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Further information: Vegetation type
A map of different terrestrial biomes around the world.

A biome /ˈbm/ is a formation of plants and animals that have common characteristics due to similar climates and can be found over a range of continents.[1] Spanning continents, Biomes are distinct biological communities that have formed in response to a shared physical climate.[2] Biomes are distinct from habitats, because any biome can comprise a variety of habitats.[citation needed]

A biome contrasts with a microbiome. A microbiome is also a mix of organisms that coexist in a defined space, but on a much smaller scale. For example, the human microbiome is the collection of bacteria, viruses, and other microorganisms that are present on a human.[citation needed]

History of the concept[edit]

The term was suggested in 1916 by Clements, originally as a synonym for biotic community of Möbius (1877).[3] Later, it gained its current definition, based on earlier concepts of phytophysiognomy, formation and vegetation (used in opposition to flora), with the inclusion of the animal element and the exclusion of the taxonomic element of species composition.[4][5] In 1935, Tansley would add the climatic and soil aspects to the idea, calling it ecosystem.[6][7] The International Biological Program (1964-74) projects would popularize the concept of biome.[8]

However, in some contexts, the term biome is used in a different manner. In German literature, particularly in the Walter terminology, the term is used similarly as biotope (a concrete geographycal unit), while the biome definition used in this article is used as a international, non-regional, terminology - irrespectively of the continent in which an area is present, it takes the same biome name - and corresponds to his "zonobiome", "orobiome" and "pedobiome" (biomes determinated by climate zone, altitude or soil).[9]

In Brazilian literature, the term "biome" is sometimes used as synonym of "biogeographic province", an area based on species composition (the term "floristic province" being used when plant species are considered), or also as synonym of the "morphoclimatic and phytogeographical domain" of Ab'Sáber, a geographic space with subcontinental dimensions, with the predominance of similar geomorphologic and climatic characteristics, and of a certain vegetation form. Both includes many biomes in fact.[10][4][11]


Biomes are defined by climate regimes and biogeography.

A 1978 study on North American grasslands[12] found a positive logistic correlation between evapotranspiration in mm/yr and above-ground net primary production in g/m2/yr. The general results from the study were that precipitation and water use led to above-ground primary production, while solar irradiation and temperature lead to below-ground primary production (roots), and temperature and water lead to cool and warm season growth habit.[13] These findings help explain the categories used in Holdridge’s bioclassification scheme (see below), which were then later simplified by Whittaker. The number of classification schemes and the variety of determinants used in those schemes, however, should be taken as strong indicators that biomes do not fit perfectly into the classification schemes created.

Holdridge (1947, 1964) life zones[edit]

Main article: Holdridge life zones

Holdridge classified climates based on the biological effects of temperature and rainfall on vegetation under the assumption that these two abiotic factors are the largest determinants of the types of vegetation found in a habitat. Holdridge uses the four axes to define 30 so-called "humidity provinces", which are clearly visible in his diagram. While this scheme largely ignores soil and sun exposure, Holdridge acknowledged that these were important.

Allee (1949) biome-types[edit]

The principal biome-types by Allee (1949):[14]

  • Tundra
  • Taiga
  • Deciduous forest
  • Grasslands
  • Desert
  • High plateaus
  • Tropical forest
  • Minor terrestrial biomes

Kendeigh (1961) biomes[edit]

The principal biomes of the world by Kendeigh (1961):[15]

  • Terrestrial
    • Temperate deciduous forest
    • Coniferous forest
    • Woodland
    • Chaparral
    • Tundra
    • Grassland
    • Desert
    • Tropical savanna
    • Tropical forest
  • Marine
    • Oceanic plankton and nekton
    • Balanoid-gastropod-thallophyte
    • Pelecypod-annelid
    • Coral reef

Whittaker (1962, 1970, 1975) biome-types[edit]

The distribution of vegetation types as a function of mean annual temperature and precipitation.

Whittaker classified biomes using two abiotic factors: precipitation and temperature. His scheme can be seen as a simplification of Holdridge's; more readily accessible, but missing Holdridge's greater specificity.

