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By Keneisha Boozer
PDF generated using the open source mwlib toolkit. See for more information.
PDF generated at: Wed, 02 May 2012 18:34:00 UTC
Biome types
What is happening to our planet
Global warming
Article Sources and Contributors
Image Sources, Licenses and Contributors
Article Licenses
Biome types
Biomes are climatically and geographically defined as similar
climatic conditions on the Earth, such as communities of plants,
animals, and soil organisms,[1] and are often referred to as
ecosystems. Some parts of the earth have more or less the same
kind of abiotic and biotic factors spread over a large area, creating
a typical ecosystem over that area. Such major ecosystems are
termed as biomes. Biomes are defined by factors such as plant
structures (such as trees, shrubs, and grasses), leaf types (such as
broadleaf and needleleaf), plant spacing (forest, woodland,
savanna), and climate. Unlike ecozones, biomes are not defined by
genetic, taxonomic, or historical similarities. Biomes are often
identified with particular patterns of ecological succession and
climax vegetation (quasiequilibrium state of the local ecosystem).
An ecosystem has many biotopes and a biome is a major habitat
type. A major habitat type, however, is a compromise, as it has an
intrinsic inhomogeneity.
The planet Earth
The biodiversity characteristic of each extinction, especially the diversity of fauna and subdominant plant forms, is a
function of abiotic factors and the biomass productivity of the dominant vegetation. In terrestrial biomes, species
diversity tends to correlate positively with net primary productivity, moisture availability, and temperature.[2]
Ecoregions are grouped into both biomes and ecozones.
A fundamental classification of biomes is:
1. Terrestrial (land) biomes
2. Aquatic biomes (including freshwater biomes and marine biomes)
Biomes are often known in English by local names. For example, a temperate grassland or shrubland biome is
known commonly as steppe in central Asia, prairie in North America, and pampas in South America. Tropical
grasslands are known as savanna in Australia, whereas in southern Africa it is known as certain kinds of veld (from
Sometimes an entire biome may be targeted for protection, especially under an individual nation's biodiversity action
Climate is a major factor determining the distribution of terrestrial biomes. Among the important climatic factors are:
• Latitude: Arctic, boreal, temperate, subtropical, tropical
• Humidity: humid, semihumid, semiarid, and arid
• seasonal variation: Rainfall may be distributed evenly throughout the year or be marked by seasonal variations.
• dry summer, wet winter: Most regions of the earth receive most of their rainfall during the summer months;
Mediterranean climate regions receive their rainfall during the winter months.
• Elevation: Increasing elevation causes a distribution of habitat types similar to that of increasing latitude.
The most widely used systems of classifying biomes correspond to latitude (or temperature zoning) and humidity.
Biodiversity generally increases away from the poles towards the equator and increases with humidity.
Biome classification schemes
In this scheme, climates are classified 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 type of vegetation found in an area.
Holdridge uses the four axes to define 30 so-called "humidity provinces", which are clearly visible in the Holdridge
diagram. While his scheme largely ignores soil and sun exposure, Holdridge did acknowledge that these, too, were
important factors in biome determination.
Holdridge scheme
Biomes are classification schemes which define biomes using climatic parameters. Particularly in the 1970s and
1980s, there was a significant push to understand the relationships between these climatic parameters and properties
of ecosystem energetics because such discoveries would enable the prediction of rates of energy capture and transfer
among components within ecosystems. Such a study was conducted by Sims et al. (1978) on North American
grasslands. The study found a positive logistic correlation between evapotranspiration in mm/yr and above-ground
net primary production in g/m^2/yr. More general results from the study were that precipitation and water use lead to
above-ground primary production, solar radiation and temperature lead to belowground primary production (roots),
and temperature and water lead to cool and warm season growth habit.[3] These findings help explain the categories
used in Holdridge’s bioclassification scheme, which were then later simplified in Whittaker’s. 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 all fit perfectly into the classification schemes created.
Whittaker's biome-type classification scheme
Whittaker appreciated biome-types as a representation of the great diversity of the living world, and saw the need to
establish a simple way to classify them. He based his classification scheme on two abiotic factors: precipitation and
temperature. His scheme can be seen as a simplification of Holdridge's, one more readily accessible, but perhaps
missing the greater specificity that Holdridge's provides.
Whittaker based his representation of global biomes on both previous theoretical assertions and an ever-increasing
empirical sampling of global ecosystems. He was in a unique position to make such a holistic assertion because he
had previously compiled a review of biome classification.[4]
Key definitions for understanding Whittaker's scheme
• Physiognomy: The apparent characteristics, outward features, or appearance of ecological communities or
• 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.[5]
Whittaker's parameters for classifying biome-types
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.[6]
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 the favorability of the environment (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
• 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.
Walter system
The Heinrich Walter classification scheme, developed by Heinrich Walter, a German ecologist, differs from both the
Whittaker and Holdridge schemes because it takes into account the seasonality of temperature and precipitation. The
system, also based on precipitation and temperature, finds 9 major biomes, with the important climate traits and
vegetation types summarized in the accompanying table. 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
• I: Equatorial
• Always moist and lacking temperature seasonality
• Evergreen tropical rain forest
• II: Tropical
• Summer rainy season and cooler “winter” dry season
• Seasonal forest, scrub, or savanna
• III: Subtropical
• Highly seasonal, arid climate
• Desert vegetation with considerable exposed surface
• IV: Mediterranean
• Winter rainy season and summer drought
• Sclerophyllous (drought-adapted), frost-sensitive shrublands and woodlands
• V: Warm temperate
• Occasional frost, often with summer rainfall maximum
• Temperate evergreen forest, somewhat frost-sensitive
• VI: Nemoral
• Moderate climate with winter freezing
• Frost-resistant, deciduous, temperate forest
• VII: Continental
• Arid, with warm or hot summers and cold winters
• Grasslands and temperate deserts
• VIII: Boreal
• Cold temperate with cool summers and long winters
• Evergreen, frost-hardy, needle-leaved forest (taiga)
• IX: Polar
• Very short, cool summers and long, very cold winters
• Low, evergreen vegetation, without trees, growing over permanently frozen soils
Bailey system
Robert G. Bailey almost developed a biogeographical classification system for the United States in a map published
in 1976. He subsequently expanded the system to include the rest of South 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).[7]
• 100 Polar Domain
• 120 Tundra Division
• M120 Tundra Division - Mountain Provinces
• 130 Subarctic Division
• M130 Subarctic Division - Mountain Provinces
• 200 Humid Temperate Domain
• 210 Warm Continental Division
• M210 Warm Continental Division - Mountain Provinces
• 220 Hot Continental Division
• M220 Hot Continental Division - Mountain Provinces
• 230 Subtropical Division
• M230 Subtropical Division - Mountain Provinces
• 240 Marine Division
• M240 Marine Division - Mountain Provinces
• 250 Prairie Division
• 260 Mediterranean Division
• M260 Mediterranean Division - Mountain Provinces
• 300 Dry Domain
• 310 Tropical/Subtropical Steppe Division
• M310 Tropical/Subtropical Steppe Division - Mountain Provinces
WWF system
A team of biologists convened by the World Wide Fund for Nature (WWF) developed an ecological land
classification system that identified fourteen biomes,[8] called major habitat types, and further divided the world's
land area into 867 terrestrial ecoregions. Each terrestrial ecoregion has a specific EcoID, fomat XXnnNN (XX is the
ecozone, nn is the biome number, NN is the individual number). This classification is used to define the Global 200
list of ecoregions identified by the WWF as priorities for conservation. The WWF major habitat types are:
01 Tropical and subtropical moist broadleaf forests (tropical and subtropical, humid)
02 Tropical and subtropical dry broadleaf forests (tropical and subtropical, semihumid)
03 Tropical and subtropical coniferous forests (tropical and subtropical, semihumid)
04 Temperate broadleaf and mixed forests (temperate, humid)
05 Temperate coniferous forests (temperate, humid to semihumid)
06 Boreal forests/taiga (subarctic, humid)
07 Tropical and subtropical grasslands, savannas, and shrublands (tropical and subtropical, semiarid)
08 Temperate grasslands, savannas, and shrublands (temperate, semiarid)
09 Flooded grasslands and savannas (temperate to tropical, fresh or brackish water inundated)
10 Montane grasslands and shrublands (alpine or montane climate)
11 Tundra (Arctic)
• 12 Mediterranean forests, woodlands, and scrub or sclerophyll forests (temperate warm, semihumid to semiarid
with winter rainfall)
• 13 Deserts and xeric shrublands (temperate to tropical, arid)
• 14 Mangrove (subtropical and tropical, salt water inundated)
Freshwater biomes
According to the WWF, the following are classified as freshwater biomes:[9]
Large lakes
Temperate upland rivers
Large river deltas
Tropical and subtropical coastal rivers
Polar freshwaters
Tropical and subtropical floodplain rivers and wetlands
Montane freshwaters
Tropical and subtropical upland rivers
Temperate coastal rivers
Xeric freshwaters and endorheic basins
Temperate floodplain rivers and wetlands •
Oceanic islands
• Streams and rivers
Realms or ecozones (terrestrial and freshwater, WWF)
NA Nearctic
PA Palearctic
AT Afrotropic
IM Indomalaya
AA Australasia
NT Neotropic
OC Oceania
AN Antarctic
Marine biomes
Marine biomes (H) (major habitat types), Global 200 (WWF)
Biomes of the coastal and continental shelf areas (neritic zone - List of ecoregions (WWF))
• Polar
• Temperate shelves and sea
• Temperate upwelling
• Tropical upwelling
• Tropical coral[10]
Realms or ecozones (marine, WWF)
North temperate Atlantic
Western Indo-Pacific
Eastern tropical Atlantic
South temperate Indo-Pacific
Western tropical Atlantic
Southern Ocean
South temperate Atlantic
North temperate Indo-Pacific •
Central Indo-Pacific
Eastern Indo-Pacific
Other marine habitat types
Hydrothermal vents
Cold seeps
Benthic zone
Pelagic zone (trades and westerlies)
Hadal (ocean trench)
Major habitats, nonglobal 200 (WWF)
• Littoral/Intertidal zone
• Kelp forest
• Pack ice
Summary - ecological taxonomy (WWF)
• Biosphere (List of ecoregions)
• Ecozones or realms (8)
• Terrestrial biomes (major habitat types, 14)
• Ecoregions (867) vbn,
• Ecosystems (biotopes)
• Freshwater biomes (major habitat types, 12)
• Ecoregions (426)
• Ecosystems (biotopes)
• Marine ecozones or realms (13)
• Continental Shelf biomes (major habitat types, 5)
• (Marine provinces) (62)
• Ecoregions (232)
• Ecosystems (biotopes)
• Open & Deep Sea Biomes (major habitat types)
• Endolithic biome
• Biosphere
• Ecozone: Palearctic ecozone
• Terrestrial biome: temperate broadleaf and mixed forests
• Ecoregion: Dinaric Mountains mixed forests (PA0418)
• Ecosystem: Orjen, vegetation belt between 1,100- 1,450 m, Oromediterranean zone, nemoral zone
(temperate zone)
• Biotope: Oreoherzogio-Abietetum illyricae Fuk. (Plant list)
• Plant: Silver fir (Abies alba)
Anthropogenic biomes
Humans have fundamentally altered global patterns of biodiversity and ecosystem processes. As a result, vegetation
forms predicted by conventional biome systems are rarely observed across most of Earth's land surface.
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 how
best to live in and manage our biosphere and the anthropogenic biosphere we live in. The main biomes in the world
are freshwater, marine, coniferous, deciduous, ice, mountains, boreal, grasslands, tundra, and rainforests.
Major anthropogenic biomes
Dense settlements
Other biomes
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.
Map of biomes
Freshwater biomes
The drainage basins of the principal oceans and seas of the world are marked by continental divides. The grey areas
are endorheic basins that do not drain to the ocean.
[1] The World's Biomes (http:/ / www. ucmp. berkeley. edu/ exhibits/ biomes/ index. php), Retrieved August 19, 2008, from University of
California Museum of Paleontology (http:/ / www. ucmp. berkeley. edu/ index. php)
[2] Pidwirny, Michael (2006-10-16). "Biomes" (http:/ / www. eoearth. org/ article/ Biomes). In Sidney Draggan. Encyclopedia of Earth.
Washington, D.C.: Environmental Information Coalition, National Council for Science and the Environment. . Retrieved 2006-11-16.
[3] Pomeroy, Lawrence R. and James J. Alberts, editors. Concepts of Ecosystem Ecology. New York: Springer-Verlag, 1988.
[4] Whittaker, Robert H., Botanical Review, Classification of Natural Communities, Vol. 28, No. 1 (Jan-Mar 1962), pp. 1-239.
[5] Whittaker, Robert H. Communities and Ecosystems New York: MacMillan Publishing Company, Inc., 1975.
[6] Whittaker, Robert H. Communities and Ecosystems New York: MacMillan Publishing Company, Inc., 1975.
[7] http:/ / www. fs. fed. us/ land/ ecosysmgmt/ index. html Bailey System, US Forest Service
[8] Olson, David M. et al. (2001); Terrestrial Ecoregions of the World (http:/ / www. csrc. sr. unh. edu/ ~palace/ cerrado/ olson et al. (world veg
map). pdf): A New Map of Life on Earth, BioScience, Vol. 51, No. 11., pp. 933-938.
[9] "Freshwater Ecoregions of the World: Major Habitat Types" (http:/ / www. feow. org/ mht. php). Accessed May 12, 2008.
[10] WWF: Marine Ecoregions of the World (http:/ / www. worldwildlife. org/ science/ ecoregions/ marine/ item1266. html)
External links
• Biomes of the world (Missouri Botanic Garden) (
• Global Currents and Terrestrial Biomes Map (
• ( is a site covering the 5 principal world biome types:
aquatic, desert, forest, grasslands, and tundra.
• UWSP's online textbook The Physical Environment: - Earth Biomes (
•'s Habitats ( - describes the
14 major terrestrial habitats, 7 major freshwater habitats, and 5 major marine habitats.
•'s Habitats Simplified (
habitats/) - provides simplified explanations for 10 major terrestrial and aquatic habitat types.
• UCMP Berkeley's The World's Biomes ( - provides
lists of characteristics for some biomes and measurements of climate statistics.
• Gale/Cengage has an excellent Biome Overview (
overview.htm) of terrestrial, aquatic, and man-made biomes with a particular focus on trees native to each, and
has detailed descriptions of desert, rain forest, and wetland biomes.
• NASA's Earth Observatory Mission: Biomes ( gives an
exemplar of each biome that is described in great detail and provides scientific measurements of the climate
statistics that define each biome.
A habitat (which is Latin for "it inhabits") is an ecological or
environmental area that is inhabited by a particular species of animal,
plant, or other type of organism.[1][2] It is the natural environment in
which an organism lives, or the physical environment that surrounds
(influences and is utilized by) a species population.[3]
The term "population" is preferred to "organism" because, while it is
possible to describe the habitat of a single turtle, it is also possible that
one may not find any particular or individual bear but the grouping of
bears that constitute a breeding population and occupy a certain
biogeographical area. Further, this habitat could be somewhat different
from the habitat of another group or population of black bears living
elsewhere. Thus it is neither the species nor the individual for which
the term habitat is typically used.
The remaining fragmented habitats of the African
The term microhabitat is often used to describe the small-scale
physical requirements of a particular organism or population.
Monotypic habitat
A distribution map showing the range and
breeding grounds of Great Black-backed Gulls.
The monotypic habitat occurs in botanical and zoological contexts, and is a component of conservation biology. In
restoration ecology of native plant communities or habitats, some invasive species create monotypic stands that
replace and/or prevent other species, especially indigenous ones, from growing there. A dominant colonization can
occur from retardant chemicals exuded, nutrient monopolization, or from lack of natural controls such as herbivores
or climate, that keep them in balance with their native habitats. The yellow starthistle, Centaurea solstitialis, is a
botanical monotypic-habitat example of this, currently dominating over 15000000 acres (unknown operator:
u'strong' km2) in California alone.[4][5] The non-native freshwater zebra mussel, Dreissena polymorpha, that
colonizes areas of the Great Lakes and the Mississippi River watershed, without its home-range predator control, is a
zoological monotypic-habitat example.
Even though its name may seem to imply simplicity as compared with polytypic habitats, the monotypic habitat can
be complex.[6]
[1] Dickinson, C.I. 1963. British Seaweeds. The Kew Series
[2] Abercrombie, M., Hickman, C.J. and Johnson, M.L. 1966.A Dictionary of Biology. Penguin Reference Books, London
[3] "Living Things: Habitats and Ecosystems" (http:/ / www. fi. edu/ tfi/ units/ life/ habitat/ habitat. html). The Franklin Institute. . Retrieved 29
June 2011.
[4] Mount Diablo Review, Autumn 2007 (http:/ / www. mdia. org/ PDF Files/ MD Review Autumn2007. pdf)PDF (286 KiB), Mount Diablo
Interpretive Association. Retrieved on 2008-10-15.
[5] 1970 distribution of yellow starthistle in the U.S. (http:/ / wric. ucdavis. edu/ yst/ images/ none/ nc4. JPG), a map from UCD's Yellow
Starthistle Information website
[6] Theel Heather J., Dibble Eric D., Madsen John D. (1948). "Differential influence of a monotypic and diverse native aquatic plant bed on a
macroinvertebrate assemblage; an experimental implication of exotic plant induced habitat" (http:/ / cat. inist. fr/ ?aModele=afficheN&
cpsidt=20189959). Cat.Inist and Springer, Dordrecht, PAYS-BAS. . Retrieved 2011-04-17.
