Natural regeneration in Iberian pines: A review of dynamic processes and proposals for management
Abstract
Aim of study: Designing adequate silvicultural systems for natural regeneration of a forest species requires sound knowledge of the underlying ecological subprocesses: flowering and fruiting, seed dispersal and predation, seed germination, seedling emergence and seedling survival. The main objective of the present work is to carry out a review on the current knowledge about the different subprocesses governing the regeneration process for the main Iberian Pinus species, in order to propose scientifically based management schedules.
Area of study: The review focuses on the five main native Pinus species within their most representative areas in the Iberian Peninsula: Pinus nigra in Cuenca mountains, Pinus sylvestris in Sierra de Guadarrama, Pinus pinaster and Pinus pinea in the Northern Plateau and Pinus halepensis in Catalonia
Material and methods: Firstly, currently available information on spatiotemporal dynamics and influential factors is introduced for each subprocess and species. Secondly, current regeneration strategies are characterized and the main bottlenecks are identified. Finally, alternative silvicultural practices proposed on the light of the previous information are presented.
Main results: Different climate-mediated bottlenecks have been identified to limit natural regeneration of the Iberian pine species, with seed predation and initial seedling survival among the most influential. New approaches focusing on more gradual regeneration fellings, extended rotation periods, prevent big gaps and program fellings on mast years are presented.
Research highlights: Natural regeneration of the studied species exhibit an intermittent temporal pattern, which should be aggravated under drier scenarios. More flexible management schedules should fulfil these limitations.
Downloads
References
Aizen MA, Patterson WA, 1990. Acorn size and geographical range in the North American oaks (Quercus L.). J Biogeog 17: 327-332. https://doi.org/10.2307/2845128
Alejano R, González JM, Serrada R, 2008. Selvicultura de Pinus nigra Arn. subsp salzmannii (Dunal) Franco. In: Serrada R, Montero G, Reque JA (eds.). Compendio de selvicultura aplicada en España. INIA-FUCOVASA, Madrid. pp: 313-355.
Awada T, Radoglou K, Fotelli MN, Constantinidou HIA, 2003. Ecophysiology of seedlings of three Mediterranean pine species in contrasting light regimes. Tree Physiol 23 (1): 33-41. https://doi.org/10.1093/treephys/23.1.33
Barbeito I, Pardos M, Calama R, Cañellas I, 2008. Effect of stand structure on stone pine (Pinus pinea L.) regeneration dynamics. Forestry 81 (5): 617-629. https://doi.org/10.1093/forestry/cpn037
Barbeito I, Fortin MJ, Montes F, Cañellas I, 2009. Response of pine natural regeneration to small-scale spatial variation in a managed Mediterranean mountain forest. Appl Veg Sci 12: 488-503. https://doi.org/10.1111/j.1654-109X.2009.01043.x
Baskin CC, Baskin JM, 2001. Seeds. Ecology, biogeography, and evolution of dormancy and germination. Acad Press, San Diego, USA.
Beltrán M, Piqué M, Vericat P, Cervera T, 2011. Models de gestió per als boscos de pi blanc (Pinus halepensis): producció de fusta i prevenció d'incendis forestals. Sèrie: Orientacions de gestió forestal sostenible per a Catalunya (ORGEST). Centre de la Propietat Forestal. Generalitat de Catalunya.
Broncano MJ, Riba M, Retana J, 1998. Seed germination and seedling performance of two Mediterranean tree species, holm oak (Quercus ilex L.) and Aleppo pine (Pinus halepensis Mill.): a multifactor experimental approach. Plant Ecol 138: 17-26. https://doi.org/10.1023/A:1009784215900
Broncano MJ, Rodrigo A, Retana J, 2008. Post-dispersal seed predation in Pinus halepensis and consequences on seedling establishment after fire. Int J Wildl Fire 17: 407-414. https://doi.org/10.1071/WF07095
Burschel P, Huss J, 1997. Grundiβ des Waldbaus. Parey Buchverlag, Berlin. 487 pp.
