ISSN : 2287-8327
Whilst it is clear that increasing temperatures from global environmental change will impact the positions of alpine treelines, it is likely that a range of regional and local scaled factors will mediate the overall impact of global scale climate drivers. We summarized 12 categories of abiotic and biotic factors as 4 groups determining treeline positions. First, there are global factors related to climate-induced growth limitation and carbon limitation. Second, there are seven regional and local factors related to treeline dynamics including frost stress, topography, water stress, snow, wind, fire and non-fire disturbance. Third, species-specific factors can control treeline dynamics through their influence on reproduction and life history traits. Fourth, there are positive feedbacks in structuring the dynamics of treelines. Globally, the com¬monly accepted growth limitation hypothesis is that growth at a treeline is limited by temperature. Meanwhile, positive feedbacks between canopy cover and tree establishment are likely to control the spatial pattern and temporal dynamics of many treelines. The presence of non-linear dynamics at treelines has implications for the use of treelines as barometers of climate change because the lagged responses and abrupt shifts inherent in non-equilibrium systems may combine to mask the overall climate trend.
Alftine KJ, Malanson GP. 2004. Directional positive feedback and pattern at an alpine tree line. J Veg Sci 15: 3-12.
Anadon-Rosell A, Rixen C, Cherubini P, Wipf S, Hagedorn F, Dawes MA. 2014. Growth and phenology of three dwarf shrub species in a six-year soil warming experiment at the Alpine treeline. PLoS ONE 9: e100577. DOI 10.1371/journal.pone.0100577.
Aulitzky H. 1961. Die Bodentemperaturen in der Kampfzone oberhalb der Waldgrenze und imsubalpinen Zirben-Lärchenwald. Mitt Forstl Bund Mariabrunn 59: 153-208.
Autio J. 2006. Environmental Factors Controlling the Position of the Actual Timberline and Treeline on the Fells of Finnish Lapland. Oulu University Press, Oulu.
Autio J, Colpaert A. 2005. The impact of elevation, topography and snow load damage of trees on the position of the actual timberline on the fells in central Finnish Lapland. Fennia183: 15-36.
Bader MY. 2007. Tropical Alpine Treelines: how ecological processes control vegetation patterning and dynamics. PhD Dissertation. Wageningen University, Wageningen, The Netherlands.
Bader MY, Rietkerk M, Bregt AK. 2008. A simple spatial model exploring positive feedbacks at tropical alpine treelines. Arct Antarct Alp Res 40: 269-278.
Bader MY, Ruijten JJA. 2008. A topography-based model of forest cover at the alpine tree line in the tropical Andes. J Biogeogr 35: 711–723.
Baig MN, Tranquillini W. 1980. The effects of wind and temperature on cuticular transpiration of Picea abies and Pinus cembra and their significance in dessication damage at the alpine treeline. Oecologia 47: 252-256.
Bale JS, Masters GJ, Hodkinson ID, Awmack C, Bezemer TM, Brown VK, Butterfield J, Buse A, Coulson JC, Farrar J, Good JEG, Harrington R, Hartley S, Jones TH, Lindroth RL, Press MC, Symrnioudis I, Watt AD, Whittaker JB. 2002. Herbivory in global climate change research: direct effects of rising temperature on insect herbivores. Glob Change Biol 8: 1–16.
Ball MC, Egerton JJG, Leuning R, Cunningham RB, Dunne P. 1997. Microclimate above grass adversely affects spring growth of seedling snow gum (Eucalyptus pauciflora). Plant Cell Environ 20: 155-166.
Barbeito I, Dawes MA, Rixen C, Senn J, Bebi P. 2012. Factors driving mortality and growth at treeline: a 30-year experiment of 92000 conifers. Ecology 93: 389-401.
Batllori E, Camarero JJ, Ninot JM, Gutiérrez E. 2009. Seedling recruitment, survival and facilitation in alpine Pinus uncinata treeline ecotones. Implications and potential responses to climate warming. Glob Ecol Biogeogr 18: 460-472.
