A method for the objective selection of landscape‐scale study regions and sites at the national level

1. Ecological processes operating on large spatio-temporal scales are difficult to disentangle with traditional empirical approaches. Alternatively, researchers can take advantage of ‘natural’ experiments, where experimental control is exercised by careful site selection. Recent advances in developing protocols for designing these ‘pseudo-experiments’ commonly do not consider the selection of the focal region and predictor variables are usually restricted to two. Here, we advance this type of site selection protocol to study the impact of multiple landscape scale factors on pollinator abundance and diversity across multiple regions. 2. Using datasets of geographic and ecological variables with national coverage, we applied a novel hierarchical computation approach to select study sites that contrast asmuch as possible in four key variables,while attempting tomaintain regional comparability and national representativeness. There were three main steps to the protocol: (i) selection of six 100 9 100 km2 regions that collectively provided land cover representative of the national land average, (ii) mapping of potential sites into a multivariate space with axes representing four key factors potentially influencing insect pollinator abundance, and (iii) applying a selection algorithm which maximized differences between the four key variables, while controlling for a set of external constraints. 3. Validation data for the site selection metrics were recorded alongside the collection of data on pollinator populations during two field campaigns. While the accuracy of the metric estimates varied, the site selection succeeded in objectively identifying field sites that differed significantly in values for each of the four key variables. Between-variable correlations were also reduced or eliminated, thus facilitating analysis of their separate effects. 4. This study has shown that national datasets can be used to select randomized and replicated field sites objectively within multiple regions and alongmultiple interacting gradients. Similar protocols could be used for studying a range of alternative research questions related to land use or other spatially explicit environmental variables, and to identify networks of field sites for other countries, regions, drivers and response taxa in a wide range of scenarios.

[1]  S. Erasmi,et al.  Landscape composition and configuration differently affect trap-nesting bees, wasps and their antagonists , 2014 .

[2]  D. Goulson,et al.  Evidence for competition between honeybees and bumblebees; effects on bumblebee worker size , 2009, Journal of Insect Conservation.

[3]  Ralph T. Clarke,et al.  ITE Merlewood Land Classification of Great Britain , 1996 .

[4]  J. Diamond Ecology: Laboratory, field and natural experiments , 1983, Nature.

[5]  Pat Willmer,et al.  LINKING BEES AND FLOWERS: HOW DO FLORAL COMMUNITIES STRUCTURE POLLINATOR COMMUNITIES? , 2003 .

[6]  Mark J. Bailey,et al.  Clarity or confusion?: problems in attributing large-scale ecological changes to anthropogenic drivers , 2012 .

[7]  Teja Tscharntke,et al.  How do landscape composition and configuration, organic farming and fallow strips affect the diversity of bees, wasps and their parasitoids? , 2010, The Journal of animal ecology.

[8]  D. King,et al.  Optimizing landscape selection for estimating relative effects of landscape variables on ecological responses , 2013, Landscape Ecology.

[9]  Carsten Thies,et al.  Mixed effects of landscape complexity and farming practice on weed seed removal , 2011 .

[10]  J. Biesmeijer,et al.  Pervasiveness of Parasites in Pollinators , 2012, PloS one.

[11]  A. Ellison,et al.  Quantifying the impact of an extreme climate event on species diversity in fragmented temperate forests: the effect of the October 1987 storm on British broadleaved woodlands , 2014 .

[12]  Theodora Petanidou,et al.  MEASURING BEE DIVERSITY IN DIFFERENT EUROPEAN HABITATS AND BIOGEOGRAPHICAL REGIONS , 2008 .

[13]  Marco Vighi,et al.  Impacts of a pesticide on pollinator species richness at different spatial scales , 2010 .

[14]  A. Holzschuh,et al.  Trait-Specific Responses of Wild Bee Communities to Landscape Composition, Configuration and Local Factors , 2014, PloS one.

[15]  I. Fries,et al.  Seed coating with a neonicotinoid insecticide negatively affects wild bees , 2015, Nature.

[16]  Using historical woodland creation to construct a long‐term, large‐scale natural experiment: the WrEN project , 2016, Ecology and evolution.

[17]  F. Ratnieks,et al.  Long-range foraging by the honey-bee, Apis mellifera L. , 2000 .

[18]  Tim G Benton,et al.  Scale matters: the impact of organic farming on biodiversity at different spatial scales. , 2010, Ecology letters.

[19]  Thomas E. Dilts,et al.  The Landscape Similarity Toolbox: new tools for optimizing the location of control sites in experimental studies , 2010 .

[20]  G. Arnold,et al.  Modes of honeybees exposure to systemic insecticides: estimated amounts of contaminated pollen and nectar consumed by different categories of bees , 2005 .

[21]  T. Tscharntke,et al.  Bumblebees experience landscapes at different spatial scales: possible implications for coexistence , 2006, Oecologia.

[22]  Richard A. Wadsworth,et al.  Final Report for LCM2007 - the new UK land cover map. Countryside Survey Technical Report No 11/07 , 2011 .

[23]  J. Biesmeijer,et al.  The effect of proximity to a honeybee apiary on bumblebee colony fitness, development, and performance , 2014, Apidologie.

[24]  Teja Tscharntke,et al.  Insect pollinated plants benefit from organic farming , 2007 .

[25]  Aníbal Pauchard,et al.  Frontiers inEcology and the Environment Observational approaches in ecology open new ground in a changing world , 2009 .

[26]  Lenore Fahrig,et al.  Functional landscape heterogeneity and animal biodiversity in agricultural landscapes. , 2011, Ecology letters.

[27]  Haldre S. Rogers,et al.  Accidental experiments: ecological and evolutionary insights and opportunities derived from global change , 2013 .

[28]  L. Fahrig,et al.  Sub-optimal study design has major impacts on landscape-scale inference , 2011 .

[29]  Thomas J. Herbert,et al.  Comparisons of forager distributions from matched honey bee colonies in suburban environments , 1994, Behavioral Ecology and Sociobiology.

[30]  Gorm E. Shackelford,et al.  Comparison of pollinators and natural enemies: a meta‐analysis of landscape and local effects on abundance and richness in crops , 2013, Biological reviews of the Cambridge Philosophical Society.

[31]  John Pickering,et al.  Pseudoreplication: a sine qua non for regional ecology , 1992, Landscape Ecology.

[32]  Nigel D. Boatman,et al.  Historical nectar assessment reveals the fall and rise of Britain in bloom , 2015, Nature.