Abstract
Recurrent anthropogenic and natural perturbations, resource limitations and heterogeneous environments contribute to the maintenance of a remarkable biodiversity in Mediterranean plant communities. Yet, the essential mechanisms of community assembly in these systems remain largely unexplored. In the current paper we investigate the coexistence of Aleppo pine (Pinus halepensis Mill.) and Holm oak (Quercus ilex L.), two of the most widely distributed species in the Iberian Peninsula, in relation to gradients in water availability and disturbance. A spatial model of landscape forest dynamics was implemented, calibrated with experimental data, with stands arranged on a heterogeneous lattice and coupled by dispersal. It was found that pine population persistence in the landscape can be explained independently by a competition-colonization tradeoff (in mesic homogeneous environments) and by a tradeoff between shade and drought tolerance (in heterogeneous low disturbed environments). Both mechanisms reinforce mutually to maintain a shifting mosaic of both taxa along disturbance and aridity gradients. This view is consistent with palynological, historical and forest inventory records. In turn, equilibrium theories of vegetation dynamics (phytosociological), that neglect the role of heterogeneity and disturbances may be inadequate for Mediterranean forests, and have been shown to result in mismanagement practices. We claim that biologically informed models of forest dynamics are desperately needed as diagnosis tools for sustainable forest management.
Similar content being viewed by others
References
Acherar M., Lepart J. and Debussche M. 1984. La colonisation des friches par le pin d’Alep (Pinus halepensis Miller) en Languedoc méditerranéen. Acta Oecologica / Oecologia Plantarum 5: 179-189.
Arianoutsou M. and Groves R. H. (eds) 1994. Plant-Animal interactions in Mediterranean-type ecosystems. Kluwer Academic Publishers, Dordrecht.
Blanco Castro E. et al. (eds) 1996. Los bosques ibéricos. Una interpretación geobotánica. Planeta S.A., Barcelona.
Blondel J. and Vigne J. D. 1993. Space, time, and man as determinants of diversity of birds and mammals in the Mediterranean region. In: Ricklefs R. E. and Schluter D. (eds), Species diversity in ecological communities. The University of Chicago Press, Chicago, pp. 135-146.
Blondel J. and Aronson J. 1995. Biodiversity and ecosystem function in the Mediterranean basin: human and non-human determinants. In: Davis G. W. and Richardson, D. M. (eds), Mediterranean-type ecosystems: the function of biodiversity. Springer-Verlag, Berlin, pp. 43-119.
Bossema I. 1979. Jays and oaks: an eco-ethological study of a symbiosis. Behavior 70: 1-117.
Botkin D. D. 1993. Forest Dynamics: An Ecological Model. Oxford University Press, U.K.
Broncano M. J., Riba M. and Retana J. 1998. Seed germination and seedling performance of two Mediterranean tree species, holm oak (Quercus ilex) and Aleppo pine (Pinus halepensis): a multifactor experimental approach. Plant Ecology 1: 17-26.
Carrión J. S., Andrade A., Bennett K. D., Navarro C. and Munuera M. 2001. Crossing forest thresholds: inertia and collapse in a Holocene sequence from south-central Spain. The Holocene 11: 635-653.
Costa M., García M., Morla C. and Sainz H. 1990. La evolución de los bosques de la Península Ibérica: una interpretación basada en datos paleobiogeográficos. Ecología 1: 31-58.
Cowling R. M., Rundel P. W., Lamont B. B., Arroyo M. K. and Arianoutsou M. 1996. Plant diversity in Mediterranean climateregion. Trends in Ecology and Evolution 11: 362-366.
Crawley, M. (ed.) 1997. Dynamics of plant communities. Plant Ecology. Blackwell Scientific, Oxford.
Espelta J. M., Riba M. and Retana J. 1995. Patterns of seedling recruitment in West-Mediterranean Quercus ilex forests influenced by canopy development. Journal of Vegetation Science 6: 465-472.
Espelta, J.M. 1996. La regeneració de boscos d’alzina (Quercus ilex L.) y pi blanc (Pinus halepensis Mill.): estudi experimental de la resposta de les plàntules a la intensitat de llum i a la disponibilitat d’aigua. Ph.D. Thesis. Universidad Autónoma de Barcelona, Barcelona.
Gil L., Gordo J., Alia R., Catalán G. and Pardos J. A. 1990. Pinus pinaster Aiton en el paisaje vegetal de la Península Ibérica. Ecología 1: 469-496.
