On the evolution of clonal plant life histories

  • Markus FischerEmail author
  • Mark Van Kleunen


Clonal plant life histories are special in at least four respects: (1) Clonal plants can also reproduce vegetatively, (2) vegetative reproduction can be realised with short or long spacers, (3) and it may allow to plastically place vegetative offspring in benign patches. (4) Moreover, ramets of clonal plants may remain physically and physiologically integrated. Because of the apparent utility of such traits and because ecological patterns of distribution of clonal and non-clonal plants differ, adaptation is a tempting explanation of observed clonal life-history variation. However, adaptive evolution requires (1) heritable genetic variation and (2) a trait effect on fitness, and (3) it may be constrained if other evolutionary forces are overriding selection or by constraints, costs and trade-offs. (1) The few studies undertaken so far reported broad-sense heritability for clonal traits. Variation in selectively neutral genetic markers appears as pronounced in populations of clonal as non-clonal plants. However, neutral markers may not reflect heritable variation of life-history traits. Moreover, clonal plants may have been sampled at larger spatial scales. Empirical information on the contribution of somatic mutations to heritable variation is lacking. (2) Clonal life-history traits were found to affect fitness. However, much of this evidence stems from artificial rather than natural environments. (3) The relative importance of gene flow, inbreeding, and genetic drift, compared with selection, in the evolution of clonal life histories is hardly explored. Benefits of clonal life-history traits were frequently studied and found. However, there is also evidence for constraints, trade-offs, and costs. In conclusion, though it is very likely, that clonal life-history traits are adaptive, it is neither clear to which degree this is the case, nor which clonal life-history traits constitute adaptations to which environmental factors. Moreover, evolutionary interactions among clonal life-history traits and between clonal and non-clonal ones, such as the mating system, are not well explored. There remains much interesting work to be done in this field — which will be particularly interesting if it is done in the field.

Key words

adaptive evolution clonal integration constraint environmental heterogeneity foraging genetic variation guerilla and phalanx strategy natural selection phenotypic plasticity sexual and vegetative reproduction trade-off 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Alpert, P. (1995) Does clonal growth increase plant performance in natural communities? Abstr. Bot. 19, 11–16.Google Scholar
  2. Alpert, P. (1999) Clonal integration in Fragaria chiloensis differs between populations: ramets from grassland are selfish. Oecologia 120, 69–76.CrossRefGoogle Scholar
  3. Alpert, P. and Stuefer, J.F. (1997) Division of labour in clonal plants. In H. de Kroon and J. van Groenendael (eds) The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, pp. 137–154.Google Scholar
  4. Andrews, J.H. (1998) Bacteria as modular organisms. Ann. Rev. Microbial. 52, 105–126.CrossRefGoogle Scholar
  5. Bach, C.E. (2000) Effects of clonal integration on response to sand burial and defoliation by the dune plant Ipomoea pescaprae (Convolvulaceae). Aust. J. Bot. 48, 159–166.CrossRefGoogle Scholar
  6. Bell, A.D. (1984) Dynamic morphology: a contribution to plant population ecology. In R. Dirzo and J. Sarukhan (eds) Perspectives on Plant Population Ecology. Sinauer, Sunderland, pp. 48–65.Google Scholar
  7. Bradshaw, A.D. (1965) Evolutionary significance of phenotypic plasticity in plants. Adv. Genet. 13, 115–155.CrossRefGoogle Scholar
