Plant and Soil

, Volume 365, Issue 1–2, pp 127–140 | Cite as

Spatial patterns of total and available N and P at alpine treeline

  • Daniel Liptzin
  • Robert L. SanfordJr.
  • Timothy R. Seastedt
Regular Article


Background and aims

Vegetation can have direct and indirect effects on soil nutrients. To test the effects of trees on soils, we examined the patterns of soil nutrients and nutrient ratios at two spatial scales: at sites spanning the alpine tundra/subalpine forest ecotone (ecotone scale), and beneath and beyond individual tree canopies within the transitional krummholz zone (tree scale).


Soils were collected and analyzed for total carbon (C), nitrogen (N), and phosphorus (P) as well as available N and P on Niwot Ridge in the Colorado Rocky Mountains.


Total C, N, and P were higher in the krummholz zone than the forest or tundra. Available P was also greatest in the krummholz zone while available N increased from the forest to the tundra. Throughout the krummholz zone, total soil nutrients and available P were higher downwind compared to upwind of trees.


The krummholz zone in general, and downwind of krummholz trees in particular, are zones of nutrient accumulation. This pattern indicates that the indirect effects of trees on soils are more important than the direct effects. The higher N:P ratios in the tundra suggest nutrient dynamics differ from the lower elevation sites. We propose that evaluating soil N and P simultaneously in soils may provide a robust assay of ecosystem nutrient limitation.


Treeline Nitrogen Phosphorus fractions Biosequence Nutrient ratios 



Thanks to Nate Wojcik, Carrie Renaud, and Sheena Anderson for help with sampling and with laboratory analyses at the University of Denver. We appreciate the comments from Alan Townsend on an early version of this manuscript as well as providing unpublished data on phosphorus concentrations. Support was provided by National Science Foundation grants to support the Niwot Ridge Long-Term Ecological Research program. In addition, DL was supported by grants from the University of Colorado, the Colorado Mountain Club, and the Marr Foundation as well as a National Science Foundation Doctoral Dissertation grant.


