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Accelerated increase in plant species richness on mountain summits is linked to warming


Globally accelerating trends in societal development and human environmental impacts since the mid-twentieth century1,2,3,4,5,6,7 are known as the Great Acceleration and have been discussed as a key indicator of the onset of the Anthropocene epoch6. While reports on ecological responses (for example, changes in species range or local extinctions) to the Great Acceleration are multiplying8, 9, it is unknown whether such biotic responses are undergoing a similar acceleration over time. This knowledge gap stems from the limited availability of time series data on biodiversity changes across large temporal and geographical extents. Here we use a dataset of repeated plant surveys from 302 mountain summits across Europe, spanning 145 years of observation, to assess the temporal trajectory of mountain biodiversity changes as a globally coherent imprint of the Anthropocene. We find a continent-wide acceleration in the rate of increase in plant species richness, with five times as much species enrichment between 2007 and 2016 as fifty years ago, between 1957 and 1966. This acceleration is strikingly synchronized with accelerated global warming and is not linked to alternative global change drivers. The accelerating increases in species richness on mountain summits across this broad spatial extent demonstrate that acceleration in climate-induced biotic change is occurring even in remote places on Earth, with potentially far-ranging consequences not only for biodiversity, but also for ecosystem functioning and services.

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Fig. 1: Geographical and temporal distribution of studied summits and surveys.
Fig. 2: Average species richness change on mountain summits over time compared to mean annual temperature over time.
Fig. 3: Rate of species richness change over time.
Fig. 4: Rate of species richness change related to the rate of temperature change and precipitation change across all sampled mountains in Europe.


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We thank D. Barolin, J. Birks, A. Björken, C. Björken, S. Dahle, U. Deppe, G. Dussassois, J. V. Ferrández, T. Gassner, S. Giovanettina, F. Giuntoli, Ø. Lunde Heggebø, K. Herz, A. Jost, K. Kallnik, W. Kapfer, T. Kronstad, H. Laukeland, S. Nießner, M. Olson, P. Roux-Fouillet, K. Schofield, M. Suen, D. Watson, J. Wells Abbott, J. Zaremba and numerous additional helpers for fieldwork support; P. Barancˇ ok, J. L. Benito Alonso, M. Camenisch, G. Coldea, J. Dick, M. Gottfried, G. Grabherr, J. I. Holten, J. Kollár, P. Larsson, M. Mallaun, O. Michelsen, U. Molau, M. Pus¸  cas¸ , T. Scheurer, P. Unterluggauer, L. Villar, G.-R. Walther, and numerous helpers for data originating from the GLORIA network13; C. Jenks for linguistic support; and the following institutions for funding. M.J.S.: Danish Carlsbergfondet (CF14-0148), EU Marie Sklodowska-Curie action (grant 707491). C.R., V.S., S.W.: Velux Foundation, Switzerland. C.R., V.S., S.W., J.-P.T., P.V.: Swiss Federal Office for the Environment (FOEN). A.K.: Swiss National Science Foundation (31003A_144011 to C.R.), Basler Stiftung für biologische Forschung, Switzerland. J.K.: Fram Centre, Norway (362202). J.K., J.-A.G., P.C., B.J.: Polish-Norwegian Research Programme of the Norwegian National Centre for Research and Development (Pol-Nor/196829/87/2013). O.F.-A., M.J.H., S.P.: Instituto de Estudios Altoaragoneses (Huesca, Spain). S.D.: Austrian Climate Research Programme (ACRP, project 368575: DISEQU-ALP). F.J.: Botanical Society of Britain & Ireland; Alpine Garden Society, UK. M.J.H.: Felix de Azara research grant (IBERSUMIT project, DPH, Spain). R.K.: Slovak Research and Development Agency (APVV 0866-12). S.N., D.G.: VILLUM Foundation’s Young Investigator Programme (VKR023456; Denmark). S.P.: Ramón y Cajal fellowship (RYC-2013-14164, Ministerio de Economía y Competitividad, Spain). J.-C.S.: European Research Council (ERC-2012-StG-310886-HISTFUNC); VILLUM Investigator project (VILLUM FONDEN grant 16549; Denmark). S.W.: WSL internal grant (201307N0678, Switzerland); EU FP7 Interact Transnational Access (AlpFlor Europe). S.W., S.B., F.J., M.J.H.: Swiss Botanical Society Alpine Flower Fund. Time and effort was supported by sDiv, the Synthesis Centre of iDiv, Germany (DFG FZT 118, sUMMITDiv working group).

