View More View Less
  • 1 Department of Plant Taxonomy and Ecology, L. Eötvös Unversity H-1117 Budapest, Pázmány P. sétány 1/C
  • 2 Department of Plant Taxonomy and Ecology, L. Eötvös Unversity H-1117 Budapest, Pázmány P. sétány 1/C
Restricted access

Clonal populations are hierarchically organized: genetic individuals (genets) can consist of many physiological individuals (ramets). Each ramet takes up resources from its local environment, but the resource pattern can be reorganized within the clone by transport between ramets. Thus, an integrated clone is not directly subject to the pattern of resource availability in its habitat. Local shortages can be compensated, hence, the clone can buffer itself against spatio-temporal heterogeneity in the habitat. We modelled a series of habitat types, assuming that one limiting resource was patchily distributed in space, and could fluctuate over time. Habitat types differed in the density, size and persistence of resource patches, and in the contrast between resource-rich patches and the resource-poor background. We applied an individual-based, spatially explicit population dynamic model to compare the performance of two plant strategies in these habitat types. In the Integrator, ramets that were interconnected distributed the resource evenly. In the Splitter, no resource translocation occurred. First we observed population growth of the two strategies separately, then in competition. We found a range of habitat types, where none of the strategies was viable, because of the scarcity of resource patches. As the density of resource patches was increased, first only the Integrator could persist. Then, at intermediate densities of resource patches, the Splitter became viable, and, being a stronger competitor, excluded the Integrator. Finally, at high resource-patch densities, the Integrator occupied the area again. Since the Splitter was viable at high density of resource patches when growing alone, its disappearance is more due to spontaneous extinction, due to competitive exclusion by the Integrator. We predict, therefore, the dominance of integrated clones both in extremely unproductive and productive environments, but for different reasons. It is important to note that this trend was observable only at high spatial and temporal variation in resource availability. Less contrast between patches of different quality, smaller patch sizes, or longer persistence of patches facilitated the dominance of the Splitter. Thus, buffering is advantageous in many but not all habitat types.

  • Wijesinghe, D.K. 1994. Temporal and structural components of ramet independence in the clonal perennial herb, Potentilla simplex. J. Ecol. 82:13-20.

    'Temporal and structural components of ramet independence in the clonal perennial herb ' () 82 Potentilla simplex. J. Ecol. : 13 -20.

    • Search Google Scholar
  • Hutchings, M.J. and D.K. Wijesinghe. 1997. Patchy habitats, division of labour and growth dividends in clonal plants. TREE 12:390-394.

    'Patchy habitats, division of labour and growth dividends in clonal plants ' () 12 TREE : 390 -394.

    • Search Google Scholar
  • Slade, A.J. and M.J. Hutchings. 1987. Clonal integration and plasticity in foraging behavior in Glechoma hederacea. J. Ecol. 75:1023-1036.

    'Clonal integration and plasticity in foraging behavior ' () 75 Glechoma hederacea. J. Ecol. : 1023 -1036.

    • Search Google Scholar
  • Schmid, B. and F.A. Bazzaz. 1987. Clonal integration and population structure in perennials: effects of severing rhizome connections.Ecology 68:2016-2022.

    'Clonal integration and population structure in perennials: effects of severing rhizome connections ' () 68 Ecology : 2016 -2022.

    • Search Google Scholar
  • Schmid, B. 1986. Spatial dynamics and integration within clones of grassland perennials with different growth form. Proc. Roy. Soc. Lond.B 228:173-186.

    'Spatial dynamics and integration within clones of grassland perennials with different growth form ' () 228 Proc. Roy. Soc. Lond. B : 173 -186.

    • Search Google Scholar
  • Alpert, P. 1996. Does clonal growth increase plant performance in natural communities? In: B. Oborny and J. Podani (eds). Clonality in Plant Communities. Opulus Press, Uppsala, pp. 11-16.

    Does clonal growth increase plant performance in natural communities , () 11 -16.

  • Alpert, P. 1999. Clonal integration in Fragaria chiloensis differs between populations: ramets from grassland are selfish. Oecologia 120:69-76.

