Vertebrate Herbivory and Its Ecosystem Consequences


Herbivory is a process where animals obtain energy and nutrients from vegetative plant parts (leaves, stems, etc.). Herbivory is called grazing, browsing or folivory, depending on the size of the herbivore and the type of plant tissue consumed. The consumption is actually performed by microbes in the digestive system. The role of the animal is to prepare the material and to provide a good environment for the microbes. This requires specialisations, making herbivores a distinct animal group. Vertebrate herbivory can change forests to heathlands or grasslands and influence the composition and species richness of the herbaceous vegetation. By maintaining open habitats, herbivorous vertebrates can increase the fraction of solar radiation reflected back to the space, especially at high latitudes and altitudes. Thus, vertebrate herbivory can also counteract global warming.

Key Concepts

  • Herbivory is the collective term for consumption of plants or vegetative plant parts by animals.
  • Browsing is utilisation of woody plants by animals, normally referring to vertebrates.
  • Grazing is utilisation of herbaceous plants by animals, referring to vertebrates and big, mobile invertebrates.
  • Folivory is utilisation of photosynthetic plant tissues by small animals, normally referring to relatively immobile insect larvae.
  • Heathland, in narrow sense, refers to treeless vegetation, dominated by ericaceous plants; in broad sense, it refers to all kinds of vegetation dominated by prostrate dicotyledonous plants.
  • Grassland is treeless vegetation dominated by plants belonging to the families Poaceae (grasses), Cyperaceae (sedges and their relatives) or Juncaceae (rushes and woodrushes).

Keywords: competition; consumption; plants; species richness; vegetation; mammals; tundra

Figure 1. Tall, scattered trees in subalpine (a, from French Alps) and subarctic (b, from northern Norway) landscapes, shaped out by centuries of intense herbivory by cattle, sheep and goats (a) or reindeer (b). Photos were taken by Lauri and Tarja Oksanen.
Figure 2. The contrast between summer range, subjected to periodically intense reindeer grazing in late summer (right) and lightly grazed autumn range (left) on the mountains of northern Norway mountains. The fence across which the picture is taken is 40 years old. Photos were taken by Lauri and Tarja Oksanen.
Figure 3. Two variants of moist and nutrient‐rich low alpine meadow. Panel (a) shows an area abandoned 40 years ago when the border fence of the legal summer range was constructed; panel (b) is taken from the intensely grazed summer range. Photos were taken by Lauri and Tarja Oksanen.
Figure 4. The vegetation of a moist and nutrient‐rich low Arctic willow scrubland in the mainland (a) and of an initially similar habitat on an isolated island, where grey‐sided voles were introduced in 1991 (b; photo taken in July 2008). Photos were taken by Lauri and Tarja Oksanen.


Aunapuu M, Dahlgren J, Oksanen T, et al. (2008) Spatial patterns and dynamic responses of arctic food webs corroborate the exploitation ecosystems hypothesis (EEH). American Naturalist 171: 249–262.

Bakker ES, Olff H, Boekhoff M, et al. (2004) Impact of herbivores on nitrogen cycling: contrasting effects of small and large species. Oecologia 138: 91–101.

Bakker ES, Ritchie ME, Olff H, Milchunas DG and Knops JMH (2006) Herbivore impact on grasland plant diversity depends on habitat productivity and herbivore size. Ecology Letters 9: 780–788.

Biuw M, Jepsen JU, Cohen J, et al. (2014) Long‐term impacts of contrasting management of large ungulates in the arctic tundra‐forest ecotone: ecosystem structure and climate feedback. Ecosystems 17: 890–905.

Bond WJ, Woodward FI and Midgley GG (2005) The global distribution of ecosystems in a world without fire. New Phytologist 165: 525–538.

Bråthen KA, Ims RA, Yoccoz NG, et al. (2007) Induced shift in ecosystem productivity? Extensive scale effects of abundant large herbivores. Ecosystems 10: 773–789.

Brown JS (1999) Vigilance, patch use and habitat selection: foraging under predation risk. Evolutionary Ecology Research 1: 49–71.

Chapin FS, Sturm M, Serreze MC, et al. (2005) Role of land‐surface changes in arctic summer warming. Science 310: 657–660.

Chouinard A and Filion L (2005) Impact of introduced white‐tailed deer and native insect defoliators on the density and growth of conifer saplings on Anticosti Island, Quebec. Ecoscience 12: 506–518.

Connell JH (1978) Diversity in tropical rain forest and coral reefs. Science 199: 1302–1310.

Crête M (1999) The distribution of deer biomass supports the hypothesis of exploitation ecosystems. Ecology Letters 2: 223–227.

Crête M and Doucet DJ (1998) Persistent suppression in dwarf birch after release from heavy summer browsing by caribou. Arctic and Alpine Research 30: 126–132.

