Abstract
Seedling recruitment in many highly serotinous populations of Pinus coulteri on California's central coast depends almost entirely on periodic, stand-replacing fire. Compared to serotinous pines of the Mediterranean Basin, little detailed information is available on the postfire demography of California closed-cone pines, including P. coulteri. In September 1996 a wildfire burned the 760-ha American Canyon Research Natural Area (RNA). Using aerial photography, we mapped burn severity of P. coulteri-chaparral woodlands and forests within the RNA. From May to September of 1997, we also quantified seedling establishment and mortality in relation to biophysical site characteristics including fire severity. Seventy-six percent of P. coulteri forests and woodlands experienced high-severity burns, 9% moderate-severity burns, and 15% low-severity or unburned. Of the 53 plots used for seedling counts, 70% were high-severity, 26% moderate-severity, and 4% low-severity. Seedling densities 13 months postfire were low (0.21 m−2), but seedling mortality also was low (8.4%). Aerial seed bank size increased from north-facing to south-facing slopes and from high-severity to low-severity burns. Seedling recruitment was unrelated to burn severity and increased with the size of the canopy seed bank (cone density). Many seedlings established from rodent seed caches; 23% of the seedlings established in clumps from seeds cached by Dipodomys agilis, Chaetodipus californicus and Peromyscus maniculatus. Pinus coulteri seeds have low potential for dispersal by wind, but secondary dispersal by rodents moves seeds away from source trees and into neighboring chaparral. We discuss the potential importance of rodent seed caching to postfire demography of California and Mediterranean serotinous pines.
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References
Barbero M., Bonin G., Loisel R., Miglioretti and Quezel F. 1987. Incidence of exogenous factors on the regeneration of Pinus halepensis after fires. Ecologia Mediterranea 23: 51–56.
Benkman C.W. 1995. Wind dispersal capacity of pine seeds and the evolution of different seed dispersal modes in pines. Oikos 73: 221–224.
Borchert M. 1985. Serotiny and cone-habit variation in populations of Pinus coulteri (Pinaceae) in the southern Coast Ranges of California. Madroño 32: 29–48.
Borchert M., Lopez A., Bauer C. and Knowd T. 2000. Field guide to coastal sage scrub and chaparral series of Los Padres National Forest., 249 p. (unpublished).
Chappell C.B. and Agee J.K. 1996. Fire severity and tree seedling establishment in Abies magnifica forests, southern Cascades, Oregon. Ecological Applications 6: 628–640.
Daskalakou E.N. and Thanos C.A. 1996. Aleppo pine (Pinus halepensis) postfire regeneration: the role of canopy and soil seed banks. Int. J. Wildland Fire 6: 59–66.
Despland E. and Houle G. 1997. Aspect influences cone abundance within the crown of Pinus banksiana Lamb. trees at the limit of the species distribution in northern Quebec (Canada). Ecoscience 4: 521–525.
Engstrom F.B. and Mann D.H. 1991. Fire ecology of red pine (Pinus resinosa) in northern Vermont, USA. Canadian Journal of Forest Research 21: 882–889.
Greene D.F. and Johnson E.A. 1993. Seed mass and dispersal capacity in wind-dispersed diaspores. Oikos 67: 69–74.
Haim A. and Izhaki I. 1994. Changes in rodent community during recovery from fire: relevance to conservation. Biodiversity and Conservation 3: 573–585.
Haim A., Izhaki I. and Golan A. 1996. Rodent species diversity in pine forests recovering from fire. Israel Journal of Zoology 42: 353–359.
Herranz J.M., Martinez-Sanchez J.J., Marin A. and Ferrandis P. 1997. Postfire regeneraton of Pinus halepensis Miller in a semiarid area in Albacete province (southeastern Spain). Ecoscience 4: 86–90.
Izhaki I., Haim A. and Zohar O. 1993. Rodent populations recovering from fire in an east Mediterranean woodland. Water Science Technology 27: 539–545.
Jenkins S.H. and Peters R.A. 1992. Spatial patterns of food storage by Merriam's kangaroo rats. Behavioral Ecology 3: 60–65.
Jenkins S.J., Rothstein A. and Green W.C.H. 1995. Food hoarding by Merriam's kangaroo rats: a test of alternative hypotheses.Ecology 76: 2470–2481.
Jensen T.S. 1985. Seed-seed predator interactions of European beech, Fagus silvatica and forest rodents, Clethrionomys glareolus and Apodemus flavicollis. Oikos 44: 149–156.
Jensen T.S. and Nielsen O.F. 1986. Rodents as seed dispersers in a heath-oak wood succession. Oecologia 70: 214–221.
Johnson K.A., Morrison D.A. and Goldsack G. 1994. Post-fire flowering patterns in Blandfordia nobilis (Liliaceae). Australian Journal of Botany 42: 49–60.
