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Unsourced material may be challenged and removed.<br /><small><span class="plainlinks"><i>Find sources:</i> <a rel="nofollow" class="external text" href="https://www.google.com/search?as_eq=wikipedia&q=%22Alternative+stable+state%22">"Alternative stable state"</a> – <a rel="nofollow" class="external text" href="https://www.google.com/search?tbm=nws&q=%22Alternative+stable+state%22+-wikipedia&tbs=ar:1">news</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?&q=%22Alternative+stable+state%22&tbs=bkt:s&tbm=bks">newspapers</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.google.com/search?tbs=bks:1&q=%22Alternative+stable+state%22+-wikipedia">books</a> <b>·</b> <a rel="nofollow" class="external text" href="https://scholar.google.com/scholar?q=%22Alternative+stable+state%22">scholar</a> <b>·</b> <a rel="nofollow" class="external text" href="https://www.jstor.org/action/doBasicSearch?Query=%22Alternative+stable+state%22&acc=on&wc=on">JSTOR</a></span></small></span> <span class="date-container"><i>(<span class="date">July 2009</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p>In <a href="/wiki/Ecology" title="Ecology">ecology</a>, the theory of <b>alternative stable states</b> (sometimes termed <b>alternate stable states</b> or <b>alternative stable equilibria</b>) predicts that <a href="/wiki/Ecosystems" class="mw-redirect" title="Ecosystems">ecosystems</a> can exist under multiple "states" (sets of unique biotic and abiotic conditions). These alternative states are non-transitory and therefore considered stable over ecologically-relevant timescales. Ecosystems may transition from one stable state to another, in what is known as a state shift (sometimes termed a <a href="/wiki/Phase_(waves)" title="Phase (waves)">phase shift</a> or <a href="/wiki/Regime_shift" title="Regime shift">regime shift</a>), when <a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">perturbed</a>. Due to ecological feedbacks, ecosystems display resistance to state shifts and therefore tend to remain in one state unless perturbations are large enough. Multiple states may persist under equal environmental conditions, a phenomenon known as <a href="/wiki/Hysteresis" title="Hysteresis">hysteresis</a>. <b>Alternative stable state</b> theory suggests that discrete states are separated by <a href="/wiki/Ecological_threshold" title="Ecological threshold">ecological thresholds</a>, in contrast to ecosystems which change smoothly and continuously along an <a href="/wiki/Environmental_gradient" title="Environmental gradient">environmental gradient</a>. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Theory">Theory</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=1" title="Edit section: Theory"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Alternative stable state theory was first proposed by <a href="/wiki/Richard_Lewontin" title="Richard Lewontin">Richard Lewontin</a> (1969), but other early key authors include <a href="/wiki/C._S._Holling" title="C. S. Holling">Holling</a> (1973), Sutherland (1974), <a href="/wiki/R._M._May" class="mw-redirect" title="R. M. May">May</a> (1977), and <a href="/wiki/Marten_Scheffer" title="Marten Scheffer">Scheffer</a> et al. (2001). In the broadest sense, alternative stable state theory proposes that a change in ecosystem conditions can result in an abrupt shift in the state of the ecosystem, such as a change in population (Barange, M. et al. 2008) or community composition. Ecosystems can persist in states that are considered stable (i.e., can exist for relatively long periods of time). Intermediate states are considered unstable and are, therefore, transitory. Because ecosystems are resistant to state shifts, significant perturbations are usually required to overcome ecological thresholds and cause shifts from one stable state to another. The resistance to state shifts is known as "<a href="/wiki/Ecological_resilience" title="Ecological resilience">resilience</a>" (Holling 1973). </p><p>State shifts are often illustrated heuristically by the ball-in-cup model (Holling, C.S. et al., 1995) Biodiversity in the functioning of ecosystems: an ecological synthesis. In Biodiversity Loss, Ecological and Economical Issues (Perrings, C.A. et al., eds), pp. 44–83, Cambridge University Press). A ball, representing the ecosystem, exists on a surface where any point along the surface represents a possible state. In the simplest model, the landscape consists of two valleys separated by a hill. When the ball is in a valley, or