Ecosystem

From Wikipedia, the free encyclopedia

Coral reefs are a highly productive marine ecosystem.
[1]
Rainforest ecosystems are rich in biodiversity. his is the !ambia River in "enegal#s
$iokolo%&oba $ational 'ark.
(n ecosystem is a community of living organisms )plants, animals and microbes* in
con+unction ,ith the nonliving components of their environment )things like air, ,ater
and mineral soil*, interacting as a system.
[-]
hese biotic and abiotic components are
regarded as linked together through nutrient cycles and energy flo,s.
[.]
(s ecosystems are
defined by the net,ork of interactions among organisms, and bet,een organisms and
their environment,
[/]
they can be of any si0e but usually encompass specific, limited
spaces
[1]
)although some scientists say that the entire planet is an ecosystem*.
[2]
3nergy, ,ater, nitrogen and soil minerals are other essential abiotic components of an
ecosystem. he energy that flo,s through ecosystems is obtained primarily from the sun.
4t generally enters the system through photosynthesis, a process that also captures carbon
from the atmosphere. 5y feeding on plants and on one another, animals play an important
role in the movement of matter and energy through the system. hey also influence the
6uantity of plant and microbial biomass present. 5y breaking do,n dead organic matter,
decomposers release carbon back to the atmosphere and facilitate nutrient cycling by
converting nutrients stored in dead biomass back to a form that can be readily used by
plants and other microbes.
[7]
3cosystems are controlled both by e8ternal and internal factors. 38ternal factors such as
climate, the parent material ,hich forms the soil and topography, control the overall
structure of an ecosystem and the ,ay things ,ork ,ithin it, but are not themselves
influenced by the ecosystem.
[9]
:ther e8ternal factors include time and potential biota.
3cosystems are dynamic entities;invariably, they are sub+ect to periodic disturbances
and are in the process of recovering from some past disturbance.
[<]
3cosystems in similar
environments that are located in different parts of the ,orld can have very different
characteristics simply because they contain different species.
[9]
he introduction of non%
native species can cause substantial shifts in ecosystem function. 4nternal factors not only
control ecosystem processes but are also controlled by them and are often sub+ect to
feedback loops.
[9]
While the resource inputs are generally controlled by e8ternal processes
like climate and parent material, the availability of these resources ,ithin the ecosystem
is controlled by internal factors like decomposition, root competition or shading.
[9]
:ther
internal factors include disturbance, succession and the types of species present. (lthough
humans e8ist and operate ,ithin ecosystems, their cumulative effects are large enough to
influence e8ternal factors like climate.
[9]
5iodiversity affects ecosystem function, as do the processes of disturbance and
succession. 3cosystems provide a variety of goods and services upon ,hich people
depend= the principles of ecosystem management suggest that rather than managing
individual species, natural resources should be managed at the level of the ecosystem
itself. Classifying ecosystems into ecologically homogeneous units is an important step
to,ards effective ecosystem management, but there is no single, agreed%upon ,ay to do
this.
Contents
1 >istory and development
- 3cosystem processes
o -.1 'rimary production
o -.- 3nergy flo,
o -.. ?ecomposition
o -./ $utrient cycling
o -.1 Function and biodiversity
o -.2 3cosystem goods and services
o -.7 3cosystem management
. 3cosystem dynamics
o ..1 3cosystem ecology
/ Classification
o /.1 ypes
1 (nthropogenic threats
2 "ee also
7 $otes
9 References
< @iterature cited
1A 38ternal links
History and development
he term BecosystemB ,as first used in a publication by 5ritish ecology (rthur ansley.
[fn
1][1A]
ansley devised the concept to dra, attention to the importance of transfers of
materials bet,een organisms and their environment.
[11]
>e later refined the term,
describing it as Bhe ,hole system, ... including not only the organism%comple8, but also
the ,hole comple8 of physical factors forming ,hat ,e call the environmentB.
[1-]
ansley
regarded ecosystems not simply as natural units, but as mental isolates.
[1-]
ansley later
[1.]

defined the spatial e8tent of ecosystems using the term ecotope.
!. 3velyn >utchinson, a pioneering limnologist ,ho ,as a contemporary of ansley#s,
combined Charles 3lton#s ideas about trophic ecology ,ith those of Russian geochemist
Cladimir Cernadsky to suggest that mineral nutrient availability in a lake limited algal
production ,hich ,ould, in turn, limit the abundance of animals that feed on algae.