Johnson M., Borchert M. and Vander Wall S.B. 2002. A comparative analysis of seed and cone characteristics and seed-dispersal strategies of three pines in the subsection Sabinianae. Plant Ecology (in press).
Keeley J. 1992. Temporal and spatial dispersal syndromes. In: Plant-Animal Interactions in Mediterranean-type Ecosystems.MEDECOS VI Proceedings. University of Athens, Athens, Greece, pp. 251–256.
Keeley J.E. and Zedler P.H. 1998. Evolution of life histories in Pinus.In: Richardson D.M. (ed.), Ecology and Biogeography of Pinus. Cambridge University Press, Cambridge, UK, pp. 219–249.
Kerley G.I.H. and Erasmus T. 1991. What do mice select for in seeds? Oecologia 86: 261–267.
Lawrence G.E. 1966. Ecology of vertebrate animals in relation to chaparral fire in the Sierra Nevada foothills. Ecology 47: 278–291.
Linhart Y.B. 1978. Maintenance of variation in cone morphology in California closed-cone pines: the roles of fire, squirrels and seed output. The Southwesten Naturalist 23: 29–40.
Martinez-Sanchez J.J., Ferrandis P., de las Heras J. and Herranz J.M. 1999. Effect of burnt wood removal on the natural regeneration of Pinus halepensis after fire in a pine forest in Tus valley (SE Spain). Forest Ecology and Management 123: 1–10.
McAuliffe J.R. 1990. Paloverdes, pocket mice, and bruchid beetles: interrelationships of seeds, dispersers, and seed predators. The Southwestern Naturalist 35: 329–337.
McMaster G.S. and Zedler P.H. 1981. Delayed seed dispersal in Pinus torreyana (Torrey pine). Oecologia 51: 62–66.
Moravec J. 1990. Regeneration of N.W. African Pinus halepensis forests following fire. Vegetatio 87: 29–36.
Ne'eman G., Lahav H. and Izhaki I. 1992. Spatial patterns one 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 on pine trees in Aleppo pine forest. Ecography 21: 535–542.
Ne'eman G., Fotheringham C.J. and Keeley J.E. 1999. Patch to landscape patterns in post fire recruitment of a serotinous conifer. Plant Ecology 145: 235–242.
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 Forest Ecosystems in the Mediterranean Basin. Backhuys, Leiden, pp. 303–319.
Ne'eman G. and Trabaud L. 2000. Ecology, Biogeography and Management of Pinus halepensis and P. brutia Forest Ecosystems in the Mediterranean Basin. Backhuys, Leiden.
Quinn R.D. 1990. Habitat Preferences and Distribution of Mammals in California chaparral. PSW-202. USDA Forest Service, Pacific Southwest Forest and Range Experiment Station, 11 p.
Quinn R.D. 1994. Animals, fire, and vertebrate herbivory in Californian chaparral and other Mediterranean ecosystems. In: Moreno J.M. and Oechel W.C. (eds), The Role of Fire in Mediterranean-type Ecosystems. Springer-Verlag, New York, pp. 46–78.
Saracino A. and Leone V. 1993. Ecological features and dynamics of seed dispersal after fire in a Pinus halepensis Miller forest. In: Trabaud L. and Prodon R. (eds), Fire in Mediterranean Ecosystems. Commission of the European Communities, Banyuls-Sur-Mer., pp. 151–159.
Simpson J.D. and Powell G.R. 1981. Some factors influencing cone production on young black spruce in New Brunswick. The Forestry Chronicle 57: 267–269.
Schwilk D.W. and Keeley J.E. 1998. Rodent populations after a large wildfire in California chaparral and coastal sage scrub.The Southwestern Naturalist 43: 480–483.
Shiller G., Ne'eman G. and Korol L. 1997. Post-fire vegetation dynamics in a native Pinus halepensis Mill, forest on Mt. Carmel, Israel. Israel Journal of Plant Sciences 45: 297–308.
Sullivan T.P. 1978. Lack of caching of direct-seeded Douglas-fir seeds by deer mice. Can. J. Zool. 56: 1214–1216.
ter Braak C.J.F. 1987. The analysis of vegetation-environment relationships by canonical correspondence analysis. Vegetatio 69: 69–77.
Thanos C.A., Marcou S., Christodoulakis D. and Yannitsaros A. 1989. Early post-fire regeneration in Pinus brutia forest ecosystems of Samos island (Greece). Acta Oecologica 10:79–94.
Thanos C.A. and Marcou S. 1991. Post-fire regeneration in Pinus brutia forest ecosystems of Samos island (Greece): 6 years after. Acta Oecologica 12: 633–642.
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.
Thanos C.A. and Doussi M.A. 2000. Post-fire regeneration of Pinus brutia forests. 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, Leiden, pp. 291–301.
