Oaks Physiological Ecology. Exploring the Functional Diversity of Genus Quercus L.

  1. Escudero A. 1
  2. Mediavilla S. 1
  3. Olmo M 2
  4. Villar R. 2
  5. Merino J. 3
  1. 1 Departamento de Ecología, Facultad de BiologíaUniversidad de SalamancaSalamancaSpain
  2. 2 Área de Ecología, Facultad de CienciasUniversidad de CórdobaCórdobaSpain
  3. 3 Departamento Sistemas Físicos, Químicos y NaturalesUniversidad de Pablo OlavideSevillaSpain
Libro:
Oaks Physiological Ecology. Exploring the Functional Diversity of Genus Quercus L.

ISSN: 1568-2544

ISBN: 9783319690988

Año de publicación: 2017

Volumen: 7

Número: 6

Páginas: 195-237

Tipo: Capítulo de Libro

DOI: HTTPS://DOI.ORG/10.1007/978-3-319-69099-5_6 GOOGLE SCHOLAR

Resumen

The geographic distribution of deciduous versus evergreen woody species has been intensively investigated, but the ecological significance of both leaf habits is still far from being fully understood. The purpose of this chapter is to review the factors that are related with the carbon gain of deciduous and evergreen oak species under Mediterranean environmental conditions. We will focus on the morphological, anatomical and chemical adaptations of evergreens necessary to guarantee leaf survival during the unfavorable part of the year. We will review the information available about the construction and maintenance costs associated with the leaf traits of deciduous and evergreen oak species. Moreover, we will compare these traits with those of non-Mediterranean oaks and species belonging to other families. One central leaf trait is the leaf mass per area (LMA), which depends on the leaf anatomy and chemical composition. Differences in LMA are related to photosynthesis and the costs of construction and maintenance. We will assess the differences in these traits between deciduous and evergreen oaks, the aim being to understand the coexistence of both leaf habits in certain environments.

