Evolutionary Ecology

, Volume 17, Issue 3, pp 277–292

Are seed set and speciation rates always low among species that resprout after fire, and why?



A general dichotomy in response to catching alight during fire is for perennial plants to die (nonsprouters, N) or to regrow via dormant buds (resprouters, R). Contrasting effects on other life-history traits, especially those relating to sexual reproduction, can be expected. We hypothesized that fecundity should be lower in R. Our meta-analysis of 33 case studies of co-occurring generic pairs of N and R showed that viable seed and fruit set relative to inflorescence, flower and ovule production were lower for R than N in 30 cases. Three mechanisms have been invoked to explain these trends: resource competition between vegetative and reproductive growth, an outbreeding (R)/inbreeding (N) dichotomy, and higher genetic load among R due to their accumulation of deleterious somatic mutations over many fire cycles. Though poorly researched, we conclude that the high genetic load of R in association with strong self-incompatibility (xenogamy) can best explain the dichotomy in lifeform-fecundity as well as the exceptions. Over the long-term, we suggest that marked xenogamy-longevity in association with frequent axillary branching (induced by recurrent fire, herbivory and drought) via copious stored buds within R might favour expression among their genets and ramets of beneficial somatic mutations present in their meristematic tissues. These conditions would favour ecotypic differentiation and speciation among R that has contributed, along with N, to the exceptional species richness of several fire-prone mediterranean regions.

breeding system disturbance fire response genetic load resource allocation speciation 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Anderson, G.J. and Hill, J.D. (2002) Many to flower, few to fruit: the reproductive biology of Hamamelis virginiana (Hamamelidaceae). Am. J. Bot. 89, 67-78.Google Scholar
  2. Auld, T.D. (1987) Population dynamics of thr shrub Acacia suaveolens (Sm.) Willd.: survivorship throughout the life cycle, a synthesis. Aust. J. Ecol. 12, 139-151.Google Scholar
  3. Barker, W.R., Barker, R.M. and Haegi, L. (1999) Introduction to Hakea. Flora Aust. 17B, 1-30.Google Scholar
  4. Barrett, G.J. and Lamont, B.B. (1986) Conservation status and reproductive biology of two rare Banksia species. Report to World Wildlife Fund Australia. Environmental Biology, Curtin University, Perth. 135 pp.Google Scholar
  5. Barrett, L. and Lamont, B.B. (2002) Population attributes of resprouting and nonsprouting subspecies of Hakea petiolaris. Report to Conservation and Land Management. Environmental Biology, Curtin University, Perth. 45 pp.Google Scholar
  6. Bell, D.T. (2001) Ecological response syndromes in the flora of southwestern Australia: fire resprouters versus reseeders. Bot. Rev. 67, 417-440.Google Scholar
  7. Bell, T.L. and Ojeda, F. (1999) Underground starch storage in Erica species of the Cape Floristic Region-differences between seeders and resprouters. New Phytol. 144, 143-152.Google Scholar
  8. Bell, T.L. and Pate, J.S. (1993) Morphometric differentiation in the south-western Australian restiad Lyginea barbata (Restionaceae). Aust. J. Bot. 41, 91-104.Google Scholar
  9. Bond, W.J. and Midgley, J.J. (2001) Ecology of sprouting in woody plants: the persistence niche. Trends Ecol. Evol. 16, 45-51.Google Scholar
