Plant Ecology

, Volume 213, Issue 6, pp 955–965 | Cite as

Microtopographic heterogeneity constrains alpine plant diversity, Glacier National Park, MT



Theoretical and empirical evidence exists for a positive relationship between environmental heterogeneity and species diversity. Alpine plant communities can exhibit exceptional diversity at a fine scale, which niche theory would suggest is the result of fine scale spatial heterogeneity of the environment. To test if species diversity of alpine plants is driven by environmental heterogeneity, we sampled vascular plant species composition, microtopography, and ground cover within 1 m2 plots with and without solifluction forms in Glacier National Park, MT. We analyzed the relationship between microtopographic heterogeneity and species richness at the plot and sub-plot scale with linear and quantile regression, respectively. Species richness does not differ between the plots varying in cover type. Species richness is negatively related to the fractal dimension (D) of the ground surface and non-vegetated ground cover within 1 m2 plots. At a finer scale, the standard deviation of elevation and slope appear to impose a limit on species richness such that more variable sub-plots have lower species richness. Contrary to our expectations, microtopographic heterogeneity does not promote the diversity of alpine plants. The negative relationship between topographic heterogeneity and species richness is contrary to the theoretical prediction that environmental heterogeneity generally results in greater species diversity. It is possible that microtopographic variability represents a measure of soil disturbance, which would be expected to have a negative effect on species diversity in alpine tundra due to its low productivity.


Alpine Diversity Hierarchical partitioning Microtopography Quantile regression Tundra 


