Skip to main content
Log in

Reproductive traits of Pinus halepensis in the light of fire – a critical review

  • Published:
Plant Ecology Aims and scope Submit manuscript

Abstract

Fire is known to be a major factor in shaping plants and vegetation worldwide. Many plant traits have been described as adaptations for surviving fire, or regenerating after it. However, many of the traits are also advantageous for overcoming other disturbances. The fact that fire in the Mediterranean Basin has been almost exclusively of anthropogenic origin, and thus is of short duration in an evolutionary time scale, cast doubt on the possibility that fire can act as a selective force in the Mediterranean Basin. Our aim here is to review the ecological advantages of Pinus halepensis traits and their possibility to be selected by fire. The non-self pruning of cones and branches, and the high resin content increase the probability of canopy fires and consequent death of P. halepensis trees. Post-fire regeneration of P. halepensis depends totally upon its canopy-stored seed bank. The seedlings grow quickly and they first reproduce at an early age. Young reproductive trees function first as females with a high percentage of serotinous cones. Thus, young P. halepensistrees allocate many resources to seed production, reducing their `immaturity risk' in a case of an early successive fire. The proportion of serotinous cones is higher in post-fire naturally regenerating stands than in unburned stands, and seeds from serotinous cones germinate better under simulated post-fire conditions. The extremely high pH of the ash-bed under the burned canopies creates the post-fire regeneration niche of P. halepensis exactly under their parent trees. All these traits are advantageous for post-fire regeneration, but could they also be selected during the time scale of anthropogenic fires in the Mediterranean Basin? Pinus halepensis is a relatively short living tree with almost no recruitment under forest canopy. The longest estimated fire-return interval and generation length are about 125 years. The earliest solid evidence for the first hominid-controlled fire in the Mediterranean basin is 780,000 years ago, and thus the estimated number of post-fire generations is 6240. We suggest that such a number of generations is sufficient for the selection and radiation of fire adaptive traits in P. halepensis.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Institutional subscriptions

Similar content being viewed by others

References

  • Agee J.K. 1998. Fire and pine ecosystems. In: Richardson D.M. (ed.), Ecology and Biogeography of Pinus. Cambridge University Press, Cambridge, pp. 193-218.

    Google Scholar 

  • Arianoutsou M. and Ne'eman G. 2000. Post-fire regeneration of natural Pinus halepensis forests in the east Mediterranean. 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, pp. 269-289.

    Google Scholar 

  • Bar-Yosef O. and Goren-Inbar N. 1993. The Lithic Assemblages of Ubeidiya. Qedem 34, Institute of Archaeology, Hebrew University, Jerusalem.

    Google Scholar 

  • Bond M.J. and van Wilgen B. 1996. Fire and Plants. Chapman and Hall, London.

    Google Scholar 

  • Bond W.J. and Midgley J.J. 1995. Kill thy neighbour: an individualistic argument for the evolution of flammability. Oikos 73: 79-85.

    Google Scholar 

  • Charnov E.L. 1982. The Theory of Sex-Allocation. Princeton University Press, Princeton.

    Google Scholar 

  • Daskalakou E.N. and Thanos C.A. 1996. Aleppo pine (Pinus halepensis) post-fire regeneration: the role of canopy and soil seed banks. International Journal of Wildland Fire 6: 59-66.

    Article  Google Scholar 

  • Dimitrakopoulos A.P. 1994. Methodology for measuring ignition point temperatures of plant species. In: Viegas, D.X. (ed.), Proceedings of 2nd International Conference on Forest Fire Research. Coimbra, Portugal, pp. 435-443.

  • Doussi M.A. and Thanos C.A. 2002. The mechanism of cone opening in Pinus halepensis and P. brutia - morphology and physiology. Abstract in MEDPINE 2 International Conference ‘Conservation, Regeneration and Restoration of Mediterranean Pines and their Ecosystems’. Chania, Greece, p. 20.