  10. Bond, W.J. and Midgley, J.J. (2003) The evolutionary ecology of sprouting in woody plants. Int. J. Plant Se. 164 (in press).Google Scholar
  11. Bowen, B.J. and Pate, J.S. (1993) The significance of root starch in post-fire shoot recovery of the resprouter Stirlingia latifolia R. Br. (Proteaceae). Ann. Bot. 29, 521-531.Google Scholar
  12. Bradstock, R.A. and O'connell, M.A. (1988) Demography of woody plants in relation to fire: Banksia ericifolia L.f. and Petrophile pulchella (Shrad.) R.Br. Aust. J. Ecol. 13, 505-518.Google Scholar
  13. Burgman, M.A. and Lamont, B.B. (1992) A stochastic model for the viability of Banksia cuneata populations: environmental, demographic and genetic effects. J. Appl. Ecol. 29, 719-727.Google Scholar
  14. Burrows, G.F. (2002) Epicormic strand structure in Angophora, Eucalyptus and Lophostemon (Myrtaceae)-implications for fire resistance and recovery. New Phytol. 153, 111-131.Google Scholar
  15. Carpenter, F.L. and Recher, H.F. (1979) Pollination, reproduction, and fire. Am. Nat. 113, 871-879.Google Scholar
  16. Clarke, P.J. (2002) Habitat insularity and fire response traits: evidence from a sclerophyll archipelago. Oecologia 132, 582-591.Google Scholar
  17. Clarke, P.J. and Knox, K.J.E. (2002) Post-fire response of shrubs in the tablelands of eastern Australia: do existing models explain habitat differences? Aust. J. Bot. 50, 53-62.Google Scholar
  18. Copland, B.J. and Whelan, R.J. (1989) Seasonal variation in flowering intensity and pollination limitation of fruit set in four co-occurring Banksia species. J. Ecol. 77, 509-523.Google Scholar
  19. Cowling, R.M. and Lamont, B.B. (1987) Post-fire recruitment of four co-occurring Banksia species. J. Appl. Ecol. 24, 645-658.Google Scholar
  20. Cowling, R.M. and Lombard, A.T. (2002) Heterogeneity, speciation/extinction history and climate: explaining regional plant diversity in the Cape Floristic Region. Div. Distr. 8, 163-179.Google Scholar
  21. Cowling, R.M., Lamont, B.B. and Pierce, S.M. (1987) Seed bank dynamics of four co-occurring Banksia species. J. Ecol. 75, 289-302.Google Scholar
  22. Cowling, R.M., Rundel, P.W., Lamont, B.B., Arroyo, M.K. and Adrianoutsou, M. (1996) Plant diversity in Mediterranean-climate regions. Trends Ecol. Evol. 11, 362-366.Google Scholar
  23. Cruz, A. and Moreno, J.M. (2001) No allocation trade-offs between flowering and sprouting in the lignotuberous, Mediterranean shrub Erica australis. Acta Oecol. 22, 121-127.Google Scholar
  24. Davis, S.D., Ewers, F.W., Wood, J., Reeves, J.J. and Kolb, K.J. (1999) Differential susceptibility to xylem cavitation among three pairs of Ceanothus species in the Transverse Mountain Ranges of southern California. Ecoscience 6, 180-186.Google Scholar
  25. Deacon, H.J., Jury, M.R. and Ellis, F. (1992) Selective regime and time. In R.M. Cowling (ed.) The Ecology of Fynbos: Nutrients, Fire and Ecology. Oxford, Cape Town, pp. 6-22.Google Scholar
  26. Drechsler, M., Lamont, B.B., Burgman, M.A., Akçakaya, H.R., Witkowski, E.T.F. and Supriyadi (1999) Modelling the persistence of an apparently immortal Banksia species after fire and land clearing. Biol. Conserv. 88, 249-259.Google Scholar
  27. England, P.R., Beynon, F., Ayre, D.J. and Whelan, R.J. (2001) A molecular genetic assessment of mating-system variation in a naturally bird-pollinated shrub: contributions from birds and introduced honeybees. Conserv. Biol. 15, 1645-1655.Google Scholar
