, Volume 19, Issue 7, pp 1240–1254 | Cite as

Herbaceous Understorey: An Overlooked Player in Forest Landscape Dynamics?

  • Timothy Thrippleton
  • Harald Bugmann
  • Kathrin Kramer-Priewasser
  • Rebecca S. Snell


Dense herbaceous understorey layers can impact tree regeneration and thereby affect forest succession. However, the implications of this interaction on large spatial and temporal scales are not well understood. To analyse the role of overstorey–understorey interactions for forest dynamics, we implemented an understorey layer (composed of the plant functional types grasses, forbs, ferns, herbs and shrubs) in the forest landscape model LandClim, focusing on competition for light as the main mode of interaction. The model was used to simulate post-disturbance dynamics over an elevational gradient of 560–2800 m a.s.l. in Central Europe. Simulation results showed strong impacts of the herbaceous understorey on tree regeneration within the first decades, but generally little effect on late-successional forests, i.e. not providing any evidence for ‘arrested’ succession. The results also demonstrated varying overstorey–understorey interactions across the landscape: strongest effects were found at low to mid elevations of the study landscapes, where tree establishment was substantially delayed. At high elevations, tree growth and establishment were more limited by low temperatures, and the effect of light competition from the understorey was negligible. Although the inclusion of large windthrow disturbances increased the biomass of herbaceous understorey across the landscape, this had only a small impact on the overstorey due to the presence of advance regeneration of trees. Overall, our results demonstrate that the herbaceous understorey can have a significant impact for forest landscape dynamics through light competition, and that non-woody plants should not be neglected in forest modelling.


herbaceous vegetation overstorey–understorey interaction arrested succession Black Forest Dischma valley central Alps dynamic vegetation model 



We gratefully acknowledge the support by Dominic Michel in all IT-related questions, as well as Laura Schuler and Nica Huber for helpful comments on the discussion section. Furthermore, two anonymous reviewers are gratefully acknowledged for providing helpful comments on an earlier version of the manuscript. Funding for R.S.S. was provided by the EU FP7 project “IMPRESSIONS”, Grant No. 603416.

Supplementary material

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Supplementary material 1 (DOCX 2665 kb)