External links
Human habitats
What is happening to our planet
The scientific discipline of ecology addresses the full scale of life, from tiny bacteria to processes that span the entire planet. Ecologists study many
diverse and complex relations among species, such as predation and pollination. The diversity of life is organized into different habitats, from
terrestrial (middle) to aquatic ecosystems.
Ecology (from Greek: οἶκος, "house"; -λογία, "study of") is the scientific study of the relations that living organisms
have with respect to each other and their natural environment. Variables of interest to ecologists include the
composition, distribution, amount (biomass), number, and changing states of organisms within and among
ecosystems. Ecosystems are hierarchical systems that are organized into a graded series of regularly interacting and
semi-independent parts (e.g., species) that aggregate into higher orders of complex integrated wholes (e.g.,
communities). Ecosystems are sustained by the biodiversity within them. Biodiversity is the full-scale of life and its
processes, including genes, species and ecosystems forming lineages that integrate into a complex and regenerative
spatial arrangement of types, forms, and interactions. Ecosystems create biophysical feedback mechanisms between
living (biotic) and nonliving (abiotic) components of the planet. These feedback loops regulate and sustain local
communities, continental climate systems, and global biogeochemical cycles.
Ecology is an interdisciplinary branch of biology, the study of life. The word "ecology" ("Ökologie") was coined in
1866 by the German scientist Ernst Haeckel (1834–1919). Ancient philosophers of Greece, including Hippocrates
and Aristotle, were among the earliest to record observations and notes on the natural history of plants and animals.
Modern ecology branched out of natural history and matured into a more rigorous science in the late 19th century.
Charles Darwin's evolutionary treatise including the concept of adaptation, as it was introduced in 1859, is a pivotal
cornerstone in modern ecological theory. Ecology is not synonymous with environment, environmentalism, natural
history or environmental science. It is closely related to physiology, evolutionary biology, genetics and ethology. An
understanding of how biodiversity affects ecological function is an important focus area in ecological studies.
Ecologists seek to explain:
Life processes and adaptations
Distribution and abundance of organisms
The movement of materials and energy through living communities
The successional development of ecosystems, and
The abundance and distribution of biodiversity in context of the environment.
Ecology is a human science as well. There are many practical applications of ecology in conservation biology,
wetland management, natural resource management (agriculture, forestry, fisheries), city planning (urban ecology),
community health, economics, basic and applied science and human social interaction (human ecology). Ecosystems
sustain every life-supporting function on the planet, including climate regulation, water filtration, soil formation
(pedogenesis), food, fibers, medicines, erosion control, and many other natural features of scientific, historical or
spiritual value.[1][2][3]
Integrative levels, scope, and scale of organization
The scope of ecology covers a wide
array of interacting levels of
organization spanning micro-level
(e.g., cells) to planetary scale (e.g.,
ecosphere) phenomena. Ecosystems,
for example, contain populations of
Ecosystems regenerate after a disturbance such as fire, forming mosaics of different age
groups structured across a landscape. Pictured are different seral stages in forested
individuals that aggregate into distinct
ecosystems starting from pioneers colonizing a disturbed site and maturing in
ecological communities. It can take
successional stages leading to old-growth forests.
thousands of years for ecological
processes to mature through and until
the final successional stages of a forest. The area of an ecosystem can vary greatly from tiny to vast. A single tree is
of little consequence to the classification of a forest ecosystem, but critically relevant to organisms living in and on
it.[4] Several generations of an aphid population can exist over the lifespan of a single leaf. Each of those aphids, in
turn, support diverse bacterial communities.[5] The nature of connections in ecological communities cannot be
explained by knowing the details of each species in isolation, because the emergent pattern is neither revealed nor
predicted until the ecosystem is studied as an integrated whole. Some ecological principles, however, do exhibit
collective properties where the sum of the components explain the properties of the whole, such as birth rates of a
population being equal to the sum of individual births over a designated time frame.[6]
Hierarchical ecology
System behaviours must first be arrayed into levels of organization. Behaviors corresponding to higher levels occur at slow rates.
Conversely, lower organizational levels exhibit rapid rates. For example, individual tree leaves respond rapidly to momentary
changes in light intensity, CO2 concentration, and the like. The growth of the tree responds more slowly and integrates these
short-term changes.
The scale of ecological dynamics can operate like a closed island with respect to local site variables, such as aphids
migrating on a tree, while at the same time remain open with regard to broader scale influences, such as atmosphere
or climate. Hence, ecologists have devised means of hierarchically classifying ecosystems by analyzing data
collected from finer scale units, such as vegetation associations, climate, and soil types, and integrate this
information to identify larger emergent patterns of uniform organization and processes that operate on local to
regional, landscape, and chronological scales.
To structure the study of ecology into a manageable framework of understanding, the biological world is
conceptually organized as a nested hierarchy of organization, ranging in scale from genes, to cells, to tissues, to
organs, to organisms, to species and up to the level of the biosphere.[8] Together these hierarchical scales of life form
a panarchy[9][10] and they exhibit non-linear behaviours; "nonlinearity refers to the fact that effect and cause are
disproportionate, so that small changes in critical variables, such as the numbers of nitrogen fixers, can lead to
disproportionate, perhaps irreversible, changes in the system properties."[11]:14
Biodiversity is the variety of life and its processes. It includes the variety of living organisms, the genetic differences among them,
the communities and ecosystems in which they occur, and the ecological and evolutionary processes that keep them functioning, yet
ever changing and adapting.
Biodiversity (an abbreviation of biological diversity) describes the diversity of life from genes to ecosystems and
spans every level of biological organization. Biodiversity means different things to different people and there are
many ways to index, measure, characterize, and represent its complex organization.[13][14] Biodiversity includes
species diversity, ecosystem diversity, genetic diversity and the complex processes operating at and among these
respective levels.[14][15][16] Biodiversity plays an important role in ecological health as much as it does for human
health.[17][18] Preventing or prioritizing species extinctions is one way to preserve biodiversity, but populations, the
genetic diversity within them and ecological processes, such as migration, are being threatened on global scales and
disappearing rapidly as well. Conservation priorities and management techniques require different approaches and
considerations to address the full ecological scope of biodiversity. Populations and species migration, for example,
are more sensitive indicators of ecosystem services that sustain and contribute natural capital toward the well-being
of humanity.[19][20][21][22] An understanding of biodiversity has practical application for ecosystem-based
conservation planners as they make ecologically responsible decisions in management recommendations to
consultant firms, governments and industry.[23]
The habitat of a species describes the environment over which a species is known to occur and the type of
community that is formed as a result.[24] More specifically, "habitats can be defined as regions in environmental
space that are composed of multiple dimensions, each representing a biotic or abiotic environmental variable; that is,
any component or characteristic of the environment related directly (e.g. forage biomass and quality) or indirectly
(e.g. elevation) to the use of a location by the animal."[25]:745 For example, the habitat might refer to an aquatic or
terrestrial environment that can be further categorized as montane or alpine ecosystems. Habitat shifts provide
important evidence of competition in nature where one population changes relative to the habitats that most other
individuals of the species occupy. One population of a species of tropical lizards (Tropidurus hispidus), for example,
has a flattened body relative to the main populations that live in open savanna. The population that lives in an
isolated rock outcrop hides in crevasses where its flattened body may improve its performance. Habitat shifts also
occur in the developmental life history of amphibians and many insects that transition from aquatic to terrestrial
habitats. Biotope and habitat are sometimes used interchangeably, but the former applies to a communities
environment, whereas the latter applies to a species' environment.[24][26][27]
Biodiversity of a coral reef. Corals adapt and
modify their environment by forming calcium
carbonate skeletons that provide growing
conditions for future generations and form habitat
for many other species.
There are many definitions of the niche dating back to 1917,[31] but G.
Evelyn Hutchinson made conceptual advances in 1957[32][33] and
introduced a widely accepted definition: "the set of biotic and abiotic
conditions which a species is able to persist and maintain stable
population sizes."[31]:519 The ecological niche is a central concept in
the ecology of organisms and is sub-divided into the fundamental and
the realized niche. The fundamental niche is the set of environmental
conditions under which a species is able to persist. The realized niche
is the set of environmental plus ecological conditions under which a
species persists.[31][33][34] The Hutchisonian niche is defined more
technically as an "Euclidean hyperspace whose dimensions are defined
as environmental variables and whose size is a function of the number
of values that the environmental values may assume for which an
organism has positive fitness."[35]:71
Biogeographical patterns and range distributions are explained or
predicted through knowledge and understanding of a species traits and
niche requirements.[36] Species have functional traits that are uniquely
Termite mounds with varied heights of chimneys
regulate gas exchange, temperature and other
adapted to the ecological niche. A trait is a measurable property,
environmental parameters that are needed to
phenotype, or characteristic of an organism that influences its
sustain the internal physiology of the entire
performance. Genes play an important role in the development and
expression of traits.
Resident species evolve traits that are fitted to
their local environment. This tends to afford them a competitive
advantage and discourages similarly adapted species from having an overlapping geographic range. The competitive
exclusion principle suggests that two species cannot coexist indefinitely by living off the same limiting resource.
When similarly adapted species are found to overlap geographically, closer inspection reveals subtle ecological
differences in their habitat or dietary requirements.[38] Some models and empirical studies, however, suggest that
disturbances can stabilize the coevolution and shared niche occupancy of similar species inhabiting species rich
communities.[39] The habitat plus the niche is called the ecotope, which is defined as the full range of environmental
and biological variables affecting an entire species.[24]
Niche construction
Organisms are subject to environmental pressures, but they are also modifiers of their habitats. The regulatory
feedback between organisms and their environment can modify conditions from local (e.g., a beaver pond) to global
scales (e.g., Gaia), over time and even after death, such as decaying logs or silica skeleton deposits from marine
organisms.[40] The process and concept of ecosystem engineering has also been called niche construction. Ecosystem
engineers are defined as: "...organisms that directly or indirectly modulate the availability of resources to other
species, by causing physical state changes in biotic or abiotic materials. In so doing they modify, maintain and create
The ecosystem engineering concept has stimulated a new appreciation for the degree of influence that organisms
have on the ecosystem and evolutionary process. The terms niche construction are more often used in reference to
the under appreciated feedback mechanism of natural selection imparting forces on the abiotic niche.[29][42] An
example of natural selection through ecosystem engineering occurs in the nests of social insects, including ants, bees,
wasps, and termites. There is an emergent homeostasis or homeorhesis in the structure of the nest that regulates,
maintains and defends the physiology of the entire colony. Termite mounds, for example, maintain a constant
internal temperature through the design of air-conditioning chimneys. The structure of the nests themselves are
subject to the forces of natural selection. Moreover, the nest can survive over successive generations, which means
that ancestors inherit both genetic material and a legacy niche that was constructed before their time.[6][29][30][43]
Biomes are larger units of organization that categorize regions of the Earth's ecosystems mainly according to the
structure and composition of vegetation.[44] Different researchers have applied different methods to define
continental boundaries of biomes dominated by different functional types of vegetative communities that are limited
in distribution by climate, precipitation, weather and other environmental variables. Examples of biome names
include: tropical rainforest, temperate broadleaf and mixed forests, temperate deciduous forest, taiga, tundra, hot
desert, and polar desert.[45] Other researchers have recently started to categorize other types of biomes, such as the
human and oceanic microbiomes. To a microbe, the human body is a habitat and a landscape.[46] The microbiome
has been largely discovered through advances in molecular genetics that have revealed a hidden richness of
microbial diversity on the planet. The oceanic microbiome plays a significant role in the ecological biogeochemistry
of the planet's oceans.[47]
Ecological theory has been used to explain self-emergent regulatory phenomena at the planetary scale. The largest
scale of ecological organization is the biosphere: the total sum of ecosystems on the planet. Ecological relationships
regulate the flux of energy, nutrients, and climate all the way up to the planetary scale. For example, the dynamic
history of the planetary CO2 and O2 composition of the atmosphere has been largely determined by the biogenic flux
of gases coming from respiration and photosynthesis, with levels fluctuating over time and in relation to the ecology
and evolution of plants and animals.[48] When sub-component parts are organized into a whole there are oftentimes
emergent properties that describe the nature of the system. The Gaia hypothesis is an example of holism applied in
ecological theory.[49] The ecology of the planet acts as a single regulatory or holistic unit called Gaia. The Gaia
hypothesis states that there is an emergent feedback loop generated by the metabolism of living organisms that
maintains the temperature of the Earth and atmospheric conditions within a narrow self-regulating range of
Population ecology
The population is the unit of analysis in population ecology. A population consists of individuals of the same species
that live, interact and migrate through the same niche and habitat.[51] A primary law of population ecology is the
Malthusian growth model.[52] This law states that:
"...a population will grow (or decline) exponentially as long as the environment experienced by all individuals in the
population remains constant."[52]:18
This Malthusian premise provides the basis for formulating predictive theories and tests that follow. Simplified
population models usually start with four variables including death, birth, immigration, and emigration.
Mathematical models are used to calculate changes in population demographics using a null model. A null model is
used as a null hypothesis for statistical testing. The null hypothesis states that random processes create observed
patterns. Alternatively the patterns differ significantly from the random model and require further explanation.
Models can be mathematically complex where "...several competing hypotheses are simultaneously confronted with
the data."[53] An example of an introductory population model describes a closed population, such as on an island,
where immigration and emigration does not take place. In these island models the rate of population change is
described by:
where N is the total number of individuals in the population, B is the number of births, D is the number of deaths, b
and d are the per capita rates of birth and death respectively, and r is the per capita rate of population change. This
formula can be read out as the rate of change in the population (dN/dT) is equal to births minus deaths (B – D).[52][54]
Using these modelling techniques, Malthus' population principle of growth was later transformed into a model
known as the logistic equation:
where N is the number of individuals measured as biomass density, a is the maximum per-capita rate of change, and
K is the carrying capacity of the population. The formula can be read as follows: the rate of change in the population
(dN/dT) is equal to growth (aN) that is limited by carrying capacity (1 – N/K). The discipline of population ecology
builds upon these introductory models to further understand demographic processes in real study populations and
conduct statistical tests. The field of population ecology often uses data on life history and matrix algebra to develop
projection matrices on fecundity and survivorship. This information is used for managing wildlife stocks and setting
harvest quotas.[54][55]
Metapopulations and migration
Populations are also studied and modeled according to the metapopulation concept. The metapopulation concept was
introduced in 1969:[56] "as a population of populations which go extinct locally and recolonize."[57]:105
Metapopulation ecology is another statistical approach that is often used in conservation research.[58]
Metapopulation research simplifies the landscape into patches of varying levels of quality.[59] Metapopulations are
linked by the migratory behaviours of organisms. Animal migration is set apart from other kinds of movement
because it involves the seasonal departure and return of individuals from one habitat to another.[60] Migration is also
a population level phenomenon, such as the migration routes followed by plants as they occupied northern
post-glacial environments. Plant ecologists rely on pollen records that accumulate and stratify in wetlands to
reconstruct the timing of plant migration and dispersal relative to historic and contemporary climates. These
migration routes involved an expansion of the range as plant populations expanded from one area to another. There
is a larger taxonomy of movement, such as commuting, foraging, territorial behaviour, stasis, and ranging. Dispersal
is usually distinguished from migration because it involves the one way permanent movement of individuals from
their birth population into another population.[61][]
In metapopulation terminology there are emigrants (individuals that leave a patch), immigrants (individuals that
move into a patch) and sites are classed either as sources or sinks. A site is a generic term that refers to places where
ecologists sample populations, such as ponds or defined sampling areas in a forest. Source patches are productive
sites that generate a seasonal supply of juveniles that migrate to other patch locations. Sink patches are unproductive
sites that only receive migrants and will go extinct unless rescued by an adjacent source patch or environmental
conditions become more favorable. Metapopulation models examine patch dynamics over time to answer questions
about spatial and demographic ecology. The ecology of metapopulations is a dynamic process of extinction and
colonization. Small patches of lower quality (i.e., sinks) are maintained or rescued by a seasonal influx of new
immigrants. A dynamic metapopulation structure evolves from year to year, where some patches are sinks in dry
years and become sources when conditions are more favorable. Ecologists use a mixture of computer models and
field studies to explain metapopulation structure.[62][63]
Community ecology
Community ecology examines how interactions among species and their
environment affect the abundance, distribution and diversity of species within
Johnson & Stinchcomb
Community ecology is the study of the interactions among a collection
of interdependent species that cohabitate the same geographic area. An
example of a study in community ecology might measure primary
production in a wetland in relation to decomposition and consumption
rates. This requires an understanding of the community connections
Interspecific interactions such as predation are a
key aspect of community ecology.
between plants (i.e., primary producers) and the decomposers (e.g.,
fungi and bacteria).
or the analysis of predator-prey dynamics
affecting amphibian biomass.[66] Food webs and trophic levels are two widely employed conceptual models used to
explain the linkages among species.[67][68]
Ecosystem ecology
These ecosystems, as we may call them, are of the most various kinds and sizes. They form one category of the multitudinous
physical systems of the universe, which range from the universe as a whole down to the atom.
