A Piroecologia do Pinheiro Bravo

Paulo Fernandes*¹, Hermínio Botelho** e Francisco Rego***

*Investigador Auxiliar

**Professor Associado

Centro de Estudos em Gestão de Ecossistemas. Departamento Florestal. Universidade de Trás-os-Montes e Alto Douro. Quinta de Prados, 5000-911 VILA REAL

***Professor Associado com Agregação

Centro de Ecologia Aplicada Prof. Baeta Neves. Instituto Superior de Agronomia. Tapada da Ajuda, 1349-017 LISBOA

Sumário. O pinheiro bravo apresenta características interpretáveis como adaptações evolutivas ao fogo e que são compatíveis com as estratégias de resistência (casca espessa), que responde adequadamente à ocorrência de fogos de severidade ecológica reduzida, e de evasão (pinhas com deiscência térmica), correspondente a fogos de severidade elevada. Embora esta simultaneidade de estratégias pressuponha a capacidade de persistência face a regimes de fogo variáveis ou mistos, o grau de desenvolvimento e de coexistência das características em causa é bastante variável entre as populações ibéricas da espécie. Assim, três regimes de fogo são distinguíveis: fogo de substituição, de severidade elevada e ocorrência imprevisível; fogo de desbaste, de severidade reduzida ou moderada; e fogo de substituição, de severidade elevada.

Palavras-chave: Pinus pinaster; ecologia do fogo; efeitos do fogo; regime de fogo; dinâmica da vegetação

The Fire Ecology of Maritime Pine

Abstract. Maritime pine has characteristics that can be interpreted as evolutionary adaptations to fire and are consistent with the strategies of resistance (thick bark), an adequate response to low-severity fire occurrence, and evasion (cone serotiny), correspondent to high-severity fire. Such concurrent strategies should enable persistence under a variable or mixed fire regime, but the development and coexistence of the traits involved varies among Iberian populations of the species. Three distinct fire regimes can therefore be distinguished: high-severity, stand-replacement, unpredictable fire; thinning fire of low to moderate severity; and high-severity, stand-replacement fire.

Key words: Pinus pinaster; fire ecology; fire effects; fire regime; vegetation dynamics

L'Écologie du Feu du Pin Maritime

Résumé. Le pin maritime présente des caractéristiques qui peuvent être interprétées comme des adaptations évolutives au feu et qui sont compatibles avec les stratégies de résistance (écorce épaisse), qui répondent de façon appropriée à l'occurrence de feux de sévérité écologique réduite, et d'évasion (cônes avec déhiscence thermique), correspondant à des feux de sévérité élevée. De telles stratégies concourantes devraient permettre la persistance sous un régime de feu variable ou mixte, mais le développement et la coexistence des caractéristiques impliquées change parmi les populations ibériennes de l'espèce. Trois régimes distincts du feu peuvent donc être distingués: feu de substitution, de sévérité élevée et d'occurrence imprévisible; feu d'éclaircie, de sévérité réduite ou modérée; et feu de substitution, de sévérité élevée.

Mots clés: Pinus pinaster; écologie du feu; effets du feu; régime de feu; dynamique de la végétation

Texto completo disponivel apenas em PDF.

Full text only available in PDF format.

Bibliografia

AGEE, J.K., 1993. Fire ecology of Pacific Northwest forests. Island Press, Washington D.C.        [ Links ]

AGEE, J.K., 1998. Fire and pine ecosystems. In Ecology and biogeography of Pinus (ed. D.M. Richardson), Cambridge University Press, Cambridge, pp. 193-218.        [ Links ]

ALÍA, R., MARTÍN, S., DE MIGUEL, J., GALERA, R., AGÚNDEZ, D., GORDO, J., SALVADOR, L., CATALÁN, G., GIL, L., 1996. Regiones de procedencia Pinus pinaster Aiton. Ministerio de Medio Ambiente, Organismo Autónomo Parques Nacionales, Madrid.        [ Links ]

