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Ecology

Ecology is the study of the relationships between living organisms and their environment, encompassing various levels of organization from individuals to the biosphere. It has practical applications in conservation, resource management, and urban planning, and is closely related to other biological sciences. The field examines biodiversity, ecosystem dynamics, and the interactions between species and their habitats, highlighting the importance of ecological processes in sustaining life on Earth.
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0% found this document useful (0 votes)
49 views54 pages

Ecology

Ecology is the study of the relationships between living organisms and their environment, encompassing various levels of organization from individuals to the biosphere. It has practical applications in conservation, resource management, and urban planning, and is closely related to other biological sciences. The field examines biodiversity, ecosystem dynamics, and the interactions between species and their habitats, highlighting the importance of ecological processes in sustaining life on Earth.
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© © All Rights Reserved
We take content rights seriously. If you suspect this is your content, claim it here.
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Ecology

Ecology (from Ancient Greek οἶκος (oîkos) 'house' and


Ecology
-λογία (-logía) 'study of')[A] is the natural science of
the relationships among living organisms and their
environment. Ecology considers organisms at the
individual, population, community, ecosystem, and
biosphere levels. Ecology overlaps with the closely
related sciences of biogeography, evolutionary biology,
genetics, ethology, and natural history.

Ecology is a branch of biology, and is the study of


abundance, biomass, and distribution of organisms in
the context of the environment. It encompasses life
processes, interactions, and adaptations; movement of
materials and energy through living communities; Ecology addresses the full scale of life, from
successional development of ecosystems; cooperation, tiny bacteria to processes that span the entire
competition, and predation within and between planet. Ecologists study many diverse and
species; and patterns of biodiversity and its effect on complex relations among species, such as
ecosystem processes. predation and pollination. The diversity of life is
organized into different habitats, from
Ecology has practical applications in conservation terrestrial to aquatic ecosystems.
biology, wetland management, natural resource
management (agroecology, agriculture, forestry,
agroforestry, fisheries, mining, tourism), urban planning (urban ecology), community health, economics,
basic and applied science, and human social interaction (human ecology).

The word ecology (German: Ökologie) was coined in 1866 by the German scientist Ernst Haeckel. The
science of ecology as we know it today began with a group of American botanists in the 1890s.[1]
Evolutionary concepts relating to adaptation and natural selection are cornerstones of modern ecological
theory.

Ecosystems are dynamically interacting systems of organisms, the communities they make up, and the
non-living (abiotic) components of their environment. Ecosystem processes, such as primary production,
nutrient cycling, and niche construction, regulate the flux of energy and matter through an environment.
Ecosystems have biophysical feedback mechanisms that moderate processes acting on living (biotic) and
abiotic components of the planet. Ecosystems sustain life-supporting functions and provide ecosystem
services like biomass production (food, fuel, fiber, and medicine), the regulation of climate, global
biogeochemical cycles, water filtration, soil formation, erosion control, flood protection, and many other
natural features of scientific, historical, economic, or intrinsic value.

Levels, scope, and scale of organization


The scope of ecology contains a wide array of interacting levels of organization spanning micro-level
(e.g., cells) to a planetary scale (e.g., biosphere) phenomena. Ecosystems, for example, contain abiotic
resources and interacting life forms (i.e., individual organisms that aggregate into populations which
aggregate into distinct ecological communities). Because ecosystems are dynamic and do not necessarily
follow a linear successional route, changes might occur quickly or slowly over thousands of years before
specific forest successional stages are brought about by biological processes. An ecosystem's area can
vary greatly, from tiny to vast. A single tree is of little consequence to the classification of a forest
ecosystem, but is critically relevant to organisms living in and on it.[2] Several generations of an aphid
population can exist over the lifespan of a single leaf. Each of those aphids, in turn, supports diverse
bacterial communities.[3] 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.[4] 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.[5]

The main subdisciplines of ecology, population (or community) ecology and ecosystem ecology, exhibit a
difference not only in scale but also in two contrasting paradigms in the field. The former focuses on
organisms' distribution and abundance, while the latter focuses on materials and energy fluxes.[6]

Hierarchy
The scale of ecological dynamics can operate like a closed system, such as aphids migrating on a single
tree, while at the same time remaining open about broader scale influences, such as atmosphere or
climate. Hence, ecologists classify ecosystems hierarchically by analyzing data collected from finer scale
units, such as vegetation associations, climate, and soil types, and integrate this information to identify
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 conceptually manageable framework, the biological world is
organized into a nested hierarchy, ranging in scale from genes, to cells, to tissues, to organs, to organisms,
to species, to populations, to guilds, to communities, to ecosystems, to biomes, and up to the level of the
biosphere.[8] This framework forms a panarchy[9] and exhibits non-linear behaviors; this means that
"effect and cause are disproportionate, so that small changes to critical variables, such as the number of
nitrogen fixers, can lead to disproportionate, perhaps irreversible, changes in the system
properties."[10]: 14