The concept of the ecosystem was fully
synthesized in 1935 to describe habitats within
biomes that form an integrated whole and a
dynamically responsive system having both
physical and biological complexes. However, the
underlying concept can be traced back to 1864 in
the published work of George Perkins Marsh
("Man and Nature").[70][71] Within an ecosystem
there are inseparable ties that link organisms to
the physical and biological components of their
environment to which they are adapted.[69]
Ecosystems are complex adaptive systems where
Figure 1. A riparian forest in the White Mountains, New Hampshire (USA),
the interaction of life processes form
an example of ecosystem ecology
self-organizing patterns across different scales of
time and space.[72] terrestrial, freshwater,
atmospheric, and marine ecosystems very broadly cover the major types. Differences stem from the nature of the
unique physical environments that shapes the biodiversity within each. A more recent addition to ecosystem ecology
are the novel technoecosystems of the anthropocene.[6]
Food webs
A food web is the archetypal ecological network. Plants capture and convert solar energy into the biomolecular
bonds of simple sugars during photosynthesis. This food energy is transferred through a series of organisms starting
with those that feed on plants and are themselves consumed. The simplified linear feeding pathways that move from
a basal trophic species to a top consumer is called the food chain. The larger interlocking pattern of food chains in an
ecological community creates a complex food web. Food webs are a type of concept map or a heuristic device that is
used illustrate and study pathways of energy and material flows.[7][73][74]
Generalized food web of waterbirds from
Chesapeake Bay
Food webs are often limited relative to the real world. Complete
empirical measurements are generally restricted to a specific habitat,
such as a cave or a pond. Principles gleaned from food web microcosm
studies are used to extrapolate smaller dynamic concepts to larger
systems.[75] Feeding relations require extensive investigations into the
gut contents of organisms, which can be very difficult to decipher, or
(more recently) stable isotopes can be used to trace the flow of nutrient
diets and energy through a food web.[76] While food webs often give
an incomplete measure of ecosystems, they are nonetheless a valuable
tool in understanding community ecosystems.[77]
Food-webs exhibit principals of ecological emergence through the
nature of trophic entanglement, where some species have many weak feeding links (e.g., omnivores) while some are
more specialized with fewer stronger feeding links (e.g., primary predators). Theoretical and empirical studies
identify non-random emergent patterns of few strong and many weak linkages that serve to explain how ecological
communities remain stable over time.[78] Food-webs have compartments, where the many strong interactions create
subgroups among some members in a community and the few weak interactions occur between these subgroups.
These compartments increase the stability of food-webs.[79] As plants grow, they accumulate carbohydrates and are
eaten by grazing herbivores. Step by step lines or relations are drawn until a web of life is illustrated.[74][80][81][82]
Trophic levels
The Greek root of the word troph,
τροφή, trophē, means food or feeding.
Links in food-webs primarily connect
feeding relations or trophism among
ecosystems can be organized into
vertical and horizontal dimensions.
The vertical dimension represents
feeding relations that become further
A trophic pyramid (a) and a food-web (b) illustrating ecological relationships among
removed from the base of the food
creatures that are typical of a northern Boreal terrestrial ecosystem. The trophic pyramid
chain up toward top predators. A
roughly represents the biomass (usually measured as total dry-weight) at each level.
Plants generally have the greatest biomass. Names of trophic categories are shown to the
trophic level is defined as "a group of
right of the pyramid. Some ecosystems, such as many wetlands, do not organize as a strict
organisms acquiring a considerable
pyramid, because aquatic plants are not as productive as long-lived terrestrial plants such
majority of its energy from the
as trees. Ecological trophic pyramids are typically one of three kinds: 1) pyramid of
adjacent level nearer the abiotic
numbers, 2) pyramid of biomass, or 3) pyramid of energy.
dimension represents the abundance or biomass at each level.
When the relative abundance or biomass of each
functional feeding group is stacked into their respective trophic levels they naturally sort into a 'pyramid of
Functional groups are broadly categorized as autotrophs (or primary producers), heterotrophs (or consumers), and
detrivores (or decomposers). Autotrophs are organisms that can produce their own food (production is greater than
respiration) and are usually plants or cyanobacteria that are capable of photosynthesis but can also be other
organisms such as bacteria near ocean vents that are capable of chemosynthesis. Heterotrophs are organisms that
must feed on others for nourishment and energy (respiration exceeds production).[6] Heterotrophs can be further
sub-divided into different functional groups, including: primary consumers (strict herbivores), secondary consumers
(carnivorous predators that feed exclusively on herbivores) and tertiary consumers (predators that feed on a mix of
herbivores and predators).[86] Omnivores do not fit neatly into a functional category because they eat both plant and
animal tissues. It has been suggested that omnivores have a greater functional influence as predators because relative
to herbivores they are comparatively inefficient at grazing.[87]
Trophic levels are part of the holistic or complex systems view of ecosystems.[88][89] Each trophic level contains
unrelated species that grouped together because they share common ecological functions. Grouping functionally
similar species into a trophic system gives a macroscopic image of the larger functional design.[90] While the notion
of trophic levels provides insight into energy flow and top-down control within food webs, it is troubled by the
prevalence of omnivory in real ecosystems. This has lead some ecologists to "reiterate that the notion that species
clearly aggregate into discrete, homogeneous trophic levels is fiction."[91]:815 Nonetheless, recent studies have
shown that real trophic levels do exist, but "above the herbivore trophic level, food webs are better characterized as a
tangled web of omnivores."[92]:612
Keystone species
A keystone species is a species that is disproportionately connected to more species in the food-web. Keystone
species have lower levels of biomass in the trophic pyramid relative to the importance of their role. The many
connections that a keystone species holds means that it maintains the organization and structure of entire
communities. The loss of a keystone species results in a range of dramatic cascading effects that alters trophic
dynamics, other food-web connections and can cause the extinction of other species in the community.[93][94]
Sea otters (Enhydra lutris) are commonly cited as an example of a keystone species because they limit the density of
sea urchins that feed on kelp. If sea otters are removed from the system, the urchins graze until the kelp beds
disappear and this has a dramatic effect on community structure.[95] Hunting of sea otters, for example, is thought to
have indirectly led to the extinction of the Steller's Sea Cow (Hydrodamalis gigas).[96] While the keystone species
concept has been used extensively as a conservation tool, it has been criticized for being poorly defined from an
operational stance. It is very difficult to experimentally determine in each different ecosystem what species may hold
a keystone role. Furthermore, food-web theory suggests that keystone species may not be all that common. It is
therefore unclear how generally the keystone species model can be applied.[95][97]
Soil is the living top layer of mineral and organic dirt that covers the surface of the planet, it is the chief organizing
centre of most ecosystem functions, and it is of critical importance in agricultural science and ecology. The
decomposition of dead organic matter, such as leaves falling on the forest floor, turns into soils containing minerals
and nutrients that feed into plant production. The total sum of the planet's soil ecosystems is called the pedosphere
where a very large proportion of the Earth's biodiversity sorts into other trophic levels. Invertebrates that feed and
shred larger leaves, for example, create smaller bits for smaller organisms in the feeding chain. Collectively, these
are the detrivores that regulate soil formation. [98][99][100][101] Tree roots, fungi, bacteria, worms, ants, beetles,
centipedes, spiders, mammals, birds, reptiles, amphibians and other less familiar creatures all work to create the
trophic web of life in soil ecosystems. As organisms feed and migrate through soils they physically displace
materials, which is an important ecological process called bioturbation. Bioturbation helps to aerate the soils, thus
stimulating hetertrophic growth and production. Biomass of soil microorganisms are influenced by and feed back
into the trophic dynamics of the exposed solar surface ecology. Paleoecological studies of soils places the origin for
bioturbation to a time before the Cambrian period. Other events, such as the evolution of trees and amphibians
moving into land in the Devonian period played a significant role in the development of the ecological trophism in
Ecological complexity
Complexity is easily understood as a large computational effort needed to piece together numerous interacting parts
exceeding the iterative memory capacity of the human mind. Global patterns of biological diversity are complex.
This biocomplexity stems from the interplay among ecological processes that operate and influence patterns at
different scales that grade into each other, such as transitional areas or ecotones spanning landscapes.[103]
Complexity stems from the interplay among levels of biological organization as energy and matter is integrated into
larger units that superimpose onto the smaller parts. "What were wholes on one level become parts on a higher
one."[104]:209 Small scale patterns do not necessarily explain large scale phenomena, otherwise captured in the
expression (coined by Aristotle) 'the sum is greater than the parts'.[105][106]
"Complexity in ecology is of at least six distinct types: spatial, temporal, structural, process, behavioral, and
geometric."[107]:3 Out of these principles, ecologists have identified emergent and self-organizing phenomena that
operate at different environmental scales of influence, ranging from molecular to planetary, and these require
different sets of scientific explanation at each integrative level.[50][108] Ecological complexity relates to the dynamic
resilience of ecosystems that transition to multiple shifting steady-states directed by random fluctuations of
history.[9][109] Long-term ecological studies provide important track records to better understand the complexity and
resilience of ecosystems over longer temporal and broader spatial scales. The International Long Term Ecological
Network[110] manages and exchanges scientific information among research sites. The longest experiment in
existence is the Park Grass Experiment that was initiated in 1856.[111] Another example includes the Hubbard Brook
study in operation since 1960.[112]
The biological organization of life self-organizes into layers of emergent whole systems that function according to
nonreducible properties called holism. This means that higher order patterns of a whole functional system, such as an
ecosystem, cannot be predicted or understood by a simple summation of the parts. "New properties emerge because
the components interact, not because the basic nature of the components is changed."[6]:8
Ecological studies are necessarily holistic as opposed to reductionistic.[108][113] Holism has three scientific meanings
or uses that identify with: 1) the mechanistic complexity of ecosystems, 2) the practical description of patterns in
quantitative reductionist terms where correlations may be identified but nothing is understood about the causal
relations without reference to the whole system, which leads to 3) a metaphysical hierarchy whereby the causal
relations of larger systems are understood without reference to the smaller parts. An example of the metaphysical
aspect to holism is identified in the trend of increased exterior thickness in shells of different species. The reason for
a thickness increase can be understood through reference to principals of natural selection via predation without need
to reference or understand the biomolecular properties of the exterior shells.[114]
Relation to evolution
Ecology and evolution are considered sister disciplines of the life sciences. Natural selection, life history,
development, adaptation, populations, and inheritance are examples of concepts that thread equally into ecological
and evolutionary theory. Morphological, behavioral and/or genetic traits, for example, can be mapped onto
evolutionary trees to study the historical development of a species in relation to their functions and roles in different
ecological circumstances. In this framework, the analytical tools of ecologists and evolutionists overlap as they
organize, classify and investigate life through common systematic principals, such as phylogenetics or the Linnaean
system of taxonomy.[115] The two disciplines often appear together, such as in the title of the journal Trends in
Ecology and Evolution.[116] There is no sharp boundary separating ecology from evolution and they differ more in
their areas of applied focus. Both disciplines discover and explain emergent and unique properties and processes
operating across different spatial or temporal scales of organization.[50][117][118] While the boundary between
ecology and evolution is not always clear, it is understood that ecologists study the abiotic and biotic factors that
influence the evolutionary process.[119][120]
Behavioral ecology
Social display and color variation in differently adapted species of chameleons
(Bradypodion spp.). Chameleons change their skin color to match their background as a
behavioral defense mechanism and also use color to communicate with other members
of their species, such as dominant (left) versus submissive (right) patterns shown in the
three species (A-C) above.
All organisms are motile to some
extent. Even plants express complex
behavior, including memory and
ecology is the study of an organism's
behavior in its environment and its
implications. Ethology is the study of
observable movement or behavior in
investigations of motile sperm of plants,
mobile phytoplankton, zooplankton
swimming toward the female egg, the
cultivation of fungi by weevils, the
mating dance of a salamander, or social
Adaptation is the central unifying concept in behavioral ecology.[128] Behaviors can be recorded as traits and
inherited in much the same way that eye and hair color can. Behaviors evolve and become adapted to the ecosystem
because they are subject to the forces of natural selection.[15] Hence, behaviors can be adaptive, meaning that they
evolve functional utilities that increases reproductive success for the individuals that inherit such traits.[129] This is
also the technical definition for fitness in biology, which is a measure of reproductive success over successive
Predator-prey interactions are an introductory concept into food-web studies as well as behavioral ecology.[130] Prey
species can exhibit different kinds of behavioral adaptations to predators, such as avoid, flee or defend. Many prey
species are faced with multiple predators that differ in the degree of danger posed. To be adapted to their
environment and face predatory threats, organisms must balance their energy budgets as they invest in different
aspects of their life history, such as growth, feeding, mating, socializing, or modifying their habitat. Hypotheses
posited in behavioral ecology are generally based on adaptive principals of conservation, optimization or
efficiency.[34][119][131] For example,
"The threat-sensitive predator avoidance hypothesis predicts that prey should assess the degree of threat posed by
different predators and match their behavior according to current levels of risk."[132]
"The optimal flight initiation distance occurs where expected postencounter fitness is maximized, which depends on
the prey's initial fitness, benefits obtainable by not fleeing, energetic escape costs, and expected fitness loss due to
predation risk."[133]
Elaborate sexual displays and posturing are encountered in the
behavioral ecology of animals. The birds of paradise, for example,
display elaborate ornaments and song during courtship. These displays
serve a dual purpose of signaling healthy or well-adapted individuals
and desirable genes. The elaborate displays are driven by sexual
selection as an advertisement of quality of traits among male
Social ecology
Social ecological behaviors are notable in the social insects, slime
moulds, social spiders, human society, and naked mole rats where
eusocialism has evolved. Social behaviors include reciprocally
beneficial behaviors among kin and nest mates.[15][125][136] Social
behaviors evolve from kin and group selection. Kin selection explains
altruism through genetic relationships, whereby an altruistic behavior
leading to death is rewarded by the survival of genetic copies
Symbiosis: Leafhoppers (Eurymela fenestrata)
distributed among surviving relatives. The social insects, including
protected by ants (Iridomyrmex purpureus) in
ants, bees and wasps are most famously studied for this type of
a symbiotic relationship. The ants protect the
relationship because the male drones are clones that share the same
leafhoppers from predators and in return the
genetic make-up as every other male in the colony.[15] In contrast,
leafhoppers feeding on plants exude honeydew
from their anus that provides energy and nutrients
group selectionists find examples of altruism among non-genetic
to tending ants.
relatives and explain this through selection acting on the group,
whereby it becomes selectively advantageous for groups if their
members express altruistic behaviors to one another. Groups that are predominantly altruists beat groups that are
predominantly selfish.[15][137]
Ecological interactions can be divided into host and associate relationships. A host is any entity that harbors another
that is called the associate.[138] Host and associate relationships among species that are mutually or reciprocally
beneficial are called mutualisms. If the host and associate are physically connected, the relationship is called
symbiosis. Approximately 60% of all plants, for example, have a symbiotic relationship with arbuscular mycorrhizal
fungi. Symbiotic plants and fungi exchange carbohydrates for mineral nutrients.[139] Symbiosis differs from indirect
mutualisms where the organisms live apart. For example, tropical rainforests regulate the Earth's atmosphere. Trees
living in the equatorial regions of the planet supply oxygen into the atmosphere that sustains species living in distant
polar regions of the planet. This relationship is called commensalism because many other host species receive the
benefits of clean air at no cost or harm to the associate tree species supplying the oxygen.[140] The host and associate
relationship is called parasitism if one species benefits while the other suffers. Competition among species or among
members of the same species is defined as reciprocal antagonism, such as grasses competing for growth space.[141]
Popular ecological study systems for
mutualism include, fungus-growing ants
employing agricultural symbiosis, bacteria
living in the guts of insects and other
organisms, the fig wasp and yucca moth
pollination complex, lichens with fungi and
photosynthetic algae, and corals with
photosynthetic algae.[142][143] Nevertheless,
many organisms exploit host rewards
without reciprocating and thus have been
branded with a myriad of not-very-flattering
names such as 'cheaters', 'exploiters',
'robbers', and 'thieves'. Although cheaters
impose several host cots (e.g., via damage to
their reproductive organs or propagules,
denying the services of a beneficial partner),
their net effect on host fitness is not
necessarily negative and, thus, becomes
difficult to forecast.[144][145]
Parasites: A harvestman arachnid is parasitized by mites. This is parasitism
because the harvestman is being consumed as its juices are slowly sucked out
while the mites gain all the benefits traveling on and feeding off of their host.
The word biogeography is an amalgamation
of biology and geography. Biogeography is
the comparative study of the geographic distribution of organisms and the corresponding evolution of their traits in
space and time.[146] The Journal of Biogeography was established in 1974.[147] Biogeography and ecology share
many of their disciplinary roots. For example, the theory of island biogeography, published by the mathematician
Robert MacArthur and ecologist Edward O. Wilson in 1967[148] is considered one of the fundamentals of ecological
Biogeography has a long history in the natural sciences where questions arise concerning the spatial distribution of
plants and animals. Ecology and evolution provide the explanatory context for biogeographical studies.[146]
Biogeographical patterns result from ecological processes that influence range distributions, such as migration and
dispersal.[149] and from historical processes that split populations or species into different areas.[150] The
biogeographic processes that result in the natural splitting of species explains much of the modern distribution of the
Earth's biota. The splitting of lineages in a species is called vicariance biogeography and it is a sub-discipline of
biogeography.[150][151][152] There are also practical applications in the field of biogeography concerning ecological
systems and processes. For example, the range and distribution of biodiversity and invasive species responding to
climate change is a serious concern and active area of research in context of global warming.[20][153]
r/K-Selection theory
A population ecology concept (introduced in MacArthur and Wilson's (1967) book, The Theory of Island
Biogeography) is r/K selection theory, one of the first predictive models in ecology used to explain life-history
evolution. The premise behind the r/K selection model is that natural selection pressures change according to
population density. For example, when an island is first colonized, density of individuals is low. The initial increase
in population size is not limited by competition, leaving an abundance of available resources for rapid population
growth. These early phases of population growth experience density-independent forces of natural selection, which is
called r-selection. As the population becomes more crowded, it approaches the island's carrying capacity, thus
forcing individuals to compete more heavily for fewer available resources. Under crowded conditions the population
experiences density-dependent forces of natural selection, called K-selection.[154]
In the r/K-selection model, the first variable r is the intrinsic rate of natural increase in population size and the
second variable K is the carrying capacity of a population.[34] Different species evolve different life-history strategies
spanning a continuum between these two selective forces. An r-selected species is one that has high birth rates, low
levels of parental investment, and high rates of mortality before individuals reach maturity. Evolution favors high
rates of fecundity in r-selected species. Many kinds of insects and invasive species exhibit r-selected characteristics.