Referencias bibliográficas

  • 1. Acherar M, Rambal S (1992) Comparative water relations of four Mediterranean oak species. Vegetatio 99–100:177–184CrossRef
  • 2. Acherar M, Rambal S, Lepart J (1991) Évolution du potentiel hydrique foliaire et de la conductance stomatique de quatre chênes méditerranéens lors d’une période de dessèchement. Ann des Sci For 48:561–573
  • 3. Ackerly DD, Bazzaz IFA (1995) Leaf dynamics, self-shading and carbon gain in seedlings of a tropical pioneer tree. Oecologia 101:289–298
  • 4. Allard V, Ourcival J, Rambal S, Joffre R, Rocheteau A (2008) Seasonal and annual variation of carbon exchange in an evergreen Mediterranean forest in southern France. Global Change Biol 14:714–725
  • 5. Asensio D, Peñuelas J, Ogaya R, Llusià J (2007) Seasonal soil and leaf CO2 exchange rates in a Mediterranean holm oak forest and their responses to drought conditions. Atmos Environ 41:2447–2455
  • 6. Baldocchi DD, Ma S, Rambal S, Misson L, Ourcival JM, Limousin JM, Pereira J, Papale D (2010) On the differential advantages of evergreenness and deciduousness in mediterranean oak woodlands: a flux perspective. Ecol Appl 20:1583–1597
  • 7. Bartlett MK, Scoffoni C, Sack L (2012) The determinants of leaf turgor loss point and prediction of drought tolerance of species and biomes: a global meta-analysis. Ecol Lett 15:393–405
  • 8. Bassow S, Bazzaz FA (1998) How environmental conditions affect canopy leaf-level photosynthesis in four deciduous tree species. Ecology 79:2660–2675
  • 9. Blumler MA (2015) Deciduous woodlands in Mediterranean California. In: Box EO, Fujiwara K (eds) Warm-temperate deciduous forests around the Northern Hemisphere. Springer, Switzerland, pp 257–266
  • 10. Bongers FJ, Olmo M, Lopez-Iglesias B, Anten N, Villar R (2017) Drought responses, phenotypic plasticity and survival of Mediterranean species in two different microclimatic sites. Plant Biol 19:386–395
  • 11. Canadell J, Jackson R, Ehleringer J, Mooney HA, Sala OE, Schulze E-D (1996) Maximum rooting depth of vegetation types at the global scale. Oecologia 108:583–595
  • 12. Castro-Díez P, Villar-Salvador P, Perez-Rontome P, Maestro-Martinez M, Montserrat-Marti G (1997) Leaf morphology and leaf chemical composition in three Quercus (Fagaceae) species along a rainfall gradient in NE Spain. Trees 11:127–134
  • 13. Cavender-Bares J, Kitajima K, Bazzaz FA (2004) Multiple trait associations in relation to habitat differentiation among 17 floridian oak species. Ecol Monogr 74:635–662
  • 14. Challabathula D, Vishwakarma A, Raghavendra AS, Padmasree K (2016) Alternative oxidase pathway optimizes photosynthesis during osmotic and temperature stress by regulating cellular ROS, malate valve and antioxidative systems. Front Plant Sci 7: 68. Published online 2016 Feb 9. doi: 10.3389/fpls.2016.00068
  • 15. Chapin FS III (1989) The cost of tundra plant structures: evaluation of concepts and currencies. Amer Nat 133:1–19 16. Cornelissen J, Cerabolini B, Castro-Díez P, Villar-Salvador P, Montserrat-Martí G, Puyravaud JP, Maestro M, Werger MJA, Aerts R (2003) Functional traits of woody plants: correspondence of species rankings between field adults and laboratory-grown seedlings? J Veg Sci 14:311–322
  • 17. Daas C, Montpied P, Hanchi B, Dreyer E (2008) Responses of photosynthesis to high temperatures in oak saplings assessed by chlorophyll-a fluorescence: inter-specific diversity and temperature-induced plasticity. Ann For Sci 65:305
  • 18. David TS, Ferreira MI, Cohen S, Pereira JS, David JS (2004) Constraints on transpiration from an evergreen oak tree in southern Portugal. Agric For Meteorol 122:193–205
  • 19. de la Riva EG, Olmo M, Poorter H, Ubera JL, Villar R (2016) Leaf Mass per Area (LMA) and its relationship with leaf structure and anatomy in 34 Mediterranean woody species along a water availability gradient. PLoS ONE. doi: 10.1371/journal.pone.0148788