  1. Aarssen LW, Schamp BS, Pither J (2006) Why are there so many small plants? Implications for species coexistence. J Ecol 94:569–580CrossRefGoogle Scholar
  2. Beatty SW (1984) Influence of microtopography and canopy species on spatial patterns of forest understory plants. Ecology 65:1406–1419CrossRefGoogle Scholar
  3. Bertness MD, Callaway R (1994) Positive interactions in communities. Trends Ecol Evol 9:191–193PubMedCrossRefGoogle Scholar
  4. Billings WD (1973) Arctic and alpine vegetations: similarities, differences, and susceptibility to disturbance. Bioscience 23:697–704CrossRefGoogle Scholar
  5. Billings WD, Bliss LC (1959) An alpine snowbank environment and its effects on vegetation, plant development, and productivity. Ecology 40:388–397CrossRefGoogle Scholar
  6. Billings WD, Mooney HA (1968) The ecology of arctic and alpine plants. Biol Rev 43:481–529CrossRefGoogle Scholar
  7. Boorman LA, Woodell SRJ (1966) The topograph, an instrument for measuring microtopography. Ecology 47:869–870CrossRefGoogle Scholar
  8. Bowman WD, Theodose TA, Schardt JC, Conant RT (1993) Constraints of nutrient availability on primary production in two alpine tundra communities. Ecology 74:2085–2097CrossRefGoogle Scholar
  9. Bratton SP (1976) Resource division in an understory herb community: responses to temporal and microtopographic gradients. Am Nat 110:679–693CrossRefGoogle Scholar
  10. Brown RL, Peet RK (2003) Diversity and invasibility of southern Appalachian plant communities. Ecology 84:32–39CrossRefGoogle Scholar
  11. Bruland GL, Richardson CJ (2005) Hydrologic, edaphic, and vegetative responses to microtopographic reestablishment in a restored wetland. Restor Ecol 13:515–523CrossRefGoogle Scholar
  12. Burnett MR, August PV, Brown JH, Killingbeck KT (1998) The influence of geomorphological heterogeneity on biodiversity. I. A patch-scale perspective. Conserv Biol 12:363–370CrossRefGoogle Scholar
  13. Burrough PA (1981) Fractal dimensions of landscapes and other environmental data. Nature 294:240–242CrossRefGoogle Scholar
  14. Butler DR, Malanson GP (1989) Periglacial patterned-ground, Waterton–Glacier International Peace Park, Canada and USA. Zeitschrift Fur Geomorphologie 33:43–57Google Scholar
  15. Butler DR, Malanson GP (1999) Site locations and characteristics of miniature patterned ground, eastern Glacier National Park, Montana, USA. Landform Analysis 2:45–49Google Scholar
  16. Butler DR, Malanson GP, Resler LM (2004) Turf-banked treads and risers, turf exfoliation, and possible relationships with advancing treeline. Catena 58:259–274CrossRefGoogle Scholar
  17. Cade BS, Noon BR (2003) A gentle introduction to quantile regression for ecologists. Front Ecol Environ 1:412–420CrossRefGoogle Scholar
  18. Cade BS, Terrell JW, Schroeder RL (1999) Estimating effects of limiting factors with regression quantiles. Ecology 80:311–323CrossRefGoogle Scholar
  19. Callaway RM, Brooker RW, Choler P, Kikividze Z, Lortie CJ, Michalet R, Paolini L, Pugnaire FI, Newingham B, Aschehoug ET, Armas C, Kikodze D, Cook BJ (2002) Positive interactions among alpine plants increase with stress. Nature 417:844–848PubMedCrossRefGoogle Scholar
  20. Chevan A, Sutherland M (1991) Hierarchical partitioning. Am Stat 45:90–96Google Scholar
  21. Choler P, Michalet R, Callaway RM (2001) Facilitation and competition on gradients in alpine plant communities. Ecology 82:3295–3308CrossRefGoogle Scholar
  22. Damm C (2001) A phytosociological study of Glacier National Park, Montana, U.S.A., with notes on the syntaxonomy of alpine vegetation in western North America. PhD Thesis, Georg-August-Universität zu Göttingen, GöttingenGoogle Scholar
  23. Fisk MC, Schmidt SK, Seastedt TR (1998) Topographic patterns of above- and belowground production and nitrogen cycling in alpine tundra. Ecology 79:2253–2266CrossRefGoogle Scholar
  24. Forbis TA (2003) Seedling demography in an alpine ecosystem. Am J Bot 90:1197–1206PubMedCrossRefGoogle Scholar
  25. Fox JF (1981) Intermediate levels of soil disturbance maximize alpine plant diversity. Nature 293:564–565CrossRefGoogle Scholar