Raymond @indeman took these ideas one step further to suggest that the flo, of energy
through a lake ,as the primary driver of the ecosystem. >utchinson#s students, brothers
>o,ard . :dum and 3ugene '. :dum, further developed a Bsystems approachB to the
study of ecosystems, allo,ing them to study the flo, of energy and material through
ecological systems.
[11]
Ecosystem processes
3nergy and carbon enter ecosystems through photosynthesis, are incorporated into living
tissue, transferred to other organisms that feed on the living and dead plant matter, and
eventually released through respiration.
[1/]
Dost mineral nutrients, on the other hand, are
recycled ,ithin ecosystems.
[11]
3cosystems are controlled both by e8ternal and internal factors. 38ternal factors, also
called state factors, control the overall structure of an ecosystem and the ,ay things ,ork
,ithin it, but are not themselves influenced by the ecosystem. he most important of
these is climate.
[9]
Climate determines the biome in ,hich the ecosystem is embedded.
Rainfall patterns and temperature seasonality determine the amount of ,ater available to
the ecosystem and the supply of energy available )by influencing photosynthesis*.
[9]

'arent material, the underlying geological material that gives rise to soils, determines the
nature of the soils present, and influences the supply of mineral nutrients. opography
also controls ecosystem processes by affecting things like microclimate, soil development
and the movement of ,ater through a system. his may be the difference bet,een the
ecosystem present in ,etland situated in a small depression on the landscape, and one
present on an ad+acent steep hillside.
[9]
:ther e8ternal factors that play an important role in ecosystem functioning include time
and potential biota. 3cosystems are dynamic entities;invariably, they are sub+ect to
periodic disturbances and are in the process of recovering from some past disturbance.
[<]

ime plays a role in the development of soil from bare rock and the recovery of a
community from disturbance.
[9]
"imilarly, the set of organisms that can potentially be
present in an area can also have a ma+or impact on ecosystems. 3cosystems in similar
environments that are located in different parts of the ,orld can end up doing things very
differently simply because they have different pools of species present.
[9]
he introduction
of non%native species can cause substantial shifts in ecosystem function.
Enlike e8ternal factors, internal factors in ecosystems not only control ecosystem
processes, but are also controlled by them. Conse6uently, they are often sub+ect to
feedback loops.
[9]
While the resource inputs are generally controlled by e8ternal processes
like climate and parent material, the availability of these resources ,ithin the ecosystem
is controlled by internal factors like decomposition, root competition or shading.
[9]
:ther
factors like disturbance, succession or the types of species present are also internal
factors. >uman activities are important in almost all ecosystems. (lthough humans e8ist
and operate ,ithin ecosystems, their cumulative effects are large enough to influence
e8ternal factors like climate.
[9]
Primary production
!lobal oceanic and terrestrial phototroph abundance, from "eptember 1<<7 to (ugust
-AAA. (s an estimate of autotroph biomass, it is only a rough indicator of primary
production potential, and not an actual estimate of it. 'rovided by the "eaWiF" 'ro+ect,
$("(F!oddard "pace Flight Center and :R54D(!3.
Dain articleG 'rimary production
'rimary production is the production of organic matter from inorganic carbon sources.
:ver,helmingly, this occurs through photosynthesis. he energy incorporated through
this process supports life on earth, ,hile the carbon makes up much of the organic matter
in living and dead biomass, soil carbon and fossil fuels. 4t also drives the carbon cycle,
,hich influences global climate via the greenhouse effect.
hrough the process of photosynthesis, plants capture energy from light and use it to
combine carbon dio8ide and ,ater to produce carbohydrates and o8ygen. he
photosynthesis carried out by all the plants in an ecosystem is called the gross primary
production )!''*.
[12]
(bout /9H2AI of the !'' is consumed in plant respiration. he
remainder, that portion of !'' that is not used up by respiration, is kno,n as the net
primary production )$''*.
[1/]
otal photosynthesis is limited by a range of environmental
factors. hese include the amount of light available, the amount of leaf area a plant has to
capture light )shading by other plants is a ma+or limitation of photosynthesis*, rate at
,hich carbon dio8ide can be supplied to the chloroplasts to support photosynthesis, the
availability of ,ater, and the availability of suitable temperatures for carrying out
photosynthesis.