Biodiversity
Biodiversity (an abbreviation of "biological diversity") describes the diversity of life from genes to
ecosystems and spans every level of biological organization. The term has several interpretations, and
there are many ways to index, measure, characterize, and represent its complex organization.[12][13][14]
Biodiversity includes species diversity, ecosystem diversity, and genetic diversity and scientists are
interested in the way that this diversity affects the complex ecological processes operating at and among
these respective levels.[13][15][16] Biodiversity plays an important role in ecosystem services which by
definition maintain and improve human quality of life.[14][17][18] Conservation priorities and management
techniques require different approaches and considerations to address the full ecological scope of
biodiversity. Natural capital that supports populations is critical for
maintaining ecosystem services[19][20] and species migration (e.g., riverine System behaviors
fish runs and avian insect control) has been implicated as one mechanism must first be arrayed
by which those service losses are experienced.[21] An understanding of into different levels of
biodiversity has practical applications for species and ecosystem-level the organization.
conservation planners as they make management recommendations to Behaviors
consulting firms, governments, and industry.[22] corresponding to
higher levels occur at
slow rates.
Habitat Conversely, lower
The habitat of a species describes the environment over which a species is organizational levels
known to occur and the type of community that is formed as a result.[24] exhibit rapid rates. For
More specifically, "habitats can be defined as regions in environmental example, individual
space that are composed of multiple dimensions, each representing a biotic tree leaves respond
or abiotic environmental variable; that is, any component or characteristic rapidly to momentary
of the environment related directly (e.g. forage biomass and quality) or changes in light
indirectly (e.g. elevation) to the use of a location by the animal."[25]: 745 For intensity, CO2
example, a habitat might be an aquatic or terrestrial environment that can be concentration, and the
further categorized as a montane or alpine ecosystem. Habitat shifts provide like. The growth of the
important evidence of competition in nature where one population changes tree responds more
relative to the habitats that most other individuals of the species occupy. For slowly and integrates
example, one population of a species of tropical lizard (Tropidurus hispidus) these short-term
has a flattened body relative to the main populations that live in open changes.
savanna. The population that lives in an isolated rock outcrop hides in
crevasses where its flattened body offers a selective advantage. Habitat O'Neill et al. (1986)[7]: 76
shifts also occur in the developmental life history of amphibians, and in
insects that transition from aquatic to terrestrial habitats. Biotope and
habitat are sometimes used interchangeably, but the former applies to a Biodiversity refers to
community's environment, whereas the latter applies to a species' the variety of life and
environment.[24][26][27] its processes. It
includes the variety of
living organisms, the
Niche genetic differences
Definitions of the niche date back to 1917,[30] but G. Evelyn Hutchinson among them, the
made conceptual advances in 1957[31][32] by introducing a widely adopted communities and
definition: "the set of biotic and abiotic conditions in which a species is able ecosystems in which
to persist and maintain stable population sizes."[30]: 519 The ecological niche they occur, and the
is a central concept in the ecology of organisms and is sub-divided into the ecological and
fundamental and the realized niche. The fundamental niche is the set of evolutionary processes
environmental conditions under which a species is able to persist. The that keep them
realized niche is the set of environmental plus ecological conditions under functioning, yet ever-
which a species persists.[30][32][33] The Hutchinsonian niche is defined more changing and
technically as a "Euclidean hyperspace whose dimensions are defined as adapting.
environmental variables and whose size is a function of the number of
values that the environmental values may assume for which an organism Noss & Carpenter
(1994)[11]: 5
has positive fitness."[34]: 71
Biogeographical patterns and range distributions are explained or
predicted through knowledge of a species' traits and niche
requirements.[35] Species have functional traits that are uniquely adapted
to the ecological niche. A trait is a measurable property, phenotype, or
characteristic of an organism that may influence its survival. Genes play
an important role in the interplay of development and environmental
expression of traits.[36] Resident species evolve traits that are fitted to the
selection pressures of 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 states that two species cannot coexist indefinitely by living off
the same limiting resource; one will always out-compete the other. When Biodiversity of a coral reef.
similarly adapted species overlap geographically, closer inspection reveals Corals adapt to and modify
subtle ecological differences in their habitat or dietary requirements.[37] their environment by
Some models and empirical studies, however, suggest that disturbances forming calcium carbonate
can stabilize the co-evolution and shared niche occupancy of similar skeletons. This provides
growing conditions for
species inhabiting species-rich communities.[38] The habitat plus the niche
future generations and
is called the ecotope, which is defined as the full range of environmental forms a habitat for many
and biological variables affecting an entire species.[24] other species.[23]

Niche construction
Organisms are subject to environmental pressures, but they also
modify their habitats. The regulatory feedback between organisms
and their environment can affect conditions from local (e.g., a
beaver pond) to global scales, over time and even after death, such
as decaying logs or silica skeleton deposits from marine
organisms.[39] The process and concept of ecosystem engineering
are related to niche construction, but the former relates only to the Long-tailed broadbill building its nest
physical modifications of the habitat whereas the latter also
considers the evolutionary implications of physical changes to the
environment and the feedback this causes on the process of natural selection. 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 habitats."[40]: 373

The ecosystem engineering concept has stimulated a new appreciation for the influence that organisms
have on the ecosystem and evolutionary process. The term "niche construction" is more often used in
reference to the under-appreciated feedback mechanisms of natural selection imparting forces on the
abiotic niche.[28][41] 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 is subject to
the forces of natural selection. Moreover, a nest can survive over
successive generations, so that progeny inherit both genetic material and a
legacy niche that was constructed before their time.[5][28][29]

Biome
Biomes are larger units of organization that categorize regions of the
Earth's ecosystems, mainly according to the structure and composition of
vegetation.[42] There are different methods to define the 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. Biomes include
Termite mounds with varied
tropical rainforest, temperate broadleaf and mixed forest, temperate
heights of chimneys
deciduous forest, taiga, tundra, hot desert, and polar desert.[43] Other
regulate gas exchange,
researchers have recently categorized other biomes, such as the human temperature and other
and oceanic microbiomes. To a microbe, the human body is a habitat and a environmental parameters
landscape.[44] Microbiomes were discovered largely through advances in that are needed to sustain
molecular genetics, which have revealed a hidden richness of microbial the internal physiology of
diversity on the planet. The oceanic microbiome plays a significant role in the entire colony.[28][29]
the ecological biogeochemistry of the planet's oceans.[45]

Biosphere
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 atmosphere's CO2 and O2 composition has been
affected by the biogenic flux of gases coming from respiration and photosynthesis, with levels fluctuating
over time in relation to the ecology and evolution of plants and animals.[46] Ecological theory has also
been used to explain self-emergent regulatory phenomena at the planetary scale: for example, the Gaia
hypothesis is an example of holism applied in ecological theory.[47] The Gaia hypothesis states that there
is an emergent feedback loop generated by the metabolism of living organisms that maintains the core
temperature of the Earth and atmospheric conditions within a narrow self-regulating range of
tolerance.[48]

Population ecology
Population ecology studies the dynamics of species populations and how these populations interact with
the wider environment.[5] A population consists of individuals of the same species that live, interact, and
migrate through the same niche and habitat.[49]

A primary law of population ecology is the Malthusian growth model[50] which states, "a population will
grow (or decline) exponentially as long as the environment experienced by all individuals in the
population remains constant."[50]: 18 Simplified population models usually starts with four variables:
death, birth, immigration, and emigration.
An example of an introductory population model describes a closed population, such as on an island,
where immigration and emigration does not take place. Hypotheses are evaluated with reference to a null
hypothesis which states that random processes create the observed data. In these island models, the rate of
population change is described by:

where N is the total number of individuals in the population, b and d are the per capita rates of birth and
death respectively, and r is the per capita rate of population change.[50][51]

Using these modeling techniques, Malthus' population principle of growth was later transformed into a
model known as the logistic equation by Pierre Verhulst:

where N(t) is the number of individuals measured as biomass density as a function of time, t, r is the
maximum per-capita rate of change commonly known as the intrinsic rate of growth, and is the
crowding coefficient, which represents the reduction in population growth rate per individual added. The
formula states that the rate of change in population size ( ) will grow to approach equilibrium,
where ( ), when the rates of increase and crowding are balanced, . A common,
analogous model fixes the equilibrium, as K, which is known as the "carrying capacity."

