Plant Ecology

, Volume 212, Issue 12, pp 1959–1966 | Cite as

Leaf physiological traits in relation to resprouter ability in the Mediterranean Basin

  • E. I. Hernández
  • J. G. Pausas
  • A. Vilagrosa


In Mediterranean ecosystems, fire is a strong selective agent among plants, and the different post-fire regeneration strategies (e.g. resprouting and non-resprouting) have implications for other plant traits. Because young plants of non-resprouters need to grow quickly and mature well before the next fire, we predict that they should possess leaf traits related to increased efficiency in growth and resource acquisition compared with resprouter species. To test this hypothesis, we measured specific leaf area, leaf nitrogen and carbon concentrations and leaf physiological traits, including gas exchange parameters and chlorophyll fluorescence, in 19 Mediterranean species cultivated in a common garden. Both cross-species and phylogenetically informed analyses suggest that non-resprouters have better physiological performance at the leaf level (i.e. higher photosynthetic capacity) than resprouters. All these results suggest that non-resprouter species are able to take greater advantage for vegetative growth and carbon fixation than resprouters during periods when water is readily available. The contrasted physiological differences between resprouters and non-resprouters reinforce the idea that these two syndromes are functionally different (i.e. they are functional types).


Functional traits Leaf nitrogen and carbon Non-resprouters Photosynthetic capacity PSII photochemical efficiency Resprouters Specific leaf area 



This study was financed by the Spanish projects SINREG (REN2003-07198-C02-02/GLO), PERSIST (CGL2006-07126/BOS), ESTRES (063/SGTB/2007/7.1) and FUME (GA-243888). E.I. Hernández thank the University of Alicante for her FPU research fellowship. We are all grateful to M. Llorca for assistance with the experiment, to J. Scheiding for language corrections on the manuscript, and to S. Paula for corrections and suggestions. CEAM is partly supported by Generalitat Valenciana, Fundación Bancaja, and the projects GRACCIE (Consolider-Ingenio 2010) and FEEDBACKS (Prometeo–Generalitat Valenciana). CIDE is supported by CSIC, Generalitat Valenciana and Universitat de Valencia.

Supplementary material

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Supplementary material 1 (DOC 51 kb)