Dahl E (1957) Rondane: mountain vegetation in South Norway and its relation to the environment. Skrifter av den Norske Videnskaps‐Akademin, I: Matematisk‐Naturvidenskaplig Klasse 1956 (3): 1–347.

Dahlgren J, Oksanen L, Olofsson J and Oksanen T (2009) Plant defenses at no cost? The recovery of tundra scrubland following heavy grazing by gray‐sided voles (Myodes rufocanus). Evolutionary Ecology Research 11: 1205–1216.

Ehrlich PR and Raven PH (1964) Butterflies and plants: a study in coevolution. Evolution 18: 586–608.

Ellenberg H (1988) Vegetation Ecology of Central Europe, 4th edn. Cambridge, UK: Cambridge University Press.

Eskelinen A (2007) Herbivore and neighbour effects on tundra plants depend on species identity, nutrient availability and local environmental conditions. Journal of Ecology 96: 155–165.

Estes J (ed) (2009) The Science of Trophic Cascades. Washington, DC: Island Press.

Fortin D, Beyer HL, Boyce MS, et al. (2005) Wolves influence elk movements: behavior shapes a trophic cascade in Yellowstone National Park. Ecology 86: 1320–1330.

Gimingham CH (1972) Ecology of Heathlands. London: Chapman & Hall.

Grime JP (1973) Competitive exclusion in herbaceous vegetation. Nature 242: 344–347.

Grime JP (1979) Plant Strategies and Vegetation Processes. Chichester: John Wiley & Sons, Ltd.

Hairston NG, Smith FE and Slobodkin LB (1960) Community structure, population control, and competition. American Naturalist 149: 421–425.

Harsh MA, Hulme PE, McGlone MS and Duncan RP (2009) Are treelines advancing? A global meta‐analysis of treeline response to climate warming. Ecology Letters 12: 1040–1049.

Hansen BB, Henriksen S, Aanes R and Sæther B‐E (2007) Ungulate impact on vegetation in a two trophic level system. Polar Biology 30: 549–558.

Hansson L (1985) Damage by wildlife, especially small rodents, to North American Pinus contorta provenances introduced into Sweden. Canadian Journal of Forest Research 15: 1167–1171.

Holt RD (1977) Predation, apparent competition, and the structure of prey communities. Theoretical Population Biology 12: 276–290.

Hoset KH, Ruffino L, Tuomi M, et al. (2017) Changes in the spatial configuration and strength of trophic control across a productivity gradient during a massive rodent outbreak. Ecosystems 20: 1421–1435. 10.1007/s10021-017-0124-1.

Janzen DH (1970) Herbivores and the number of tree species in tropical forests. American Naturalist 104: 501–528.

Jepsen JU, Biuw M, Ims RA, et al. (2013) Ecosystem impacts of a range expanding forest defoliator at the forest‐tundra ecotone. Ecosystems 16: 561–575.

Kallio P and Lehtonen J (1975) On the ecocatastrophe of birch forests caused by Oporinia autumnata. In: Wielgolaski FE (ed) Fennoscandian Tundra Ecosystems. Part 2. Animals and Systems Analysis (Ecological Studies 17), pp. 174–180. Berlin: Springer‐Verlag.

Körner C (2012) Alpine Treelines: Functional Ecology of the Global High Elevation Tree Limits. Basel: Springer Verlag.

Kurtén B (1971) The Age of Mammals. London: Weidenfeld & Nicholson.

Murdoch WW (1966) Community structure, population control, and competition – a critique. American Naturalist 100: 219–226.

Nilsson M‐C, Zackrisson O, Sterner O and Wallstedt A (2000) Characterisation of the differential interference effects of two boreal dwarf shrub species. Oecologia 123: 122–128.

Oksanen L, Fretwell SD, Arruda J and Niemelä P (1981) Exploitation ecosystems in gradients of primary productivity. American Naturalist 118: 240–261.

Oksanen L (1990a) Predation, herbivory, and plant strategies along gradients of primary productivity. In: Tilman D and Grace J (eds) Perspectives on Plant Consumption, pp. 445–474. New York: Academic Press.

Oksanen T (1990b) Exploitation ecosystems in heterogeneous habitat complexes. Evolutionary Ecology 4: 220–234.

Oksanen L and Oksanen T (2000) The logic and realism of the hypothesis of exploitation ecosystems. American Naturalist 155: 703–723.

Oksanen L, Oksanen T, Dahlgren J, et al. (2010) Islands as tests of the green world hypothesis. In: Terborgh J and Estes JA (eds) Trophic Cascades: Predators, Prey, and the Changing Dynamics of Nature, pp. 163–177. Washington, DC: Island Press.

