Plant Ecology

, Volume 169, Issue 2, pp 195–204 | Cite as

Germination response to fire-related factors of seeds from non-serotinous and serotinous cones

  • S. GoubitzEmail author
  • M.J.A. Werger
  • G. Ne'eman


Pinus halepensis, a Mediterranean pine tree, is a partially serotinous species: individual trees of this species carry both non-serotinous and serotinous cones. Serotinous cones open mainly after fire, whereas non-serotinous cones open in absence of fire. In this study we addressed the question, whether or not this cone response is linked with the germination response of seeds to fires. Two main factors associated with fire are heating of seeds and soil pH. A combination of high heat and high pH simulates a scenario with fire, whereas low heat and low pH simulates a scenario without fire. We assessed the separate and combined effects of heat and pH on the germination rate and the percentage of germination of seeds from non-serotinous cones and two age classes of serotinous cones of P. halepensis. Heat had no effect on the percentage of germination of seeds from any of the cone types, but did positively affect the germination rates of seeds from both age-classes of serotinous cones. High pH negatively affected the germination rate of seeds from all cone types as well as the percentage of germination of seeds from non-serotinous cones. The combinations of heat and pH had different effects on the three cone types: percentage of germination and rate of germination of seeds from non-serotinous cones was higher in the combination high heat-high pH than in the combination low heat-low pH. In the combination high heat-high pH, seeds from serotinous cones germinated better than seeds from non-serotinous cones. The different germination responses of seeds from non-serotinous and serotinous cones could not be attributed to differences in cone age. Our results indicate that the cone response is linked to the germination response of the seeds in P. halepensis, with seeds from serotinous cones being more tolerant to fire related factors.