  1. Arthur MA, Fahey TJ (1992) Biomass and nutrients in a Picea Englemannii/Abies lasiocarpa forest in north-central Colorado: pools, annual production, and nutrient cycling. Can J For Res 22:315–325CrossRefGoogle Scholar
  2. Baron JS (2006) Hindcasting nitrogen deposition to determine an ecological critical load. Ecol Appl 16:433–439PubMedCrossRefGoogle Scholar
  3. Baron JS, Ojima DS, Holland EA, Parton WJ (1994) Analysis of nitrogen saturation potential in Rocky Mountain tundra and forest: implications for aquatic systems. Biogeochemistry 27:61–82CrossRefGoogle Scholar
  4. Barrick KA, Schoettle AW (1996) A comparison of the foliar nutrient status of elfinwood and symmetrically formed tall trees, Colorado Front Range, USA. Can J Bot 74:1461–1475CrossRefGoogle Scholar
  5. Barry RG (1973) A climatological transect on the east slope of the Front Range, Colorado. Arctic Alpine Res 5:89–110CrossRefGoogle Scholar
  6. Bedford BL, Walbridge MR, Aldous A (1999) Patterns in nutrient availability and plant diversity of temperate North American wetlands. Ecology 80:2151–2169CrossRefGoogle Scholar
  7. Bonanza Creek LTER database (2006) Accessed 9/24/2006
  8. BOREAS database (2006) Accessed 7/14/2006
  9. Bowman WD (1992) Inputs and storage of nitrogen in winter snowpack in an alpine ecosystem. Arctic Alpine Res 24:211–215CrossRefGoogle Scholar
  10. Bowman WD (1994) Accumulation and use of nitrogen and phosphorus following fertilization in 2 alpine tundra communities. Oikos 70:261–270CrossRefGoogle Scholar
  11. Bowman WD, Theodose TA, Schardt JC, Conant RT (1993) Constraints of nutrient availability on primary production in 2 Alpine Tundra communities. Ecology 74:2085–2097CrossRefGoogle Scholar
  12. Burns SF (1980) Alpine soil distribution and development, Indian Peaks, Colorado Front Range. Ph.D. University of Colorado, BoulderGoogle Scholar
  13. Cairns DM (1999) Multi-scale analysis of soil nutrients at alpine treeline in Glacier National Park, Montana. Phys Geogr 20:256–271Google Scholar
  14. Cleveland CC, Liptzin D (2007) C:N:P stoichiometry in soil: is there a “Redfield ratio” for the microbial biomass? Biogeochemistry 85:235–252CrossRefGoogle Scholar
  15. Crews TE, Kitayama K, Fownes JH, Riley RH, Herbert DA, Muellerdombois D, Vitousek PM (1995) Changes in soil–phosphorus fractions and ecosystem dynamics across a long chronosequence in Hawaii. Ecology 76:1407–1424CrossRefGoogle Scholar
  16. Cross AF, Schlesinger WH (1995) A literature-review and evaluation of the Hedley fractionation—applications to the biogeochemical cycle of soil–phosphorus in natural ecosystems. Geoderma 64:197–214CrossRefGoogle Scholar
  17. Daly C (1984) Snow distribution patterns in the alpine Krummholz zone. Prog Phys Geogr 8:157–175CrossRefGoogle Scholar
  18. Daubenmire R (1954) Alpine timberlines in the Americas and their interpretation. Butler Univ Bot Stud 11:119–136Google Scholar
  19. Elliott GP, Kipfmueller KF (2011) Multi-scale influences of climate on upper treeline dynamics in the Southern Rocky Mountains, USA: evidence of intra-regional variability and bioclimatic thresholds in response to 20th century climate. Ann Assoc Am Geogr 101:1181–1203CrossRefGoogle Scholar
  20. Elser JJ, Bracken ME, Cleland EE, Gruner DS, Harpole WS, Hillebrand H, Ngai JT, Seabloom EW, Shurin JB, Smith JE (2007) Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine and terrestrial ecosystems. Ecol Lett 10:1135–1142PubMedCrossRefGoogle Scholar
  21. Gable DJ, Madole RF (1976) Geologic map of the Ward quadrangle, Boulder County, Colorado. Map GQ-1277, US Geological Survey, Washington, D.C.Google Scholar
  22. Gannett H (1899) The timber-line. Am Geogr Soc J 31:118–122Google Scholar
  23. Gold WG, Glew KA, Dickson LG (2001) Functional influences of cryptobiotic surface crusts in an alpine tundra basin of the Olympic Mountains, Washington, USA. Northwest Sci 75:315–326Google Scholar
  24. Harrington RA, Fownes JH, Vitousek PM (2001) Production and resource use efficiencies in N- and P-limited tropical forests: a comparison of responses to long-term fertilization. Ecosystems 4:646–657CrossRefGoogle Scholar
  25. Hedin LO (2004) Global organization of terrestrial plant–nutrient interactions. Proc Natl Acad Sci U S A 101:10849–10850PubMedCrossRefGoogle Scholar
  26. Hedin LO, Vitousek PM, Matson PA (2003) Nutrient losses over four million years of tropical forest development. Ecology 84:2231–2255CrossRefGoogle Scholar
  27. Hedley MJ, Stewart JWB, Chauhan BS (1982) Changes in inorganic and organic soil phosphorus fractions induced by cultivation practices and by laboratory incubations. Soil Sci Soc Am J 46:970–976CrossRefGoogle Scholar
  28. Hobbie SE, Gough L (2002) Foliar and soil nutrients in tundra on glacial landscapes of contrasting ages in northern Alaska. Oecologia 131:453–462CrossRefGoogle Scholar