Oksanen T, Oksanen L, Dahlgren J and Olofsson J (2008) Arctic lemmings, Lemmus spp. and Dicrostonyx spp.: integrating ecological and evolutionary perspectives. Evolutionary Ecology Research 10: 415–434.

Olff H, De Leeuw J, Bakker JP, et al. (1997) Vegetation succession and herbivory on a salt marsh: changes induced by sea level rise and silt deposition along an elevational gradient. Journal of Ecology 85: 799–814.

Olofsson J, Hulme PE, Oksanen L and Suominen O (2004a) Importance of large and small mammalian herbivores for the plant community structure in the forest tundra ecotone. Oikos 106: 324–334.

Olofsson J, Kitti H, Rautiainen P, Stark S and Oksanen L (2001) Impact of summer grazing by reindeer on vegetation structure, productivity and nutrient cycling in the North Fennoscandian tundra. Ecography 24: 13–24.

Olofsson J and Oksanen L (2005) Effects of reindeer density on plant diversity in the Fennoscandian mountain chain. In: Haugerud RE (ed) Proceedings of eleventh Arctic Ungulate Congress, Saariselkä, Finland, 2003 (Rangifer 25). pp. 5–18.

Olofsson J, Oksanen L, Callaghan T, et al. (2009) Herbivores inhibit climate driven shrub expansion on the tundra. Global Change Biology 15: 2681–2693.

Olofsson J, Oksanen L, Oksanen T, et al. (2014) Long‐term experiments reveal strong interactions between lemmings and plants in the Fennoscandian highland tundra. Ecosystems 17: 606–615.

Olofsson J, Stark S and Oksanen L (2004b) Herbivore influence on ecosystem processes in tundra. Oikos 105: 386–396.

Olofsson J, Tømmervik H and Callaghan TV (2012) Vole and lemming activity observed from space. Nature Climate Change 2: 880–883.

Post E and Pedersen C (2008) Opposing plant community responses to warming with and without herbivores. PNAS 105: 12353–12358.

Ripple WJ and Beschta RL (2003) Wolf reintroduction, predation risk, and cottonwood recovery in Yellowstone National Park. Forest Ecology and Management 184: 299–313.

Ripple WJ and Beschta RL (2006) Linking a cougar decline, trophic cascade and catastrophic regime shift in Zion National Park. Biological Conservation 133: 397–408.

Ripple WJ, Rooney TP and Beschta RL (2010) Large predators, deer, and trophic cascades in boreal and temperate ecosystems. In: Terborgh J and Estes JA (eds) Trophic Cascades, pp. 141–161. Washington, DC: Island Press.

Roem WJ and Berendse F (2000) Soil acidity and nutrient supply ratio as possible factors determining changes in plant species diversity in grassland and heathland communities. Biological Conservation 92: 151–161.

Rosén E (1982) Vegetation development and sheep grazing in limestone grasslands of South Öland, Sweden. Acta Phytogeographica Suecica 72: 1–104.

Ruffino L, Oksanen T, Hoset KS, et al. (2016) Predator‐rodent‐plant interactions along a coast‐inland gradient in Fennoscandian tundra. Ecography 39: 871–883. DOI: 10.1111/ecog.01758.

Saccone P and Virtanen R (2016) Extrapolating multi‐decadal plant community changes based on medium‐term experiments can be risky: evidence from high‐latitude tundra. Oikos 125: 76–85.

Schrama MJJ, Veen GF, Bakke ES, et al. (2013) An integrated perspective to explain nitrogen mineralization in grazed ecosystems. Perspectives in Plant Ecology, Evolution and Systematics 15: 32–44.

Siira J (1985) Saline soils and their vegetation on the coast of the Gulf of Bothnia, Finland. Annales Botanici Fennici 22: 63–90.

Speed JDM, Martinsen V, Mysterud A, et al. (2014) Long‐term increase in aboveground carbon stocks following exclusion of grazers and forest establishment in an alpine ecosystem. Ecosystems 17: 1138–1150. DOI: 10.1007/s10021-014-9784-2.

Stebbins GL (1981) Coevolution of grasses and herbivores. Annals of the Missouri Botanical Garden 68: 75–86.

Tansley AG (1939) The British Islands and their Vegetation. Cambridge, UK: Cambridge University Press.

Tape K, Sturm M and Racine C (2006) The evidence for shrub expansion in northern Alaska and the Pan‐Arctic. Global Change Biology 12: 686–702.

Tenow O (1972) The outbreaks of Oporinia autumnata (Bhk.) and Opherophtera spp. (Lep., Geometridae) in the Scandinavian mountain chain and northern Finland 1862–1968. Zoologiska Bidrag från Uppsala Universitet 2 (suppl): 1–107.

