Impact of wild herbivorous mammals and birds on the altitudinal and northern treeline ecotones


  • Friedrich-Karl Holtmeier Institute of Landscape Ecology, University of Münster, Robert-Koch-Straße 28, D-48149 Münster, Germany



Browsing, Burrowing, Over-grazing, Seed dispersal, Trampling, Treeline dynamics, Treeline spatial pattern, Wallowing


Wild herbivorous mammals may damage treeline vegetation an cause soil erosion at a local scale. In many high mountain areas of Europe and North America, large numbers of red deer have become a threat to the maintenance of high-elevation forests and attempts to restore the climatic treeline. In northern Fennoscandia, overgrazing by reindeer in combination with mass outbreaks of the autumnal moth are influencing treeline dynamics. Moose are also increasingly involved damaging treeline forest. In the Alps, the re-introduction of ibex is causing local damage to subalpine forests and tree establishment above the forest limit as well as aggravating soil erosion. High-elevation forests and treeline in Europe are susceptible to the deleterious impact of wild ungulate populations because of former extensive pastoral use. Rodents may damage tree seedlings and saplings by girdling, root cutting, bark stripping and burrowing. Hares damage young trees by gnawing. Large numbers of small rodents may occasionally impede tree regeneration by depleting the seed sources. Rodents do not contribute to forest expansion beyond the current treeline. Among birds, nutcrackers are highly effective in influencing tree distribution patterns and treeline dynamics. Without the nutcracker caching of stone pine seeds any upward advance of the trees in response to climatic warming would be impossible. Some bird species such as black grouse, willow grouse and ptarmigan can impair tree growth by feeding on buds, catkins and fresh terminal shoots.


Aalborg, T. (1955). Skador på Tallplantor Inom Ett Starkt Renbetat Område i Hedesocken,

Adams, L. A., & Bailey, J. A. (1983). Winter forages of mountain goats in central colorado. Journal of Wildlife Management, 47, 1237-1243.

Anschlag, K., Broll, G., & Holtmeier, F. -. (2008). Mountain birch seedlings in the treeline ecotone, subarctic finland: Variation in above- and below-ground growth depending on microtopography. Arctic, Antarctic, and Alpine Research, 40(4), 609-616. doi:10.1657/1523-0430(07-087)[ANSCHLAG]2.0.CO;2

Asakawa, S. (1957). Studies on hastening the germination of the seeds of five-leaved pines. Bulletin of the Governmental Forest Experiment Station, 10, 41-54.

Aune, S., Hofgaard, A., & Söderström, L. (2011). Contrasting climate- and land-use-driven tree encroachment patterns of subarctic tundra in northern norway and the kola peninsula. Canadian Journal of Forest Research, 41(3), 437-449. doi:10.1139/X10-086

Barclay, A. M., & Crawford, R. M. M. (1984). Seedling emergence in the rowan ( sorbus aucuparia) from an altitudinal gradient. Journal of Ecology, 72(2), 627-636. doi:10.2307/2260072

Bäumler, W., & Hohenadl, W. (1980). The influence of small mammals upon regeneration of a mountain forest in the alias (chiemgau). [Über den Einfluß alpiner Kleinsäuger auf die Verjüngung in einem Bergmischwald der Chiemgauer Alpen] Forstwissenschaftliches Centralblatt, 99(1), 207-221. doi:10.1007/BF02770941

Bavier, G. (1976). Graubündens waldwirtschaft. ein vergleich und ausblick. Bündner Wald, 29(6), 220-248.

Beachman, J. J., Cameron, P., Collins, M. S., & Reynolds, T. D. (2007).

Bekker, M. F., Clark, J. T., & Jackson, M. W. (2009). Landscape metrics indicate differences in patterns and dominant controls of ribbon forests in the rocky mountains, USA. Applied Vegetation Science, 12(2), 237-249. doi:10.1111/j.1654-109X.2009.01021.x

Belnap, J. (2003). Biological soil crusts and wind erosion. Biological Soil Crusts: Structure, Function, and Management, 150, 339-347.

Belnap, J., Büdel, B., & Lange, O. (2001). Biological Crusts: Characteristics and Distribution, 150, 1-30.

Bergström, R., & Hjeljord, O. (1987). Moose and vegetation interactions in northwestern europe and poland. Swedish Wildlife Research, 1(SUPPL. 1), 213-228.