Whittaker based his approach on theoretical assertions and empirical sampling. He was in a unique position to make such a holistic assertion because he had previously compiled a review of biome classifications.[16]

Key definitions for understanding Whittaker's scheme[edit]

  • Physiognomy: the apparent characteristics, outward features, or appearance of ecological communities or species.
  • Biome: a grouping of terrestrial ecosystems on a given continent that are similar in vegetation structure, physiognomy, features of the environment and characteristics of their animal communities.
  • Formation: a major kind of community of plants on a given continent.
  • Biome-type: grouping of convergent biomes or formations of different continents, defined by physiognomy.
  • Formation-type:a grouping of convergent formations.

Whittaker's distinction between biome and formation can be simplified: formation is used when applied to plant communities only, while biome is used when concerned with both plants and animals. Whittaker's convention of biome-type or formation-type is simply a broader method to categorize similar communities.[17]

Whittaker's parameters for classifying biome-types[edit]

Whittaker, seeing the need for a simpler way to express the relationship of community structure to the environment, used what he called "gradient analysis" of ecocline patterns to relate communities to climate on a worldwide scale. Whittaker considered four main ecoclines in the terrestrial realm.[17]

  1. Intertidal levels: The wetness gradient of areas that are exposed to alternating water and dryness with intensities that vary by location from high to low tide
  2. Climatic moisture gradient
  3. Temperature gradient by altitude
  4. Temperature gradient by latitude

Along these gradients, Whittaker noted several trends that allowed him to qualitatively establish biome-types:

  • The gradient runs from favorable to extreme, with corresponding changes in productivity.
  • Changes in physiognomic complexity vary with how favorable of an environment exists (decreasing community structure and reduction of stratal differentiation as the environment becomes less favorable).
  • Trends in diversity of structure follow trends in species diversity; alpha and beta species diversities decrease from favorable to extreme environments.
  • Each growth-form (i.e. grasses, shrubs, etc.) has its characteristic place of maximum importance along the ecoclines.
  • The same growth forms may be dominant in similar environments in widely different parts of the world.

Whittaker summed the effects of gradients (3) and (4) to get an overall temperature gradient, and combined this with gradient (2), the moisture gradient, to express the above conclusions in what is known as the Whittaker classification scheme. The scheme graphs average annual precipitation (x-axis) versus average annual temperature (y-axis) to classify biome-types.


  • 1. Tropical rainforest
  • 2. Tropical seasonal rainforest
    • deciduous
    • semideciduous
  • 3. Temperate giant rainforest
  • 4. Montane rainforest
  • 5. Temperate deciduous forest
  • 6. Temperate evergreen forest
    • needleleaf
    • sclerophyll
  • 7. Subarctic-subalpin needle-leaved forests (taiga)
  • 8. Elfin woodland
  • 9. Thorn forests and woodlands
  • 10. Thorn scrub
  • 11. Temperate woodland
  • 12. Temperate shrublands
    • deciduous
    • heath
    • sclerophyll
    • subalpine-needleleaf
    • subalpine-broadleaf
  • 13. Savanna
  • 14. Temperate grassland
  • 15. Alpine grasslands
  • 16. Tundra
  • 17. Tropical desert
  • 18. Warm-temperate desert
  • 19. Cool temperate desert scrub
  • 20. Arctic-alpine desert
  • 21. Bog
  • 22. Tropical fresh-water swamp forest
  • 23. Temperate fresh-water swamp forest
  • 24. Mangroove swamp
  • 25. Saltmarsh[18]

Goodall (1974-) ecosystem types[edit]

The multiauthored series Ecosystems of the world, edited by David W. Goodall, provides a comprehensive coverage of the major "ecosystem types or biomes" on earth:[19]