Terborgh J, Lopez L, Nuňez PV, et al. (2001) Ecological meltdown in predator‐free forest fragments. Science 294: 1923–1926.

Terborgh J, Feeley K, Silman M, Nuňez PV and Balukjan B (2006) Vegetation dynamics on predator‐free land bridge islands. Journal of Ecology 94: 253–263.

Terborgh J and Feeley K (2010) Propagation of trophic cascades via multiple pathways in tropical forests. In: Terborgh J and Estes JA (eds) Trophic Cascades: Predators, Prey, and the Changing Dynamics of Nature, pp. 125–140. Washington DC: Island Press.

Tihomirov BA (1959) Vzajmosvjazi životnogo mira i rastitel'nogo pokrova tundry. Moscow: Botaničeskij Institut VL Komarova, Akademija Nauk SSSR.

Tilman D (1988) Plant Strategies and the Structure and Dynamics of Plant Communities. Princeton, NJ: Princeton University Press.

Turchin P, Oksanen L, Ekerholm P, Oksanen T and Henttonen H (2000) Are lemmings prey or predators? Nature 405: 562–565.

Virtanen RJ (1998) Impact of grazing and neighbour removal on a heath plant community transplanted onto a snowbed site, NW Finnish Lapland. Oikos 81: 359–367.

Virtanen RJ (2000) Effects of grazing on above‐ground biomass on a mountain snowbed, NW Finland. Oikos 90: 295–300.

Virtanen RJ, Henttonen H and Laine K (1997) Lemming grazing and structure of a snowbed plant community: a long‐term experiment at Kilpisjärvi, Finnish Lapland. Oikos 79: 155–166.

Virtanen R, Oksanen L, Oksanen T, et al. (2016) Where do the treeless tundra areas of northern highlands fit in the global biome system: toward an ecologically natural subdivision of the tundra biome. Ecology and Evolution 6: 143–158.

Walter H (1964) Die Vegetation der Erde in öko‐physiologischer Betrachtung: I Die tropischen und subtropischen Zonen. Jena: Gustav Fischer Verlag.

Walter H (1968) Die Vegetation der Erde in öko‐physiologischer Betrachtung. II Die gemäßigten und arktischen Zonen. Jena: Gustav Fischer Verlag.

Welch DB, Staines W, Scott D, French DD and Catt DC (1991) Leader browsing by red and roe deer on young Sitka spruce trees in western Scotland. 1. Damage rates and the influence of habitat factors. Forestry 64: 61–82.

Ylänne H, Olofsson J, Oksanen L and Stark S (2018) Consequences of grazer‐induced vegetation transitions on ecosystem carbon storage in the tundra. Functional Ecology. DOI: 10.1111/1365-2435.13029.

Zimov SA, Chuprynin VI, Oreshko AP, et al. (1995) Steppe‐tundra transition: a herbivore‐driven biome shift at the end of the Pleistocene. American Naturalist 146: 765–794.

Further Reading

Crawley MJ (1986) Herbivory: The Dynamics of Plant Animal Interactions. Oxford, UK: Blackwell Science.

Crawley MJ (1997) Plant‐herbivore dynamics. In: Crawley MJ (ed) Plant Ecology, pp. 284–324. Oxford, UK: Blackwell Science.

Hartley SE (2001) Plant interactions with biotic factors. Encyclopedia of Life Sciences. Chichester: John Wiley & Sons, Ltd. [DOI: 10.1038/npg.els.0003203]

Hartley SE and Jones CG (1997) Plant chemistry and herbivory, or why the world is green. In: Crawley MJ (ed) Plant Ecology, pp. 284–324. Oxford, UK: Blackwell Science.

Huntly N (1991) Herbivores and the dynamics of communities and ecosystems. Annual Review of Ecology and Systematics 22: 477–503.

Olff H and Ritchie ME (1998) Effects of herbivores on grassland plant diversity. Trends in Ecology and Evolution 13: 261–265.

Schardl CL (2001) Plant defences against herbivore and insect attack. Encyclopedia of Life Sciences. Chichester: John Wiley & Sons, Ltd. [DOI: 10.1038/npg.els.0001324]

Whitney HM and Beverley GJ (2007) Coevolution: plant–insect. Encyclopedia of Life Sciences. Chichester: John Wiley & Sons, Ltd. [DOI: 10.1002/9780470015902.a0001762]

Wink M (2001) Secondary Metabolites: dettering herbivores. Encyclopedia of Life Sciences. Chichester: John Wiley & Sons, Ltd. [DOI: 10.1038/npg.els.0000918]

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Oksanen, Lauri, and Olofsson, Johan(Apr 2018) Vertebrate Herbivory and Its Ecosystem Consequences. In: eLS. John Wiley & Sons Ltd, Chichester. [doi: 10.1002/9780470015902.a0003283.pub2]