Community Ecology

, Volume 17, Issue 2, pp 205–215 | Cite as

Vegetative sprouting as an additional pathway for a seed size-number trade-off: a field-parameterised simulation approach

  • J. DoudaEmail author
  • J. Hulík
  • J. Doudová


Studies of perennial plants generally search for a seed size vs. seed number trade-off. Surprisingly, the fact that perennials may replace an investment in large seeds by the allocation to vegetative propagation has not yet been investigated as an additional pathway enabling species coexistence. We focused on the mechanisms of coexistence in Carex elata and C. elongata, two co-occurring clonal sedges dominant in European swamp alder forests. We asked the following questions: i) Is the number of germinated seeds a better predictor of species coexistence than the total number of seeds? ii) What recruitment conditions and competition rules determine vegetative sprouting to be an alternative to large, competitively superior seeds? We measured several species functional traits related to the colonisation and fitness of perennials. To examine the competitive hierarchy between species and microsite species preferences, we analysed the effects of environmental factors and plant densities on fitness-related traits using Structural Equation Modelling (SEM). Then, using a series of spatially explicit simulations partly parameterised based on the field measurement, we evaluated the importance of seed and ramet propagation and recruitment conditions for long-term species coexistence. SEM indicated a competitive hierarchy and a large overlap in microsite preferences between species. As a response to our initial questions we found that: i) Only differences in the numbers of germinated seeds, allowed the two species to coexist. If we consider only differences in the total number of seeds, the superior competitor (Carex elata) outcompeted the inferior competitor (C. elongata) in all scenarios. This is because the former produced about three-times as many seeds as the latter. ii) We show that vegetative sprouting represents an additional pathway for the seed size-number trade-off when the competitive superiority of species is attributed to vegetative propagation. This is another way that a species deals with the omnipresent seeds of other species. Taken together, our study demonstrates that differences in seed performance, coupled with differences in vegetative propagation related to competitive ability, are an additional mechanism allowing the coexistence of perennial plants.


Carex elata Carex elongata Competition-colonisation trade-off Functional traits Sedges Spatially explicit models Structural equation models Wetland forests 



Comparative Fit Index


Generalised Linear Mixed Model


Maximum Likelihood Method


Root Mean Square Error of Approximation Index


Structural Equation Modelling


Spatial Eigenvalue Vector Mapping


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

Supplementary material

42974_2016_1702205_MOESM1_ESM.pdf (106 kb)
Appendix 1 Fitness-related data measured in field study. C. elo, Carex elongata ; C. ela, C. elata .