Beschta, R. L., & Ripple, W. J. (2010). Recovering riparian plant communities with wolves in northern yellowstone, U.S.A. Restoration Ecology, 18(3), 380-389. doi:10.1111/j.1526-100X.2008.00450.x

Bichler, C. (1997). Auswirkungen Von Huftieren Auf Die Wiederbewaldungsdynamik Auf Drei Subalpinen Weiden Im Schweizerischen Nationalpark,

Bisaz, O. (1968). Das lawinenverbauungs- und aufforstungsprojekt munt da la bês-cha (schafberg) der gemeinde pontresina. Bündnerwald, 21(5), 201-207.

Bjørneraas, K., Herfindal, I., Solberg, E. J., Sæther, B. -., van Moorter, B., & Rolandsen, C. M. (2012). Habitat quality influences population distribution, individual space use and functional responses in habitat selection by a large herbivore. Oecologia, 168(1), 231-243. doi:10.1007/s00442-011-2072-3

Blüthgen, J. (1942). Die polare baumgrenze. Die Polare Baumgrenze.Veröffentlichungen Des Deutschen Wissen-Schaftlichen,

Bodenmann, A., & Eiberle, K. (1967). Über die auswirkungen des verbisses der gemse im aletschwald. Schweizerische Zeitschrift Für Forstwesen, 118, 461-470.

Bradshaw, R. H. W., Hannon, G. E., & Lister, A. M. (2003). A long-term perspective on ungulate-vegetation interactions. Forest Ecology and Management, 181(1-2), 267-280. doi:10.1016/S0378-1127(03)00138-5

Brandborg, S. M. (1955). Life history and management of the mountain goat in idaho. Idaho Wildlife Bulletin, 2, 1-142.

Broll, G., Holtmeier, F. -., Anschlag, K., Brauckmann, H. -., Wald, S., & Drees, B. (2007). Landscape mosaic in the treeline ecotone on mt rodjanoaivi. Subarctic Finland.Fennia, 185(2), 88-105.

Buckner, D. L. (1977). Ribbon forest development and maintenance in the central rocky mountains of colorado. Ribbon Forest Development and Maintenance in the Central Rocky Mountains of Colorado,

Burgess, P. (1999). Human Environmental Interactions in Upper Lapland, Finland,

Butler, D. R. (1993). The impact of mountain goat migration on unconsolidated slopes in glacier national park, montana. The Geographical Bulletin, 35(2), 98-106.

Butler, D. R. (1995). Zoogeomorphology: Animals as geomorphic agents. Zoogeomorphology: Animals as Geomorphic Agents,

Butler, D. R., Malanson, G. P., & Resler, L. M. (2004). Turf-banked terrace treads and risers, turf exfoliation and possible relationships with advancing treeline. Catena, 58(3), 259-274. doi:10.1016/j.catena.2004.05.003

Butler, D. R., Malanson, G. P., Resler, L. M., Walsh, S. J., Wilkerson, F. D., Schmid, G. L., & Sawyer, C. F. (2009). Geomorphic patterns and processes at alpine treeline. The Changing Alpine Treeline, 12, 63-84.

Cairns, D. M., & Moen, J. (2004). Herbivory influences tree lines. Journal of Ecology, 92(6), 1019-1024. doi:10.1111/j.1365-2745.2004.00945.x

Campell, E. (1955). Der lärchen-arvenwald. Bündner Wald, 5, 14-26.

Campell, E. (1950). Der tannen- oder nusshäher und die arvenverbreitung. Bündnerwald, 4(1), 3-7.

Campell, E. (1944). Der wald des oberengadins im wandel der zeiten. Festschrift Zur Jahresversammlung Der Schweizerischen Naturforschenden Gesellschaft, 124, 93-102.

Campell, E. (1958). Steinwild im engadin. Bündner Wald, 11, 287-294.

Chadwick, D. H. (1983). A beast the color of winter: The mountain goat observed. A Beast the Color of Winter,

Chase, A. (1986). Playing God in Yellowstone: The Destruction of America's First National Park,

Clarke, J. A., & Johnson, R. E. (2005). Comparisons and Contrasts between the Foraging Behaviors of Two White-Tailed Ptarmigan (Lagopus Leucurus) Populations, 37(2), 176-191.