    'Clonal integration in Fragaria chiloensis differs between populations: ramets from grassland are selfish ' () 120 Oecologia : 69 -76.

    • Search Google Scholar
  • Alpert, P., C. Holzapfel and J.M. Benson. 2002. Hormonal control of resource sharing in the clonal plant Fragaria chiloensis. Funct. Ecol. 16:191-197.

    'Hormonal control of resource sharing in the clonal plant ' () 16 Fragaria chiloensis. Funct. Ecol. : 191 -197.

    • Search Google Scholar
  • Alpert, P., C. Holzapfel and C. Slominski. 2003. Differences in performance between genotypes of Fragaria chiloensis with different degrees of resource sharing. J. Ecol. 91:27-35.

    'Differences in performance between genotypes of Fragaria chiloensis with different degrees of resource sharing ' () 91 J. Ecol. : 27 -35.

    • Search Google Scholar
  • Alpert, P. and H.A. Mooney. 1986. Resource sharing among ramets in the clonal herb, Fragaria chiloensis. Oecologia 70:227-233.

    'Resource sharing among ramets in the clonal herb ' () 70 Fragaria chiloensis. Oecologia : 227 -233.

    • Search Google Scholar
  • Alpert, P. and E.L. Simms. 2002. The relative advantages of plasticity and fixity in diffe environments: when is it good for a plant to adjust. Evol. Ecol. 16:285-297.

    'The relative advantages of plasticity and fixity in diffe environments: when is it good for a plant to adjust ' () 16 Evol. Ecol. : 285 -297.

    • Search Google Scholar
  • Apert, P. and J. Stuefer. 1997. Division of labour in clonal plants. In: H. de Kroon and J. van Groenendael (eds). The Evolution and Ecology of Clonal Plants. Backhuys Publishers, Leiden, pp. 137-154.

    Division of labour in clonal plants , () 137 -154.

  • Birch, C.P.D. and MJ. Hutchings. 1994. Exploitation of patchily distributed soil resources by the clonal herb Glechoma hederacea. J. Ecol. 82:653-664.

    'Exploitation of patchily distributed soil resources by the clonal herb ' () 82 Glechoma hederacea. J. Ecol. : 653 -664.

    • Search Google Scholar
  • Cain, M.L., D. Dudle and J.P. Evans. 1996. Spatial models of foraging in clonal plant species. Am. J. Bot. 83:76-85.

    'Spatial models of foraging in clonal plant species ' () 83 Am. J. Bot. : 76 -85.

  • Caraco, T. and C.K. Kelly. 1991. On the adaptive value of physiological integration in clonal plants. Ecology 72:81-93.

    'On the adaptive value of physiological integration in clonal plants ' () 72 Ecology : 81 -93.

    • Search Google Scholar
  • 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, Connecticut, USA. pp. 259-296.

    Growth and development in clonal plant populations , () 259 -296.

  • Crawley, MJ. 1997. Life history and environment. In: MJ. Crawley (ed.), Plant Ecology. Blackwell Science Ltd., pp. 73-131.

    Life history and environment , () 73 -131.

  • de Kroon, H. and F. Schieving. 1990. Resource partitioning in relation to clonal growth strategy. In: J. van Groenendael and H. de Kroon (eds). Clonal Growth in Plants: Regulation and Function. SPB Academic Publ., Leiden, pp. 113-130.

    Resource partitioning in relation to clonal growth strategy , () 113 -130.

  • de Kroon, H. and J. van Groenendael. 1990. Regulation and function of clonal growth in plants: an evaluation. In: J. van Groenendael and H. de Kroon (eds). Clonal Growth in Plants: Regulation and Function. SPB Academic Publishing, The Hague, pp. 177-186.

    Regulation and function of clonal growth in plants: an evaluation , () 177 -186.

  • D'Hertefeldt, T. and W.H. van der Putten. 1998. Physiological integration of the clonal plant Carex arenaria and its response to siol-borne pathogens. Oikos 81:229-237.

    'Physiological integration of the clonal plant Carex arenaria and its response to siol-borne pathogens ' () 81 Oikos : 229 -237.

