title: An evolutionary perspective on leaf economics: phylogenetics of leaf mass per area in vascular plants.
abstract: In plant leaves, resource use follows a trade-off between rapid resource capture and conservative storage. This "worldwide leaf economics spectrum" consists of a suite of intercorrelated leaf traits, among which leaf mass per area, LMA, is one of the most fundamental as it indicates the cost of leaf construction and light-interception borne by plants. We conducted a broad-scale analysis of the evolutionary history of LMA across a large dataset of 5401 vascular plant species. The phylogenetic signal in LMA displayed low but significant conservatism, that is, leaf economics tended to be more similar among close relatives than expected by chance alone. Models of trait evolution indicated that LMA evolved under weak stabilizing selection. Moreover, results suggest that different optimal phenotypes evolved among large clades within which extremes tended to be selected against. Conservatism in LMA was strongly related to growth form, as were selection intensity and phenotypic evolutionary rates: woody plants showed higher conservatism in relation to stronger stabilizing selection and lower evolutionary rates compared to herbaceous taxa. The evolutionary history of LMA thus paints different evolutionary trajectories of vascular plant species across clades, revealing the coordination of leaf trait evolution with growth forms in response to varying selection regimes.
authors.affiliation: CNRS, Centre d'Écologie Fonctionnelle et Évolutive (CEFE), UMR 5175 1919 route de Mende, 34293, Montpellier Cedex 5, France ; UMR PVMBT, Université de la Réunion, CIRAD 7 chemin de l'IRAT, 94710, Saint-Pierre, France.
authors.forename: Olivier
authors.initials: O
authors.lastname: Flores
authors.affiliation: CNRS, Centre d'Écologie Fonctionnelle et Évolutive (CEFE), UMR 5175 1919 route de Mende, 34293, Montpellier Cedex 5, France.
authors.forename: Eric
authors.initials: E
authors.lastname: Garnier
authors.affiliation: Department of Biological Sciences, Macquarie University New South Wales, 2109, Australia.
authors.forename: Ian J
authors.initials: IJ
authors.lastname: Wright
authors.affiliation: Department of Forest Resources and Institute on the Environment, University of Minnesota St Paul, Minnesota ; Hawkesbury Institute for the Environment, University of Western Sydney Hawkesbury, New South Wales, Australia.
authors.forename: Peter B
authors.initials: PB
authors.lastname: Reich
authors.affiliation: Department of Plant Production, University of Milan via Celoria 2, I-20133, Milan, Italy.
authors.forename: Simon
authors.initials: S
authors.lastname: Pierce
authors.affiliation: Instituto Multidisciplinario de Biología Vegetal (CONICET - UNC) and FCEFyN, Universidad Nacional de Córdoba Casilla de Correo 495, Vélez Sársfield 299, 5000, Córdoba, Argentina.
authors.forename: Sandra
authors.initials: S
authors.lastname: Dìaz
authors.affiliation: James Hutton Institute Craigiebuckler, Aberdeen, AB15 8QH, UK.
authors.forename: Robin J
authors.initials: RJ
authors.lastname: Pakeman
authors.affiliation: Norwegian Institute for Nature Research Tungasletta 2, 7485, Trondheim, Norway.
authors.forename: Graciela M
authors.initials: GM
authors.lastname: Rusch
authors.affiliation: CNRS, Centre d'Écologie Fonctionnelle et Évolutive (CEFE), UMR 5175 1919 route de Mende, 34293, Montpellier Cedex 5, France.
authors.forename: Maud
authors.initials: M
authors.lastname: Bernard-Verdier
authors.affiliation: CNRS, Centre d'Écologie Fonctionnelle et Évolutive (CEFE), UMR 5175 1919 route de Mende, 34293, Montpellier Cedex 5, France.
authors.forename: Baptiste
authors.initials: B
authors.lastname: Testi
authors.affiliation: Community and Conservation Ecology Group PO Box 14, 9750, AA Haren, The Netherlands.
authors.forename: Jan P
authors.initials: JP
authors.lastname: Bakker
authors.affiliation: Community and Conservation Ecology Group PO Box 14, 9750, AA Haren, The Netherlands.
authors.forename: Renée M
authors.initials: RM
authors.lastname: Bekker
authors.affiliation: DBSF, Università degli Studi dell'Insubria Via J.H. Dunant 3, I- 21100, Varese, Italy.
authors.forename: Bruno E L
authors.initials: BE
authors.lastname: Cerabolini
authors.affiliation: Centro Flora Autoctona, c/o Consorzio Parco Monte Barro via Bertarelli 11, I-23851, Galbiate (LC), Italy.