González Bernáldez F. 1990. Consideraciones ecológico-políticas acerca de la conservación y regeneración de la cubierta vegetal en España. Ecología 1: 439-445.
Gracia C. 1992. Inventari ecològic i forestal de Catalunya. CREAF Publ, Barcelona, Spain.
Hanes T. L. 1971. Succession after fire in the chaparral of southern California. Ecological. Monographs 41: 27-52.
Hastings A. 1980. Disturbance, coexistence, history and competition for space. Theoretical Population Biology 18: 363-373.
Holmgren M., Scheffer M. and Huston M. A. 1997. The interplay of facilitation and competition in plant communities. Ecology 78: 1966-1975.
Horn H. S. and MacArthur R. H. 1972. Competition among fugitive species in a harlequin environment. Ecology 53: 749-752.
Horn H. S. 1974. The ecology of secondary succession. Annual Review of Ecology and Systematics 5: 25-37.
Hubbell S. P. 2001. A unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton, New Jersey.
Hunter M. L. 1990. Wildlife, Forests and Forestry: Principles of managing forests for biological diversity. Prentice Hall, Englewood Cliffs, New Jersey.
Huston M. A. 1994. Biological Diversity. Cambridge University Press, Cambridge.
ICONA. 1995. Segundo Inventario Forestal Nacional. Ministerio de Agricultura, Pesca y Alimentación, Madrid.
Kadmon R. and Harari-Kremer R. 1999. Landscape-scale regeneration dynamics of disturbed Mediterranean maquis. Journal of Vegetation Science 10: 393-402.
Kazanis D. and Arianoutsou M. 1996. Vegetation structure in a post-fire successional gradient of Pinus halepensis in Attica. International Journal of Wildfire 6: 83-91.
Keeley J. 1986. Resilience of Mediterranean shrub communities to fires. In: Dell B., Hopkins A. J. M. and Lamont B. B. (eds), Resilience inMediterranean-type ecosystems. DrW. Junk Publishers, Dordrecht, The Netherlands, pp. 95-112.
Keeley J. E., Fotheringham C. J. and Ne'eman G. 1999. Immaturity risk in a fire dependent pine. Journal of Mediterranean Ecology 1: 41-48.
Levin S. A. 1974. Dispersion and population interactions. American Naturalist 114: 103-114.
Lookingbill T. and Zavala M. A. 2000. Spatial pattern of Quercus ilex and Quercus pubescens recruitment in Pinus halepensis dominated woodlands. Journal of Vegetation Science 11: 607-612.
Malanson G. P. and O'Leary J. F. 1995. The coastal sage scrubchaparral boundary and responses to global climate change. In: Moreno J. M. and Oechel W. C. (eds), Global Change in Mediterranean-Type Ecosystems. Springer-Verlag, Berlin, pp. 203-224.
Marañón T. and Ojeda J. F. 1998. Ecology and history of a wooded landscape in Southern Spain. In: Kirby K. J. and Watkins C. (eds), The ecological history of European forests. CAB International. Wallingford, pp. 107-116.
May R. M. 1973. Stability and Complexity in Model Ecosystems. Princeton University Press. Princeton, New Jersey.
Mooney H. A. 1988. Lessons from Mediterranean-climate regions. In: Wilson, E. O. (ed.), Biodiversity. National Academy Press, Washington D.C, pp. 157-165.
Morla C. 1993. Significación de los pinares en el paisaje vegetal de la Península Ibérica. In: Silva J. and Vega G. (eds.), Actas I Congreso Forestal Español. Ponencias y Comunicaciones. Pontevedra, Spain, pp. 361-370.
Nathan R., Safriel U. N., Noy-Meir I. and Schiller G. 2000. Spatiotemporal variation in seed dispersal and recruitment near and far from Pinus halepensis trees. Ecology 81: 2156-2169.
Nathan R., Safriel U. N. and Noy-Meir I. 2001. Field validation and sensitivity analyses of a mechanistic model for tree dispersal by wind. Ecology 82: 374-388.
Naveh Z. 1971. The conservation of ecological diversity of Mediterranean ecosystems through ecological management. In: Duffey E. and Watt A. S. (eds), The scientific management of animal and plant communities for conservation. Blackwell, London. pp. 605-622.