  28. Enright, N.J. and Goldblum, D. (1999) Demography of a non-sprouting and resprouting Hakea species (Proteaceae) in fire-prone Eucalyptus woodlands of southeastern Australia in relation to stand age, drought and disease. Plant Ecol. 144, 71-82.Google Scholar
  29. Enright, N.J. and Lamont, B.B. (1989) Seed banks, fire season, safe sites and seedling recruitment in five co-occurring Banksia species. J. Ecol. 77, 1111-1122.Google Scholar
  30. Enright, N.J. and Lamont, B.B. (1992) Recruitment variability in the resprouting shrub Banksia attenuata and non-sprouting congeners in the northern sandplain heaths of south-western Australia. Acta Oecol. 13, 727-741.Google Scholar
  31. 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 nonsprouting shrubs. J. Ecol. 86, 946-959.Google Scholar
  32. Flannery, T.F. (1994) The Future Eaters. Reed, Chaswood. 423 pp.Google Scholar
  33. Frazer, J.M. and Davis, S.D. (1988) differential survival of chaparral seedlings during the first summer drought after wildfire. Oecologia 90, 215-221.Google Scholar
  34. Fulton, R.E. and Carpenter, F.L. (1979) Pollination, reproduction and fire in California Arctostaphylos. Oecologia 38, 147-157.Google Scholar
  35. Gill, D.E. (1986) Individual plants as genetic mosaics: ecological organisms versus evolutionary individuals. In M.J. Crawley (ed.) Plant Ecology. Blackwell, Oxford, pp. 321-343.Google Scholar
  36. Gill, D.E. and Halverson, T.G. (1984) Fitness variation among branches within trees. In B. Shorrocks (ed.) Evolutionary Ecology. Blackwell, Oxford, pp. 105-116.Google Scholar
  37. Groom, P.K. and Lamont, B.B. (1996) Reproductive ecology of non-sprouting and re-sprouting Hakea species (Proteaceae) in southwestern Australia. In S.D. Hopper, M. Harvey, J. Chappill and A.S. George (eds) Gondwanan Heritage. Surrey Beatty, Chipping Norton, pp. 239-248.Google Scholar
  38. Groom, P.K., Lamont, B.B. and Wright, I. (2001) Lottery (stochastic) and non-lottery (biological) processes explain recruitment patterns among eight congeneric shrubs in southwestern Australia. J. Medit. Ecol. 2, 1-14.Google Scholar
  39. Haegi, L. and Barker, W.R. (1985) Taxonomy of the South Australian species allied to Hakea ulicina R. Br. (Proteaceae). J. Adelaide Bot. Gard. 7, 249-271.Google Scholar
  40. Hansen, A., Pate, J.S. and Hansen, A.P. (1991) Growth and reproductive performance of a seeder and a resprouter species of Bossiaea as a function of plant age after fire. Ann. Bot. 67, 497-509.Google Scholar
  41. Head, M.J. and Lacey, C.J. (1988) Radiocarbon age determinations from lignotubers. Aust. J. Bot. 36, 93-100.Google Scholar
  42. Heiken, J.L. (1958) Aberrant types in the potato. Acta Agric. Scand. 8, 319-358.Google Scholar
  43. Hopper, S.D., Harvey, M.S., Chappill, J.A., Main, A.R. and York Main, B. (1996) The Western Australian biota as gonwanan heritage-a review. In S.D. Hopper, M. Harvey, J. Chappill and A.S. George, (eds) Gondwanan Heritage: Evolution and Conservation of the Western Australian Biota. Surrey Beatty, Chipping Norton, pp. 1-46.Google Scholar
  44. James, S. (1984) Lignotubers and burls-their structure, function and ecological significance in mediterranean ecosystems. Bot. Rev. 50, 225-266.Google Scholar
  45. Keeley, J.E. (1977) Seed production, seed populations in soil, and seedling production after fire for two congeneric pairs of sprouting and non-sprouting chaparral shrubs. Ecology 58, 820-829.Google Scholar