  • Enright N.J., Marsula L., Lamont B.B. and Wissel C. 1999a. The ecological significance of canopy seed storage in fire-prone environments: a model for non-resprouting shrubs. Journal of Ecology 86: 946-959.

    Article  Google Scholar 

  • Enright N.J., Marsula L., Lamont B.B. and Wissel C. 1999b. The ecological significance of canopy seed storage in fire-prone environments: a model for resprouting shrubs. Journal of Ecology 86: 960-973.

    Article  Google Scholar 

  • Eshel A., Henig-Sever N. and Ne'eman G. 2000. Spatial variation of seedling distribution in an east Mediterranean pine woodland at the beginning of post-fire succession. Plant Ecology 148: 175-182.

    Article  Google Scholar 

  • FAO 1999. Forest Fire Statistics. Timber Bulletin Vol. LII No. 4. Freeman D.C., Drant McArthur E., Harper K.T. and Blauer A.C. 1981. Influence of environment on the floral sex ratio of monoecious plants. Evolution 35: 194-197.

    Google Scholar 

  • Garrod D.A.E. and Bate D.M.A. 1937. The Stone Age of Mt Carmel: Vol 1. Excavations at the Wadi El-Mughara. Clarendon Press, Oxford.

    Google Scholar 

  • Gauthier S., Bergeron Y. and Simon J-P. 1996. Effects of fire regime on the serotiny level of jack pine. Journal of Ecology 84: 539-548.

    Article  Google Scholar 

  • Goubitz S. 2001. Reproduction Ecology of Pinus halepensis, a Monoecious, Wind-Pollinated and Partially Serotinous Mediterranean Pine Tree. Ph.D. dissertation, Utrecht University, Utrecht.

    Google Scholar 

  • Goubitz S., Werger M. and Ne'eman G. In press. Canopy seed bank structure in relation to fire tree size and density. Plant Ecology.

  • Goubitz S., Werger M. and Ne'eman G. 2003. Germination response to fire-related factors of seeds from non-serotinous and serotinous cones. Plant Ecology 169: 195-204.

    Article  Google Scholar 

  • Habrouk A., Retana J. and Espelta J.M. 1999. The role of heat tolerance and cone protection of seeds in the response of three pine species to wildfires. Plant Ecology 145: 91-99.

    Article  Google Scholar 

  • Henig-Sever N., Eshel A. and Ne'eman G. 1996. pH and osmotic potential of pine ash as post-fire germination inhibitors. Physiologia Plantarum 96: 71-76.

    Article  CAS  Google Scholar 

  • Henig-Sever N., Eshel A. and Ne'eman G. 2000. Regulation of the germination of Aleppo pine (Pinus halepensis) by nitrate, ammonium, and gibelrellin, and its role in post-fire forest regeneration. Physiologia Plantarum 108: 390-397.

    Article  CAS  Google Scholar 

  • Higgins S.I. and Richardson D.M. 1998. Pine invasions in the southern hemisphere: modeling interactions between organisms, environment and disturbance. Plant Ecology 135: 79-93.

    Article  Google Scholar 

  • Izhaki I. and Ne'eman G. 2000. Soil seed banks in east Mediterranean pine forests. 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, pp. 167-182.

    Google Scholar 

  • Keeley J.E. 1993. Smoke-induced flowering in the fire-lily Cyrtanthus ventricosus. South African Journal of Botany 59: 638.

    Google Scholar 

  • Keeley J.E. and Fotheringham C.J. 1997. Gas emissions in smoke-induced seed germination. Science 276: 1248-1251.

    Article  CAS  Google Scholar 

  • Keeley J.E. and Zedler P.H. 1998. Life history evolution in pines. In: Richardson D.M. (ed.), Ecology and Biogeography of Pinus. Cambridge University Press, Cambridge, pp. 219-250.

    Google Scholar 

  • Keidar G. 2001. Causes and Factors in Forest Fires in Israel. Thesis submitted for MA degree, Department of Geography, University of Haifa, Haifa, Israel (In Hebrew with English abstract).