Fire Germination Heat pH Pinus halepensis Serotiny 


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  1. Beaufait W.R. 1960. Some effects of high temperatures on the cones and seeds of jack pine. Forest Science 6: 194–199.Google Scholar
  2. Bond W.J. and van Wilgen B.W. 1996. Fire and plants. In: Usher M.B., DeAngelis D.L. and Manly B.F.J. (eds). Chapman and Hall, London.Bradstock R.A., Gill A.M., Hastings S.M. and Moore P.H.R. 1994.Survival of serotinous seedbanks during bushfires: comparative studies of Hakea species from southeastern Australia. Australian Journal of Ecology 19: 276–282.Google Scholar
  3. Broncano M.J., Riba M. and Retana J. 1998. Seed germination and seedling performance of two Mediterranean tree species, Holm oak (Quercus ilex L.) and Aleppo pine (P. halepensis Mill.): a multifactor experimental approach. Plant Ecology 138: 17–26.CrossRefGoogle Scholar
  4. Cowling R.M. and Lamont B.B. 1985. variation in serotiny of three Banksia species along a climatic gradient. Australian Journal of Ecology 10: 345–350.CrossRefGoogle Scholar
  5. Cowling R.M., Lamont B.B. and Pierce S.M. 1987. Seed bank dynamics of four co-occuring Banksia species. Journal of Ecology 75: 289–302.CrossRefGoogle Scholar
  6. Daskalakou E.N. and Thanos C.A. 1996. Aleppo pine (Pinus halepensis) postfire regeneration: the role of canopy and soil seed banks. International Journal of Wildland fire 6: 59–66.CrossRefGoogle Scholar
  7. Despain D.G., Clark D.L. and Reardon J.J. 1996. Simulation of crown fire effects on canopy seed bank in Lodgepole pine. International Journal of Wildland Fire 6: 45–49.CrossRefGoogle Scholar
  8. Enright N.J., Marsula R., Lamont B.B. and Wissel C. 1998. The ecological significance of canopy seed storage in fire-prone environments: a model for non-sprouting shrubs. Journal of Ecology 86: 946–959.CrossRefGoogle Scholar
  9. Escudero A., Sanz M.V., Pita J.M. and Perez-Garcia F. 1999. Probability of germination after heat treatment of native Spanish pines. Ann. For. Sci. 56: 511–520.Google Scholar
  10. Fitter A.H. and Hay R.K.M. 1987. Environmental Physiology of Plants. Academic Press, London.Google Scholar
  11. Fox G.A. 1993. Failure time analysis: emergence, flowering, survivorship, and other waiting times. In: Scheiner S.M. and Gurevitch J. (eds), Design and analysis of ecological experiments.Chapman & Hall, New York, pp. 253–289.Google Scholar
  12. Fraver S. 1992. The insulating value of serotinous cones in protecting pitch pine (Pinus rigida) seeds from high temperatures.Journal of the Pennsylvania Academy of Science 65: 112–116.Google Scholar
  13. Gauthier S., Bergeron Y. and Simon J.P. 1993. Cone serotiny in jack pine: ontogenetic, positional and environmental effects.Canadian Journal of Forestry Research 23: 394–401.Google Scholar
  14. 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.CrossRefGoogle Scholar
  15. Gutterman Y. 2001. Regeneration of plants in arid ecosystems resulting from patch disturbance. Geobotany 27, Kluwer Academic Publishers, The Hague, pp. 1–232.Google Scholar
  16. Habrouk A., Retana J. and Espelta J.M. 1999. Role of heat tolerance and cone protection of seeds in the response of three pine species to wildfires. Plant Ecology 145: 91–99.CrossRefGoogle Scholar
  17. Hanley M.E. and Fenner M. 1998. Pre-germination temperature and the survivorship and onward growth of Mediterranean firefollowing plant species. Acta Oecologica 19: 181–187.CrossRefGoogle Scholar
  18. Hanley M.E. and Lamont B.B. 2000. Heat pre-treatment and the germination of soil and canopy-stored seeds of south-western australian species. Acta Oecologica-International Journal of Ecology 21: 315–321.CrossRefGoogle Scholar
  19. Hellum A.K. and Pelchat M. 1978. Temperature and time affect the release and quality of seed from cones of lodgepole pine from Alberta. Canadian Journal of Forestry research 9: 154–159.Google Scholar
  20. 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.CrossRefGoogle Scholar
  21. Henig-Sever N., Eshel A. and Ne'eman G. 2000. Regulation of the germination of Aleppo pine (Pinus halepensis) by nitrate, ammonium, and gibberellin, and its role in post-fire forest regeneration.Physiologia Plantarum 108: 390–397.CrossRefGoogle Scholar
  22. Herranz J.M., Ferrandis P. and Martinez-Sanches J.J. 1998. Influence of heat on seed germination of seven Mediterranean Luguminosae species. Plant Ecology 136: 95–103.CrossRefGoogle Scholar
  23. Izhaki I., Henig-Sever N. and Ne'eman G. 2000. Soil seed banks in Mediterranean Aleppo pine forests: the effect of heat, cover and ash on seedling emergence. Journal of Ecology 88: 667– 675.CrossRefGoogle Scholar
  24. Johnson E.A. and Gutsell S.L. 1993. Heat budget and fire behaviour associated with the opening of serotinous cones in two Pinus species. Journal of Vegetation Science 4: 745–750.CrossRefGoogle Scholar
  25. Keeley J.E. 1987. Role of fire in seed germination of woody taxa in California chaparral. Ecology 68: 434–443.CrossRefGoogle Scholar
  26. Keeley J.E. 1994. Seed-germination patterns in fire-prone Mediterranean-climate regions. In: Arroyo M.T.K., Zedler P.H. and Fox M.D. (eds), Ecology and Biogeography of Mediterranean ecosystems in Chile, California and Australia. Springer-Verlag, New York, pp. 239–273.Google Scholar
