide ecosystem functions not seen in bees, such as crop protection from pests, recycling of organic matter and long-distance pollen transfer. Migratory species, in particular, can be hugely abundant and unlike many insect pollinators, do not yet appear to be in serious decline. In this review, we contrast the roles of hoverflies and bees as pollinators, discuss the need for research and monitoring of different pollinator responses to anthropogenic change and examine emerging research into large populations of migratory hoverflies, the threats they face and how they might be used to improve sustainable agriculture.

Effects of future agricultural change scenarios on beneficial insects.

Insects provide vital ecosystem services to agricultural systems in the form of pollination and natural pest control. However, there are currently widespread declines in the beneficial insects which deliver these services (i.e. pollinators and 'natural enemies' such as predators and parasitoids). Two key drivers of these declines have been the expansion of agricultural land and intensification of agricultural production. With an increasing human population requiring additional sources of food, further changes in agricultural land use appear inevitable. Identifying likely trajectories of change and predicting their impacts on beneficial insects provides a scientific basis for making informed decisions on the policies and practices of sustainable agriculture. We created spatially explicit, exploratory scenarios of potential changes in the extent and intensity of agricultural land use across Great Britain (GB). Scenarios covered 52 possible combinations of change in agricultural land cover (i.e. agricultural expansion or grassland restoration) and intensity (i.e. crop type and diversity). We then used these scenarios to predict impacts on beneficial insect species richness and several metrics of functional diversity at a 10km (hectad) resolution. Predictions were based on species distribution models derived from biological records, comprising data on 116 bee species (pollinators) and 81 predatory beetle species (natural enemies). We identified a wide range of possible consequences for beneficial insect species richness and functional diversity as result of future changes in agricultural extent and intensity. Current policies aimed at restoring semi-natural grassland should result in increases in the richness and functional diversity of both pollinators and natural enemies, even if agricultural practices remain intensive on cropped land (i.e. land-sparing). In contrast, any expansion of arable land is likely to be accompanied by widespread declines in richness of beneficial insects, even if cropping practices become less intensive (i.e. land-sharing), although effects of functional diversity are more mixed.

Landscape simplification weakens the association between terrestrial producer and consumer diversity in Europe.

Land-use change is one of the primary drivers of species loss, yet little is known about its effect on other components of biodiversity that may be at risk. Here, we ask whether, and to what extent, landscape simplification, measured as the percentage of arable land in the landscape, disrupts the functional and phylogenetic association between primary producers and consumers. Across seven European regions, we inferred the potential associations (functional and phylogenetic) between host plants and butterflies in 561 seminatural grasslands. Local plant diversity showed a strong bottom-up effect on butterfly diversity in the most complex landscapes, but this effect disappeared in simple landscapes. The functional associations between plant and butterflies are, therefore, the results of processes that act not only locally but are also dependent on the surrounding landscape context. Similarly, landscape simplification reduced the phylogenetic congruence among host plants and butterflies indicating that closely related butterflies become more generalist in the resources used. These processes occurred without any detectable change in species richness of plants or butterflies along the gradient of arable land. The structural properties of ecosystems are experiencing substantial erosion, with potentially pervasive effects on ecosystem functions and future evolutionary trajectories. Loss of interacting species might trigger cascading extinction events and reduce the stability of trophic interactions, as well as influence the longer term resilience of ecosystem functions. This underscores a growing realization that species richness is a crude and insensitive metric and that both functional and phylogenetic associations, measured across multiple trophic levels, are likely to provide additional and deeper insights into the resilience of ecosystems and the functions they provide.

We need to talk about nonprobability samples.

In most circumstances, probability sampling is the only way to ensure unbiased inference about population quantities where a complete census is not possible. As we enter the era of 'big data', however, nonprobability samples, whose sampling mechanisms are unknown, are undergoing a renaissance. We explain why the use of nonprobability samples can lead to spurious conclusions, and why seemingly large nonprobability samples can be (effectively) very small. We also review some recent controversies surrounding the use of nonprobability samples in biodiversity monitoring. These points notwithstanding, we argue that nonprobability samples can be useful, provided that their limitations are assessed, mitigated where possible and clearly communicated. Ecologists can learn much from other disciplines on each of these fronts.

An operational workflow for producing periodic estimates of species occupancy at national scales.