  • 20. Díaz S, Kattge J, Cornelissen JHC, Wright IJ, Lavorel S, Dray S, Reu B, Kleyer M, Wirth C, Colin Prentice I, Garnier E, Bönisch G, Westoby M, Poorter H, Reich PB, Moles AT, Dickie J, Gillison AN, Zanne AE, Chave J, Joseph Wright S, Sheremet’ev SN, Jactel H, Baraloto C, Cerabolini B, Pierce S, Shipley B, Kirkup D, Casanoves F, Joswig JS, Günther A, Falczuk V, Rüger N, Mahecha MD, Gorné LD (2016) The global spectrum of plant form and function. Nature 529:167–171
  • 21. Duhme F, Hinckley TM (1992) Daily and seasonal variation in water relations of macchia shrubs and trees in France (Montpellier) and Turkey (Antalya). Vegetatio 99–100:185–198
  • 22. Eamus D, Prichard H (1998) A cost-benefit analysis of leaves of four Australian savanna species. Tree Physiol 18:537–545
  • 23. Edwards EJ, Chatelet DS, Sack L, Donoghue MJ (2014) Leaf life span and the leaf economic spectrum in the context of whole plant architecture. J Ecol 102:328–336
  • 24. Flexas J, Bota J, Cifre J, Escalona JM, Galmés J, Gulías J, Lefi EK, Martínez-Cañellas SF, Moreno MT, Ribas-Carbó M, Riera D, Sampol B, Medrano H (2004) Understanding down-regulation of photosynthesis under water stress: future prospects and searching for physiological tools for irrigation management. Ann Appl Biol 144:273–283
  • 25. Flexas J, Barbour MM, Brendel O, Cabrera HM, Carriquí M, Díaz-Espejo A, Douthe C, Dreyer E, Ferrio JP, Gago J, Gallé A, Galmés J, Kodama N, Medrano H, Niinemets Ü, Peguero-Pina JJ, Pou A, Ribas-Carbó M, Tomás M, Tosens T, Warren CR (2012) Mesophyll diffusion conductance to CO2: an unappreciated central player in photosynthesis. Plant Sci 193–194:70–84
  • 26. Flexas J, Diaz-Espejo A, Gago J, Gallé A, Galmés J, Gulías J, Medrano H (2014) Photosynthetic limitations in Mediterranean plants: a review. Environ Exp Bot 103:12–23
  • 27. Florez-Sarasa ID, Bouma TJ, Medrano H et al (2007) Contribution of the cytochrome and alternative pathways to growth respiration and maintenance respiration in Arabidopsis thaliana. Physiol Plant 129:143–151
  • 28. Gallardo A, Merino J (1993) Leaf decomposition in two Mediterranean ecosystems of southwest Spain: influence of substrate quality. Ecology 74:152–161
  • 29. Galmés J, Medrano H, Flexas J (2007) Photosynthetic limitations in response to water stress and recovery in Mediterranean plants with different growth forms. New Phytol 175:81–93
  • 30. Garbulsky MF, Peñuelas J, Papale D, Filella I (2008) Remote estimation of carbon dioxide uptake by a Mediterranean forest. Global Change Biol 14:2860–2867
  • 31. García Nogales A, Linares JC, Laureano RG, Seco JI, Merino J (2016) Range-wide variation in life-history phenotypes: spatio temporal plasticity across the latitudinal gradient of the evergreen oak Quercus ilex. J Biogeogr 43:2366–2379
  • 32. Givnish TJ (2002) Adaptive significance of evergreen vs. deciduous leaves: solving the triple paradox. Silva Fenn 36:703–743
  • 33. González-Zurdo P, Escudero A, Babiano J, García-Ciudad A, Mediavilla S (2016a) Costs of leaf reinforcement in response to winter cold in evergreen species. Tree Physiol 36:273–286
  • 34. González-Zurdo P, Escudero A, Nuñez R, Mediavilla S (2016b) Losses of leaf area owing to herbivory and early senescence in three tree species along a winter temperature gradient. Int J Biometeorol 60:1661–1674
  • 35. Gulías J, Cifre J, Jonasson S, Medrano H, Flexas J (2009) Seasonal and inter-annual variations of gas exchange in thirteen woody species along a climatic gradient in the Mediterranean island of Mallorca. Flora 204:169–181
  • 36. Günthardt-Goerg MS, Kuster TM, Arend M, Vollenweider P (2013) Foliage response of young central European oaks to air warming, drought and soil type. Plant Biol 15:185–197
  • 37. Hallik L, Niinemets Ü, Wright IJ (2009) Are species shade and drought tolerance reflected in leaf-level structural and functional differentiation in Northern Hemisphere temperate woody flora? New Phytol 184:257–274
  • 38. Harley PC, Tenhunen JD, Beyschlag W, Lange OL (1987) Seasonal changes in net photosynthesis rates and photosynthetic capacity in leaves of Cistus salvifolius, a European mediterranean semi-deciduous shrub. Oecologia 74:380–388
  • 39. Hollinger DY (1992) Leaf and simulated whole-canopy photosynthesis in two co-occurring tree species. Ecology 73:1–14