  27. Keeley J.E. and Bond W.J. 1997. Convergent seed germination in South African fynbos and Californian chaparral. Plant Ecology 133: 153–167.CrossRefGoogle Scholar
  28. Keeley J.E. and Fotheringham C.J. 1998. Smoke induced seed germination in California chaparral. Ecology 79: 2320–2336.CrossRefGoogle Scholar
  29. Keeley J.E. and Zedler P.H. 1998. Evolution of life histories in pines. In: Richardson D.M. (ed.), Ecology and biogeography of Pinus. Cambridge University Press, Cambridge, pp. 219–249.Google Scholar
  30. Lamont B.B. 1991. Canopy seed storage and release-what's in a name? Oikos 60: 266–268.Google Scholar
  31. Lamont B.B., Le Maitre D.C., Cowling R.M. and Enright N.J. 1991. Canopy seed storage in woody plants. Botanical Review 57: 277–317.CrossRefGoogle Scholar
  32. Lamont B.B., Witkowski E.T.F. and Enright N.J. 1993. Postfire litter microsites-safe for seeds, unsafe for seedlings. Ecology 74: 501–512.CrossRefGoogle Scholar
  33. Leone V., Logiurato A. and Saracino A. 1999. Serotiny in Pinus halepensis Mill: recent issues. Abstract in MEDPINE, International workshop on Mediterranean pines, February 1999, Beit Oren, Israel.Google Scholar
  34. Martínez-Sánchez J.J., Maran 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
  35. Mayer A.M. and Poljakoff-Mayber A. 1982. The Germination of Seeds. Pergamon Press, Oxford.Google Scholar
  36. Midgley J. 2000. What are the relative costs, limits and correlates of increased degree of serotiny? Austral Ecology 25: 65–68.CrossRefGoogle Scholar
  37. 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.CrossRefGoogle Scholar
  38. 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 forest ecosystems in the Mediterranean basin. Backhuys publishers, Leiden, pp. 105–118.Google Scholar
  39. Ne'eman G. 1997. Regeneration of natural pine forest-review of work done after the 1989 fire in Mount Carmel, Israel. International Journal of Wildland Fire 7: 295–306.CrossRefGoogle Scholar
  40. Ne'eman G., Meir I. and Ne'eman R. 1993. The effect of ash on the germination and early growth of shoots and roots of Pinus, Cistus and annuals. Seed Science and Technology 21: 339–349.Google Scholar
  41. Ne'eman G., Henig-Sever N. and Eshel A. 1999. Regulation of the germination of Rhus coriaria, a post-fire pioneer, by heat, ask, pH and ethylene. Physiologia Plantarum 106: 47–52.CrossRefGoogle Scholar
  42. Nunez M.R. and Calvo L. 2000. Effect of high temperatures on seed germination of Pinus sylvestris and Pinus halepensis. Forest Ecology and Management 131: 183–190.CrossRefGoogle Scholar
  43. Panetsos K.P. 1981. Monograph of Pinus halepensis and Pinus brutia. Annales Forestales 9: 39–77.Google Scholar
  44. Reyes O. and Casal M. 1995. Germination behaviour of 3 species of the genus Pinus in relation to high temperatures suffered during forest fires. Annales des Sciences Forestieres 52: 385– 392.Google Scholar
  45. Reyes O. and Casal M. 1998. Germination of Pinus pinaster, Pradiata, and Ecalyptus globules in relation to the amount of ash produced in forest fires. Annales des Sciences Forestieres 55: 837–845.Google Scholar
  46. Richardson D.M. 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 forest ecosystems in the Mediterranean basin.Backhuys Publishers, Leiden, pp. 131–142.Google Scholar
  47. Tang C., Robson A.D., Longnecker N.E. and Greenway H. 1993.Physiological responses of lupin root to high pH. Plant Soil 155: 509–512.CrossRefGoogle Scholar
  48. Thanos C.A. 2000. Ecophysiology of seed germination of Pinus halepensis and P. brutia. In: Ne'eman G. and Trabaud L. (eds), Ecology, Biogeography and management of P. halepensis and P. brutia forest systems in the Mediterranean Basin. Backhuys Publishers, Leiden, pp. 37–50.Google Scholar
  49. Thanos C.A. and Skordilis A. 1987. The effects of light, temperature and osmotic stress on the germination of Pinus halepensis and P. brutia. Seed Science and Technology 15: 163–174.Google Scholar
  50. Tinker D.B., Romme W.H., Hargrove W.W., Gardner R.H. and Turner M.G. 1994. Landscape-scale heterogeneity in lodgepole pine serotiny. Canadian Journal of Forest Research.Google Scholar
  51. Trabaud L. 1987. Fire and survival traits of plants. In: Trabaud L. (ed.), The role of fire in ecological systems. SPB Academic Publishing, The Hague, pp. 65–91.Google Scholar
  52. Trabaud L. and Renard P. 1999. Do light and litter influence the recruitment of Cistus spp. stands? Israel Journal of Plant Sciences 47: 1–9.Google Scholar
  53. Underwood A.J. 1997. Experiments in Ecology. Their logical design and interpretation using analysis of variance. Cambridge University Press, Cambridge.Google Scholar
  54. Whelan R.J. 1995. The Ecology of Fire. Cambridge University Press, Cambridge.Google Scholar
  55. Whelan R. and Brown C.L. 1998. The role of Callistemon fruits and infructescences in protecting seeds from heat in fires. Australian Journal of Botany 46: 235–239.CrossRefGoogle Scholar
  56. Whelan R.J., de Jong N.H. and Von Der Burg S. 1998. Variation in bradspory and seedling recruitment without fire among populations of Banksia serrata (Proteaceae). Australian Journal of Ecology 23: 121–128.CrossRefGoogle Scholar

Copyright information

© Kluwer Academic Publishers 2003

Authors and Affiliations

  1. 1.Department of Plant EcologyUtrecht UniversityUtrechtThe Netherlands
  2. 2.Department of BiologyHaifa University at OranimHaifaIsrael

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