Batllori E, Gutiérrez E. 2008. Regional treeline dynamics in response to global change in the Pyrenees. J Ecol 96: 1275-1288.
Bebi P, Kulakowski D, Rixen C. 2009. Snow avalanche disturbances in forest ecosystems—State of research and implications for management. For Ecol Manage 257: 1883-1892.
Bekker MF. 2005. Positive feedback between tree establishment and patterns of subalpine forest advancement, Glacier National Park, Montana, USA. Arct Antarc Alp Res 37: 97-107.
Berdanier AB. 2010. Global treeline position. Nat Educ Knowl 3: 11.
Black RA, Bliss LC. 1980. Reproductive ecology of Picea mariana (Mill.) BSP., at the tree line near Inuvik, Northwest Territories, Canada. Ecol Monogr 50: 331-354.
Bolli J, Rigling A, Bugmann H. 2007. Regeneration dynamics of Norway spruce (Picea abies L.) on a subalpine meadow near the treeline in Sedrun, Kt. Graubünden, Switzerland. Silva Fenn 41: 55-70.
Brown CD. 2010. Tree-line dynamics: adding fire to climate change prediction. Arctic 63: 488-492.
Brown DG. 1994a. Comparison of vegetation-topography relationships at the alpine treeline ecotone. Phys Geogr 15: 125-145.
Brown DG. 1994b. Predicting vegetation types at treeline using topography and biophysical disturbance variables. J Veg Sci 5: 641-656.
Butler DR, Hill C, Malanson GP, Cairns DM. 1994. Stability of alpine treeline in Glacier National Park, Montana, USA. Phytocoenologia 22: 485-500.
Butler DR, Malanson GP, Walsh SJ, Fagre DB. 2007. Influences of geomorphology and geology on alpine treeline in the American West - more important than climatic influences? Phys Geogr 28: 434-450.
Cairns DM. 1999. Multi-scale analysis of soil nutrients at alpine treeline in Glacier National Park, Montana. Phys Geogr 20: 256-271.
Cairns DM, Malanson GP. 1998. Environmental variables influencing the carbon balance at the alpine treeline: a modeling approach. J Veg Sci 9: 679-692.
Cairns DM, Moen J. 2004. Herbivory influences tree lines. J Ecol 92: 1019-1024.
Camarero JJ, Gutiérrez E. 2004. Pace and pattern of recent treeline dynamics: response of ecotones to climatic variability in the Spanish Pyrenees. Clim Chang 63: 181-200.
Carlson BZ, Randin CF, Boulangeat I, Lavergne S, Thuiller W, Choler P. 2013. Working toward integrated models of alpine plant distribution. Alp Bot 123:41-53.
Case BS, Duncan RP. 2014. A novel framework for disentangling the scale-dependent influences of abiotic factors on alpine treeline position. Ecography 37: 838-851.
Chang NK, Shim KC, Lee HU, Kang KM, So KH. 1998. The theory of boundary distribution of plant and wave character of the timber line on Mt. Paektu. Korean J Ecol 21: 491-499.
Chauchard S, Beilhe F, Denis N, Carcaillet C. 2010. An increase in the upper tree-limit of silver fir (Abies alba Mill.) in the Alps since the mid-20th century: a land-use change phenomenon. For Ecol Manage 259: 1406-1415.
Cho DS. 1994. Community structure, and size and age distribution of conifers in subalpine Korean Fir (Abies koreana) Forest in Mt. Chiri. Korean J Ecol 17: 415-424.
Cochrane PM, Slatyer RO. 1988. Water relations of Eucalyptus pauciflora near the alpine tree line in winter. Tree Physiol 4: 45-52.
Cogbill CV, White PS. 1991. The latitude-elevation relationship for spruce-fir forest and treeline along the Appalachian mountain chain. Vegetatio 94: 153-175.