  • 40. Kikuzawa K (1995) Leaf phenology as an optimal strategy for carbon gain in plants. Can J Bot 73:158–163
  • 41. Kikuzawa K, Onoda Y, Wright IJ, Reich PB (2013) Mechanisms underlying global temperature-related patterns in leaf longevity. Glob Ecol Biogeogr 22:982–993
  • 42. Kok B (1948) A critical consideration of the quantum yield of Chlorella-photosynthesis. Enzimologia 13:1–56
  • 43. Kuglitsch FG, Reichstein M, Beer C, Carrara A, Ceulemans R, Granier A, Janssens IA, Koestner B, Lindroth A, Loustau D, Matteucci G, Montagnani L, Moors EJ, Papale D, Pilegaard K, Rambal S, Rebmann C, Schulze ED, Seufert G, Verbeeck H, Vesala T, Aubinet M, Bernhofer C, Foken T, Grünwald T, Heinesch B, Kutsch W, Laurila T, Longdoz B, Miglietta F, Sanz MJ, Valentini R (2008) Characterisation of ecosystem water-use efficiency of european forests from eddy covariance measurements. Biogeosciences Discuss 5:4481–4519
  • 44. Kurz-Besson C, Otieno D, Lobo Do Vale R, Siegwolf R, Schmidt M, Herd A, Nogueira C, David TS, David JS, Tenhunen J, Pereira JS, Chaves M (2006) Hydraulic lift in cork oak trees in a savannah-type Mediterranean ecosystem and its contribution to the local water balance. Plant Soil 282:361–378
  • 45. Laureano RG, García-Nogales A, Seco JI, Linares JC, Martínez F, Merino J (2016) Plant maintenance and environmental stress. Summarising the effects of contrasting elevation, soil, and latitude on Quercus ilex respiration rates. Plant Soil 409:389–403
  • 46. Lloyd J, Syvertsen JP, Kriedemann PE, Farquhar GD (1992) Low conductances for CO2 diffusion from stomata to the sites of carboxylation in leaves of woody species. Plant Cell Environ 15:873–899
  • 47. Lo Gullo MA, Salleo S, Rosso R, Trifilò P (2003) Drought resistance of 2-year-old saplings of Mediterranean forest trees in the field: relations between water relations, hydraulics and productivity. Plant Soil 250:259–272
  • 48. Lucas PW, Turner IM, Dominy TN, Yamasitha N (2000) Mechanical defences to herbivory. Ann Bot 86:913–920
  • 49. Ma S, Baldocchi DD, Xu L, Hehn T (2007) Inter-annual variability in carbon dioxide exchange of an oak/grass savanna and open grassland in California. Agric For Meteorol 147:157–171
  • 50. Ma S, Baldocchi DD, Mambelli S, Dawson TE (2011) Are temporal variations of leaf traits responsible for seasonal and inter-annual variability in ecosystem CO2 exchange? Funct Ecol 25:258–270
  • 51. Martín Vicente A, Infante JM, García Gordo J, Merino J, Fernández Alés R (1998) Producción de bellotas en montes y dehesas del suroeste español. Pastos 28:237–248
  • 52. Martínez F, Merino O, Martín A, García Martín D, Merino J (1998) Belowground structure and production in a Mediterranean scrub community. Plant Soil 201:209–216
  • 53. Martinez F, Lazo YO, Fernández-Galiano RM, Merino J (2002a) Chemical composition and construction cost for roots of Mediterranean trees, shrub species and grasslands communities. Plant Cell Environ 25:601–608
  • 54. Martinez F, Lazo YO, Fernández-Galiano JM, Merino J (2002b) Root respiration and associated costs in deciduous and evergreen species of Quercus. Plant Cell Environ 25:1271–1278
  • 55. Martínez F, Laureano RG, Merino J (2003) Alternative respiration in seven Quercus spp of SW Spain. J Mediterr Ecol 4:9–14
  • 56. Maselli F, Barbati A, Chiesi M, Chirici G, Corona P (2006) Use of remotely sensed and ancillary data for estimating forest gross primary productivity in Italy. Remote Sens Environ 100:563–575
  • 57. Mediavilla S, Escudero A (2003) Photosynthetic capacity, integrated over the lifetime of a leaf, is predicted to be independent of leaf longevity in some tree species. New Phytol 159:203–211
  • 58. Mediavilla S, Garcia-Ciudad A, Garcia-Criado B, Escudero A (2008) Testing the correlations between leaf life span and leaf structural reinforcement in 13 species of European Mediterranean woody plants. Funct Ecol 22:787–793
  • 59. Mediavilla S, Herranz M, González-Zurdo P, Escudero A (2014) Ontogenetic transition in leaf traits: a new cost associated with the increase in leaf longevity. J Plant Ecol 7:567–575