[12]
Energy flow
Left: 3nergy flo, diagram of a frog. he frog represents a node in an e8tended food
,eb. he energy ingested is utili0ed for metabolic processes and transformed into
biomass. he energy flo, continues on its path if the frog is ingested by predators,
parasites, or as a decaying carcass in soil. his energy flo, diagram illustrates ho,
energy is lost as it fuels the metabolic process that transforms the energy and nutrients
into biomass.
Right: (n e8panded three link energy food chain )1. plants, -. herbivores, .. carnivores*
illustrating the relationship bet,een food flo, diagrams and energy transformity. he
transformity of energy becomes degraded, dispersed, and diminished from higher 6uality
to lesser 6uantity as the energy ,ithin a food chain flo,s from one trophic species into
another. (bbreviationsG 4Jinput, (Jassimilation, RJrespiration, $EJnot utili0ed,
'Jproduction, 5Jbiomass.
[17]
Dain articleG 3nergy flo, )ecology*
"ee alsoG Food ,eb and rophic level
he carbon and energy incorporated into plant tissues )net primary production* is either
consumed by animals ,hile the plant is alive, or it remains uneaten ,hen the plant tissue
dies and becomes detritus. 4n terrestrial ecosystems, roughly <AI of the $'' ends up
being broken do,n by decomposers. he remainder is either consumed by animals ,hile
still alive and enters the plant%based trophic system, or it is consumed after it has died,
and enters the detritus%based trophic system. 4n a6uatic systems, the proportion of plant
biomass that gets consumed by herbivores is much higher.
[19]
4n trophic systems
photosynthetic organisms are the primary producers. he organisms that consume their
tissues are called primary consumers or secondary producers;herbivores. :rganisms
,hich feed on microbes )bacteria and fungi* are termed microbivores. (nimals that feed
on primary consumers;carnivores;are secondary consumers. 3ach of these constitutes
a trophic level.
[19]
he se6uence of consumption;from plant to herbivore, to carnivore;
forms a food chain. Real systems are much more comple8 than this;organisms ,ill
generally feed on more than one form of food, and may feed at more than one trophic
level. Carnivores may capture some prey ,hich are part of a plant%based trophic system
and others that are part of a detritus%based trophic system )a bird that feeds both on
herbivorous grasshoppers and earth,orms, ,hich consume detritus*. Real systems, ,ith
all these comple8ities, form food ,ebs rather than food chains.
[19]
Decomposition
"ee alsoG ?ecomposition
he carbon and nutrients in dead organic matter are broken do,n by a group of processes
kno,n as decomposition. his releases nutrients that can then be re%used for plant and
microbial production, and returns carbon dio8ide to the atmosphere )or ,ater* ,here it
can be used for photosynthesis. 4n the absence of decomposition, dead organic matter
,ould accumulate in an ecosystem and nutrients and atmospheric carbon dio8ide ,ould
be depleted.
[1<]
(ppro8imately <AI of terrestrial $'' goes directly from plant to
decomposer.
[19]
?ecomposition processes can be separated into three categories;leaching, fragmentation
and chemical alteration of dead material. (s ,ater moves through dead organic matter, it
dissolves and carries ,ith it the ,ater%soluble components. hese are then taken up by
organisms in the soil, react ,ith mineral soil, or are transported beyond the confines of
the ecosystem )and are considered BlostB to it*.
[1<]
$e,ly shed leaves and ne,ly dead
animals have high concentrations of ,ater%soluble components, and include sugars,
amino acids and mineral nutrients. @eaching is more important in ,et environments, and
much less important in dry ones.
[1<]
Fragmentation processes break organic material into smaller pieces, e8posing ne,
surfaces for coloni0ation by microbes. Freshly shed leaf litter may be inaccessible due to
an outer layer of cuticle or bark, and cell contents are protected by a cell ,all. $e,ly
dead animals may be covered by an e8oskeleton. Fragmentation processes, ,hich break
through these protective layers, accelerate the rate of microbial decomposition.
[1<]

(nimals fragment detritus as they hunt for food, as does passage through the gut. Free0e%
tha, cycles and cycles of ,etting and drying also fragment dead material.
