Plant Ecology

, Volume 135, Issue 2, pp 215–228 | Cite as

Species-environment relationships and vegetation patterns: effects of spatial scale and tree life-stage

  • Thomas J. Stohlgren
  • Richard R. Bachand
  • Yasuhiro Onami
  • Dan Binkley
Article

Abstract

Do relationships between species and environmental gradients strengthen or weaken with tree life-stage (i.e., small seedlings, large seedlings, saplings, and mature trees)? Strengthened relationships may lead to distinct forest type boundaries, or weakening connections could lead to gradual ecotones and heterogeneous forest landscapes. We quantified the changes in forest dominance (basal area of tree species by life-stage) and environmental factors (elevation, slope, aspect, intercepted photosynthetically active radiation (PAR), summer soil moisture, and soil depth and texture) across 14 forest ecotones (n = 584, 10 m #x00D7; 10 m plots) in Rocky Mountain National Park, Colorado, U.S.A. Local, ecotone-specific species-environment relationships, based on multiple regression techniques, generally strengthened from the small seedling stage (multiple R2 ranged from 0.00 to 0.26) to the tree stage (multiple R2 ranged from 0.20 to 0.61). At the landscape scale, combined canonical correspondence analysis (CCA) among species and for all tree life-stages suggested that the seedlings of most species became established in lower-elevation, drier sites than where mature trees of the same species dominated. However, conflicting evidence showed that species-environment relationships may weaken with tree life-stage. Seedlings were only found in a subset of plots (habitats) occupied by mature trees of the same species. At the landscape scale, CCA results showed that species-environment relationships weakened somewhat from the small seedling stage (86.4% of the variance explained by the first two axes) to the tree stage (76.6% of variance explained). The basal area of tree species co-occurring with Pinus contorta Doug. ex. Loud declined more gradually than P. contorta basal area declined across ecotones, resulting in less-distinct forest type boundaries. We conclude that broad, gradual ecotones and heterogeneous forest landscapes are created and maintained by: (1) sporadic establishment of seedlings in sub-optimal habitats; (2) survivorship of saplings and mature trees in a wider range of environmental conditions than seedlings presently endure; and (3) the longevity of trees and persistence of tree species in a broad range of soils, climates, and disturbance regimes.

Colorado Canonical correspondence analysis Ecotones Heterogeneous forest landscapes Rocky Mountain National Park 

Preview

Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.