BERBIGIER, P., BONNEFOND, J.M., 1995. Measurement and modelling of radiation transmission within a stand of maritime pine (Pinus pinaster Ait). Ann. Sci. For. 52 : 23-42.        [ Links ]

BOND, W.J., VAN WILGEN, B.W., 1996. Fire and plants. Population and Community Biology Series, Kluwer Academic Publishers, New York.        [ Links ]

BOTELHO, H.S., 1996. Efeitos do fogo controlado em árvores de povoamentos jovens de Pinus pinaster Ait. Dissertação de Doutoramento, UTAD, Vila Real.        [ Links ]

BOTELHO, H., FERNANDES, P., RUAS, L., 1998a. Modeling Pinus pinaster trees damage induced by up-slope wind-driven prescribed fires in Northern Portugal. In Proc. 13th conf. on fire and forest meteorology, IAWF, pp. 473-476.        [ Links ]

BOTELHO, H.S., FERNANDES, P., LOUREIRO, C., 1998c. Growth response of maritime pine (Pinus pinaster) trees to high-intensity prescribed fires. In Proc. 3rd int. conf. on forest fire research & 14th fire and forest meteorology Conf. (ed. D.X. Viegas), ADAI, Coimbra, pp. 1863-1873.        [ Links ]

BOTELHO, H.S., REGO, F.C., RYAN, K.C., 1998b. Tree mortality models for Pinus pinaster of Northern Portugal. In Proc. 13th conf. on fire and forest meteorology, IAWF, pp. 235-240.        [ Links ]

BROWN, J.K., 2000. Introduction and fire regimes. In Wildland fire in ecosystems: effects of fire on flora. USDA For. Serv. Gen. Tech. Rep. RMRS-GTR-42-vol. 2, Rocky Mte. Res. Stn., Fort Collins, pp. 1-8.        [ Links ]

BURROWS, N., WARD, B., ROBINSON, A., 2000. Behaviour and some impacts of a large wildfire in the Gnangara maritime pine (Pinus pinaster) plantation, Western Australia. CALMScience 3(2) : 251-260.        [ Links ]

BYRAM, G.M., 1948. Vegetation temperature and fire damage in southern pines. Fire Control Notes 9 : 34-36.        [ Links ]

BYRAM, G.M., 1958. Some basic thermal processes controlling the effects of fire on living vegetation. USDA For. Serv. Res. Note 114, Southeast. For. Exp. Stn., Asheville.        [ Links ]

CARCAILLET, C., BARAKAT, H.N., PANAIOTIS, C., LOISEL, R., 1997. Fire and late-Holocene expansion of Quercus ilex and Pinus pinaster on Corsica. J. Veg. Sci. 8(1) : 85-94.        [ Links ]

CARRIÓN, J.S., NAVARRO, C., NAVARRO, J., MUNUERA, M., 2000. The distribution of cluster pine (Pinus pinaster) in Spain as derived from palaeocological data: relationships with phytosociological classification. Holocene 10(2) : 243-252.        [ Links ]

CARRIÓN, J.S., VAN GEEL, B., 1999. Fine-resolution Upper Weichselian and Holocene palynological record from Navarrés (Valencia, Spain) and a discussion about factors of Mediterranean forest succession. Rev. Palaeobot. Palyno. 106 : 209-236.        [ Links ]

CASTRO, E.B., GONZÁLEZ, M.C., TENORIO, M.C., BOMBÍN, R.E., ANTÓN, M.G., FUSTER, M.G., MANZANEQUE, A.G., MANZANEQUE, F.G., SAIZ, J.M., JUARISTI, C.M., PAJARES, P.R., OLLERO, H.S., 1997. Los bosques ibéricos, una interpretación geobotánica. Editorial Planeta, S.A., Barcelona.        [ Links ]

CASTRO, J.F., BENTO, J., REGO, F., 1990. Regeneration of Pinus pinaster forests after wildfire. In Fire in ecosystem dynamics, Proc. 3rd int. symp. fire ecology (eds. J.G. Goldhammer, M.J. Jenkins), SPB Academic Publishing, The Hague., pp. 71-75.        [ Links ]