[1<]
he chemical alteration of dead organic matter is primarily achieved through bacterial
and fungal action. Fungal hyphae produce en0ymes ,hich can break through the tough
outer structures surrounding dead plant material. hey also produce en0ymes ,hich break
do,n lignin, ,hich allo,s to them access to both cell contents and to the nitrogen in the
lignin. Fungi can transfer carbon and nitrogen through their hyphal net,orks and thus,
unlike bacteria, are not dependent solely on locally available resources.
[1<]
?ecomposition rates vary among ecosystems. he rate of decomposition is governed by
three sets of factors;the physical environment )temperature, moisture and soil
properties*, the 6uantity and 6uality of the dead material available to decomposers, and
the nature of the microbial community itself.
[-A]
emperature controls the rate of
microbial respiration= the higher the temperature, the faster microbial decomposition
occurs. 4t also affects soil moisture, ,hich slo,s microbial gro,th and reduces leaching.
Free0e%tha, cycles also affect decomposition;free0ing temperatures kill soil
microorganisms, ,hich allo,s leaching to play a more important role in moving nutrients
around. his can be especially important as the soil tha,s in the "pring, creating a pulse
of nutrients ,hich become available.
[-A]
?ecomposition rates are lo, under very ,et or very dry conditions. ?ecomposition rates
are highest in ,et, moist conditions ,ith ade6uate levels of o8ygen. Wet soils tend to
become deficient in o8ygen )this is especially true in ,etlands*, ,hich slo,s microbial
gro,th. 4n dry soils, decomposition slo,s as ,ell, but bacteria continue to gro, )albeit at
a slo,er rate* even after soils become too dry to support plant gro,th. When the rains
return and soils become ,et, the osmotic gradient bet,een the bacterial cells and the soil
,ater causes the cells to gain ,ater 6uickly. Ender these conditions, many bacterial cells
burst, releasing a pulse of nutrients.
[-A]
?ecomposition rates also tend to be slo,er in
acidic soils.
[-A]
"oils ,hich are rich in clay minerals tend to have lo,er decomposition
rates, and thus, higher levels of organic matter.
[-A]
he smaller particles of clay result in a
larger surface area that can hold ,ater. he higher the ,ater content of a soil, the lo,er
the o8ygen content
[-1]
and conse6uently, the lo,er the rate of decomposition. Clay
minerals also bind particles of organic material to their surface, making them less
accessibly to microbes.
[-A]
"oil disturbance like tilling increase decomposition by
increasing the amount of o8ygen in the soil and by e8posing ne, organic matter to soil
microbes.
[-A]
he 6uality and 6uantity of the material available to decomposers is another ma+or factor
that influences the rate of decomposition. "ubstances like sugars and amino acids
decompose readily and are considered BlabileB. Cellulose and hemicellulose, ,hich are
broken do,n more slo,ly, are Bmoderately labileB. Compounds ,hich are more resistant
to decay, like lignin or cutin, are considered BrecalcitrantB.
[-A]
@itter ,ith a higher
proportion of labile compounds decomposes much more rapidly than does litter ,ith a
higher proportion of recalcitrant material. Conse6uently, dead animals decompose more
rapidly than dead leaves, ,hich themselves decompose more rapidly than fallen
branches.
[-A]
(s organic material in the soil ages, its 6uality decreases. he more labile
compounds decompose 6uickly, leaving and increasing proportion of recalcitrant
material. Dicrobial cell ,alls also contain a recalcitrant materials like chitin, and these
also accumulate as the microbes die, further reducing the 6uality of older soil organic
matter.
[-A]
Nutrient cycling
"ee alsoG $utrient cycle, 5iogeochemical cycle and $itrogen cycle
5iological nitrogen cycling
3cosystems continually e8change energy and carbon ,ith the ,ider environment=
mineral nutrients, on the other hand, are mostly cycled back and forth bet,een plants,
animals, microbes and the soil. Dost nitrogen enters ecosystems through biological
nitrogen fi8ation, is deposited through precipitation, dust, gases or is applied as fertili0er.
[11]
"ince most terrestrial ecosystems are nitrogen%limited, nitrogen cycling is an
important control on ecosystem production.
[11]
Entil modern times, nitrogen fi8ation ,as the ma+or source of nitrogen for ecosystems.