Population ecology builds upon these introductory models to further understand demographic processes
in real study populations. Commonly used types of data include life history, fecundity, and survivorship,
and these are analyzed using mathematical techniques such as matrix algebra. The information is used for
managing wildlife stocks and setting harvest quotas.[51][52] In cases where basic models are insufficient,
ecologists may adopt different kinds of statistical methods, such as the Akaike information criterion,[53]
or use models that can become mathematically complex as "several competing hypotheses are
simultaneously confronted with the data."[54]

Metapopulations and migration


The concept of metapopulations was defined in 1969[55] as "a population of populations which go extinct
locally and recolonize".[56]: 105 Metapopulation ecology is another statistical approach that is often used
in conservation research.[57] Metapopulation models simplify the landscape into patches of varying levels
of quality,[58] and 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 a habitat.[59] Migration is also a population-level phenomenon, as with the migration
routes followed by plants as they occupied northern post-glacial environments. Plant ecologists use 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 behavior, 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.[60][61]
In metapopulation terminology, migrating individuals are classed as emigrants (when they leave a region)
or immigrants (when they enter a region), 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; the
population at the site will disappear unless rescued by an adjacent source patch or environmental
conditions become more favorable. Metapopulation models examine patch dynamics over time to answer
potential 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 are 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 is the study of
Community ecology the interactions among a collection
examines how
of species that inhabit the same
interactions among
geographic area. Community
species and their ecologists study the determinants of
environment affect the
patterns and processes for two or
abundance, more interacting species. Research in
distribution and community ecology might measure Interspecific interactions such as
diversity of species
species diversity in grasslands in predation are a key aspect of
within communities. relation to soil fertility. It might also community ecology.
include the analysis of predator-prey
Johnson & Stinchcomb
dynamics, competition among
(2007)[64]: 250
similar plant species, or mutualistic interactions between crabs and corals.

Ecosystem ecology
Ecosystems may be habitats within
These ecosystems, as biomes that form an integrated
we may call them, are
whole and a dynamically responsive
of the most various system having both physical and
kinds and sizes. They biological complexes. Ecosystem
form one category of ecology is the science of determining
the multitudinous the fluxes of materials (e.g. carbon,
physical systems of
phosphorus) between different pools A riparian forest in the White
the universe, which (e.g., tree biomass, soil organic Mountains, New Hampshire (US) is
range from the material). Ecosystem ecologists an example of ecosystem ecology
universe as a whole attempt to determine the underlying
down to the atom. causes of these fluxes. Research in
ecosystem ecology might measure primary production (g C/m^2) in a
Tansley (1935)[65]: 299
wetland in relation to decomposition and consumption rates (g C/m^2/y).
This requires an understanding of the community connections between
plants (i.e., primary producers) and the decomposers (e.g., fungi and bacteria).[66]
The underlying concept of an ecosystem can be traced back to 1864 in the published work of George
Perkins Marsh ("Man and Nature").[67][68] Within an ecosystem, organisms are linked to the physical and
biological components of their environment to which they are adapted.[65] Ecosystems are complex
adaptive systems where the interaction of life processes form self-organizing patterns across different
scales of time and space.[69] Ecosystems are broadly categorized as terrestrial, freshwater, atmospheric, or
marine. Differences stem from the nature of the unique physical environments that shapes the
biodiversity within each. A more recent addition to ecosystem ecology are technoecosystems, which are
affected by or primarily the result of human activity.[5]

Food webs
A food web is the archetypal ecological
network. Plants capture solar energy and use
it to synthesize simple sugars during
photosynthesis. As plants grow, they
accumulate nutrients and are eaten by
grazing herbivores, and the energy is
transferred through a chain of organisms by
consumption. The simplified linear feeding
pathways that move from a basal trophic
species to a top consumer is called the food
chain. Food chains in an ecological
community create a complex food web.
Food webs are a type of concept map that is
used to illustrate and study pathways of
Generalized food web of waterbirds from Chesapeake Bay
energy and material flows.[7][70][71]

Empirical measurements are generally


restricted to a specific habitat, such as a cave or a pond, and principles gleaned from small-scale studies
are extrapolated to larger systems.[72] Feeding relations require extensive investigations, e.g. into the gut
contents of organisms, which can be difficult to decipher, or stable isotopes can be used to trace the flow
of nutrient diets and energy through a food web.[73] Despite these limitations, food webs remain a
valuable tool in understanding community ecosystems.[74]

Food webs illustrate important principles of ecology: some species have many weak feeding links (e.g.,
omnivores) while some are more specialized with fewer stronger feeding links (e.g., primary predators).
Such linkages explain how ecological communities remain stable over time[75][76] and eventually can
illustrate a "complete" web of life.[71][77][78][79]

The disruption of food webs may have a dramatic impact on the ecology of individual species or whole
ecosystems. For instance, the replacement of an ant species by another (invasive) ant species has been
shown to affect how elephants reduce tree cover and thus the predation of lions on zebras.[80][81]

Trophic levels
A trophic level (from Greek
troph, τροφή, trophē, meaning
"food" or "feeding") is "a
group of organisms acquiring
a considerable majority of its
energy from the lower
adjacent level (according to
ecological pyramids) nearer
the abiotic source."[82]: 383
Links in food webs primarily A trophic pyramid (a) and a food-web (b) illustrating ecological relationships
connect feeding relations or among creatures that are typical of a northern boreal terrestrial ecosystem.
trophism among species. The trophic pyramid roughly represents the biomass (usually measured as
Biodiversity within total dry-weight) at each level. Plants generally have the greatest biomass.
ecosystems can be organized Names of trophic categories are shown to the right of the pyramid. Some
ecosystems, such as many wetlands, do not organize as a strict pyramid,
into trophic pyramids, in
because aquatic plants are not as productive as long-lived terrestrial plants
which the vertical dimension
such as trees. Ecological trophic pyramids are typically one of three kinds:
represents feeding relations 1) pyramid of numbers, 2) pyramid of biomass, or 3) pyramid of
that become further removed energy.[5]: 598
from the base of the food
chain up toward top predators,
and the horizontal dimension represents the abundance or biomass at each level.[83] When the relative
abundance or biomass of each species is sorted into its respective trophic level, they naturally sort into a
'pyramid of numbers'.[84]

Species are broadly categorized as autotrophs (or primary producers), heterotrophs (or consumers), and
Detritivores (or decomposers). Autotrophs are organisms that produce their own food (production is
greater than respiration) by photosynthesis or chemosynthesis. Heterotrophs are organisms that must feed
on others for nourishment and energy (respiration exceeds production).[5] 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).[85] 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 compared to herbivores, they are relatively inefficient at
grazing.[86]

Trophic levels are part of the holistic or complex systems view of ecosystems.[87][88] Each trophic level
contains unrelated species that are grouped together because they share common ecological functions,
giving a macroscopic view of the system.[89] 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 led some ecologists to "reiterate that the notion that species clearly aggregate into
discrete, homogeneous trophic levels is fiction."[90]: 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."[91]: 612