In contrast, a K-selected species has low rates of fecundity, high levels of parental investment in the young, and low
rates of mortality as individuals mature. Humans and elephants are examples of species exhibiting K-selected
characteristics, including longevity and efficiency in the conversion of more resources into fewer offspring.[148][155]
Molecular ecology
The important relationship between ecology and genetic inheritance predates modern techniques for molecular
analysis. Molecular ecological research became more feasible with the development of rapid and accessible genetic
technologies, such as the polymerase chain reaction (PCR). The rise of molecular technologies and influx of research
questions into this new ecological field resulted in the publication Molecular Ecology in 1992.[156] Molecular
ecology uses various analytical techniques to study genes in an evolutionary and ecological context. In 1994, John
Avise also played a leading role in this area of science with the publication of his book, Molecular Markers, Natural
History and Evolution.[157] Newer technologies opened a wave of genetic analysis into organisms once difficult to
study from an ecological or evolutionary standpoint, such as bacteria, fungi and nematodes. Molecular ecology
engendered a new research paradigm for investigating ecological questions considered otherwise intractable.
Molecular investigations revealed previously obscured details in the tiny intricacies of nature and improved
resolution into probing questions about behavioral and biogeographical ecology.[157] For example, molecular
ecology revealed promiscuous sexual behavior and multiple male partners in tree swallows previously thought to be
socially monogamous.[158] In a biogeographical context, the marriage between genetics, ecology and evolution
resulted in a new sub-discipline called phylogeography.[159]
Human ecology
Human ecology is the interdisciplinary investigation into the ecology of our species. "Human ecology may be
defined: (1) from a bio-ecological standpoint as the study of man as the ecological dominant in plant and animal
communities and systems; (2) from a bio-ecological standpoint as simply another animal affecting and being affected
by his physical environment; and (3) as a human being, somehow different from animal life in general, interacting
with physical and modified environments in a distinctive and creative way. A truly interdisciplinary human ecology
will most likely address itself to all three."[160] The term human ecology was formally introduced in 1921, but many
sociologists, geographers, psychologists, and other disciplines were interested in human relations to natural systems
centuries prior, especially in the late 19th century.[160][161] Some authors have identified a new unifying science in
coupled human and natural systems that builds upon, but moves beyond the field human ecology.[162] Ecology is as
much a biological science as it is a human science.[6] "Perhaps the most important implication involves our view of
human society. Homo sapiens is not an external disturbance, it is a keystone species within the system. In the long
term, it may not be the magnitude of extracted goods and services that will determine sustainability. It may well be
our disruption of ecological recovery and stability mechanisms that determines system collapse."[71]:3282
Relation to the environment
The environment is dynamically interlinked, imposed upon and constrains organisms at any time throughout their
life cycle.[163] Like the term ecology, environment has different conceptual meanings and to many these terms also
overlap with the concept of nature. Environment "...includes the physical world, the social world of human relations
and the built world of human creation."[164]:62 The environment in ecosystems includes both physical parameters and
biotic attributes. The physical environment is external to the level of biological organization under investigation,
including abiotic factors such as temperature, radiation, light, chemistry, climate and geology. The biotic
environment includes genes, cells, organisms, members of the same species (conspecifics) and other species that
share a habitat.[165] The laws of thermodynamics applies to ecology by means of its physical state. Armed with an
understanding of metabolic and thermodynamic principles a complete accounting of energy and material flow can be
traced through an ecosystem.[166]
Environmental and ecological relations are studied through reference to conceptually manageable and isolated parts.
Once the effective environmental components are understood they conceptually link back together as a
holocoenotic[167] system. In other words, the organism and the environment form a dynamic whole (or
umwelt).[168]:252 Change in one ecological or environmental factor can concurrently affect the dynamic state of an
entire ecosystem.[169][170]
Disturbance and resilience
Ecosystems are regularly confronted with natural environmental variations and disturbances over time and
geographic space. A disturbance is any process that removes living biomass from a community, such as a fire, flood,
drought, or predation.[171] Fluctuations causing disturbance occur over vastly different ranges in terms of magnitudes
as well as distances and time periods.[172] Disturbances, such as fire, are both cause and product of natural
fluctuations in death rates, species assemblages, and biomass densities within an ecological community. These
disturbances create places of renewal where new directions emerge out of the patchwork of natural experimentation
and opportunity.[171][173] [174] Ecological resilience is a cornerstone theory in ecosystem management. Biodiversity
fuels the resilience of ecosystems acting as a kind of regenerative insurance.[174]
Metabolism and the early atmosphere
Metabolism – the rate at which energy and material resources are taken up from the environment, transformed within an organism,
and allocated to maintenance, growth and reproduction – is a fundamental physiological trait.
Ernest et al.
The Earth formed approximately 4.5 billion years ago[176] and environmental conditions were too extreme for life to
form for the first 500 million years. During this early Hadean period, the Earth started to cool, allowing a crust and
oceans to form. Environmental conditions were unsuitable for the origins of life for the first billion years after the
Earth formed. The Earth's atmosphere transformed from being dominated by hydrogen, to one composed mostly of
methane and ammonia. Over the next billion years the metabolic activity of life transformed the atmosphere to
higher concentrations of carbon dioxide, nitrogen, and water vapor. These gases changed the way that light from the
sun hit the Earth's surface and greenhouse effects trapped heat. There were untapped sources of free energy within
the mixture of reducing and oxidizing gasses that set the stage for primitive ecosystems to evolve and, in turn, the
atmosphere also evolved.[177]
Throughout history, the Earth's atmosphere and biogeochemical cycles
have been in a dynamic equilibrium with planetary ecosystems. The
history is characterized by periods of significant transformation
followed by millions of years of stability.[178] The evolution of the
earliest organisms, likely anaerobic methanogen microbes, started the
process by converting atmospheric hydrogen into methane (4H2 + CO2
→ CH4 + 2H2O). Anoxygenic photosynthesis converting hydrogen
sulfide into other sulfur compounds or water (for example 2H2S + CO2
+ hv → CH2O + H2O + 2S), as occurs in deep sea hydrothermal vents
The leaf is the primary site of photosynthesis in
today, reduced hydrogen concentrations and increased atmospheric
most plants.
methane. Early forms of fermentation also increased levels of
atmospheric methane. The transition to an oxygen dominant
atmosphere (the Great Oxidation) did not begin until approximately 2.4-2.3 billion years ago, but photosynthetic
processes started 0.3 to 1 billion years prior.[178][179]
Radiation: heat, temperature and light
The biology of life operates within a certain range of temperatures. Heat is a form of energy that regulates
temperature. Heat affects growth rates, activity, behavior and primary production. Temperature is largely dependent
on the incidence of solar radiation. The latitudinal and longitudinal spatial variation of temperature greatly affects
climates and consequently the distribution of biodiversity and levels of primary production in different ecosystems or
biomes across the planet. Heat and temperature relate importantly to metabolic activity. Poikilotherms, for example,
have a body temperature that is largely regulated and dependent on the temperature of the external environment. In
contrast, homeotherms regulate their internal body temperature by expending metabolic energy.[119][120][166]
There is a relationship between light, primary production, and ecological energy budgets. Sunlight is the primary
input of energy into the planet's ecosystems. Light is composed of electromagnetic energy of different wavelengths.
Radiant energy from the sun generates heat, provides photons of light measured as active energy in the chemical
reactions of life, and also acts as a catalyst for genetic mutation.[119][120][166] Plants, algae, and some bacteria absorb
light and assimilate the energy through photosynthesis. Organisms capable of assimilating energy by photosynthesis
or through inorganic fixation of H2S are autotrophs. Autotrophs—responsible for primary production—assimilate
light energy that becomes metabolically stored as potential energy in the form of biochemical enthalpic
Physical environments
Wetland conditions such as shallow water, high plant productivity, and anaerobic substrates provide a suitable environment for
important physical, biological, and chemical processes. Because of these processes, wetlands play a vital role in global nutrient and
element cycles.:29
The rate of diffusion of carbon dioxide and oxygen is approximately 10,000 times slower in water than it is in air.
When soils become flooded, they quickly lose oxygen and transform into a low-concentration (hypoxic - O2
concentration lower than 2 mg/liter) environment and eventually become completely (anoxic) environment where
anaerobic bacteria thrive among the roots. Water also influences the spectral composition and amount of light as it
reflects off the water surface and submerged particles.[180] Aquatic plants exhibit a wide variety of morphological
and physiological adaptations that allow them to survive, compete and diversify these environments. For example,
the roots and stems develop large air spaces (Aerenchyma) that regulate the efficient transportation gases (for
example, CO2 and O2) used in respiration and photosynthesis. In drained soil, microorganisms use oxygen during
respiration. In aquatic environments, anaerobic soil microorganisms use nitrate, manganese ions, ferric ions, sulfate,
carbon dioxide and some organic compounds. The activity of soil microorganisms and the chemistry of the water
reduces the oxidation-reduction potentials of the water. Carbon dioxide, for example, is reduced to methane (CH4)
by methanogenic bacteria. Salt water plants (or halophytes) have specialized physiological adaptations, such as the
development of special organs for shedding salt and osmo-regulate their internal salt (NaCl) concentrations, to live in
estuarine, brackish, or oceanic environments.[180] The physiology of fish is also specially adapted to deal with high
levels of salt through osmoregulation. Their gills form electrochemical gradients that mediate salt excresion in saline
environments and uptake in fresh water.[181]
The shape and energy of the land is affected to a large degree by gravitational forces. On a larger scale, the
distribution of gravitational forces on the earth are uneven and influence the shape and movement of tectonic plates
as well as having an influence on geomorphic processes such as orogeny and erosion. These forces govern many of
the geophysical properties and distributions of ecological biomes across the Earth. On a organism scale, gravitational
forces provide directional cues for plant and fungal growth (gravitropism), orientation cues for animal migrations,
and influence the biomechanics and size of animals.[119] Ecological traits, such as allocation of biomass in trees
during growth are subject to mechanical failure as gravitational forces influence the position and structure of
branches and leaves.[182] The cardiovascular systems of all animals are functionally adapted to overcome pressure
and gravitational forces that change according to the features of organisms (e.g., height, size, shape), their behavior
(e.g., diving, running, flying), and the habitat occupied (e.g., water, hot deserts, cold tundra).[183]
Climatic and osmotic pressure places physiological constraints on organisms, such as flight and respiration at high
altitudes, or diving to deep ocean depths. These constraints influence vertical limits of ecosystems in the biosphere as
organisms are physiologically sensitive and adapted to atmospheric and osmotic water pressure differences.[119]
Oxygen levels, for example, decrease with increasing pressure and are a limiting factor for life at higher
altitudes.[184] Water transportation through trees is another important ecophysiological parameter where osmotic
pressure gradients factor in.[185][186][187] Water pressure in the depths of oceans requires that organisms adapt to
these conditions. For example, mammals, such as whales, dolphins and seals are specially adapted to deal with
changes in sound due to water pressure differences.[188] Different species of hagfish provide another example of
adaptation to deep-sea pressure through specialized protein adaptations.[189]
Wind and turbulence
Turbulent forces in air and water have significant effects on the
environment and ecosystem distribution, form and dynamics. On a
planetary scale, ecosystems are affected by circulation patterns in the
global trade winds. Wind power and the turbulent forces it creates can
influence heat, nutrient, and biochemical profiles of ecosystems.[119]
For example, wind running over the surface of a lake creates
turbulence, mixing the water column and influencing the
environmental profile to create thermally layered zones, partially
governing how fish, algae, and other parts of the aquatic ecology are
structured.[192][193] Wind speed and turbulence also exert influence on
rates of evapotranspiration rates and energy budgets in plants and
The architecture of inflorescence in grasses is
animals.[180][194] Wind speed, temperature and moisture content can
subject to the physical pressures of wind and
shaped by the forces of natural selection
vary as winds travel across different landfeatures and elevations. The
facilitating wind-pollination (or
westerlies, for example, come into contact with the coastal and interior
mountains of western North America to produce a rain shadow on the
leeward side of the mountain. The air expands and moisture condenses
as the winds move up in elevation which can cause precipitation; this is called orographic lift. This environmental
process produces spatial divisions in biodiversity, as species adapted to wetter conditions are range-restricted to the
coastal mountain valleys and unable to migrate across the xeric ecosystems of the Columbia Basin to intermix with
sister lineages that are segregated to the interior mountain systems.[195][196]
Forest fires modify the land by leaving behind an environmental mosaic that diversifies the landscape into different seral stages
and habitats of varied quality (left). Some species are adapted to forest fires, such as pine trees that open their cones only after
fire exposure (right).
Plants convert carbon dioxide into biomass and emit oxygen into the atmosphere.[197] Approximately 350 million
years ago (near the Devonian period) the photosynthetic process brought the concentration of atmospheric oxygen
above 17%, which allowed combustion to occur.[198] Fire releases CO2 and converts fuel into ash and tar. Fire is a
significant ecological parameter that raises many issues pertaining to its control and suppression in management.[199]
While the issue of fire in relation to ecology and plants has been recognized for a long time,[200] Charles Cooper
brought attention to the issue of forest fires in relation to the ecology of forest fire suppression and management in
the 1960s.[201][202]
Fire creates environmental mosaics and a patchiness to ecosystem age and canopy structure. Native North Americans
were among the first to influence fire regimes by controlling their spread near their homes or by lighting fires to
stimulate the production of herbaceous foods and basketry materials.[203] The altered state of soil nutrient supply and
cleared canopy structure also opens new ecological niches for seedling establishment.[204][205] Most ecosystem are
adapted to natural fire cycles. Plants, for example, are equipped with a variety of adaptations to deal with forest fires.
Some species (e.g., Pinus halepensis) cannot germinate until after their seeds have lived through a fire. This
environmental trigger for seedlings is called serotiny.[206] Some compounds from smoke also promote seed
germination.[207] Fire plays a major role in the persistence and resilience of ecosystems.[173]
Ecologists study and measure nutrient budgets to understand how these materials are regulated, flow, and recycled
through the environment.[119][120][166] This research has led to an understanding that there is a global feedback
between ecosystems and the physical parameters of this planet including minerals, soil, pH, ions, water and
atmospheric gases. There are six major elements, including H (hydrogen), C (carbon), N (nitrogen), O (oxygen), S
(sulfur), and P (phosphorus) that form the constitution of all biological macromolecules and feed into the Earth's
geochemical processes. From the smallest scale of biology the combined effect of billions upon billions of ecological
processes amplify and ultimately regulate the biogeochemical cycles of the Earth. Understanding the relations and
cycles mediated between these elements and their ecological pathways has significant bearing toward understanding
global biogeochemistry.[208]
The ecology of global carbon budgets gives one example of the linkage between biodiversity and biogeochemistry.
For starters, the Earth's oceans are estimated to hold 40,000 gigatonnes (Gt) carbon, vegetation and soil is estimated
to hold 2070 Gt carbon, and fossil fuel emissions are estimated to emit an annual flux of 6.3 Gt carbon.[209] At
different times in the Earth's history there has been major restructuring in these global carbon budgets that was
regulated to a large extent by the ecology of the land. For example, through the early-mid Eocene volcanic
outgassing, the oxidation of methane stored in wetlands, and seafloor gases increased atmospheric CO2 (carbon
dioxide) concentrations to levels as high as 3500 ppm.[210] In the Oligocene, from 25 to 32 million years ago, there
was another significant restructuring in the global carbon cycle as grasses evolved a special type of C4
photosynthesis and expanded their ranges. This new photosynthetic pathway evolved in response to the drop in
atmospheric CO2 concentrations below 550 ppm.[211] These kinds of ecosystem functions feed back significantly
into global atmospheric models for carbon cycling. Loss in the abundance and distribution of biodiversity causes
global carbon cycle feedbacks that are expected to increase rates of global warming in the next century.[212] The
effect of global warming melting large sections of permafrost creates a new mosaic of flooded areas where
decomposition results in the emission of methane (CH4). Hence, there is a relationship between global warming,
decomposition and respiration in soils and wetlands producing significant climate feedbacks and altered global
biogeochemical cycles.[213][214] There is concern over increases in atmospheric methane in the context of the global
carbon cycle, because methane is also a greenhouse gas that is 23 times more effective at absorbing long-wave
radiation than CO2 on a 100 year time scale.[215]
Early beginnings
Ecology has a complex origin due in large part to its interdisciplinary nature.[216] Ancient philosophers of Greece,
including Hippocrates and Aristotle were among the first to record their observations on natural history. However,
philosophers in ancient Greece viewed life as a static element that did not require an understanding of adaptation, a
modern cornerstone of ecological theory.[217] Topics more familiar in the modern context, including food chains,
population regulation, and productivity, did not develop until the 1700s through the published works of microscopist
Antoni van Leeuwenhoek (1632–1723) and botanist Richard Bradley (1688?-1732).[6] Biogeographer Alexander von
Humbolt (1769–1859) was another early pioneer in ecological thinking and was among the first to recognize
ecological gradients. Humbolt alluded to the modern ecological law of species to area relationships.[218][219]
In the early 20th century, ecology was an analytical form of natural history.[220] Following in the traditions of
Aristotle, the descriptive nature of natural history examined the interaction of organisms with both their environment
and their community. Natural historians, including James Hutton and Jean-Baptiste Lamarck, contributed significant
works that laid the foundations of the modern ecological sciences.[221] The term "ecology" (German: Oekologie) is
of a more recent origin and was first coined by the German biologist Ernst Haeckel in his book Generelle
Morphologie der Organismen (1866). Haeckel was a zoologist, artist, writer, and later in life a professor of
comparative anatomy.[222][223]
By ecology we mean the body of knowledge concerning the economy of nature-the investigation of the total relations of the animal
both to its inorganic and its organic environment; including, above all, its friendly and inimical relations with those animals and
plants with which it comes directly or indirectly into contact-in a word, ecology is the study of all those complex interrelations
referred to by Darwin as the conditions of the struggle of existence.
