Folia Geobotanica

, Volume 41, Issue 1, pp 95–106 | Cite as

Demographic response to shading and defoliation in two woodland orchids

  • Richard P. Shefferson
  • Tiiu Kull
  • Kadri Tali


Many woodland herbs are long-lived, clonal geophytes that have evolved life histories favoring survival over reproduction. We examined the life history responses of natural populations of two woodland orchid species,Cypripedium calceolus andCephalanthera longifolia to defoliation and heavy shading conducted early in the growing seasons of 2002 and 2003. We asked whether, in view of the importance of growth for the survival of geophytes, treated plants were more likely to exhibit reduced flowering than reduced vegetative growth in the seasons following treatment. We also asked whether plants would suffer reduced ramet performance. Both treatments led to significant declines in flower number per ramet, number of leaves per ramet, and mean ramet height relative to controls inCypripedium. However, inCephalanthera, only shaded plants exhibited significant declines in flower number per ramet, and only defoliated plants exhibited declines in mean ramet height. The number of ramets per plant did not decline relative to controls in either species. Thus, these orchids, especiallyCypripedium, appeared to allocate resources preferentially to vegetative growth functions over sexual reproduction. Per-plant variation in leaf and flower number per ramet, as well as in mean ramet height, consistently declined in response to treatment, significantly so in the case of mean ramet height, suggesting that ramets became more similar within genets. These results suggest both similarities and differences in the ways in whichCephalanthera andCypripedium mobilize resources in response to stress.


Cephalanthera longifolia Cypripedium calceolus Geophytes Life history trade-offs Perennials Orchidaceae Resource allocation 


Tutin T.G. et al. (1968–1993) 