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Nature thanks J. Alexander, A. Hester and K. Verheyen for their contribution to the peer review of this work.

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Authors and Affiliations



S.W., M.J.S., J.-A.G., G.J., A.K., C.R., A.D.B., S.H., J.K., K.K., I.K., J.L., S.N., H.P., P.V. and M.W. elaborated the concept; A.K., S.W., M.B.-D., H.P., C.R., P.V. and M.W. organized and harmonized data; M.J.S. implemented the data analyses with support from other authors, particularly S.W., J.-A.G., J.L., J.-C.S., S.D. and D.G.; E.B., S.B., F.T.B., P.C., M.A.D., A.D., B.E., V.A.F., O.F.-A., K.F.F., D.G.-G., E.T.G., J.-A.G., S.V.H., H.H., M.J.H., B.J., F.J., R.K., K.K., J.K., A.L., M.M., U.M.d.C., A.O., S.L.O., S.P., H.P., M.P., V.P., B.S., K.S., V.S., C.R., G.T., J.-P.T., P.V., S.J.W., S.W and N.E.Z. contributed data. M.J.S. led the manuscript writing, with contributions from all authors.

Corresponding authors

Correspondence to Manuel J. Steinbauer or Sonja Wipf.

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Extended data figures and tables

Extended Data Fig. 1 Visualizing richness change.

This conceptual figure shows the approach implemented in the main text to visualize richness change over time based on the raw data (Figs. 2, 3). a, The mean richness change per year (ΔSR/∆t = (SRt2 − SRt1)/(t2t1)) across all summits was calculated (Fig. 3). b, The mean richness change per year accumulates with time to yield absolute changes in species richness (black line in Fig. 2). c, d, Variability in the absolute change in species richness was visualized by randomly sampling ΔSR from all mountains available each year, but adding the s.d. within a region and year. The displayed range in Fig. 2 illustrates the 5th and 95th percentiles of the resulting richness change values from 1,000 runs (orange shading in Fig. 2). This approach reveals changes in variability among mountains over time while also showing overall variability for time steps where only a few summits were sampled (particularly in early time periods).

Extended Data Fig. 2 Relationship between rates of changes in species richness across Europe and rates of increase in temperature (left column), rates of change in precipitation (middle column) and accumulated nitrogen deposition (right column).

Trend lines are interpolated from a simple linear model and are in many cases not significant. Species richness was quantified as the difference between vegetation surveys from the same summit at different times (Extended Data Fig. 1). No nitrogen data were available for Svalbard. The number of observations (comparison of survey and resurveys) are: Svalbard, 7; Northern Scandes, 54; Southern Scandes, 27; Scotland, 7; NW Carpathians, 16; Eastern Alps, 122; Western Alps, 48; SE Carpathians, 9; Pyrenees, 12 (see Fig. 1 for more details).

Extended Data Fig. 3 Historical and recent species richness versus sampling area.

Historical species richness was exceeded within a small sampling area during recent resurveys. Species richness of the historical survey (yellow) contrasted with a species richness accumulation curve of the recent surveys on summits where the highest occurrence of each recent species was estimated to the nearest 1-m elevation. The number of species found historically within the uppermost 10 m of a summit was exceeded within the uppermost 5 m in the most recent resurveys. This analysis includes all 157 European summits for which such data are available, regardless of whether the historical species number was reached in recent times. The blue circle visualizes average species richness of the recent surveys within the uppermost 10 m.

Extended Data Table 1 Increase in species richness with time
Extended Data Table 2 Acceleration of the increase in species richness over time
Extended Data Table 3 Explanatory variables for velocity in species richness changes
Extended Data Table 4 Explanatory variables for species richness changes
Extended Data Table 5 Model evaluation for different explanatory variables and time periods
Extended Data Table 6 Model evaluation for different time lags
Extended Data Table 7 Trait differences between colonizing and old-established species

Supplementary information

Supplementary Information

This file contains Data Table 1: Mountain summits used in the analyses with resurvey dates and references. Details on the references are listed in the table.

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Steinbauer, M.J., Grytnes, JA., Jurasinski, G. et al. Accelerated increase in plant species richness on mountain summits is linked to warming. Nature 556, 231–234 (2018).

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