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Constraints and trade-offs in Mediterranean plant communities: The case of holm oak-Aleppo pine forests

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Abstract

In this paper we review those aspects that are relevant to the development of a mechanistic ecological theory to account for the structure and dynamics of Mediterranean forests, focusing our attention on mixed forests of holm oak (Quercus ilex L.), a shade-tolerant, slowgrowing species that resprouts vigorously after disturbance, and Aleppo pine (Pinus halepensis M.), a fast-growing, nonresprouting, shade-intolerant species. The main objectives of this report are: to introduce some of the primary features of these forests, showing their structural complexity and historical peculiarities; to show that much of this complexity can be conceptually reduced to two main factors of variation, soil-moisture gradients and a complex interaction of historical management and disturbance regimes; and to contrast the unique features of Mediterranean systems with other communities that have inspired generalization in ecology.

Plants in Mediterranean-climate regions must face several environmental constraints during their life cycle: water limitation, competition for light, and a complex set of disturbance regimes, mainly fire, herbivory, and human exploitation. The response of co-occurring species to a given set of environmental constraints depends on a combination of physiological and morphological traits. In holm oak-Aleppo pine forests, the lower limit of distribution along a soil-moisture gradient appears to be controlled by dry-season water stress on seedling performance, and the upper limit seems to be controlled by shade tolerance relative to competitors. The processes that generate and maintain these patterns are related to the responses of the two species to the water and light environments that result from interacting gradients of disturbance and resource availability.

The dynamics of mixed holm oak-Aleppo pine forests may be represented along two major environmental axes: water availability and light intensity; namely, time since last disturbance. At the regional scale, the presence of holm oak and Aleppo pine is expected to be driven mainly by the precipitation regime, with the proportion of Aleppo pine increasing toward the driest border and with holm oak being the dominant species in areas with higher precipitation. Changes of dominance of holm oak and Aleppo pine also respond to water availability at the local scale. In this case, variations between species depend on different factors in a complex way, because reduced soil-moisture levels may result either from low precipitation or from topography and edaphic features. The dynamics of holm oak-Aleppo pine forests are also determined by temporal changes in canopy closure; that is, forest recovery after disturbance. In this case, the proportion of Aleppo pine would increase in recently disturbed stands (i.e., with high light intensity reaching the forest floor), whereas regeneration of holm oak would be dominant under partially closed canopies.

Theories of forest dynamics developed in humid regions may apply only poorly to Mediterranean plant communities, where vegetation change is qualitatively or quantitatively different. Thus, succession in temperate forests appears to be driven by differences in light availability and shade tolerance; but in Mediterranean plant communities, water limitation is of greater importance for the distribution of forest species. In Mediterranean landscapes the interaction of life-history strategies with changing environments is difficult to infer from observational and experimental studies. A mechanistic approach, in which competition or plant performance is measured as a function of resource availability, seems more feasible. The idea should be to develop multispecies models calibrated specifically for Mediterranean forests in a combined program of modeling, field research, and experimentation.

Resumen

En este trabajo se revisan los aspectos más relevantes en el desarrollo de una teoría ecológica mecanicista sobre la estructura y dinámica de los bosques mediterráneos, en concreto sobre los bosques mixtos de encina (Quercus ilex L.), una especie tolerante a la sombra, de crecimiento lento, y que rebrota vigorosamente después de una perturbación, y pino carrasco (Pinus halepensis M.), una especie de crecimiento rápido, no rebrotadora, e intolerante a la sombra. Los principales objetivos de este estudio son: introducir algunas de las principales características de estos bosques, mostrando su complejidad estructural y sus peculiaridades históricas; mostrar que gran parte de esta complejidad puede reducirse conceptualmente a dos principales factores de variación, la disponibilidad de agua en el suelo y una compleja interaction de regímenes de gestión y perturbación; y contrastar las características únicas de los sistemas mediterráneos con las de otros sistemas en los que se han basados gran parte de las generalizaciones que se han hecho en ecologia.

En las regiones de clima mediterráneo, las plantas deben enfrentarse a diferentes restricciones: limitación hídrica, competencia por la luz, y un conjunto complejo de regímenes de perturbación, principalmente fuego, herbivoría, y explotación humana. La respuesta de las especies a un conjunto de limitaciones ambientales depende de la combinación de caracteres fisiológicos y morfológicos. En los bosques de pino carrasco y encina, el límite inferior de distribución de las especies viene controlado por el estrés hídrico de la estación, mientras que el límite superior viene controlado por la tolerancia a la sombra. Los procesos que generan y mantienen estos patrones están relacionados con las respuestas de ambas especies a lo largo de su ciclo de vida a los ambientes de luz y agua que se generan como resultado de la interacción de los gradientes de perturbación y disponibilidad de recursos.

La dinámica de los bosques mixtos de pino carrasco y encina se puede representar a lo largo de dos principales ejes ambientales: disponibilidad de agua e intensidad de luz; o, lo que es lo mismo, tiempo desde la última perturbación. A una escala regional, la presencia de pino carrasco y encina viene determinada por el régimen de precipitación: el pino carrasco aumenta hacia el extremo más seco, mientras que la encina se hace dominante en áreas con mayor precipitación. A escala local, los cambios en la dominancia de ambas especies también responden a la disponibilidad de agua. En este caso las variaciones entre especies dependen de diferentes factores, ya que los niveles bajos de humedad del suelo pueden ser debidos a la baja precipitación, pero también a la topografía o a las características edáficas. La dinámica de los bosques de pino y encina también depende de los cambios temporales en el cierre de la cubierta, es decir, de la recuperación del bosque después de la perturbación. En este caso, la proporción de pino carrasco se incrementa en los rodales recién perturbados (en los cuales llega mucha luz al suelo del bosque), mientras que la regeneración de encina domina bajo cubiertas parcialmente cerradas.

Las teorías de dinámica forestal desarrolladas en las regiones húmedas se aplican con dificultad en las comunidades de plantas mediterráneas, donde los cambios de la vegetación suelen ser cualitativamente y cuantitativamente distintos. Así, en los bosques templados la sucesión viene determinada por diferencias en la disponibilidad de luz y la tolerancia a la sombra de las diferentes especies, mientras que en las comunidades mediterráneas la limitación hídrica es más importante a la hora de explicar la distribución de las especies forestales. En los ecosistemas mediterráneos, la interacción de las estrategias del ciclo vital con los cambios en el ambiente dificilmente pueden deducirse a partir de estudios observacionales y experimentales. Se requiere una aproximación mecanicista, en la cual la competencia o el éxito de las plantas se pueda medir en función de la disponibilidad de recursos. La idea sería desarrollar modelos multiespecíficos calibrados específicamente para los bosques mediterráneos, en los que se combinara la modelización con los estudios de campo y la experimentación.

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  1. Department of Ecology and Evolutionary Biology, Princeton University, 161 Guyot Hall, 08544, Princeton, NJ, USA

    M. A. Zavala

  2. CREAF, Autonomous University of Barcelona, E-08193, Bellaterra (Barcelona), Spain

    J. M. Espelta

  3. CREAF, Autonomous University of Barcelona, E-08193, Bellaterra (Barcelona), Spain

    Javier Retana

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Zavala, M.A., Espelta, J.M. & Retana, J. Constraints and trade-offs in Mediterranean plant communities: The case of holm oak-Aleppo pine forests. Bot. Rev 66, 119–149 (2000). https://doi.org/10.1007/BF02857785

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