Unveiling the factors shaping the distribution of widely distributed alpine vertebrates, using multi-scale ecological ni

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Unveiling the factors shaping the distribution of widely distributed alpine vertebrates, using multi-scale ecological niche modelling of the bat Plecotus macrobullaris Antton Alberdi, Ostaizka Aizpurua, Joxerra Aihartza and Inazio Garin*

Abstract Several alpine vertebrates share a distribution pattern that extends across the South-western Palearctic but is limited to the main mountain massifs. Although they are usually regarded as cold-adapted species, the range of many alpine vertebrates also includes relatively warm areas, suggesting that factors beyond climatic conditions may be driving their distribution. In this work we first recognize the species belonging to the mentioned biogeographic group and, based on the environmental niche analysis of Plecotus macrobullaris, we identify and characterize the environmental factors constraining their ranges. Distribution overlap analysis of 504 European vertebrates was done using the Sorensen Similarity Index, and we identified four birds and one mammal that share the distribution with P. macrobullaris. We generated 135 environmental niche models including different variable combinations and regularization values for P. macrobullaris at two different scales and resolutions. After selecting the best models, we observed that topographic variables outperformed climatic predictors, and the abruptness of the landscape showed better predictive ability than elevation. The best explanatory climatic variable was mean summer temperature, which showed that P. macrobullaris is able to cope with mean temperature ranges spanning up to 16°C. The models showed that the distribution of P. macrobullaris is mainly shaped by topographic factors that provide rock-abundant and open-space habitats rather than climatic determinants, and that the species is not a cold-adapted, but rather a cold-tolerant eurithermic organism. P. macrobullaris shares its distribution pattern as well as several ecological features with five other alpine vertebrates, suggesting that the conclusions obtained from this study might be extensible to them. We concluded that rock-dwelling and open-space foraging vertebrates with broad temperature tolerance are the best candidates to show wide alpine distribution in the Western Palearctic. Keywords: Alpine distribution, Alpine long-eared bat, Biogeography, Chiroptera, Distribution, Modelling, Mountain long-eared bat, Zoogeography

Introduction Studying the causes shaping distribution patterns is one of the major tasks in biogeography [1]. Many organisms exhibit similar distributions that despite their current ranges could be the result of different historical, ecological and physiological processes. Identifying the common factors constraining their range is key to understanding these geographic analogies. The joint characteristic attribute of * Correspondence: [email protected] Department of Zoology and Animal Cell Biology, Faculty of Science and Technology, University of The Basque Country UPV/EHU, Sarriena auzoa s/n, 48940 Leioa, The Basque Count