  • 60. Merino J, García Novo F, Sánchez Díaz M (1976) Annual fluctuation of water potential in the xerophytic shrub of the Doñana biological reserve. Oecologia Plant 11:1–11
  • 61. Millenaar FF, Lambers H (2003) The alternative oxidase: in vivo regulation and fuctions. Plant Biol 5:2–15
  • 62. Mitrakos K (1980) A theory for Mediterranean plant life. Acta Oecologica 1:245–252
  • 63. Ne’eman G (1993) Variation in leaf phenology and habit in Quercus ithaburensis, a Mediterranean deciduous tree. J Ecol 81:627–634
  • 64. Ne’eman G, Goubitz S (2000) Phenology of east-Mediterranean vegetation. In: Trabaud L (ed) Life and environment in the Mediterranean. WIT Press, UK, pp 155–202
  • 65. Niinemets Ü (2001) Global-scale climatic controls of leaf dry mass per area, density, and thickness in trees and shrubs. Ecology 82:453–469
  • 66. Niinemets Ü (2016) Does the touch of cold make evergreen leaves tougher? Tree Physiol 36:267–272
  • 67. Niinemets Ü, Díaz-Espejo A, Flexas J, Galmés J, Warren CR (2009) Role of mesophyll diffusion conductance in constraining potential photosynthetic productivity in the field. J Exp Bot 60:2249–2270
  • 68. Ogaya R, Peñuelas J (2007) Leaf mass per area ratio in Quercus ilex leaves under a wide range of climatic conditions. The importance of low temperatures. Acta Oecologica 31:168–173
  • 69. Penning de Vries FWT, Brusting AHM, Van Laar HH (1974) Products, requirements and efficiency of biosynthesis: a quantitative approach. J Theor Biol 45:339–377
  • 70. Pereira JS, Mateus JA, Aires LM, Pita G, Pio C, David JS, Andrade V, Banza J, David TS, Paço TA, Rodrigues A (2007) Net ecosystem carbon exchange in three contrasting Mediterranean ecosystems—the effect of drought. Biogeosciences 4:791–802
  • 71. Poorter HA, Jong R (1999) A comparison of specific leaf area, chemical composition and leaf construction cost of field plants from 15 habitats differing in productivity. New Phytol 143:163–176
  • 72. Poorter H, Niinemets Ü, Poorter L, Wright IJ, Villar R (2009) Causes and consequences of variation in leaf mass per area (LMA): a meta-analysis. New Phytol 82:565–588
  • 73. Prieto I, Armas C, Pugnaire FI (2012) Water release through plant roots: new insights into its consequences at the plant and ecosystem level. New Phytol 193:830–841
  • 74. Prior LD, Eamus D, Bowman DMJS (2003) Leaf attributes in the seasonally dry tropics: a comparison of four habitats in northern Australia. Funct Ecol 17:504–515
  • 75. Quero JL, Villar R, Marañon T, Zamora R (2006) Interactions of drought and shade effects on seedlings of four Quercus species: physiological and structural leaf responses. New Phytol 170:819–834
  • 76. Reich PB (2014) The world-wide “fast-slow” plant economics spectrum: a traits manifesto. J Ecol 102:275–301
  • 77. Reich PB, Walters MB, Ellsworth DS (1992) Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems. Ecol Monogr 62:365–392
  • 78. Reich PB, Walters MB, Ellsworth DS (1997) From tropics to tundra: global convergence in plant functioning. Proc Natl Acad Sci U S A 94:13730–13734
  • 79. Reich PB, Ellsworth DS, Walters MB, Vose JM, Gresham C, Volin JC, Bowman WD (1999) Generality of leaf trait relationships: a test across six biomes. Ecology 80:1955–1969
  • 80. Reich PB, Falster DS, Ellsworth DS, Wright IJ, Westoby M, Oleksyn J, Lee TD (2009) Controls on declining carbon balance with leaf age among 10 woody species in Australian woodland: do leaves have zero daily net carbon balances when they die? New Phytol 183:153–166
  • 81. Ribas-Carbo M, Taylor NL, Giles L, Busquets S, Finnegan PM, Day DA, Lambers H, Medrano H, Berry JA, Flexas J (2005) Effects of water stress on respiration in soybean leaves 1. Plant Physiol 139:466–473
  • 82. Shipley B, Lechowicz MJ, Wright I, Reich PB (2006) Fundamental tradeoffs generating the worldwide leaf economics spectrum. Ecology 87:535–541
  • 83. Takashima T, Hikosaka K, Hirose T (2004) Photosynthesis or persistence: nitrogen allocation in leaves of evergreen and deciduous Quercus species. Plant Cell Environ 27:1047–1054