$itrogen fi8ing bacteria either live symbiotically ,ith plants, or live freely in the soil.
he energetic cost is high for plants ,hich support nitrogen%fi8ing symbionts;as much
as -1I of !'' ,hen measured in controlled conditions. Dany members of the legume
plant family support nitrogen%fi8ing symbionts. "ome cyanobacteria are also capable of
nitrogen fi8ation. hese are phototrophs, ,hich carry out photosynthesis. @ike other
nitrogen%fi8ing bacteria, they can either be free%living or have symbiotic relationships
,ith plants.
[11]
:ther sources of nitrogen include acid deposition produced through the
combustion of fossil fuels, ammonia gas ,hich evaporates from agricultural fields ,hich
have had fertili0ers applied to them, and dust.
[11]
(nthropogenic nitrogen inputs account
for about 9AI of all nitrogen flu8es in ecosystems.
[11]
When plant tissues are shed or are eaten, the nitrogen in those tissues becomes available
to animals and microbes. Dicrobial decomposition releases nitrogen compounds from
dead organic matter in the soil, ,here plants, fungi and bacteria compete for it. "ome soil
bacteria use organic nitrogen%containing compounds as a source of carbon, and release
ammonium ions into the soil. his process is kno,n as nitrogen minerali0ation. :thers
convert ammonium to nitrite and nitrate ions, a process kno,n as nitrification. $itric
o8ide and nitrous o8ide are also produced during nitrification.
[11]
Ender nitrogen%rich and
o8ygen%poor conditions, nitrates and nitrites are converted to nitrogen gas, a process
kno,n as denitrification.
[11]
:ther important nutrients include phosphorus, sulfur, calcium, potassium, magnesium
and manganese.
[--]
'hosphorus enters ecosystems through ,eathering. (s ecosystems age
this supply diminishes, making phosphorus%limitation more common in older landscapes
)especially in the tropics*.
[--]
Calcium and sulfur are also produced by ,eathering, but
acid deposition is an important source of sulfur in many ecosystems. (lthough
magnesium and manganese are produced by ,eathering, e8changes bet,een soil organic
matter and living cells account for a significant portion of ecosystem flu8es. 'otassium is
primarily cycled bet,een living cells and soil organic matter.
[--]
Function and iodiversity
"ee alsoG 5iodiversity and 3cosystem engineer
@och @omond in "cotland forms a relatively isolated ecosystem. he fish community of
this lake has remained stable over a long period until a number of introductions in the
1<7As restructured its food ,eb.
[-.]
"piny forest at 4faty, Dadagascar, featuring various Adansonia )baobab* species,
Alluaudia procera )Dadagascar ocotillo* and other vegetation.
3cosystem processes are broad generali0ations that actually take place through the
actions of individual organisms. he nature of the organisms;the species, functional
groups and trophic levels to ,hich they belong;dictates the sorts of actions these
individuals are capable of carrying out, and the relative efficiency ,ith ,hich they do so.
hus, ecosystem processes are driven by the number of species in an ecosystem, the e8act
nature of each individual species, and the relative abundance organisms ,ithin these
species.
[-/]
5iodiversity plays an important role in ecosystem functioning.
[-1]
3cological theory suggests that in order to coe8ist, species must have some level of
limiting similarity;they must be different from one another in some fundamental ,ay,
other,ise one species ,ould competitively e8clude the other.
[-2]
?espite this, the
cumulative effect of additional species in an ecosystem is not linear;additional species
may enhance nitrogen retention, for e8ample, but beyond some level of species richness,
additional species may have little additive effect.
[-/]
he addition )or loss* of species
,hich are ecologically similar to those already present in an ecosystem tends to only have
a small effect on ecosystem function. 3cologically distinct species, on the other hand,
have a much larger effect. "imilarly, dominant species have a large impact on ecosystem
function, ,hile rare species tend to have a small effect. &eystone species tend to have an
effect on ecosystem function that is disproportionate to their abundance in an ecosystem.
[-/]
Ecosystem goods and services
Dain articlesG 3cosystem services and 3cological goods and services
"ee alsoG 3cosystem valuation and 3cological yield
3cosystems provide a variety of goods and services upon ,hich people depend.
[-7]

3cosystem goods include the Btangible, material productsB
[-9]
of ecosystem processes;
food, construction material, medicinal plants;in addition to less tangible items like
tourism and recreation, and genes from ,ild plants and animals that can be used to
improve domestic species.
[-7]
3cosystem services, on the other hand, are generally
Bimprovements in the condition or location of things of valueB.