    • Search Google Scholar
  • Dickmann, U., R. Law and J.A J. Metz. 2000. The Geometry of Ecological Interactions. Cambridge University Press, Cambridge.

    The Geometry of Ecological Interactions. , ().

  • Eriksson, O. 1997. Clonal life histories and the evolution of seed recruitment. In: J. van Groenendael and H. de Kroon (eds). The Ecology and Evolution of Clonal Plants. Backhuys Publishing, Leiden, The Netherlands, pp. 211-226.

    Clonal life histories and the evolution of seed recruitment , () 211 -226.

  • Eriksson, O. and L. Jerling. 1990. Hierarchical selection and risk spreading in clonal plants. In: J. van Groenendael and H. de Kroon (eds). Clonal Growth in Plants: Regulation and Function. SPB Academic Publ., The Hague, pp. 79-94.

    Hierarchical selection and risk spreading in clonal plants , () 79 -94.

  • Evans, J.P. and M.L. Cain. 1995. A spatially explicit test of foraging behavior in a clonal plant. Ecology 76:1147-1155.

    'A spatially explicit test of foraging behavior in a clonal plant ' () 76 Ecology : 1147 -1155.

    • Search Google Scholar
  • Harper, J.L. 1978. The demography of plants with clonal growth. In: A.H.J. Freysen and J.W. Woldendorps (eds). Structure and Functioning of Plant Population. North Holland, Amsterdam. pp. 27-45.

    The demography of plants with clonal growth , () 27 -45.

  • Harper, J.L. 1985. Modules, branches, and the capture of resources. 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, Connecticut, pp. 1-33.

    Modules, branches, and the capture of resources , () 1 -33.

  • Harper, J.L. and A.D. Bell. 1979. The population dynamics of growth form in organisms with modular construction. In: R.M. Anderson, B.D. Turner and L.R. Taylor (eds). Population Dynamics. Blackwell Scientific Publ., Oxford. pp. 29-52.

    The population dynamics of growth form in organisms with modular construction , () 29 -52.

    • Search Google Scholar
  • Hartnett, CD. and F.A. Bazzaz. 1985. The genet and ramet population dynamics of Solidago canadensis in an abandoned field. J. Ecol. 73:407-413.

    'The genet and ramet population dynamics of Solidago canadensis in an abandoned field ' () 73 J. Ecol. : 407 -413.

    • Search Google Scholar
  • Herben, T., T. Hara, C. Marshall and L. Soukupova. 1994. Plant Clonality: biology and diversity. In: L. Soukupova, C. Marshall, T. Hara and T. Herben (eds). Plant Clonality: Biology and Diversity. Opulus Press, Uppsala, pp. 7-16.

    Plant Clonality: biology and diversity , () 7 -16.

  • Herben, T. and J. Suzuki. 2002. A simulation study of the effects of architectural constrains and resource translocation on population structure and competition in clonal plant. Evol. Ecol. 15:403-423.

    'A simulation study of the effects of architectural constrains and resource translocation on population structure and competition in clonal plant ' () 15 Evol. Ecol. : 403 -423.

    • Search Google Scholar
  • Hutchings, M.J. and I.K. Bradbury. 1986. Ecological perspectives on clonal perennial herbs. BioScience 36:178-182.

    'Ecological perspectives on clonal perennial herbs ' () 36 BioScience : 178 -182.

  • Hutchings, M.J. and H. de Kroon. 1994. Foraging in plants: the role of morphological plasticity in resource acquisition. Adv. Ecol. Research 25:159-238.

    'Foraging in plants: the role of morphological plasticity in resource acquisition ' () 25 Adv. Ecol. Research : 159 -238.

    • Search Google Scholar
  • Hutchings, M.J., D.K. Wijesinghe and E. John. 2000. The effect of heterogeneous nutrient supply on plant performance: a survey of responses, with special reference to clonal herbs. In: M.J. Hutchings, E. John and A.J. Stewart (eds). The Ecological Consequences of Environmental Heterogeneity. Blackwell, Oxford, pp. 91-110.

    The effect of heterogeneous nutrient supply on plant performance: a survey of responses, with special reference to clonal herbs , () 91 -110.