a "domain of attraction", it exists in a stable state and must be perturbed to move from this state. In the absence of perturbations, the ball will always roll downhill and therefore will tend to stay in the valley (or stable state). State shifts can be viewed from two different viewpoints, the "community perspective" and the "ecosystem perspective". The ball can only move between stable states in two ways: (1) moving the ball or (2) altering the landscape. The community perspective is analogous to moving the ball, while the ecosystem perspective is analogous to altering the landscape. </p><p>These two viewpoints consider the same phenomenon with different mechanisms. The community perspective considers ecosystem variables (which change relatively quickly and are subject to feedbacks from the system), whereas the ecosystem perspective considers ecosystem parameters (which change relatively slowly and operate independently of the system). The community context considers a relatively constant environment in which multiple stable states are accessible to populations or communities. This definition is an extension of stability analysis of populations (e.g., Lewontin 1969; Sutherland 1973) and communities (e.g., Drake 1991; Law and Morton 1993). The ecosystem context focuses on the effect of exogenic "drivers" on communities or ecosystems (e.g., May 1977; Scheffer et al. 2001; Dent et al. 2002). Both definitions are explored within this article. </p> <div class="mw-heading mw-heading2"><h2 id="Ecological_community">Ecological community</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=2" title="Edit section: Ecological community"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Ecosystems can shift from one state to another via a significant perturbation directly to <a href="/wiki/State_variable" title="State variable">state variables</a>. State variables are quantities that change quickly (in ecologically-relevant time scales) in response to feedbacks from the system (i.e., they are dependent on system feedbacks), such as <a href="/wiki/Population_density" title="Population density">population densities</a>. This perspective requires that different states can exist simultaneously under equal environmental conditions, since the ball moves only in response to a state variable change. </p><p>For example, consider a very simple system with three microbial species. It may be possible for the system to exist under different community structure regimes depending on <a href="/wiki/Initial_value_problem" title="Initial value problem">initial conditions</a> (e.g., population densities or spatial arrangement of individuals) (Kerr et al. 2002). Perhaps under certain initial densities or spatial configurations, one species dominates over all others, while under different initial conditions all species can mutually coexist. Because the different species interact, changes in populations affect one another synergistically to determine community structure. Under both states the environmental conditions are identical. Because the states have resilience, following small perturbations (e.g., changes to <a href="/wiki/Population_size" title="Population size">population size</a>) the community returns to the same configuration while large perturbations may induce a shift to another configuration. </p><p>The community perspective requires the existence of alternative stable states (i.e., more than one valley) before the perturbation, since the landscape is not changing. Because communities have some level of <a href="/wiki/Change_management" title="Change management">resistance to change</a>, they will stay in their domain of attraction (or stable state) until the perturbation is large enough to force the system into another state. In the ball-and-cup model, this would be the energy required to push the ball up and over a hill, where it would fall downhill into a different valley. </p> <div class="mw-heading mw-heading2"><h2 id="Ecosystem">Ecosystem</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=3" title="Edit section: Ecosystem"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>It is also possible to cause state shifts in another context, by indirectly affecting <a href="/wiki/State_variable" title="State variable">state variables</a>. This is known as the ecosystem perspective. This perspective requires a change in environmental parameters that affect the behavior of state variables. For example, <a href="/wiki/Birth_rate" title="Birth rate">birth rate</a>, <a href="/wiki/Mortality_rate" title="Mortality