  46. Keeley, J.E. (1986) Resilience of Mediterranean shrub communities to fires. In B. Dell, A.J. Hopkins and B.B. Lamont (eds) Resilience in Mediterranean-type Ecosystems. W. Junk, The Hague, pp. 95-112.Google Scholar
  47. Klekowski, E.J. (1988) Mutation, Developmental Selection, and Plant Evolution. Columbia University Press, New York.Google Scholar
  48. Klekowski, E.J. and Godfrey, P.J. (1989) Ageing and mutation in plants. Nature 340, 389-391.Google Scholar
  49. Lamont, B. (1985) The comparative reproductive biology of three Leucospermum species (Proteaceae) in relation to fire responses and breeding system. Aust. J. Bot. 33, 139-145.Google Scholar
  50. Lamont, B.B. (1988) Sexual versus vegetative reproduction in Banksia elegans. Bot. Gaz. 149, 370-375.Google Scholar
  51. Lamont, B.B. and Barker, M.J. (1988) Seed bank dynamics of a serotinous, fire-sensitive Banksia species. Aust. J. Bot. 36, 193-203.Google Scholar
  52. Lamont, B.B. and Barrett, G.J. (1988) Constraints on seed production and storage in a rootsuckering Banksia. J. Ecol. 76, 1069-1082.Google Scholar
  53. Lamont, B.B. and Bergl, S.M. (1991) Water relations, shoot and root architecture, and phenology of three co-occurring Banksia species: no evidence for niche differentiation in the pattern of water use. Oikos 60, 291-298.Google Scholar
  54. Lamont, B.B. and Downes, S. (1979) The age, flowering and fire history of the grass trees Xanthorrhoea preissii and Kingia australis. J. Appl. Ecol. 16, 893-899.Google Scholar
  55. Lamont, B.B. and Markey, A. (1995) Biogeography of fire-killed and resprouting Banksia species in southwestern Australia. Aust. J. Bot. 48, 283-303.Google Scholar
  56. Lamont, B.B. and Markey, A.S. (1996) Does breeding system depend on fire response among banksias? In Proceedings of International Symposium on the Biology of Proteaceae. University of Melbourne. p. 32.Google Scholar
  57. Lamont, B.B. and van Leeuwen, S.J. (1988) Seed production and mortality in a rare Banksia species. J. Appl. Ecol. 25, 551-559.Google Scholar
  58. Lamont, B.B., Connell, S. and Bergl, S.M. (1991) Population and seed bank dynamics of Banksia cuneata: the role of time, fire and moisture. Bot. Gaz. 152, 114-122.Google Scholar
  59. Lamont, B.B., Klinkhamer, P.G. and Witkowski, E.T.F. (1993) Population fragmentation may reduce fertility to zero in Banksia goodii-a demonstration of the Allee effect. Oecologia 94, 446-450.Google Scholar
  60. Lamont, B.B., Olesen, J.M. and Briffa, P. (1998) Seed production, pollinator attractants and breeding systems of two species pairs in relation to fire response-are there reproductive syndromes? Aust. J. Bot. 46, 377-385.Google Scholar
  61. Lamont, B.B., Rees, R., Witkowski, E.T.F. and Whitten, V. (1994a) Comparative size, fecundity and ecophysiology of roadside plants of Banksia hookeriana. J. Appl. Ecol. 31, 137-144.Google Scholar
  62. Lamont, B.B., Whitten, V., Witkowski, E.T.F., Rees, R. and Enright, N.J. (1994b) Regional and local (road verge) effects on size and fecundity in Banksia menziesii. Aust. J. Ecol. 19, 197-205.Google Scholar
  63. Le Maitre, D.C. and Midgley, J.J. (1992) Plant reproductive biology. In R.M. Cowling (ed.) The Ecology of Fynbos: Nnutrients, Fire and Ecology. Oxford, Cape Town, pp. 135-174.Google Scholar
  64. Lee, H.M. (1984) The biology of Hakea ulicina R.Br and H. repullulans H.M. Lee (Proteaceae). Aust. J. Bot. 32, 679-699.Google Scholar
