The functional trait spectrum of European temperate grasslands

This is the accepted, but not typeset version of the paper: 1 Ladouceur E., Bonomi C., Bruelheide H., Klimešová J., Burrascano S., Poschlod P., Tudela‐ 2 Isanta M., Iannetta P., Mondoni A., Amiaud B., Cerabolini B.E.L., Cornelissen J.H., Craine J., 3 Louault F., Minden V., Öllerer K., Onipchenko V., Soudzilovskaia N.A., Jiménez‐Alfaro B. 4 2019. The functional trait spectrum of European temperate grasslands. 5 Journal of Vegetation Science 30: 777‐788. https://doi.org/10.1111/jvs.12784 6 7 Title 8 The functional trait spectrum of European temperate grasslands 9 10 Running head 11 Functional traits of European grasslands 12 13 Authors and affiliations 14 Emma Ladouceur, Costantino Bonomi, Helge Bruelheide, Jitka Klimešová, Sabina 15

Burrascano, Peter Poschlod _{, Maria Tudela‐Isanta, Pietro Iannetta , Andrea Mondoni,} 16 Bernard Amiaud, Bruno E. L. Cerabolini, J. Hans C. Cornelissen, Joseph Craine, 17 Frédérique Louault, Vanessa Minden, Kinga Öllerer, Vladimir Onipchenko, Nadejda A. 18 Soudzilovskaia & Borja Jiménez‐Alfaro 19

Ladouceur, E.1,2,3_{*, Bonomi, C. (costantino.bonomi@muse.it)}2_{, Bruelheide, H.} 20

(helge.bruelheide@botanik.uni‐halle.de)4,1_{, Klimešová, J. (jitka.klimesova@ibot.cas.cz)}5_, 21

Burrascano, S. (sabina.burrascano@uniroma1.it)6_{, Poschlod, P.} 22

(peter.poschlod@ur.de)7_{, Tudela‐Isanta, M. (mariatudelaa@gmail.com)}3_{, Iannetta, P.} 23

(pete.iannetta@hutton.ac.uk)8_{, Mondoni, A. (andrea.mondoni@unipv.it)}3_{, Amiaud, B.} 24

(bernard.amiaud@univ‐lorraine.fr)9_{, Cerabolini, B.E.L.} 25

(bruno.cerabolini@uninsubria.it )10_{, Cornelissen, J.H. (j.h.c.cornelissen@vu.nl )}11_{, Craine,} 26

J. (josephmcraine@gmail.com)12_{, Louault, F. (frederique.louault@inra.fr)}13_{, Minden, V.} 27

(vanessa.minden@uni‐oldenburg.de)14,15_{, Öllerer, K. (kinga.ollerer@gmail.com)}16,17_, 28

Onipchenko, V. (vonipchenko@mail.ru)18_{, Soudzilovskaia, N.A.} 29

(n.a.soudzilovskaia@cml.leidenuniv.nl)19_{, Jiménez‐Alfaro, B.} 30

1_{German Centre for Integrative Biodiversity Research (iDiv) Halle‐Jena‐Leipzig,} 32 Deutscher Platz 5e, 04103 Leipzig, Germany. 33 2_{Botany Department, Museo delle Scienze (MUSE), Corso Del Lavoro 3, 38122 Trento,} 34 Italy 35 3_{Department of Earth and Environmental Sciences, University of Pavia, Via S. Epifanio 14,} 36 27100 Pavia, Italy 37 4_{Geobotany and Botanical Garden, Institute of Biology, Martin Luther University, Halle‐} 38 Wittenberg, Am Kirchtor 1, 06108, Halle, Germany 39 5_{Institute of Botany, Academy of Sciences of the Czech Republic, Dukelská 135, CZ‐379} 40 82 , Třeboň, Czech Republic 41 6_{Department of Environmental Biology, Sapienza University of Rome, P.le Aldo Moro 5,} 42 00185, Rome, Italy 43