  • 84. Tausz M, Grulke NE, Wieser G (2007) Defense and avoidance of ozone under global change. Environ Pollut 147:525–531
  • 85. Tretiach M (1993) Photosynthesis and transpiration of evergreen Mediterranean and deciduous trees in an ecotone during a growing season. Acta Oecologica 14:341–360
  • 86. Turner IM (1994) Sclerophylly: primarily protective? Funct Ecol 8:669–675
  • 87. Tyree MT, Cochard H (1996) Summer and winter embolism in oak: impact on water relations. Ann des Sci For 53:173–180
  • 88. Valladares F, Vilagrosa A, Peñuelas J, Ogaya R, Julio J, Corcuera L, Sisó S (2004) Estrés hídrico: ecofisiología y escalas de la sequía. In: Valladares F (ed) Ecología del bosque mediterráneo en un mundo cambiante, Ministerio de Medio Ambiente, EGRAF, S. A., Madrid, pp 163–190
  • 89. Van Ommen Kloeke AEE, Douma JC, Ordoñez JC, Reich PB, Van Bodegom PM (2012) Global quantification of contrasting leaf life span strategies for deciduous and evergreen species in response to environmental conditions. Global Ecol Biogeogr 21:224–235
  • 90. Villar R, Merino J (1995) Dark leaf respiration in light and darkness of an evergreen and a deciduous plant species. Plant Physiol 107:421–427
  • 91. Villar R, Merino J (2001) Comparison of leaf construction costs in woody species with differing leaf life-spans in contrasting ecosystems. New Phytol 151:213–226
  • 92. Villar R, Robleto JR, De Jong Y, Poorter H (2006) Differences in construction costs and chemical composition between deciduous and evergreen woody species are small as compared to differences among families. Plant Cell Environ 29:1629–1643
  • 93. Villar R, Ruíz-Robleto J, Ubera JL, Poorter H (2013) Exploring variation in leaf mass per area (LMA) from leaf to cell: an anatomical analysis of 26 woody species. Am J Bot 100:1969–1980
  • 94. Vitousek PM, Field CB, Matson PA (1990) Variation in foliar δ13C in Hawaiian Metrosideros polymorpha: a case of internal resistance? Oecologia 84:362–370
  • 95. Warren CR, Adams MA (2000) Trade-offs between the persistence of foliage and productivity in two Pinus species. Oecologia 124:487–494
  • 96. Warton D, Wright I, Falster D, Westoby M (2006) Bivariate line-fitting methods for allometry. Biol Rev 81:259–291
  • 97. Watanabe CKA, Yamori W, Takahashi S, Terashima I, Noguchi KXLM (2016) Mitochondrial alternative pathway-associated photoprotection of Photosystem II is related to the photorespiratory pathway. Plant Cell Physiol 57:1426–1431
  • 98. Westoby M, Warton D, Reich PB (2000) The time value of leaf area. Am Nat 155:649–656
  • 99. Williams K, Field CB, Mooney HA (1989) Relationships among leaf construction cost, leaf longevity, and light environment in rain-forest plants of the genus Piper. Am Nat 133:198–211
  • 100. Witkowski ETF, Lamont BB (1991) Leaf specific mass confounds leaf density and thickness. Oecologia 88:486–493
  • 101. Wright IJ, Cannon K (2001) Relationships between leaf lifespan and structural defences in a low-nutrient, sclerophyll flora. Funct Ecol 15:351–359
  • 102. Wright JP, Sutton-Grier A (2012) Does the leaf economic spectrum hold within local species pools across varying environmental conditions? Funct Ecol 26:1390–1398
  • 103. Wright IJ, Westoby M, Reich PB (2002) Convergence towards higher leaf mass per area in dry and nutrient-poor habitats has different consequences for leaf life span. J Ecol 90:534–543
  • 104. Wright IJ, Westoby M, Reich PB, Oleksyn J, Ackerly DD, Baruch Z, Bongers F, Cavender-Bares J, Chapin T, Cornellissen JHC, Diemer M, Flexas J, Gulias J, Garnier E, Navas ML, Roumet C, Groom PK, Lamont BB, Hikosaka K, Lee T, Lee W, Lusk C, Midgley JJ, Niinemets Ü, Osada H, Poorter H, Pool P, Veneklaas EJ, Prior L, Pyankov VI, Thomas SC, Tjoelker MG, Villar R (2004) The worldwide leaf economics spectrum. Nature 428:821–827
  • 105. Wright IJ, Reich PB, Cornelissen JHC, Falster DS, Groom PK, Hikosaka K, Lee W, Lusk CH, Niinemets Ü, Oleksyn J, Osada N, Poorter H, Warton DI, Westoby M (2005) Modulation of leaf economic traits and trait relationships by climate. Glob Ecol Biogeogr 14:411–421