  1. Abrahamson, W.A. 1980. Demography and vegetative reproduction. In: O.T. Solbrig (ed.), Demography and Evolution in Plant Populations. University of California Press, Los Angeles. pp. 89–106.Google Scholar
  2. Baskin, C.C. and Baskin, J.M. 2014. Seeds: Ecology, Biogeography, and Evolution of Dormancy and Germination. 2nd ed. Elsevier/ Academic Press, San Diego.Google Scholar
  3. Beatty, S.W. 1984. Influence of microtopography and canopy species on spatial patterns of forest understory plants. Ecology 65:1406–1419.CrossRefGoogle Scholar
  4. Ben-Hur, E., Fragman-Sapir, O., Hadas, R., Singer, A. and Kadmon, R. 2012. Functional trade-offs increase species diversity in experimental plant communities. Ecol. Lett. 15:1276–1282.CrossRefPubMedPubMedCentralGoogle Scholar
  5. Benot, M.-L., Bittebiere, A.-K., Ernoult, A., Clément, B. and Mony, C. 2013. Fine-scale spatial patterns in grassland communities depend on species clonal dispersal ability and interactions with neighbours. J. Ecol. 101:626–636.CrossRefGoogle Scholar
  6. Bierzychudek, P. 1982. Life histories and demography of shade-tolerant temperate forest herbs: a review. New Phytol. 90:757–776.CrossRefGoogle Scholar
  7. Bollen, K.A. and Long, J.S. 1993. Testing Structural Equation Models. Sage Publications, Newbury Park, CA.Google Scholar
  8. Bruelheide, H. and Udelhoven, P. 2005. Correspondence of the fine-scale spatial variation in soil chemistry and the herb layer vegetation in beech forests. For. Ecol. Manage. 210:205–223.CrossRefGoogle Scholar
  9. Bullock, J.M., Hill, B.C., Silvertown, J. and Sutton, M. 1995. Gap colonization as a source of grassland community change: effects of gap size and grazing on the rate and mode of colonization by different species. Oikos 72:273–282.CrossRefGoogle Scholar
  10. Cadotte, M.W. 2007. Concurrent niche and neutral processes in the competition-colonization model of species coexistence. Proc. R. Soc. B-Biol. Sci. 274:2739–2744.CrossRefGoogle Scholar
  11. Canham, C.D. 1988. An index for understory light levels in and around canopy gaps. Ecology 69:1634–1638.CrossRefGoogle Scholar
  12. 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.CrossRefPubMedPubMedCentralGoogle Scholar
  13. Cheplick, G.P. 1995. Life history trade-offs in Amphibromus scabrivalvis (Poaceae): allocation to clonal growth, storage, and cleis-togamous reproduction. Am. J. Bot. 82:621–629.CrossRefGoogle Scholar
  14. Cheplick, G.P. 1997. Responses to severe competitive stress in a clonal plant: differences between genotypes. Oikos 79:581–591.CrossRefGoogle Scholar
  15. Chesson, P. and Huntly, N. 1997. The roles of harsh and fluctuating conditions in the dynamics of ecological communities. Am. Nat. 150:519–553.CrossRefPubMedPubMedCentralGoogle Scholar
  16. Douda, J. 2010. The role of landscape configuration in plant composition of floodplain forests across different physiographic areas. J. Veg. Sci. 21:1110–1124.CrossRefGoogle Scholar
  17. Douda, J., Boublík, K., Slezák, M., Biurrun, I., Nociar, J., Havrdová, A., Doudová, J., Aćić, S., Brisse, H., Brunet, J., Chytrý, M., Claessens, H., Csiky, J., Didukh, Y., Dimopoulos, P., Dullinger, S., FitzPatrick, Ú., Guisan, A., Horchler, P.J., Hrivnák, R., Jandt, U., Kącki, Z., Kevey, B., Landucci, F., Lecomte, H., Lenoir, J., Paal, J., Paternoster, D., Pauli, H., Pielech, R., Rodwell, J.S., Roelandt, B., Svenning, J.-C., Šibík, J., Šilc, U., Škvorc, Ž., Tsiripidis, I., Tzonev, R.T., Wohlgemuth, T. and Zimmermann, N.E. 2016. Vegetation classification and biogeography of European floodplain forests and alder carrs. Appl. Veg. Sci. 19:147–163.CrossRefGoogle Scholar
  18. Douda, J., Doudová-Kochánková, J., Boublík, K. and Drašnarová, A. 2012. Plant species coexistence at local scale in temperate swamp forest: test of habitat heterogeneity hypothesis. Oecologia 169:523–534.CrossRefPubMedPubMedCentralGoogle Scholar
  19. Dormann, C.F., McPherson, J.M., Araújo, M.B., Bivand, R., Bolliger, J., Carl, G., Davies, R.G., Hirzel, A., Jetz, W., Kissling, W.D., Kühn, I., Ohlemüller, R., Peres-Neto, P.R., Reineking, B., Schröder, B., Schurr, F.M. and Wilson, R. 2007. Methods to account for spatial autocorrelation in the analysis of species distributional data: a review. Ecography 30:609–628.CrossRefGoogle Scholar