Cuevas JG. 2000. Tree recruitment at the Nothofagus pumilio alpine timberline in Tierra del Fuego, Chile. J Ecol 88: 840-855.
Cullen LE, Stewart GH, Duncan RP, Palmer JG. 2001. Disturbance and climate warming influences on New Zealand Nothofagus tree-line population dynamics. J Ecol 89: 1061-1071.
Daly C, Shankman D. 1985. Seedling establishment by conifers above the tree limit on Niwot Ridge, Front Range, Colorado, USA. Arct Alp Res 17: 389-400.
Danby RK, Hik DS. 2007. Variability, contingency and rapid change in recent subarctic alpine tree line dynamics. J Ecol 95: 352-363.
Dang H, Zhang K, Zhang Y, Tan S, Jiang M, Zhang Q. 2009. Treeline dynamics in relation to climatic variability in the Shennongjia Mountains, Central China. Can J For Res 39: 1848-1858.
Daniels LD, Veblen TT. 2003. Regional and local effects of disturbance and climate on altitudinal treelines in northern Patagonia. J Veg Sci 14: 733-742.
Devi N, Hagedorn F, Moiseev P, Bugmann H, Shiyatov S, Mazepa V, Rigling A. 2008. Expanding forests and changing growth forms in Siberian larch at the Polar Urals treeline during the 20th century. Glob Change Biol 14: 1581-1591.
Díaz-Varela RA, Colombo R, Meroni M, Calvo-Iglesias MS, Buffoni A, Tagliaferri A. 2010. Spatio-temporal analysis of alpine ecotones: a spatial explicit model targeting altitudinal vegetation shifts. Ecol Model 221: 621-633.
Dullinger S, Dirnbock T, Grabherr G. 2004. Modelling climate change-driven treeline shifts: relative effects of temperature increase, dispersal and invisibility. J Ecol 92: 241-252.
Fajardo A, Piper FI, Cavieres LA. 2011. Distinguishing local from global climate influences in the variation of carbon status with altitude in a tree line species. Glob Ecol Biogeogr 20: 307-318.
Fajardo A, Piper FI, Pfund L, Körner C, Hoch G. 2012. Variation of mobile carbon reserves in trees at the alpine treeline ecotone is under environmental control. New Phytol 195: 794-802.
Feeley KJ, Silman MR, Bush MB, Farfan W, Cabrera KG, Malhi Y, Meir P, Revilla NS, Quisiyupanqui MNR, Saatchi S. 2011. Upslope migration of Andean trees. J Biogeogr 38: 783-791.
Fletcher MS, Wood SW, Haberle SG. 2014. A fire driven shift from forest to non-forest: evidence for alternative stable states? Ecology 95: 2504-2513.
Gamache I, Payette S. 2004. Height growth response of tree line black spruce to recent climate warming across the forest-tundra of eastern Canada. J Ecol 92: 835-845.
Gehrig-Fasel J, Guisan A, Zimmermann NE. 2007. Tree line shifts in the Swiss Alps: climate change or land abandonment? J Veg Sci 18: 571-582.
Germino MJ, Smith WK. 1999. Sky exposure, crown architecture, and low‐temperature photoinhibition in conifer seedlings at alpine treeline. Plant Cell Environ 22: 407-415.
Germino MJ, Smith WK, Resor AC. 2002. Conifer seedling distribution and survival in an alpine-treeline ecotone. Plant Ecol 162: 157-168.
Gottfried M, Hantel M, Maurer C, Toechterle R, Pauli H, Grabherr G. 2011. Coincidence of the alpine–nival ecotone with the summer snowline. Environ Res Lett 6: 014013.
Gottfried M, Pauli H, Reiter K, Grabherr G. 1999. A finescaled predictive model for changes in species distributions patterns of high mountain plants induced by climate warming. Divers Distrib 5: 241-252.