  1. Ackerly DD (2004) Functional strategies of chaparral shrubs in relation to seasonal water deficit and disturbance. Ecol Monogr 74:25–44CrossRefGoogle Scholar
  2. Bond WJ, Midgley JJ (2001) Ecology of sprouting in woody plants: the persistence niche. Trends Ecol Evol 16:45–51PubMedCrossRefGoogle Scholar
  3. Bond WJ, van Wilgen BW (1996) Fire and plants. Chapman and Hall, LondonCrossRefGoogle Scholar
  4. Bowen BJ, Pate JS (1993) The significance of root starch in postfire shoot recovery of the resprouter Stirlingia latifolia R. Br. (Proteaceae). Ann Bot 72:7–16CrossRefGoogle Scholar
  5. Felsenstein J (1985) Phylogenies and the comparative method. Am Nat 125:1–15CrossRefGoogle Scholar
  6. 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 175:81–93PubMedCrossRefGoogle Scholar
  7. Gulías J, Flexas J, Mus M, Cifre J, Lefi E, Medrano H (2003) Relationship between maximum leaf photosynthesis, nitrogen content and specific leaf area in Balearic endemic and non-endemic Mediterranean species. Ann Bot 92:215–222PubMedCrossRefGoogle Scholar
  8. Guzmán J, Vargas P (2009) Long-distance colonization of the Western Mediterranean by Cistus ladanifer (Cistaceae) despite the absence of special dispersal mechanisms. J Biogeogr 36:954–968CrossRefGoogle Scholar
  9. Hernández EI, Vilagrosa A, Luis VC, Llorca M, Chirino E, Vallejo VR (2009) Root hydraulic conductance, gas exchange and leaf water potential in seedlings of Pistacia lentiscus L. and Quercus suber L. grown under different fertilization and light regimes. Environ Exp Bot 67:269–276CrossRefGoogle Scholar
  10. Hernández EI, Vilagrosa A, Pausas JG, Bellot J (2010) Morphological traits and water use strategies in seedlings of Mediterranean coexisting species. Plant Ecol 207:233–244CrossRefGoogle Scholar
  11. Jacobsen AL, Pratt RB, Davis SD, Ewers FW (2008) Comparative community physiology: nonconvergence in water relations among three semi-arid shrub communities. New Phytol 180:100–113PubMedCrossRefGoogle Scholar
  12. Keeley JE (1986) Resilience of Mediterranean shrub communities to fire. In: Dell B, Hopkins AJM, Lamont BB (eds) Resilience in Mediterranean-type ecosystems. Dr. W. Junk, Dordrecht, pp 95–112CrossRefGoogle Scholar
  13. Keeley JE, Pausas JG, Rundel PW, Bond WJ, Bradstock RA (2011) Fire as an evolutionary pressure shaping plant traits. Trends Plant Sci 16:406–411PubMedCrossRefGoogle Scholar
  14. Lamont BB, Wiens D (2003) Are seed set and speciation rates always low among species that resprout after fire, and why? Evol Ecol 17:277–292CrossRefGoogle Scholar
  15. Lamont BB, Groom PK, Cowling RM (2002) High leaf mass per area of related species assemblages may reflect low rainfall and carbon isotope discrimination rather than low phosphorus and nitrogen concentrations. Funct Ecol 16:403–412CrossRefGoogle Scholar
  16. Larcher W (1995) Physiological plant ecology: ecophysiology and stress physiology of functional groups, 3rd edn. Springer, BerlinGoogle Scholar
  17. Levitt J (1980) Response of plants to environmental stresses, vol 1, 2nd edn. Academic Press, New YorkGoogle Scholar
  18. Maxwell K, Johnson N (2000) Chlorophyll fluorescence—a practical guide. J Exp Bot 51:659–668PubMedCrossRefGoogle Scholar
  19. Medrano H, Flexas J, Galmés J (2009) Variability in water use efficiency at the leaf level among Mediterranean plants with different growth forms. Plant Soil 317:17–29CrossRefGoogle Scholar
  20. Meziane D, Shipley B (2001) Direct and indirect relationships between specific leaf area, leaf nitrogen and leaf gas exchange. Effects of irradiance and nutrient supply. Ann Bot 88:915–927CrossRefGoogle Scholar
  21. Montès N, Ballini C, Bonin G, Faures J (2004) A comparative study of aboveground biomass of three Mediterranean species in a post-fire sucession. Acta Oecol 25:1–6CrossRefGoogle Scholar
  22. Moreira B, Torno J, Estrelles E, Pausas JG (2010) Disentangling the role of heat and smoke as germination cues in Mediterranean Basin flora. Ann Bot 105:627–635PubMedCrossRefGoogle Scholar
  23. Naveh Z (1975) The evolutionary significance of fire in the Mediterranean region. Vegetatio 29:199–208CrossRefGoogle Scholar
  24. Paradis E, Claude J (2002) Analysis of comparative data using generalized estimating equations. J Theor Biol 218:175–185PubMedCrossRefGoogle Scholar
  25. Paradis E, Claude J, Strimmer K (2004) APE: analyses of phylogenetics and evolution in R language. Bioinformatics 20:289–290PubMedCrossRefGoogle Scholar
  26. Pate JS, Froend RH, Bowen BJ, Hansen A, Kuo J (1990) Seedling growth and storage characteristics of seeder and resprouter species of Mediterranean-type ecosystems of S.W. Australia. Ann Bot 65:585–601Google Scholar
  27. Paula S, Pausas JG (2006) Leaf traits and resprouting ability in the Mediterranean basin. Funct Ecol 20:941–947CrossRefGoogle Scholar
  28. Paula S, Pausas JG (2008) Burning seeds: germinative response to heat treatments in relation to resprouting ability. J Ecol 96:543–552CrossRefGoogle Scholar
  29. Paula S, Pausas JG (2011) Root traits explain different foraging strategies between resprouting abilities. Oecologia 165:321–331PubMedCrossRefGoogle Scholar
  30. Paula S, Arianoutsou M, Kazanis D, Tavsanoglu Ç, Lloret F, Buhk C, Ojeda F, Luna B, Moreno JM, Rodrigo A, Espelta JM, Palacio S, Fernández-Santos B, Fernandes PM, Pausas JG (2009) Fire-related traits for plant species of the Mediterranean Basin. Ecology 90:1420CrossRefGoogle Scholar