rate">death rate</a>, migration, and density-dependent predation indirectly alter the ecosystem state by changing population density (a state variable). Ecosystem parameters are quantities that are unresponsive (or respond very slowly) to feedbacks from the system (i.e., they are independent of system feedbacks). The stable state landscape is changed by environmental drivers, which may result in a change in the quantity of stable states and the relationship between states. </p><p>By the ecosystem perspective, the landscape of the ecological states is changed, which forces a change in the ecosystem state. Changing the landscape can modify the number, location, and resilience of stable states, as well as the unstable intermediate states. By this view, the topography in the ball-and-cup model is not static, as it is in the community perspective. This is a fundamental difference between the two perspectives. </p><p>Although the mechanisms of community and ecosystem perspectives are different, the <a href="/wiki/Empirical_research" title="Empirical research">empirical evidence</a> required for documentation of alternative stable states is the same. In addition, state shifts are often a combination of internal processes and external forces (Scheffer et al. 2001). For example, consider a stream-fed lake in which the pristine state is dominated by benthic vegetation. When upstream construction releases soils into the stream, the system becomes turbid. As a result, benthic vegetation cannot receive light and decline, increasing nutrient availability and allowing <a href="/wiki/Phytoplankton" title="Phytoplankton">phytoplankton</a> to dominate. In this state shift scenario the state variables changing are the populations of benthic vegetation and phytoplankton, and the ecosystem parameters are turbidity and nutrient levels. So, whether identifying mechanisms of variables or parameters is a matter of formulation (Beisner et al. 2003). </p> <div class="mw-heading mw-heading2"><h2 id="Hysteresis_effects">Hysteresis effects</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=4" title="Edit section: Hysteresis effects"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Hysteresis" title="Hysteresis">Hysteresis</a> is an important concept in alternative stable state theory. In this ecological context, hysteresis refers to the existence of different stable states under the same variables or parameters. Hysteresis can be explained by "path-dependency", in which the <a href="/wiki/Equilibrium_point" class="mw-redirect" title="Equilibrium point">equilibrium point</a> for the trajectory of "A → B" is different from for "B → A". In other words, it matters which way the ball is moving across the landscape. Some ecologists (e.g., Scheffer et al. 2001) argue that hysteresis is a prerequisite for the existence of alternative stable states. Others (e.g., Beisner et al. 2003) claim that this is not so; although shifts often involve hysteresis, a system can show alternative stable states yet have equal paths for "A → B" and "B → A". </p><p>Hysteresis can occur via changes to variables or parameters. When variables are changed the ball is pushed from one domain of attraction to another, yet the same push from the other direction cannot return the ball to the original domain of attraction. When parameters are changed a modification to the landscape results in a state shift, but reversing the modification does not result in a reciprocal shift. </p><p>A real-world example of hysteresis is helpful to illustrate the concept. <a href="/wiki/Coral_reef" title="Coral reef">Coral reef</a> systems can dramatically shift from pristine coral-dominated systems to degraded <a href="/wiki/Algae" title="Algae">algae</a>-dominated systems when populations grazing on algae decline. The 1983 crash of <a href="/wiki/Sea_urchin" title="Sea urchin">sea urchin</a> populations in <a href="/wiki/Caribbean" title="Caribbean">Caribbean</a> reef systems released algae from top-down (<a href="/wiki/Herbivory" class="mw-redirect" title="Herbivory">herbivory</a>) control, allowing them to overgrow corals and resulting in a shift to a <a href="/wiki/Environmental_degradation" title="Environmental degradation">degraded</a> state. When urchins rebounded, the high (pre-crash) coral cover levels did not return, indicating hysteresis (Mumby et al. 2007). </p><p>In some cases, state shifts under hysteresis may be irreversible. For example, tropical <a href="/wiki/Cloud_forest" title="Cloud forest">cloud forests</a> require high moisture