Haeckel's definition quoted in Esbjorn-Hargens
Ernst Haeckel (left) and Eugenius Warming (right), two founders of ecology
Opinions differ on who was the founder of modern ecological theory. Some mark Haeckel's definition as the
beginning,[225] others say it was Eugenius Warming with the writing of Oecology of Plants: An Introduction to the
Study of Plant Communities (1895).[226] Ecology may also be thought to have begun with Carl Linnaeus' research
principals on the economy of nature that matured in the early 18th century.[80][227] He founded an early branch of
ecological study he called the economy of nature.[80] The works of Linnaeus influenced Darwin in The Origin of
Species where he adopted the usage of Linnaeus' phrase on the economy or polity of nature.[222] Linnaeus was the
first to frame the balance of nature as a testable hypothesis. Haeckel, who admired Darwin's work, defined ecology
in reference to the economy of nature which has led some to question if ecology is synonymous with Linnaeus'
concepts for the economy of nature.[227]
The modern synthesis of ecology is a young science, which first attracted substantial formal attention at the end of
the 19th century (around the same time as evolutionary studies) and become even more popular during the 1960s
environmental movement,[221] though many observations, interpretations and discoveries relating to ecology extend
back to much earlier studies in natural history. For example, the concept on the balance or regulation of nature can
be traced back to Herodotos (died c. 425 BC) who described an early account of mutualism along the Nile river
where crocodiles open their mouths to beneficially allow sandpipers safe access to remove leeches.[216] In the
broader contributions to the historical development of the ecological sciences, Aristotle is considered one of the
earliest naturalists who had an influential role in the philosophical development of ecological sciences. One of
Aristotle's students, Theophrastus, made astute ecological observations about plants and posited a philosophical
stance about the autonomous relations between plants and their environment that is more in line with modern
ecological thought. Both Aristotle and Theophrastus made extensive observations on plant and animal migrations,
biogeography, physiology, and their habits in what might be considered an analog of the modern ecological
niche.[228][229] Hippocrates, another Greek philosopher, is also credited with reference to ecological topics in its
earliest developments.[6]
From Aristotle to Darwin the natural world was predominantly
considered static and unchanged since its original creation. Prior to The
Origin of Species there was little appreciation or understanding of the
dynamic and reciprocal relations between organisms, their adaptations
and their modifications to the environment.[232][224] While Charles
Darwin is most notable for his treatise on evolution,[233] he is also one
of the founders of soil ecology.[234] In The Origin of Species Darwin
also made note of the first ecological experiment that was published in
1816.[230] In the science leading up to Darwin the notion of evolving
species was gaining popular support. This scientific paradigm changed
the way that researchers approached the ecological sciences.[235]
The layout of the first ecological experiment,
noted by Charles Darwin in The Origin of
Species, was studied in a grass garden at Woburn
Abbey in 1817. The experiment studied the
performance of different mixtures of species
planted in different kinds of soils.
Nowhere can one see more clearly illustrated what may be called the sensibility of such an organic complex,--expressed by the fact
that whatever affects any species belonging to it, must speedily have its influence of some sort upon the whole assemblage. He will
thus be made to see the impossibility of studying any form completely, out of relation to the other forms,--the necessity for taking a
comprehensive survey of the whole as a condition to a satisfactory understanding of any part.
Stephen Forbes (1887)
After the turn of 20th century
Some suggest that the first ecological text (Natural History of Selborne) was published in 1789, by Gilbert White
(1720–1793).[237] The first American ecology book was published in 1905 by Frederic Clements.[238] In his book,
Clements forwarded the idea of plant communities as a superorganism. This publication launched a debate between
ecological holism and individualism that lasted until the 1970s. The Clements superorganism concept proposed that
ecosystems progress through regular and determined stages of seral development that are analogous to
developmental stages of an organism whose parts function to maintain the integrity of the whole. The Clementsian
paradigm was challenged by Henry Gleason.[239] According to Gleason, ecological communities develop from the
unique and coincidental association of individual organisms. This perceptual shift placed the focus back onto the life
histories of individual organisms and how this relates to the development of community associations.[240]
The Clementsian superorganism theory has not been completely rejected, but some suggest it was an overextended
application of holism.[114] Holism remains a critical part of the theoretical foundation in contemporary ecological
studies.[162] Holism was first introduced in 1926 by a polarizing historical figure, a South African General named
Jan Christian Smuts. Smuts was inspired by Clement's superorganism theory as he developed and published on the
concept of holism, which contrasts starkly against his racial political views as the father of apartheid.[241] Around the
same time, Charles Elton pioneered the concept of food chains in his classical book "Animal Ecology".[85] Elton[85]
defined ecological relations using concepts of food chains, food cycles, food size, and described numerical relations
among different functional groups and their relative abundance. Elton's 'food cycle' was replaced by 'food web' in a
subsequent ecological text.[242]
Ecology has developers in many nations, including Russia's Vladimir Vernadsky and his founding of the biosphere
concept in the 1920s[243] or Japan's Kinji Imanishi and his concepts of harmony in nature and habitat segregation in
the 1950s.[244] The scientific recognition or importance of contributions to ecology from other cultures is hampered
by language and translation barriers.[243]
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fmns. rug. nl/ pdf/ marbee/ 2006-Meysman-TREE. pdf). TRENDS in Ecology and Evolution 21 (22): 688–695. doi:10.1016/j.tree.2006.08.002.
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comunidades/ pdf/ pdf curso posgrado Elena/ Tema 1/ gleason1926. pdf). Bulletin of the Torrey Botanical Club 53 (1): 7–26.
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Further reading
• Beman, J. (2010). "Energy economics in ecosystems" (
energy-economics-in-ecosystems-13254442). Nature Education Knowledge 1 (8): 22.
• Bryant, P. J.. Biodiversity and Conservation. A Hypertext Book. (
• Cleland, E. E. (2011). "Biodiversity and Ecosystem Stability" (
library/biodiversity-and-ecosystem-stability-17059965). Nature Education Knowledge 2 (1): 2.
• Costanza, R.; Cumberland, J. H.; Daily, H.; Goodland, R.; Norgaard, R. B. (2007). An Introduction to Ecological
Economics (e-book). (
St. Lucie Press and International Society for Ecological Economics.
• "Ecosystem Services: A Primer." ( Ecological
Society of America. 2000.
• Farabee, M. J.. The Online Biology Book. (
biobooktoc.html). Avondale, Arizona: Estrella Mountain Community College.
• Forseth, I. (2010). "Terrestrial Biomes" (
terrestrial-biomes-13236757). Nature Education Knowledge 1 (8): 12.
• Henkel, T. P. (2010). "Coral reefs" (
coral-reefs-15786954). Nature Education Knowledge 1 (11): 5.
• McCabe, D. J. (2010). "Rivers and streams: Life in flowing water" (
knowledge/library/rivers-and-streams-life-in-flowing-water-16819919). Nature Education Knowledge 1 (12): 4.
• Odum, H. (1973). "Energy, ecology, and economics" (
EnergÃa,+economÃa+y+redistribución.pdf) (PDF). Ambio 2 (6): 220–227.
• Stevens, A. (2010). "Earth's varying climate" (
earth-s-varying-climate-13368032). Nature Education Knowledge 1 (8): 45.
• Stevens, A. (2010). "Predation, herbivory, and parasitism" (
library/predation-herbivory-and-parasitism-13261134). Nature Education Knowledge 1 (8): 38.
• Brand, F. S.; Jax, K. (2007). "Focusing the meaning(s) of resilience: resilience as a descriptive concept and a
boundary object" ( Ecology and Society 12 (1): 23.
• Carpenter, S. R.; Mooney, H. A.; Agard, J.; Capistrano, D.; DeFries, R. S.; Díaz, S.; Dietz, T.; Duraiappah, A. K.
et al (2009). "Science for managing ecosystem services: Beyond the Millennium Ecosystem Assessment" (http://
pdf) (PDF). Proceedings of the National Academy of Sciences 106 (5): 1305–1312.
Bibcode 2009PNAS..106.1305C. doi:10.1073/pnas.0808772106.
• Ettema, C.H.; Wardle, D.A. (2002). "Spatial soil ecology" (
ettema.pdf) (PDF). Trends in Ecology & Evolution 17 (4): 177–183. doi:10.1016/S0169-5347(02)02496-5.
• Getz, W.M. (2009). "Disease and the dynamics of food webs" (
citationList.action?articleURI=info:doi/10.1371/journal.pbio.1000209). PLoS Biol 7 (9): e1000209.
• Gotelli, N.J.; Ellison, A.M. (2006). "Food-Web models predict species abundances in response to habitat change"
( PLoS Biol 4 (10): e324.
• Green, J.L.; Hastings, A.; Arzberger, P.; Ayala, F.; Cottingham, K.L.; Cuddington, K.; Davis, F.; Dunne, J.A. et al
(2005). "Complexity in ecology and conservation: mathematical, statistical, and computational challenges" (http:/
/ et al.2005 Bioscience.pdf) (PDF). BioScience 55 (6):
501–510. doi:10.1641/0006-3568(2005)055[0501:CIEACM]2.0.CO;2. ISSN 0006-3568.
Hanski, I. (1998). "Metapopulation dynamics" ( 1998 Hanski.
pdf) (PDF). Nature 396 (6706): 41–49. Bibcode 1998Natur.396...41H. doi:10.1038/23876.
Heneghan, L.; Coleman, D. C.; Zou, X.; Crossley, D. A.; Haines, B. L. (1999). "Soil microarthropod
contributions to decomposition dynamics: Tropical-temperate comparisons of a single substrate" (http://cwt33. (PDF). Ecology 80:
Laland, K. N.; Odling-Smee, J.; Feldman, M. W. (2000). "Niche construction, biological evolution, and cultural
change" (
pdf) (PDF). Behavioral and Brain Sciences 23: 131–175. doi:10.1017/S0140525X00002417.
Magurran, A. E., & Henderson, P. A. 2010. Temporal turnover and the maintenance of diversity in ecological
assemblages. Philosophical Transactions of the Royal Society B: Biological Sciences, 365(1558):3611-3620
Peterson, G.; Allen, C. R.; Holling, C. S. (1998). "Ecological resilience, biodiversity, and scale" (http://www. (PDF). Ecosystems 1: 6–18.
• Quinn, J. F.; Dunham, A. E. (1983). "On hypothesis testing in ecology and evolution" (http://www.usm.maine.
edu/bio/courses/bio621/on_hypothesis_testing.pdf) (PDF). The American Naturalist 122 (5): 602–617.
• Saccheri, I.; Hanski, I. (2006). "Natural selection and population dynamics" (
Articles/TREE 2006 Saccheri &Hanski.pdf) (PDF). Trends in Ecology and Evolution 21 (6): 341–347.
doi:10.1016/j.tree.2006.03.018. PMID 16769435.
• Simberloff, D. S. (1974). "Equilibrium theory of island biogeography and ecology" (
110174.pdf) (PDF). Annual Review of Ecology and Systematics 5 (1): 161–182.
• Wiens, J. J.; Donoghue, M. J. (2004). "Historical biogeography, ecology and species richness" ( (PDF). Trends in Ecology & Evolution
19 (12): 639–644. doi:10.1016/j.tree.2004.09.011. PMID 16701326.
• Womack, A. M.; Bohannan, B. J. M.; Green, J. L. (2010). "Biodiversity and biogeography of the atmosphere"
( Philosophical Transactions
of the Royal Society B: Biological Sciences 365 (1558): 3645–3653. doi:10.1098/rstb.2010.0283.
External links
• Ecology (Stanford Encyclopedia of Philosophy) (
• The Nature Education Knowledge Project: Ecology (
• Ecology Journals List of ecological scientific journals (
• Ecology Dictionary - Explanation of Ecological Terms (
• Canadian Society for Ecology and Evolution (
• Ecological Society of America (
• Ecology Global Network (
• Ecological Society of Australia (
• British Ecological Society (
• Ecological Society of China (
• International Society for Ecological Economics (
( European Ecological Federation
UN Millennium Ecosystem Assessment (
The Encyclopedia of Earth – Wilderness: Biology & Ecology (
Ecology and Society - A journal of integrative science for resilience and sustainability (http://www.
• National Pesticide Information Center (United States)- Follow these steps to report an environmental incident
(wildlife, air, soil or water) (
• Science Aid: Ecology ( U.K. High School (GCSE,
Alevel) Ecology
Global warming
Global mean land-ocean temperature change from 1880–2011, relative to the 1951–1980 mean. The black line is the annual mean and the red line is
the 5-year running mean. The green bars show uncertainty estimates. Source: NASA GISS
The map shows the 10-year average (2000–2009) global mean temperature anomaly relative to the 1951–1980 mean. The largest temperature
increases are in the Arctic and the Antarctic Peninsula. Source: NASA Earth Observatory
Fossil fuel related CO2 emissions compared to five of IPCC's emissions scenarios. The dips are related to global recessions. Data from IPCC SRES
scenarios ; Data spreadsheet included with International Energy Agency's "CO2 Emissions from Fuel Combustion 2010 – Highlights" ; and
Supplemental IEA data . Image source: Skeptical Science
Global warming is the rising average temperature of Earth's atmosphere and oceans since the late 19th century and
its projected continuation. Since the early 20th century, Earth's average surface temperature has increased by about
0.8 °C (unknown operator: u'strong' °F), with about two thirds of the increase occurring since 1980.[6] Warming
of the climate system is unequivocal, and scientists are more than 90% certain that most of it is caused by increasing
concentrations of greenhouse gases produced by human activities such as deforestation and the burning of fossil
Global warming
fuels.[7][8][9][10] These findings are recognized by the national science academies of all major industrialized
Climate model projections are summarized in the 2007 Fourth Assessment Report (AR4) by the Intergovernmental
Panel on Climate Change (IPCC). They indicate that during the 21st century the global surface temperature is likely
to rise a further 1.1 to 2.9 °C (2 to 5.2 °F) for their lowest emissions scenario and 2.4 to 6.4 °C (4.3 to 11.5 °F) for
their highest.[12] The ranges of these estimates arise from the use of models with differing sensitivity to greenhouse
gas concentrations.[13][14]
An increase in global temperature will cause sea levels to rise and will change the amount and pattern of
precipitation, and a probable expansion of subtropical deserts.[15] Warming is expected to be strongest in the Arctic
and would be associated with continuing retreat of glaciers, permafrost and sea ice. Other likely effects of the
warming include more frequent occurrence of extreme-weather events including heat waves, droughts and heavy
rainfall, species extinctions due to shifting temperature regimes, and changes in crop yields. Warming and related
changes will vary from region to region around the globe, with projections being more robust in some areas than
others.[16] If global mean temperature increases to 4 °C (unknown operator: u'strong' °F) above preindustrial
levels, the limits for human adaptation are likely to be exceeded in many parts of the world, while the limits for
adaptation for natural systems would largely be exceeded throughout the world. Hence, the ecosystem services upon
which human livelihoods depend would not be preserved.[17]
Most countries are parties to the United Nations Framework Convention on Climate Change (UNFCCC),[18] whose
ultimate objective is to prevent dangerous anthropogenic (i.e., human-induced) climate change.[19] Parties to the
UNFCCC have adopted a range of policies designed to reduce greenhouse gas emissions[20]:10[21][22][23]:9 and to
assist in adaptation to global warming.[20]:13[23]:10[24][25] Parties to the UNFCCC have agreed that deep cuts in
emissions are required,[26] and that future global warming should be limited to below 2.0 °C (unknown operator:
u'strong' °F) relative to the pre-industrial level.[26][B] A 2011 report of analyses by the United Nations Environment
Programme[27] and International Energy Agency[28] suggest that efforts as of the early 21st century to reduce
emissions may be inadequately stringent to meet the UNFCCC's 2 °C target.
Observed temperature changes
Evidence for warming of the climate system includes observed
increases in global average air and ocean temperatures, widespread
melting of snow and ice, and rising global average sea level.[29][30][31]
The Earth's average surface temperature, expressed as a linear trend,
rose by 0.74 ± 0.18 °C over the period 1906–2005. The rate of
warming over the last half of that period was almost double that for the
period as a whole (0.13 ± 0.03 °C per decade, versus 0.07 ± 0.02 °C
per decade). The urban heat island effect is very small, estimated to
account for less than 0.002 °C of warming per decade since 1900.[32]
Temperatures in the lower troposphere have increased between 0.13
and 0.22 °C (0.22 and 0.4 °F) per decade since 1979, according to
satellite temperature measurements. Climate proxies show the
temperature to have been relatively stable over the one or two thousand
years before 1850, with regionally varying fluctuations such as the
Medieval Warm Period and the Little Ice Age.[33]
Two millennia of mean surface temperatures
according to different reconstructions from
climate proxies, each smoothed on a decadal
scale, with the instrumental temperature record
overlaid in black.