References

  1. Allen, R. B., Peet, R. K. & Baker, W. L. 1991. Gradient analysis of latitudinal variation in southern Rocky Mountain forests. J. Biogeog. 18: 123-139.Google Scholar
  2. Barbour, M.G., Burk, J.H. & Pitts, W.D. 1987. Terrestrial plant ecology. Second edition. Benjamin/Cummings, Menlo Park, California.Google Scholar
  3. Clements, F. E. 1916. Plant succession: an analysis of the development of Vegetation. Publication 242. Carnegie Institution of Washington. Washington, D.C.Google Scholar
  4. Cochran, P. H. & Berntsen, C. M. 1973. Tolerance of lodgepole and ponderosa pine seedlings to low night temperatures. Forest Sci. 19: 272-280.Google Scholar
  5. Cornelius, J. M. & Reynolds, J. F. 1991. On determining the statistical significance of discontinuities within ordered ecological data. Ecology 72: 2057-2070.Google Scholar
  6. Day, R. J. 1972. Stand structure, succession, and use of Southern Alberta's Rocky Mountain forest. Ecology 53: 472-478.Google Scholar
  7. DeLucia, E. H. & Smith, W. K. 1987. Air and soil temperature limitations on photosynthesis in Engelmann spruce during summer. Can. J Forest Res. 17: 527-533.Google Scholar
  8. Dewey, D. R. & Lu, K. H. 1959. A correlation and path-coefficient analysis of components of crested wheatgrass seed production. Agron. J. 51: 515-518.Google Scholar
  9. Gee, G. W. & Bauder, J. W. 1986. Pp. 383-411. In: Klute, A. (ed.), Methods of soil analysis. Part 1, Physical and mineralogical methods. Second edition. American Society of Agronomy, Madison, Wisconsin.Google Scholar
  10. Gleason, H. A. 1926. The individualistic concept of the plant association. Bull. Torrey Bot. Club 53: 1-20.Google Scholar
  11. Good, R. E. 1931. A theory of plant geography. The New Phytol. 30: 149-203.Google Scholar
  12. Good, R. E. 1953. The geography of the flowering plants. Second edition. Longmans, Green, and Co., New York, New York.Google Scholar
  13. Gosz, J. R. 1993. Ecotone hierarchies. Ecol. Appl. 3: 369-376.Google Scholar
  14. Hadley, K. S. & Veblen, T. T. 1993. Stand response to western spruce budworm and Douglas-fir bark beetle outbreaks, Colorado front range. Can. J. Forest Res. 23: 479-491.Google Scholar
  15. Hansen, A. J. & di Castri, F. (eds). 1992. Landscape boundaries: consequences for biotic diversity and landscape flows. Ecological Studies 92. Springer-Verlag, New York, New York.Google Scholar
  16. Holland, M. M., Risser, P. G. & Naiman, R. J. (eds). 1991. Ecotones. Chapman and Hall, New York, New York.Google Scholar
  17. Knapp, A. K. & Smith, W. K. 1982. Factors influencing understory seedling establishment of Engelmann spruce (Picea engelmannii) and subalpine fir (Abies lasiocarpa) in southeast Wyoming. Can. J. Bot. 60: 2753-2761.Google Scholar
  18. Knight, D. H. 1994. Mountains and Plains: the Ecology of Wyoming Landscapes. Yale University Press, Thomson-Shore, Dexter, Michigan.Google Scholar
  19. Madole, R. F. 1976. Glacial geology of the front range, Colorado. Pp. 319-351. In: Mahaney, W. C. (ed.), Quaternary stratigraphy of North America. Dowden, Hutchinson, and Ross, Stroudsburg, Pennsylvania.Google Scholar
  20. Neter, J., Wasserman, W. & Kutner, M. H. 1990. Applied linear statistical models: regression, analysis of variance, and experimental designs. Third edition. Irwin, Homewood, Illinois.Google Scholar
  21. Palmer, M. W. 1993. Putting things in even better order: the advantages of canonical correspondence analysis. Ecology 74: 2215-2230.Google Scholar
  22. Peet, R. K. 1981. Forest vegetation of the Colorado front range. Vegetatio 45: 3-75.Google Scholar
  23. Peet, R. K. 1988. Forests of the Rocky Mountains. Pp. 63-101. In: Barbour, M. G. & Billings, W. D. (eds), North American terrestrial vegetation. Cambridge University Press, New York, New York.Google Scholar
  24. Pickett, S. T. A. 1989. Space-for-time substitutions as an alternative to long-term studies. Pp. 110-135. In: Likens, G. E. (ed.), Long-term studies in ecology. Springer-Verlag, New York, New York.Google Scholar
  25. Reed, R. A., Peet, R. K., Palmer, M. W. & White, P. S. 1993. Scale dependence of vegetation-environment correlations: a case study of a North Carolina piedmont woodland. J. Veg. Sci. 4: 329-340.Google Scholar