  8. Butlin, R.K. and Tregenza, T. (1998) Levels of genetic polymorphism: marker loci vs. quantitative traits. Phil. Trans. R. Soc. Lon. B 353, 187–198.PubMedCrossRefGoogle Scholar
  9. Byers, D.L. and Waller, D.M. (1999) Do plant populations purge their genetic load? Effects of population size and mating history on inbreeding depression. Ann. Rev. Ecol. Syst. 30, 479–513.CrossRefGoogle Scholar
  10. Cain, M.L. (1994) Consequences of foraging in clonal plant species. Ecology 75, 933–944.CrossRefGoogle Scholar
  11. Cain, M.L., Dudle, D.A. and Evans, J.P. (1996) Spatial models of foraging in clonal plant species. Am. J. Bot. 83, 76–85.CrossRefGoogle Scholar
  12. Callaghan, T.V. (1988) Physiological and demographic implications of modular construction in cold environments. In A.J. Davy, M.J. Hutchings and A.R. Watkinson (eds) Plant Population Ecology. Blackwell Scientific, Oxford, pp. 111–135.Google Scholar
  13. Caraco, T. and Kelly, C.K. (1991) On the adaptive value of physiological integration in clonal plants. Ecology 72, 81–93.CrossRefGoogle Scholar
  14. Charlesworth, B. (1992) Evolutionary rates in partially self-fertilizing species. Am. Nat. 140, 126148.Google Scholar
  15. Charpentier A., Grillas, P. and Thompson, J.D. (2000) The effects of population size limitation on fecundity in mosaic populations of the clonal macrophyte Scirpus maritimus (Cyperaceae). Am. J. Bot. 87, 502–507.PubMedCrossRefGoogle Scholar
  16. Charpentier A., Mesleard, F. and Thompson, J.D. (1998) The effects of rhizome severing on the clonal growth and clonal architecture of Scirpus maritimus. Oikos 83, 107–116.CrossRefGoogle Scholar
  17. Cheplick, G.P. (1995) Genotypic variation and plasticity of clonal growth in relation to nutrient availability in Amphibromus scabrivalvis. J. Ecol. 83, 459–468.CrossRefGoogle Scholar
  18. Cheplick, G.P. (1997) Responses to severe competitive stress in a clonal plant: differences between genotypes. Oikos 79, 581–591.CrossRefGoogle Scholar
  19. Cheplick, G.P. and Gutierrez, C.M. (2000) Clonal growth and storage in relation to competition in genets of the rhizomatous perennial Amphibromus scabrivalvis. Can. J. Bot. 78, 537–546.Google Scholar
  20. Cook, R.E. (1985) Growth and development in clonal plant populations. In J.B.C. Jackson, L.W. Buss and R.E. Cook (eds) Population Biology and Evolution of Clonal Organisms. Yale University Press, New Haven, pp. 259–296.Google Scholar
  21. de Kroon, H. and Hutchings, M.J. (1995) Morphological plasticity in clonal plants: the foraging concept reconsidered. J. Ecol. 83, 143–152.CrossRefGoogle Scholar
  22. de Kroon, H., Stuefer, J.F., Dong, M. and During, H.J. (1994) On plastic and non-plastic variation in clonal plant morphology and its ecological significance. Folio Geobot. and Phytotax. 29, 123–138.CrossRefGoogle Scholar
  23. DeWitt, T.J., Sih, A. and Wilson, D.S. (1998) Costs and limits of phenotypic plasticity. Trends Ecol. Evol. 13, 77–81.PubMedCrossRefGoogle Scholar
  24. D’Hertefeldt, T. and van der Putten, W.H. (1998) Physiological integration of the clonal plant Carex arenaria and its response to soil-borne pathogens. Oikos 81, 229–237.CrossRefGoogle Scholar
  25. Dolch, R. and Tscharntke, T. (2000) Defoliation of alders (Alnus glutinosa) affects herbivory by leaf beetles on undamaged neighbours. Oecologia 125, 504–511.CrossRefGoogle Scholar
  26. Eckert, C.G. (2000) Contributions of autogamy and geitonogamy to self-fertilization in a mass-flowering, clonal plant. Ecology 81, 532–542.CrossRefGoogle Scholar
  27. Ellstrand, N.C. and Roose, M.L. (1987) Patterns of genotypic diversity in clonal plant species. Am. J. Bot. 74, 123–131.CrossRefGoogle Scholar
  28. Epstein, E. (1972) Mineral Nutrition of Plants: Principles and Perspectives. John Wiley and Sons, New York.Google Scholar