Recent estimates by NASA's Goddard Institute for Space Studies (GISS) and the National Climatic Data Center
show that 2005 and 2010 tied for the planet's warmest year since reliable, widespread instrumental measurements
became available in the late 19th century, exceeding 1998 by a few hundredths of a degree.[34][35][36] Estimates by
Global warming
the Climatic Research Unit (CRU) show 2005 as the second warmest year, behind 1998 with 2003 and 2010 tied for
third warmest year, however, "the error estimate for individual years ... is at least ten times larger than the
differences between these three years."[37] The World Meteorological Organization (WMO) statement on the status
of the global climate in 2010 explains that, "The 2010 nominal value of 0.53 °C ranks just ahead of those of 2005
(0.52 °C) and 1998 (0.51 °C), although the differences between the three years are not statistically significant..."[38]
Temperatures in 1998 were unusually warm because global
temperatures are affected by the El Niño-Southern Oscillation (ENSO),
and the strongest El Niño in the past century occurred during that
year.[39] Global temperature is subject to short-term fluctuations that
overlay long term trends and can temporarily mask them. The relative
stability in temperature from 2002 to 2009 is consistent with such an
episode.[40][41] 2010 was also an El Niño year. On the low swing of the
oscillation, 2011 as an La Niña year was cooler but it was still the 11th
warmest year since records began in 1880. Of the 13 warmest years
since 1880, 11 were the years from 2001 to 2011. Over the more recent
record, 2011 was the warmest La Niña year in the period from 1950 to
2011, and was close to 1997 which was not at the lowest point of the
NOAA graph of Global Annual Temperature
Anomalies 1950–2011, showing the El
Niño-Southern Oscillation
Temperature changes vary over the globe. Since 1979, land temperatures have increased about twice as fast as ocean
temperatures (0.25 °C per decade against 0.13 °C per decade).[43] Ocean temperatures increase more slowly than
land temperatures because of the larger effective heat capacity of the oceans and because the ocean loses more heat
by evaporation.[44] The Northern Hemisphere warms faster than the Southern Hemisphere because it has more land
and because it has extensive areas of seasonal snow and sea-ice cover subject to ice-albedo feedback. Although more
greenhouse gases are emitted in the Northern than Southern Hemisphere this does not contribute to the difference in
warming because the major greenhouse gases persist long enough to mix between hemispheres.[45]
The thermal inertia of the oceans and slow responses of other indirect effects mean that climate can take centuries or
longer to adjust to changes in forcing. Climate commitment studies indicate that even if greenhouse gases were
stabilized at 2000 levels, a further warming of about 0.5 °C (0.9 °F) would still occur.[46]
Initial causes of temperature changes (external forcings)
Greenhouse effect schematic showing energy flows between space, the atmosphere, and earth's surface. Energy exchanges are expressed in watts
per square meter (W/m2).
Global warming
This graph, known as the Keeling Curve, shows the increase of atmospheric carbon dioxide (CO2) concentrations from 1958–2008. Monthly CO2
measurements display seasonal oscillations in an upward trend; each year's maximum occurs during the Northern Hemisphere's late spring, and
declines during its growing season as plants remove some atmospheric CO2.
External forcing refers to processes external to the climate system (though not necessarily external to Earth) that
influence climate. Climate responds to several types of external forcing, such as radiative forcing due to changes in
atmospheric composition (mainly greenhouse gas concentrations), changes in solar luminosity, volcanic eruptions,
and variations in Earth's orbit around the Sun.[47]:0 Attribution of recent climate change focuses on the first three
types of forcing. Orbital cycles vary slowly over tens of thousands of years and at present are in an overall cooling
trend which would be expected to lead towards an ice age, but the 20th century instrumental temperature record
shows a sudden rise in global temperatures.[48]
Greenhouse gases
The greenhouse effect is the process by which absorption and emission of infrared radiation by gases in the
atmosphere warm a planet's lower atmosphere and surface. It was proposed by Joseph Fourier in 1824 and was first
investigated quantitatively by Svante Arrhenius in 1896.[49]
Naturally occurring amounts of greenhouse gases have a mean warming effect of about 33 °C (unknown operator:
u'strong' °F).[50][C] The major greenhouse gases are water vapor, which causes about 36–70% of the greenhouse
effect; carbon dioxide (CO2), which causes 9–26%; methane (CH4), which causes 4–9%; and ozone (O3), which
causes 3–7%.[51][52][53] Clouds also affect the radiation balance through cloud forcings similar to greenhouse gases.
Human activity since the Industrial Revolution has increased the amount of greenhouse gases in the atmosphere,
leading to increased radiative forcing from CO2, methane, tropospheric ozone, CFCs and nitrous oxide. The
concentrations of CO2 and methane have increased by 36% and 148% respectively since 1750.[54] These levels are
much higher than at any time during the last 800,000 years, the period for which reliable data has been extracted
from ice cores.[55][56][57][58] Less direct geological evidence indicates that CO2 values higher than this were last seen
about 20 million years ago.[59] Fossil fuel burning has produced about three-quarters of the increase in CO2 from
human activity over the past 20 years. The rest of this increase is caused mostly by changes in land-use, particularly
Per capita greenhouse gas emissions in 2005, including land-use change.
Total greenhouse gas emissions in 2005, including land-use change.
Over the last three decades of the 20th century, gross domestic product per capita and population growth were the
main drivers of increases in greenhouse gas emissions.[61] CO2 emissions are continuing to rise due to the burning of
fossil fuels and land-use change.[62][63]:71 Emissions can be attributed to different regions. The two figures opposite
show annual greenhouse gas emissions for the year 2005, including land-use change. Attribution of emissions due to
land-use change is a controversial issue.[64][65]:289
Global warming
Emissions scenarios, estimates of changes in future emission levels of greenhouse gases, have been projected that
depend upon uncertain economic, sociological, technological, and natural developments.[66] In most scenarios,
emissions continue to rise over the century, while in a few, emissions are reduced.[67][68] Fossil fuel reserves are
abundant, and will not limit carbon emissions in the 21st century.[69] Emission scenarios, combined with modelling
of the carbon cycle, have been used to produce estimates of how atmospheric concentrations of greenhouse gases
might change in the future. Using the six IPCC SRES "marker" scenarios, models suggest that by the year 2100, the
atmospheric concentration of CO2 could range between 541 and 970 ppm.[70] This is an increase of 90–250% above
the concentration in the year 1750.
The popular media and the public often confuse global warming with ozone depletion, i.e., the destruction of
stratospheric ozone by chlorofluorocarbons.[71][72] Although there are a few areas of linkage, the relationship
between the two is not strong. Reduced stratospheric ozone has had a slight cooling influence on surface
temperatures, while increased tropospheric ozone has had a somewhat larger warming effect.[73]
Particulates and soot
Ship tracks over the Atlantic Ocean on the east
coast of the United States. The climatic impacts
from particulate forcing could have a large effect
on climate through the indirect effect.
Global dimming, a gradual reduction in the amount of global direct
irradiance at the Earth's surface, was observed from 1961 until at least
1990.[74] The main cause of this dimming is particulates produced by
volcanoes and human made pollutants, which exerts a cooling effect by
increasing the reflection of incoming sunlight. The effects of the
products of fossil fuel combustion – CO2 and aerosols – have largely
offset one another in recent decades, so that net warming has been due
to the increase in non-CO2 greenhouse gases such as methane.[75]
Radiative forcing due to particulates is temporally limited due to wet
deposition which causes them to have an atmospheric lifetime of one
week. Carbon dioxide has a lifetime of a century or more, and as such,
changes in particulate concentrations will only delay climate changes
due to carbon dioxide.[76]
In addition to their direct effect by scattering and absorbing solar radiation, particulates have indirect effects on the
radiation budget.[77] Sulfates act as cloud condensation nuclei and thus lead to clouds that have more and smaller
cloud droplets. These clouds reflect solar radiation more efficiently than clouds with fewer and larger droplets,
known as the Twomey effect.[78] This effect also causes droplets to be of more uniform size, which reduces growth
of raindrops and makes the cloud more reflective to incoming sunlight, known as the Albrecht effect.[79] Indirect
effects are most noticeable in marine stratiform clouds, and have very little radiative effect on convective clouds.
Indirect effects of particulates represent the largest uncertainty in radiative forcing.[80]
Soot may cool or warm the surface, depending on whether it is airborne or deposited. Atmospheric soot directly
absorb solar radiation, which heats the atmosphere and cools the surface. In isolated areas with high soot production,
such as rural India, as much as 50% of surface warming due to greenhouse gases may be masked by atmospheric
brown clouds.[81] When deposited, especially on glaciers or on ice in arctic regions, the lower surface albedo can
also directly heat the surface.[82] The influences of particulates, including black carbon, are most pronounced in the
tropics and sub-tropics, particularly in Asia, while the effects of greenhouse gases are dominant in the extratropics
and southern hemisphere.[83]
Global warming
Solar activity
Solar variations causing changes in solar radiation energy reaching the
Earth have been the cause of past climate changes.[84] The effect of
changes in solar forcing in recent decades is uncertain, but small, with
some studies showing a slight cooling effect,[85] while others studies
suggest a slight warming effect.[47][86][87][88]
Greenhouse gases and solar forcing affect temperatures in different
Satellite observations of Total Solar Irradiance
ways. While both increased solar activity and increased greenhouse
from 1979–2006.
gases are expected to warm the troposphere, an increase in solar
activity should warm the stratosphere while an increase in greenhouse
gases should cool the stratosphere.[47] Radiosonde (weather balloon) data show the stratosphere has cooled over the
period since observations began (1958), though there is greater uncertainty in the early radiosonde record. Satellite
observations, which have been available since 1979, also show cooling.[89]
A related hypothesis, proposed by Henrik Svensmark, is that magnetic activity of the sun deflects cosmic rays that
may influence the generation of cloud condensation nuclei and thereby affect the climate.[90] Other research has
found no relation between warming in recent decades and cosmic rays.[91][92] The influence of cosmic rays on cloud
cover is about a factor of 100 lower than needed to explain the observed changes in clouds or to be a significant
contributor to present-day climate change.[93]
Studies in 2011 have indicated that solar activity may be slowing, and that the next solar cycle could be delayed. To
what extent is not yet clear; Solar Cycle 25 is due to start in 2020, but may be delayed to 2022 or even longer. It is
even possible that Sol could be heading towards another Maunder Minimum. While there is not yet a definitive link
between solar sunspot activity and global temperatures, the scientists conducting the solar activity study believe that
global greenhouse gas emissions would prevent any possible cold snap.[94]
“The fact we still see a positive imbalance despite the prolonged solar minimum isn't a surprise given what we've learned about the
climate system...But it's worth noting, because this provides unequivocal evidence that the sun is not the dominant driver of global
In line with other details mentioned above, director of NASA's Goddard Institute for Space Studies James Hansen
says that the sun is not nearly the biggest factor in global warming. Discussing the fact that low amounts of solar
activity between 2005 and 2010 had hardly any effect on global warming, Hansen says it is more evidence that
greenhouse gases are the largest culprit; that is, he supports the theory advanced by "nearly all climate scientists"
including the IPCC.[95]
Feedback is a process in which changing one quantity changes a second quantity, and the change in the second
quantity in turn changes the first. Positive feedback increases the change in the first quantity while negative feedback
reduces it. Feedback is important in the study of global warming because it may amplify or diminish the effect of a
particular process.
The main positive feedback in the climate system is the water vapor feedback. The main negative feedback is
radiative cooling through the Stefan–Boltzmann law, which increases as the fourth power of temperature. Positive
and negative feedbacks are not imposed as assumptions in the models, but are instead emergent properties that result
from the interactions of basic dynamical and thermodynamic processes.
A wide range of potential feedback processes exist, such as Arctic methane release and ice-albedo feedback.
Consequentially, potential tipping points may exist, which may have the potential to cause abrupt climate change.[96]
Global warming
For example, the "emission scenarios" used by IPCC in its 2007 report primarily examined greenhouse gas emissions
from human sources. In 2011, a joint study by the US National Snow and Ice Data Center and National Oceanic and
Atmospheric Administration calculated the additional greenhouse gas emissions that would emanate from melted
and decomposing permafrost, even if policymakers attempt to reduce human emissions from the A1FI scenario to the
A1B scenario.[97] The team found that even at the much lower level of human emissions, permafrost thawing and
decomposition would still result in 190 Gt C of permafrost carbon being added to the atmosphere on top of the
human sources. Importantly, the team made three extremely conservative assumptions: (1) that policymakers will
embrace the A1B scenario instead of the A1FI scenario, (2) that all of the carbon would be released as carbon
dioxide instead of methane, which is more likely and over a 20 year lifetime has 72x the greenhouse warming power
of CO2, and (3) their model did not project additional temperature rise caused by the release of these additional
gases.[97][98] These very conservative permafrost carbon dioxide emissions are equivalent to about 1/2 of all carbon
released from fossil fuel burning since the dawn of the Industrial Age,[99] and is enough to raise atmospheric
concentrations by an additional 87 ± 29 ppm, beyond human emissions. Once initiated, permafrost carbon forcing
(PCF) is irreversible, is strong compared to other global sources and sinks of atmospheric CO2, and due to thermal
inertia will continue for many years even if atmospheric warming stops.[97] A great deal of this permafrost carbon is
actually being released as highly flammable methane instead of carbon dioxide.[100] IPCC 2007's temperature
projections did not take any of the permafrost carbon emissions into account and therefore underestimate the degree
of expected climate change.[97][98]
Other research published in 2011 found that increased emissions of methane could instigate significant feedbacks
that amplify the warming attributable to the methane alone. The researchers found that a 2.5-fold increase in methane
emissions would cause indirect effects that increase the warming 250% above that of the methane alone. For a
5.2-fold increase, the indirect effects would be 400% of the warming from the methane alone.[101]
Climate models
Calculations of global warming prepared in or before 2001 from a range of climate models under the SRES A2 emissions scenario, which assumes
no action is taken to reduce emissions and regionally divided economic development.
The geographic distribution of surface warming during the 21st century calculated by the HadCM3 climate model if a business as usual scenario is
assumed for economic growth and greenhouse gas emissions. In this figure, the globally averaged warming corresponds to 3.0 °C (5.4 °F).
A climate model is a computerized representation of the five components of the climate system: Atmosphere,
hydrosphere, cryosphere, land surface, and biosphere.[102] Such models are based on physical principles including
fluid dynamics, thermodynamics and radiative transfer. There can be components which represent air movement,
temperature, clouds, and other atmospheric properties; ocean temperature, salt content, and circulation; ice cover on
land and sea; the transfer of heat and moisture from soil and vegetation to the atmosphere; chemical and biological
processes; and others.
Although researchers attempt to include as many processes as possible, simplifications of the actual climate system
are inevitable because of the constraints of available computer power and limitations in knowledge of the climate
system. Results from models can also vary due to different greenhouse gas inputs and the model's climate sensitivity.
For example, the uncertainty in IPCC's 2007 projections is caused by (1) the use of multiple models with differing
Global warming
sensitivity to greenhouse gas concentrations, (2) the use of differing estimates of humanities' future greenhouse gas
emissions, (3) any additional emissions from climate feedbacks that were not included in the models IPCC used to
prepare its report, i.e., greenhouse gas releases from permafrost.[97]
The models do not assume the climate will warm due to increasing levels of greenhouse gases. Instead the models
predict how greenhouse gases will interact with radiative transfer and other physical processes. One of the
mathematical results of these complex equations is a prediction whether warming or cooling will occur.[103]
Recent research has called special attention to the need to refine models with respect to the effect of clouds[104] and
the carbon cycle.[105][106][107]
Models are also used to help investigate the causes of recent climate change by comparing the observed changes to
those that the models project from various natural and human-derived causes. Although these models do not
unambiguously attribute the warming that occurred from approximately 1910 to 1945 to either natural variation or
human effects, they do indicate that the warming since 1970 is dominated by man-made greenhouse gas
The physical realism of models is tested by examining their ability to simulate contemporary or past climates.[108]
Climate models produce a good match to observations of global temperature changes over the last century, but do
not simulate all aspects of climate.[109] Not all effects of global warming are accurately predicted by the climate
models used by the IPCC. Observed Arctic shrinkage has been faster than that predicted.[110] Precipitation increased
proportional to atmospheric humidity, and hence significantly faster than global climate models predict.[111][112]
Expected environmental effects
"Detection" is the process of demonstrating that climate has changed in some defined statistical sense, without
providing a reason for that change. Detection does not imply attribution of the detected change to a particular cause.
"Attribution" of causes of climate change is the process of establishing the most likely causes for the detected change
with some defined level of confidence.[113] Detection and attribution may also be applied to observed changes in
physical, ecological and social systems.[114]
Natural systems
Global warming has been detected in a
number of systems. Some of these changes,
e.g., based on the instrumental temperature
record, have been described in the section
on temperature changes. Rising sea levels
and observed decreases in snow and ice
extent are consistent with warming.[115]
Most of the increase in global average
temperature since the mid-20th century is,
with high probability,[D] attributable to
human-induced changes in greenhouse gas
Sparse records indicate that glaciers have been retreating since the early 1800s. In
the 1950s measurements began that allow the monitoring of glacial mass balance,
reported to the World Glacier Monitoring Service (WGMS) and the National Snow
and Ice Data Center (NSIDC)
Even with policies to reduce emissions,
global emissions are still expected to
continue to grow over time.[117]
Global warming
In the IPCC Fourth Assessment Report, across a range of future emission scenarios, model-based estimates of sea
level rise for the end of the 21st century (the year 2090–2099, relative to 1980–1999) range from 0.18 to 0.59 m.