Policy makers require high-level summaries of biodiversity change. However, deriving such summaries from raw biodiversity data is a complex process involving several intermediary stages. In this paper, we describe an operational workflow for generating annual estimates of species occupancy at national scales from raw species occurrence data, which can be used to construct a range of policy-relevant biodiversity indicators. We describe the workflow in detail: from data acquisition, data assessment and data manipulation, through modelling, model evaluation, application and dissemination. At each stage, we draw on our experience developing and applying the workflow for almost a decade to outline the challenges that analysts might face. These challenges span many areas of ecology, taxonomy, data science, computing and statistics. In our case, the principal output of the workflow is annual estimates of occupancy, with measures of uncertainty, for over 5000 species in each of several defined 'regions' (e.g. countries, protected areas, etc.) of the UK from 1970 to 2019. This data product corresponds closely to the notion of a species distribution Essential Biodiversity Variable (EBV). Throughout the paper, we highlight methodologies that might not be applicable outside of the UK and suggest alternatives. We also highlight areas where the workflow can be improved; in particular, methods are needed to mitigate and communicate the risk of bias arising from the lack of representativeness that is typical of biodiversity data. Finally, we revisit the 'ideal' and 'minimal' criteria for species distribution EBVs laid out in previous contributions and pose some outstanding questions that should be addressed as a matter of priority. Going forward, we hope that this paper acts as a template for research groups around the world seeking to develop similar data products.

ROBITT: A tool for assessing the risk-of-bias in studies of temporal trends in ecology.

Aggregated species occurrence and abundance data from disparate sources are increasingly accessible to ecologists for the analysis of temporal trends in biodiversity. However, sampling biases relevant to any given research question are often poorly explored and infrequently reported; this can undermine statistical inference. In other disciplines, it is common for researchers to complete 'risk-of-bias' assessments to expose and document the potential for biases to undermine conclusions. The huge growth in available data, and recent controversies surrounding their use to infer temporal trends, indicate that similar assessments are urgently needed in ecology.We introduce ROBITT, a structured tool for assessing the 'Risk-Of-Bias In studies of Temporal Trends in ecology'. ROBITT has a similar format to its counterparts in other disciplines: it comprises signalling questions designed to elicit information on the potential for bias in key study domains. In answering these, users will define study inferential goal(s) and relevant statistical target populations. This information is used to assess potential sampling biases across domains relevant to the research question (e.g. geography, taxonomy, environment), and how these vary through time. If assessments indicate biases, then users must clearly describe them and/or explain what mitigating action will be taken.Everything that users need to complete a ROBITT assessment is provided: the tool, a guidance document and a worked example. Following other disciplines, the tool and guidance document were developed through a consensus-forming process across experts working in relevant areas of ecology and evidence synthesis.We propose that researchers should be strongly encouraged to include a ROBITT assessment when publishing studies of biodiversity trends, especially when using aggregated data. This will help researchers to structure their thinking, clearly acknowledge potential sampling issues, highlight where expert consultation is required and provide an opportunity to describe data checks that might go unreported. ROBITT will also enable reviewers, editors and readers to establish how well research conclusions are supported given a dataset combined with some analytical approach. In turn, it should strengthen evidence-based policy and practice, reduce differing interpretations of data and provide a clearer picture of the uncertainties associated with our understanding of reality.

occAssess: An R package for assessing potential biases in species occurrence data.

Species occurrence records from a variety of sources are increasingly aggregated into heterogeneous databases and made available to ecologists for immediate analytical use. However, these data are typically biased, i.e. they are not a probability sample of the target population of interest, meaning that the information they provide may not be an accurate reflection of reality. It is therefore crucial that species occurrence data are properly scrutinised before they are used for research. In this article, we introduce occAssess, an R package that enables straightforward screening of species occurrence data for potential biases. The package contains a number of discrete functions, each of which returns a measure of the potential for bias in one or more of the taxonomic, temporal, spatial, and environmental dimensions. Users can opt to provide a set of time periods into which the data will be split; in this case separate outputs will be provided for each period, making the package particularly useful for assessing the suitability of a dataset for estimating temporal trends in species' distributions. The outputs are provided visually (as ggplot2 objects) and do not include a formal recommendation as to whether data are of sufficient quality for any given inferential use. Instead, they should be used as ancillary information and viewed in the context of the question that is being asked, and the methods that are being used to answer it. We demonstrate the utility of occAssess by applying it to data on two key pollinator taxa in South America: leaf-nosed bats (Phyllostomidae) and hoverflies (Syrphidae). In this worked example, we briefly assess the degree to which various aspects of data coverage appear to have changed over time. We then discuss additional applications of the package, highlight its limitations, and point to future development opportunities.

Enriching the Shared Socioeconomic Pathways to co-create consistent multi-sector scenarios for the UK.