Grabherr G, Gottfried M, Pauli H. 1994. Climate effects on mountain plants. Nature 369: 448.
Grace J. 1977. Plant Responses to Wind. Academic Press, London.
Grace J. 1989. Tree lines. Phil Trans R Soc Lond B 324: 233-245.
Grace J, Berninger F, Nagy L. 2002. Impacts of climate change on the tree line. Ann Bot 90: 537-544.
Green K. 2009. Causes of stability in the alpine treeline in the Snowy Mountains of Australia–a natural experiment. Aust J Bot 57: 171-179.
Green K, Venn SE. 2012. Tree-limit ribbons in the snowy mountains, Australia: characterization and recent seedling establishment. Arct Antarc Alp Res 44: 180-187.
Guisan A, Theurillat JP. 2000. Assessing alpine plant vulnerability to climate change: a modeling perspective. Integr Assess 1: 307-320.
Gworek JR, Vander Wall SB, Brussard PF. 2007. Changes in biotic interactions and climate determine recruitment of Jeffrey pine along an elevation gradient. For Ecol Manage 239: 57-68.
Hagedorn F, Shiyatov SG, Mazepa VS, Devi NM, Grigor’ev AA, Bartysh AA, Fomin VV, Kapralov DS, Terent’ev M, Bugman H, Rigling A, Moiseev PA. 2014. Treeline advances along the Urals mountain range - driven by improved winter conditions? Glob Change Biol 20: 3530-3543.
Handa IT, Körner C, Hättenschwiler S. 2005. A test of the treeline carbon limitation hypothesis by in situ CO2 enrichment and defoliation. Ecology 86: 1288-1300.
Harsch MA, Bader MY. 2011. Treeline form - a potential key to understanding treeline dynamics. Glob Ecol Biogeogr 20: 582-596.
Harsch MA, Hulme PE, McGlone MS, Duncan RP. 2009. Are treelines advancing? A meta-analysis of treeline response to climate warming. Ecol Lett 12: 1040-1049.
Harvell CD, Mitchell CE, Ward JR, Altizer S, Dobson AP, Ostfeld RS, Samuel MD. 2002. Climate warming and disease risks for terrestrial and marine biota. Science 296: 2158-2162.
Hättenschwiler S, Körner C. 1995. Responses to recent climate warming of Pinus sylvestris and Pinus cembra within their montane transition zone in the Swiss Alps. J Veg Sci 6: 357-368.
Hättenschwiler S, Smith WK. 1999. Seedling occurrence in alpine treeline conifers: a case study from the central Rocky Mountains, USA. Acta Oecol 20: 219-224.
Heikkinen O, Tuovinen M, Autio J. 2002. What determines the timberline? Fennia 180: 67-74.
Hemstrom MA, Franklin JF. 1982. Fire and other disturbances of the forests in Mount Rainier National Park. Quat Res 18: 32-51.
Herrero A, Zamora R, Castro J, Hódar JA. 2011. Limits of pine forest distribution at the treeline: herbivory matters. Plant Ecol 213: 459-469.
Hertel D, Schöling D. 2011. Below-ground response of Norway spruce to climate conditions at Mt. Brocken (Germany)-A re-assessment of Central Europe’s northernmost treeline. Flora 206: 127-135.
Hirota M, Holmgren M, Van Nes EH, Scheffer M. 2011. Global resilience of tropical forest and savanna to critical transitions. Science 334: 232-235.
Hoch G, Körner C. 2003. The carbon charging of pines at the climatic treeline: a global comparison. Oecologia 135: 10-21.
Hoch G, Körner C. 2005. Growth, demography and carbon relations of Polylepis trees at the world’s highest treeline. Funct Ecol 19: 941-951.
Hoch G, Körner C. 2009. Growth and carbon relations of tree line forming conifers at constant vs. variable low temperatures. J Ecol 97: 57-66.