  29. Eriksson, O. (1989) Seedling dynamics and life histories in clonal plants. Oikos 55, 231–238.CrossRefGoogle Scholar
  30. Eriksson, O. (1997) Clonal life histories and the evolution of seed recruitment. In H. de Kroon and J. van Groenendael (eds) The Ecology und Evolution of Clonal Plants. Backhuys Publishers, Leiden, pp. 211–226.Google Scholar
  31. Fahrig, L., Coffin, D.P., Lauenroth, W.K., and Shugart, H.H. (1994) The advantage of long-distance clonal spreading in highly disturbed habitats. Evol. Ecol. 8, 172–187.CrossRefGoogle Scholar
  32. Falconer, D.S. and Mackay, T.F.C. (1996) Introduction to Quantitative Genetics. Longman, Essex. Fischer, M. and Stöcklin, J. (1997) Local extinctions of plants in remnants of extensively used calcareous grasslands 1950–1985. Cons. Biol. 11, 727–737.Google Scholar
  33. Fischer, M., Husi, R., Prati. D., Peintinger, M., van Kleunen, M. and Schmid, B. (2000) RAPD variation among and within small and large populations of the rare clonal plant Ranunculus reptans (Ranunculaceae). Am. J. Bot. 87, 1128–1137.PubMedGoogle Scholar
  34. Friedman, D. and Alpert, P. (1991) Reciprocal transport between ramets increases growth of Fragaria chiloensis when light and nitrogen occur in separate patches but only if patches are rich. Oecologia 86, 76–80.CrossRefGoogle Scholar
  35. Gardner, S.N. and Mangel, M. (1999) Modeling investments in seeds, clonal offspring, and translocation in a clonal plant. Ecology 80, 1202–1220.CrossRefGoogle Scholar
  36. Geber, M.A., Watson, M.A. and Furnish, R. (1992) Genetic-differences in clonal demography in Eichhornia-crassipes. J. Ecol. 80, 329–341.CrossRefGoogle Scholar
  37. Gliddon, C., Belhassen, E., and Gouyon, P.H. (1987) Genetic neighborhoods in plants with diverse systems of mating and different patterns of growth. Heredity 59, 29–32.CrossRefGoogle Scholar
  38. Gliddon, C. and Saleem (1985) The adaptive significance of clonal reproduction in angiosperms–an aquatic perspective. Aqua. Bot. 44, 159–180.Google Scholar
  39. Handel, S.N. (1985) The intrusion of clonal growth-patterns on plant-breeding systems. Am. Nat. 125, 367–384.CrossRefGoogle Scholar
  40. Harada, Y., Kawano, S. and Iwasa, Y. (1997) Clonal identity probability: inferring the relative success of sexual vs. clonal reproduction from spatial genetic patterns. J. Ecol. 85, 591–600.CrossRefGoogle Scholar
  41. Harper, J.L. (1981) The concept of populations in modular organisms. In R.M. May (ed.) Theoretical Ecology: Principles and Applications. Blackwell Scientific, Oxford, pp. 53–77.Google Scholar
  42. Hard, D.L. and Clark, A.G. (1994) Principles of Population Genetics. Sinauer, Sunderland.Google Scholar
  43. Holler, L.C. and Abrahamson, W.G. (1977) Seed and vegetative reproduction in relation to density in Fragaria virginiana (Rosaceae). Am. J. Bot. 64, 1003–1007.CrossRefGoogle Scholar
  44. Huber, H. and Wiggerman, L. (1997) Shade avoidance in the clonal herb Trifolium fragiferum: a field study with experimentally manipulated vegetation height. Plant Ecol. 130, 53–62.CrossRefGoogle Scholar
  45. Humphrey, L.D. and Pyke, D.A. (1997) Clonal foraging in perennial wheatgrasses: a strategy for exploiting patchy soil nutrients. J. Ecol. 85, 601–610.CrossRefGoogle Scholar
  46. Humphrey, L.D. and Pyke, D.A. (1998) Demographic and growth responses of a guerrilla and a phalanx perennial grass in competitive mixtures. J. Ecol. 86, 854–865.CrossRefGoogle Scholar
  47. Hutchings, M.J. and de Kroon, H. (1994) Foraging in plants: the role of morphological plasticity in resource acquisition. Adv. Ecol. Res. 25, 159–238.CrossRefGoogle Scholar
  48. Hutchings, M.J. and Wijesinghe, D.K. (1997) Patchy habitats, division of labour and growth dividends in clonal plants. Trends Ecol. Evol. 12, 390–394.PubMedCrossRefGoogle Scholar