authors.forename: Roberta M
authors.initials: RM
authors.lastname: Ceriani
authors.affiliation: UR B&SEF CIRAD, TA C-105/D, Campus International de Baillarguet 34398, Montpellier Cedex 5, France.
authors.forename: Guillaume
authors.initials: G
authors.lastname: Cornu
authors.affiliation: INRA UMR 1248 AGIR, Equipe ORPHEE BP 52627 - Auzeville, 31326, Castanet-Tolosan, France.
authors.forename: Pablo
authors.initials: P
authors.lastname: Cruz
authors.affiliation: UR B&SEF CIRAD, TA C-105/D, Campus International de Baillarguet 34398, Montpellier Cedex 5, France.
authors.forename: Matthieu
authors.initials: M
authors.lastname: Delcamp
authors.affiliation: Institute of Botany, Academy of Sciences of the Czech Republic Dukelská 135, CZ-37982, Třeboň, Czech Republic.
authors.forename: Jiri
authors.initials: J
authors.lastname: Dolezal
authors.affiliation: Department of Botany, Stockholm University Stockholm, 106 91, Sweden.
authors.forename: Ove
authors.initials: O
authors.lastname: Eriksson
authors.affiliation: UR B&SEF CIRAD, TA C-105/D, Campus International de Baillarguet 34398, Montpellier Cedex 5, France.
authors.forename: Adeline
authors.initials: A
authors.lastname: Fayolle
authors.affiliation: Centre for Functional Ecology, University of Coimbra Coimbra, Portugal.
authors.forename: Helena
authors.initials: H
authors.lastname: Freitas
authors.affiliation: Department of Molecular Biology and Ecology of Plants, Faculty of Life Sciences, Tel Aviv University Tel Aviv, 69978, Israel.
authors.forename: Carly
authors.initials: C
authors.lastname: Golodets
authors.affiliation: UR B&SEF CIRAD, TA C-105/D, Campus International de Baillarguet 34398, Montpellier Cedex 5, France.
authors.forename: Sylvie
authors.initials: S
authors.lastname: Gourlet-Fleury
authors.affiliation: Department of Archaeology, The University Sheffield, S1 4ET, UK.
authors.forename: John G
authors.initials: JG
authors.lastname: Hodgson
authors.affiliation: DBSF, Università degli Studi dell'Insubria Via J.H. Dunant 3, I- 21100, Varese, Italy.
authors.forename: Guido
authors.initials: G
authors.lastname: Brusa
authors.affiliation: Landscape Ecology Group, Carl von Ossietzky University of Oldenburg P.O. Box 2503, 26111, Oldenburg, Germany.
authors.forename: Michael
authors.initials: M
authors.lastname: Kleyer
authors.affiliation: Landscape Ecology Group, Carl von Ossietzky University of Oldenburg P.O. Box 2503, 26111, Oldenburg, Germany ; Landscape Ecology & Consulting Lerchenstrasse 20, 26215, Wiefelstede, Germany.
authors.forename: Dieter
authors.initials: D
authors.lastname: Kunzmann
authors.affiliation: Laboratoire d'Écologie Alpine (CNRS UMR 5553) and Station Alpine Joseph Fourier (UMS-UJF-CNRS 2925), Université Joseph Fourier BP 53, F-38042, Grenoble, Cedex 09, France.
authors.forename: Sandra
authors.initials: S
authors.lastname: Lavorel
authors.affiliation: Laboratory of Rangeland Ecology, Aristotle University of Thessaloniki 54124, Thessaloniki, Greece.
authors.forename: Vasilios P
authors.initials: VP
authors.lastname: Papanastasis
authors.affiliation: Instituto Multidisciplinario de Biología Vegetal (CONICET - UNC) and FCEFyN, Universidad Nacional de Córdoba Casilla de Correo 495, Vélez Sársfield 299, 5000, Córdoba, Argentina.
authors.forename: Natalia
authors.initials: N
authors.lastname: Pérez-Harguindeguy
authors.affiliation: Instituto Multidisciplinario de Biología Vegetal (CONICET - UNC) and FCEFyN, Universidad Nacional de Córdoba Casilla de Correo 495, Vélez Sársfield 299, 5000, Córdoba, Argentina.
authors.forename: Fernanda
authors.initials: F
authors.lastname: Vendramini
authors.affiliation: Department of Biology, University of Wisconsin-Eau Claire Phillips Hall 353, Eau Claire, Wisconsin, 4702-4004.
authors.forename: Evan
authors.initials: E
authors.lastname: Weiher
keywords: Brownian model; Ornstein–Uhlenbeck model; functional trait; phenotypic evolution