[-9]
hese include things
like the maintenance of hydrological cycles, cleaning air and ,ater, the maintenance of
o8ygen in the atmosphere, crop pollination and even things like beauty, inspiration and
opportunities for research.
[-7]
While ecosystem goods have traditionally been recogni0ed
as being the basis for things of economic value, ecosystem services tend to be taken for
granted.
[-9]
While !retchen ?aily#s original definition distinguished bet,een ecosystem
goods and ecosystem services, Robert Costan0a and colleagues# later ,ork and that of the
Dillennium 3cosystem (ssessment lumped all of these together as ecosystem services.
[-9]
Ecosystem management
Dain articleG 3cosystem management
"ee alsoG 3cological economics, "ustainability and "ustainable development
When natural resource management is applied to ,hole ecosystems, rather than single
species, it is termed ecosystem management.
[-<]
( variety of definitions e8istG F. "tuart
Chapin and coauthors define it as Bthe application of ecological science to resource
management to promote long%term sustainability of ecosystems and the delivery of
essential ecosystem goods and servicesB,
[.A]
,hile $orman Christensen and coauthors
defined it as Bmanagement driven by e8plicit goals, e8ecuted by policies, protocols, and
practices, and made adaptable by monitoring and research based on our best
understanding of the ecological interactions and processes necessary to sustain ecosystem
structure and functionB
[-7]
and 'eter 5russard and colleagues defined it as Bmanaging
areas at various scales in such a ,ay that ecosystem services and biological resources are
preserved ,hile appropriate human use and options for livelihood are sustainedB.
[.1]
(lthough definitions of ecosystem management abound, there is a common set of
principles ,hich underlie these definitions.
[.A]
( fundamental principle is the long%term
sustainability of the production of goods and services by the ecosystem=
[.A]

Bintergenerational sustainability [is] a precondition for management, not an afterthoughtB.
[-7]
4t also re6uires clear goals ,ith respect to future tra+ectories and behaviors of the
system being managed. :ther important re6uirements include a sound ecological
understanding of the system, including connectedness, ecological dynamics and the
conte8t in ,hich the system is embedded. :ther important principles include an
understanding of the role of humans as components of the ecosystems and the use of
adaptive management.
[-7]
While ecosystem management can be used as part of a plan for
,ilderness conservation, it can also be used in intensively managed ecosystems
[-7]
)see,
for e8ample, agroecosystem and close to nature forestry*.
Ecosystem dynamics
emperate rainforest on the :lympic 'eninsula in Washington state.
he >igh 'eaks Wilderness (rea in the 2,AAA,AAA%acre )-,/AA,AAA ha* (dirondack 'ark
is an e8ample of a diverse ecosystem.
3cosystems are dynamic entities;invariably, they are sub+ect to periodic disturbances
and are in the process of recovering from some past disturbance.
[<]
When an ecosystem is
sub+ect to some sort of perturbation, it responds by moving a,ay from its initial state.
he tendency of a system to remain close to its e6uilibrium state, despite that disturbance,
is termed its resistance. :n the other hand, the speed ,ith ,hich it returns to its initial
state after disturbance is called its resilience.
[<]
From one year to another, ecosystems e8perience variation in their biotic and abiotic
environments. ( drought, an especially cold ,inter and a pest outbreak all constitute
short%term variability in environmental conditions. (nimal populations vary from year to
year, building up during resource%rich periods and crashing as they overshoot their food
supply. hese changes play out in changes in $'', decomposition rates, and other
ecosystem processes.
[<]
@onger%term changes also shape ecosystem processes;the forests
of eastern $orth (merica still sho, legacies of cultivation ,hich ceased -AA years ago,
,hile methane production in eastern "iberian lakes is controlled by organic matter ,hich
accumulated during the 'leistocene.
[<]
?isturbance also plays an important role in ecological processes. F. "tuart Chapin and
coauthors define disturbance as Ba relatively discrete event in time and space that alters
the structure of populations, communities and ecosystems and causes changes in
resources availability or the physical environmentB.
[.-]
his can range from tree falls and
insect outbreaks to hurricanes and ,ildfires to volcanic eruptions and can cause large
changes in plant, animal and microbe populations, as ,ell soil organic matter content.