Hoch G, Popp M, Körner C. 2002. Altitudinal increase of mobile carbon pools in Pinus cembra suggests sink limitation of growth at the Swiss treeline. Oikos 98: 361-374.
Holtmeier FK. 2003. Mountain Timberlines: Ecology, Patchiness and Dynamics. Kluwer Academic Publishers, Dordrecht.
Holtmeier FK, Broll G. 2005. Sensitivity and response of northern hemisphere altitudinal and polar treelines to environmental change at landscape and local scales. Glob Ecol Biogeogr 14: 395-410.
Holtmeier FK, Broll G. 2007. Treeline advance – driving processes and adverse factors. Landsc Online 1: 1-33.
Holtmeier FK, Broll G. 2010. Wind as an ecological agent at treelines in North America, the Alps, and the European Subarctic. Phys Geogr 31: 203-233.
Ihm BS, Lee JS, Kim JW. 2012. Global warming and biodiversity model projections. J Ecol Field Biol 35: 157-166.
Ihm BS, Lee JS, Kim JW, Kim JH. 2007. Relationship between global warming and species richness of vascular plants. J Plant Biol 50: 321-324.
IPCC. 2007. Climate Change 2007: Synthesis Report. Contribution of Working Groups I, II and III to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change. IPCC, Geneva.
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. IPCC, Geneva.
Jeffers ES, Bonsall MB, Brooks SJ, Willis KJ. 2011. Abrupt environmental changes drive shifts in tree–grass interaction outcomes. J Ecol 99: 1063-1070.
Jobbagy EG, Jackson RB. 2000. Global controls of forest line elevation in the northern and southern hemispheres. Glob Ecol Biogeogr 9: 253-268.
Johnson DM, Germino MJ, Smith WK. 2004. Abiotic factors limiting photosynthesis in Abies lasiocarpa and Picea engelmannii seedlings below and above the alpine timberline. Tree Physiol 24: 377-386.
Kim JH. 2012. Global Warming. Seoul National University Press, Seoul.
Knox KJE, Clarke PJ. 2012. Fire severity, feedback effects and resilience to alternative community states in forest assemblages. For Ecol Manage 265: 47-54.
Kong WS. 1999. The vertical distribution of air temperature and thermal amplitude of alpine plants on Mt. Halla, Cheju Island, Korea. J Korean Geo Soc 34: 385-393.
Kong WS. 2000. Geoecology on the subalpine vegetation and landscape of Mt. Sorak. J Korean Geo Soc 35: 177-187.
Kong WS, Watts D. 1993. The Plant Geography of Korea with an Emphasis on the Alpine Zones. Kluwer Academic Publishers, Dordrecht.
Körner C. 1998. A re-assessment of high-elevation treeline positions and their explanation. Oecologia 115: 445-459.
Körner C. 2003. Carbon limitation in trees. J Ecol 91: 4-17.
Körner C. 2012. Alpine Treelines. Springer, Berlin.
Körner C, Paulsen J. 2004. A world-wide study of high altitude treeline temperatures. J Biogeogr 31: 713-732.
Kreyling J, Wana D, Beierkuhnlein C. 2010. Potential consequences of climate warming for tropical plant species in high mountains of southern Ethiopia. Divers Distrib 16: 593-605.
Kullman L. 2001. 20th century climate warming and treelimit rise in the southern Scandes of Sweden. Ambio 30: 72-80.
Kullman L. 2002. Rapid recent range-margin rise of tree and shrub species in the Swedish Scandes. J Ecol 90: 68-77.
Kullman L. 2005. Wind-conditioned 20th century decline of birch treeline vegetation in the Swedish Scandes. Arctic 58: 286-294.
Kullman L. 2007. Tree line population monitoring of Pinus sylvestris in the Swedish Scandes, 1973–2005: implications for tree line theory and climate change ecology. J Ecol 95: 41-52.