Coleman, M. D., Hinckley, T. M., McNaughton, G., & Smit, B. A. (1992). Root cold hardiness and native distribution of subalpine conifers. Canadian Journal of Forest Research, 22(7), 932-938. doi:10.1139/x92-124

Conrads, K., & Balda, R. P. (1979). Überwinterungschancen sibirischer tannenhäher (nucifraga caryocatactes macrorhynchos) im invasionsgebiet. Ber.Naturwiss.Ver.Bielefeld, 24, 115-137.

Crocq, C. (1978). Écologie Du Casse-Noix (Nucifraga Caryocatactes L.) Dans Les Alpes Francaises due Sud; Ses Relations Avec l"Arolle (Pinus Cembra L.),

Danell, Ö., Staaland, H., & Nieminen, M. (1999). Rennäring i nordvest-europa. Reindrift i Nordvest-Europa 1998: Biologiske Muligheter Och Begrensininger, , 19-30.

Danilkin, A., & Hewison, A. J. M. (1996). Behavioural Ecology of Siberian and European Roe Deer,

DeLella-Benedict, A. (1991). The Southern Rockies.the Rocky Moutain Regions of Southern Wyoming, Colorado, and Northern New Mexico,

den Herder, M., Virtanen, R., & Roininen, H. (2008). Reindeer herbivory reduces willow growth and grouse forage in a forest-tundra ecotone. Basic and Applied Ecology, 9(3), 324-331. doi:10.1016/j.baae.2007.03.005

Donaubauer, E. (1963). Über die schneeschütte-krankheit (phacidium infestans karst.) der zirbe (pinus cembra L.) und einige begleitpilze. Mitt.Forstl.Bundesversuchsanst.Mariabrunn, 60, 575-600.

Eiberle, K. (1975). Ergebnisse einer simulation des wildverbisses durch den triebschnitt. Schweiz.Z.Forstwes., 126, 821-839.

Eiberle, K. (1978). Folgewirkungen eines simulierten wildverbisses auf die entwicklung junger waldbäume. Schweiz.Z.Forstwes., 129(9), 757-768.

Eiberle, K. (1980). Methodische möglichkeiten zum verständnis der waldbaulich tragbaren verbissbelastung. Schweiz.Z.Forstwes., 131(4), 311-326.

Eiberle, K. (1974). Waldkundliche aspekte der forschung an rauhfusshühnern. Schweiz.Z.Forstwirtsch., 125, 147-170.

Ericsson, A., Hellqvist, C., Langstrom, B., Larsson, S., & Tenow, O. (1985). Effects on growth of simulated and induced shoot pruning by tomicus piniperda as related to carbohydrate and nitrogen dynamics in scots pine. Journal of Applied Ecology, 22(1), 105-124. doi:10.2307/2403331

Esslinger, H. (0000). Ski- Heil.ARD Sendereihe Globus, Beiheft 1 Zur Sendung Vom 10., , 1988.

Estes, J. A., Terborgh, J., Brashares, J. S., Power, M. E., Berger, J., Bond, W. J., . . . Wardle, D. A. (2011). Trophic downgrading of planet earth. Science, 333(6040), 301-306. doi:10.1126/science.1205106

Evans, R. (1995). Impacts of reindeer grazing on soils and vegetation in finnmark. Impacts of Reindeer Grazing on Soils and Vegetation in Finnmark,

Feuerstein, G. C. (1997). Analyse Von Stammverletzungen Durch Alpensteinböcke (Capra i.Ibex) in Einem Subalpinen Lärchen-Árvenwald,

Formozov, A. N. (1946). Snow cover as an integral factor of the environment and its importance in the ecology of mammals and birds. New Series, Zoology, 5(20), 1-152.

Fox, J. L., & Smith, C. A. (1988). Winter mountain goat diets in southeast alaska. Journal of Wildlife Management, 52(2), 362-365. doi:10.2307/3801249

Franklin, J. F., & Dyrness, C. T. (1973). Natural vegetation of oregon and washington. Natural Vegetation of Oregon and Washington,

French, D. D., Miller, G. R., & Cummins, R. P. (1997). Recent development of high-altitude pinus sylvestris scrub in the northern cairngorm mountains, scotland. Biological Conservation, 79(2-3), 133-144. doi:10.1016/S0006-3207(96)00104-8

Furnier, G. R., Knowles, P., Clyde, M. A., & Dancik, B. P. (1987). Effects of avian seed dispersal on the genetic structure of whitebark pine populations. Evolution, 41(3), 607-612. doi:10.1111/j.1558-5646.1987.tb05831.x