levels, provided by clouds that are intercepted by the canopy (via condensation). When <a href="/wiki/Deforested" class="mw-redirect" title="Deforested">deforested</a>, moisture delivery ceases. Therefore, reforestation is often unsuccessful because conditions are too dry to allow the trees to grow (<a href="#CITEREFWilsonAgnew1992">Wilson & Agnew 1992</a>). Even in cases where there is no obvious barrier to recovery, alternative states can be remarkably persistent: an experimental grassland heavily fertilized for 10 years lost much of its biodiversity, and was still in this state 20 years later (<a href="#CITEREFIsbellTilmanPolaskyBinder2013">Isbell et al. 2013</a>). </p> <div class="mw-heading mw-heading2"><h2 id="Resilience">Resilience</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=5" title="Edit section: Resilience"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>By their very nature, basins of attraction display <a href="/wiki/Ecological_resilience" title="Ecological resilience">resilience</a>. Ecosystems are resistant to state shifts – they will only undergo shifts under substantial perturbations – but some states are more resilient than others. In the ball-and-cup model, a valley with steep sides has greater resilience than a shallow valley, since it would take more force to push the ball up the hill and out of the valley. </p><p>Resilience can change in stable states when environmental parameters are shifted. Often, humans influence stable states by reducing the resilience of basins of attraction. There are at least three ways in which anthropogenic forces reduce resilience (Folke et al. 2004): (1) Decreasing diversity and <a href="/wiki/Functional_group" title="Functional group">functional groups</a>, often by top-down effects (e.g., overfishing); (2) altering the physico-chemical environment (e.g., <a href="/wiki/Climate_change" title="Climate change">climate change</a>, <a href="/wiki/Pollution" title="Pollution">pollution</a>, <a href="/wiki/Fertilizer" title="Fertilizer">fertilization</a>); or (3) modifying disturbance regimes to which organisms are adapted (e.g., <a href="/wiki/Bottom_trawling" title="Bottom trawling">bottom trawling</a>, <a href="/wiki/Coral_mining" class="mw-redirect" title="Coral mining">coral mining</a>, etc.). When the resilience is decreased, ecosystems can be pushed into alternative, and often less-desirable, stable states with only minor perturbations. When hysteresis effects are present, the return to a more-desirable state is sometimes impossible or impractical (given management constraints). Shifts to less-desirable states often entail a loss of <a href="/wiki/Ecosystem_service" title="Ecosystem service">ecosystem service</a> and function, and have been documented in an array of terrestrial, marine, and freshwater environments (reviewed in Folke et al. 2004). </p> <div class="mw-heading mw-heading2"><h2 id="Evidence">Evidence</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=6" title="Edit section: Evidence"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most work on alternative stable states has been theoretical, using <a href="/wiki/Mathematical_model" title="Mathematical model">mathematical models</a> and simulations to test ecological hypotheses. Other work has been conducted using empirical evidence from surveying, <a href="/wiki/History" title="History">historical records</a>, or comparisons across <a href="/wiki/Spatial_scale" title="Spatial scale">spatial scales</a>. There has been a lack of direct, manipulative experimental tests for alternative stable states. This is especially true for studies outside of controlled laboratory conditions, where state shifts have been documented for cultures of <a href="/wiki/Microorganism" title="Microorganism">microorganisms</a>. </p><p>Verifying the existence of alternative stable states carries profound implications for <a href="/wiki/Ecosystem_management" title="Ecosystem management">ecosystem management</a>. If stable states exist, gradual changes in <a href="/wiki/Environmental_factor" title="Environmental factor">environmental factors</a> may have little effect on a system until a threshold is reached, at which point a catastrophic state shift may occur. Understanding the nature of these thresholds will help inform the design of monitoring programs, ecosystem restoration, and other management decisions. Managers are particularly interested in the potential of hysteresis, since it may be difficult to recover from a state shift (Beisner et al. 2003). The mechanisms of <a