  26. Gonzalez-Espinosa M, Rey-Benayas JM, Ramirez-Marcial N, Huston MA, Golicher D (2004) Tree diversity in the northern Neotropics: regional patterns in highly diverse Chiapas, Mexico. Ecography 27:741–756CrossRefGoogle Scholar
  27. Gotelli NJ, Colwell RK (2001) Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecol Lett 4:379–391CrossRefGoogle Scholar
  28. Gough L, Osenberg CW, Gross KL, Collins SL (2000) Fertilization effects on species density and primary productivity in herbaceous plant communities. Oikos 89:428–439CrossRefGoogle Scholar
  29. Hubbell SP (2001) The unified neutral theory of biodiversity and biogeography. Princeton University Press, PrincetonGoogle Scholar
  30. Hubbell SP (2006) Neutral theory and the evolution of ecological equivalence. Ecology 87:1387–1398PubMedCrossRefGoogle Scholar
  31. Hubbell SP, Foster RB (1986) Biology, chance, and history and the structure of tropical rain forest tree communities. In: Diamond J, Case TJ (eds) Community ecology. Harper & Row, New York, pp 314–329Google Scholar
  32. Huston MA (1994) Biological diversity: the coexistence of species on changing landscapes. Cambridge University Press, CambridgeGoogle Scholar
  33. Johnson PL, Billings WD (1962) The alpine vegetation of the Beartooth Plateau in relation to cryopedogenic processes and patterns. Ecol Monogr 32:105–135CrossRefGoogle Scholar
  34. Kikvidze Z, Pugnaire FI, Brooker RW, Choler P, Lortie CJ, Michalet R, Callaway RM (2005) Linking patterns and processes in alpine plant communities: a global study. Ecology 86:2047–2054CrossRefGoogle Scholar
  35. Klanderud K, Totland O (2005) Simulated climate change altered dominance hierarchies and diversity of an alpine biodiversity hotspot. Ecology 86:2047–2054CrossRefGoogle Scholar
  36. Klinkenberg B (1992) Fractals and morphometric measures: is there a relationship? Geomorphology 5:5–20CrossRefGoogle Scholar
  37. Körner C (1995) Alpine plant diversity: a global survey and functional interpretations. In: Chapin FS, Körner C (eds) Arctic and alpine biodiversity: patterns, causes and ecosystem consequences. Springer, New York, pp 45–62CrossRefGoogle Scholar
  38. Körner C (2002) Mountain biodiversity, its causes and function: an overview. In: Körner C, Spehn EM (eds) Mountain biodiversity: a global assessment. Parthenon, New York, pp 3–20Google Scholar
  39. Körner C (2003) Alpine plant life: functional plant ecology of high mountain ecosystems, 2nd edn. Springer, New YorkGoogle Scholar
  40. Lechowicz MJ, Bell G (1991) The ecology and genetics of fitness in forest plants. II. Microspatial heterogeneity of the edaphic environment. J Ecol 79:687–696CrossRefGoogle Scholar
  41. Lesica P (2002) Flora of glacier national park. Oregon State University Press, CorvallisGoogle Scholar
  42. Loffler J, Pape R (2008) Diversity patterns in relation to the environment in alpine tundra ecosystems of northern Norway. Arct Antarct Alp Res 40:373–381CrossRefGoogle Scholar
  43. Loneragan WA, del Moral R (1984) The influence of microrelief on community structure of subalpine meadows. Bull Torrey Bot Club 111:209–216CrossRefGoogle Scholar
  44. Lundholm JT (2009) Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses. J Veg Sci 20:377–391CrossRefGoogle Scholar
  45. Lundholm JT, Larson DW (2003) Relationships between spatial environmental heterogeneity and plant species diversity on a limestone pavement. Ecography 26:715–722CrossRefGoogle Scholar
  46. Mac Nally R (2000) Regression and model-building in conservation biology, biogeography, and ecology: the distinction between—and reconciliation of—‘predictive’ and ‘explanatory’ models. Biodivers Conserv 9:655–671CrossRefGoogle Scholar
  47. Mac Nally R (2002) Multiple regression and inference in ecology and conservation biology: further comments on identifying important predictor variables. Biodivers Conserv 11:1387–1401CrossRefGoogle Scholar
  48. Malanson GP, Butler DR, Cairns DM, Welsh TE, Resler LM (2002) Variability in an edaphic indicator in alpine tundra. Catena 49:203–215CrossRefGoogle Scholar
  49. Malanson GP, Rose JP, Schroeder PJ, Fagre DB (2011) Contexts for change in alpine tundra. Phys Geogr 32:97–113CrossRefGoogle Scholar
  50. Malanson GP, Bengtson L, Fagre D (2012) Geomorphic determinants of species composition of alpine tundra, Glacier National Park, USA. Arctic Arct Alp Res. (in press)Google Scholar