Kullman L. 2010a. A richer, greener and smaller alpine world: review and projection of warming-induced plant cover change in the Swedish Scandes. Ambio 39: 159-169.
Kullman L. 2010b. One century of treeline change and stability - experiences from the Swedish Scandes. Landsc Online 17: 1-31.
Lenoir J, Gegout JC, Marquet PA, de Ruffray P, Brisse H. 2008. A significant upward shift in plant species optimum elevation during the 20th century. Science 320: 1768-1771.
Leonelli G, Pelfini M, di Cella UM. 2009. Detecting climatic treelines in the Italian Alps: The influence of geomorphological factors and of human impacts. Phys Geogr 30: 338-352.
Leonelli G, Pelfini M, di Cella UM, Garavaglia V. 2011. Climate warming and the recent treeline shift in the European Alps: the role of geomorphological factors in highaltitude sites. Ambio 40: 264-273.
Li MH, Xiao WF, Wang SG, Cheng GW, Cherubini P, Cal XH, Liu XL, Wang XD, Zhu WZ. 2008. Mobile carbohydrates in Himalayan treeline trees I. Evidence for carbon gain limitation but not for growth limitation. Tree Physiol 28: 1287-1296.
Lloyd AH, Fastie CL. 2003. Recent changes in treeline forest distribution and structure in interior Alaska. Ecoscience 10: 176-185.
MacDonald GM, Kremenetski KV, Beilman DW. 2008. Climate change and the northern Russian treeline zone. Phil Trans R Soc B 363: 2285-2299.
Macek P, Macková J, de Bello F. 2009. Morphological and ecophysiological traits shaping altitudinal distribution of three Polylepis treeline species in the dry tropical Andes. Acta Oecol 35: 778-785.
Malanson GP. 2001. Complex responses to global change at alpine treeline. Phys Geogr 22: 333-342.
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: 378-396.
Malanson GP, Resler LM, Bader MY, Holtmeier FK, Butler DR, Weiss DJ, Daniels LD, Fagre DB. 2011. Mountain treelines: a roadmap for research orientation. Arct Antarc Alp Res 43: 167-177.
Marchand PJ, Chabot BF. 1978. Winter water relations of tree-line plant species on Mt. Washington, New Hampshire. Arct Alp Res 10: 105-116.
Mayer AL, Khalyani AH. 2011. Grass trumps trees with fire. Science 334: 188-189.
Moen J, Aune K, Edenius L, Angerbjörn A. 2004. Potential effects of climate change on treeline position in the Swedish mountains. Ecol Soc 9: 16.
Moen J, Cairns DM, Lafon CW. 2008. Factors structuring the treeline ecotone in Fennoscandia. Plant Ecol Divers 1: 77-87.
Motta R, Nola P. 2001. Growth trends and dynamics in subalpine forest stands in the Varaita Valley (Piedmont, Italy) and their relationships with human activities and global change. J Veg Sci 12: 219-230.
Murphy BP, Bowman DMJS. 2012. What controls the distribution of tropical forest and savanna? Ecol Lett 15: 748-758.
Noble IR. 1980. Interactions between tussock grass (Poa spp.) and Eucalyptus pauciflora seedlings near treeline in South-Eastern Australia. Oecologia 45: 350-353.
Noble IR. 1993. A model of the responses of ecotones to climate change. Ecol Appl 3: 396-403.
Odion DC, Moritz MA, DellaSala DA. 2010. Alternative community states maintained by fire in the Klamath Mountains, USA. J Ecol 98: 96-105.
Odland A. 2009. Interpretation of altitudinal gradients in South Central Norway based on vascular plants as environmental indicators. Ecol Indic 9: 409-421.
Pauli H, Gottfried M, Grabherr G. 1996. Effects of climate change on mountain ecosystems - upward shifting of alpine plants. World Resource Review 8: 382-390.