  20. Eckert, C.G. 2002. The loss of sex in clonal plants. Evol. Ecol. 15:501–520.CrossRefGoogle Scholar
  21. Egler, F.E. 1954. Vegetation science concepts I. Initial floristic composition, a factor in old-field vegetation development. Vegetatio 4:412–417.Google Scholar
  22. Elias, R.B. and Dias, E. 2009. The effects of landslides on the mountain vegetation of Flores Island, Azores. J. Veg. Sci. 20: 706–717.CrossRefGoogle Scholar
  23. Eriksson, O. 2005. Game theory provides no explanation for seed size variation in grasslands. Oecologia 144:98–105.CrossRefPubMedPubMedCentralGoogle Scholar
  24. Eriksson, O. 2011. Niche shifts and seed limitation as mechanisms behind seedling recruitment patterns in clonal plants. Preslia 83:301–314.Google Scholar
  25. Eriksson, O. and Jakobsson, A. 1998. Abundance, distribution and life histories of grassland plants: a comparative study of 81 species. J. Ecol. 86:922–933.CrossRefGoogle Scholar
  26. Fang, X., Yuan, J., Wang, G. and Zhao, Z. 2006. Fruit production of shrub, Caragana korshinskii, following above-ground partial shoot removal: mechanisms underlying compensation. Plant Ecol. 187:213–225.CrossRefGoogle Scholar
  27. Fischer, M. and van Kleunen, M. 2002. On the evolution of clonal plant life histories. Evol. Ecol. 15:565–582.CrossRefGoogle Scholar
  28. Fukami, T. 2004. Assembly history interacts with ecosystem size to influence species diversity. Ecology 85:3234–3242.CrossRefGoogle Scholar
  29. Fox, J.W. 2013. The intermediate disturbance hypothesis should be abandoned. Trends Ecol. Evol. 28:86–92.CrossRefPubMedPubMedCentralGoogle Scholar
  30. Frazer, G.W., Canham, C.D. and Lertzman, K.P. 1999. Gap Light Analyzer (GLA), Version 2.0: Imaging software to extract canopy structure and gap light transmission indices from true-colour fisheye photographs, users manual and program documentation. Burnaby, British Columbia, Canada: Simon Fraser University and Millbrook, New York, NY: the Institute of Ecosystem Studies.Google Scholar
  31. Geritz, S.A., van der Meijden, E. and Metz, J.A. 1999. Evolutionary dynamics of seed size and seedling competitive ability. Theor. Popul. Biol. 55:324–343.CrossRefPubMedPubMedCentralGoogle Scholar
  32. Gilbert, B. and Lechowicz, M.J. 2004. Neutrality, niches, and dispersal in a temperate forest understory. PNAS 101:7651–7656.CrossRefPubMedPubMedCentralGoogle Scholar
  33. Grace, J.B. 2006. Structural Equation Modeling and Natural Systems. Cambridge University Press, Cambridge.Google Scholar
  34. Grime, J.P. 1998. Benefits of plant diversity to ecosystems: immediate, filter and founder effects. J. Ecol. 86:902–910.CrossRefGoogle Scholar
  35. Grman, E. and Suding, K.N. 2010. Within-year soil legacies contribute to strong priority effects of exotics on native California grassland communities. Rest. Ecol. 18:664–670.CrossRefGoogle Scholar
  36. Guo, Q., Brown, J.H., Valone, T.J. and Kachman, S.D. 2000. Constraints of seed size on plant distribution and abundance. Ecology 81:2149–2155.CrossRefGoogle Scholar
  37. Handel, S.N. 1985. The intrusion of clonal growth patterns on plant breeding systems. Am. Nat. 125:367–384.CrossRefGoogle Scholar
  38. Harper, J.L. 1967. A Darwinian approach to plant ecology. J. Ecol. 55:247–270.CrossRefGoogle Scholar
  39. Herben, T., Nováková, Z., Klimešová, J. and Hrouda, L. 2012. Species traits and plant performance: functional trade-offs in a large set of species in a botanical garden. J. Ecol. 100:1522–1533.CrossRefGoogle Scholar
  40. Huston, M.A. 1979. A general hypothesis of species diversity. Am. Nat. 113:81–101.CrossRefGoogle Scholar
  41. Lande, R.1993. Risks of population extinction from demographic and environmental stochasticity and random catastrophes. Am. Nat. 142:911–927CrossRefPubMedPubMedCentralGoogle Scholar
  42. Lee, T.D. 1988. Patterns of fruit and seed production. In: J. Lovett Doust and L. Lovett Doust (eds.), Plant Reproductive Ecology: Patterns and Strategies. Oxford University Press, New York. pp. 179–202.Google Scholar
  43. Leishman, M.R. 2001. Does the seed size/number trade-off model determine plant community structure? An assessment of the model mechanisms and their generality. Oikos 93:294–302.CrossRefGoogle Scholar
  44. Meekins, J.F. and McCarthy, B.C. 2000. Responses of the biennial forest herb Alliaria petiolata to variation in population density, nutrient addition and light availability. J. Ecol. 88:447–463.CrossRefGoogle Scholar