These estimates, however, were not given a likelihood due to a lack of scientific understanding, nor was an upper
bound given for sea level rise. On the timescale of centuries to millennia, the melting of ice sheets could result in
even higher sea level rise. Partial deglaciation of the Greenland ice sheet, and possibly the West Antarctic Ice Sheet,
could contribute 4–6 metres (13 to 20 ft) or more to sea level rise.[118]
Changes in regional climate are expected to include greater warming over land, with most warming at high northern
latitudes, and least warming over the Southern Ocean and parts of the North Atlantic Ocean.[117] Snow cover area
and sea ice extent are expected to decrease, with the Arctic expected to be largely ice-free in September by 2037.[119]
The frequency of hot extremes, heat waves, and heavy precipitation will very likely increase.
Ecological systems
In terrestrial ecosystems, the earlier timing of spring events, and poleward and upward shifts in plant and animal
ranges, have been linked with high confidence to recent warming.[115] Future climate change is expected to
particularly affect certain ecosystems, including tundra, mangroves, and coral reefs.[117] It is expected that most
ecosystems will be affected by higher atmospheric CO2 levels, combined with higher global temperatures.[120]
Overall, it is expected that climate change will result in the extinction of many species and reduced diversity of
Dissolved CO2 increases ocean acidity. This decreases the amount of carbonate ions, which organisms at the base of
the marine food chain, such as foraminifera, use to make structures they need to survive. The current rate of
acidification is many times faster than at least the past 300 million years, which included four mass extinctions that
involved rising ocean acidity. By the end of the century, acidity changes since the industrial revolution would match
the Palaeocene-Eocene Thermal Maximum, which occurred over 5000 years and killed 35-50% of benthic
Expected social system effects
Vulnerability of human societies to climate change mainly lies in the effects of extreme-weather events rather than
gradual climate change.[123] Impacts of climate change so far include adverse effects on small islands,[124] adverse
effects on indigenous populations in high-latitude areas,[125] and small but discernable effects on human health.[126]
Over the 21st century, climate change is likely to adversely affect hundreds of millions of people through increased
coastal flooding, reductions in water supplies, increased malnutrition and increased health impacts.[127] Most
economic studies suggest losses of world gross domestic product (GDP) for this magnitude of warming.[128][129]
Food security
Under present trends, by 2030, rice, millet and maize in South Asia could decrease by up to 10% while maize
production in Southern Africa.[130] By 2100, rice and maize yields in the tropics are expected to decrease by 20-40%
because of higher temperatures while the population of three billion is expected to double. This does not account for
the decrease in yields as a result of soil moisture and water supplies stressed by rising temperatures. [131]
Future warming of around 3 °C (by 2100, relative to 1990–2000) could result in increased crop yields in mid- and
high-latitude areas, but in low-latitude areas, yields could decline, increasing the risk of malnutrition.[124] A similar
regional pattern of net benefits and costs could occur for economic (market-sector) effects.[126] Warming above 3 °C
could result in crop yields falling in temperate regions, leading to a reduction in global food production.[132]
Global warming
Habitat inundation
In small islands and megadeltas, inundation as a result of sea level rise is expected to threaten vital infrastructure and
human settlements. [133] [134] This could lead to issues of statelessness for population from countries including the
Maldives and Tuvalu[135] and homelessness in countries with low lying areas such as Bangladesh.
Responses to global warming
Reducing the amount of future climate change is called mitigation of climate change. The IPCC defines mitigation as
activities that reduce greenhouse gas (GHG) emissions, or enhance the capacity of carbon sinks to absorb GHGs
from the atmosphere.[136] Many countries, both developing and developed, are aiming to use cleaner, less polluting,
technologies.[63]:192 Use of these technologies aids mitigation and could result in substantial reductions in CO2
emissions. Policies include targets for emissions reductions, increased use of renewable energy, and increased energy
efficiency. Studies indicate substantial potential for future reductions in emissions.[137]
In order to limit warming to within the lower range described in the IPCC's "Summary Report for Policymakers"[138]
it will be necessary to adopt policies that will limit greenhouse gas emissions to one of several significantly different
scenarios described in the full report.[139] This will become more and more difficult with each year of increasing
volumes of emissions and even more drastic measures will be required in later years to stabilize a desired
atmospheric concentration of greenhouse gases. Energy-related carbon-dioxide (CO2) emissions in 2010 were the
highest in history, breaking the prior record set in 2008.[140]
Since even in the most optimistic scenario, fossil fuels are going to be used for years to come, mitigation may also
involve carbon capture and storage, a process that traps CO2 produced by factories and gas or coal power stations
and then stores it, usually underground.[141]
Other policy responses include adaptation to climate change. Adaptation to climate change may be planned, either in
reaction to or anticipation of climate change, or spontaneous, i.e., without government intervention.[142] The ability
to adapt is closely linked to social and economic development.[137] Even societies with high capacities to adapt are
still vulnerable to climate change. Planned adaptation is already occurring on a limited basis. The barriers, limits, and
costs of future adaptation are not fully understood.
Views on global warming
There are different views over what the appropriate policy response to climate change should be.[143] These
competing views weigh the benefits of limiting emissions of greenhouse gases against the costs. In general, it seems
likely that climate change will impose greater damages and risks in poorer regions.[144]
Global warming controversy
The global warming controversy refers to a variety of disputes, significantly more pronounced in the popular media
than in the scientific literature,[145][146] regarding the nature, causes, and consequences of global warming. The
disputed issues include the causes of increased global average air temperature, especially since the mid-20th century,
whether this warming trend is unprecedented or within normal climatic variations, whether humankind has
contributed significantly to it, and whether the increase is wholly or partially an artifact of poor measurements.
Additional disputes concern estimates of climate sensitivity, predictions of additional warming, and what the
consequences of global warming will be.
Global warming
In the scientific literature, there is a strong consensus that global surface temperatures have increased in recent
decades and that the trend is caused mainly by human-induced emissions of greenhouse gases. No scientific body of
national or international standing disagrees with this view,[147][148] though a few organisations hold non-committal
From 1990–1997 in the United States, conservative think tanks mobilized to undermine the legitimacy of global
warming as a social problem. They challenged the scientific evidence; argued that global warming will have
benefits; and asserted that proposed solutions would do more harm than good.[149]
Most countries are Parties to the United Nations Framework
Convention on Climate Change (UNFCCC).[152] The ultimate
objective of the Convention is to prevent dangerous human
interference of the climate system.[153] As is stated in the Convention,
this requires that GHG concentrations are stabilized in the atmosphere
at a level where ecosystems can adapt naturally to climate change, food
production is not threatened, and economic development can proceed
in a sustainable fashion.[154] The Framework Convention was agreed in
1992, but since then, global emissions have risen.[155] During
negotiations, the G77 (a lobbying group in the United Nations
representing 133 developing nations)[156]:4 pushed for a mandate
requiring developed countries to "[take] the lead" in reducing their
emissions.[157] This was justified on the basis that: the developed
world's emissions had contributed most to the stock of GHGs in the
atmosphere; per-capita emissions (i.e., emissions per head of
population) were still relatively low in developing countries; and the
emissions of developing countries would grow to meet their
development needs.[65]:290 This mandate was sustained in the Kyoto
Protocol to the Framework Convention,[65]:290 which entered into legal
effect in 2005.[158]
Article 2 of the UN Framework Convention
refers explicitly to "stabilization of greenhouse
gas concentrations."
In order to stabilize the
atmospheric concentration of CO2, emissions
worldwide would need to be dramatically reduced
from their present level.
In ratifying the Kyoto Protocol, most developed countries accepted legally binding commitments to limit their
emissions. These first-round commitments expire in 2012.[158] US President George W. Bush rejected the treaty on
the basis that "it exempts 80% of the world, including major population centers such as China and India, from
compliance, and would cause serious harm to the US economy."[156]:5
At the 15th UNFCCC Conference of the Parties, held in 2009 at Copenhagen, several UNFCCC Parties produced the
Copenhagen Accord.[159] Parties associated with the Accord (140 countries, as of November 2010)[160]:9 aim to limit
the future increase in global mean temperature to below 2 °C.[161] A preliminary assessment published in November
2010 by the United Nations Environment Programme (UNEP) suggests a possible "emissions gap" between the
voluntary pledges made in the Accord and the emissions cuts necessary to have a "likely" (greater than 66%
probability) chance of meeting the 2 °C objective.[160]:10–14 The UNEP assessment takes the 2 °C objective as being
measured against the pre-industrial global mean temperature level. To having a likely chance of meeting the 2 °C
objective, assessed studies generally indicated the need for global emissions to peak before 2020, with substantial
declines in emissions thereafter.
The 16th Conference of the Parties (COP16) was held at Cancún in 2010. It produced an agreement, not a binding
treaty, that the Parties should take urgent action to reduce greenhouse gas emissions to meet a goal of limiting global
warming to 2 °C above pre-industrial temperatures. It also recognized the need to consider strengthening the goal to
Global warming
a global average rise of 1.5 °C.[162]
Public opinion
In 2007–2008 Gallup Polls surveyed 127 countries. Over a third of the
world's population was unaware of global warming, with people in
developing countries less aware than those in developed, and those in
Africa the least aware. Of those aware, Latin America leads in belief
that temperature changes are a result of human activities while Africa,
parts of Asia and the Middle East, and a few countries from the Former
Soviet Union lead in the opposite belief.[164] In the Western world,
opinions over the concept and the appropriate responses are divided.
Nick Pidgeon of Cardiff University said that "results show the different
Based on Rasmussen polling of 1,000 adults in
the USA conducted 29–30 July 2011.
stages of engagement about global warming on each side of the
Atlantic", adding, "The debate in Europe is about what action needs to
be taken, while many in the US still debate whether climate change is happening."[165][166] A 2010 poll by the Office
of National Statistics found that 75% of UK respondents were at least "fairly convinced" that the world's climate is
changing, compared to 87% in a similar survey in 2006.[167] A January 2011 ICM poll in the UK found 83% of
respondents viewed climate change as a current or imminent threat, while 14% said it was no threat. Opinion was
unchanged from an August 2009 poll asking the same question, though there had been a slight polarisation of
opposing views.[168]
A survey in October, 2009 by the Pew Research Center for the People & the Press showed decreasing public
perception in the US that global warming was a serious problem. All political persuasions showed reduced concern
with lowest concern among Republicans, only 35% of whom considered there to be solid evidence of global
warming.[169] The cause of this marked difference in public opinion between the US and the global public is
uncertain but the hypothesis has been advanced that clearer communication by scientists both directly and through
the media would be helpful in adequately informing the American public of the scientific consensus and the basis for
it.[170] The US public appears to be unaware of the extent of scientific consensus regarding the issue, with 59%
believing that scientists disagree "significantly" on global warming.[171]
By 2010, with 111 countries surveyed, Gallup determined that there was a substantial decrease in the number of
Americans and Europeans who viewed Global Warming as a serious threat. In the US, a little over half the
population (53%) now viewed it as a serious concern for either themselves or their families; this was 10% below the
2008 poll (63%). Latin America had the biggest rise in concern, with 73% saying global warming was a serious
threat to their families.[172] That global poll also found that people are more likely to attribute global warming to
human activities than to natural causes, except in the USA where nearly half (47%) of the population attributed
global warming to natural causes.[173]
On the other hand, in May 2011 a joint poll by Yale and George Mason Universities found that nearly half the
people in the USA (47%) attribute global warming to human activities, compared to 36% blaming it on natural
causes. Only 5% of the 35% who were "disengaged", "doubtful", or "dismissive" of global warming were aware that
97% of publishing US climate scientists agree global warming is happening and is primarily caused by humans.[174]
Researchers at the University of Michigan have found that the public's belief as to the causes of global warming
depends on the wording choice used in the polls.[175]
In the United States, according to the Public Policy Institute of California's (PPIC) eleventh annual survey on
environmental policy issues, 75% said they believe global warming is a very serious or somewhat serious threat to
the economy and quality of life in California.[176]
Global warming
A July 2011 Rasmussen Reports poll found that 69% of adults in the USA believe it is at least somewhat likely that
some scientists have falsified global warming research.[163]
A September 2011 Angus Reid Public Opinion poll found that Britons (43%) are less likely than Americans (49%)
or Canadians (52%) to say that "global warming is a fact and is mostly caused by emissions from vehicles and
industrial facilities." The same poll found that 20% of Americans, 20% of Britons and 14% of Canadians think
"global warming is a theory that has not yet been proven."[177]
Other views
Most scientists agree that humans are contributing to observed climate change.[62][178] National science academies
have called on world leaders for policies to cut global emissions.[179] However, some scientists and non-scientists
question aspects of climate-change science.[178][180][181]
Organizations such as the libertarian Competitive Enterprise Institute, conservative commentators, and some
companies such as ExxonMobil have challenged IPCC climate change scenarios, funded scientists who disagree with
the scientific consensus, and provided their own projections of the economic cost of stricter
controls.[182][183][184][185] In the finance industry, Deutsche Bank has set up an institutional climate change
investment division (DBCCA),[186] which has commissioned and published research[187] on the issues and debate
surrounding global warming.[188] Environmental organizations and public figures have emphasized changes in the
climate and the risks they entail, while promoting adaptation to changes in infrastructural needs and emissions
reductions.[189] Some fossil fuel companies have scaled back their efforts in recent years,[190] or called for policies to
reduce global warming.[191]
The term global warming was probably first used in its modern sense on 8 August 1975 in a science paper by Wally
Broecker in the journal Science called "Are we on the brink of a pronounced global warming?".[192][193][194]
Broecker's choice of words was new and represented a significant recognition that the climate was warming;
previously the phrasing used by scientists was "inadvertent climate modification," because while it was recognized
humans could change the climate, no one was sure which direction it was going.[195] The National Academy of
Sciences first used global warming in a 1979 paper called the Charney Report, which said: "if carbon dioxide
continues to increase, [we find] no reason to doubt that climate changes will result and no reason to believe that these
changes will be negligible."[196] The report made a distinction between referring to surface temperature changes as
global warming, while referring to other changes caused by increased CO2 as climate change.[195]
Global warming became more widely popular after 1988 when NASA climate scientist James Hansen used the term
in a testimony to Congress.[195] He said: "global warming has reached a level such that we can ascribe with a high
degree of confidence a cause and effect relationship between the greenhouse effect and the observed warming."[197]
His testimony was widely reported and afterward global warming was commonly used by the press and in public
A. ^ The 2001 joint statement was signed by the national academies of science of Australia, Belgium, Brazil,
Canada, the Caribbean, the People's Republic of China, France, Germany, India, Indonesia, Ireland, Italy,
Malaysia, New Zealand, Sweden, and the UK.[198] The 2005 statement added Japan, Russia, and the U.S. The
2007 statement added Mexico and South Africa. The Network of African Science Academies, and the Polish
Academy of Sciences have issued separate statements. Professional scientific societies include American
Astronomical Society, American Chemical Society, American Geophysical Union, American Institute of Physics,
American Meteorological Society, American Physical Society, American Quaternary Association, Australian
Global warming
Meteorological and Oceanographic Society, Canadian Foundation for Climate and Atmospheric Sciences,
Canadian Meteorological and Oceanographic Society, European Academy of Sciences and Arts, European
Geosciences Union, European Science Foundation, Geological Society of America, Geological Society of
Australia, Geological Society of London-Stratigraphy Commission, InterAcademy Council, International Union
of Geodesy and Geophysics, International Union for Quaternary Research, National Association of Geoscience
Teachers [199], National Research Council (US), Royal Meteorological Society, and World Meteorological
B. ^ Earth has already experienced almost 1/2 of the 2.0 °C (unknown operator: u'strong' °F) described in the
Cancun Agreement. In the last 100 years, Earth's average surface temperature increased by about 0.8 °C
(unknown operator: u'strong' °F) with about two thirds of the increase occurring over just the last three
C. ^ Note that the greenhouse effect produces an average worldwide temperature increase of about 33 °C
(unknown operator: u'strong' °F) compared to black body predictions without the greenhouse effect, not an
average surface temperature of 33 °C (unknown operator: u'strong' °F). The average worldwide surface
temperature is about 14 °C (unknown operator: u'strong' °F).
D. [50]
E. ^ In the IPCC Fourth Assessment Report, published in 2007, this attribution is given a probability of greater than
90%, based on expert judgement.[200] According to the US National Research Council Report – Understanding
and Responding to Climate Change – published in 2008, "[most] scientists agree that the warming in recent
decades has been caused primarily by human activities that have increased the amount of greenhouse gases in the
[1] http:/ / data. giss. nasa. gov/ gistemp/
[2] 2009 Ends Warmest Decade on Record (http:/ / earthobservatory. nasa. gov/ IOTD/ view. php?id=42392). NASA Earth Observatory Image of
the Day, 22 January 2010.
[3] http:/ / www. ipcc. ch/ ipccreports/ sres/ emission/ data/ allscen. xls
[4] http:/ / www. iea. org/ co2highlights/ co2Highlights. XLS
[5] http:/ / www. guardian. co. uk/ environment/ 2011/ may/ 29/ carbon-emissions-nuclearpower
[6] America's Climate Choices (http:/ / www. nap. edu/ openbook. php?record_id=12781& page=1). Washington, D.C.: The National Academies
Press. 2011. p. 15. ISBN 978-0-309-14585-5. . "The average temperature of the Earth’s surface increased by about 1.4 °F (0.8 °C) over the
past 100 years, with about 1.0 °F (0.6 °C) of this warming occurring over just the past three decades"
[7] "Warming of the climate system is unequivocal, as is now evident from observations of increases in global average air and ocean
temperatures, widespread melting of snow and ice and rising global average sea level." IPCC, Synthesis Report (http:/ / www. ipcc. ch/
publications_and_data/ ar4/ syr/ en/ main. html), Section 1.1: Observations of climate change (http:/ / www. ipcc. ch/ publications_and_data/
ar4/ syr/ en/ mains1. html), in IPCC AR4 SYR 2007.