[<]

?isturbance is follo,ed by succession, a Bdirectional change in ecosystem structure and
functioning resulting from biotically driven changes in resources supply.B
[.-]
he fre6uency and severity of disturbance determines the ,ay it impacts ecosystem
function. Da+or disturbance like a volcanic eruption or glacial advance and retreat leave
behind soils that lack plants, animals or organic matter. 3cosystems that e8perience
disturbances that undergo primary succession. @ess severe disturbance like forest fires,
hurricanes or cultivation result in secondary succession.
[<]
Dore severe disturbance and
more fre6uent disturbance result in longer recovery times. 3cosystems recover more
6uickly from less severe disturbance events.
[<]
he early stages of primary succession are dominated by species ,ith small propagules
)seed and spores* ,hich can be dispersed long distances. he early coloni0ers;often
algae, cyanobacteria and lichens;stabili0e the substrate. $itrogen supplies are limited in
ne, soils, and nitrogen%fi8ing species tend to play an important role early in primary
succession. Enlike in primary succession, the species that dominate secondary
succession, are usually present from the start of the process, often in the soil seed bank.
4n some systems the successional path,ays are fairly consistent, and thus, are easy to
predict. 4n others, there are many possible path,ays;for e8ample, the introduced
nitrogen%fi8ing legume, Myrica faya, alter successional tra+ectories in >a,aiian forests.
[<]
he theoretical ecologist Robert Elano,ic0 has used information theory tools to describe
the structure of ecosystems, emphasi0ing mutual information )correlations* in studied
systems. ?ra,ing on this methodology and prior observations of comple8 ecosystems,
Elano,ic0 depicts approaches to determining the stress levels on ecosystems and
predicting system reactions to defined types of alteration in their settings )such as
increased or reduced energy flo,, and eutrophication.
[..]
Ecosystem ecology
Dain articleG 3cosystem ecology
"ee alsoG 3cosystem model
( hydrothermal vent is an ecosystem on the ocean floor. )he scale bar is 1 m.*
3cosystem ecology studies Bthe flo, of energy and materials through organisms and the
physical environmentB. 4t seeks to understand the processes ,hich govern the stocks of
material and energy in ecosystems, and the flo, of matter and energy through them. he
study of ecosystems can cover 1A orders of magnitude, from the surface layers of rocks to
the surface of the planet.
[./]
here is no single definition of ,hat constitutes an ecosystem.
[.1]
!erman ecologist 3rnst%
?etlef "chul0e and coauthors defined an ecosystem as an area ,hich is Buniform
regarding the biological turnover, and contains all the flu8es above and belo, the ground
area under consideration.B hey e8plicitly re+ect !ene @ikens# use of entire river
catchments as Btoo ,ide a demarcationB to be a single ecosystem, given the level of
heterogeneity ,ithin such an area.
[.2]
:ther authors have suggested that an ecosystem can
encompass a much larger area, even the ,hole planet.
[2]
"chul0e and coauthors also
re+ected the idea that a single rotting log could be studied as an ecosystem because the
si0e of the flo,s bet,een the log and its surroundings are too large, relative to the
proportion cycles ,ithin the log.
[.2]
'hilosopher of science Dark "agoff considers the
failure to define Bthe kind of ob+ect it studiesB to be an obstacle to the development of
theory in ecosystem ecology.
[.1]
3cosystems can be studied through a variety of approaches;theoretical studies, studies
monitoring specific ecosystems over long periods of time, those that look at differences
bet,een ecosystems to elucidate ho, they ,ork and direct manipulative e8perimentation.
[.7]
"tudies can be carried out at a variety of scales, from microcosms and mesocosms
,hich serve as simplified representations of ecosystems, through ,hole%ecosystem
studies.
[.9]
(merican ecologist "tephen R. Carpenter has argued that microcosm
e8periments can be Birrelevant and diversionaryB if they are not carried out in con+unction
,ith field studies carried out at the ecosystem scale, because microcosm e8periments
often fail to accurately predict ecosystem%level dynamics.
[.<]
he >ubbard 5rook 3cosystem "tudy, established in the White Dountains, $e,
>ampshire in 1<2., ,as the first successful attempt to study an entire ,atershed as an
ecosystem. he study used stream chemistry as a means of monitoring ecosystem
properties, and developed a detailed biogeochemical model of the ecosystem.
[/A]
@ong%
term research at the site led to the discovery of acid rain in $orth (merica in 1<7-, and
,as able to document the conse6uent depletion of soil cations )especially calcium* over
the ne8t several decades.