Keystone species
A keystone species is a species that is connected to a
disproportionately large number of other 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 Sea otters, an example of a
keystone species results in a range of dramatic cascading effects keystone species
(termed trophic cascades) that alters trophic dynamics, other food
web connections, and can cause the extinction of other
species.[92][93] The term keystone species was coined by Robert Paine in 1969 and is a reference to the
keystone architectural feature as the removal of a keystone species can result in a community collapse
just as the removal of the keystone in an arch can result in the arch's loss of stability.[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 led indirectly 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 difficult
to experimentally determine what species may hold a keystone role in each ecosystem. Furthermore, food
web theory suggests that keystone species may not be common, so it is unclear how generally the
keystone species model can be applied.[95][97]

Complexity
Complexity is 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. 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."[98]: 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'.[99][100][E]

"Complexity in ecology is of at least six distinct types: spatial, temporal, structural, process, behavioral,
and geometric."[101]: 3 From 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 explanations at each integrative level.[48][102] Ecological complexity
relates to the dynamic resilience of ecosystems that transition to multiple shifting steady-states directed
by random fluctuations of history.[9][103] Long-term ecological studies provide important track records to
better understand the complexity and resilience of ecosystems over longer temporal and broader spatial
scales. These studies are managed by the International Long Term Ecological Network (LTER).[104] The
longest experiment in existence is the Park Grass Experiment, which was initiated in 1856.[105] Another
example is the Hubbard Brook study, which has been in operation since 1960.[106]

Holism
Holism remains a critical part of the theoretical foundation in contemporary ecological studies. Holism
addresses the biological organization of life that self-organizes into layers of emergent whole systems that
function according to non-reducible properties. 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.[107] "New properties emerge because the components interact, not because the basic nature of the
components is changed."[5]: 8

Ecological studies are necessarily holistic as opposed to reductionistic.[36][102][108] Holism has three
scientific meanings or uses that identify with ecology: 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. Scientific holism differs from mysticism that has appropriated the
same term. An example of metaphysical 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
principles of natural selection via predation without the need to reference or understand the biomolecular
properties of the exterior shells.[109]

Relation to evolution
Ecology and evolutionary biology 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, behavioural, and 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 principles, such as phylogenetics or the Linnaean system of taxonomy.[110]
The two disciplines often appear together, such as in the title of the journal Trends in Ecology and
Evolution.[111] 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.[36][48] While the boundary
between ecology and evolution is not always clear, ecologists study the abiotic and biotic factors that
influence evolutionary processes,[112][113] and evolution can be rapid, occurring on ecological timescales
as short as one generation.[114]

Behavioural ecology
All organisms can exhibit behaviours. Even plants express complex behaviour, including memory and
communication.[116] Behavioural ecology is the study of an organism's behaviour in its environment and
its ecological and evolutionary implications. Ethology is the study of observable movement or behaviour
in animals. This could include 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 gatherings of
amoeba.[117][118][119][120][121]

Adaptation is the central unifying concept in


Social display and colour variation in differently adapted behavioural ecology.[122] Behaviours can be
species of chameleons (Bradypodion spp.). Chameleons recorded as traits and inherited in much the
change their skin colour to match their background as a same way that eye and hair colour can.
behavioural defence mechanism and also use colour to Behaviours can evolve by means of natural
communicate with other members of their species, such selection as adaptive traits conferring
as dominant (left) versus submissive (right) patterns
functional utilities that increases reproductive
shown in the three species (A-C) above.[115]
fitness.[123][124]

Predator-prey interactions are an introductory concept into food-web


studies as well as behavioural ecology.[126] Prey species can exhibit
different kinds of behavioural 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
behavioural ecology are generally based on adaptive principles of
conservation, optimization, or efficiency.[33][112][127] For example, "[t]he
threat-sensitive predator avoidance hypothesis predicts that prey should
assess the degree of threat posed by different predators and match their
behaviour according to current levels of risk"[128] or "[t]he optimal flight
Mutualism: Leafhoppers
initiation distance occurs where expected postencounter fitness is (Eurymela fenestrata) are
maximized, which depends on the prey's initial fitness, benefits obtainable protected by ants
by not fleeing, energetic escape costs, and expected fitness loss due to (Iridomyrmex purpureus) in
predation risk."[129] a mutualistic relationship.
The ants protect the
Elaborate sexual displays and posturing are encountered in the leafhoppers from predators
behavioural ecology of animals. The birds-of-paradise, for example, sing and stimulate feeding in the
and display elaborate ornaments during courtship. These displays serve a leafhoppers, and in return,
the leafhoppers feeding on
dual purpose of signalling healthy or well-adapted individuals and
plants exude honeydew
desirable genes. The displays are driven by sexual selection as an from their anus that
advertisement of quality of traits among suitors.[130] provides energy and
nutrients to tending
ants.[125]
Cognitive ecology
Cognitive ecology integrates theory and observations from evolutionary
ecology and neurobiology, primarily cognitive science, in order to understand the effect that animal
interaction with their habitat has on their cognitive systems and how those systems restrict behavior
within an ecological and evolutionary framework.[131] "Until recently, however, cognitive scientists have
not paid sufficient attention to the fundamental fact that cognitive traits evolved under particular natural
settings. With consideration of the selection pressure on cognition, cognitive ecology can contribute
intellectual coherence to the multidisciplinary study of cognition."[132][133] As a study involving the
'coupling' or interactions between organism and environment, cognitive ecology is closely related to
enactivism,[131] a field based upon the view that "...we must see the organism and environment as bound
together in reciprocal specification and selection...".[134]

Social ecology
Social-ecological behaviours are notable in the social insects, slime moulds, social spiders, human
society, and naked mole-rats where eusocialism has evolved. Social behaviours include reciprocally
beneficial behaviours among kin and nest mates[119][124][135] and evolve from kin and group selection.
Kin selection explains altruism through genetic relationships, whereby an altruistic behaviour leading to
death is rewarded by the survival of genetic copies distributed among surviving relatives. The social
insects, including ants, bees, and wasps are most famously studied for this type of relationship because
the male drones are clones that share the same genetic make-up as every other male in the colony.[124] In
contrast, group selectionists find examples of altruism among non-genetic relatives and explain this
through selection acting on the group; whereby, it becomes selectively advantageous for groups if their
members express altruistic behaviours to one another. Groups with predominantly altruistic members
survive better than groups with predominantly selfish members.[124][136]

Coevolution
Ecological interactions can be classified broadly into a host and an
associate relationship. A host is any entity that harbours another
that is called the associate.[137] Relationships between species that
are mutually or reciprocally beneficial are called mutualisms.
Examples of 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
Bumblebees and the flowers they
with photosynthetic algae.[138][139] If there is a physical
pollinate have coevolved so that
connection between host and associate, the relationship is called both have become dependent on
symbiosis. Approximately 60% of all plants, for example, have a each other for survival.
symbiotic relationship with arbuscular mycorrhizal fungi living in
their roots forming an exchange network of carbohydrates for
mineral nutrients.[140]