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  1. Amundsen T. &Slagsvold T. (1998): Hatching asynchrony in great tits: a bet-hedging strategy?Ecology 79: 295–304.CrossRefGoogle Scholar
  2. Baltzer J.L., Reekie E.G., Hewlin H.L., Taylor P.D. &Boates J.S. (2002): Impact of flower harvesting on the salt marsh plantLimonium carolinianum.Canad. J. Bot. 80: 841–851.CrossRefGoogle Scholar
  3. Callaghan T.V., Carlsson B.Å., Jónsdóttir I.S., Svensson B.M. &Jonasson S. (1992): Clonal plants and environmental change: introduction to the proceedings and summary.Oikos 63: 341–347.CrossRefGoogle Scholar
  4. Chesson P. &Peterson A.G. (2002): The quantitative assessment of the benefits of physiological integration in clonal plants.Evol. Ecol. Res. 4: 1153–1176.Google Scholar
  5. Collins B.S. &Pickett S.T.A. (1988a): Demographic responses of herb layer species to experimental canopy gaps in a northern hardwoods forest.J. Ecol. 76: 437–450.CrossRefGoogle Scholar
  6. Collins B.S. &Pickett S.T.A. (1988b): Response of herb layer cover to experimental canopy gaps.Amer. Midl. Naturalist 119: 282–290.CrossRefGoogle Scholar
  7. Curtis J.T. (1943): Germination and seedling development in five species ofCypripedium LAmer. J. Bot. 30: 199–206.CrossRefGoogle Scholar
  8. Curtis J.T. (1954): Annual fluctuation in rate of flower production by native Cypripediums during two decades.Bull. Torrey Bot. Club 81: 340–352.CrossRefGoogle Scholar
  9. Curtis J.T. (1959):The vegetation of Wisconsin: an ordination of plant communities. University of Wisconsin Press, Madison.Google Scholar
  10. Dafni A. &Ivri Y. (1981): The flower biology ofCephalanthera longifolia (Orchidaceae) - pollen limitation and facultative floral mimicry.Pl. Syst. Evol. 137: 229–240.CrossRefGoogle Scholar
  11. de Jong G. (1993): Covariances between traits deriving from successive allocations of a resource.Funct. Ecol. 7: 75–83.CrossRefGoogle Scholar
  12. de Kroon H., Whigham D.F. &Watson M.A. (1991): Developmental ecology of mayapple: effects of rhizome severing, fertilization and timing of shoot senescence.Funct. Ecol. 5: 360–368.CrossRefGoogle Scholar
  13. Derner J.D. &Briske D.D. (1998): An isotopic (N-15) assessment of intraclonal regulation in C-4 perennial grasses: ramet interdependence, independence or both?J. Ecol. 86: 305–314.CrossRefGoogle Scholar
  14. Fuller A.M. (1933): Studies on the flora of Wisconsin. Part I: The orchids;Orchidaceae. Bull. Public Mus. Milwaukee 14: 1–284.Google Scholar
  15. Gaillard J.-M. &Yoccoz N.G. (2003): Temporal variation in survival of mammals: a case of environmental canalization?Ecology 84: 3294–3306.CrossRefGoogle Scholar
  16. Garcia M.B. &Ehrlén J. (2002): Reproductive effort and herbivory timing in a perennial herb: fitness components at the individual and population levels.Amer. J. Bot. 89: 1295–1302.Google Scholar
  17. Glazier D.S. (2002): Resource-allocation rules and the heritability of traits.Evolution 56: 1696–1700.PubMedGoogle Scholar
  18. Hutchings M. (1999): Clonal plants as cooperative systems: benefits in heterogeneous environments.Pl. Spec. Biol. 14: 1–10.CrossRefGoogle Scholar
  19. Kull T. (1987): Population ecology ofCypripedium calceolus L. In:Laasimer L. &Kull T. (eds.),The plant cover of the Estonian SSR: flora, vegetation, and ecology, Valgus, Tallinn, pp. 77–83.Google Scholar
  20. Kull T. (1999):Cypripedium calceolus L.J. Ecol. 87: 913–924.CrossRefGoogle Scholar
  21. Kull T. (2002): Population dynamics of north temperate orchids. In:Kull T. &Arditti J. (eds.),Orchid biology, reviews and perspecitves 8, Kluwer Academic Publishers, Dordrecht, pp. 139–165.Google Scholar
  22. Kull T. &Kull K. (1991): Preliminary results from a study of populations ofCypripedium calceolus in Estonia. In:Wells T.C.E. &Willems J.H. (eds.),Population ecology of terrestrial orchids, SPB Academic Publishing, The Hague, pp. 69–76.Google Scholar
  23. Kull T. &Tuulik T. (1994): Orchid studies on permanent plots. In:Kull T. (eds.),Orchid ecology and protection in Estonia, Tartu University Press, Tartu, pp. 35–42Google Scholar
  24. Lesica P. &Steele B.M. (1994): Prolonged dormancy in vascular plants and implications for monitoring studies.Natural Areas J. 14: 209–212.Google Scholar