  45. Muller-Landau, H.C. 2010. The tolerance-fecundity trade-off and the maintenance of diversity in seed size. PNAS 107:4242–4247.CrossRefPubMedPubMedCentralGoogle Scholar
  46. Platt, W.J. 1975. The colonization and formation of equilibrium plant species associations on badger disturbances in a tall-grass prairie. Ecol. Monog. 45:285–305.CrossRefGoogle Scholar
  47. Rangel, T.F., Diniz-Filho, J.A.F. and Bini, L.M. 2010. SAM: a comprehensive application for Spatial Analysis in Macroecology. Ecography 33:46–50.CrossRefGoogle Scholar
  48. R Core Team 2015. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing. Vienna, Austria: R Foundation for Statistical Computing.Google Scholar
  49. Rees, M. 1995. Community structure in sand dune annuals: is seed weight a key quantity? J. Ecol. 83:857–863.CrossRefGoogle Scholar
  50. Reynolds, H.L., Mittelbach, G.G., Darcy-Hall, T.L., Houseman, G.R. and Gross, K.L. 2007. No effect of varying soil resource heterogeneity on plant species richness in a low fertility grassland. J. Ecol. 95:723–733.CrossRefGoogle Scholar
  51. Rosseel, Y. 2012. lavaan: An R package for structural equation modeling. J. Stat. Softw. 48:1–36.CrossRefGoogle Scholar
  52. Scheller, R.M. and Mladenoff, D.J. 2002. Understory species patterns and diversity in old-growth and managed northern hardwood forests. Ecol. Appl. 12:1329–1343.CrossRefGoogle Scholar
  53. Schütz, W. 2000. Ecology of seed dormancy and germination in sedges (Carex). Perspect. Plant. Ecol. Evol. Syst. 3:67–89.CrossRefGoogle Scholar
  54. Schütz, W. and Rave, G. 2003. Variation in seed dormancy of the wetland sedge, Carex elongata, between populations and individuals in two consecutive years. Seed. Sci. Res. 13:315–322.CrossRefGoogle Scholar
  55. Shaffer, M. 1981. Minimum population sizes for species conservation. BioScience 31:131–134.CrossRefGoogle Scholar
  56. Shipley, B. 2004. Analysing the allometry of multiple interacting traits. Perspect. Plant. Ecol. Evol. Syst. 6:235–241.CrossRefGoogle Scholar
  57. Soetaert, K. and Herman, P.M.J. 2009. A Practical Guide to Ecological Modelling. Using R as a Simulation Platform. Springer, Dordrecht.Google Scholar
  58. Thompson, F.L. and Eckert, C.G. 2004. Trade-offs between sexual and clonal reproduction in an aquatic plant: experimental manipulations vs. phenotypic correlations. J. Evol. Biol. 17:581–592.CrossRefPubMedPubMedCentralGoogle Scholar
  59. Tilman, D. 1994. Competition and biodiversity in spatially structured habitats. Ecology 75:2–16.CrossRefGoogle Scholar
  60. van de Koppel, J. and Crain, C.M. 2006. Scale-dependent inhibition drives regular tussock spacing in a freshwater marsh. Am. Nat. 168:136–147.CrossRefGoogle Scholar
  61. van Drunen, W.E. and Dorken, M.E. 2012. Trade-offs between clonal and sexual reproduction in Sagittaria latifolia (Alismataceae) scale up to affect the fitness of entire clones. New Phytol. 196:606–616.CrossRefPubMedPubMedCentralGoogle Scholar
  62. Weiner, J., Campbell, L.G., Pino, J. and Echarte, L. 2009. The allometry of reproduction within plant populations. J. Ecol. 97:1220– 1233.CrossRefGoogle Scholar
  63. Weppler, T., Stoll, P. and Stöcklin, J. 2006. The relative importance of sexual and clonal reproduction for population growth in the long-lived alpine plant Geum reptans. J. Ecol. 94:869–879.CrossRefGoogle Scholar
  64. Wijesinghe, D.K., John, E.A. and Hutchings, M.J. 2005. Does pattern of soil resource heterogeneity determine plant community structure? An experimental investigation. J. Ecol. 93:99–112.CrossRefGoogle Scholar
  65. Williams, R.D., Quimby, Jr. P.C. and Frick, K.E. 1977. Intraspecific competition of purple nutsedge (Cyperus rotundus) under greenhouse conditions. Weed Sci. 25:477–481.CrossRefGoogle Scholar

Copyright information

© Akadémiai Kiadó, Budapest 2016

This article is distributed under the terms of the Creative Commons Attribution 4.0 International License (, which permits unrestricted use, distribution, and reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made.

Authors and Affiliations

  1. 1.Faculty of Environmental SciencesCzech University of Life Sciences PraguePraha 6 — SuchdolCzech Republic

Personalised recommendations