  49. Jonsdottir, I.S. and Watson, M.A. (1997) Extensive physiological integration: an adaptive trait in resource-poor environments? In H. de Kroon and J. van Groenendael (eds) The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, pp. 109–136.Google Scholar
  50. Jordan, N. (1991) Multivariate analysis of selection in experimental populations of two ecotypes of the annual plant Diodia teres W. (Rubiaceae). Evolution 45: 1760–1772.CrossRefGoogle Scholar
  51. Keller, L. (1999) Levels of Selection in Evolution. Princeton University Press, Princeton. Klekowski, E.J. (1997) Somatic mutation theory of clonality. In H. de Kroon and J. van Groenendael (eds) The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, pp. 227–241.Google Scholar
  52. Klimes, L., Klimesovd, J., Hendriks, R. and van Groenendael, J. (1997) Clonal plant architecture: a comparative analysis of form and function. In H. de Kroon and J. van Groenendael (eds) The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, pp. 1–30.Google Scholar
  53. Lande, R. and Arnold, S.J. (1983) The measurement of selection on correlated characters. Evolution 37, 1210–1226.CrossRefGoogle Scholar
  54. Lechowicz, M.J. and Bell, G. (1991) The ecology and genetics of fitness in forest plants. 2. Microspatial heterogeneity of the edaphic environment. J. Ecol. 79, 687–696.CrossRefGoogle Scholar
  55. Lovett Doust, L. (1981) Population dynamics and local specialization in a clonal perennial (Ranunculus repens). I. The dynamics of ramets in contrasting habitats. J. Ecol. 69, 743–755.CrossRefGoogle Scholar
  56. Lovett Doust, L. and Lovett Doust, J. (1982) The battle strategies of plants. New Sci. 8, 81–84.Google Scholar
  57. Lynch M. (1999) Estimating genetic correlations in natural populations. Gen. Res. 74, 255–264.CrossRefGoogle Scholar
  58. Lynch, M. and Walsh, B. (1998) Genetics and Analysis of Quantitative Traits. Sinauer, Sunderland.Google Scholar
  59. Mazer, S.J. (1998) Alternative approaches to the analysis of comparative data: compare and contrast. Am. J. Bot. 85, 1194–1199.CrossRefGoogle Scholar
  60. McCrea, K.D. and Abrahamson, W.G. 1985. Evolutionary impacts of the goldenrod ball gall-maker on Solidago altissima clones. Oecologia 68, 20–22.CrossRefGoogle Scholar
  61. McLellan, A.J., Prati, D., Kaltz, O. and Schmid, B. (1997) Structure and analysis of phenotypic and genetic variation in clonal plants. In H. de Kroon and J. van Groenendael (eds) The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, pp. 185–210.Google Scholar
  62. Mitton, J.B. and Grant, M.C. (1996) Genetic variation and the natural history of quaking aspen. Bioscience 46, 25–31.CrossRefGoogle Scholar
  63. Mousseau, T.A. and Roff, D.A. (1987) Natural selection and the heritability of fitness components. Heredity 59, 181–197.PubMedCrossRefGoogle Scholar
  64. Muirhead C.A. and Lande, R. (1997) Inbreeding depression under joint selfing, outcrossing, and asexuality. Evolution 51, 1409–1415.CrossRefGoogle Scholar
  65. Niklas, K.J. (1988) Biophysical limitations on plant form and evolution. In L.D. Gottlieb and S.K.Google Scholar
  66. Jain (eds) Plant Evolutionary Ecology. Chapman and Hall, London, pp. 185–220.Google Scholar
  67. Oborny, B. (1994) Growth rules in clonal plants and environmental predictability–a simulation study. J. Ecol. 82, 341–351.CrossRefGoogle Scholar
  68. Oborny B., Kun, A., Czaran, T. and Bokros, S. (2000) The effect of clonal integration on plant competition for mosaic habitat space. Ecology 81, 3291–3304.CrossRefGoogle Scholar
  69. Orive, M.E. (1993) Effective population size in organisms with complex life histories. Theor. Popul. Biol. 44, 316–340.PubMedCrossRefGoogle Scholar
  70. Orive, M.E. (2001) Somatic mutations in organisms with complex life histories. Theor. Popul. Biol. 59, 235–249.PubMedCrossRefGoogle Scholar