  1. Alaback PB. 1982. Dynamics of understory biomass in Sitka spruce-western hemlock forests of southeast Alaska. Ecology 63:1932–48.CrossRefGoogle Scholar
  2. Alaback PB. 1984. Plant succession following logging in the Sitka spruce—western hemlock forests of southeast Alaska: implications for management. USDA Forest Service General Technical Report. Oregon State University.Google Scholar
  3. Allen CD. 2007. Interactions across spatial scales among forest dieback, fire, and erosion in northern New Mexico landscapes. Ecosystems 10:797–808.CrossRefGoogle Scholar
  4. Balandier P, Collet C, Miller JH, Reynolds PE, Zedaker SM. 2006. Designing forest vegetation management strategies based on the mechanisms and dynamics of crop tree competition by neighbouring vegetation. Forestry 79:3–27.CrossRefGoogle Scholar
  5. Bogenrieder A, Rasbach H. 1982. Der Feldberg im Schwarzwald: subalpine Insel im Mittelgebirge. Karlsruhe: Landesanstalt für Umweltschutz Baden-Württemberg; Institut für Oekologie und Naturschutz.Google Scholar
  6. Bugmann H. 2001. A review of forest gap models. Clim Change 51:259–305.CrossRefGoogle Scholar
  7. Coomes DA, Grubb PJ. 2000. Impacts of root competition in forests and woodlands: a theoretical framework and review of experiments. Ecol Monogr 70:171–207.CrossRefGoogle Scholar
  8. Dale VH, Crisafulli CM, Swanson FJ. 2005. 25 years of ecological change at Mount St. Helens. Science 308:961–2.CrossRefPubMedGoogle Scholar
  9. de la Cretaz AL, Kelty MJ. 1999. Establishment and control of hay-scented fern: a native invasive species. Biol Invasions 1:223–36.CrossRefGoogle Scholar
  10. de la Cretaz AL, Kelty MJ. 2002. Development of tree regeneration in fern-dominated forest understories after reduction of deer browsing. Restor Ecol 10:416–26.CrossRefGoogle Scholar
  11. De Long JR, Kardol P, Sundqvist MK, Veen GF, Wardle DA. 2015. Plant growth response to direct and indirect temperature effects varies by vegetation type and elevation in a subarctic tundra. Oikos 124:772–83.CrossRefGoogle Scholar
  12. DenOuden J. 2000. The role of bracken (Pteridium aquilinum) in forest dynamics. Dissertation for the degree of Doctor of Sciences, Ph.D. Netherlands: Wageningen University.Google Scholar
  13. Dolling AHU. 1996. Interference of bracken (Pteridium aquilinum L Kuhn) with Scots pine (Pinus sylvestris L) and Norway spruce (Picea abies L Karst) seedling establishment. For Ecol Manag 88:227–35.CrossRefGoogle Scholar
  14. Ellenberg H. 1996. Vegetation Mitteleuropas mit den Alpen in ökologischer, dynamischer und historischer Sicht. Stuttgart: Ulmer. p 1096p.Google Scholar
  15. Fischer A. 1999. Die Entwicklung von Wald-Biozönosen nach Sturmwurf. Weinheim: Wiley.CrossRefGoogle Scholar
  16. Frehner M, Wasser B, Schwitter R. 2005. Nachhaltigkeit im Schutzwald (Projekt NaiS) - Bundesamt für Umwelt BAFU. Bern.Google Scholar
  17. George LO, Bazzaz FA. 1999a. The fern understory as an ecological filter: emergence and establishment of canopy-tree seedlings. Ecology 80:833–45.CrossRefGoogle Scholar
  18. George LO, Bazzaz FA. 1999b. The fern understory as an ecological filter: growth and survival of canopy-tree seedlings. Ecology 80:846–56.CrossRefGoogle Scholar
  19. George LO, Bazzaz FA. 2003. The herbaceous layer as a filter determining spatial pattern in forest tree regeneration. In: Gilliam FSR, Eds. The herbaceous layer in forests of eastern North America. New York: Oxford University Press, pp 265–82.Google Scholar
  20. Gilliam FS. 2007. The ecological significance of the herbaceous layer in temperate forest ecosystems. Bioscience 57:845–58.CrossRefGoogle Scholar
  21. Hart SA, Chen HYH. 2006. Understory vegetation dynamics of North American boreal forests. Crit Rev Plant Sci 25:381–97.CrossRefGoogle Scholar
  22. Henne PD, Elkin C, Colombaroli D, Samartin S, Bugmann H, Heiri O, Tinner W. 2013. Impacts of changing climate and land use on vegetation dynamics in a Mediterranean ecosystem: insights from paleoecology and dynamic modeling. Landsc Ecol 28:819–33.CrossRefGoogle Scholar
  23. Hickler T, Vohland K, Feehan J, Miller PA, Smith B, Costa L, Giesecke T, Fronzek S, Carter TR, Cramer W, Kuhn I, Sykes MT. 2012. Projecting the future distribution of European potential natural vegetation zones with a generalized, tree species-based dynamic vegetation model. Glob Ecol Biogeogr 21:50–63.CrossRefGoogle Scholar
  24. Hiltbrunner E, Aerts R, Bühlmann T, Huss-Danell K, Magnusson B, Myrold DD, Reed SC, Sigurdsson BD, Körner C. 2014. Ecological consequences of the expansion of N-2-fixing plants in cold biomes. Oecologia 176:11–24.CrossRefPubMedGoogle Scholar
  25. Horsley S. 1993. Mechanisms of interference between hay-scented fern and black-cherry. Can J For Res 23:2059–69.CrossRefGoogle Scholar
