, Volume 173, Issue 4, pp 1613–1624 | Cite as

Growth and stable isotope signals associated with drought-related mortality in saplings of two coexisting pine species

  • Asier Herrero
  • Jorge Castro
  • Regino Zamora
  • Antonio Delgado-Huertas
  • José I. Querejeta
Global change ecology - Original research


Drought-induced events of massive tree mortality appear to be increasing worldwide. Species-specific vulnerability to drought mortality may alter patterns of species diversity and affect future forest composition. We have explored the consequences of the extreme drought of 2005, which caused high sapling mortality (approx. 50 %) among 10-year-old saplings of two coexisting pine species in the Mediterranean mountains of Sierra Nevada (Spain): boreo-alpine Pinus sylvestris and Mediterranean P. nigra. Sapling height growth, leaf δ13C and δ18O, and foliar nitrogen concentration in the four most recent leaf cohorts were measured in dead and surviving saplings. The foliar isotopic composition of dead saplings (which reflects time-integrated leaf gas-exchange until mortality) displayed sharp increases in both δ13C and δ18O during the extreme drought of 2005, suggesting an important role of stomatal conductance (gs) reduction and diffusional limitations to photosynthesis in mortality. While P. nigra showed decreased growth in 2005 compared to the previous wetter year, P. sylvestris maintained similar growth levels in both years. Decreased growth, coupled with a sharper increase in foliar δ18O during extreme drought in dead saplings, indicate a more conservative water use strategy for P. nigra. The different physiological behavior of the two pine species in response to drought (further supported by data from surviving saplings) may have influenced 2005 mortality rates, which contributed to 2.4-fold greater survival for P. nigra over the lifespan of the saplings. This species-specific vulnerability to extreme drought could lead to changes in dominance and distribution of pine species in Mediterranean mountain forests.


Drought stress Tree mortality Leaf δ13Leaf δ18Mediterranean mountain 



The authors acknowledge Christian Körner, Dan Yakir and three anonymous reviewers for insightful comments that improved the manuscript. We thank the Consejería de Medioambiente (Andalusian Government) and the direction of Sierra Nevada National Park for facilities and support to carry out the experiment. We also wish to thank Ignacio Villegas, Jose Antonio Hódar, and Ana Mellado for field and lab assistance. David Nesbitt checked the English of this paper. This study was supported by the coordinated Ministerio de Educación y Ciencia (Spanish Government) projects CGL2008-04794, CSD2008-00040, CGL2008-01671 and by grant FPU-MEC (AP2005-1561) to A.H.

Supplementary material

442_2013_2707_MOESM1_ESM.doc (153 kb)
Supplementary material 1 (DOC 153 kb)


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Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  • Asier Herrero
    • 1
  • Jorge Castro
    • 1
  • Regino Zamora
    • 1
  • Antonio Delgado-Huertas
    • 2
  • José I. Querejeta
    • 3
  1. 1.Grupo de Ecología Terrestre, Departamento de Ecología, Facultad de CienciasUniversidad de GranadaGranadaSpain
  2. 2.Grupo de Biogeoquímica de Isótopos EstablesInstituto Andaluz de Ciencias de la Tierra (CSIC-UGR)ArmillaSpain
  3. 3.Centro de Edafología y Biología Aplicada del Segura (CEBAS-CSIC)Campus Universitario de EspinardoMurciaSpain

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