Naveh Z. 1987. Landscape ecology, management and conservation of European and Levant Mediterranean uplands. In: Tenhunen J. D., Catarino F. M., Lange O. L. and Oechel W. C. (eds), Plant response to stress: functional analysis in Mediterranean ecosystems. Springer-Verlag, Berlin, pp. 641-657.
Naveh Z. and Dan J. 1973. The human degradation ofMediterranean landscapes in Israel. In: Di Castri, F. and Mooney H. A. (eds), Mediterranean type ecosystems: origin and structure. Springer-Verlag, Berlin, pp. 370-390.
Naveh Z. and Whittaker R. H. 1979. Structural and floristic diversity of shrublands and woodlands in northern Israel and other Mediterranean areas. Vegetatio 41: 171-190.
Naveh Z. and Lieberman A.S. 1984. Landscape ecology. Theory and application. Springer-Verlag, Berlin.
Ne'eman G., Lahav H. and Izhaki I. 1992. Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest. Oecologia 91: 365-370.
Ne'eman G. and Izhaki I. 1998. Stability of pre-and post-fire spatial structure of pine trees in Aleppo pine forest. Ecography 21: 535-542.
Ne'eman G., Fotheringham C. J. and Keeley J. E. 1999. Patch to landscape pattern in post-fire recruitment of a serotinous conifer. Plant ecology 145: 235-242.
Ne'eman H. and Trabaud L. (eds) 2000. The Ecology and Management of Pinus halepensis and P. Brutia ecosystems. Backhuys Publishers, Leiden, The Netherlands.
Pacala S. W. 1997. Dynamics of plant communities. In: Crawley M. C. (ed.), Plant Ecology. Blackwell Scientific, Oxford. pp. 532-555.
Pacala S. W. and Tilman D. 1994. Limiting similarity in mechanistic and spatial models of plant competition in heterogeneous environment. American Naturalist 143: 222-257.
Pacala S. W., Canham C. D., Saponara J., Silander J. A., Kobe R. K. and Ribbens E. 1996. Forest models defined by field measurements: estimation, error analysis and dynamics. Ecological Monographs 66: 1-43.
Pausas G. J. 2001. Resprouting vs seeding-a Mediterranean perspective. Oikos 94: 193-194.
Pigott C. D. and Pigott S. 1993. Water as a determinant of the distribution of trees at the boundary of the Mediterranean zone. Journal of Ecology 81: 557-566.
Rees M, Grubb P. J. and Kelly D. 1996. Quantifying the impact of competition and spatial heterogeneity on the structure and dynamics of a four species guild of winter annuals. American Naturalist 147: 1-32.
Retana J., Riba M., Castell C. and Espelta J. M. 1992. Regeneration by sprouting of holm oak (Quercus ilex L.) stands exploited by selection thinning. Vegetatio 99-100: 355-364.
Retana J., Espelta J. M. and Gracia M. 1996. Caracterización de masas mixtas de pino carrasco y encina en el Montseny (Nordeste de la Península Ibérica). Cuadernos de la Sociedad Española de Ciencias Forestales 3: 167-179.
Retana J., Espelta J. M, Gracia M. and Riba M. 1999. Seedling recruitment. In: Rodà F., Retana J., Gracia C. and Bellot J. (eds), Ecology of Mediterranean evergreen oak forests. Springer-Verlag, Berlin, pp. 89-103.
Rey Benayas J. M. 1998. Growth and survival in Quercus ilex L. seedlings after irrigation y artificial shading on Mediterranean set-aside agricultural lands. Annales des Sciences Forestières 55: 801-807.
Rivas-Martínez S. 1987. Mapa de Series de Vegetación de España. Memoria del Mapa de Series de Vegetación de España. ICONA, Madrid.
Rodà F., Retana J., Gracia C. and Bellot J. (eds) 1999. Ecology of Mediterranean evergreen oak forests. Springer-Verlag, Berlin, Germany.
Romane F., Bacillieri R., Bran D. and Bouchet M. A. 1992. Natural degenerate Mediterranean forests: which future? The examples of holm oak (Quercus ilex) and chestnut (Castanea sativa) coppice stands. In: Teller A., Mathy, P. and Jeffers J. N. R. (eds), Responses of forest ecosystems to environmental changes. Elsevier Applied Science, London, pp. 374-380.
Ruiz de la Torre J. 1973. Significación de los pinares xerófilos. Vida Silvestre 6: 108-113.