  26. IPCC. 2013. Climate Change 2013: The Physical Science Basis. Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. In: Stocker TF, Qin D, Plattner G-K, Tignor M, Allen SK, Boschung J, Nauels A, Xia Y, Bex V, Midgley PM, Eds. Cambridge, United Kingdom and New York, NY, USA: Cambridge University Press.Google Scholar
  27. Jules MJ, Sawyer JO, Jules ES. 2008. Assessing the relationships between stand development and understory vegetation using a 420-year chronosequence. For Ecol Manag 255:2384–93.CrossRefGoogle Scholar
  28. Kellomäki S, Väisänen H. 1991. Application of a gap model for the simulation of forest ground vegetation in boreal conditions. For Ecol Manag 42:35–47.CrossRefGoogle Scholar
  29. Koop H, Hilgen P. 1987. Forest dynamics and regeneration mosaic shifts in unexploited beech (Fagus-Sylvatica) stands at Fontainebleau (France). For Ecol Manag 20:135–50.CrossRefGoogle Scholar
  30. Kowalenko CG. 2003. An evaluation of estimating and indexing methods to simplify the determination of management treatment effects on raspberry yields. Can J Plant Sci 83:141–7.CrossRefGoogle Scholar
  31. Kramer K, Brang P, Bachofen H, Bugmann H, Wohlgemuth T. 2014. Site factors are more important than salvage logging for tree regeneration after wind disturbance in Central European forests. For Ecol Manage 331:116–28.CrossRefGoogle Scholar
  32. Kupferschmid AD, Bugmann H. 2005. Effect of microsites, logs and ungulate browsing on Picea abies regeneration in a mountain forest. For Ecol Manag 205:251–65.CrossRefGoogle Scholar
  33. Kupferschmid AD, Wasem U, Bugmann H. 2015. Browsing regime and growth response of Abies alba saplings planted along light gradients. Eur J Forest Res 134:75–87.CrossRefGoogle Scholar
  34. Laurent JM, Bar-Hen A, Francois L, Ghislain M, Cheddadi R. 2004. Refining vegetation simulation models: From plant functional types to bioclimatic affinity groups of plants. J Veg Sci 15:739–46.CrossRefGoogle Scholar
  35. Li MH, Du Z, Pan HL, Yan CF, Xiao WF, Lei JP. 2012. Effects of neighboring woody plants on target trees with emphasis on effects of understorey shrubs on overstorey physiology in forest communities: a mini-review. Commun Ecol 13:117–28.CrossRefGoogle Scholar
  36. Lieffers V, Macdonald S, Hogg E. 1993. Ecology of and control strategies for Calamagrostis canadensis in boreal forest sites. Can J For Res 23:2070–7.CrossRefGoogle Scholar
  37. Maguire DA, Forman RTT. 1983. Herb cover effects on tree seedling patterns in a mature hemlock-hardwood forest. Ecology 64:1367–80.CrossRefGoogle Scholar
  38. Maher EL, Germino MJ, Hasselquist NJ. 2005. Interactive effects of tree and herb cover on survivorship, physiology, and microclimate of conifer seedlings at the alpine tree-line ecotone. Can J For Res 35:567–74.CrossRefGoogle Scholar
  39. McCarthy BC. 2003. The herbaceous layer of eastern old-growth deciduous forests. In: Gilliam FS, Roberts MR, Eds. The herbaceous layer in forests of eastern North America. New York: Oxford University Press, pp 163–76.Google Scholar
  40. McKenzie D, Raymond CL, Cushman SA. 2009. Modeling understory vegetation and its response to fire. In: Millspaugh JJ, Thompson FR, Eds. Models for planning wildlife conservation in large landscapes. San Diego: Academic Press, pp 391–414.Google Scholar
  41. Mladenoff DJ. 2004. LANDIS and forest landscape models. Ecol Model 180:7–19.CrossRefGoogle Scholar
  42. Monsi M, Saeki T. 2005. On the factor light in plant communities and its importance for matter production. Ann Bot 95:549–67.CrossRefPubMedPubMedCentralGoogle Scholar
  43. Nabuurs GJ. 1996. Quantification of herb layer dynamics under tree canopy. For Ecol Manag 88:143–8.CrossRefGoogle Scholar
  44. Niering W, Goodwin R. 1974. Creation of relatively stable shrublands with herbicides—arresting succession on rights-of-way and pastureland. Ecology 55:784–95.CrossRefGoogle Scholar
  45. Nieto-Lugilde D, Lenoir J, Abdulhak S, Aeschimann D, Dullinger S, Gegout JC, Guisan A, Pauli H, Renaud J, Theurillat JP, Thuiller W, Van Es J, Vittoz P, Willner W, Wohlgemuth T, Zimmermann NE, Svenning JC. 2015. Tree cover at fine and coarse spatial grains interacts with shade tolerance to shape plant species distributions across the Alps. Ecography 38:578–89.CrossRefPubMedPubMedCentralGoogle Scholar
  46. Nilsson M-C, Wardle DA. 2005. Understory vegetation as a forest ecosystem driver: evidence from the northern Swedish boreal forest. Front Ecol Environ 3:421–8.CrossRefGoogle Scholar
  47. Oliver C. 1981. Forest development in north-America following major disturbances. For Ecol Manag 3:153–68.CrossRefGoogle Scholar
  48. Ott E, Frehner M, Frey HU, Lüscher P. 1997. Gebirgsnadelwälder: ein praxisorientierter Leitfaden für eine standortgerechte Waldbehandlung. Bern: P. Haupt Verlag. p 288p.Google Scholar