  • I. Terrestrial Ecosystems
    • A. Natural Terrestrial Ecosystems
      • 1. Wet Coastal Ecosystems
      • 2. Dry Coastal Ecosystems
      • 3. Polar and Alpine Tundra
      • 4. Mires: Swamp, Bog, Fen and Moor
      • 5. Temperate Deserts and Semi-Deserts
      • 6. Coniferous Forests
      • 7. Temperate Deciduous Forests
      • 8. Natural Grasslands
      • 9. Heathlands and Related Shrublands
      • 10. Temperate Broad-Leaved Evergreen Forests
      • 11. Mediterranean-Type Shrublands
      • 12. Hot Deserts and Arid Shrublands
      • 13. Tropical Savannas
      • 14. Tropical Rain Forest Ecosystems
      • 15. Wetland Forests
      • 16. Ecosystems of Disturbed Ground
    • B. Managed Terrestrial Ecosystems
      • 17. Managed Grasslands
      • 18. Field Crop Ecosystems
      • 19. Tree Crop Ecosystems
      • 20. Greenhouse Ecosystems
      • 21. Bioindustrial Ecosystems
  • II. Aquatic Ecosystems
    • A. Inland Aquatic Ecosystems
      • 22. River and Stream Ecosystems
      • 23. Lakes and Reservoirs
    • B. Marine Ecosystems
      • 24. Intertidal and Littoral Ecosystems
      • 25. Coral Reefs
      • 26. Estuaries and Enclosed Seas
      • 27. Ecosystems of the Continental Shelves
      • 28. Ecosystems of the Deep Ocean
    • C. Managed Aquatic Ecosystems
      • 29. Managed Aquatic Ecosystems
  • III. Underground Ecosystems
    • 30. Cave Ecosystems

Walter (1976, 2002) zonobiomes[edit]

The eponymously-named Heinrich Walter classification scheme considers the seasonality of temperature and precipitation. The system, also assessing precipitation and temperature, finds nine major biome types, with the important climate traits and vegetation types. The boundaries of each biome correlate to the conditions of moisture and cold stress that are strong determinants of plant form, and therefore the vegetation that defines the region. Extreme conditions, such as flooding in a swamp, can create different kinds of communities within the same biome.[20][21][9]

Zonobiome Zonal soil type Zonal vegetation type
ZB I. Equatorial, always moist, little temperature seasonality Equatorial brown clays Evergreen tropical rainforest
ZB II. Tropical, summer rainy season and cooler “winter” dry season Red clays or red earths Tropical seasonal forest, seasonal dry forest, scrub, or savanna
ZB III. Subtropical, highly seasonal, arid climate Serosemes, sierozemes Desert vegetation with considerable exposed surface
ZB IV. Mediterranean, winter rainy season and summer drought Mediterranean brown earths Sclerophyllous (drought-adapted), frost-sensitive shrublands and woodlands
ZB V. Warm temperate, occasional frost, often with summer rainfall maximum Yellow or red forest soils, slightly podsolic soils Temperate evergreen forest, somewhat frost-sensitive
ZB VI. Nemoral, moderate climate with winter freezing Forest brown earths and grey forest soils Frost-resistant, deciduous, temperate forest
ZB VII. Continental, arid, with warm or hot summers and cold winters Chernozems to serozems Grasslands and temperate deserts
ZB VIII. Boreal, cold temperate with cool summers and long winters Podsols Evergreen, frost-hardy, needle-leaved forest (taiga)
ZB IX. Polar, short, cool summers and long, cold winters Tundra humus soils with solifluction (permafrost soils) Low, evergreen vegetation, without trees, growing over permanently frozen soils

Schültz (1988) ecozones[edit]

Schültz (1988) defined nine ecozones (note that his concept of ecozone is more similar to the concept of biome used in this article than to the concept of ecozone of BBC):[22]

  • 1. polar/subpolar zone
  • 2. boreal zone
  • 3. humid mid-latitudes
  • 4. arid mid-latitudes
  • 5. tropical/subtropical arid lands
  • 6. Mediterranean-type subtropics
  • 7. seasonal tropics
  • 8. humid subtropics
  • 9. humid tropics

Bailey (1989) ecoregions[edit]

Robert G. Bailey nearly developed a biogeographical classification system of ecoregions for the United States in a map published in 1976. He subsequently expanded the system to include the rest of North America in 1981, and the world in 1989. The Bailey system, based on climate, is divided into seven domains (polar, humid temperate, dry, humid, and humid tropical), with further divisions based on other climate characteristics (subarctic, warm temperate, hot temperate, and subtropical; marine and continental; lowland and mountain).[23][24]