href="/wiki/Feedback" title="Feedback">feedback loops</a> that maintain stable states are important to understand if we hope to effectively manage an ecosystem with alternative stable states. </p> <div class="mw-heading mw-heading3"><h3 id="Empirical_evidence">Empirical evidence</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=7" title="Edit section: Empirical evidence"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Empirical evidence for the existence of alternative stable states is vital to advancing the idea beyond theory. Schröder et al. (2005) reviewed the current ecological literature for alternative stable states and found 35 direct experiments, of which only 21 were deemed valid. Of these, 62% (14) showed evidence for and 38% (8) showed no evidence for alternative stable states. However, the Schröder et al. (2005) analysis required evidence of hysteresis, which is not necessarily a prerequisite for alternative stable states. Other authors (e.g., Scheffer et al. 2001; Folke et al. 2004) have had less-stringent requirements for the documentation of alternative stable states. </p><p>State shifts via the community perspective have been induced experimentally by the addition or removal of predators, such as in Paine's (1966) work on <a href="/wiki/Keystone_predator" class="mw-redirect" title="Keystone predator">keystone predators</a> (i.e., predators with disproportionate influence on community structure) in the <a href="/wiki/Intertidal_zone" title="Intertidal zone">intertidal zone</a> (although this claim is refuted by Schröder et al. 2005). Also, Beisner et al. (2003) suggest that commercially exploited fish populations can be forced between alternative stable states by <a href="/wiki/Fishing_pressure" class="mw-redirect" title="Fishing pressure">fishing pressure</a> due to <a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a> that work at very low population sizes. Once a fish population falls below a certain threshold, it will inevitably go extinct when low population densities make replacement of adults impossible due to, for example, the inability to find mates or density-dependent <a href="/wiki/Mortality_rate" title="Mortality rate">mortality</a>. Since populations cannot return from extinction, this is an example of an irreversible state shift. </p><p>Although alternative stable state theory is still in its infancy, empirical evidence has been collected from a variety of <a href="/wiki/Biome" title="Biome">biomes</a>: </p> <ul><li><a href="/wiki/Sahara" title="Sahara">The Sahara</a> region, once vegetated, suddenly collapsed into a desert about 5,000 to <a href="/wiki/4th_millennium_BC" title="4th millennium BC">6,000 years ago</a> due to climate change (Scheffer and Carpenter 2003). Recent alteration of disturbance states via <a href="/wiki/Fire_fighting" class="mw-redirect" title="Fire fighting">fire suppression</a> has caused state shifts in fire-adapted ecosystems of the <a href="/wiki/Southeastern_United_States" title="Southeastern United States">southeastern United States</a> (<a href="#CITEREFPeterson2002">Peterson 2002</a>).</li> <li>In the <a href="/wiki/Northeastern_United_States" title="Northeastern United States">northeastern United States</a> alternative stable states have been experimentally documented in <a href="/wiki/Temperate_hardwood_forest" class="mw-redirect" title="Temperate hardwood forest">hardwood forest</a> <a href="/wiki/Food_chain" title="Food chain">food webs</a> (Schmitz 2004). Spiders exhibit top-down control on generalist herbivore grasshoppers, causing them to hide in and forage on a competitively dominant plant species. Herbivory on the superior plant competitor reduces dominance, thereby increasing plant diversity. Under experimental removal of spiders, grasshoppers were released from predation and consumed plants without preference for the competitively-superior species. This top competitor outcompeted other species, resulting in lower overall plant diversity. Furthermore, reintroductions of spiders failed to return the system to high plant diversity, indicating ecological thresholds.</li> <li>Ponds, lakes, and other small <a href="/wiki/Body_of_water" title="Body of water">bodies of water</a> are ideal for studying alternative stable states because they are relatively self-contained (Holling 1973). There has been great interest in state shifts in shallow temperate lakes between clear-water and turbid-water states. Nutrient loading can shift clear lakes to turbid systems by causing phytoplankton blooms (Scheffer 1997). Feedbacks maintaining the turbid state include shading out benthic macrophytes, which otherwise stabilize sediments and reduce rapid <a href="/wiki/Nutrient_cycling" class="mw-redirect" title="Nutrient cycling">nutrient cycling</a>. Hysteresis may occur if reductions in nutrient input do not return the clear-water state.