  51. Marchand PJ, Roach DA (1980) Reproductive strategies of pioneering alpine species: seed production, dispersal, and germination. Arct Alp Res 12:137–146CrossRefGoogle Scholar
  52. Matsuoka N (2001) Solifluction rates, processes and landforms: a global review. Earth Sci Rev 55:107–134CrossRefGoogle Scholar
  53. McGill BJ, Maurer BA, Weiser MD (2006) Empirical evaluation of neutral theory. Ecology 87:1411–1423PubMedCrossRefGoogle Scholar
  54. Miriti MN (2006) Ontogenetic shift from facilitation to competition in a desert shrub. J Ecol 94:973–979CrossRefGoogle Scholar
  55. Nichols WF, Killingbeck KT, August PV (1998) The influence of geomorphological heterogeneity on biodiversity: II. A landscape perspective. Conserv Biol 12:371–379CrossRefGoogle Scholar
  56. Oksanen J (1996) Is the humped relationship between species richness and biomass an artifact due to plot size? J Ecol 84:293–295CrossRefGoogle Scholar
  57. Olofsson J, Shams H (2007) Determinants of plant species richness in an alpine meadow. J Ecol 95:916–925CrossRefGoogle Scholar
  58. Onipchenko VG, Blinnikov MS, Gerasimova MA, Volkova EV, Cornelissen JHC (2009) Experimental comparison of competition and facilitation in alpine communities varying in productivity. J Veg Sci 20:718–727CrossRefGoogle Scholar
  59. Pellissier L, Fournier B, Guisan A, Vittoz P (2010) Plant traits co-vary with altitude in grasslands and forests in the European Alps. Plant Ecol 211:351–365CrossRefGoogle Scholar
  60. Perry GLW, Groeneveld J, Miller BP (2010) Development, application and place of neutral theory in ecology and biogeography. Geography Compass 4:904–918CrossRefGoogle Scholar
  61. Ricklefs RE (2006) The unified neutral theory of biodiversity: do the numbers add up? Ecology 87:1424–1431PubMedCrossRefGoogle Scholar
  62. Ryel RJ, Caldwell MM, Manwaring JH (1996) Temporal dynamics of soil spatial heterogeneity in sage-brush-wheatgrass steppe during a growing season. Plant Soil 184:299–309CrossRefGoogle Scholar
  63. Scharf FS, Juanes F, Sutherland M (1998) Inferring ecological relationships from the edges of scatter diagrams: comparison of regression techniques. Ecology 79:448–460CrossRefGoogle Scholar
  64. Schmitz M, Platt W, DeCoster J (2002) Substrate heterogeneity and number of plant species in Everglades savannas (Florida, USA). Plant Ecol 160:137–148CrossRefGoogle Scholar
  65. Shmida A, Ellner S (1984) Coexistence of plant species with similar niches. Vegetatio 58:29–55Google Scholar
  66. Stöcklin J, Bäumler E (1996) Seed rain, seedling establishment and clonal growth strategies on a glacier foreland. J Veg Sci 7:45–56CrossRefGoogle Scholar
  67. Theodose TA, Bowman WD (1997) Nutrient availability, plant abundance, and species diversity in two alpine tundra communities. Ecology 78:1861–1872CrossRefGoogle Scholar
  68. Vivian-Smith G (1997) Microtopographic heterogeneity and floristic diversity in experimental wetland communities. J Ecol 85:71–82CrossRefGoogle Scholar
  69. Walker DA, Halfpenny JC, Walker MD, Wessman CA (1993) Long-term studies of snow-vegetation interactions. Bioscience 43:287–301CrossRefGoogle Scholar
  70. Walker MD, Webber PJ, Arnold EH, Ebert-May D (1994) Effects of interannual climate variation on aboveground phytomass in alpine vegetation. Ecology 75:393–408CrossRefGoogle Scholar
  71. Walsh C, Mac Nally R (2003) The hier.part package. Hierarchical partitioning. R project for statistical computing. URL: Accessed 1 Feb 2010
  72. Zak DR, Tilman D, Parmenter RR, Rice CW, Fisher FM, Vose J, Milchunas D, Martin CW (1994) Plant production and soil microorganisms in late-successional ecosystems: a continental-scale study. Ecology 75:2333–2347CrossRefGoogle Scholar
  73. Zedler JB, Zedler PH (1969) Association of species and their relationship to microtopography within old fields. Ecology 50:432–442CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media B.V. 2012

Authors and Affiliations

  1. 1.Department of Wildlife, Fish, & Conservation BiologyUniversity of California, DavisDavisUSA
  2. 2.Department of GeographyUniversity of IowaIowa CityUSA

Personalised recommendations