Payette S, Filion L, Delwaide A, Begin C. 1989. Reconstruction of tree-line vegetation response to long-term climate change. Nature 341: 429-432.
Rada F, Azócar A, Briceño B, González J, García-Núñez C. 1996. Carbon and water balance in Polylepis sericea, a tropical treeline species. Trees 10: 218-222.
Resler LM. 2006. Geomorphic controls of spatial pattern and process at alpine treeline. Prof Geogr 58: 124-138.
Richardson AD, Friedland AJ. 2009. A review of the theories to explain Arctic and alpine treelines around the world. J Sustainable For 28: 218-242.
Rixen C, Dawes MA, Wipf S, Hagedorn F. 2012. Evidence of enhanced freezing damage in treeline plants during six years of CO2 enrichment and soil warming. Oikos 121: 1532-1543.
Sakai A. 1970. Mechanism of desiccation damage of conifers wintering in soil-frozen areas. Ecology 51: 657-664.
Scheffer M, Carpenter S, Foley JA, Folke C, Walker B. 2001. Catastrophic shifts in ecosystems. Nature 413: 591-596.
Scherrer D, Körner C. 2011. Topographically controlled thermal-habitat differentiation buffers alpine plant diversity against climate warming. J Biogeogr 38: 406-416.
Schulze ED, Mooney HA, Dunn EL. 1967. Wintertime photosynthesis of bristlecone pine (Pinus aristata) in White Mountains of California. Ecology 48: 1044-1047.
Scott AC, Bowman DMJS, Bond WJ, Pyne SJ, Alexander ME. 2014. Fire on Earth: An Introduction. John Wiley & Sons, Hoboken, NJ.
Scott PA, Hansell RIC, Erickson WR. 1993. Influences of wind and snow on northern tree-line environments at Churchill, Manitoba, Canada. Arctic 46: 316-323.
Shankman D, Daly C. 1988. Forest regeneration above tree limit depressed by fire in the Colorado Front Range. B Torrey Bot Club 115: 272-279.
Shi P, Körner C, Hoch G. 2006. End of season carbon supply status of woody species near the treeline in western China. Basic Appl Ecol 7: 370-377.
Shi P, Körner C, Hoch G. 2008. A test of the growth-limitation theory for alpine tree line formation in evergreen and deciduous taxa of the eastern Himalayas. Funct Ecol 22: 213-220.
Singh CP, Panigrahy S, Thapliyal A, Kimothi MM, Soni P, Parihar JS. 2012. Monitoring the alpine treeline shift in parts of the Indian Himalayas using remote sensing. Curr Sci 102: 559-562.
Smith WK, Germino MJ, Hancock TE, Johnson DM. 2003. Another perspective on altitudinal limits of alpine timberlines. Tree Physiol 23: 1101-1112.
Smith WK, Germino MJ, Johnson DM, Reinhardt K. 2009. The altitude of alpine treeline: a bellwether of climate change effects. Bot Rev 75: 163-190.
Smith-McKenna EK, Malanson GP, Resler LM, Carstensen LW, Prisley SP, Tomback DF. 2014. Cascading effects of feedbacks, disease, and climate change on alpine treeline dynamics. Environ Model Softw 62: 85-96.
Speed JDM, Austrheim G, Hester AJ, Mysterud A. 2010. Experimental evidence for herbivore limitation of the treeline. Ecology 91: 3414-3420.
Staver AC, Archibald S, Levin SA. 2011. The global extent and determinants of savanna and forest as alternative biome states. Science 334: 230-232.
Stevens GC, Fox JF. 1991. The causes of treeline. Annu Rev Ecol Syst 22: 177-191.
Stueve KM, Cerney DL, Rochefort RM, Kurth LL. 2009. Postfire tree establishment patterns at the alpine treeline ecotone: Mount Rainier National Park, Washington, USA. J Veg Sci 20: 107-120.