</li> <li>Alternative stable states have also been shown for freshwater faunal communities. Initial conditions, such as densities of snails, can influence the movement of an ecosystem into one of several alternative stable equilibria (Chase 2003).</li> <li>Additionally, alternative stable states have been documented experimentally in coastal <a href="/wiki/Marine_ecosystem" title="Marine ecosystem">marine ecosystems</a>. Arctic <a href="/wiki/Salt_marshes" class="mw-redirect" title="Salt marshes">salt marshes</a> of <a href="/wiki/Hudson_Bay" title="Hudson Bay">Hudson Bay</a>, Manitoba have been shown to shift from a vegetated state to an unvegetated state due to <a href="/wiki/Overgrazing" title="Overgrazing">overgrazing</a> by <a href="/wiki/Snow_goose" title="Snow goose">snow geese</a> (<a href="#CITEREFHandaHarmsenJefferies2002">Handa, Harmsen & Jefferies 2002</a>). When geese were experimentally removed, recovery of the system only occurred under ideal soil conditions, indicating hysteresis.</li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Alternative_stable_state&action=edit&section=8" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><style data-mw-deduplicate="TemplateStyles:r1238218222">.mw-parser-output cite.citation{font-style:inherit;word-wrap:break-word}.mw-parser-output .citation q{quotes:"\"""\"""'""'"}.mw-parser-output .citation:target{background-color:rgba(0,127,255,0.133)}.mw-parser-output .id-lock-free.id-lock-free a{background:url("//upload.wikimedia.org/wikipedia/commons/6/65/Lock-green.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-limited.id-lock-limited a,.mw-parser-output .id-lock-registration.id-lock-registration a{background:url("//upload.wikimedia.org/wikipedia/commons/d/d6/Lock-gray-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .id-lock-subscription.id-lock-subscription a{background:url("//upload.wikimedia.org/wikipedia/commons/a/aa/Lock-red-alt-2.svg")right 0.1em center/9px no-repeat}.mw-parser-output .cs1-ws-icon a{background:url("//upload.wikimedia.org/wikipedia/commons/4/4c/Wikisource-logo.svg")right 0.1em center/12px no-repeat}body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-free a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-limited a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-registration a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .id-lock-subscription a,body:not(.skin-timeless):not(.skin-minerva) .mw-parser-output .cs1-ws-icon a{background-size:contain;padding:0 1em 0 0}.mw-parser-output .cs1-code{color:inherit;background:inherit;border:none;padding:inherit}.mw-parser-output .cs1-hidden-error{display:none;color:var(--color-error,#d33)}.mw-parser-output .cs1-visible-error{color:var(--color-error,#d33)}.mw-parser-output .cs1-maint{display:none;color:#085;margin-left:0.3em}.mw-parser-output .cs1-kern-left{padding-left:0.2em}.mw-parser-output .cs1-kern-right{padding-right:0.2em}.mw-parser-output .citation .mw-selflink{font-weight:inherit}@media screen{.mw-parser-output .cs1-format{font-size:95%}html.skin-theme-clientpref-night .mw-parser-output .cs1-maint{color:#18911f}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .cs1-maint{color:#18911f}}</style><cite id="CITEREFBarangeBeaugrandHarrisPerry2008" class="citation journal cs1">Barange, M.; Beaugrand, G.; Harris, R.; Perry, R.; Perry, M.; Werner, F. 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(2002). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1693037">"Complex numerical responses to top-down and bottom-up processes in vertebrate populations"</a>. <i>Philosophical Transactions of the Royal Society B</i>. <b>357</b> (1425): 1221–1231. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frstb.2002.1123">10.1098/rstb.2002.1123</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1693037">1693037</a></span>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12396514">12396514</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Philosophical+Transactions+of+the+Royal+Society+B&rft.atitle=Complex+numerical+responses+to+top-down+and+bottom-up+processes+in+vertebrate+populations&rft.volume=357&rft.issue=1425&rft.pages=1221-1231&rft.date=2002&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1693037%23id-name%3DPMC&rft_id=info%3Apmid%2F12396514&rft_id=info%3Adoi%2F10.1098%2Frstb.2002.1123&rft.aulast=Sinclaire&rft.aufirst=A.