  26. Risser, P. G. 1993. Ecotones: ecotones at local to regional scales from around the world. Ecol. Appl. 3: 367-368.Google Scholar
  27. Risser, P. G. 1995. The status of the science of examining ecotones. BioScience 45: 318-325.Google Scholar
  28. Rusek, J. 1993. Air-pollution-mediated changes in alpine ecosystems and ecotones. Ecol. Appl. 3: 406-416.Google Scholar
  29. SAS Institute Inc. 1990. SAS version 6. Third edition. Cary, North Carolina.Google Scholar
  30. Shelford, V. E. 1913. Animal communities in temperate America. University of Chicago Press, Chicago, Illinois.Google Scholar
  31. Stohlgren, T. J. 1992. Resilience of a heavily logged grove of giant sequoia (Sequoiadendron giganteum) in Kings Canyon National Park, California. Forest Ecol. Manag. 54: 115-140.Google Scholar
  32. Stohlgren, T. J. 1993. Spatial patterns of giant sequoia (Sequoiadendron giganteum) in two sequoia groves in Sequoia Canyon National Park, California. Can. J. Forest Res. 23: 120-132.Google Scholar
  33. Stohlgren, T. J. 1997. The Rocky Mountains. In: National Status and Trends Report. U.S. Geological Survey, Biological Resources Division, Washington, D.C. (in press).Google Scholar
  34. Stohlgren, T. J. & Bachand, R. R. 1997. Lodgepole pine (Pinus contorta) ecotones in Rocky Mountain National Park, Colorado, USA. Ecology 78: 632-641.Google Scholar
  35. Stohlgren, T. J., Binkley, D., Veblen, T. T. & Baker, W. L. 1995. Attributes of landscape-scale, long-term studies: malpractice insurance for landscape ecologists. Env. Monit. Assessment 36: 1-25.Google Scholar
  36. Stohlgren, T. J., Chase, T. N., Pielke, R. A., Kittel, T. G. F. and Baron, J. 1998. Evidence that local land use practices influence regional climate, vegetation, and stream flow patterns in adjacent natural areas. Global Change Biology (in press).Google Scholar
  37. ter Braak, C. J. F. 1986. Canonical correspondence analysis: a new eigen vector technique for multivariate direct gradient analysis. Ecology 67: 1167-1179.Google Scholar
  38. ter Braak, C. J. F. 1987a. The analysis of vegetation-environment relationships by Canonical Correspondence Analysis. Vegetatio 69: 69-77.Google Scholar
  39. ter Braak, C. J. F. 1987b. CANOCO-a FORTRAN program for canonical community ordination by [partial] [detrended] [canonical] correspondence analysis, principal components analysis and redundancy analysis (version 2.1). TNO Institute of Applied Computer Science, Wageningen, The Netherlands.Google Scholar
  40. ter Braak, C. J. F. 1991. CANOCO version 3.12. Agricultural Mathematics Group, Wageningen, The Netherlands.Google Scholar
  41. Veblen, T. T., Hadley, K. S., Reid, M. S. & Rebertus, A. J. 1991. Stand response to spruce beetle outbreak in Colorado subalpine forests. Ecology 72: 213-231.Google Scholar
  42. Veblen, T. T. & Lorenz, D. C. 1991. The Colorado front range: a century of ecological change. University of Utah Press, Salt Lake City, Utah.Google Scholar
  43. Weisberg, P. J. & Baker, W. L. 1995. Spatial variation in tree regeneration in the forest-tundra ecotone, Rocky Mountain National Park, Colorado. Can. J. Forest Res. 25: 1326-1339.Google Scholar
  44. Wesser, S. D. & Armbruster, W. S. 1991. Species distribution controls across a forest-steppe transition: a causal model and experimental test. Ecol. Monog. 61: 323-342.Google Scholar
  45. Whittaker, R. H. 1967. Gradient analysis of vegetation. Biol. Rev. 42: 207-264.Google Scholar
  46. Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall Inc., Englewood Cliffs, New Jersey.Google Scholar

Copyright information

© Kluwer Academic Publishers 1998

Authors and Affiliations

  • Thomas J. Stohlgren
    • 1
  • Richard R. Bachand
    • 1
  • Yasuhiro Onami
    • 2
    • 4
  • Dan Binkley
    • 3
    • 4
  1. 1.Midcontinent Ecological Science Center, U.S. Geological SurveyColorado State UniversityFort CollinsUSA
  2. 2.Natural Resource Ecology LaboratoryColorado State UniversityFort CollinsUSA
  3. 3.Department of Forest SciencesColorado State UniversityFort CollinsUSA
  4. 4.Graduate Degree Program in EcologyColorado State UniversityFort CollinsUSA

Personalised recommendations