Trabaud L., Michels C. and Grosman J. 1985. Recovery of burnt Pinus halepensis Mill. Forests. II. Pine reconstitution after wildfire. Forest Ecology and Management 13: 167–179.
Tsitoni T. 1997. Conditions determining natural regeneration after wildfires in the Pinus halepensis (Miller, 1768) forests of Kassandra Peninsula (North Greece). Forest Ecology and Management 92: 199–208.
Vander Wall S.B. and Balda R.P. 1981. Ecology and evolution of food-storage behavior of conifer-seed-caching corvids. Z. Tierpsychol. 56: 217–242.
Vander Wall S. 1992. The role of animals in dispersing a "winddispersed" pine. Ecology 73: 614–621.
Vander Wall S.B. 1993. Cache site selection by chipmunks (Tamias spp.) and its influence on the effectiveness of seed dispersal in Jeffrey pine (Pinus jeffreyi). Oecologia 9: 246–252.
Vander Wall S.B. 1994. Removal of wind-dispersed pine seeds by ground-foraging vertebrates. Oikos 69: 125–132.
Vander Wall S.B. 1997. Dispersal of singleleaf pinon pine (Pinus monophylla) by seed-caching rodents. Journal of Mammology 78: 181–191.
Vander Wall S.B. 2002. Secondary dispersal of Jeffrey pine seeds by rodent scatter hoarders: the roles of pilfering, recaching, and a variable environment. In: Levey D.J., Silva W.R. and Galetti M. (eds), Seed Dispersal and Frugivory: Ecology, Evolution and Conservation. CABI Publishing, Wallingford, Oxfordshire, UK, pp. 193–208.
Vogl R.J. 1968. Fire adaptations of some southern California plants. Proceedings of the Tall Timbers Fire Ecology Conference 8: 79–110.
Vogl R.J. 1973. Ecology of P. attenuata pine in the Santa Ana Mountains, California. Ecological Monographs 43: 125–143.
Vogl R.J., Armstrong W.O., White K.L. and Cole K.L. 1977. The closed cone pite and cypressus. In: Barbour M.G. and Major J. (eds), Terrestrial Vegetation of California.Wiley, New York, pp. 295–358.
Wada N. and Uemura S. 1994. Seed dispersal and predation by small rodents on the herbaceous understory plant Symplocarpus renifolius. American Midland Naturalist 132: 320–327.
Waldrop T.A. and Brose P.H. 1999. A comparison of fire intensity levels for stand replacement of table mountain pine (Pinus pungens Lamb.). Forest Ecology and Management 113: 155–166.
Warren R. and Fordham A.J. 1978. The fire pines. Arnoldia 38: 1–11.
Wirtz W.O. II 1977. Vertebrate post-fire succession. In: Mooney H.A. and Conrad C.E. (eds), Proceedings of the Symposium on The Environmental Consequences of Fire and Fuel Management in Mediterranean Ecosystems. Gen. Tech. Rep. WO-3. USDA Forest Service, pp. 46–47.
Wirtz W.O. II, Hoekman D., Muhm J.R. and Souza S.L. 1988. Postfire rodent succession following prescribed fire in southern California chaparral. In: Szaro R.C., Severson K.E. and Patton D.R. (eds), Proceedings of the Symposium on Management of Amphibians, Reptiles and Small Mammals in North America. Gen. Tech. Rep. RM-166. USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, pp. 333–339.
Wirtz W.O. II 1995. Responses of rodent populations to wildfire and prescribed fire in southern California chaparral. In: Keeley J.E. and Scott T. (eds), Brushfire in California: Ecology and Resource Management. International Association of Wildland Fire, Fairfield, Washington, USA, pp. 63–67.
Zedler P.H. 1977. Life history attributes and the fire cycle: a case study in chaparral dominated by Cupressus forbesii. Gen. Tech. Rep. WO-3. In: Mooney H.A. and Conrad C.E. (eds), Proceedings of the Symposium on the Environmental Consequences of Fire and Fuel Management in Mediterranean Ecosystems. USDA Forest Service, pp. 451–458.
Zedler P.H. 1981. Vegetation change in chaparral and desert communities in San Diego County, California. In: West D.C., Shugart H.H. and Botkin D. (eds), Forest Succession: Concepts and Applications. Spinger-Verlag, New York, pp. 406–430.
Zedler P.H. 1995. Fire frequency in southern California shrublands: biological effects and management options. In: Keeley J.E. and Scott T. (eds), Brushfires in California: Ecology and Resource Management. International Association of Wildland Fire, Fair-field, WA, USA, pp. 101–112.
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Borchert, M., Johnson, M., Schreiner, D.S. et al. Early postfire seed dispersal, seedling establishment and seedling mortality of Pinus coulteri (D. Don) in central coastal California, USA. Plant Ecology 168, 207–220 (2003). https://doi.org/10.1023/A:1024447811238
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DOI: https://doi.org/10.1023/A:1024447811238