Cabrera M, Donés J, 2010. Proyecto de la 7ª Revisión de la Ordenación del Monte de U. P. Nº 2 de la provincia de Segovia "Pinar" de Valsaín, perteneciente al OAPN, sito en el término municipal de San Ildefonso de La Granja. MAPAMA. http://www.mapama.gob.es/es/parques-nacionales-oapn/centros-fincas/valsain/ordenaciones.aspx .
Calama R, Finat L, Gordo FJ, Bachiller A, Ruíz-Peinado R, Montero G, 2005. Estudio comparativo de la producción de madera y piña en masas regulares e irregulares de Pinus pinea en la provincia de Valladolid. IV Cong For Esp (SECF). Zaragoza, Sept.
Calama R, Mutke S, Tomé JA, Gordo FJ, Montero G, Tomé M, 2011. Modelling spatial and temporal variability in a zero-inflated variable: the case of stone pine (Pinus pinea L.) cone production. Ecol Model 222: 606-618. https://doi.org/10.1016/j.ecolmodel.2010.09.020
Calama R, Mutke S, Pardos M, Morales L, Gordo Fj, Montero G, Finat L, 2012. La producción de piña y piñón de Pinus pinea L. en la Meseta Norte; Gordo FJ, et al. (eds.): La regeneración natural de los pinares en los arenales de la meseta castellana: 81-98, iuFOR-INIA-JCyL.
Calama R, Manso R, Barbeito I, Madrigal G, Garriga E, Gordo FJ, Montero G, Cañellas I, Pardos M, 2015a. Do inter-specific differences in seed size determine natural regeneration traits in Pinus pinea and Pinus sylvestris? Appl Ecol Environ Res 13 (2): 387-404.
Calama R, Manso R, Barbeito I, Cañellas I, Pardos M, 2015b. Is natural regeneration a bottleneck in the performance of Pinus sylvestris mountain forests? Mountain Forest Management in a Changing World, Smokovce, 6-9 July.
Calama R, Puértolas J, Manso R, Pardos M, 2015c. Defining the optimal regeneration niche for Pinus pinea L. through physiology-based models for seedling survival and carbon assimilation. Trees 29 (6): 1761-1771. https://doi.org/10.1007/s00468-015-1257-5
Calama R, Gordo FJ, Madrigal G, Mutke S, Conde M, Montero G, Pardos M, 2016. Enhanced tools for predicting annual stone pine (Pinus pinea L.) cone production at tree and forest scale in Inner Spain. Forest Syst 25(3): e079 . https://doi.org/10.5424/fs/2016253-09671
Calama R, Fortin M, Pardos M, Manso R, 2017. Modelling spatiotemporal dynamics of Pinus pinea cone infestation by Dioryctria mendacella. For Eco Manage 389: 136-148.
Cañellas I, Montes F, del Río M, Pardos M, Montero G, 2004. The effect of different soil substrates in the germination and survival of seed in Pinus sylvestris L. regeneration mountain forests. In: Ammer Ch, Weber M, Mosandl R (eds), Conf Proc. Kloster Seeon, Germany, p 83.
Castro J, Gómez JM, García D, Zamora R, Hódar JA, 1999. Seed predation and dispersal in relict Scots pine forests in southern Spain. Plant Ecol 145 (1): 115-123. https://doi.org/10.1023/A:1009865703906
de Las Heras J, Moya D, Vega JA, Daskalakou E, Vallejo R, Grigoriadis N, Tsitsoni T, Baeza J, Valdecantos A, Fernández C, Espelta JM, Fernandes P, 2012. Post-fire management of serotinous pine forests. In: Post-Fire Management and Restoration of Southern European Forests, vol 24; Moreira F et al. (eds.), pp: 121-150. Springer, Dordrecht, Netherlands. https://doi.org/10.1007/978-94-007-2208-8_6
del Peso C, Bravo F, Ruano I, Pando V, 2012. Patrones de diseminación y nascencia de Pinus pinaster en Mesta castellana. In: La regeneración natural de los pinares en los arenales de la Meseta Castellana; Gordo FJ, et al. (eds.). pp 161-174. iuFOR-INIA-JCyL.
del Río M, Calama R, Montero G, 2008. Selvicultura de Pinus halepensis Mill. In: Compendio de selvicultura aplicada en España; Serrada R, Montero G, Reque JA (eds.), pp: 289-312. INIA-FUCOVASA, Madrid.