  • 100 Polar Domain
    • 120 Tundra Division (Köppen: Ft)
    • M120 Tundra Division – Mountain Provinces
    • 130 Subarctic Division (Köppen: E)
    • M130 Subarctic Division – Mountain Provinces
  • 200 Humid Temperate Domain
    • 210 Warm Continental Division (Köppen: portion of Dcb)
    • M210 Warm Continental Division – Mountain Provinces
    • 220 Hot Continental Division (Köppen: portion of Dca)
    • M220 Hot Continental Division – Mountain Provinces
    • 230 Subtropical Division (Köppen: portion of Cf)
    • M230 Subtropical Division – Mountain Provinces
    • 240 Marine Division (Köppen: Do)
    • M240 Marine Division – Mountain Provinces
    • 250 Prairie Division (Köppen: arid portions of Cf, Dca, Dcb)
    • 260 Mediterranean Division (Köppen: Cs)
    • M260 Mediterranean Division – Mountain Provinces
  • 300 Dry Domain
    • 310 Tropical/Subtropical Steppe Division
    • M310 Tropical/Subtropical Steppe Division – Mountain Provinces
    • 320 Tropical/Subtropical Desert Division
    • 330 Temperate Steppe Division
    • 340 Temperate Desert Division
  • 400 Humid Tropical Domain
    • 410 Savanna Division
    • 420 Rainforest Division

Olson & Dinerstein (1998) biomes for WWF / Global 200[edit]

Main article: Global 200

A team of biologists convened by the World Wildlife Fund (WWF) developed a scheme that divided the world's land area into biogeographic realms (called "ecozones" in a BBC scheme), and these into ecoregions (Olson & Dinerstein, 1998, etc.). Each ecoregion is characterized by a main biome (also called major habitat type).[25][26]

This classification is used to define the Global 200 list of ecoregions identified by the WWF as priorities for conservation.[25]

For the terrestrial ecoregions, there is a specific EcoID, format XXnnNN (XX is the biogeographic realm, nn is the biome number, NN is the individual number).

Biogeographic realms (terrestrial and freshwater)[edit]

It should be noted, however, that the applicability of the realms scheme above - based on Udvardy (1975) - to most freshwater taxa is unresolved.[27]

Biogeographic realms (marine)[edit]

Biomes (terrestrial)[edit]

Biomes (freshwater)[edit]

According to the WWF, the following are classified as freshwater biomes:[29]

Biomes (marine)[edit]

Biomes of the coastal and continental shelf areas (neritic zone):

Summary of the scheme[edit]


Other biomes[edit]

Marine biomes[edit]

Pruvot (1896) zones or "systems":[31]

Longhurst (1998) biomes:[32]

  • Coastal
  • Polar
  • Trade wind
  • Westerly

Other marine habitat types (not covered yet by the Global 200/WWF scheme):[citation needed]

Anthropogenic biomes[edit]

Humans have altered global patterns of biodiversity and ecosystem processes. As a result, vegetation forms predicted by conventional biome systems can no longer be observed across much of Earth's land surface as they have been replaced by crop and rangelands or cities. Anthropogenic biomes provide an alternative view of the terrestrial biosphere based on global patterns of sustained direct human interaction with ecosystems, including agriculture, human settlements, urbanization, forestry and other uses of land. Anthropogenic biomes offer a new way forward in ecology and conservation by recognizing the irreversible coupling of human and ecological systems at global scales and moving us toward an understanding of how best to live in and manage our biosphere and the anthropogenic biomes we live in.

