Abstract
California chaparral and South African fynbos are fire-prone communities dominated by species exhibiting remarkable similarities in germination response. In both regions there are a substantial number of species with germination stimulated chemically by charred wood and smoke. This type of germination behaviour has arisen independently in distantly related families and is interpreted as convergent evolution. Heat-shock is also an important germination trigger that is widespread, although in both regions it is most common in the same families. Phylogeney may play an important role in the presence of this postfire germination cue in both regions, but a much more rigorous analysis is required to show that this trait represents a single unique event in each lineage. In both regions, germination response is not randomly distributed across growth forms and there are marked regional similiarities in the type of germination behaviour associated with certain growth forms. Geophytes largely lack refractory seeds, which require fire-type cues for germination, but the presence of fire-stimulated flowering of bulbs and corms may time recruitment to subsequent postfire years. Annuals that cue germination to postfire conditions are predominantly triggered by chemicals from smoke and/or charred wood.
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Anderson, I. C., Levine, J. S., Poth, M. A. & Riggan, P. J. 1988. Enhanced biogenic emissions of nitric oxide and nitrous oxide following surface biomass burning. J. Geophys. Res. 93: 3893- 3898.
Arroyo, M. T. K., Cavieres, L., Marticorena, C. & Munoz-Schick, M. 1994. Convergence in the mediterranean floras in central Chile and California: insights from comparative biogeography. Pp. 43-88. In: M. T. K. Arroyo, P. H. Zedler, & M. D. Fox (eds), Ecology and biogeography of mediterranean ecosystems in Chile, California, and Australia, Springer-Verlag, New York.
Atwater, B. R. 1980. Germination, dormancy and morphology of the seeds of herbaceous ornamental plants. Seed Sci. Techn. 8: 523-573.
Baldwin, I. T., Staszak-Kozinski, L. & Davidson, R. 1994. Up in smoke. 1. Smoke-derived germination cues for postfire annual, Nicotiana attentuataTorr ex Watson. J. Chem. Ecol. 20: 2345-2372.
Ballard, L. A. T. 1973. Physical barriers to germination. Seed Sci. Techn. 1: 285-303.
Barbour, M. G. & Minnich, R. A. 1990. The myth of chaparral convergence. Israel J. Bot. 39: 435-463.
Baxter, B. J. M., Granger, J. E., & van Staden, J. 1995. Plant-derived smoke and seed germination: is all smoke good smoke? That is the burning question. S. Afr. J. Bot. 61: 275-278.
Bell, D. T., Plummer, J. A. & Taylor, S. K. 1993. Seed germination ecology in southwestern Western Australia. Bot. Rev. 59: 24-73.
Bond, W. J. 1985. Canopy-stored seed reserves (serotiny) in Cape Proteaceae. S. Afr. J. Bot. 39: 435-463.
Bond, W. J., Vlok, J. & Viviers, M. 1984. Variation in seedling recruitment of Cape Proteaceae after fire. J. Ecol. 72: 209-221.
Bond, W. J., Roux, D. L. & Erntzen, R. 1990. Fire intensity and regeneration of myrmecochorous Proteaceae. S. Afr. J. Bot. 56: 326-330.
Bond, P.& Goldblatt, P. 1984. Plants of the Cape flora. A descriptive catalogue. J. S. Afr. Bot. Suppl. No. 13.
Brits, G. J. 1987. Germination depth vs. temperature requirements in naturally dispersed seeds of Leucospermum cordifoliumand L. cuneiforme(Proteaceae). S. Afr. J. Bot. 53: 119-124.
Brits, G. J., Calitz, F. J., Brown, N. A. C. & Manning, J. C. 1993. Desiccation as the active principle in heat-stimulated seed germination of LeucospermumR. Br. (Proteaceae) in fynbos. New Phytol. 125: 397-403.
Brown, N. A. C. 1993a. Promotion of germination of fynbos seeds by plant-derived smoke. New Phytol. 123: 575-584.
Brown, N. A. C. 1993b. Seed germination in the fynbos fire ephemeral, Syncarpha vestita(L.) B. Nord. is promoted by smoke, aqueous extracts of smoke and charred wood derived from burning the ericoid-leaved shrub, Passerina vulgarisThoday. Inter. J. Wildland Fire 3: 203-206.
Brown, N. A. C., Jamieson, H. & Botha, P. A. 1995. Stimulation of seed germination in South African species of Restionaceae by plant-derived smoke. Pl. Grow. Regul. 15: 93-100.
Brown, N. A. C., Kotze, G. & Botha, P. A. 1993. The promotion of seed germination of Cape Ericaspecies by plant-derived smoke. Seed Sci. Techn. 21: 573-580.
Brown, P. J., Manders, P. T., Bands, D. P., Kruger, F. J. & Andrag, R. H. 1991. Prescribed burning as a conservation management practice: a case history from the Cedarberg mountins, Cape Province, South Africa. Biol. Conserv. 56: 133-150.
Bullock, S. H. 1976. Comparison of the distribution of seed and parent-plant populations. Southw. Nat. 21: 383-389.