  71. Otto, S.P. and Orive, M.E. (1995) Evolutionary consequences of mutation and selection within an individual. Genetics 141, 1173–1187.PubMedGoogle Scholar
  72. Ouborg, N.J., Piquot, Y. and van Groenendael, J.M. (1999) Population genetics, molecular markers and the study of dispersal in plants. J. Ecol. 87, 551–568.CrossRefGoogle Scholar
  73. Pennings, S.C. and Callaway, R.M. (2000) The advantages of clonal integration under different ecological conditions: A community-wide test. Ecology 81, 709–716.CrossRefGoogle Scholar
  74. Pineda-Krch, M. (2001) Evolution in multicellular mitotic lineages. PhD Thesis, University of Lund.Google Scholar
  75. Piqueras, J. (1999) Infection of Trientalis europaea by the systemic smut fungus Urocystis trientalis: disease incidence, transmission and effects on performance of host ramets. J. Ecol. 87, 995–1004.CrossRefGoogle Scholar
  76. Piquot, Y., Petit, D., Valero, M., Cuguen, J., de Laguerie, P. and Vernet, P. (1998) Variation in sexual and asexual reproduction among young and old populations of the perennial macrophyte Sparganium erectum. Oikos 82, 139–148.CrossRefGoogle Scholar
  77. Pitelka, L.F. and Ashmun, J.W. (1985) Physiology and integration of ramets in clonal plants. In J.B.C. Jackson, L.W. Buss and R.E. Cook (eds) Population Biology and Evolution of Cloned Organisms. Yale University Press, New Haven, pp. 399–435.Google Scholar
  78. Prati, D. (1998) The genetics and life-history evolution of the clonal plant Ranunculus reptans. PhD Thesis, University of Zürich.Google Scholar
  79. Prati, D. and Schmid, B. (2000) Genetic differentiation of life-history traits within populations of the clonal plant Ranunculus reptans. Oikos 90, 442–456.CrossRefGoogle Scholar
  80. Rausher, M.D. (1992) The measurement of selection on quantitative traits: biases due to environmental covariances between traits and fitness. Evolution 46, 616–626.CrossRefGoogle Scholar
  81. Ritland, K. (2000) Detecting inheritance with inferred relatedness in nature. In T.A. Mousseau, B. Sinervo and J. Endler (eds) Adaptive Genetic Variation in the Wild. Oxford University Press, New York, pp. 187–199.Google Scholar
  82. Roff, D.A. and Simon, A.M. (1997) The quantitative genetics of wing dimorphism under laboratory and `field’ conditions in the cricket Gryllus pennsylvanicus. Heredity 78, 235–240.CrossRefGoogle Scholar
  83. Sackville Hamilton, N.R., Schmid, B. and Harper, J.L. (1987) Life-history concepts and the population biology of clonal organisms. Proc. R. Soc. Lond. 232, 35–57.CrossRefGoogle Scholar
  84. Saikkonen, K., Koivunen, S., Vuorisalo, T. and Mutikainen, P. (1998) Interactive effects of pollination and heavy metals on resource allocation in Potentilla anserina L. Ecology 79, 1620–1629.Google Scholar
  85. Salzman, A.G. and Parker, M.A. (1985) Neighbors ameliorate local salinity stress for a rhizomatous plant in a heterogeneous environment. Oecologia 65, 273–277.CrossRefGoogle Scholar
  86. Schaffer, R., Landgraf, J., Perez-Amador, M. and Wisman, E. (2000) Monitoring genome-wide expression in plants. Curr. Opin. Biotech. 11, 162–167.PubMedCrossRefGoogle Scholar
  87. Scheiner, S.M. (1993) Genetics and evolution of phenotypic plasticity. Ann. Rev. Ecol. Syst. 24, 35–68.CrossRefGoogle Scholar
  88. Scheiner, S.M. and Lyman, R.F. (1991) The genetics of phenotypic plasticity. 11. Response to selection. J. Evol. Biol. 4, 23–50.CrossRefGoogle Scholar
  89. Schena, M., Shalon, D., Davis, R.W. and Brown, P.O. (1995) Quantitative monitoring of gene expression patterns with a complementary DNA microarray. Science 270, 467–470.PubMedCrossRefGoogle Scholar
  90. Schenk, P.M., Kazan, K., Wilson, I., Anderson, J.P., Richmond, T., Somerville, S.C. and Manners, J.M. (2000) Coordinated plant defense responses in Arahidopsis revealed by microarray analysis. Proc. Nat. Acad. Sei. USA 97, 11655–11660.CrossRefGoogle Scholar