[8] "Three different approaches are used to describe uncertainties each with a distinct form of language. * * * Where uncertainty in specific
outcomes is assessed using expert judgment and statistical analysis of a body of evidence (e.g. observations or model results), then the
following likelihood ranges are used to express the assessed probability of occurrence: virtually certain >99%; extremely likely >95%; very
likely >90%......" IPCC, Synthesis Report (http:/ / www. ipcc. ch/ publications_and_data/ ar4/ syr/ en/ main. html), Treatment of Uncertainty
(http:/ / www. ipcc. ch/ publications_and_data/ ar4/ syr/ en/ mainssyr-introduction. html), in IPCC AR4 SYR 2007.
[9] IPCC, Synthesis Report (http:/ / www. ipcc. ch/ publications_and_data/ ar4/ syr/ en/ main. html), Section 2.4: Attribution of climate change
(http:/ / www. ipcc. ch/ publications_and_data/ ar4/ syr/ en/ mains2-4. html), in IPCC AR4 SYR 2007.
[10] America's Climate Choices: Panel on Advancing the Science of Climate Change; National Research Council (2010). Advancing the Science
of Climate Change (http:/ / www. nap. edu/ catalog. php?record_id=12782). Washington, D.C.: The National Academies Press.
ISBN 0-309-14588-0. . "(p1) ... there is a strong, credible body of evidence, based on multiple lines of research, documenting that climate is
changing and that these changes are in large part caused by human activities. While much remains to be learned, the core phenomenon,
scientific questions, and hypotheses have been examined thoroughly and have stood firm in the face of serious scientific debate and careful
evaluation of alternative explanations. * * * (p21-22) Some scientific conclusions or theories have been so thoroughly examined and tested,
and supported by so many independent observations and results, that their likelihood of subsequently being found to be wrong is vanishingly
small. Such conclusions and theories are then regarded as settled facts. This is the case for the conclusions that the Earth system is warming
and that much of this warming is very likely due to human activities."
[11] "Joint Science Academies' Statement" (http:/ / nationalacademies. org/ onpi/ 06072005. pdf) (PDF). . Retrieved 9 August 2010.
Global warming
[12] Meehl et al., Chap. 10: Global Climate Projections (http:/ / www. ipcc. ch/ publications_and_data/ ar4/ wg1/ en/ ch10. html), Sec. 10.ES:
Mean Temperature (http:/ / www. ipcc. ch/ publications_and_data/ ar4/ wg1/ en/ ch10s10-es-1-mean-temperature. html), in IPCC AR4 WG1
[13] Schneider Von Deimling, Thomas; Held, Ganopolski, Rahmstorf (2006). "Climate sensitivity estimated from ensemble simulations of
glacial climate" (http:/ / citeseerx. ist. psu. edu/ viewdoc/ download?doi=10. 1. 1. 172. 3264& rep=rep1& type=pdf). Climate Dynamics. .
Retrieved 20 July 2011.
[14] Meehl et al., Chap. 10: Global Climate Projections (http:/ / www. ipcc. ch/ publications_and_data/ ar4/ wg1/ en/ ch10. html), Section 10.5:
Quantifying the Range of Climate Change (http:/ / www. ipcc. ch/ publications_and_data/ ar4/ wg1/ en/ ch10s10-5. html), in IPCC AR4 WG1
[15] Lu, Jian; Vechhi, Gabriel A.; Reichler, Thomas (2007). "Expansion of the Hadley cell under global warming" (http:/ / www. atmos.
berkeley. edu/ ~jchiang/ Class/ Spr07/ Geog257/ Week10/ Lu_Hadley06. pdf) (PDF). Geophysical Research Letters 34 (6): L06805.
Bibcode 2007GeoRL..3406805L. doi:10.1029/2006GL028443. .
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• IPCC AR4 SYR (2007), Core Writing Team; Pachauri, R.K; and Reisinger, A., ed., 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, ISBN 92-9169-122-4
• IPCC AR4 WG1 (2007), Solomon, S.; Qin, D.; Manning, M.; Chen, Z.; Marquis, M.; Averyt, K.B.; Tignor, M.;
and Miller, H.L., ed., Climate Change 2007: The Physical Science Basis (
publications_and_data/ar4/wg1/en/contents.html), Contribution of Working Group I to the Fourth Assessment
Report of the Intergovernmental Panel on Climate Change, Cambridge University Press,
ISBN 978-0-521-88009-1 (pb: 978-0-521-70596-7)
• IPCC AR4 WG2 (2007), Parry, M.L.; Canziani, O.F.; Palutikof, J.P.; van der Linden, P.J.; and Hanson, C.E., ed.,
Climate Change 2007: Impacts, Adaptation and Vulnerability (
wg2/en/contents.html), Contribution of Working Group II to the Fourth Assessment Report of the
Intergovernmental Panel on Climate Change, Cambridge University Press, ISBN 978-0-521-88010-7 (pb:
• IPCC AR4 WG3 (2007), Metz, B.; Davidson, O.R.; Bosch, P.R.; Dave, R.; and Meyer, L.A., ed., Climate Change
2007: Mitigation of Climate Change (,
Contribution of Working Group III to the Fourth Assessment Report of the Intergovernmental Panel on Climate
Change, Cambridge University Press, ISBN 978-0-521-88011-4 (pb: 978-0-521-70598-1)
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Second Assessment Synthesis of Scientific-Technical Information Relevant to Interpreting Article 2 of the UN Framework Convention on
Climate Change" and the Summaries for Policymakers of the three Working Groups.
• IPCC SAR WG3 (1996), Bruce, J.P.; Lee, H.; and Haites, E.F., ed., Climate Change 1995: Economic and Social
Dimensions of Climate Change, Contribution of Working Group III to the Second Assessment Report of the
Intergovernmental Panel on Climate Change, Cambridge University Press, ISBN 0-521-56051-9 (pb:
0-521-56854-4) pdf (
• IPCC TAR WG1 (2001), Houghton, J.T.; Ding, Y.; Griggs, D.J.; Noguer, M.; van der Linden, P.J.; Dai, X.;
Maskell, K.; and Johnson, C.A., ed., Climate Change 2001: The Scientific Basis (
publications/other/ipcc_tar/?src=/climate/ipcc_tar/wg1/index.htm), Contribution of Working Group I to the
Third Assessment Report of the Intergovernmental Panel on Climate Change, Cambridge University Press,
ISBN 0-521-80767-0 (pb: 0-521-01495-6)
• IPCC TAR WG2 (2001), McCarthy, J. J.; Canziani, O. F.; Leary, N. A.; Dokken, D. J.; and White, K. S., ed.,
Climate Change 2001: Impacts, Adaptation and Vulnerability (
ipcc_tar/?src=/climate/ipcc_tar/wg2/index.htm), Contribution of Working Group II to the Third Assessment
Report of the Intergovernmental Panel on Climate Change, Cambridge University Press, ISBN 0-521-80768-9
(pb: 0-521-01500-6)
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(, Contribution of
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Further reading
• Association of British Insurers (2005–06) (PDF). Financial Risks of Climate Change (http://www.climatewise.
• Ammann, Caspar; et al. (2007). "Solar influence on climate during the past millennium: Results from transient
simulations with the NCAR Climate Simulation Model" (
(PDF). Proceedings of the National Academy of Sciences of the United States of America 104 (10): 3713–3718.
Bibcode 2007PNAS..104.3713A. doi:10.1073/pnas.0605064103. PMC 1810336. PMID 17360418. "Simulations
with only natural forcing components included yield an early 20th century peak warming of 0.2 °C (≈1950 AD),
which is reduced to about half by the end of the century because of increased volcanism"
• Barnett, TP; Adam, JC; Lettenmaier, DP; Adam, J. C.; Lettenmaier, D. P. (17 November 2005). "Potential
impacts of a warming climate on water availability in snow-dominated regions" (
nature/journal/v438/n7066/abs/nature04141.html) (abstract). Nature 438 (7066): 303–309.
Bibcode 2005Natur.438..303B. doi:10.1038/nature04141. PMID 16292301.
• Behrenfeld, MJ; O'malley, RT; Siegel, DA; Mcclain, CR; Sarmiento, JL; Feldman, GC; Milligan, AJ; Falkowski,
PG et al; et al. (7 December 2006). "Climate-driven trends in contemporary ocean productivity" (http://www. (PDF). Nature 444 (7120): 752–755.
Bibcode 2006Natur.444..752B. doi:10.1038/nature05317. PMID 17151666.
• Choi, Onelack; Fisher, Ann (May 2005). "The Impacts of Socioeconomic Development and Climate Change on
Severe Weather Catastrophe Losses: Mid-Atlantic Region (MAR) and the U.S" (
content/m6308777613702q0/). Climate Change 58 (1–2): 149–170. doi:10.1023/A:1023459216609.
• Dyurgerov, Mark B.; Meier, Mark F. (2005) (PDF). Glaciers and the Changing Earth System: a 2004 Snapshot
( Institute of Arctic and Alpine
Research Occasional Paper #58. ISSN 0069-6145.
• Emanuel, K (4 August 2005). "Increasing destructiveness of tropical cyclones over the past 30 years" (ftp:// (PDF). Nature 436 (7051): 686–688.
Bibcode 2005Natur.436..686E. doi:10.1038/nature03906. PMID 16056221.
• Hansen, James; et al. (3 June 2005). "Earth's Energy Imbalance: Confirmation and Implications" (http://pangea.'s Energy Balance.pdf) (PDF). Science 308
(5727): 1431–1435. Bibcode 2005Sci...308.1431H. doi:10.1126/science.1110252. PMID 15860591.
• Hinrichs, Kai-Uwe; Hmelo, Laura R.; Sylva, Sean P. (21 February 2003). "Molecular Fossil Record of Elevated
Methane Levels in Late Pleistocene Coastal Waters". Science 299 (5610): 1214–1217.
Bibcode 2003Sci...299.1214H. doi:10.1126/science.1079601. PMID 12595688.
• Hirsch, Tim (11 January 2006). "Plants revealed as methane source" (
nature/4604332.stm). BBC.
• Hoyt, Douglas V.; Schatten, Kenneth H. (1993–11). "A discussion of plausible solar irradiance variations,
1700–1992". Journal of Geophysical Research 98 (A11): 18,895–18,906. Bibcode 1993JGR....9818895H.
• Karnaukhov, A. V. (2001). "Role of the Biosphere in the Formation of the Earth's Climate: The Greenhouse
Catastrophe" ( (PDF). Biophysics 46 (6).
• Kenneth, James P.; et al. (14 February 2003). Methane Hydrates in Quaternary Climate Change: The Clathrate
Gun Hypothesis ( American Geophysical
Global warming
• Keppler, Frank; et al. (18 January 2006). "Global Warming – The Blame Is not with the Plants" (http://www.
index.html). Max Planck Society.
• Lean, Judith L.; Wang, Y.M.; Sheeley, N.R. (2002–12). "The effect of increasing solar activity on the Sun's total
and open magnetic flux during multiple cycles: Implications for solar forcing of climate" (abstract). Geophysical
Research Letters 29 (24): 2224. Bibcode 2002GeoRL..29x..77L. doi:10.1029/2002GL015880.
• Lerner, K. Lee; Lerner, K. Lee; Wilmoth, Brenda (26 July 2006). Environmental issues: essential primary
sources. Thomson Gale. ISBN 1-4144-0625-8.
• McKibben, Bill (2011). The Global Warming Reader ( OR Books.
ISBN 978-1-935928-36-2.
• Muscheler, Raimund, R; Joos, F; Müller, SA; Snowball, I; et al. (28 July 2005). "Climate: How unusual is today's
solar activity?" (
(PDF). Nature 436 (7012): 1084–1087. Bibcode 2005Natur.436E...3M. doi:10.1038/nature04045.
PMID 16049429.
• Oerlemans, J. (29 April 2005). "Extracting a Climate Signal from 169 Glacier Records" (
abstracts/EGU05/04572/EGU05-J-04572.pdf) (PDF). Science 308 (5722): 675–677.
Bibcode 2005Sci...308..675O. doi:10.1126/science.1107046. PMID 15746388.
• Purse, BV; Mellor, PS; Rogers, DJ; Samuel, AR; Mertens, PP; Baylis, M; et al. (February 2005). "Climate change
and the recent emergence of bluetongue in Europe" (
nrmicro1090_fs.html) (abstract). Nature Reviews Microbiology 3 (2): 171–181. doi:10.1038/nrmicro1090.
PMID 15685226.
• Revkin, Andrew C (5 November 2005). "Rise in Gases Unmatched by a History in Ancient Ice" (http://www.
adxnnl=1&partner=rssuserland&emc=rss). The New York Times.
• Royal Society (2005). "Joint science academies' statement: Global response to climate change" (http:// Retrieved 19 April
• Ruddiman, William F. (15 December 2005). Earth's Climate Past and Future (
ruddiman/). New York: Princeton University Press. ISBN 0-7167-3741-8.
• Ruddiman, William F. (1 August 2005). Plows, Plagues, and Petroleum: How Humans Took Control of Climate.
New Jersey: Princeton University Press. ISBN 0-691-12164-8.
• Solanki, SK; Usoskin, IG; Kromer, B; Schüssler, M; Beer, J; et al. (23 October 2004). "Unusual activity of the
Sun during recent decades compared to the previous 11,000 years" (
nature02995.pdf) (PDF). Nature 431 (7012): 1084–1087. Bibcode 2004Natur.431.1084S.
doi:10.1038/nature02995. PMID 15510145.
• Solanki, Sami K.; et al. (28 July 2005). "Climate: How unusual is today's solar activity? (Reply)" (http://cc. (PDF). Nature 436 (7050): E4–E5.
Bibcode 2005Natur.436E...4S. doi:10.1038/nature04046.
• Sowers, Todd (10 February 2006). "Late Quaternary Atmospheric CH4 Isotope Record Suggests Marine
Clathrates Are Stable". Science 311 (5762): 838–840. Bibcode 2006Sci...311..838S.
doi:10.1126/science.1121235. PMID 16469923.
• Svensmark, Henrik; et al. (8 February 2007). "Experimental evidence for the role of ions in particle nucleation
under atmospheric conditions". Proceedings of the Royal Society A (FirstCite Early Online Publishing) 463
(2078): 385–396. Bibcode 2007RSPSA.463..385S. doi:10.1098/rspa.2006.1773.(online version requires
• Walter, KM; Zimov, SA; Chanton, JP; Verbyla, D; Chapin Fs, 3rd; et al. (7 September 2006). "Methane bubbling
from Siberian thaw lakes as a positive feedback to climate warming". Nature 443 (7107): 71–75.
Global warming
Bibcode 2006Natur.443...71W. doi:10.1038/nature05040. PMID 16957728.
• Wang, Y.-M.; Lean, J.L.; Sheeley, N.R. (20 May 2005). "Modeling the sun's magnetic field and irradiance since
1713" ( (PDF). Astrophysical Journal 625
(1): 522–538. Bibcode 2005ApJ...625..522W. doi:10.1086/429689.
External links
• NASA Goddard Institute for Space Studies ( – Global change research
• NOAA State of the Climate Report ( – U.S. and global
monthly state of the climate reports
• Climate Change at the National Academies (
Reports-Academies-Findings) – repository for reports
• Nature Reports Climate Change ( – free-access web resource
• Met Office: Climate change ( – UK National Weather Service
• Global Science and Technology Sources on the Internet ( –
commented list
• Educational Global Climate Modelling ( (EdGCM) – research-quality climate
change simulator
• DISCOVER ( – satellite-based ocean and climate data since 1979 from NASA
• Global Warming Art ( – collection of figures and images
• What Is Global Warming? (
html) – by National Geographic
• Global Climate Change Indicators ( – from NOAA
• NOAA Climate Services ( – from NOAA
• Global Warming Frequently Asked Questions (
html) – from NOAA
• Understanding Climate Change – Frequently Asked Questions (
climatechange/faqs.jsp) – from UCAR
• Global Warming: Center for Global Studies at the University of Illinois (
• Global Climate Change: NASA's Eyes on the Earth ( – from NASA's JPL and
• OurWorld 2.0 ( – from the United Nations University
• Center for Climate and Energy Solutions ( – business and politics
• Best Effort Global Warming Trajectories – Wolfram Demonstrations Project (http://demonstrations.wolfram.
com/BestEffortGlobalWarmingTrajectories/) – by Harvey Lam
• Koshland Science Museum – Global Warming Facts and Our Future (http://www.koshland-science-museum.
org/exhibitgcc/) – graphical introduction from National Academy of Sciences
• Climate Change: Coral Reefs on the Edge ( – A video presentation by
Prof. Ove Hoegh-Guldberg, University of Auckland
• Climate Change Indicators in the United States ( Report
by United States Environmental Protection Agency, 80 pp.
• Global Warming (
• Video on the effects of global warming on St. Lawrence Island in the Bering Sea (
Article Sources and Contributors
Article Sources and Contributors
Biome Source: Contributors: 0x6D667061, 12 Noon,, 149AFK,,, 18taniry, 1exec1,
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