Cody, M. L. & Mooney, H. A. 1978. Convergence versus nonconvergence in Mediterranean-climate ecosystems. Ann. Rev. Ecol. Syst. 9: 265-321.
Corner, E. J. H. 1976. The seeds of dicotyledons. Volume 1. Cambridge University Press, New York.
Cowling, R. M. & Campbell, B. M. 1980. Convergence in vegetation structure in the Mediterranean communities of California, Chile, and South Africa. Vegetatio 43: 191-197.
Dafni, A., Cohen, D. & Noy-Meir, I. 1981. Life-cycle variation in geophytes. Ann. Mo. Bot. Gard. 68: 652-660.
de Lange, J. H. & Boucher, C. 1990. Autecological studies on Audouinia capitata(Bruniaceae). I. Plant-derived smoke as a seed germination cue. S. Afr. J. Bot. 56: 700-703.
de Lange, J. H. & Boucher, C. 1993. Autecological studies on Audouinia capitata(Bruniaceae). 8. Role of fire in regeneration S. Afr. J. Bot. 59: 188-202.
di Castri, F. & Mooney, H. A. (eds.). 1973. Mediterranean ecosystems: origin and structure. Springer-Verlag, New York.
Dixon, K. W., Roche, S. & Pate, J. S. 1995. The promotive effect of smoke derived from burnt native vegetation on seed germination of Western Australian plants. Oecologia 101: 185-192.
Donoghue, M. J. 1989. Phylogenies and the analysis of evolutionary sequences, with examples from seed plants. Evolution 43: 1137-1156.
Eggleton, P. & Vane-Wright, R. I. (eds). 1994. Phylogenetics and ecology. Academic Press, New York.
Egley, G. H. 1989. Water-impermeable seed coverings as barriers to germination. Pp. 207-223. In: R. B. Taylorson (ed.), Recent advances in the development and germination of seeds, Plenum Press, New York.
Gill, A. M. 1981. Adaptive responses of Australian vascular plant species to fires. Pp. 243-272. In: A. M. Gill, R. H. Groves, & I. R. Noble (eds), Fire and the Australian biota, Australian Academy of Science, Canberra, Australian Capital Territory.
Hickman, J. C. (ed.). 1993. The Jepson manual. Higher plants of California. Univ. Calif. Press, Los Angeles.
J¨ager, A. K., Rabe, T., van Staden, J. 1996. Food-flavouring smoke extracts promote seed germination. S. Afr. J. Bot. 62: 282-284.
Jefferey, D. J., Holmes, P. M. & Rebelo, A. G. 1988. Effects of dry heat on seed germination in selected indigenous and alien legume species in South Africa. S. Afr. J. Bot. 54: 28-34.
Jones, C. S. & Schlesinger, W. H. 1980. Emmenanthe penduli flora(Hydrophyllaceae): further consideration of germination response. Madroño 27: 122-125.
Keeley, J. E. 1982. Distribution of lightning and man-caused wild-fires in California. Pp. 431-437. In: C. E. Conrad & W. C. Oechel (eds), Proceedings of the symposium on dynamics and management of mediterranean-type ecosystems, USDA For. Serv., Pac. Southw. For. Range Exp. Stat., Gen. Techn. Rep. PSW-58.
Keeley, J. E. 1986. Resilience of mediterranean shrub communities to fire. Pp. 95-112. In: B. Dell, A. J. M. Hopkins, & B. B. Lamont (eds), Resilience in editerranean-type ecosystems, Dr. W. Junk, Dordrecht, The Netherlands.
Keeley, J. E. 1987. Role of fire in seed germination of woody taxa in California chaparral. Ecology 68: 434-443.
Keeley, J. E. 1991. Seed germination and life history syndromes in the California chaparral. Bot. Rev. 57: 81-116.
Keeley, J. E. 1992. A Californian's view of fynbos. Pp. 372-388. In: R. M. Cowling (ed.), The ecology of fynbos, Oxford Univ. Press, Cape Town.
Keeley, J. E. 1993. Smoke-induced flowering in the fire-lily Cyrtanthus ventricosus. S. Afr. J. Bot. 59: 638.
Keeley, J. E. 1994. Seed-germination patterns in fire-prone mediterranean-climate regions. Pp. 239-273. In: T. K. Arroyo, P. H. Zedler, & M. D. Fox (eds), Ecology and biogeography of mediterranean ecosystems in Chile, California, and Australia, Springer-Verlag, New York.
Keeley, J. E. & Fotheringham, C. J. 1997a. Trace gas emissions in smoke-induced seed germination. Science 276: 1248-1251.
Keeley, J. E. & Fotheringham, C. J. 1997b. Smoke-induced seed germination in Californian chaparral. Ecology (in submission).
Keeley, J. E. & Keeley, S. C. 1984. Postfire recovery of California coastal sage scrub. Am. Midl. Nat. 111: 105-117.
Keeley, J. E., Morton, B. A., Pedrosa, A., & Trotter, P. 1985. Role of allelopathy, heat, and charred wood in the germination of chaparral herbs and suffrutescents. J. Ecol. 73: 445-458.