+R.+E.&rft.au=Krebs%2C+C.+J.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1693037&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAlternative+stable+state" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSutherland1974" class="citation journal cs1">Sutherland, J. 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"Multiple stable points in natural communities". <i>The American Naturalist</i>. <b>108</b> (964): 859–873. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1086%2F282961">10.1086/282961</a>. <a href="/wiki/JSTOR_(identifier)" class="mw-redirect" title="JSTOR (identifier)">JSTOR</a> <a rel="nofollow" class="external text" href="https://www.jstor.org/stable/2459615">2459615</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:85014132">85014132</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+American+Naturalist&rft.atitle=Multiple+stable+points+in+natural+communities&rft.volume=108&rft.issue=964&rft.pages=859-873&rft.date=1974&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A85014132%23id-name%3DS2CID&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F2459615%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.1086%2F282961&rft.aulast=Sutherland&rft.aufirst=J.+P.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAlternative+stable+state" class="Z3988"></span></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWilsonAgnew1992" class="citation book cs1">Wilson, J. Bastow; Agnew, Andrew D.Q. (1992). "Positive-feedback Switches in Plant Communities". <i>Advances in Ecological Research Volume 23</i>. Vol. 23. pp. 263–336. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0065-2504%2808%2960149-X">10.1016/S0065-2504(08)60149-X</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-013923-1" title="Special:BookSources/978-0-12-013923-1"><bdi>978-0-12-013923-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Positive-feedback+Switches+in+Plant+Communities&rft.btitle=Advances+in+Ecological+Research+Volume+23&rft.pages=263-336&rft.date=1992&rft_id=info%3Adoi%2F10.1016%2FS0065-2504%2808%2960149-X&rft.isbn=978-0-12-013923-1&rft.aulast=Wilson&rft.aufirst=J.+Bastow&rft.au=Agnew%2C+Andrew+D.Q.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAlternative+stable+state" class="Z3988"></span></li></ul> </div> <div 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title="Ecosystem model">Modelling ecosystems</a>: <a href="/wiki/Trophic_level" title="Trophic level">Trophic</a> components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">General</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiotic_component" title="Abiotic component">Abiotic component</a></li> <li><a href="/wiki/Abiotic_stress" title="Abiotic stress">Abiotic stress</a></li> <li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behaviour</a></li> <li><a href="/wiki/Biogeochemical_cycle" title="Biogeochemical cycle">Biogeochemical cycle</a></li> <li><a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li> <li><a href="/wiki/Biotic_component" class="mw-redirect" title="Biotic component">Biotic component</a></li> <li><a href="/wiki/Biotic_stress" title="Biotic stress">Biotic stress</a></li> <li><a href="/wiki/Carrying_capacity" title="Carrying capacity">Carrying capacity</a></li> <li><a href="/wiki/Competition_(biology)" title="Competition (biology)">Competition</a></li> <li><a href="/wiki/Ecosystem" title="Ecosystem">Ecosystem</a></li> <li><a href="/wiki/Ecosystem_ecology" title="Ecosystem ecology">Ecosystem ecology</a></li> <li><a href="/wiki/Ecosystem_model" title="Ecosystem model">Ecosystem model</a></li> <li><a href="/wiki/Green_world_hypothesis" title="Green world hypothesis">Green world hypothesis</a></li> <li><a href="/wiki/Keystone_species" title="Keystone species">Keystone species</a></li> <li><a href="/wiki/List_of_feeding_behaviours" title="List of feeding behaviours">List of feeding behaviours</a></li> <li><a href="/wiki/Metabolic_theory_of_ecology" title="Metabolic theory of ecology">Metabolic theory of ecology</a></li> <li><a href="/wiki/Productivity_(ecology)" title="Productivity (ecology)">Productivity</a></li> <li><a href="/wiki/Resource_(biology)" title="Resource (biology)">Resource</a></li> <li><a href="/wiki/Restoration_ecology" class="mw-redirect" title="Restoration ecology">Restoration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Autotroph" title="Autotroph">Producers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Autotroph" title="Autotroph">Autotrophs</a></li> <li><a