Major anthropogenic biomes:

Microbial biomes[edit]

Further information: Microhabitats

Endolithic biomes[edit]

The endolithic biome, consisting entirely of microscopic life in rock pores and cracks, kilometers beneath the surface, has only recently been discovered, and does not fit well into most classification schemes.[citation needed]

Dermal biome[edit]

The dermal biome is the living ecosystem that animals (including humans) have evolved, that permits them to live symbiotically and in balance with the microbes on and in them (the microbiome). This ecosystem consists of skin, follicles, hair, sebaceous glands, sweat glands, arrector pili muscles, peptides, proteins, lipids and its associated microbiota. A healthy dermal biome has several functions: it resists infection of pathogens, protects against moisture loss and water damage, dynamically regulates body temperature and supports the healthy renewal of skin through the epidermal cell life cycle.[citation needed]

  • Infection Resistance: Commensal microbiota assist the dermal biome resist infection for pathogenic bacteria by i] out-competing pathogens for resources, ii] training or stimulating the host’s immune system to defeat the pathogen, or iii] expressing substances that are directly hostile to the pathogen.
  • Water-barrier regulation: The dermal biome regulates the water barrier – preventing moisture from escaping (except when expressed as sweat) and preventing environmental water from permeating the skin. Because environmental water can have a chilling effect to mammals and warm-blooded animals when kept in close proximity to the epidermis, the dermal biome also produces hydrophobic lipids that repel water.
  • Temperature regulation: The dermal biome is responsible for thermoregulation. To regulate excess heat, the dermal biome activates the sweat glands, allowing for evaporative cooling as sweat evaporates. To regulate cooler temperatures, the arrector pili muscles contract, causing hairs to “stand up” (goosebumps), and thereby trap an insulating blanket of air close to the skin.
  • Skin renewal: a healthy biome supports the replacement of skin through the life cycle of epidermal cells as they proliferate in the basal layers of the epidermis until they die and are shed (desquamation).

See also[edit]