Espelta JM, 1996. La regeneración de bosques de encina (Quercus ilex L.) y pino carrasco (Pinus halepensis Mill.): Estudio experimental de la respuesta de las plántulas a la intensidad de luz y disponibilidad hídrica. Tesis doctoral, UAB, 187 pp.
Espelta JM, Verkaik I, Eugenio M, Lloret F, 2008. Recurrent wildfires constrain long-term reproduction ability in Pinus halepensis Mill. Int J Wildl Fire 17: 579-585. https://doi.org/10.1071/WF07078
Espelta JM, Arnan X, Rodrigo A, 2011. Non-fire induced seed release in a weakly serotinous pine: Climatic factors, maintenance costs or both? Oikos 120: 1752-1760. https://doi.org/10.1111/j.1600-0706.2011.19570.x
Eugenio M, Verkaik I, Lloret F, Espelta JM, 2006. Recruitment and growth decline in Pinus halepensis populations after recurrent wildfires in Catalonia (NE Iberian Peninsula). For Eco Manage 231: 47-54.
Gordo FJ, 1999. Ordenación y selvicultura de Pinus pinea en L. en la provincia de Valladolid. In Ciencias y Técnicas Forestales. 150 años de aportaciones de los ingenieros de montes; Madrigal A (Ed.). pp: 79-100. FUCOVASA. Madrid.
Gordo FJ, Montero G, Gil L, 2012a. La problemática de la regeneración natural de los pinares en los arenales de la Meseta Castellana. In: La regeneración natural de los pinares en los arenales de la Meseta Castellana; Gordo FJ et al. (eds.). pp 11-16. iuFOR-INIA-JCyL.
Gordo FJ, Rojo LI, Calama R, Mutke S, Martín R, García M, 2012b. Selvicultura de regeneración natural de Pinus pinea L. en montes públicos de la provincia de Valladolid. In: La regeneración natural de los pinares en los arenales de la Meseta Castellana; Gordo FJ et al. (eds.). pp: 145-159. iuFOR-INIA-JCyL.
Greene DF, Zasada JC, Sioris L, Kneeshawd, Morin H, Charron I, Simard MJ, 1999. A review of the regeneration of boreal forest trees. Can J For Res 29: 824-839. https://doi.org/10.1139/x98-112
Herrero C, Bravo F, San Martin R, 2004. Modelo de probabilidad de germinación de pino negral (Pinus pinaster Aiton) tras incendio. Actas de la Reunión de Modelización Forestal. Cuad Soc Esp Cienc For 18: 57-63.
Hille M, den Ouden J, 2004. Improved recruitment and early growth of Scots pine (Pinus sylvestris L.) seedlings after fire and soil scarification. Eur J For Res 123 (3): 213-218. https://doi.org/10.1007/s10342-004-0036-4
Hilli A, Hokkanen T, Hyvonen J, 2008: Long-term variation in Scots pine seed crop size and quality in northern Finland. Scan J For Res 23 (5): 395-403. https://doi.org/10.1080/02827580802334217
Howe HF, Smallwood J, 1982. Ecology of seed dispersal. Annu Rev Ecol Syst 13: 201-228. https://doi.org/10.1146/annurev.es.13.110182.001221
Hulme PE, 1997. Post-dispersal seed predation and the establishment of vertebrate dispersed plants in Mediterranean scrublands. Oecologia 111: 91-98. https://doi.org/10.1007/s004420050212
Janzen DH, 1976. Why bamboos wait so long to flower. Ann Rev Ecol Syst 7: 347-391. https://doi.org/10.1146/annurev.es.07.110176.002023
Jordano P, 1992. Fruits and frugivory. In: Seeds: the ecology of regeneration in natural plant communities; Fenner M (ed). CAB Int, Wallingford.
Juez L, González-Martínez SC, Nanos N, Lucas AI, Ordoñez C, del Peso C, Bravo F, 2014. Can seed production and restricted dispersal limit recruitment in Pinus pinaster Ait from the Spanish Northern Plateau? For Eco Manage 313: 329-339.