  91. Schläpfer, F. and Fischer, M. (1998) An isozyme study of clone diversity and relative importance of sexual and vegetative reproduction in the grass Brachypodium pinnatum. Ecography 21, 351–360.CrossRefGoogle Scholar
  92. Schlichting, C.D. and Pigliucci, M. (1998) Phenotypic Evolution: A Reaction Norm Perspective. Sinauer, Sunderland.Google Scholar
  93. Schmid, B. (1985) Clonal growth in grassland perennials 11I. Genetic variation and plasticity between and within populations of Bellis perennis and Prunella vulgaris. J. Ecol. 73, 819–830.CrossRefGoogle Scholar
  94. Schmid, B. (1990) Some ecological and evolutionary consequences of modular organization and clonal growth in plants. Evol. Trends Plants 4, 25–34.Google Scholar
  95. Schmid, B. and Bazzaz, F.A. (1990) Plasticity in plant size and architecture in rhizome-derived vs. seed-derived Solidago and Aster. Ecology 71, 523–535.CrossRefGoogle Scholar
  96. Schmid, B. and Bazzaz, F.A. (1991) Growth of transplanted and native shoots in perennials with contrasting genet architecture. Flora 185, 335–344.Google Scholar
  97. Schmid, B. and Harper, J.L. (1985) Clonal growth in grassland perennials. Il. Growth form and fine-scale colonization ability. J. Ecol. 73, 809–818.CrossRefGoogle Scholar
  98. Schmid, B. and Weiner, J. (1993) Plastic relationships between reproductive and vegetative mass in Solidago altissima. Evolution 47, 61 74.Google Scholar
  99. Schmid, B., Puttick, G.M., Burgess, K.H. and Bazzaz, F.A. (1988) Clonal integration and effects of simulated herbivory in old-field perennials. Oecologia 75, 465–471.CrossRefGoogle Scholar
  100. Schmitt, J., Dudley, S.A. and Pigliucci, M. (1999) Manipulative approaches to testing adaptive plasticity: phytochrome-mediated shade-avoidance responses in plants. Am. Nat. 154, S43 - S54.CrossRefGoogle Scholar
  101. Seskar, M., Shulaev. V., and Raskin, 1. (1998) Endogenous methyl salicylate in pathogen-inoculated tobacco plants. Plant Physiol. 116, 387–392.Google Scholar
  102. Silander, J.A. (1985) Microevolution in clonal plants. In J.B.C. Jackson, L.W. Buss and R.E. Cook (eds) Population Biology and Evolution of Clonal Organisms. Yale University Press, New Haven, pp. 107–152.Google Scholar
  103. Skalova, H., Krahulec, F., During, H.J., Hadincova, V., Pechackova, S., and Herben, T. (1999) Grassland canopy composition and spatial heterogeneity in the light quality. Plant Ecol. 143, 129–139.CrossRefGoogle Scholar
  104. Skâlovâ, H., Pechäíckovâ, S., Suzuki, J., Herben, T., Hara, T., Hadincovâ, V. and Krahulec, F. (1997) Within population genetic differentiation in traits affecting clonal growth: Festuca rubra in a mountain grassland. J. Evol. Biol. 10, 383–406.CrossRefGoogle Scholar
  105. Stearns, S.C. (1992) The evolution of life-histories. Oxford University Press, Oxford, UK.Google Scholar
  106. Stehlik, I. and Holderegger, R. (2000) Spatial genetic structure and clonal diversity of Anemone nemorosa in late successional deciduous woodlands of Central Europe. J. Ecol. 88, 424–435.CrossRefGoogle Scholar
  107. Stöcklin, J. (1992) Environment, morphology and growth of clonal plants–an overview. Bot. Heiv. 102, 3–21 (in German).Google Scholar
  108. Stöcklin, J. and Favre, P. (1999) Differences in life history traits of related Epilobium species. Clonality, seed size and seed number. Folio Geobot. 34, 7–18.CrossRefGoogle Scholar
  109. Stuefer, J.F., During, H.J. and de Kroon, H. (1994) High benefits of clonal integration in two stoloniferous species, in response to heterogeneous light environment. J. Ecol. 82, 511–518.CrossRefGoogle Scholar
  110. Sutherland, W.J. and Stillman, R.A. (1988) The foraging tactics of plants. Oikos 52, 239–244.CrossRefGoogle Scholar