  31. Pausas JG (1999) Mediterranean vegetation dynamics: modelling problems and functional types. Plant Ecol 140:27–39CrossRefGoogle Scholar
  32. Pausas JG (2004) Changes in fire and climate in the eastern Iberian Peninsula (Mediterranean basin). Clim Change 63:337–350CrossRefGoogle Scholar
  33. Pausas JG, Keeley JE (2009) A burning story: the role of fire in the history of life. Bioscience 59:593–601CrossRefGoogle Scholar
  34. Pausas JG, Verdú M (2005) Plant persistence traits in fire-prone ecosystems of the Mediterranean basin: a phylogenetic approach. Oikos 109:196–292CrossRefGoogle Scholar
  35. Pausas JG, Bradstock RA, Keith DA, Keeley JE, The GCTE (Global change of terrestrial ecosystems) fire network (2004) Plant functional traits in relation to fire in crown-fire ecosystems. Ecology 85:1085–1100CrossRefGoogle Scholar
  36. Piñol J, Terradas J, Lloret F (1998) Climate warming, wildfire hazard, and wildfire occurrence in coastal eastern Spain. Clim Change 38:345–357CrossRefGoogle Scholar
  37. Poorter H, Evans JR (1998) Photosynthetic nitrogen-use efficiency of species that differ inherently in specific leaf area. Oecologia 116:26–37CrossRefGoogle Scholar
  38. Pratt RB, Jacobsen AL, Golgotiu KA, Sperry JS, Ewers FW, Davis SD (2007) Life history type and water stress tolerance in nine California chaparral species (Rhamnaceae). Ecol Monogr 77:239–253CrossRefGoogle Scholar
  39. Pratt RB, North GB, Jacobsen AL, Ewers FW, Davis SD (2009) Xylem root and shoot hydraulics is linked to life history type in chaparral seedlings. Funct Ecol 24:70–81CrossRefGoogle Scholar
  40. Pugnaire FI, Chapin FS III, Hardig TM (2006) Evolutionary changes in correlations among functional traits in Ceanothus in response to Mediterranean conditions. Web Ecology 6:17–26Google Scholar
  41. Reich PB, Kloeppel BD, Ellsworth DS, Walters MB (1995) Different photosynthesis-nitrogen relations in deciduous hardwood and evergreen coniferous tree species. Oecologia 104:24–30CrossRefGoogle Scholar
  42. 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 80:1955–1969CrossRefGoogle Scholar
  43. Schutz AEN, Bond WJ, Cramer MD (2009) Juggling carbon: allocation patterns of a dominant tree in a fire-prone savanna. Oecologia 160:235–246PubMedCrossRefGoogle Scholar
  44. Schwilk DW, Ackerly DD (2005) Is there a cost to resprouting? Seedling growth rate and drought tolerance in sprouting and nonsprouting Ceanothus (Rhamnaceae). Am J Bot 92:404–410PubMedCrossRefGoogle Scholar
  45. Valladares F (2008) A mechanistic view of the capacity of forest to cope with climate change. In: Bravo F, May VL, Jandl R, Gadow Kv (eds) Managing forest ecosystems: the challenge of climate change. Springer–Verlag, Berlin, pp 15–40CrossRefGoogle Scholar
  46. Valladares F, Sánchez-Gómez D (2006) Ecophysiological traits associated with drought in Mediterranean tree seedlings: individual responses versus interspecific trends in eleven species. Plant Biol 8:688–697PubMedCrossRefGoogle Scholar
  47. Valladares F, Skillman JB, Pearcy RW (2002) Convergence in light capture efficiencies among tropical forest understory plants with contrasting crown architectures: a case of morphological compensation. Am J Bot 89:1275–1284PubMedCrossRefGoogle Scholar
  48. Verdú M (2000) Ecological and evolutionary differences between Mediterranean seeders and resprouters. J Veg Sci 11:265–268CrossRefGoogle Scholar
  49. Verdú M, Pausas JG (2007) Fire drives phylogenetic clustering in Mediterranean Basin woody plant communities. J Ecol 95:1316–1323CrossRefGoogle Scholar
  50. Vilagrosa A, Vallejo VR, Bellot J, Gil-Pelegrín E (2003) Cavitation, stomatal conductance, and leaf dieback in seedlings of two co-occurring Mediterranean shrubs during an intense drought. J Exp Bot 54:2015–2024PubMedCrossRefGoogle Scholar
  51. Vilagrosa A, Morales F, Abadia A, Bellot J, Cochard H, Gil-Pelegrín E (2010) Are symplast tolerance to intense drought conditions and xylem vulnerability to cavitation coordinated? An integrated analysis of photosynthetic, hydraulic and leaf-level processes in two Mediterranean drought-resistant species. Environ Exp Bot 69:233–242. doi: 10.1016/j.envexpbot.2010.04.013 CrossRefGoogle Scholar
  52. Webb CO, Ackerly DD, Kembel SW (2008) Phylocom: software for the analysis of phylogenetic community structure and trait evolution. Bioinformatics 24:2098–2100PubMedCrossRefGoogle Scholar
  53. Wright IJ, Westoby M (2001) Understanding seedling growth relationships through specific leaf area and leaf nitrogen concentration: generalizations across growth forms and growth irradiance. Oecologia 127:21–29CrossRefGoogle Scholar
  54. Wright IJ, Reich PB, Westoby M (2001) Strategy shifts in leaf physiology, structure and nutrient content between species of high- and low-rainfall and high- and low-nutrient habitats. Funct Ecol 15:423–434CrossRefGoogle Scholar
  55. Wright IJ, Groom PK, Lamont BB, Poot P, Prior LD, Reich PB, Schulze ED, Veneklaas EJ, Westoby M (2004) Leaf trait relationships in Australian plant species. Funct Plant Biol 31:551–558CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2011

Authors and Affiliations

  • E. I. Hernández
    • 1
    • 2
  • J. G. Pausas
    • 3
  • A. Vilagrosa
    • 2
  1. 1.Departamento de EcologíaUniversidad de AlicanteAlicanteSpain
  2. 2.CEAM Centro de Estudios Ambientales del MediterráneoUniversidad de AlicanteAlicanteSpain
  3. 3.CIDE, CSICMontcadaSpain

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