    Google Scholar 

  • Laland K.N., Odling-Smee F.J. and Feldman M.W. 1996. The evolutionary consequences of niche construction; a theoretical investigation using two-locus theory. Journal of Evolutionary Biology 9: 293-316.

    Article  Google Scholar 

  • Lamont B.B., LeMaitre D.C., Cowling R.M. and Enright N.J. 1991. Seed canopy storage in woody plants. Botanical Review 57: 277-317.

    Article  Google Scholar 

  • Le Houérou H.N. 1974. Fire and vegetation in the Mediterranean basin. Annual Tall Timber Fire Ecology Conference 13: 237-277.

    Google Scholar 

  • Leone V., Borghetti M. and Saracino A. 2000. Ecology of post-fire recovery in Pinus halepensis in southern Italy. In: Trabaud L. (ed.), Life and Environment in Mediterranean Ecosystems. WIT Press, Southampton, pp. 129-154.

    Google Scholar 

  • Lloret F. and Mari G. 2001. A comparison of the medieval and the current fire regimes in managed pine forests of Catalunia (NE Spain). Forest Ecology and Management 141: 155-163.

    Article  Google Scholar 

  • Lotan J.E. 1976. Cone serotiny fire relationships in lodgepole pine. Annual Tall Timber Fire Ecology Conference 13: 267-278.

    Google Scholar 

  • Martinez-Sanchez J.J., Marin A. Herranz J.M., Ferrandis P. and de las Heras J. 1995. Effects of high temperatures on germination of Pinus halepensis Mill. and P. pinaster Aiton subsp. pinaster seeds in southeast Spain. Vegetatio 116: 69-72.

    Google Scholar 

  • Mercier N., Valladas H., Valladas G., Reyss J.-L., Jelinek A., Meignen L. and Joron J.-L. 1995. TL dates of burnt flint from Jelink’s excavations at Tabun and their implications. Journal of Archaeological Science 22: 495-509.

    Article  Google Scholar 

  • Mirov N.T. 1967. The Genus Pinus. Ronald Press, New York.

    Google Scholar 

  • Nathan R. and Ne'eman G. 2000. Serotiny, seed dispersal and seed predation in Pinus halepensis. 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, pp. 105-118.

    Google Scholar 

  • Nathan R., Safriel U. N., Noy-Meir I. and Schiller G. 1999. Seed release without fire in Pinus halepensis, a Mediterranean serotinous, wind-dispersed tree. Journal of Ecology 87: 659-669.

    Article  Google Scholar 

  • 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.

    Article  Google Scholar 

  • Naveh Z. 1974. Effects of fire in the Mediterranean region. In: Kozlowski T.T. and Ahlgren C.E. (eds), Fire and Ecosystems. Academic Press, New York, pp. 401-434.

    Google Scholar 

  • Naveh Z. 1990. Fire in the Mediterranean - a landscape ecological perspective. In: Goldammer J.G. and Jenkins M.J. (eds), Fire ecosystem dynamics. SPB Academic Publishing, The Hague, pp. 1-20.

    Google Scholar 

  • 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.

    Google Scholar 

  • Ne'eman G. and Trabaud L. (eds) 2000. Ecology, Biogeography and Management of Pinus halepensis and P. brutia Ecosystems in the Mediterranean Basin. Backhuys Publishers, Leiden.

  • Ne'eman G. 2000. The effect of burned pine trees on post-fire regeneration. 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, pp. 303-320.

  • Odling-Smee F.J. Laland K.N. and Feldman M.W. 1996. Niche construction. American Naturalist 147: 641-648.

    Article  Google Scholar 

  • Perry D.A. and Lotan J.E. 1979. A model of fire selection for serotiny in lodgepole pine. Evolution 33: 958-968.

    Article  Google Scholar 

  • Phillips M.A. and Croteau R.B. 1999. Resin based defenses in conifers. Trends in Plant Science 4: 184-190.

    Article  PubMed  Google Scholar 

  • Prodon R., Fons R. and Athias-Binche F. 1987. The impact of fire on animal communities in Mediterranean area. In: Trabaud L. (ed.), The Role of Fire in Ecological Systems. SPB Academic Publishers, The Hague, pp. 121-157.