DE RONDE, C., 1982. The resistance of Pinus species to fire damage. S. Afr. For. J. 122 : 22-25.        [ Links ]

DE RONDE, C., 1983. Controlled burning in pine stands in the Cape: the influence of crown scorch on tree growth and litterfall. S. Afr. For. J. 123 : 39-41.        [ Links ]

DE RONDE, C., GOLDAMMER, J.G., WADE, D., SOARES, R.V., 1990. Prescribed fire in industrial pine plantations. In Fire in the tropical biota: ecosystem processes and global challenges (ed. J. Goldammer), Springer-Verlag, Berlin, pp. 216-272.        [ Links ]

DUHOUX, F., 1994. Sensibilites de plusieurs especes arborees mediterraneennes aux stress thermiques et prediction de la mortalite apres incendie du pin d'alep et du pin pignon. Mémoire de fin d'études, ETSIPA/INRA, Avignon.        [ Links ]

ESCUDERO, A., SANZ, M.V., PITA, J.M., PÉREZ-GARCÍA, F., 1999. Probability of germination after heat treatment of native Spanish pines. Ann. For. Sci. 56 : 511-520.        [ Links ]

ESCUDERO, A., BARRERO, S., PITA, J.M., 1997. Effects of high temperatures and ash on seed germination of two Iberian pines (Pinus nigra ssp. salzmannii, P. sylvestris var. iberica). Ann. Sci. For. 54 : 553-562.        [ Links ]

ESCUDERO, A., NÚÑEZ, Y., PÉREZ-GARCÍA, F., 2000. Is fire a selective force of seed size in pine species? Acta Oecol. 21(4-5) : 245-256.        [ Links ]

FAHNESTOCK, G.R., HARE, R.C., 1964. Heating of tree trunks in surface fires. J. For. 62 : 799-805.        [ Links ]

FERNANDES, P.M., 2002. Desenvolvimento de relações preditivas para uso no planeamento de fogo controlado em povoamentos de Pinus pinaster Ait. Dissertação de Doutoramento, UTAD, Vila Real.        [ Links ]

FIGUEIRAL, I., 1995. Charcoal analysis and the history of Pinus pinaster (cluster pine) in Portugal. Rev. Palaeobot. Palyno. 89 : 441-454.        [ Links ]

FISCHER, W.C., MILLER, M., JOHNSTON, C.M., SMITH, J.K., SIMMERMAN, D.G., BROWN, J.K., 1996. Fire effects information system: user's guide. USDA For. Serv. Gen. Tech. Rep. INT-GTR-327, Intermt. Res. Stn., Ogden.        [ Links ]

FONDA, R.W., 2001. Burning characteristics of needles from eight pine species. For. Sci. 47(2) : 390-396.        [ Links ]

FREITAS, H., 1995. Efeitos do fogo nos processos regenerativos do Pinus pinaster. Rel. Fin. de estágio da Lic. em Engª Florestal, UTAD, Vila Real.        [ Links ]

GAUTHIER, S., BERGERON, Y., SIMON, J.P., 1993. Cone serotiny in jack pine: ontogenetic, positional, and environmental effects. Can. J. For. Res. 23 : 394-401.        [ Links ]

GAUTHIER, S., BERGERON, Y., SIMON, J.P., 1996. Effects of fire regime on the serotiny level of jack pine. J. Ecol. 84 : 539-548.        [ Links ]

HARE, R.C., 1961. Heat effects on living plants. USDA For. Serv. Occas. Pap. 183, South. For. Exp. Stn., Asheville.        [ Links ]

HARE, R.C., 1965. Contribution of bark to fire resistance of southern trees. J. For. 63 : 248-251.        [ Links ]

HARMON, M.E., 1984. Survival of trees after low-intensity surface fires in Great Smoky Mountains National Park. Ecology 65(3) : 796-802.        [ Links ]

HENGST, G.E., DAWSON, J.O., 1994. Bark properties and fire resistance of selected tree species from the central hardwood region of North America. Can. J. For. Res. 24 : 688-696.        [ Links ]