Indirect mutualisms occur where the organisms live apart. For example, 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 others receive the benefits
of clean air at no cost or harm to trees supplying the oxygen.[5][141] If the associate benefits while the host
suffers, the relationship is called parasitism. Although parasites impose a cost to their host (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.[142][143] Co-
evolution is also driven by competition among species or among members of the same species under the
banner of reciprocal antagonism, such as grasses competing for growth
space. The Red Queen Hypothesis, for example, posits that parasites track
down and specialize on the locally common genetic defense systems of its
host that drives the evolution of sexual reproduction to diversify the
genetic constituency of populations responding to the antagonistic
pressure.[144][145]

Biogeography
Biogeography (an amalgamation of biology and geography) is the
Parasitism: A harvestman
comparative study of the geographic distribution of organisms and the
arachnid being parasitized
corresponding evolution of their traits in space and time.[146] The Journal by mites. The harvestman
of Biogeography was established in 1974.[147] Biogeography and ecology is being consumed, while
share many of their disciplinary roots. For example, the theory of island the mites benefit from
biogeography, published by the Robert MacArthur and Edward O. Wilson traveling on and feeding off
in 1967[148] is considered one of the fundamentals of ecological of their host.
theory.[149]

Biogeography has a long history in the natural sciences 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. The biogeographic processes that result in the natural splitting of species explain 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] 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 the context of global warming.[151][152]

r/K selection theory


A population ecology concept is r/K selection theory,[D] 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.[153]

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.[33] 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 favours 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][154]

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 the influx of research questions into this new ecological field resulted in the publication
Molecular Ecology in 1992.[155] 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.[156] 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 behavioural and biogeographical ecology.[156] For example,
molecular ecology revealed promiscuous sexual behaviour and multiple male partners in tree swallows
previously thought to be socially monogamous.[157] In a biogeographical context, the marriage between
genetics, ecology, and evolution resulted in a new sub-discipline called phylogeography.[158]

Human ecology
Ecology is as much a biological science as it is a human science.[5] Human
The history of life on ecology is an interdisciplinary investigation into the ecology of our species.
Earth has been a "Human ecology may be defined: (1) from a bioecological standpoint as the
history of interaction study of man as the ecological dominant in plant and animal communities
between living things and systems; (2) from a bioecological standpoint as simply another animal
and their affecting and being affected by his physical environment; and (3) as a
surroundings. To a human being, somehow different from animal life in general, interacting
large extent, the with physical and modified environments in a distinctive and creative way.
physical form and the A truly interdisciplinary human ecology will most likely address itself to all
habits of the earth's three."[160]: 3 The term was formally introduced in 1921, but many
vegetation and its sociologists, geographers, psychologists, and other disciplines were
animal life have been interested in human relations to natural systems centuries prior, especially in
molded by the the late 19th century.[160][161]
environment.
Considering the whole The ecological complexities human beings are facing through the
span of earthly time, technological transformation of the planetary biome has brought on the
the opposite effect, in Anthropocene. The unique set of circumstances has generated the need for a
which life actually new unifying science called coupled human and natural systems that builds
modifies its upon, but moves beyond the field of human ecology.[107] Ecosystems tie
surroundings, has been into human societies through the critical and all-encompassing life-
relatively slight. Only supporting functions they sustain. In recognition of these functions and the
within the moment of incapability of traditional economic valuation methods to see the value in
time represented by ecosystems, there has been a surge of interest in social-natural capital,
the present century has which provides the means to put a value on the stock and use of information
one species man and materials stemming from ecosystem goods and services. Ecosystems
acquired significant produce, regulate, maintain, and supply services of critical necessity and
power to alter the beneficial to human health (cognitive and physiological), economies, and
nature of his world. they even provide an information or reference function as a living library
giving opportunities for science and cognitive development in children
Rachel Carson, "Silent engaged in the complexity of the natural world. Ecosystems relate
[159]
Spring" importantly to human ecology as they are the ultimate base foundation of
global economics as every commodity, and the capacity for exchange
ultimately stems from the ecosystems on Earth.[107][162][163][164]

Restoration Ecology
Ecology is an employed science of restoration, repairing disturbed sites
through human intervention, in natural resource management, and in Ecosystem
environmental impact assessments. Edward O. Wilson predicted in 1992 management is not
just about science nor
that the 21st century "will be the era of restoration in ecology".[166]
Ecological science has boomed in the industrial investment of restoring is it simply an
extension of
ecosystems and their processes in abandoned sites after disturbance. Natural
traditional resource
resource managers, in forestry, for example, employ ecologists to develop,
adapt, and implement ecosystem based methods into the planning, management; it offers
a fundamental
operation, and restoration phases of land-use. Another example of
reframing of how
conservation is seen on the east coast of the United States in Boston, MA.
humans may work
The city of Boston implemented the Wetland Ordinance,[167] improving the
with nature.
stability of their wetland environments by implementing soil amendments
that will improve groundwater storage and flow, and trimming or removal
Grumbine (1994)[165]: 27
of vegetation that could cause harm to water quality. Ecological science is
used in the methods of sustainable harvesting, disease, and fire outbreak
management, in fisheries stock management, for integrating land-use with protected areas and
communities, and conservation in complex geo-political landscapes.[22][165][168][169]

Relation to the environment


The environment of ecosystems includes both physical parameters and biotic attributes. It is dynamically
interlinked and contains resources for organisms at any time throughout their life cycle.[5][170] Like
ecology, the term environment has different conceptual meanings and overlaps with the concept of nature.
Environment "includes the physical world, the social world of human relations and the built world of
human creation."[171]: 62 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.[172]

The distinction between external and internal environments, however, is an abstraction parsing life and
environment into units or facts that are inseparable in reality. There is an interpenetration of cause and
effect between the environment and life. The laws of thermodynamics, for example, apply to ecology by
means of its physical state. With an understanding of metabolic and thermodynamic principles, a
complete accounting of energy and material flow can be traced through an ecosystem. In this way, the
environmental and ecological relations are studied through reference to conceptually manageable and
isolated material parts. After the effective environmental components are understood through reference to
their causes; however, they conceptually link back together as an integrated whole, or holocoenotic
system as it was once called. This is known as the dialectical approach to ecology. The dialectical
approach examines the parts but integrates the organism and the environment into a dynamic whole (or
umwelt). Change in one ecological or environmental factor can concurrently affect the dynamic state of
an entire ecosystem.[36][173]

Disturbance and resilience


A disturbance is any process that changes or removes biomass from a community, such as a fire, flood,
drought, or predation.[174] Disturbances are both the cause and product of natural fluctuations within an
ecological community.[175][174][176][177] Biodiversity can protect ecosystems from disturbances.[177]

The effect of a disturbance is often hard to predict, but there are numerous examples in which a single
species can massively disturb an ecosystem. For example, a single-celled protozoan has been able to kill
up to 100% of sea urchins in some coral reefs in the Red Sea and Western Indian Ocean. Sea urchins
enable complex reef ecosystems to thrive by eating algae that would otherwise inhibit coral growth.[178]
Similarly, invasive species can wreak havoc on ecosystems. For instance, invasive Burmese pythons have
caused a 98% decline of small mammals in the Everglades.[179]