[/1]
Classification
"ee alsoG 3cosystem diversity, 3coregion, 3cological land classification and 3cotope
Flora of 5a+a California ?esert, CataviKa region, De8ico.
Classifying ecosystems into ecologically homogeneous units is an important step to,ards
effective ecosystem management.
[/-]
( variety of systems e8ist, based on vegetation
cover, remote sensing, and bioclimatic classification systems.
[/-]
(merican geographer
Robert 5ailey defines a hierarchy of ecosystem units ranging from microecosystems
)individual homogeneous sites, on the order of 1A s6uare kilometres )/ s6 mi* in area*,
through mesoecosystems )landscape mosaics, on the order of 1,AAA s6uare kilometres
)/AA s6 mi** to macroecosystems )ecoregions, on the order of 1AA,AAA s6uare kilometres
)/A,AAA s6 mi**.
[/.]
5ailey outlined five different methods for identifying ecosystemsG gestalt )Ba ,hole that
is not derived through considerable of its partsB*, in ,hich regions are recogni0ed and
boundaries dra,n intuitively= a map overlay system ,here different layers like geology,
landforms and soil types are overlain to identify ecosystems= multivariate clustering of
site attributes= digital image processing of remotely sensed data grouping areas based on
their appearance or other spectral properties= or by a Bcontrolling factors methodB ,here a
subset of factors )like soils, climate, vegetation physiognomy or the distribution of plant
or animal species* are selected from a large array of possible ones are used to delineate
ecosystems.
[//]
4n contrast ,ith 5ailey#s methodology, 'uerto Rico ecologist (riel @ugo
and coauthors identified ten characteristics of an effective classification systemG that it be
based on georeferenced, 6uantitative data= that it should minimi0e sub+ectivity and
e8plicitly identify criteria and assumptions= that it should be structured around the factors
that drive ecosystem processes= that it should reflect the hierarchical nature of
ecosystems= that it should be fle8ible enough to conform to the various scales at ,hich
ecosystem management operates= that it should be tied to reliable measures of climate so
that it can Banticipat[e] global climate change= that it be applicable ,orld,ide= that it
should be validated against independent data= that it take into account the sometimes
comple8 relationship bet,een climate, vegetation and ecosystem functioning= and that it
should be able to adapt and improve as ne, data become availableB.
[/-]
!ypes
(6uatic ecosystem
o Darine ecosystem
@arge marine ecosystem
o Fresh,ater ecosystem
@ake ecosystem
River ecosystem
Wetland
errestrial ecosystem
o Forest
o !reater Lello,stone 3cosystem
o @ittoral 0one
o Riparian 0one
o "ubsurface lithoautotrophic microbial ecosystem
o Erban ecosystem
o Dovile Cave
o ?esert
( fresh,ater ecosystem in !ran Canaria, an island of the Canary 4slands.
"nthropogenic threats
"ee alsoG 'lanetary boundaries
(s human populations gro,, so do the resource demands imposed on ecosystems and the
impacts of the human ecological footprint. $atural resources are not invulnerable and
infinitely available. he environmental impacts of anthropogenic actions, ,hich are
processes or materials derived from human activities, are becoming more apparent H air
and ,ater 6uality are increasingly compromised, oceans are being overfished, pests and
diseases are e8tending beyond their historical boundaries, and deforestation is
e8acerbating flooding do,nstream. 4t has been reported that appro8imately /A%1AI of
3arthMs ice%free land surface has been heavily transformed or degraded by anthropogenic
activities, 22I of marine fisheries are either overe8ploited or at their limit, atmospheric
C:- has increased more than .AI since the advent of industriali0ation, and nearly -1I
of 3arthMs bird species have gone e8tinct in the last t,o thousand years.
[/1]
"ociety is
increasingly becoming a,are that ecosystem services are not only limited, but also that
they are threatened by human activities. he need to better consider long%term ecosystem
health and its role in enabling human habitation and economic activity is urgent. o help
inform decision%makers, many ecosystem services are being assigned economic values,
often based on the cost of replacement ,ith anthropogenic alternatives. he ongoing
challenge of prescribing economic value to nature, for e8ample through biodiversity
banking, is prompting transdisciplinary shifts in ho, ,e recogni0e and manage the
environment, social responsibility, business opportunities, and our future as a species.