  1. ^ The World's Biomes, Retrieved August 19, 2008, from University of California Museum of Paleontology
  2. ^ Cain, Michael; Bowman, William; Hacker, Sally (2014). Ecology (Third Edition ed.). Massachusetts: Sinauer. p. 51. ISBN 9780878939084. 
  3. ^ Clements, F. E. 1917. The development and structure of biotic communities. J. Ecology 5:120–121. Abstract of a talk in 1916, [1].
  4. ^ a b Coutinho, L. M. (2006). O conceito de bioma. Acta Bot. Bras. 20(1): 13-23, [2].
  5. ^ Martins, F. R. & Batalha, M. A. (2011). Formas de vida, espectro biológico de Raunkiaer e fisionomia da vegetação. In: Felfili, J. M., Eisenlohr, P. V.; Fiuza de Melo, M. M. R.; Andrade, L. A.; Meira Neto, J. A. A. (Org.). Fitossociologia no Brasil: métodos e estudos de caso. Vol. 1. Viçosa: Editora UFV. p. 44-85. [3]. Earlier version, 2003, [4].
  6. ^ Cox, C. B., Moore, P.D. & Ladle, R. J. 2016. Biogeography: an ecological and evolutionary approach. 9th edition. John Wiley & Sons: Hoboken, p. 20, [5].
  7. ^ Tansley, A.G. (1935). The use and abuse of vegetational terms and concepts. Ecology 16 (3): 284–307, [6].
  8. ^ Box, E.O. & Fujiwara, K. (2005). Vegetation types and their broad-scale distribution. In: van der Maarel, E. (ed.). Vegetation ecology. Blackwell Scientific, Oxford. pp 106–128, [7].
  9. ^ a b Walter, H. & Breckle, S-W. (2002). Walter's Vegetation of the Earth: The Ecological Systems of the Geo-Biosphere. New York: Springer-Verlag, p. 86, [8].
  10. ^ Batalha, M.A. (2011). The Brazilian cerrado is not a biome. Biota Neotrop. 11:21–4, [9].
  11. ^ Fiaschi, P.; Pirani, J.R. 2009. Review of plant biogeographic studies in Brazil. Journal of Systematics and Evolution, v. 47, p. 477-496. Disponível em: <>.
  12. ^ Sims, Phillip L.; Singh, J.S. (July 1978). "The Structure and Function of Ten Western North American Grasslands: III. Net Primary Production, Turnover and Efficiencies of Energy Capture and Water Use". Journal of Ecology. British Ecological Society. 66 (2): 573–597. doi:10.2307/2259152. 
  13. ^ Pomeroy, Lawrence R. and James J. Alberts, editors. Concepts of Ecosystem Ecology. New York: Springer-Verlag, 1988.
  14. ^ Allee, W.C. (1949). Principles of animal ecology. Philadelphia,Saunders Co., [10].
  15. ^ Kendeigh, S.C. (1961). Animal ecology. Englewood Cliffs, N.J.,Prentice-Hall, [Englewood Cliffs, N.J.,Prentice-Hall,1961.].
  16. ^ Whittaker, Robert H., Botanical Review, Classification of Natural Communities, Vol. 28, No. 1 (Jan–Mar 1962), pp. 1–239.
  17. ^ a b Whittaker, Robert H. Communities and Ecosystems. New York: MacMillan Publishing Company, Inc., 1975.
  18. ^ Whittaker, R. H. (1970). Communities and Ecosystems. Toronto, p. 51–64, [11].
  19. ^ Goodall, D. W. (editor-in-chief). Ecosystems of the World. Elsevier, Amsterdam. 36 vol., 1974-, [12].
  20. ^ Walter, H. 1976. Die ökologischen Systeme der Kontinente (Biogeosphäre). Prinzipien ihrer Gliederung mit Beispielen. Stuttgart.
  21. ^ Walter, H. & Breckle, S-W. (1991). Ökologie der Erde, Band 1, Grundlagen. Stuttgart.
  22. ^ Schültz, J. Die Ökozonen der Erde, 1st ed., Ulmer, Stuttgart, Germany, 1988, 488 pp.; 2nd ed., 1995, 535 pp.; 3rd ed., 2002. Transl.: The Ecozones of the World: The Ecological Divisions of the Geosphere. Berlin: Springer-Verlag, 1995; 2nd ed., 2005, [13].
  23. ^ Bailey System, US Forest Service
  24. ^ Bailey, R. G. 1989. Explanatory supplement to ecoregions map of the continents. Environmental Conservation 16: 307-309. [With map of land-masses of the world, "Ecoregions of the Continents — Scale 1 : 30,000,000", published as a supplement.]
  25. ^ a b Olson, D. M. & E. Dinerstein (1998). The Global 200: A representation approach to conserving the Earth’s most biologically valuable ecoregions. Conservation Biol. 12:502–515, [14].
  26. ^ a b c Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N. D., Powell, G. V. N., Underwood, E. C., D'Amico, J. A., Itoua, I., Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts, T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P., Kassem, K. R. (2001). Terrestrial ecoregions of the world: a new map of life on Earth. Bioscience 51(11):933-938, [15].
  27. ^ Abell, R., M. Thieme, C. Revenga, M. Bryer, M. Kottelat, N. Bogutskaya, B. Coad, N. Mandrak, S. Contreras-Balderas, W. Bussing, M. L. J. Stiassny, P. Skelton, G. R. Allen, P. Unmack, A. Naseka, R. Ng, N. Sindorf, J. Robertson, E. Armijo, J. Higgins, T. J. Heibel, E. Wikramanayake, D. Olson, H. L. Lopez, R. E. d. Reis, J. G. Lundberg, M. H. Sabaj Perez, and P. Petry. (2008). Freshwater ecoregions of the world: A new map of biogeographic units for freshwater biodiversity conservation. BioScience 58:403-414, [16].
  28. ^ Spalding, M. D. et al. (2007). Marine ecoregions of the world: a bioregionalization of coastal and shelf areas. BioScience 57: 573-583, [17].
  29. ^ "Freshwater Ecoregions of the World: Major Habitat Types" [18]. Accessed May 12, 2008.
  30. ^ WWF: Marine Ecoregions of the World
  31. ^ Pruvot, G. Conditions générales de la vie dans les mers et principes de distribution des organismes marins: Année Biologique, vol. 2, pp. 559—587, 1896, [19].
  32. ^ Longhurst, A. 1998. Ecological Geography of the Sea. San Diego: Academic Press, [20].
  33. ^ Zimmer, Carl (March 19, 2015). "The Next Frontier: The Great Indoors". New York Times. Retrieved March 2015.  Check date values in: |access-date= (help)

External links[edit]