HERNANDO, C., GUIJARRO, M., SANTOS, J.A., 1995. Determinación de la relación superficie/volumen de las acículas muertas. Invest. Agr.: Sist. Recur. For. 4(1) : 73-85.        [ Links ]

JACKSON, J.F., ADAMS, D.C., JACKSON, U.B., 1999. Allometry of constitutive defense: a model and a comparative test with tree bark and fire regime. Am. Nat. 153(6) : 614-632.        [ Links ]

JRC, 2003. Update to the report ‘The European Forest Fires Information System (EFFIS) results on the 2003 fire season in Portugal’ (Sept. 15, 2003). European Commission, Directorate General, Institute for Environment and Sustainability, Land Management Unit, Ispra.        [ Links ]

KEELEY, J.E., ZEDLER. P.H., 1998. Evolution of life histories in Pinus. In Ecology and biogeography of Pinus (ed. D.M. Richardson), Cambridge University Press, Cambridge, pp. 219-250.        [ Links ]

LANDERS, J.L., 1991. Disturbance influences on pine traits in the Southeastern United States. In High intensity fire in wildlands: management challenges and options, Proc. 10th Tall Timbers fire ecology conf., Tall Timbers Res. Stn., Tallahassee, pp. 61-97.        [ Links ]

LANNER, R.M., 1998. Seed dispersal in Pinus. In Ecology and biogeography of Pinus (ed. D.M. Richardson), Cambridge University Press, Cambridge, pp. 281-295.        [ Links ]

LUIS-CALABUIG, E., TORRES, O., VALBUENA, L., CALVO, L., MARCOS, E., 2002. Impact of large fires on a community of Pinus pinaster. In Fire and biological processes (eds. L. Trabaud, R. Prodon), Backhuys Publishers, Leiden, pp. 1-12.        [ Links ]

MARTIN, R.E., 1963. Thermal properties of bark. Forest Products Journal 13(10) : 419-426.        [ Links ]

MARTÍN, R.T., GIL, L., 2000. Adaptación reproductiva de las especies forestales ante el fuego. In La defensa contra incendios forestales: fundamentos y experiencias. McGraw-Hill, Madrid, pp. 4.36-4.66.        [ Links ]

MARTÍNEZ, E., MADRIGAL, J., HERNANDO, C., GUIJARRO, M., VEGA, J.A., PÉREZ-GOROSTIAGA, P., FONTURBEL, M.T., CUIÑAS, P., ALONSO, M., BELOSO, M., 2002. Effect of fire intensity on seed dispersal and early regeneration in a Pinus pinaster forest. In Proc. IV int. conf. forest on fire research & 2002 wildland fire safety summit (ed. D.X. Viegas), Millpress Science Publishers, Rotherdam, CD-ROM        [ Links ]

MARTÍNEZ-SÁNCHEZ, J.J., MARÍN, A., HERRANZ, J.M., FERRANDIS, P., 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(1) : 69-72.        [ Links ]

MAUGÉ, J.P., 1987. Le pin maritime premier résineux de France. Centre de Productivité & d'Action Forestière d'Aquitaine, Institut pour le Développement Forestier, Paris.        [ Links ]

MCCORMICK, J., 1976. Recovery of maritime pine (Pinus pinaster) after severe crown scorch. Res. Pap. No. 20, Forests Dept. of West. Austr., Perth.        [ Links ]

MCRAE, D.J., TODD, B., OGILVIE, C.J., 1991. A prescribed fire ignition expert system for Canada. In Proc. 11th conf. fire and forest meteorology (eds., P.L. Andrews, D.F. Potts), SAF Pub. 91-04, Soc. Am. For., Bethesda, pp. 234-241.        [ Links ]

MOREIRA, F., REGO, F.C., FERREIRA, P.G., 2001. Temporal (1958-1995) pattern of change in a cultural landscape of northwestern Portugal: implications for fire occurrence. Landscape Ecol. 16 : 557-567.        [ Links ]

