The Limited Legacy of Post-Glacial Recolonization in the Floristic Patterns of the European Alps

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Abstract— Past glacial periods were highly disruptive to plant species distributions in mid-latitude mountain belts, such as the European Alps, and drove many species to survive in peripheral or nunatak refugia. Progressive glacial retreat during the late Pleistocene and Holocene is thought to have triggered a mass recolonization event in the European Alps. Here, we asked whether this recolonization event has left a spatial legacy in the floristic patterns of the extant Alpine flora, and if so, how strongly this determines floristic patterns in comparison to present-day landscape and climatic drivers. We built on approximately 6 million data points sourced from the Global Biodiversity Information Facility and herbaria to build plant species ranges within a 10 × 10 km grid system laid across the European Alps, and then calculated species assemblage turnover across the grid. Additionally, we used a recent species-level molecular phylogeny encompassing over 80% of the Alpine flora to calculate standardized phylogenetic diversity, phylogenetic endemism, and phylogenetic turnover between grid cells. Using spatial autoregressive and generalised dissimilarity modelling, we showed that all floristic measures vary significantly, albeit weakly, with three key post-glacial variables: climate change velocity, time since deglaciation, and distance from the nearest refugium. Unexpectedly, differences in species’ dispersal ability did not explain these patterns. The post-glacial variables have little explanatory power relative to contemporary climate and landscape drivers. We conclude that post-glacial recolonization of the Alps is largely complete for the flora as a whole, and that the spatial structure of the extant flora is primarily driven by the contemporary environmental conditions.

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Compared to species turnover, patterns of phylogenetic turnover provide extra information about the spatial structure of biodiversity, for example providing more informative comparisons between the biota of sites which share no species. To harness this information for broad‐scale spatial analysis, we present phylo‐GDM, a technique for interpolating the spatial structure of phylogenetic turnover between sampled locations in relation to environment, based on generalised dissimilarity modelling (GDM).Using a database of over 150 000 location records for 114 myobatrachid frog species in Australia, linked to a species‐level phylogeny inferred from 2467 base pairs of mitochondrial DNA, we calculated species and phylogenetic turnover between pairs of sites. We show how phylogenetic turnover extended the range of informative comparison of compositional turnover to more biologically and environmentally dissimilar sites. We generated GDM models which predict species and phylogenetic turnover across Australia, and tested the fit of models for different ages within the phylogeny to find the phylogenetic tree depth at which the relationship to current day environment is greatest. We also incorporated explanatory variables based on biogeographic patterns, to represent broad‐scale turnover resulting from divergent evolutionary histories. We found that while the predictive power of our models was lower for full phylogenetic turnover than for species turnover, models based on the more recent components of the phylogeny (closer to the tips) outperformed species models and full phylogenetic models.Phylo‐GDM has considerable potential as a method for incorporating phylogenetic relationships into biodiversity analyses in ways not previously possible. Because phylogenies do not require named taxa, phylo‐GDM may also provide a means of including lineages with poorly resolved taxonomy (e.g. from metagenomic sequencing) into biodiversity planning and phylogeographic analysis.

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Zoogeographical regions and geospatial patterns of phylogenetic diversity and endemism of New World bats
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The analysis of regional scale patterns of diversity allows insights into the processes that have shaped modern biodiversity at the macro‐scale. Previous analyses studying biogeographic regionalisation across different high‐level taxa have shown similar trends at a global scale. However, incorporating phylogenetic methods when comparing biogeographic regionalisation between subgroups facilitates identification of mechanisms leading to the biogeographic distribution of specific taxa. We analysed the spatial trends of phylogenetic diversity and phylogenetic endemism of 325 species of New World bats, using updated range maps of the modern distributions. These analyses showed phylogeographic signals that reflect the different evolutionary histories of these families. Zoogeographical zones were detected based on range‐weighted phylogenetic turnover. Values of high phylogenetic diversity and endemism were distributed differently across families, suggesting niche conservatism, but a general latitudinal trend of diversity was evident across taxa. Overall, two main bioregions were shared across New World bat taxa (Nearctic and Neotropical), with two additional subregions (Andean and La Platan). We found strong support for an additional transitional zone in the Pacific coast of South America for Emballonuridae and Molossidae. Differences in regionalisation across families indicate that niche conservatism, in situ diversification and dispersal ability are major drivers for the regionalisation of New World bats, within a dual‐centre of diversification scenario. We also found strong inter‐familial support for an independent Caribbean biogeographic region.