Keeley, J. E. & Zedler, P. H. Evolution of life histories in pines. In: D. Richardson & R. Cowling (eds), Ecology and biogeography of pines. Cambridge Univ. Press, Cambridge, U. K. (in press).
Keeley, S. C. & Pizzorno, M. 1986. Charred wood stimulated germination of two fire-following herbs of the California chaparral and the role of hemicellulose. Amer. J. Bot. 73: 1289-1297.
Kilian, D. & Cowling, R. M. 1992. Comparative seed biology and co-existence of two fynbos shrub species. J. Veg. Sci. 3: 637-646.
Kruger, F. J. 1983. Plant community diversity and dynamics in relation to fire. Pp. 446-472. In: F. J. Kruger, D. T. Mitchell, & J. U. M. Jarvis (eds), Mediterranean-type ecosystems. The role of nutrients, Springer-Verlag, New York.
Le Maitre, D. C. & Brown, P. J. 1992. Life cycles and fire-stimulated flowering in geophytes. Pp. 145-160. In: B. W. van Wilgen, D. M. Richardson, F. J. Kruger, & H. J. van Hensbergen (eds), Fire in South African mountain fynbos, Springer-Verlag, Berlin.
Le Maitre, D. C. & Midgley, J. J. 1992. Plant reproductive ecology. Pp. 135-174. In: R. M. Cowling (ed.), The ecology of fynbos, Oxford Univ. Press, Cape Town.
Losos, J. B. 1996. Phylogenetic perspectives on community ecology. Ecology 77: 1344-1354.
Martin, A. C. 1946. The comparative internal morphology of seeds. Amer. Midl. Nat. 36: 513-660.
Mazer, S. J. 1989. Ecological, taxonomic, and life history correlates of seed mass among Indiana dune angiosperms. Ecol. Monogr. 59: 153-175.
Mooney, H. A. (ed.). 1977. Convergent evolution of Chile and California Mediterranean climate ecosystems. Dowden, Hutchinson and Ross, Stroudsburg, Pennsylvania.
Mooney, H. A. & Conrad, C. E. (eds). 1977. Proceedings of the symposium on environmental consequences of fire and fuel management in Mediterranean ecosystems. USDA For. Serv., Gen. Techn. Rep. WO-3.
Musil, C. F. & de Witt, D. M. 1991. Heat-stimulated germination in two Restionaceae species. S. Afr. J. Bot. 57: 175-176.
Pagel, M. D. 1994. The adaptationist wager. Pp. 29-51. In: P. Eggleton and R. I. Vane-Wright (eds), Phylogenetics and ecology, Academic Press, San Diego.
Peet, R. K. 1978. Ecosystem convergence. Amer. Nat. 112: 441-444.
Pierce, S. M., Esler, K. & Cowling, R. M. 1995. Smoke-induced germination of succulents (Mesembryanthemaceae) from fire-prone and fire-free habitats in South Africa. Oecologia 102: 520-522.
Pierce, S. M. & Moll, E. J. 1994. Germination ecology of six shrubs in fire-prone Cape fynbos. Vegetatio 110: 25-41.
Quinlivan, B. J. 1971. Seed coat impermeability in legumes. J. Aus. Inst. Agr. Sci. 37: 283-295.
Rundel, P. W. 1996. Monocotyledonous geophytes in the California flora. Madroño 43: 355-368.
Shmida, A. & Whittaker, R. H. 1984. Convergence and nonconvergence of Mediterranean type communities in the Old and the New World. Pp. 5-11. In: N. S. Margaris, M. Arianoustou-Faraggitaki, & W. C. Oechel (eds), Being alive on land, Dr. W. Junk, The Hague, The Netherlands.
Stone, E. C. 1951. The stimulative effect of fire on the flowering of the golden brodiaea (Brodiaeae ixiodesWats. var. lugensJeps.). Ecology 32: 534-537.
Trabaud, L. & Prodon, R. 1993. Fire in mediterranean ecosystems. Commission of the European Communities, Banyuls-Sur-Mer, France. 441 pp.
van Staden, J., Drewes, F. E. & Jager, A. K. 1995. The search for germination stimulants in plant-derived smoke extracts. S. Afr. J. Bot. 61: 260-263.
van Wilgen, B. W., Richardson, D. M., Kruger, F. J. & van Hensbergen, H. J. (eds.). 1992. Fire in South African mountain fynbos. Ecosystem community and species response at Swartboskloof. Springer-Verlag, Berlin.
Wicklow, D. T. 1977. Germination response in Emmenanthe penduliflora(Hydrophyllaceae). Ecology 58: 201-205.
Young, J. A. & Young, C. G. (eds). 1992. [Revision of] Seeds of woody plants in North America. Dioscorides Press, Portland,Oregon.
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Keeley, J.E., Bond, W.J. Convergent seed germination in South African fynbos and Californian chaparral. Plant Ecology 133, 153–167 (1997). https://doi.org/10.1023/A:1009748603202
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DOI: https://doi.org/10.1023/A:1009748603202