  29. Holtmeier FK, Broll G (1992) The influence of tree islands and microtopography on pedoecological conditions in the forest-alpine tundra ecotone on Niwot Ridge, Colorado Front Range, USA. Arctic Alpine Res 24:216–228CrossRefGoogle Scholar
  30. Jahren AH (2004) Factors of soil formation: biota. In: Rosenzweig C, Powlson D, Scow K, Singer M, Sparks D (eds) Encyclopedia of soils in the environment. Academic, New York, pp 507–512CrossRefGoogle Scholar
  31. Jenny H (1941) Factors of soil formation. McGraw-Hill, New York, p 281Google Scholar
  32. Koerselman W, Meuleman AFM (1996) The vegetation N: P ratio: a new tool to detect the nature of nutrient limitation. J Appl Ecol 33:1441–1450CrossRefGoogle Scholar
  33. Körner C (1998) A re-assessment of high elevation treeline positions and their explanations. Oecologia 115:445–459CrossRefGoogle Scholar
  34. Körner C, Paulsen J (2004) A world-wide study of high altitude treeline temperatures. J Biogeogr 31:1365–2699CrossRefGoogle Scholar
  35. Lipson D, Näsholm T (2001) The unexpected versatility of plants: organic nitrogen use and availability in terrestrial ecosystems. Oecologia 128:305–316CrossRefGoogle Scholar
  36. Liptzin D, Seastedt TR (2009) Patterns of snow, deposition, and soil nutrients at multiple spatial scales at a Rocky Mountain tree line ecotone. J Geophys Res – Biogeo 114:G04002. doi: 10.1029/2009JG000941 CrossRefGoogle Scholar
  37. Litaor MI (1987) The influence of eolian dust on the genesis of alpine soils in the Front Range, Colorado. Soil Sci Soc Am J 51:142–147CrossRefGoogle Scholar
  38. Litaor MI, Seastedt TR, Walker MD, Carbone M, Townsend A (2005) The biogeochemistry of phosphorus across an alpine topographic/snow gradient. Geoderma 124:49–61CrossRefGoogle Scholar
  39. Lund LJ, Brown AD, Lueking MA, Nodvin SC, Page AL, Sposito G (1987) Soil processes at emerald lake watershed, final report. Contract Number A3-105-32 California Air Resources BoardGoogle Scholar
  40. Malanson GP, Butler DR, Fagre DB, Walsh SJ, Tomback DF, Daniels LD, Resler LM, Smith WK, Weiss DJ, Peterson DL, Bunn AG, Hiemstra CA, Liptzin D, Bourgeron PS, Shen Z, Millar CI (2007) Alpine treeline of western North America: Linking organism-to-landscape dynamics. Phys Geogr 28:378–396CrossRefGoogle Scholar
  41. Martínez I, Wiegand T, Camarero JJ, Batllori E, Gutiérrez E (2011) Disentangling the formation of contrasting tree line physiognomies combining model selection and Bayesian parameterization for simulation models. Am Nat 177:E136–E152PubMedCrossRefGoogle Scholar
  42. May DE, Webber PJ (1982) Spatial and temporal variation of the vegetation and its productivity on Niwot Ridge, Colorado. In: JC Halfpenny (ed) Ecological studies in the Colorado alpine: a Festschrift for John W. Marr. pp. 35–62. University of Colorado, Institute of Arctic and Alpine Research Occasional Paper No. 37Google Scholar
  43. McGroddy M, Daufresne T, Hedin L (2004) Scaling of C:N:P stoichiometry in forests worldwide: implications of terrestrial Redfield-type ratios. Ecology 85:2390–2401CrossRefGoogle Scholar
  44. Muhs DR, Benedict JB (2006) Eolian additions to late Quaternary alpine soils, Indian Peaks Wilderness Area, Colorado Front Range. Arctic Antarc Alp Res 38:120–130CrossRefGoogle Scholar
  45. Neff JC, Harden JW, Gleixner G (2005) Fire effects on soil organic matter content, composition, and nutrients in boreal interior Alaska. Can J For Res 35:2178–2187CrossRefGoogle Scholar
  46. Neff JC, Ballantyne AP, Farmer GL, Mahowald NM, Conroy JL, Landry CC, Overpeck JT, Painter TH, Lawrence CR, Reynolds RL (2008) Increasing eolian dust deposition in the western United States linked to human activity. Nat Geosci. doi: 10.1038/ngeo133
  47. Niwot Ridge LTER database (2007) accessed March 29, 2007
  48. Parker ER, Sanford RL (1999) The effects of mobile tree islands on soil phosphorus concentrations and distribution in an alpine ecosystem on Niwot Ridge, Colorado Front Range, USA. Arctic Alp Res 31:16–20CrossRefGoogle Scholar
  49. Pauker SJ, Seastedt TR (1996) Effects of mobile tree islands on soil carbon storage in tundra ecosystems. Ecology 77:2563–2567CrossRefGoogle Scholar
  50. Redfield AC (1958) The biological control of chemical factors in the environment. Am Sci 46:205–222Google Scholar
  51. Reich PB, Oleksyn J (2004) Global patterns of plant leaf N and P in relation to temperature and latitude. Proc Natl Acad Sci U S A 101:11001–11006PubMedCrossRefGoogle Scholar
  52. Rochelle RR (1998) Pedogenic effects of krummholz migration. M.S. Thesis. Colorado State University, Ft. Collins, COGoogle Scholar
  53. Rueth HM, Baron JS (2002) Differences in Englemann spruce forest biogeochemistry east and west of the Continental Divide in Colorado, USA. Ecosystems 5:45–57CrossRefGoogle Scholar