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Summary Understanding changes of biodiversity across the landscape underlies biogeography and ecology and is important in land management and conservation. Measures of species and phylogenetic turnover used to estimate the rate of change of assemblages between sets of locations are more often influenced by wide‐ranging taxa. Transition zones between regions that are associated with range‐restricted taxa can be obscured by wide‐ranging taxa that span them. We present a set of new range‐weighted metrics of taxon and phylogenetic turnover, as modifications of conventional metrics, where the range‐restricted components of the assemblages are assigned greater weight in the calculations. We show how these metrics are derived from weighted endemism and phylogenetic endemism and demonstrate their properties using a continent‐wide data set of Australian Acacia. The range‐weighted metrics result in better delineated transition zones between regions, in that the rate of turnover is steeper than with conventional turnover measures. These metrics provide important complementary information for the interpretation of spatial turnover patterns derived from conventional turnover metrics. Additionally, the phylogenetic variant incorporates information about phylogenetic relatedness while also not saturating at high values of turnover, thus remaining useful for comparisons over greater distances than conventional turnover metrics.

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Alpine Biodiversity in Europe
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Overview: An Outline of Europe's Alpine Areas.- A Bioclimatic Characterisation of Europe's Alpine Areas.- The High Mountain Vegetation of: the Scandes.- The High Mountain Vegetation of Scotland.- Vegetation of the Giant Mountains, Central Europe.- The Alpine Vegetation of the Alps.- The Alpine Flora and Vegetation of the South-Eastern Carpathians.- The High Mountain Flora and Vegetation of the Apennines and the Italian Alps.- The Vegetation of the Alpine Zone in the Pyrenees.- High Mountain Vegetation of the Caucasus Region.- The Vegetation of the Corsican High Mountains.- The High Mountain Vegetation of the Balkan Peninsula.- Overview: Patterns in Diversity.- TAxonomic Diversity of Vascular Plants in the European Alpine Areas.- Patterns in the Plant Species Richness of European High Mountain Vegetation.- Altitude Ranges and Spatial Patterns of Alpine Plants in Northern Europe.- Vascular Plant and Bryophyte Diversity Along Elevational Gradients in the Alps.- Assessing the Long-term Dynamics of Endemic Plants at Summit Habitats.- Mapping Alpine Vegetation.- A GIS Assessment of Alpine Biodiversity at a Range of Scales.- Overview: Invertebrate Diversity in Europe's Alpine Regions.- The Geographical Distribution of High Mountain Macrolepidoptera in Europe.- High Altitude Invertebrate Diversity in the Ural Mountains.- The Diversity of High Altitude Arachnids (Araneae. Opiliones, Pseudoscorpiones) in the Alps.- Patterns of Butterfly Diversity Above the Timberline in the Italian Alps and Apennines.- Diversity Patterns of Carabids in the Alps and the Apennines.- Overview: Alpine Vertebrates.- Breeding Bird Assemblages and Habitat Use of Alpine Areas in Scotland.- Rodents in the European Alps: Population Ecology and Potential Impacts on Ecosystems.- Large Herbivores in Continental European Alpine Ecosystems: Current Status and Challenges for the Future.- Diversity of Alpine Vertebrates in the Pyrenees and Sierra Nevada, Spain.- The Impacts of Vertebrate Grazers onVegetation in Eurpean High Mountains.- Overview: Alpine Vegetation Dynamics and Climate Change - a Synthesis of Long-term Studies and Observations.- Long-term Changes in Alpine Plant Communities in Norway and Finland.- Vegetation Dynamcis at the Treeline Ecotone in the Ural Highlands, Russia.- Recent Increases in Summit Flora Caused by Warming in the Alps.- The Piz Linard - Europe's Oldest Mountain Vegetation Study Site.- Alpine Biodiversity in Space and Time: A Synthesis

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