  65. Low, A.B. and Lamont, B.B. (1990) Aerial and below-ground phytomass of Banksia scrub-heath at Eneabba, Western Australia. Aust. J. Bot. 38, 351-359.Google Scholar
  66. Marcotrigiano, M. (2000) Herbivory could unlock mutations sequestered in stratified shoot apices of genetic mosaics. Am. J. Bot. 87, 355-361.Google Scholar
  67. Meney, K.A. and Pate, J.S. (1999) Australian Rushes. University of Western Australia Press, Nedlands, 486 pp.Google Scholar
  68. Meney, K.A., Dixon, K.W. and Pate, J.S. (1997) Reproductive potential of obligate seeded and resprouter herbaceous perennial monocots (Restionaceae, Anarthriaceae, Ecdeiocoleaceae) from south-western Australia. Aust. J. Bot. 45, 771-782.Google Scholar
  69. Moreno, J.M. and Oechel, W.C. (1992) Factors controlling post-fire seedling establishment in southern California chaparral. Oecologia 90, 50-60.Google Scholar
  70. Morgan, M.T. (2001) Consequences of life history for inbreeding depression and mating system evolution in plants. Proc. Roy. Soc. Lond. 268, 1824-1847.Google Scholar
  71. Owen-Smith, N. and Danckwerts, J.E. (1997) Herbivory. In R.M. Cowling, D.M. Richardson and S.M. Pierce (eds) Vegetation of Southern Africa. Cambridge University, Cambridge, pp. 397-420.Google Scholar
  72. Pate, J.S., Casson, N.E., Rullo, J. and Kuo, J. (1985) Biology of the fire ephemeral of the sandplains of the kwongan of south-western Australia. Aust. J. Plant Physiol. 12, 641-655.Google Scholar
  73. Pate, J.S., Meney, K.A. and Dixon, K.W. (1991) Contrasting growth and morphological characteristics of fire-sensitive (obligate seeder) and fire-resistant (resprouter) species of Restionaceae (S. Hemisphere restiads) from south-western Western Australia. Aust. J. Bot. 39, 505-525.Google Scholar
  74. Peck, J.R. (1994) A ruby in the rubbish: beneficial mutations, deleterious mutations and the evolution of sex. Genetics 137, 597-606.Google Scholar
  75. Radtke, K.A., Arndt, A. and Wakimoto, R. (1982) Fire history of the Santa Monica Mountains. In C. Conrad and W. Oechel (eds) Proceedings of the Symposium on Dynamics and Management of Mediterranean-type Ecosystems. USDA Forest Service, Gen. Tech. Rep PSW-58, Berkley, pp. 438-443.Google Scholar
  76. Ramsey, M. and Vaughton, G. (1991) Self-incompatibility, protandry, pollen production and pollen longevity in Banksia menziesii. Aust. J. Bot. 39, 497-504.Google Scholar
  77. Rice, W.R. (1989) Analysing tables of statistical tests. Evolution 43, 223-225.Google Scholar
  78. Richards, M.B. and Lamont, B.B. (1996) Post-fire mortality and water relations of three congeneric shrub species under extreme water stress-a trade-o. with fecundity? Oecologia 102, 53-60.Google Scholar
  79. Sampson, J.F., Coates, D.J. and van Leeuwen, S.J. (1996) Mating system variation in animalpollinated rare and endangered plant populations in Western Australia. In S.D. Hopper, M. Harvey, J. Chappill and A.S. George (eds) Gondwanan Heritage. Surrey Beatty, Chipping Norton, pp. 187-195.Google Scholar
  80. Scott, J.K. (1980) Estimation of the outcrossing rate of Banksia attenuata R.Br. and Banksia menziesii R.Br. (Proteaceae). Aust. J. Bot. 28, 53-59.Google Scholar
  81. Silander, J.A. (1985) Microevolution in clonal plants. In J.B. Jackson, L.W. Buss and R. Look (eds.) The population Biology and Evolution of Clonal Organisms. Yale University Press, Yale, pp. 107-152.Google Scholar