  25. Li R., Werger M.J.A., de Kroon H., During H.J. &Zhong Z.C. (2000): Interactions between shoot age structure, nutrient availability and physiological integration in the giant bambooPhyllostachys pubescens.Pl. Biol. 2: 437–446.CrossRefGoogle Scholar
  26. Marshall C. &Price E.A.C. (1997): Sectoriality and its implications for physiological integration. In:de Kroon H. &van Groenendael J. (eds.),The ecology and evolution of clonal plants, Backhuys Publishers, Leiden, pp. 79–107.Google Scholar
  27. McKendrick S.L. (1996): The effects of shading on seedlings ofOrchis morio andDactylorhiza fuchsii in chalk and clay soil.New Phytol. 134: 343–352.CrossRefGoogle Scholar
  28. Metzger F. &Schultz J. (1984): Understory response to 50 years of management of a northern hardwood forest in upper Michigan.Amer. Midl. Naturalist 112: 209–223.CrossRefGoogle Scholar
  29. Primack R.B., Miao S.L. &Becker K.R. (1994): Costs of reproduction in the pink lady’s slipper orchid (Cypripedium acaule): defoliation, increased fruit production, and fire.Amer. J. Bot. 81: 1083–1090.CrossRefGoogle Scholar
  30. Reekie E.G., Budge S. &Baltzer J.L. (2002): The shape of the trade-off function between reproduction and future performance inPlantago major andPlantago rugelii.Canad. J. Bot. 80: 140–150.CrossRefGoogle Scholar
  31. Ruiz N., Ward D. &Saltz D. (2002): Responses ofPancratium sickenbergeri to simulated bulb herbivory: combining defence and tolerance strategies.J. Ecol. 90: 472–479.CrossRefGoogle Scholar
  32. Sather B.-E. &Bakke O. (2000): Avian life history variation and contribution of demographic traits to the population growth rate.Ecology 81: 642–653.Google Scholar
  33. Shefferson R.P., Sandercock B.K., Proper J. &Beissinger S.R. (2001): Estimating dormancy and survival of a rare herbaceous perennial using mark-recapture models.Ecology 82: 145–156.Google Scholar
  34. Silvertown J., Franco M., Pisanty I. &Mendoza A. (1993): Comparative plant demography — relative importance of life-cycle components to the finite rate of increase in woody and herbaceous perennials.J. Ecol. 81: 465–476.CrossRefGoogle Scholar
  35. Slatkin M. (1974): Hedging one’s evolutionary bets.Nature 250: 704–705.Google Scholar
  36. Stancato G.C., Mazzafera P., &Buckeridge M.S. (2001): Effect of a drought period on the mobilisation of non-structural carbohydrates, photosynthetic efficiency and water status in an epiphytic orchid.Pl. Physiol. Biochem. 39: 1009–1016.CrossRefGoogle Scholar
  37. Stowe K.A., Marquis R.J., Hochwender C.G. &Simms E.L. (2000): The evolutionary ecology of tolerance to consumer damage.Annual Rev. Ecol. Syst. 31: 565–595.CrossRefGoogle Scholar
  38. Stuefer J.F., de Kroon H. &During H.J. (1996): Exploitation of environmental heterogeneity by spatial division of labour in a clonal plant.Funct. Ecol. 10: 328–334.CrossRefGoogle Scholar
  39. Summerhayes V.S. (1968):Wild orchids of Britain. Collins, London.Google Scholar
  40. Sutherland W.J. &Stillman R.A. (1988): The foraging tactics of plants.Oikos 52: 239–244.CrossRefGoogle Scholar
  41. Tutin T.G. et al. (eds.) (1968–1993):Flora europaea Ed. 1, 2. Cambridge University Press, Cambridge.Google Scholar
  42. van der Meijden E., de Boer N.J. &van der Veen-van WijkC.A.M. (2000): Pattern of storage and regrowth in ragwort.Evol. Ecol. 14: 439–455.CrossRefGoogle Scholar
  43. Whigham D. &Chapa A. (1999): Timing and intensity of herbivory: its influence on the performance of clonal woodland herbs.Pl. Spec. Biol. 14: 29–37.CrossRefGoogle Scholar
  44. Whigham D.F. (2004): Ecology of woodland herbs in temperate deciduous forests.Annual Rev. Ecol. Evol. Syst. 35: 583–621.CrossRefGoogle Scholar
  45. Willems J.H., Balounová Z. &Kindlmann P. (2001): The effect of experimental shading on seed production and plant survival ofSpiranthes spiralis (Orchidaceae).Lindleyana 16: 31–37.Google Scholar
  46. World Conservation Union (1963):The Convention on International Trade in Endangered Species of Wild Fauna and Flora. The World Conservation Union (IUCN), Gland.Google Scholar

Copyright information

© Institute of Botany, Academy of Sciences of the Czech Republic 2006

Authors and Affiliations

  1. 1.Department of Integrative Biology, 3060 VLSB #3140University of CaliforniaBerkeleyUSA
  2. 2.Institute of Zoology and BotanyEstonian Agricultural UniversityTartuEstonia

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