    • Search Google Scholar
  • Jónsdóttir, I.S. and M.A. Watson. 1997. Extensive physiological integration: an adaptive trait in resource-poor environments? In: H. de Kroon and J. van Groenendael (eds). The Evolution and Ecology of Clonal Plants. Backhuys Publishers, Leiden, pp. 109-136.

    Extensive physiological integration: an adaptive trait in resource-poor environments , () 109 -136.

    • Search Google Scholar
  • Kelly, C.K. 1995. Thoughts on clonal integration: facing the evolutionary context. Evol. Ecol. 9:575-585.

    'Thoughts on clonal integration: facing the evolutionary context ' () 9 Evol. Ecol. : 575 -585.

    • Search Google Scholar
  • Klimes, L., J. Klimesova, R. Hendriks and J. van Groenendael. 1997. Clonal plant architecture: a comparative analysis of form and function. In: H. de Kroon and J. van Groenendael (eds). The Evolution and Ecology of Clonal Plants. Backhuys Publishers, Leiden, pp. 1-30.

    Clonal plant architecture: a comparative analysis of form and function , () 1 -30.

  • Landa, K., B. Benner, M.A. Watson and J. Gartner. 1992. Physiological integration for carbon in mayapple (Podophyllum peltatum), a clonal perennial herb. Oikos 63:348-356.

    'Physiological integration for carbon in mayapple (Podophyllum peltatum), a clonal perennial herb ' () 63 Oikos : 348 -356.

    • Search Google Scholar
  • Marshall, C. 1990. Source-sink relations of interconnected ramets. In: J. van Groenendael and H. de Kroon (eds). Clonal Growth in Plants: Regulation and Function. SPB Academic Publ., The Hague, pp. 23-42.

    Source-sink relations of interconnected ramets , () 23 -42.

  • Marshall, C. and E.A.C. Price. 1997. Sectoriality and its implications for physiological integration. In: H. de Kroon and J. van Groenendael (eds). The Evolution and Ecology of Clonal Plants. Backhuys Publishers, Leiden, pp. 79-107.

    Sectoriality and its implications for physiological integration , () 79 -107.

  • Noble, J.C. and C. Marshall. 1983. The population biology of plants with clonal growth II. The nutrient strategy and modular physiology of Carex arenaria. J. Ecol. 71:865-877.

    'The population biology of plants with clonal growth II ' () 71 The nutrient strategy and modular physiology of Carex arenaria. J. Ecol. : 865 -877.

    • Search Google Scholar
  • Oborny, B. and S. Bartha. 1996. Clonality in plant communities: an overview. In: B. Oborny and J. Podani (eds). Clonality in Plant Communities. Opulus Press, Uppsala pp. 115-127.

    Clonality in plant communities: an overview , () 115 -127.

  • Oborny, B. and M.L. Cain. 1997. Models of spatial spread and foraging in clonal plants. In: H. de Kroon and J. van Groenendael (eds). The Ecology and Evolution of Clonal Plants. Backhuys Publ., Leiden, The Netherlands, pp. 115-127.

    Models of spatial spread and foraging in clonal plants , () 115 -127.

  • Wijesinghe, D.K. and D.F. Whigham. 2001. Nutrient foraging in woodland herbs: a comparison of three species of Uvularia (Liliaceae) with contrasting belowground morphologies. Am. J. Bot. 88:1071-1079.

    'Nutrient foraging in woodland herbs: a comparison of three species of Uvularia (Liliaceae) with contrasting belowground morphologies ' () 88 Am. J. Bot. : 1071 -1079.

    • Search Google Scholar
  • Wilhalm, T. 1996. A comparative study of clonal fragmentation in tussock-forming grasses. In: B. Oborny and J. Podani (eds). Clonality in Plant Communities. Opulus Press, Uppsala, pp. 51-60.

    A comparative study of clonal fragmentation in tussock-forming grasses , () 51 -60.

  • Yu, F., Y. Chen and M. Dong. 2002. Clonal integration enhances survival and performance of Potentilla anserina, suffering from partial sand burial on Ordos plateau, China. Evol. Ecol. 15:303-318.