Kelly D, 1994. The evolutionary ecology of mast seedling. Trends Ecol Evol 9: 465-470. https://doi.org/10.1016/0169-5347(94)90310-7
Latham RE, 1992. Co-occurring tree species change rank in seedling performance with resources varied experimentally. Ecology 73 (6): 2129-2144. https://doi.org/10.2307/1941461
Lucas-Borja ME, Silva-Santos P, Fonseca T, Tíscar PA, López-Serrano FR, Andrés-Abellán M, Martínez-García E, Del Cerro-Barja A, 2010. Modelling Spanish black pine postdispersal seed predation in Central-eastern Spain. For Syst 19: 393-403.
Lucas-Borja ME, Fonseca T, Parresol B, Silva-Santos P, García-Morote FA, Tíscar-Oliver PA, 2011. Modelling Spanish black pine seedling emergence: Establishing management strategies for endangered forest areas. For Eco Manage 262: 195-202.
Lucas-Borja ME, Fidalgo-Fonseca T, Lousada JL, Silva-Santos P, Martínez-Garcia E, Andrés-Abellán M, 2012. Natural regeneration of Spanish black pine [Pinus nigra Arn. ssp. salzmannii (Dunal) Franco] at contrasting altitudes in a Mediterranean mountain area. Ecol Res 27: 913-931. https://doi.org/10.1007/s11284-012-0969-x
Lucas-Borja ME, Candel-Pérez D, García-Morote FA, Onkelinx T, Tíscar PA, Balandier P, 2016. Pinus nigra Arn. ssp. salzmannii seedling recruitment is affected by stand basal area, shrub cover and climate interactions Ann For Sci 73 (3): 649-656. https://doi.org/10.1007/s13595-016-0550-9
Manso R, Pardos M, Keyes CR, Calama R, 2012. Modelling the spatio-temporal pattern of primary dispersal in stone pine (Pinus pinea L.) stands in the Northern Plateau (Spain). Ecol Model 226: 11-21. https://doi.org/10.1016/j.ecolmodel.2011.11.028
Manso R, Calama R, Madrigal G, Pardos M, 2013a. A silviculture-oriented spatio-temporal model for germination in Pinus pinea L. in the Spanish Northern Plateau based on a direct seeding experiment. Eur J For Res 132 (5): 969-982. https://doi.org/10.1007/s10342-013-0724-z
Manso R, Fortin M, Calama R, Pardos M, 2013b. Modelling seed germination in forest tree species through survival analysis. The Pinus pinea L. case study. For Eco Manage 289: 515-524.
Manso R, Pukkala R, Pardos M, Miina J, Calama R, 2014a. Modelling Pinus pinea forest management to attain natural regeneration under present and future climatic scenarios. Can J For Res 44 (3): 250-262. https://doi.org/10.1139/cjfr-2013-0179
Manso R, Pardos M, Calama R, 2014b. Climatic factors control rodent seed predation in Pinus pinea L. stands in Central Spain. Ann For Sci 71(8): 873-883. https://doi.org/10.1007/s13595-014-0396-y
Martín-Sanz RC, Santos-del-Blanco L, Notivol E, Chambel R, Sanmartín R, Climent J, 2016. Disentangling plasticity of serotiny, a key adaptive traït in a Mediterranean conifer. Am J Bot 103: 1582-1591. https://doi.org/10.3732/ajb.1600199
Masetti C, Mencussini M, 1991. Régénération naturelle du pin pignon (Pinus pinea L.) dans la Pineta Granducale di Alberese (Parco Naturalle della Maremma, Toscana, Italie). Ecol Medit 17: 103-188.
Mast JN, Veblen TT, 1999. Tree spatial patterns and stand development along the pine-grassland ecotone in the Colorado Front Range. Can J For Res 29: 575-584. https://doi.org/10.1139/x99-025
Matias L, Jump AS, 2014. Impacts of predicted climate change on recruitment at the geographical limits of Scots pine. J Exp Bot 65 (1): 299-310. https://doi.org/10.1093/jxb/ert376
Matias L, Mendoza I, Zamora R, 2009. Consistent pattern of habitat and species selection by post-dispersal seed predators in a Mediterranean mosaic landscape. Pl Ecol 203 (1): 137-147. https://doi.org/10.1007/s11258-008-9518-7
MacArthur RH, Wilson EO, 1967. The theory of island biogeography (MPB-1). Princeton University, USA.