Metabolism and the early atmosphere


The Earth was formed approximately 4.5 billion years ago.[181] As it cooled
Metabolism – the rate
and a crust and oceans formed, its atmosphere transformed from being
at which energy and
dominated by hydrogen to one composed mostly of methane and ammonia.
Over the next billion years, the metabolic activity of life transformed the material resources are
taken up from the
atmosphere into a mixture of carbon dioxide, nitrogen, and water vapor.
environment,
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 transformed within an
energy within the mixture of reducing and oxidizing gasses that set the stage organism, and
allocated to
for primitive ecosystems to evolve and, in turn, the atmosphere also
maintenance, growth
evolved.[182]
and reproduction – is a
Throughout history, the Earth's atmosphere and biogeochemical cycles have fundamental
been in a dynamic equilibrium with planetary ecosystems. The history is physiological trait.
characterized by periods of significant transformation followed by millions
of years of stability.[183] The evolution of the earliest organisms, likely Ernest et al.[180]: 991
anaerobic methanogen microbes, started the process by converting
atmospheric hydrogen into methane (4H2 + CO2 → CH4 + 2H2O). Anoxygenic photosynthesis reduced
hydrogen concentrations and increased atmospheric methane, by converting hydrogen sulfide into water
or other sulfur compounds (for example, 2H2S + CO2 + hv → CH2O + H2O + 2S). 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.[183][184]

Radiation: heat, temperature and light


The biology of life operates within a certain range of
The leaf is the primary site of
photosynthesis in many higher
temperatures. Heat is a form of energy that regulates temperature.
plants. Heat affects growth rates, activity, behaviour, 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.[112][113][173]

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.[112][113][173] 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
which becomes metabolically stored as potential energy in the form of biochemical enthalpic
bonds.[112][113][173]

Physical environments

Water
Diffusion of carbon dioxide and oxygen is approximately 10,000 times
slower in water than in air. When soils are flooded, they quickly lose Wetland conditions
oxygen, becoming hypoxic (an environment with O2 concentration below such as shallow water,
high plant
2 mg/liter) and eventually completely anoxic where anaerobic bacteria
productivity, and
thrive among the roots. Water also influences the intensity and spectral
anaerobic substrates
composition of light as it reflects off the water surface and submerged
provide a suitable
particles.[185] Aquatic plants exhibit a wide variety of morphological and
environment for
physiological adaptations that allow them to survive, compete, and diversify
important physical,
in these environments. For example, their roots and stems contain large air
biological, and
spaces (aerenchyma) that regulate the efficient transportation of gases (for
chemical processes.
example, CO2 and O2) used in respiration and photosynthesis. Salt water
Because of these
plants (halophytes) have additional specialized adaptations, such as the
processes, wetlands
development of special organs for shedding salt and osmoregulating their
internal salt (NaCl) concentrations, to live in estuarine, brackish, or oceanic play a vital role in
environments. Anaerobic soil microorganisms in aquatic environments use global nutrient and
nitrate, manganese ions, ferric ions, sulfate, carbon dioxide, and some element cycles.
organic compounds; other microorganisms are facultative anaerobes and use
oxygen during respiration when the soil becomes drier. The activity of soil Cronk & Fennessy
microorganisms and the chemistry of the water reduces the oxidation- (2001)[185]: 29
reduction potentials of the water. Carbon dioxide, for example, is reduced to
methane (CH4) by methanogenic bacteria.[185] The physiology of fish is also specially adapted to
compensate for environmental salt levels through osmoregulation. Their gills form electrochemical
gradients that mediate salt excretion in salt water and uptake in fresh water.[186]

Gravity
The shape and energy of the land are significantly affected by gravitational forces. On a large scale, the
distribution of gravitational forces on the earth is uneven and influences the shape and movement of
tectonic plates as well as influencing geomorphic processes such as orogeny and erosion. These forces
govern many of the geophysical properties and distributions of ecological biomes across the Earth. On the
organismal 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.[112]
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.[187] The cardiovascular
systems of animals are functionally adapted to overcome the pressure and gravitational forces that change
according to the features of organisms (e.g., height, size, shape), their behaviour (e.g., diving, running,
flying), and the habitat occupied (e.g., water, hot deserts, cold tundra).[188]

Pressure
Climatic and osmotic pressure places physiological constraints on organisms, especially those that fly and
respire at high altitudes, or dive to deep ocean depths.[189] These constraints influence vertical limits of
ecosystems in the biosphere, as organisms are physiologically sensitive and adapted to atmospheric and
osmotic water pressure differences.[112] For example, oxygen levels decrease with decreasing pressure
and are a limiting factor for life at higher altitudes.[190] Water transportation by plants is another
important ecophysiological process affected by osmotic pressure gradients.[191][192][193] Water pressure in
the depths of oceans requires that organisms adapt to these conditions. For example, diving animals such
as whales, dolphins, and seals are specially adapted to deal with changes in sound due to water pressure
differences.[194] Differences between hagfish species provide another example of adaptation to deep-sea
pressure through specialized protein adaptations.[195]

Wind and turbulence


Turbulent forces in air and water affect 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.[112] 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, affecting how
fish, algae, and other parts of the aquatic ecosystem are structured.[198][199] Wind speed and turbulence
also influence evapotranspiration rates and energy budgets in plants and animals.[185][200] Wind speed,
temperature and moisture content can vary as winds travel across
different land features and elevations. For example, the westerlies
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 increase in elevation; this is called orographic lift and
can cause precipitation. 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 (e.g., of the
Columbia Basin in western North America) to intermix with sister The architecture of the inflorescence
lineages that are segregated to the interior mountain in grasses is subject to the physical
systems.[201][202] pressures of wind and shaped by
the forces of natural selection
facilitating wind-pollination
Fire (anemophily).[196][197]
Plants convert carbon dioxide into biomass and emit oxygen into

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).

the atmosphere. By approximately 350 million years ago (the end of the Devonian period),
photosynthesis had brought the concentration of atmospheric oxygen above 17%, which allowed
combustion to occur.[203] 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.[204] While the issue
of fire in relation to ecology and plants has been recognized for a long time,[205] Charles Cooper brought
attention to the issue of forest fires in relation to the ecology of forest fire suppression and management in
the 1960s.[206][207]

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.[208] Fire creates a heterogeneous ecosystem age and canopy structure, and the altered soil
nutrient supply and cleared canopy structure opens new ecological niches for seedling
establishment.[209][210] Most ecosystems 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 or been exposed to certain compounds
from smoke. Environmentally triggered germination of seeds is called serotiny.[211][212] Fire plays a
major role in the persistence and resilience of ecosystems.[176]

