, Volume 23, Issue 4, pp 787–799 | Cite as

Summer-drought constrains the phenology and growth of two coexisting Mediterranean oaks with contrasting leaf habit: implications for their persistence and reproduction

  • Gabriel Montserrat-MartíEmail author
  • Jesús Julio Camarero
  • Sara Palacio
  • Carmen Pérez-Rontomé
  • Rubén Milla
  • Jorge Albuixech
  • Melchor Maestro
Original Paper


This study analyses how coexisting evergreen and deciduous oaks adjust their phenology to cope with the stressful Mediterranean summer conditions. We test the hypothesis that the vegetative and reproductive growth of the winter deciduous (Quercus faginea Lam.) is more affected by summer drought than that of the evergreen [Quercus ilex L. subsp. ballota (Desf.) Samp.]. First, we assessed the complete aboveground phenology of both species during two consecutive years. Shoot and litter production and bud, acorn and secondary growth were monitored monthly. Second, we identified several parameters affected by summer conditions: apical bud size, individual leaf area (LA), leaf mass per area (LMA) and acorn yield in both species, and leaf-fall in Q. faginea; and analysed their variation over 10 years. Q. ilex performed up to 25% of shoot growth and most leaf development during summer, whereas Q. faginea completed most of both phenophases during spring. Secondary growth was arrested in summer under drought conditions. Approximately, 30–40% of bud and 40–50% of acorn growth was undertaken during summer in both species. Summer drought related to differences in LA, LMA and leaf senescence, but not to acorn yield. Both species had similar year-to-year patterns of acorn production, though yields were always lower in Q. faginea. Bud size decreased severely in both species during extremely dry years. In Q. ilex, bud size tended to alternate between years of large and small buds, and these patterns were followed by opposite trends in stem length. In Q. faginea, bud size was more stable through time. Q. ilex was more phenologically active during summer than Q. faginea, indicating a higher tolerance to drought. Furthermore, bud and fruit growth (the only two phenophases that both species performed during summer) were more severely affected by summer drought in Q. faginea than in the evergreen. The differential effects of summer drought on key phenophases for the persistence (bud growth) and colonization ability (fruit production) of both species may have consequences for their coexistence.


Acorn development and production Bud development Quercus ilex subsp. ballota Quercus faginea Primary and secondary growth 



We thank Pilar Castro and four anonymous referees for their valuable comments on previous versions of the manuscript, and Helena Lahoz, Patricia Fustero, María Isabel Luengo, Javier Lou, María Lázaro, Pablo Almazán, Inmaculada Benito and Carmen Arroyo for their help in data collection and analysis. Francisco Alberto kindly classified the soil. This study was possible thanks to the collaboration within the GLOBIMED network (Ministerio de Educación y Ciencia, Spain) and it was supported by the MEC-CICyT projects AGF96-0399, CGL2007-66066-C04/BOS and CGL2008-04847-C02-01, DGA projects P-038/96 and GA-LC-011/2008, and INIA projects RTA2005-00100-C02-00 and SUM2006-00025-00-00. JJC acknowledges the support of the “Fundación Aragón I+D”. SP and RM were funded by MEC by a postdoc (SEUI-FECYT) and a Juan de la Cierva contract, respectively. JA was funded by DGA.


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Copyright information

© Springer-Verlag 2009

Authors and Affiliations

  • Gabriel Montserrat-Martí
    • 1
    Email author
  • Jesús Julio Camarero
    • 1
  • Sara Palacio
    • 1
    • 2
  • Carmen Pérez-Rontomé
    • 3
  • Rubén Milla
    • 4
  • Jorge Albuixech
    • 1
  • Melchor Maestro
    • 1
  1. 1.Departamento de Conservación de Ecosistemas NaturalesInstituto Pirenaico de Ecología (CSIC)ZaragozaSpain
  2. 2.Macaulay InstituteAberdeenUK
  3. 3.Departamento de Nutrición VegetalEstación Experimental de Aula Dei (CSIC)ZaragozaSpain
  4. 4.Área de Biodiversidad y ConservaciónUniversidad Rey Juan CarlosMóstoles, MadridSpain

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