As the pressure to take action against global warming is growing in urgency, scenarios that incorporate multiple social, economic and environmental drivers become increasingly critical to support governments and other stakeholders in planning climate change mitigation or adaptation actions. This has led to the recent explosion of future scenario analyses at multiple scales, further accelerated since the development of the Intergovernmental Panel on Climate Change (IPCC) research community Shared Socioeconomic Pathways (SSPs) and Representative Concentration Pathways (RCPs). While RCPs have been widely applied to climate models to produce climate scenarios at multiple scales for investigating climate change impacts, adaptation and vulnerabilities (CCIAV), SSPs are only recently being scaled for different geographical and sectoral applications. This is seen in the UK where significant investment has produced the RCP-based UK Climate Projections (UKCP18), but no equivalent UK version of the SSPs exists. We address this need by developing a set of multi-driver qualitative and quantitative UK-SSPs, following a state-of-the-art scenario methodology that integrates national stakeholder knowledge on locally-relevant drivers and indicators with higher level information from European and global SSPs. This was achieved through an intensive participatory process that facilitated the combination of bottom-up and top-down approaches to develop a set of UK-specific SSPs that are locally comprehensive, yet consistent with the global and European SSPs. The resulting scenarios balance the importance of consistency and legitimacy, demonstrating that divergence is not necessarily the result of inconsistency, nor comes as a choice to contextualise narratives at the appropriate scale.

Widespread losses of pollinating insects in Britain.

Pollination is a critical ecosystem service underpinning the productivity of agricultural systems across the world. Wild insect populations provide a substantial contribution to the productivity of many crops and seed set of wild flowers. However, large-scale evidence on species-specific trends among wild pollinators are lacking. Here we show substantial inter-specific variation in pollinator trends, based on occupancy models for 353 wild bee and hoverfly species in Great Britain between 1980 and 2013. Furthermore, we estimate a net loss of over 2.7 million occupied 1 km2 grid cells across all species. Declines in pollinator evenness suggest that losses were concentrated in rare species. In addition, losses linked to specific habitats were identified, with a 55% decline among species associated with uplands. This contrasts with dominant crop pollinators, which increased by 12%, potentially in response agri-environment measures. The general declines highlight a fundamental deterioration in both wider biodiversity and non-crop pollination services.

Habitat availability explains variation in climate-driven range shifts across multiple taxonomic groups.

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.

Annual estimates of occupancy for bryophytes, lichens and invertebrates in the UK, 1970-2015.

Here, we determine annual estimates of occupancy and species trends for 5,293 UK bryophytes, lichens, and invertebrates, providing national scale information on UK biodiversity change for 31 taxonomic groups for the time period 1970 to 2015. The dataset was produced through the application of a Bayesian occupancy modelling framework to species occurrence records supplied by 29 national recording schemes or societies (n = 24,118,549 records). In the UK, annual measures of species status from fine scale data (e.g. 1 × 1 km) had previously been limited to a few taxa for which structured monitoring data are available, mainly birds, butterflies, bats and a subset of moth species. By using an occupancy modelling framework designed for use with relatively low recording intensity data, we have been able to estimate species trends and generate annual estimates of occupancy for taxa where annual trend estimates and status were previously limited or unknown at this scale. These data broaden our knowledge of UK biodiversity and can be used to investigate variation in and drivers of biodiversity change.

Efficient occupancy model-fitting for extensive citizen-science data.

Appropriate large-scale citizen-science data present important new opportunities for biodiversity modelling, due in part to the wide spatial coverage of information. Recently proposed occupancy modelling approaches naturally incorporate random effects in order to account for annual variation in the composition of sites surveyed. In turn this leads to Bayesian analysis and model fitting, which are typically extremely time consuming. Motivated by presence-only records of occurrence from the UK Butterflies for the New Millennium data base, we present an alternative approach, in which site variation is described in a standard way through logistic regression on relevant environmental covariates. This allows efficient occupancy model-fitting using classical inference, which is easily achieved using standard computers. This is especially important when models need to be fitted each year, typically for many different species, as with British butterflies for example. Using both real and simulated data we demonstrate that the two approaches, with and without random effects, can result in similar conclusions regarding trends. There are many advantages to classical model-fitting, including the ability to compare a range of alternative models, identify appropriate covariates and assess model fit, using standard tools of maximum likelihood. In addition, modelling in terms of covariates provides opportunities for understanding the ecological processes that are in operation. We show that there is even greater potential; the classical approach allows us to construct regional indices simply, which indicate how changes in occupancy typically vary over a species' range. In addition we are also able to construct dynamic occupancy maps, which provide a novel, modern tool for examining temporal changes in species distribution. These new developments may be applied to a wide range of taxa, and are valuable at a time of climate change. They also have the potential to motivate citizen scientists.

Publisher Correction: The global distribution of tetrapods reveals a need for targeted reptile conservation.