7_{Department of Ecology and Conservation Biology, University of Regensburg,} 44

Universitätsstraße 31, 93053 Regensburg, Germany 45

8_{Ecological Sciences, James Hutton Institute, Dundee, DD2 5DA, Scotland, United} 46

Kingdom, 47

9_{Department of Forest Ecology and Ecophysiology, University of Lorraine‐INRA, UMR} 48 1137, Vandoeuvre‐les‐Nancy, 54500, Lorraine, France 49 10_{Department of Theoretical and Applied Sciences, University of Insubria, Via J.H.} 50 Dunant 3, 21100, Varese, Italy 51 11_{Department of Ecological Science, Faculty of Earth and Life Sciences, Vrije Universiteit,} 52 De Boelelaan 1085, 1081 HV, Amsterdam, Netherlands 53 12_{Jonah Ventures, 1600 Range Street Suite 201, Boulder, Colorado, 80301, USA} 54 13_{INRA, VetAgro Sup, UMR Ecosystème Prairial, 63000, Clermont‐Ferrand, France} 55

18_{Faculty of Biology, Moscow State University, 1‐12 Leninskie Gory, 119991, Moscow,} 64 Russia, 65 19_{Department of Ecology and Environmental Sciences, Leiden University, Rapenburg 70,} 66 2311 EZ, Leiden, The Netherlands 67 20_{Research Unit of Biodiversity (CSIC/UO/PA), González Gutiérrez Quirós, Universidad} 68 de Oviedo, 33600 Mieres, Spain 69 70

Correspondence: 1_{Emma Ladouceur, Biodiversity Synthesis, German Centre for} 71

Table 1: Description of the five grassland types investigated. Altitude shows estimated ranges above the sea level. 689

Number of specialist species describes the total number of specialists identified for each grassland type. EUNIS and ANNEX 690

I habitats and their management practices according to www.eunis.eea.europa.eu/habitats. Soil pH and moisture values 691 are derived from the Ellenberg indicator values of each species (Fitter & Peat 1994; Sanda et al. 2003; Ciocârlan 2009; 692 Moretti & Legg 2009; Ellenberg 2010; Hill et al.; Kattge; Öllerer), expressed as the mean of those specialists (Table 2). VPL= 693 vegetation period length (Aeschimann et al. 2004). Data sources listed in Appendix S2. 694 Grassland type Altitude # of specialist species European Nature Information System (EUNIS) habitat classification ANNEX I habitats of EU Habitat Directive _Traditional management practice Soil pH Soil moisture VPL Meadows 0‐ 1000m 57 altitude meadow E2.3 E2.2 Low & Medium Mountain hay meadows Lowland Hay Meadows 6510 + High Altitude Hay Meadows 6520 Mowing, (Grazing) (neutral6.9 ) 4.78 March‐ October (8 months) Dry Grasslands 500‐ 1400m 77 E1.2 Perennial calcareous grassland and basic steppes Semi‐natural Dry Grasslands (Festuco‐ Brometalia) 6210 Mowing & Grazing (calcare7.3 ous) 3.2 March‐ October (8 months) Nardus‐

dominated 1800m 1400‐ 81 Mediterranean dry e1.7 Closed non‐ acid & neutral grassland Mountain acid grassland Species Rich Nardus Grasslands 6230 Grazing 4.39

(acidic) 5.8 September May‐ (5 months) Calcareous

Alpine 2700m 1800‐ 149 E4.4 Calcareous alpine and subalpine grassland Alpine & subalpine calcareous grasslands 6170 None 7.7 (calcare ous) 4.68 June‐ August (3 months) Acidic

Table 2: Loadings of the plant trait values for the first three axes of the PCA for: all traits and all species. Bold numbers 696

indicate the top three (3)‐ five (5) highest loadings on each axis. 697

a) All Traits  i) All species (Main Analysis) 

  Axis 1  Axis 2  Axis 3  Axis 4  Axis 5