  82. Specht, R.L., Rayson, P. and Jackman, M.E. (1958) Dark Island Heath (Ninety-Mile Plain, South Australia). VI. Pyric succession: changes in composition, coverage, dry weight, and mineral status. Aust. J. Bot. 6, 59-88.Google Scholar
  83. Stock, W.D., Pate, J.S., Kuo, J. and Hansen, A.P. (1989) Resource control of seed set in Banksia laricina C. Gardner (Proteaceae). Funct. Ecol. 3, 453-460.Google Scholar
  84. Thiele, K. and Ladiges, P.Y. (1996) A cladistic analysis of Banksia (Proteaceae). Aust. Syst. Bot. 9, 661-733.Google Scholar
  85. Vaughton, G. and Ramsey, M. (1998) Sources and consequences of seed variation in Banksia marginata (Proteaceae). J. Ecol. 86, 563-573.Google Scholar
  86. Wann, J.M. and Bell, D.T. (1997) Dietery preferences of the black-gloved wallaby (Macropus irma) and the western grey kangaroo (M. fuliginosus) in Whiteman Park, Perth, Western Australia. J. Roy. Soc. West. Aust. 80, 55-62.Google Scholar
  87. Ward, D.J., Lamont, B.B. and Burrows, C.L. (2001) Grasstrees reveal contrasting fire regimes in eucalypt forest before and after European settlement of southwestern Australia. For. Ecol. Manage. 150, 323-329.Google Scholar
  88. Wells, P.V. (1969) The relation between mode of reproduction and extent of speciation in woody genera of the California chaparral. Evolution 23, 264-267.Google Scholar
  89. Westwood, J. and Lamont, B.B. (1994) Reproductive ecology of co-occurring Hakea species differing in leaf morphology and regenerative mode. Report to Department of Conservation and Land Management, Environmental Biology. Curtin University, Perth, 46 pp.Google Scholar
  90. Whitham, T.G. and Slobodchiko., C.N. (1981) Evolution by individuals, plant-herbivore interactions, and mosaics of genetic variability: the adaptive significance of somatic mutations in plants. Oecologia 49, 287-292.Google Scholar
  91. Wiens, D., Calvin, C.L., Wilson, C.A., Davern, C.I., Frank, D. and Seavey, S.R. (1987) Reproductive success, spontaneous embryo abortion, and genetic load in flowering plants. Oecologia 71, 501-509.Google Scholar
  92. Williams, I.J.M. (1972) A revision of the genus Leucadendron (Proteaceae). Contr. Bolus Herb. 3, 1-425.Google Scholar
  93. Wisheu, I.C., Rosenzweig, M.L., Olsvig-whittaker, L. and Shmida, A. (2000) What makes nutrientpoor mediterranean heathlands so rich in plant diversity? Evol. Ecol. Res. 2, 935-955.Google Scholar
  94. Witkowski, E.T.F. and Lamont, B.B. (1996) Disproportionate allocation of mineral nutrients and carbon between vegetative and reproductive structures in Banksia hookeriana. Oecologia 105, 38-42.Google Scholar
  95. Witkowski, E.T.F., Lamont, B.B. and Connell, S.J. (1991) Seed bank dynamics of three co-occurring banksias in south coastal Western Australia: the role of plant age, cockatoos, senescence and interfire establishment. Aust. J. Bot. 39, 385-397.Google Scholar
  96. Wooller, S.J. and Wooller, R.D. (2001) Seed set in two sympatric banksias, Banksia attenuata and B. baxteri. Aust. J. Bot. 49, 597-602.Google Scholar
  97. Wooller, S.J., Wooller, R.D. and Brown, K.L. (2002) Regeneration by three species of Banksia on the south coast of Western Australia in relation to fire interval. Aust. J. Bot. 50, 311-317.Google Scholar

Copyright information

© Kluwer Academic Publishers 2003

Authors and Affiliations

  1. 1.Environmental BiologyCurtin UniversityPerthAustralia
  2. 2.Rancho Santa Ana Botanic GardenClaremontUSA

Personalised recommendations