In this Article originally published, owing to a technical error, the author 'Laurent Chirio' was mistakenly designated as a corresponding author in the HTML version, the PDF was correct. This error has now been corrected in the HTML version. Further, in Supplementary Table 3, the authors misspelt the surname of 'Danny Meirte'; this file has now been replaced.

The global distribution of tetrapods reveals a need for targeted reptile conservation.

The distributions of amphibians, birds and mammals have underpinned global and local conservation priorities, and have been fundamental to our understanding of the determinants of global biodiversity. In contrast, the global distributions of reptiles, representing a third of terrestrial vertebrate diversity, have been unavailable. This prevented the incorporation of reptiles into conservation planning and biased our understanding of the underlying processes governing global vertebrate biodiversity. Here, we present and analyse the global distribution of 10,064 reptile species (99% of extant terrestrial species). We show that richness patterns of the other three tetrapod classes are good spatial surrogates for species richness of all reptiles combined and of snakes, but characterize diversity patterns of lizards and turtles poorly. Hotspots of total and endemic lizard richness overlap very little with those of other taxa. Moreover, existing protected areas, sites of biodiversity significance and global conservation schemes represent birds and mammals better than reptiles. We show that additional conservation actions are needed to effectively protect reptiles, particularly lizards and turtles. Adding reptile knowledge to a global complementarity conservation priority scheme identifies many locations that consequently become important. Notably, investing resources in some of the world's arid, grassland and savannah habitats might be necessary to represent all terrestrial vertebrates efficiently.

Agricultural Management and Climatic Change Are the Major Drivers of Biodiversity Change in the UK.

Action to reduce anthropogenic impact on the environment and species within it will be most effective when targeted towards activities that have the greatest impact on biodiversity. To do this effectively we need to better understand the relative importance of different activities and how they drive changes in species' populations. Here, we present a novel, flexible framework that reviews evidence for the relative importance of these drivers of change and uses it to explain recent alterations in species' populations. We review drivers of change across four hundred species sampled from a broad range of taxonomic groups in the UK. We found that species' population change (~1970-2012) has been most strongly impacted by intensive management of agricultural land and by climatic change. The impact of the former was primarily deleterious, whereas the impact of climatic change to date has been more mixed. Findings were similar across the three major taxonomic groups assessed (insects, vascular plants and vertebrates). In general, the way a habitat was managed had a greater impact than changes in its extent, which accords with the relatively small changes in the areas occupied by different habitats during our study period, compared to substantial changes in habitat management. Of the drivers classified as conservation measures, low-intensity management of agricultural land and habitat creation had the greatest impact. Our framework could be used to assess the relative importance of drivers at a range of scales to better inform our policy and management decisions. Furthermore, by scoring the quality of evidence, this framework helps us identify research gaps and needs.

Trait correlates of distribution trends in the Odonata of Britain and Ireland.

A major challenge in ecology is understanding why certain species persist, while others decline, in response to environmental change. Trait-based comparative analyses are useful in this regard as they can help identify the key drivers of decline, and highlight traits that promote resistance to change. Despite their popularity trait-based comparative analyses tend to focus on explaining variation in range shift and extinction risk, seldom being applied to actual measures of species decline. Furthermore they have tended to be taxonomically restricted to birds, mammals, plants and butterflies. Here we utilise a novel approach to estimate occurrence trends for the Odonata in Britain and Ireland, and examine trait correlates of these trends using a recently available trait dataset. We found the dragonfly fauna in Britain and Ireland has undergone considerable change between 1980 and 2012, with 22 and 53% of species declining and increasing, respectively. Distribution region, habitat specialism and range size were the key traits associated with these trends, where habitat generalists that occupy southern Britain tend to have increased in comparison to the declining narrow-ranged specialist species. In combination with previous evidence, we conclude that the lower trend estimates for the narrow-ranged specialists could be a sign of biotic homogenization with ecological specialists being replaced by warm-adapted generalists.

Morphological and geographical traits of the british odonata.

Trait data are fundamental for many aspects of ecological research, particularly for modeling species response to environmental change. We synthesised information from the literature (mainly field guides) and direct measurements from museum specimens, providing a comprehensive dataset of 26 attributes, covering the 43 resident species of Odonata in Britain. Traits included in this database range from morphological traits (e.g. body length) to attributes based on the distribution of the species (e.g. climatic restriction). We measured 11 morphometric traits from five adult males and five adult females per species. Using digital callipers, these measurements were taken from dry museum specimens, all of which were wild caught individuals. Repeated measures were also taken to estimate measurement error. The trait data are stored in an online repository (https://github.com/BiologicalRecordsCentre/Odonata_traits), alongside R code designed to give an overview of the morphometric data, and to combine the morphometric data to the single value per trait per species data.