Ruiz de la Torre J. 1985. Conservation of plants within their native ecosystems. In: Gómez-Campo C. (ed.), Plant Conservation in the Mediterranean. Dr. J. Junk Publishers, The Hague, pp. 197-219.
Shugart H. H. 1984. A Theory of Forest Dynamics: the Ecological Implications of Forest Succession Models. Springer-Verlag, Berlin.
Smith T. M. and Urban D. L. 1988. Scale and resolution of forest structural pattern. Vegetatio 74: 143-150.
Smith T. M. and Huston M. A. 1989. A theory of the spatial and temporal dynamics of plant communities. Vegetatio 83: 49-69.
Thanos C. A., Daskalakou E. N. and Nikolaidou S. 1996. Early post-fire regeneration of a Pinus halepensis forest on Mount Parnis, Greece. Journal of Vegetation Science 7: 273-280.
Thirgood J. V. 1981. Man and the Mediterranean forest. Academic Press, London.
Tilman D. 1982. Resource competition and community structure. Princeton University Press. Princeton, New Jersey.
Tilman D. 1988. Plant strategies and the dynamics and structure of plant communities. Princeton University Press, New Jersey.
Tilman D. 1990. Constraints and tradeoffs: toward a predictive theory of competition and succession. Oikos 58: 3-15.
Tilman D. and Kareiva P. (eds) 1998. Spatial ecology: the role of space in population dynamics and interspecific interactions. Princeton University Press, Princeton, New Jersey.
Trabaud L., Michels C. and Grosman J. 1985. Recovery of burnt Pinus halepensis Mill. forests. II. Pine reconstitution after wild-fire. Forest Ecology and Management 13: 167-179.
Westman W. E. 1990. Managing for biodiversity. Bioscience 4: 26-32.
Whittaker R. H. 1975. Communities and Ecosystems. Mac-Millan. New York.
Yll E. I., Pérez-Obiol R., Pantaleón-Cano J. and Roure J. M. 1995. Dinámica del paisaje vegetal en la vertiente Mediterránea de la Península Ibérica e Islas Baleares desde el tardiglaciar hasta el presente. In: Aleixandre T. and Pérez-González A. (eds), Reconstrucción de paleoambientes y cambios climáticos durante el Cuaternario. Monografías C.S.I.C., Madrid, pp. 319-328.
Zavala M. 1999. A model of stand dynamics for holm oak-Aleppo pine forests. In: Rodà F., Retana J., Gracia C. and Bellot J. (eds), Ecology of Mediterranean evergreen oak forests. Springer-Verlag, Berlin. pp. 105–117.
Zavala M. A. 2000a. The ecology and dynamics of Mediterranean evergreen pine-oak forests. Ph.D.Thesis. Princeton University. Princeton, New Jersey.
Zavala M. A. 2000b. Aleppo pine stand dynamics in relation to water balance and disturbance: a mechanistic model. In: Ne'eman G. and Trabaud L. (eds), Ecology, Biogeography and Management of Pinus halepensis and P. Brutia Ecosystems in the Mediterranean Basin, Backhuys Publishers, Leiden, The Netherlands, pp. 153-165.
Zavala M. A. and Oria J. A. 1995. Preserving biological diversity in managed forests: a meeting point for forestry and ecology. Urban and Landscape Planning 31: 363-378.
Zavala M. A., Espelta J. M. and Retana J. 2000. Constraints and trade-offs inMediterranean plant communities: the case of mixed holm oak-Aleppo pine forests. Botanical Review 66: 119-149.
Zedler P. H. 1981. Vegetation change in chaparral and desert communities in San Diego communities in San Diego county, California. In: West, D. C., Shugart H. H. and Botkin D. B. (eds), Forest succession: concepts and applications. Springer-Verlag, Berlin, pp. 406-430.
Zedler P. H. and Zammitt C. A. 1989. A population-based critique of concepts of change in the chaparral. In: Keeley J. (ed.), The California chaparral: paradigms re-examined. Natural History Museum. Los Angeles County, California, pp. 73-83.
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
About this article
Cite this article
Zavala, M.A., Zea, E. Mechanisms maintaining biodiversity in Mediterranean pine-oak forests: insights from a spatial simulation model. Plant Ecology 171, 197–207 (2004). https://doi.org/10.1023/B:VEGE.0000029387.15947.b7
Issue Date:
DOI: https://doi.org/10.1023/B:VEGE.0000029387.15947.b7