  111. Takhtajan, A.L. (1980) Outline of the classification of flowering plants (Magnoliophyta). Bot. Rev. 46, 225–359.CrossRefGoogle Scholar
  112. Tiffney, B.H. and Niklas, K.J. (1985) Clonal growth in land plants, a paleobotanical perspective. In J.B.C. Jackson, L.W. Buss and R.E. Cook (eds) Population Biology and Evolution of Clonal Organisms. Yale University Press, New Haven, pp. 35–66.Google Scholar
  113. Tuomi, J. and Vuarisalo, T. (1989) What are the units of selection in modular organisms. Oikos 54, 227–233.CrossRefGoogle Scholar
  114. van Baalen, J., Ernst, W.H.O., van Andel, J., Janssen, D.W. and Nelissen, H.J.M. (1990) Reproductive allocation in plants of Scrophularia nodosa grown at various levels of irradiance and soil fertility. Acta Bot. Neerl. 39, 183–196.Google Scholar
  115. van Groenendael, J.M., Klimes, L., Klimesovh, J. and Hendriks, F.J.J. (1996) Comparative ecology of clonal plants. Phil. Trans. R. Soc. Land. B 351, 1331–1339.CrossRefGoogle Scholar
  116. van Kleunen, M. and Fischer, M. (2001) Adaptive evolution of plastic foraging responses in a clonal plant. Ecology 82, 3309–3319.CrossRefGoogle Scholar
  117. van Kleunen, M. and Stuefer, J.F. (1999) Quantifying the effects of reciprocal assimilate and water translocation in a clonal plant by the use of steam-girdling. Oikos 85, 135–145.CrossRefGoogle Scholar
  118. van Kleunen, M., Fischer, M. and Schmid, B. (2000a) Costs of plasticity in foraging characteristics of the clonal Ranunculus reptans. Evolution 54, 1947–1955.CrossRefGoogle Scholar
  119. van Kleunen, M., Fischer, M. and Schmid, B. (2000b). Clonal integration in Ranunculus reptans: by-product or adaptation? J. Evol. Biol. 13, 237–248.CrossRefGoogle Scholar
  120. van Tienderen, P.H. (1992) Variation in a population of Plantago lanceolata along a topographical gradient. Oikos 64, 560–572.CrossRefGoogle Scholar
  121. Via, S. (1993) Adaptive phenotypic plasticity–target or by-product of selection in a variable environment. Am. Nat. 142, 352–365.PubMedCrossRefGoogle Scholar
  122. Watson, M.A. and Casper, B.B. (1984) Morphogenetic constraints on patterns of carbon distribution in plants. An. Rev. Ecol. Syst. 15, 233–258.CrossRefGoogle Scholar
  123. Wenström, A. and Ericson, L. (1992) Environmental heterogeneity and disease transmission within clones of Lactuca sibirica. J. Ecol. 80, 71–77.Google Scholar
  124. Widén, B., Cronberg, N. and Widén, M. (1994) Genotypic diversity, molecular markers and spatial distribution of genets in clonal plants, a literature survey. Folia Geobot. Phytotax. 29, 245–263.CrossRefGoogle Scholar
  125. Wilhalm, T. (1995) A comparative study of clonal fragmentation in tussock-forming grasses. Abstr. Bot. 19, 51–60.Google Scholar
  126. Williams, G.C. (1975) Sex and Evolution. Princeton University Press, New Jersey.Google Scholar
  127. Winkler, E. and Fischer, M. (1999) Two fitness measures for clonal plants and the importance of spatial aspects. Plant Ecol. 141, 191–199.CrossRefGoogle Scholar
  128. Winkler, E. and Schmid, B. (1995) Clonal strategies of herbaceous plant species: a simulation study on population growth and competition. Abstr. Bot. 19, 17–28.Google Scholar
  129. Winkler, E., Fischer, M. and Schmid, B. (1999) Modelling the competitiveness of clonal plants by complementary analytical and simulation approaches. Oikos 85, 217–233.CrossRefGoogle Scholar
  130. Wolf, A.T., Howe, R.W. and Hamrick, J.L. (2000) Genetic diversity and population structure of the serpentine endemic Calystegia collina (Convolvulaceae) in northern California. Am. J. Bot. 87, 1138–1146.PubMedCrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media Dordrecht 2002

Authors and Affiliations

  1. 1.Institut für UmweltwissenschaftenUniversity of ZürichZürichSwitzerland

Personalised recommendations