  54. Rueth HM, Baron JS, Allstott EJ (2003) Responses of Engelmann spruce forests to nitrogen fertilization in the Colorado Rocky Mountains. Ecol Appl 13:664–673CrossRefGoogle Scholar
  55. Scholes RJ, Archer SR (1997) Tree–grass interactions in savannas. Annu Rev Ecol Syst 28:517–544CrossRefGoogle Scholar
  56. Seastedt TR, Adams GA (2001) Effects of mobile tree islands on alpine tundra soils. Ecology 82:8–17CrossRefGoogle Scholar
  57. Seastedt TR, Bowman WD, Caine TN, McKnight D, Townsend A, Williams MW (2004) The landscape continuum: a model for high-elevation ecosystems. BioScience 54:111–121CrossRefGoogle Scholar
  58. Shaver GR, Chapin FS (1995) Long-term responses to factorial, NPK fertilizer treatment by Alaskan wet and moist tundra sedge species. Ecography 18:259–275CrossRefGoogle Scholar
  59. Shiels AB, Sanford RL (2001) Soil nutrient differences between two krummholz-form tree species and adjacent alpine tundra. Geoderma 102:205–217CrossRefGoogle Scholar
  60. Sjogersten S, Wookey PA (2005) The role of soil organic matter quality and physical environment for nitrogen mineralization at the forest-tundra ecotone in Fennoscandia. Arctic Antarc Alp Res 37:118–126CrossRefGoogle Scholar
  61. Smith BFL, Bain DC (1982) A sodium hydroxide fusion method for the determination of total phosphate in soils. Commun Soil Sci Plan 13:185–190CrossRefGoogle Scholar
  62. Stark S (2007) Nutrient cycling in the tundra. In: Nutrient cycling in terrestrial ecosystems. Springer-Verlag Berlin Heidelberg. Soil Biol 10:309–331Google Scholar
  63. Sterner RW, Elser JJ (2002) Ecological stoichiometry: the biology of elements from molecules to the biosphere. Princeton University Press, PrincetonGoogle Scholar
  64. Stevens GC, Fox JF (1991) The causes of treeline. Annu Rev Ecol Syst 22:177–191CrossRefGoogle Scholar
  65. Sundqvist MK, Giesler R, Graae BJ, Wallander H, Fogelberg E, Wardle D (2011) Interactive effects of vegetation type and elevation on aboveground and belowground properties in a subarctic tundra. Oikos 120:128–142CrossRefGoogle Scholar
  66. Theodose TA, Bowman WD (1997) Nutrient availability, plant abundance, and species diversity in two alpine tundra communities. Ecology 78:1861–1872CrossRefGoogle Scholar
  67. Thorn C, Darmody R (1980) Contemporary eolian sediments in the alpine zone, Colorado Front Range. Phys Geogr 1:162–171Google Scholar
  68. Tiessen H, Moir JO (1993) Characterization of available P by sequential extraction. In: Carter MR (ed) Soil sampling and methods of analysis. Lewis Publishers, Boca Raton, pp 75–86Google Scholar
  69. Townsend AR, Cleveland CC, Asner GP, Bustamante MMC (2007) Controls of foliar N:P ratios in tropical rain forests. Ecology 88:107–118PubMedCrossRefGoogle Scholar
  70. Van Miegroet H, Hysell MT, Johnson AD (2001) Soil microclimate and chemistry of spruce-fir tree islands in Northern Utah. Soil Sci Soc Am J 64:1515–1525CrossRefGoogle Scholar
  71. Vitousek PM, Howarth RW (1991) Nitrogen limitation on land and in the sea: how can it occur? Biogeochemistry 13:87–115CrossRefGoogle Scholar
  72. Walbridge MR (1991) Phosphorus availability in acid organic soils of the lower North Carolina coastal plain. Ecology 72:2083–2100CrossRefGoogle Scholar
  73. Walker TW, Adams AFR (1958) Studies on soil organic matter. I. Influence of phosphorus content of parent materials on accumulations of carbon, nitrogen, sulfur and organic phosphorus in grassland soils. Soil Sci 85:307–318CrossRefGoogle Scholar
  74. Walker TW, Syers JK (1976) Fate of phosphorus during pedogenesis. Geoderma 15:1–19CrossRefGoogle Scholar
  75. Weiss L, Shiels AB, Walker LR (2005) Soil impacts of bristlecone pine (Pinus longaeva) tree islands on alpine tundra, Charleston Peak, Nevada. West N Am Nat 65:536–540Google Scholar
  76. Williams MW, Baron JS, Caine N, Sommerfeld R, Sanford R (1996) Nitrogen saturation in the Rocky Mountains. Environ Sci Technol 30:640–646CrossRefGoogle Scholar
  77. Williams MW, Rikkers M, Pfeffer WT (2000) Ice columns and frozen rills in a warm snowpack, Green Lakes Valley, Colorado, USA. Nord Hydrol 31:169–186Google Scholar

Copyright information

© Springer Science+Business Media B.V. 2012

Authors and Affiliations

  • Daniel Liptzin
    • 1
    • 3
  • Robert L. SanfordJr.
    • 2
    • 4
  • Timothy R. Seastedt
    • 1
  1. 1.Department of Ecology and Environmental Biology and INSTAARUniversity of ColoradoBoulderUSA
  2. 2.Department of Biological SciencesUniversity of DenverDenverUSA
  3. 3.Department of Natural Resources and the EnvironmentUniversity of New HampshireDurhamUSA
  4. 4.School of Earth Sciences and Environmental SustainabilityNorthern Arizona UniversityFlagstaffUSA

Personalised recommendations