Meiners SJ, Pickett STA, Handel SN, 2002. Probability of tree seedling establishment changes across a forest-old field edge gradient. Am J Bot 89 (3): 466-471. https://doi.org/10.3732/ajb.89.3.466
Modrý M, Hubený, Rejsêk K, 2004. Differential response of naturally regenerated European shade tolerant tree species to soil type and light availability. For Eco Manage 188: 185-195.
Montero G, del Río M, Roig S, Rojo A, 2008a. Selvicultura de Pinus sylvestris Mill. In: Compendio de selvicultura aplicada en España; Serrada R, Montero G, Reque JA (eds.). INIA-FUCOVASA, Madrid. pp: 503-534.
Montero G, Calama R, Ruiz-Peinado R, 2008b. Selvicultura de Pinus pinea L. In: Compendio de selvicultura aplicada en España; Serrada R, Montero G, Reque JA (eds.). INIA-FUCOVASA, Madrid. pp: 431-470.
Moya D, Espelta JM, Verkaik I, López-Serrano F, de Las Heras J, 2007. Tree density and site quality influence on Pinus halepensis Mill. reproductive characteristics after large fires. Ann For Sci 64: 649-656. https://doi.org/10.1051/forest:2007043
Moya D, de las Heras J, López-Serrano FR, Leone V, 2008. Optimal intensity and age of management in young aleppo pine stands for post-fire resilience. For Eco Manage 255: 3270-3280.
Muller-Landau HC, Wright SJ, Calderón O, Hubbell SP, Foster RB, 2002. Assessing recruitment limitation: concepts, methods and case-studies from a Tropical Forest. In: Seed dispersal and frugivory: Ecology, evolution and conservation; Levey DJ, et al. (eds). CAB Int,, Wallingford, pp: 35-53. https://doi.org/10.1079/9780851995250.0035
Mutke S, Gordo J, Gil L, 2005a. Variability of Mediterranean stone pine cone production: yield loss as response to climatic change. Agric For Met 132: 263-272. https://doi.org/10.1016/j.agrformet.2005.08.002
Mutke S, Calama R, González-Martinez S, Montero G, Gordo J, Bono D, Gil L, 2012. Mediterranean Stone Pine: Botany and Horticulture. Hortic Rev 39: 153-202.
Mutke S, Calama R, Gordo J, Nicolas JL, Herrero N, Roques A, 2015. Pérdida del rendimiento en piñón blanco de Pinus pinea en fábrica - Leptoglossus y la seca de la piña. III Reun Cien de San For, SECF. Madrid 7-8 oct.
Myczko L, Benkman CW, 2011. Great spotted woodpeckers Dendrocopos major exert multiple forms of phenotypic selection on Scots pine Pinus sylvestris. J Avian Biol 42 (5): 429-433. https://doi.org/10.1111/j.1600-048X.2011.05326.x
Nathan R, Ne'eman G, 2000. Serotiny, seed dispersal and seed predation in Pinus halepensis. In Ecology, biogeography and management of Pinus halepensis and Pinus brutia forest ecosystems in the Mediterranean Basin; Ne'eman G, Trabaud L. (eds). Backhuy Publ, Leiden, pp: 105-118.
Nathan R, Casagrandi R, 2004. A simple mechanistic model of seed dispersal, predation and plant establishment: Janzen-Connel and beyond. J Ecol 92 (5): 733-746. https://doi.org/10.1111/j.0022-0477.2004.00914.x
Ne'eman G, Goubitz S, Nathan R, 2004. Reproductive traits of Pinus halepensis in the light of fire - A critical review. Plant Ecol 171: 69-79. https://doi.org/10.1023/B:VEGE.0000029380.04821.99
Núñez MR, Sierra de Grado R, Alía R, Bravo F, 2013. Efecto del estrés hídrico y la oscilación de las temperaturas sobre la germinación de semillas de diversas procedencias de Pinus pinaster Ait. VI Congr For Esp, Vitoria, junio.