  49. Pitman JI. 2000. Absorption of photosynthetically active radiation, radiation use efficiency and spectral reflectance of bracken [Pteridium aquilinum (L.) Kuhn] canopies. Ann Bot 85:101–11.Google Scholar
  50. Priewasser K. 2013. Factors influencing tree regeneration after windthrow in Swiss forests. Dissertation for the degree of Doctor of Sciences, Ph.D. Thesis No. 21011. ETH Zurich.Google Scholar
  51. Provendier D, Balandier P. 2008. Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica). Ann For Sci 65:510.CrossRefGoogle Scholar
  52. R Development Core Team. 2014. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, AustriaGoogle Scholar
  53. Raich JW, Russell AE, Vitousek PM. 1997. Primary productivity and ecosystem development along an elevational gradient on Mauna Loa, Hawai’i. Ecology 78:707–21.Google Scholar
  54. Royo AA, Carson WP. 2006. On the formation of dense understory layers in forests worldwide: consequences and implications for forest dynamics, biodiversity, and succession. Can J For Res 36:1345–62.CrossRefGoogle Scholar
  55. Royo AA, Carson WP. 2008. Direct and indirect effects of a dense understory on tree seedling recruitment in temperate forests: habitat-mediated predation versus competition. Can J For Res 38:1634–45.CrossRefGoogle Scholar
  56. Sato H, Itoh A, Kohyama T. 2007. SEIB-DGVM: A new dynamic global vegetation model using a spatially explicit individual-based approach. Ecol Model 200:279–307.CrossRefGoogle Scholar
  57. Sayer U, Reif A. 1999. Entwicklung der Vegetation im überregionalen Vergleich. Fischer A editor. Die Entwicklung von Wald-Biozonosen nach Sturmwurf. Weinheim: Wiley, pp 146–68.Google Scholar
  58. Schumacher S. 2004. The role of large-scale disturbances and climate for the dynamics of forested landscapes in the European Alps. Dissertation for the degree of Doctor of Sciences, Ph.D. Thesis No. 15573. ETH Zurich.Google Scholar
  59. Schumacher S, Bugmann H, Mladenoff DJ. 2004. Improving the formulation of tree growth and succession in a spatially explicit landscape model. Ecol Model 180:175–94.CrossRefGoogle Scholar
  60. Schumacher S, Reineking B, Sibold J, Bugmann H. 2006. Modeling the impact of climate and vegetation on fire regimes in mountain landscapes. Landsc Ecol 21:539–54.CrossRefGoogle Scholar
  61. Schwitter R, Sandri A, Bebi P, Wohlgemuth T, Brang P. 2015. Lehren aus Vivian für den Gebirgswald - im Hinblick auf den nächsten Sturm. Schweizerische Zeitschrift für Forstwesen 166:159–67.CrossRefGoogle Scholar
  62. Shropshire C, Wagner RG, Bell FW, Swanton CJ. 2001. Light attenuation by early successional plants of the boreal forest. Can J For Res 31:812–23.CrossRefGoogle Scholar
  63. Smith B, Prentice IC, Sykes MT. 2001. Representation of vegetation dynamics in the modelling of terrestrial ecosystems: comparing two contrasting approaches within European climate space. Glob Ecol Biogeogr 10:621–37.CrossRefGoogle Scholar
  64. Stromayer KAK, Warren RJ. 1997. Are overabundant deer herds in the eastern United States creating alternate stable states in forest plant communities? Wildl Soc Bull 25:227–34.Google Scholar
  65. Temperli C, Bugmann H, Elkin C. 2013. Cross-scale interactions among bark beetles, climate change, and wind disturbances: a landscape modeling approach. Ecol Monogr 83:383–402.CrossRefGoogle Scholar
  66. Thrippleton T, Dolos K, Perry GLW, Groeneveld J, Reineking B. 2014. Simulating long-term vegetation dynamics using a forest landscape model: the post-Taupo succession on Mt Hauhungatahi, North Island, New Zealand. N Z J Ecol 38:26–43.Google Scholar
  67. Turner MG, Dale VH, Everham EH. 1997. Fires, hurricanes, and volcanoes: comparing large disturbances. Bioscience 47:758–68.CrossRefGoogle Scholar
  68. Turner MG, Gardner RH, O’Neill RV. 2001. Landscape ecology in theory and practice: pattern and process. New York: Springer.Google Scholar
  69. Walker LR, Wardle DA, Bardgett RD, Clarkson BD. 2010. The use of chronosequences in studies of ecological succession and soil development. J Ecol 98:725–36.CrossRefGoogle Scholar
  70. Weisberg PJ, Bugmann H. 2003. Forest dynamics and ungulate herbivory: from leaf to landscape. For Ecol Manag 181:1–12.CrossRefGoogle Scholar
  71. Weisberg PJ, Hadorn C, Bugmann H. 2003. Predicting understorey vegetation cover from overstorey attributes in two temperate mountain forests. Forstwissenschaftliches Centralblatt 122:273–86.CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media New York 2016

Authors and Affiliations

  • Timothy Thrippleton
    • 1
  • Harald Bugmann
    • 1
  • Kathrin Kramer-Priewasser
    • 2
  • Rebecca S. Snell
    • 1
  1. 1.Department of Environmental Systems Science, Forest EcologySwiss Federal Institute of Technology, ETH ZurichZürichSwitzerland
  2. 2.Swiss Federal Research Institute WSLBirmensdorfSwitzerland

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