Soils
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 (for example, leaves on the forest floor), results in
soils containing minerals and nutrients that feed into plant production. The whole of the planet's soil
ecosystems is called the pedosphere where a large biomass of the Earth's biodiversity organizes into
trophic levels. Invertebrates that feed and shred larger leaves, for example, create smaller bits for smaller
organisms in the feeding chain. Collectively, these organisms are the detritivores that regulate soil
formation.[213][214] 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. Soils form composite phenotypes where inorganic matter is enveloped into the physiology of
a whole community. As organisms feed and migrate through soils they physically displace materials, an
ecological process called bioturbation. This aerates soils and stimulates heterotrophic growth and
production. Soil microorganisms are influenced by and are fed back into the trophic dynamics of the
ecosystem. No single axis of causality can be discerned to segregate the biological from
geomorphological systems in soils.[215][216] 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 the
colonization of land in the Devonian period played a significant role in the early development of
ecological trophism in soils.[214][217][218]

Biogeochemistry and climate


Ecologists study and measure nutrient budgets to understand how these materials are regulated, flow, and
recycled through the environment.[112][113][173] This research has led to an understanding that there is
global feedback between ecosystems and the physical parameters of this planet, including minerals, soil,
pH, ions, water, and atmospheric gases. Six major elements (hydrogen, carbon, nitrogen, oxygen, sulfur,
and phosphorus; H, C, N, O, S, and P) 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.[219]

The ecology of global carbon budgets gives one example of the linkage between biodiversity and
biogeochemistry. It is estimated that the Earth's oceans hold 40,000 gigatonnes (Gt) of carbon, that
vegetation and soil hold 2070 Gt, and that fossil fuel emissions are 6.3 Gt carbon per year.[220] There
have been major restructurings in these global carbon budgets during the Earth's history, 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.[221]

In the Oligocene, from twenty-five to thirty-two million years ago, there was another significant
restructuring of the global carbon cycle as grasses evolved a new mechanism of photosynthesis, C4
photosynthesis, and expanded their ranges. This new pathway evolved in response to the drop in
atmospheric CO2 concentrations below 550 ppm.[222] The relative abundance and distribution of
biodiversity alters the dynamics between organisms and their environment such that ecosystems can be
both cause and effect in relation to climate change. Human-driven modifications to the planet's
ecosystems (e.g., disturbance, biodiversity loss, agriculture) contributes to rising atmospheric greenhouse
gas levels. Transformation of the global carbon cycle in the next century is projected to raise planetary
temperatures, lead to more extreme fluctuations in weather, alter species distributions, and increase
extinction rates. The effect of global warming is already being registered in melting glaciers, melting
mountain ice caps, and rising sea levels. Consequently, species distributions are changing along
waterfronts and in continental areas where migration patterns and breeding grounds are tracking the
prevailing shifts in climate. Large sections of permafrost are also melting to create a new mosaic of
flooded areas having increased rates of soil decomposition activity that raises methane (CH4) emissions.
There is concern over increases in atmospheric methane in the context of the global carbon cycle, because
methane is a greenhouse gas that is 23 times more effective at absorbing long-wave radiation than CO2
on a 100-year time scale.[223] Hence, there is a relationship between global warming, decomposition and
respiration in soils and wetlands producing significant climate feedbacks and globally altered
biogeochemical cycles.[107][224][225][226][227][228]

History

Early beginnings
Ecology has a complex origin, due in large part to its interdisciplinary
By ecology, we mean the
nature.[230] Ancient Greek philosophers such as Hippocrates and
whole science of the
Aristotle were among the first to record observations on natural
history. However, they viewed life in terms of essentialism, where relations of the organism to
the environment including,
species were conceptualized as static unchanging things while
in the broad sense, all the
varieties were seen as aberrations of an idealized type. This contrasts
against the modern understanding of ecological theory where varieties "conditions of existence".
Thus, the theory of
are viewed as the real phenomena of interest and having a role in the
evolution explains the
origins of adaptations by means of natural selection.[5][231][232] Early
housekeeping relations of
conceptions of ecology, such as a balance and regulation in nature can
organisms mechanistically
be traced to Herodotus (died c. 425 BC), who described one of the
as the necessary
earliest accounts of mutualism in his observation of "natural
consequences of effectual
dentistry". Basking Nile crocodiles, he noted, would open their mouths
causes; and so forms the
to give sandpipers safe access to pluck leeches out, giving nutrition to
monistic groundwork of
the sandpiper and oral hygiene for the crocodile.[230] Aristotle was an
ecology.
early influence on the philosophical development of ecology. He and
his student Theophrastus made extensive observations on plant and
Ernst Haeckel (1866)[229]: 140
animal migrations, biogeography, physiology, and their behavior, [B]
giving an early analogue to the modern concept of an ecological
niche.[233][234]

Ecological concepts such as food chains, population regulation, and


productivity were first developed in the 1700s, through the published Nowhere can one see more
works of microscopist Antonie van Leeuwenhoek (1632–1723) and clearly illustrated what may
botanist Richard Bradley (1688?–1732).[5] Biogeographer Alexander be called the sensibility of
von Humboldt (1769–1859) was an early pioneer in ecological such an organic complex, –
thinking and was among the first to recognize ecological gradients, expressed by the fact that
where species are replaced or altered in form along environmental whatever affects any
gradients, such as a cline forming along a rise in elevation. Humboldt species belonging to it,
drew inspiration from Isaac Newton, must speedily have its
as he developed a form of influence of some sort upon
"terrestrial physics". In Newtonian the whole assemblage. He
fashion, he brought a scientific will thus be made to see the
exactitude for measurement into impossibility of studying
natural history and even alluded to any form completely, out of
concepts that are the foundation of a relation to the other forms,
Ernst Haeckel (left) and Eugenius modern ecological law on species- – the necessity for taking a
Warming (right), two founders of
to-area relationships.[236][237][238] comprehensive survey of
ecology
Natural historians, such as the whole as a condition to
Humboldt, James Hutton, and Jean-Baptiste Lamarck (among others) a satisfactory understanding
laid the foundations of the modern ecological sciences.[239] The term of any part.
"ecology" (German: Oekologie, Ökologie) was coined by Ernst
Haeckel in his book Generelle Morphologie der Organismen Stephen Forbes (1887)[235]
(1866).[240] Haeckel was a zoologist, artist, writer, and later in life a
professor of comparative anatomy.[229][241]

Opinions differ on who was the founder of modern ecological theory. Some mark Haeckel's definition as
the beginning;[242] others say it was Eugenius Warming with the writing of Oecology of Plants: An
Introduction to the Study of Plant Communities (1895),[243] or Carl Linnaeus' principles on the economy
of nature that matured in the early 18th century.[244][245] Linnaeus founded an early branch of ecology
that he called the economy of nature.[244] His works influenced Charles Darwin, who adopted Linnaeus'
phrase on the economy or polity of nature in The Origin of Species.[229] 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 whether ecology and the economy of
nature are synonymous.[245]