Nyland R. 2002. Silviculture: Concepts and Applications, 2nd ed. The McGraw-Hill Companies, NY.
Pardos M, Ruiz Del Castillo J, Cañellas I, Montero G, 2005. Ecophysiology of natural regeneration of forest stands in Spain. Invest Agrar: Sist Rec For 14 (3): 434-445. https://doi.org/10.5424/srf/2005143-00939
Pardos M, Montes F, Aranda I, Cañellas I, 2007. Influence of environmental conditions on Scots pine (Pinus sylvestris L.) germinant survival and diversity in Central Spain. Eur J For Res 126 (1): 37-47. https://doi.org/10.1007/s10342-005-0090-6
Pardos M, Montes F, Cañellas I, 2008. Spatial dynamics of natural regeneration in two differently managed Pinus sylvestris stands before and after silvicultural intervention using replicated spatial point patterns. For Sci 54 (3): 260-272.
Pardos M, Climent J, Almeida H, Calama R, 2014. The role of developmental stage in frost tolerance of Pinus pinea L. seedlings and saplings. Ann For Sci 71 (5): 551-562. https://doi.org/10.1007/s13595-014-0361-9
Prévosto B, Ripert C, 2008. Regeneration of Pinus halepensis stands after partial cutting in southern France: Impacts of different ground vegetation, soil and logging slash treatments. For Eco Manage 256: 2058-2064.
Prévosto B, Amandier L, Quesney T, de Boisgelin G, Ripert C, 2012. Regenerating mature Aleppo pine stands in fire-free conditions: site preparation treatments matter. For Eco Manage 282: 70-77.
Pukkala T, Kolström T, 1992. A stochastic spatial regeneration model for Pinus sylvestris. Scan J For Res 7: 377-385. https://doi.org/10.1080/02827589209382730
Putman RJ, 1996. Ungulates in temperate forest ecosystems: perspectives and recommendations for future research. For Eco Manage 88: 205-214.
Rodrigo A, Retana J, Picó FX, 2004. Direct regeneration is not the only response of Mediterranean Forests to large fires. Ecology 85: 716-729. https://doi.org/10.1890/02-0492
Rodriguez-García E, Gratzer G, Bravo F, 2011. Climatic variability and other site factors influence on natural regeneration Pinus pinastre Ait. in Mediterranean forests. Ann For Sci 68: 811-823. https://doi.org/10.1007/s13595-011-0078-y
Rojo A, Montero G, 1996. El pino silvestre en la Sierra de Guadarrama. Ministerio de Agricultura, Pesca y Alimentación, 293 pp.
Ruano I, Pando V, Bravo F, 2009. How do light and water influence Pinus pinaster Ait. Germination and early seedling development? For Eco Manage 258: 2647-2653.
Ruano I, del Peso C, Bravo F, 2014. Post-dispersal predation of P. pinaster Aiton seeds: key factors and effects on belowground seed bank. Eur J Forest Res 134 (2): 309-318. https://doi.org/10.1007/s10342-014-0853-z
Ruano I, Manso R, Fortin M, Bravo F, 2015. Extreme climate conditions limit seed availability to succesfully attain natural regeneration of Pinus pinaster in sandy areas of Central Spain. Can J For Res 45: 1795-1802. https://doi.org/10.1139/cjfr-2015-0257
Ruiz de la Torre J, Ceballos L, 1979. Árboles y arbustos de la España peninsular. UPM, ETS Ingenieros de Montes, Madrid.
Salisbury EJ, 1942. The reproductive capacity of plants. Bell, London.
Saura S, Piqué M, 2006. Forests and forest sector in Catalonia. Agric Agri-food Prod Perspect Profile Sect Catalonia. [cd-rom].
Sayer EJ, 2006. Using experimental manipulation to assess the roles of leaf litter in the functioning of forest ecosystems. Biol Rev Camb Philos Soc 81 (1): 1-31. https://doi.org/10.1017/S1464793105006846
Scarascia-Mugnozza G, Schirone B, 1984. Effetti dell'intesitá luminosa sullo sviluppo dei semnzali di pino d'Aleppo. Ann Acad It Sci For 33: 137-153.