    Google Scholar 

  • Richardson D. 2000. Mediterranean pines as invaders in the Southern Hemisphere. 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, pp. 131-142.

    Google Scholar 

  • Roitemberg D. and Ne'eman G. 1999. Post-fire seed dispersal in Pinus halepensis. Abstract in MEDPINE, International Workshop on Mediterranean Pines. February 1999, Beit Oren, Israel, p. 46.

  • Saracino A., Pacella R., Leone V. and Borghetti M. 1997. Seed dispersal and changing seed characteristics in Pinus halepensis Mill. forest after fire. Plant Ecology 130: 13-19.

    Article  Google Scholar 

  • Schwilk D.W. and Ackerly D.D. 2001. Flammability and serotiny as strategies: correlated evolution in pines. Oioks 94: 326-336.

    Article  Google Scholar 

  • Schwilk D.W. 2002. Plant Evolution in Fire-Prone Environments. Ph. D. dissertation Stanford University, Stanford.

    Google Scholar 

  • Shmida A., Lev-Yadun S., Goubitz S. and Ne'eman G. 2000. Sexual allocation and gender segregation in Pinus halepensis, P. brutia and P. pinea. 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, pp. 91-104.

    Google Scholar 

  • Tapias R., Gil L., Fuentes-Utrilla P. and Pardos J.A. 2001. Canopy seed banks in Mediterranean pines of southeastern Spain: a comparison between Pinus halepensis Mill., P. pinaster Ait., P. nigra Arn. and P. pinea L. Journal of Ecology 89: 629-638.

    Article  Google Scholar 

  • Teich A.H. 1970. Cone serotiny and inbreeding in natural populations of Pinus banksiana and Pinus contorta. Canadian Journal of Botany 48: 1805-1809.

    Article  Google Scholar 

  • Thanos C.A. 2000. Ecophysiology of seed germination in Pinus halepensis and P. brutia. 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, pp. 37-50.

    Google Scholar 

  • Thanos C.A. and Daskalakou E.N. 2000. Reproduction in Pinus halepensis and P. brutia. 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, pp. 79-90.

    Google Scholar 

  • Thanos C.A., Georghiou K., Kadis C. and Pantazi C. 1992. Cistaceae: a family with hard seeds. Israel Journal of Botany 41: 251-263.

    Google Scholar 

  • Trabaud L. 1987. Fire and survival traits in plants. In: Trabaud L. (ed.), The Role of Fire in Ecological Systems. SPB Academic Publishers The Hague, pp. 65-90.

    Google Scholar 

  • Trabaud L. 2000. Post-fire regeneration of Pinus halepensis forests in the west Mediterranean. 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, pp. 257-268.

    Google Scholar 

  • Trabaud L., Michels C. and Grosman J. 1985. Recovery of burnt Pinus halepensis Mill. Forests. II. Pine reconstitution after fire. Forest Ecology and Management 13: 167-179.

    Article  Google Scholar 

  • Vázquez A. and Moreno J.M. 1998. Patterns of lightning-and people-caused fires in peninsular Spain. International Journal of Wildland Fire 8: 103-115.

    Article  Google Scholar 

  • Whelan R.J. 1995. The Ecology of Fire. Cambridge University Press, Cambridge.

    Google Scholar 

  • Willson M.F. 1983. Plant Reproductive Ecology. John Wiley and Sons, New York.

    Google Scholar 

Download references

Author information

Authors and Affiliations

Authors

Corresponding author

Correspondence to Gidi Ne'eman.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Ne'eman, G., Goubitz, S. & Nathan, R. Reproductive traits of Pinus halepensis in the light of fire – a critical review. Plant Ecology 171, 69–79 (2004). https://doi.org/10.1023/B:VEGE.0000029380.04821.99

Download citation

  • Issue Date:

  • DOI: https://doi.org/10.1023/B:VEGE.0000029380.04821.99

Navigation