From Aristotle until Darwin, the natural world was predominantly


considered static and unchanging. Prior to The Origin of Species,
there was little appreciation or understanding of the dynamic and
reciprocal relations between organisms, their adaptations, and the
environment.[231] An exception is the 1789 publication Natural
History of Selborne by Gilbert White (1720–1793), considered by The layout of the first ecological
some to be one of the earliest texts on ecology.[248] While Charles experiment, carried out in a grass
Darwin is mainly noted for his treatise on evolution,[249] he was garden at Woburn Abbey in 1816,
one of the founders of soil ecology,[250] and he made note of the was noted by Charles Darwin in The
first ecological experiment in The Origin of Species.[246] Origin of Species. The experiment
studied the performance of different
Evolutionary theory changed the way that researchers approached
mixtures of species planted in
the ecological sciences.[251]
different kinds of soils.[246][247]

Since 1900
Modern ecology is a young science that first attracted substantial scientific attention toward the end of the
19th century (around the same time that evolutionary studies were gaining scientific interest). The
scientist Ellen Swallow Richards adopted the term "oekology" (which eventually morphed into home
economics) in the U.S. as early as 1892.[252]
In the early 20th century, ecology transitioned from a more descriptive form of natural history to a more
analytical form of scientific natural history.[236][239][253] Frederic Clements published the first American
ecology book in 1905,[254] presenting the idea of plant communities as a superorganism. This publication
launched a debate between ecological holism and individualism that lasted until the 1970s. Clements'
superorganism concept proposed that ecosystems progress through regular and determined stages of seral
development that are analogous to the developmental stages of an organism. The Clementsian paradigm
was challenged by Henry Gleason,[255] who stated that 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.[256]

The Clementsian superorganism theory was an overextended application of an idealistic form of


holism.[36][109] The term "holism" was coined in 1926 by Jan Christiaan Smuts, a South African general
and polarizing historical figure who was inspired by Clements' superorganism concept.[257][C] Around the
same time, Charles Elton pioneered the concept of food chains in his classical book Animal Ecology.[84]
Elton[84] defined ecological relations using concepts of food chains, food cycles, and 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.[258] Alfred J. Lotka brought in
many theoretical concepts applying thermodynamic principles to ecology.

In 1942, Raymond Lindeman wrote a landmark paper on the trophic dynamics of ecology, which was
published posthumously after initially being rejected for its theoretical emphasis. Trophic dynamics
became the foundation for much of the work to follow on energy and material flow through ecosystems.
Robert MacArthur advanced mathematical theory, predictions, and tests in ecology in the 1950s, which
inspired a resurgent school of theoretical mathematical ecologists.[239][259][260] Ecology also has
developed through contributions from other nations, including Russia's Vladimir Vernadsky and his
founding of the biosphere concept in the 1920s[261] and Japan's Kinji Imanishi and his concepts of
harmony in nature and habitat segregation in the 1950s.[262] Scientific recognition of contributions to
ecology from non-English-speaking cultures is hampered by language and translation barriers.[261]

Ecology surged in popular and scientific interest during the 1960–


1970s environmental movement. There are strong historical and This whole chain of
scientific ties between ecology, environmental management, and poisoning, then, seems to
protection.[239] The historical emphasis and poetic naturalistic writings rest on a base of minute
advocating the protection of wild places by notable ecologists in the plants which must have
history of conservation biology, such as Aldo Leopold and Arthur been the original
Tansley, have been seen as far removed from urban centres where, it is concentrators. But what of
claimed, the concentration of pollution and environmental degradation the opposite end of the food
is located.[239][264] Palamar (2008)[264] notes an overshadowing by chain—the human being
mainstream environmentalism of pioneering women in the early 1900s who, in probable ignorance
who fought for urban health ecology (then called euthenics)[252] and of all this sequence of
brought about changes in environmental legislation. Women such as events, has rigged his
Ellen Swallow Richards and Julia Lathrop, among others, were fishing tackle, caught a
precursors to the more popularized environmental movements after the string of fish from the
1950s. waters of Clear Lake, and
taken them home to fry for
his supper?
In 1962, marine biologist and ecologist Rachel Carson's book Silent Rachel Carson (1962)[263]: 48
Spring helped to mobilize the environmental movement by alerting the
public to toxic pesticides, such as DDT, bioaccumulating in the
environment. Carson used ecological science to link the release of environmental toxins to human and
ecosystem health. Since then, ecologists have worked to bridge their understanding of the degradation of
the planet's ecosystems with environmental politics, law, restoration, and natural resources
management.[22][239][264][265]

See also
Biology portal
Ecology portal

Plants portal

Trees portal

Water portal

Carrying capacity Human ecology


Chemical ecology Industrial ecology
Climate justice Information ecology
Circles of Sustainability Landscape ecology
Cultural ecology Natural resource
Dialectical naturalism Normative science
Ecological death Philosophy of ecology
Ecological empathy Political ecology
Ecological overshoot Theoretical ecology
Ecological psychology Sensory ecology
Ecology movement Sexecology
Ecosophy Spiritual ecology
Ecopsychology Sustainable development

Lists

Glossary of ecology
Index of biology articles
List of ecologists
Outline of biology
Terminology of ecology

Notes

A. In Ernst Haeckel's (1866) footnote where the term ecology originates, he also gives attribute
to Ancient Greek: χώρας, romanized: khōrā, lit. 'χωρα', meaning "dwelling place, distributional
area" —quoted from Stauffer (1957).
B. This is a copy of Haeckel's original definition (Original: Haeckel, E. (1866) Generelle
Morphologie der Organismen. Allgemeine Grundzige der organischen Formen-
Wissenschaft, mechanisch begriindet durch die von Charles Darwin reformirte Descendenz-
Theorie. 2 vols. Reimer, Berlin.) translated and quoted from Stauffer (1957).
C. Foster & Clark (2008) note how Smut's holism contrasts starkly against his racial political
views as the father of apartheid.
D. First introduced in MacArthur & Wilson's (1967) book of notable mention in the history and
theoretical science of ecology, The Theory of Island Biogeography.
E. Aristotle wrote about this concept in Metaphysics (Quoted from The Internet Classics
Archive (http://classics.mit.edu/Aristotle/metaphysics.mb.txt) translation by W. D. Ross.
Book VIII, Part 6): "To return to the difficulty which has been stated with respect both to
definitions and to numbers, what is the cause of their unity? In the case of all things which
have several parts and in which the totality is not, as it were, a mere heap, but the whole is
something besides the parts, there is a cause; for even in bodies contact is the cause of
unity in some cases and in others viscosity or some other such quality."

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External links
"Ecology" (https://plato.stanford.edu/entries/ecology/) entry by Alkistis Elliott-Graves in the
Stanford Encyclopedia of Philosophy
The Nature Education Knowledge Project: Ecology (https://www.nature.com/scitable/knowle
dge/ecology-102)

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