Serrada R, 2003. Apuntes de Selvicultura. EUIT Forestal, Universidad Politécnica de Madrid, Madrid. 490 pp.
Serrada R, Domínguez S, Sánchez MI, Ruiz J, 1994. El problema de la regeneración natural del Pinus nigra Arn. Montes 36: 52-57.
Shmida A, Ne'eman G, Goubitz S, Lev-Yadun S, 2000. Sexual allocation and gender segregation in Pinus halepensis. In: Ecology, biogeography, and management of Pinus halepensis and P. brutia forest ecosystems in the Mediterranean basin; Ne'eman G, Trabaud L. (eds), pp: 91-104. Backhuys Publ, Leiden.
Smith CC, Fretwell SD, 1974. The optimal balance between size and number of offspring. Am Nat 108: 499-506. https://doi.org/10.1086/282929
Smith CS, Hamrick JL, Kramer CL, 1990. The advantage of mast years for wind pollination. Am Natur 136: 154-166. https://doi.org/10.1086/285089
Tapias R, Gil L, Fuentes-Utrilla P, Pardos JA, 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. https://doi.org/10.1046/j.1365-2745.2001.00575.x
Thanos CA, Daskalakou EN, 2000. Reproduction in Pinus halepensis and P. brutia. In: Ecology, biogeography, and management of Pinus halepensis and P. brutia forest ecosystems in the Mediterranean basin; Ne'eman G, Trabaud L. (eds), pp: 79-90. Backhuys Publ, Leiden.
Tíscar PA, 2007. Dinámica de regeneración de Pinus nigra subsp. salzmannii al sur de su área de distribución: etapas, procesos y factores implicados. Invest Agrar: Sist Rec For 16: 124-135.
Tíscar PA, Lucas-Borja ME, Candel-Pérez D, 2011. Changes in the structure and composition of two Pinus nigra subsp. salzmannii forests over a century of different silvicultural treatments. For Syst 3: 525-535.
Trabaud L, Campant C, 1991. The difficulty of natural regeneration of Sazmann pine (Pinus nigra subsp. salzmannii) following fire. Biol Conserv 58 (3): 329-343. https://doi.org/10.1016/0006-3207(91)90099-U
Verkaik I, Espelta JM, 2006. Post-fire regeneration thinning, cone production, serotiny and regeneration age in Pinus halepensis. For Eco Manage 231: 155-163.
Westoby M, Jurado E, Leishman M, 1992. Comparative evolutionary ecology of seed size. Trees 7 (11): 368-372. https://doi.org/10.1016/0169-5347(92)90006-w
Worthy FR, Law R, Hulme PE, 2006. Modelling the quantitative effects of pre- and post-dispersal seed predation in Pinus sylvestris L. J Ecol 94 (6): 1201-1213. https://doi.org/10.1111/j.1365-2745.2006.01168.x
Zamora R, Gómez JM, Hódar JA, Castro J, García D, 2001. Effect of browsing by ungulates on sapling growth of Scots pine in a Mediterranean environment: consequences for forest regeneration. For Eco Manage 144: 33-42.
Zavala MA, Espelta JM, Retana J, 2000. Constraints and trade-offs in Mediterranean plant communities: the case of holm oak - Aleppo pine forests. The Bot Rev 66: 119-149. https://doi.org/10.1007/BF02857785
Zavala MA, Espelta JM, Caspersen J, Retana J, 2011. Interspecific differences in sapling performance with respect to light and aridity gradients in Mediterranean pine-oak forests: implications for species coexistence. Can J For Res 41: 1432-1444. https://doi.org/10.1139/x11-050
© CSIC. Manuscripts published in both the printed and online versions of this Journal are the property of Consejo Superior de Investigaciones Científicas, and quoting this source is a requirement for any partial or full reproduction.
All contents of this electronic edition, except where otherwise noted, are distributed under a “Creative Commons Attribution 4.0 International” (CC BY 4.0) License. You may read here the basic information and the legal text of the license. The indication of the CC BY 4.0 License must be expressly stated in this way when necessary.
Self-archiving in repositories, personal webpages or similar, of any version other than the published by the Editor, is not allowed.