MUTCH, R.W., 1970. Wildland fires and ecosystems – a hypothesis. Ecology 51(6) : 1046-1051.        [ Links ]

NICOLAS, A., GANDULLO, J.M., 1967. Ecologia de los pinares españoles. 1. Pinus pinaster Ait. Ministerio de Agricultura, Instituto Forestal de Investigaciones y Experiencias, Madrid.        [ Links ]

OLIVEIRA, A.C., PEREIRA, J.S., CORREIA, A.V., 2000. A silvicultura do pinheiro bravo. Centro Pinus, Porto.        [ Links ]

PARK, A., 2003. Spatial segregation of pines and oaks under different fire regimes in the Sierra Madre Occidental. Plant Ecol. 169 : 1-20.        [ Links ]

PAUSAS, J.G., 1999a. Mediterranean vegetation dynamics: modelling problems and functional types. Plant Ecol. 140 : 27-39.        [ Links ]

PAUSAS, J.G. 1999b. Response of plant functional types to changes in the fire regime in Mediterranean ecosystems: a simulation approach. J. Veg. Sci. 10 : 717-722.        [ Links ]

PEET, G.B., MCCORMICK, J., 1971. Short–term responses from controlled burning and intense fires in the forests of Western Australia. For. Dept. West. Austr. Bull. No. 79, Perth.        [ Links ]

PEREIRA, J.S., CORREIA, A.V., CORREIA, A.P., BRANCO, M., BUGALHO, M., CALDEIRA, M., CRUZ, C., FREITAS, H., OLIVEIRA, A., PEREIRA, J.M., REIS, R., VASCONCELOS, M.J. 2002. Forests and biodiversity. In Climate change in Portugal. Scenarios, impacts and adaptation measures – SIAM project (eds. F.D. Santos, K. Forbes, R. Moita), Gradiva, Lisboa, pp. 367-417.        [ Links ]

PLATT, W.J., GLITZENSTEIN, J.S., STRENG, D.R., 1991. Evaluating pyrogenicity and its effects on vegetation in longleaf pine savannas. In High intensity fire in wildlands: management challenges and options, Proc. 17th Tall Timbers fire ecology conf., Tall Timbers Res. Stn., Tallahassee, pp. 61-97.        [ Links ]

REBERTUS, A.J., WILLIAMSON, G.B., MOSER, B., 1989a. Longleaf pine pyrogenicity and turkey oak mortality in Florida xeric sandhills. Ecology 70(1) : 60-70.        [ Links ]

REBERTUS, A.J., WILLIAMSON, G.B., MOSER, B., 1989b. Fire-induced changes in Quercus laevis spatial pattern in Florida sandhills. J. Ecol. 77 : 638-650.        [ Links ]

REGO, F.C., 1986. Effects of prescribed fire on vegetation and soil properties in Pinus pinaster forests of Northern Portugal. PhD thesis, University of Idaho, Moscow.        [ Links ]

REYES, O., CASAL, M., 1998a. Are the dominant species in NW Spain fire-prone? In Fire management and landscape ecology (ed. L. Trabaud), IAWF, pp. 177-188.        [ Links ]

REYES, O., CASAL, M., 1998b. Germination of Pinus pinaster, P. radiata and Eucalyptus globulus in relation to the amount of ash produced in forest fires. Ann. Sci. For. 55 : 837-845.        [ Links ]

RICHARDSON, D.M., RUNDEL, P.W., 1998. Ecology and biogeography of Pinus: an introduction. In Ecology and biogeography of Pinus (ed. D.M. Richardson), Cambridge University Press, Cambridge, pp. 3-46.        [ Links ]

RODRÍGUEZ-TREJO, D., FULÉ, P., 2003. Fire ecology of Mexican pines and a fire management proposal. Int. J. Wildland Fire 12(1) : 23-37.        [ Links ]