Susiluoto S, Peramaki M, Nikinmaa E, Berninger F. 2007. Effects of sink removal on transpiration at the treeline: implications for the growth limitation hypothesis. Environ Exp Bot 60: 334-339.
Sveinbjörnsson B. 2000. North American and European treelines: external forces and internal processes controlling position. Ambio 29: 388-395.
Sveinbjörnsson B, Kauhanen H, Nordell O. 1996. Treeline ecology of mountain birch in the Torneträsk area. Ecol Bull 45: 65-70.
Szeicz JM, MacDonald GM. 1995. Recent white spruce dynamics at the subarctic alpine treeline of north-western Canada. J Ecol 83: 873-885.
Szerencsits E. 2012. Swiss tree lines - a GIS-based approximation. Landsc Online 28: 1-18.
Telewski FW. 1995. Wind-induced physiological and developmental responses in trees. In: Wind and Trees (Coutts MP, Grace J, eds). Cambridge University Press, Cambridge, pp 237-263.
Tomback DF, Resler LM. 2007. Invasive pathogens at alpine treeline: complications and concerns. Phys Geogr 28: 397-418.
Tranquillini W. 1979. Physiological Ecology of the Alpine Timber-line.Tree Existence at High Altitudes with Special References to the European Alps (Ecological studies 31). Springer, Berlin.
Wallentin G, Tappeiner U, Strobl J, Tasser E. 2008. Understanding alpine tree line dynamics: an individual based model. Ecol Model 218: 235-246.
Walsh SJ, Butler DR, Allen TR, Malanson GP. 1994. Influence of snow patterns and snow avalanches on the alpine treeline ecotone. J Veg Sci 5: 657-672.
Walsh SJ, Butler DR, Malanson GP, Crews-Meyer KA, Messina JP, Xiao N. 2003. Mapping, modeling, and visualization of the influences of geomorphic processes on the alpine treeline ecotone, Glacier National Park, MT, USA. Geomorphology 53: 129-145.
Walther G. 2003. Plants in a warmer world. Perspect Plant Ecol Evol Syst 6: 169-185.
Walther GR, Beißner S, Burga CA. 2005. Trends in the upward shift of alpine plants. J Veg Sci 16: 541-548.
Wang T, Zhang QB, Ma KP. 2006. Treeline dynamics in relation to climatic variability in the central Tianshan Mountains, northwestern China. Global Ecol Biogeogr 15: 406-415.
Wardle P, Coleman MC. 1992. Evidence for rising upper limits of four native New Zealand forest trees. New Zeal J Bot 30: 303-314.
Warman L, Moles AT. 2009. Alternative stable states in Australia’s wet tropics: a theoretical framework for the field data and a field-case for the theory. Landsc Ecol 24: 1-13.
Wieser G, Matyssek R, Luzian R, Zwerger P, Pindur P, Oberhuber W, Gruber A. 2009. Effects of atmospheric and climate change at the timberline of the Central European Alps. Ann For Sci 66: 402-412.
Wilson JB, Agnew ADQ. 1992. Positive-feedback Switches in Plant Communities. Academic Press, London.
Wood SW, Bowman DMJS. 2012. Alternative stable states and the role of fire–vegetation–soil feedbacks in the temperate wilderness of southwest Tasmania. Landsc Ecol 27: 13-28.
Wood SW, Murphy BP, Bowman DMJS. 2011. Firescape ecology: how topography determines the contrasting distribution of fire and rain forest in the south-west of the Tasmanian Wilderness World Heritage Area. J Biogeogr 38: 1807-1820.
Xu Z, Hu T, Zhang Y. 2012. Effects of experimental warming on phenology, growth and gas exchange of treeline birch (Betula utilis) saplings, Eastern Tibetan Plateau, China. Eur J Forest Res 131: 811-819.
Zhang YJ, Dai LM, Pan J. 2001. The trend of tree line on the northern slope of Changbai Mountain. J For Res 12: 97-100.