Gampe, S. (1989). Über Sanierungsaufforstungen in Den Bayerischen Alpen,

Gill, R. (2000). The impact of deer on woodland biodiversity. The Impact of Deer on Woodland Biodiversity,

Gill, R. M. A. (1992). A review of damage by mammals in north temperate forests: 1. deer. Forestry, 65(2), 145-169. doi:10.1093/forestry/65.2.145

Gill, R. M. A. (1992). A review of damage by mammals in north temperate forests: 3. impact on trees and forests. Forestry, 65(4), 363-388. doi:10.1093/forestry/65.4.363-a

Gill, R. M. A. (1990). Monitoring the status of european and north american cervids. Monitoring the Status of European and North American Cervids,

Gill, R. M. A., & Beardall, V. (2001). The impact of deer on woodlands: The effects of browsing and seed dispersal on vegetation structure and composition. Forestry, 74(3), 209-218. doi:10.1093/forestry/74.3.209

Glutz von Blotzheim, U. N. (1993). Handbuch Der Vögel Mitteleuropas, Passeriformes, 13

Grabherr, G., Gottfried, M., & Pauli, H. (1994). Climate effects on mountain plants [7]. Nature, 369(6480), 448. doi:10.1038/369448a0

Grace, J., Berninger, F., & Nagy, L. (2002). Impacts of climate change on the tree line. Annals of Botany, 90(4), 537-544. doi:10.1093/aob/mcf222

Hagemoen, R. I. M., & Reimers, E. (2002). Reindeer summer activity pattern in relation to weather and insect harassment. Journal of Animal Ecology, 71(5), 883-892. doi:10.1046/j.1365-2656.2002.00654.x

Hall, K., Boelhouwers, J., & Driscoll, K. (1999). Animals as erosion agents in the alpine zone: Some data and observations from canada, lesotho, and tibet. Arctic, Antarctic, and Alpine Research, 31(4), 436-446. doi:10.2307/1552592

Hättenschwiler, S., & Smith, W. K. (1999). Seedling occurrence in alpine treeline conifers: A case study from the central rocky mountains, USA. Acta Oecologica, 20(3), 219-224. doi:10.1016/S1146-609X(99)80034-4

Heikkilä, R., & Härkönen, S. (1996). Moose browsing in young scots pine stands in relation to forest management. Forest Ecology and Management, 88(1-2), 179-186. doi:10.1016/S0378-1127(96)03823-6

Heikkinen, R. K., & Kalliola, R. (1989). Vegetation types and map of the kevo nature reserve, northernmost finland. Reports from the Kevo Subarctic Research Station, 11, 22-35.

Helle, R. (1966). An investigation of reindeer husbandry in finland. Acta Lapponica Fenniae, 5(5), 1-65.

Helle, T. (2001). Mountain Birch Forests and Reindeer Husbandry, , 279-291.

Helle, T., & Kojola, I. (2006). Population Trends of Semidomesticated Reindeer in Fennoscandia - Evaluation of Explanations, , 319-339.

Helle, T., & Tarvainen, L. (1984). Determination of the winter digging period of semi-domestic reindeer in relation to snow conditions and food resources. Reports - Kevo Subarctic Research Station, 19, 49-56.

Henden, J. -., Ims, R. A., Yoccoz, N. G., & Killengreen, S. T. (2011). Declining willow ptarmigan populations: The role of habitat structure and community dynamics. Basic and Applied Ecology, 12(5), 413-422. doi:10.1016/j.baae.2011.05.006

Herbold, H., Suchentrunk, F., Wagner, S., & Willing, R. (1992). The influence of anthropogenic disturbances on the heart frequency of red deer (cervus elaphus) and roe deer (capreolus capreolus). [Einfluß anthropogener Störreize auf die herzfrequenz von Rotwild (Cervus elaphus) und Rehwild (Capreolus capreolus)] Zeitschrift Für Jagdwissenschaft, 38(3), 145-159. doi:10.1007/BF02286509

Hess, A. (1916). Der Tannenhäher in Forstwirtschaftlicher Bedeutung, 67

Von Haller, H. (1996). Predation and accidents among alpine ibex capra ibex in the engadine. [Prädation und Unfälle beim Steinbock Capra ibex im Engadin] European Journal of Wildlife Research, 42(1), 26-35.





How to Cite

Holtmeier, F.-K. (2012). Impact of wild herbivorous mammals and birds on the altitudinal and northern treeline ecotones. Landscape Online, 30.



Research Article