    'Clonal integration enhances survival and performance of Potentilla anserina, suffering from partial sand burial on Ordos plateau, China ' () 15 Evol. Ecol. : 303 -318.

    • Search Google Scholar
  • Pennings, S.C. and R.M. Callaway. 2000. The advantage of clonal integration under different ecological conditions: a community-wide test. Ecology 81:709-716.

    'The advantage of clonal integration under different ecological conditions: a community-wide test ' () 81 Ecology : 709 -716.

    • Search Google Scholar
  • Peterson, C.J. and R.H. Jones. 1997. Clonality in woody plants: a review and comparison with clonal herbs. In: J. van Groenendael and H. de Kroon (eds). The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, The Netherlands, pp. 263-290

    Clonality in woody plants: a review and comparison with clonal herbs , () 263 -290.

  • Pitelka, L.F. and J.W Ashmun. 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 Clonal Organisms. Yale University Press, New Haven, Connecticut, USA. pp. 399-435

    Physiology and integration of ramets in clonal plants , () 399 -435.

  • Price, E.A.C. and C. Marshall. 1999. Clonal plants and environmental heterogeneity. Plant Ecology 141:3-7.

    'Clonal plants and environmental heterogeneity ' () 141 Plant Ecology : 3 -7.

  • Pysek, P. 1997. Clonality and plant invasion: can a trait make a difference? In: J. van Groenendael and H. de Kroon (eds). The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leidens, The Netherlands, pp. 405-427.

    Clonality and plant invasion: can a trait make a difference , () 405 -427.

  • Sachs, T. and A. Novoplansky. 1997. What does aclonal organisation suggest concerning clonal plants? In: J. van Groenendael and H. de Kroon (eds). The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, The Netherlands, pp. 55-78.

    What does aclonal organisation suggest concerning clonal plants , () 55 -78.

  • Salzman, A.G. 1985. Habitat selection in a clonal plant. Science 228:603-604.

    'Habitat selection in a clonal plant ' () 228 Science : 603 -604.

  • Stoll, P. and J. Weiner. 2000. A neighborhood view of interactions among individual plants. In: U. Dieckmann, R. Law and J.A.J. Metz (eds). The Geometry of Ecological Interactions. Cambridge University Press, pp. 11-27.

    A neighborhood view of interactions among individual plants , () 11 -27.

  • Stuefer, J. and H. Huber. 1999. The role of stolon internodes for ramet survival after clone fragmentation in Potentilla anserina. Ecology Letters 2:135-139.

    'The role of stolon internodes for ramet survival after clone fragmentation. Potentilla anserina ' () 2 Ecology Letters : 135 -139.

    • Search Google Scholar
  • Stuefer, J. and M.J. Hutchings. 1994. Environmental heterogeneity and clonal growth - a study of the capacity for reciprocal translocation in Glechoma hederacea L. Oecologia 100:302-308.

    'Environmental heterogeneity and clonal growth - a study of the capacity for reciprocal translocation in Glechoma hederacea L. ' () 100 Oecologia : 302 -308.

    • Search Google Scholar
  • Stuefer, J.F. 1996. Potential and limitations of current concepts regarding the responses of clonal plants to environmental heterogeneity. Vegetatio 127:55-70.

    'Potential and limitations of current concepts regarding the responses of clonal plants to environmental heterogeneity ' () 127 Vegetatio : 55 -70.

    • Search Google Scholar
  • Stuefer, J.F., H. De Kroon and HJ. During. 1996. Exploitation of environmental heterogeneity by spatial division of labour in a clonal plant. Funct. Ecol. 10:328-334.

    'Exploitation of environmental heterogeneity by spatial division of labour in a clonal plant ' () 10 Funct. Ecol. : 328 -334.

    • Search Google Scholar
  • Stuefer, J.F., H. J. During and H. De Kroon. 1994. High benefits of clonal integration in two stoloniferous species, in response to heterogeneous light environments. J. Ecol. 82:511-518.

    'High benefits of clonal integration in two stoloniferous species, in response to heterogeneous light environments ' () 82 J. Ecol. : 511 -518.

