Ecological processes are often spatially and temporally structured, potentially leading to autocorrelation either in environmental variables or species distribution data. Because of that, spatially-biased in-situ samples or predictors might affect the outcomes of ecological models used to infer the geographic distribution of species and diversity. There is a vast heterogeneity of methods and approaches to assess and measure spatial bias; this paper aims at addressing the spatial component of data-driven biases in species distribution modelling, and to propose potential solutions to explicitly test and account for them. Our major goal is not to propose methods to remove spatial bias from the modelling procedure, which would be impossible without proper knowledge of all the processes generating it, but rather to propose alternatives to explore and handle it. In particular, we propose and describe three main strategies that may provide a fair account of spatial bias, namely: (i) how to represent spatial bias; (ii) how to simulate null models based on virtual species for testing biogeographical and species distribution hypotheses; and (iii) how to make use of spatial bias - in particular related to sampling effort - as a leverage instead of a hindrance in species distribution modelling. We link these strategies with good practice in accounting for spatial bias in species distribution modelling.

Biogeography BayCEER University of Bayreuth Universitaetsstraße 30 95440 Bayreuth Germany

BIOME Lab Department of Biological Geological and Environmental Sciences Alma Mater Studiorum University of Bologna via Irnerio 42 40126 Bologna Italy

Center for Biodiversity and Global Change Yale University New Haven CT USA

CICGE Universidade do Porto Porto Portugal

Czech University of Life Sciences Prague Faculty of Environmental Sciences Department of Spatial Sciences Kamýcka 129 Praha Suchdol 16500 Czech Republic

Department of Biogeography and Global Change Museo Nacional de Ciencias Naturales Madrid Spain

Department of Botany Institute of Ecology and Earth Science University of Tartu J Liivi 2 50409 Tartu Estonia

Department of Computer Science and Engineering Alma Mater Studiorum University of Bologna via Irnerio 42 40126 Bologna Italy

Department of Ecology and Evolution University of Lausanne 1015 Lausanne Switzerland

Department of Life Health and Environmental Sciences University of L'Aquila Piazzale Salvatore Tommasi 1 67100 L'Aquila Italy

Department of Plant Ecology Institute of Landscape and Plant Ecology University of Hohenheim Stuttgart Germany

Dept of Ecology and Evolutionary Biology Yale University New Haven CT USA

Evolutionary Ecology and Genetics Group Earth and Life Institute UCLouvain 1348 Louvain la Neuve Belgium

Faculty of Forestry and Wood Sciences Czech University of Life Sciences Prague Prague Suchdol Czech Republic

Federal University of Goiás Campus Central Anápolis Brazil

Georges Lemaître Center for Earth and Climate Research Earth and Life Institute UCLouvain Louvain la Neuve Belgium

Institute of Earth Surface Dynamics University of Lausanne 1015 Lausanne Switzerland

Institute of Geoecology and Geoinformation Adam Mickiewicz University Krygowskiego 10 61 680 Poznan Poland

Knowledge Infrastructures Campus Fryslan University of Groningen Leeuwarden The Netherlands

MARBEC Univ Montpellier CNRS Ifremer IRD Sète France

Rui Nabeiro Biodiversity Chair MED Institute University of Évora Évora Portugal

School of Geography University of Nottingham Nottingham UK

UMR CNRS 7058 Ecologie et Dynamique des Systèmes Anthropisés Université de Picardie Jules Verne 1 Rue des Louvels 80000 Amiens France

University of Leeds Leeds UK

University of Parma Parma Italy

University of Sassari Department of Chemistry Physics Mathematics and Natural Sciences Sassari Italy

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