RYAN, K.C., 1998. Analysis of the relative value of morphological variables in predicting fire-caused tree mortality. In Proc. 3rd int. conf. on forest fire research & 14th fire and forest meteorology conf. (ed. D.X. Viegas), ADAI, Coimbra, pp. 1511-1526.        [ Links ]

RYAN, K.C., REINHARDT, E.D., 1988. Predicting postfire mortality of seven western conifers. Can. J. For. Res. 18 : 1291-1297.        [ Links ]

RYAN, K.C., RIGOLOT, E., BOTELHO, H., 1994. Comparative analysis of fire resistance and survival of Mediterranean and North-American conifers. In Proc. 12th conf. fire and forest meteorology, SAF Pub. 94-02, Soc. Am. For., Bethesda, pp. 701-708.        [ Links ]

SCHWILK, D.W., ACKERLY, D.D., 2001. Serotiny and flammability as strategies: correlated evolution in pines. Oikos 94 : 326-336.        [ Links ]

SEVA, J.P., A. VALDECANTOS, A., VILAGROSA, A., CORTINA, J., BELLOT, VALLEJO, V.R., 2000. Seedling morphology and survival in Mediterranean tree and shrub species. In Proc. Mediterranean desertification research results and policy implications (eds. P. Balabanis, D. Peter, A. Ghazi, M. Tsogas), European Commission, Directorate-General Research, pp. 397-406.        [ Links ]

TAPIAS, R., GIL, L., FUENTES-UTRILLA, P., PARDOS, J.A., 2001. Canopy seed banks in Mediterranean pines of south-eastern Spain: a comparison between Pinus halepensis Mill., P. pinaster Ait., P. nigra Arn., and P. pinea L. J. Ecol. 89 : 629-638.        [ Links ]

VALBUENA, L., CALVO, L., 1998. Seedling establishment after a wildfire in Pinus pinaster Ait. and Pinus nigra Arn. stands. In Proc. 3rd int. conf. forest fire research & 14th conf. fire and forest meteorology (ed. D.X. Viegas), ADAI, pp. 1901-1911.        [ Links ]

VAN WAGNER, C.E., 1977. Conditions for the start and spread of crown fire. Can. J. For. Res. 3 : 373-378.        [ Links ]

VÁZQUEZ, A., MORENO, J.M., 1998. Patterns of lightning-, and people-caused fires in Peninsular Spain. Int. J. Wildland Fire 8(2) : 103-115.        [ Links ]

VEGA, J.A., 1978. Utilizacion del fuego controlado en las comunidades vegetales de Galicia. Bol. Estac. Central Ecol. 7(14) : 3-19.        [ Links ]

VEGA, J.A., 2000. Resistencia vegetativa ante el fuego a través de la historia de los incendios. In La defensa contra incendios forestales: fundamentos y experiencias, McGraw-Hill, Madrid, pp. 4.66-4.85.        [ Links ]

VEGA, J.A., BARA, S., GIL, M.C., 1983. Prescribed burning in pine stands for fire prevention in the NW of Spain: some results and effects. Freiburger Waldschutz Abhandlungen 4 : 49-74.        [ Links ]

VEGA, J.A., BARA, S., GIL, M.C., 1985. Algunos efectos a corto plazo del fuego prescrito en pinares de Galicia. In Estudios sobre prevención y efectos ecológicos de los incendios forestales, MAPA, ICONA, Madrid, pp. 103-145.        [ Links ]

VEGA, J.A., CUIÑAS, P. BARÁ, S., FONTURBEL, M.T., SANTOS, J.A., ROZADOS, M.J., ALONSO, M., BELOSO, M.C., CALVO, E., 1993. Informe final del proyecto Forest Fire Prevention through Prescribed Burning: Experimental Study on Fire Effects on Litter and Soil, contract no. CE/STEP-CT90-0087, CIF, Lourizãn.        [ Links ]

WILLIAMSON, G.B., BLACK, E.M., 1981. High temperature of forest fires under pines as a selective advantage over oaks. Nature 293 : 643-644.        [ Links ]

Entregue para publicação em Janeiro de 2004

¹ 1º Autor E-mail: ffern@utad.pt