    • Search Google Scholar
  • Suzuki, J. and M.J. Hutchings. 1997. Interactions between shoots in clonal plants and the effects of stored resources on the structure of shoot populations. In: H. de Kroon and J. van Groenendael (eds). The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, pp. 311-330.

    Interactions between shoots in clonal plants and the effects of stored resources on the structure of shoot populations , () 311 -330.

    • Search Google Scholar
  • Suzuki, J. and J. Stuefer. 1999. On the ecological and evolutionary significance of storage in clonal plants. Plant Species Biology 14:11-17.

    'On the ecological and evolutionary significance of storage in clonal plants ' () 14 Plant Species Biology : 11 -17.

    • Search Google Scholar
  • van Kleunen, M., M. Fischer and B. Schmid. 2000. Clonal integration in Ranunculus reptans: by-product or adaptation? J. Evol Biol. 13:237-248.

    'Clonal integration in Ranunculus reptans: by-product or adaptation ' () 13 J. Evol Biol. : 237 -248.

    • Search Google Scholar
  • Vourisalo, T., J. Tuomi, B. Pedersen and P. Käär. 1997. Hierarchical selection in clonal plants. In: H. De Kroon and J. van Groenedael (eds), The Ecology and Evolution of Clonal Plants. Backhuys Publishers, Leiden, The Netherlands, pp. 243-262.

    Hierarchical selection in clonal plants , () 243 -262.

  • Watson, M.A. 1986. Integrated physiological units in plants. TREE 1:119-123.

    'Integrated physiological units in plants ' () 1 TREE : 119 -123.

  • Wennström, A. 1999. The effect of systematic rusts and smuts on clonal plants in natural systems. Plant Ecology 141:93-97.

    'The effect of systematic rusts and smuts on clonal plants in natural systems ' () 141 Plant Ecology : 93 -97.

    • Search Google Scholar
  • Oborny, B., T. Czárán and Á. Kun. 2001. Exploration of resource patches by clonal growth: a spatial model on the effect of transport between modules. Ecol. Model. 141:151-169.

    'Exploration of resource patches by clonal growth: a spatial model on the effect of transport between modules ' () 141 Ecol. Model. : 151 -169.

    • Search Google Scholar
  • Oborny, B. and Á. Kun. 2002. Fragmentation of clones: How does it influence dispersal and competitive ability? Evol. Ecol. 15:319-346.

    'Fragmentation of clones: How does it influence dispersal and competitive ability ' () 15 Evol. Ecol. : 319 -346.

    • Search Google Scholar
  • Oborny, B., Á. Kun, T. Czárán and S. Bokros. 2000. The effect of clonal integration on plant competition for mosaic habitat space. Ecology 81:3291-3304.

    'The effect of clonal integration on plant competition for mosaic habitat space ' () 81 Ecology : 3291 -3304.

    • Search Google Scholar
  • Wijesinghe, D.K. and S.N. Handel. 1994. Advantages of clonal growth in heterogeneous habitats: an experiment with Potentilla simplex. J. Ecol. 82:495-502.

    'Advantages of clonal growth in heterogeneous habitats: an experiment with Potentilla simplex ' () 82 J. Ecol. : 495 -502.

    • Search Google Scholar
  • Wijesinghe, D.K. and M.J. Hutchings. 1997. The effects of spatial scale of environmental heterogeneity on the growth of a clonal plant: an experimental study with Glechoma hederacea. J. Ecol. 85:17-28.

    'The effects of spatial scale of environmental heterogeneity on the growth of a clonal plant: an experimental study with Glechoma hederacea ' () 85 J. Ecol. : 17 -28.

    • Search Google Scholar
  • Wijesinghe, D.K. and M.J. Hutchings. 1999. The effect of environmental heterogeneity on the performance of Glechoma hederacea: the interactions between patch contrast and patch scale. J. Ecol. 87:860-872.

    'The effect of environmental heterogeneity on the performance of Glechoma hederacea: the interactions between patch contrast and patch scale ' () 87 J. Ecol. : 860 -872.

    • Search Google Scholar