href="/wiki/Chemosynthesis" title="Chemosynthesis">Chemosynthesis</a></li> <li><a href="/wiki/Chemotroph" title="Chemotroph">Chemotrophs</a></li> <li><a href="/wiki/Foundation_species" title="Foundation species">Foundation species</a></li> <li><a href="/wiki/Kinetotroph" title="Kinetotroph">Kinetotrophs</a></li> <li><a href="/wiki/Mixotroph" title="Mixotroph">Mixotrophs</a></li> <li><a href="/wiki/Myco-heterotrophy" title="Myco-heterotrophy">Myco-heterotrophy</a></li> <li><a href="/wiki/Mycotroph" title="Mycotroph">Mycotroph</a></li> <li><a href="/wiki/Organotroph" title="Organotroph">Organotrophs</a></li> <li><a href="/wiki/Photoheterotroph" title="Photoheterotroph">Photoheterotrophs</a></li> <li><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a></li> <li><a href="/wiki/Photosynthetic_efficiency" title="Photosynthetic efficiency">Photosynthetic efficiency</a></li> <li><a href="/wiki/Phototroph" title="Phototroph">Phototrophs</a></li> <li><a href="/wiki/Primary_nutritional_groups" title="Primary nutritional groups">Primary nutritional groups</a></li> <li><a href="/wiki/Primary_production" title="Primary production">Primary production</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Consumer_(food_chain)" title="Consumer (food chain)">Consumers</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Apex_predator" title="Apex predator">Apex predator</a></li> <li><a href="/wiki/Bacterivore" title="Bacterivore">Bacterivore</a></li> <li><a href="/wiki/Carnivore" title="Carnivore">Carnivores</a></li> <li><a href="/wiki/Chemoorganotroph" class="mw-redirect" title="Chemoorganotroph">Chemoorganotroph</a></li> <li><a href="/wiki/Foraging" title="Foraging">Foraging</a></li> <li><a href="/wiki/Generalist_and_specialist_species" title="Generalist and specialist species">Generalist and specialist species</a></li> <li><a href="/wiki/Intraguild_predation" title="Intraguild predation">Intraguild predation</a></li> <li><a href="/wiki/Herbivore" title="Herbivore">Herbivores</a></li> <li><a href="/wiki/Heterotroph" title="Heterotroph">Heterotroph</a></li> <li><a href="/wiki/Heterotrophic_nutrition" title="Heterotrophic nutrition">Heterotrophic nutrition</a></li> <li><a href="/wiki/Insectivore" title="Insectivore">Insectivore</a></li> <li><a href="/wiki/Mesopredator" title="Mesopredator">Mesopredators</a></li> <li><a href="/wiki/Mesopredator_release_hypothesis" title="Mesopredator release hypothesis">Mesopredator release hypothesis</a></li> <li><a href="/wiki/Omnivore" title="Omnivore">Omnivores</a></li> <li><a href="/wiki/Optimal_foraging_theory" title="Optimal foraging theory">Optimal foraging theory</a></li> <li><a href="/wiki/Planktivore" title="Planktivore">Planktivore</a></li> <li><a href="/wiki/Predation" title="Predation">Predation</a></li> <li><a href="/wiki/Prey_switching" title="Prey switching">Prey switching</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Decomposer" title="Decomposer">Decomposers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chemoorganoheterotrophy" class="mw-redirect" title="Chemoorganoheterotrophy">Chemoorganoheterotrophy</a></li> <li><a href="/wiki/Decomposition" title="Decomposition">Decomposition</a></li> <li><a href="/wiki/Detritivore" title="Detritivore">Detritivores</a></li> <li><a href="/wiki/Detritus" title="Detritus">Detritus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Microorganism#Habitats_and_ecology" title="Microorganism">Microorganisms</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteriophage" title="Bacteriophage">Bacteriophage</a></li> <li><a href="/wiki/Lithoautotroph" title="Lithoautotroph">Lithoautotroph</a></li> <li><a href="/wiki/Lithotroph" title="Lithotroph">Lithotrophy</a></li> <li><a href="/wiki/Marine_microorganisms" title="Marine microorganisms">Marine</a></li> <li><a href="/wiki/Microbial_cooperation" title="Microbial cooperation">Microbial cooperation</a></li> <li><a href="/wiki/Microbial_ecology" title="Microbial ecology">Microbial ecology</a></li> <li><a href="/wiki/Microbial_food_web" title="Microbial food web">Microbial food web</a></li> <li><a href="/wiki/Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li> <li><a href="/wiki/Microbial_loop" title="Microbial loop">Microbial loop</a></li> <li><a href="/wiki/Microbial_mat" title="Microbial mat">Microbial mat</a></li> <li><a href="/wiki/Microbial_metabolism" title="Microbial metabolism">Microbial metabolism</a></li> <li><a href="/wiki/Phage_ecology" title="Phage ecology">Phage ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Food_web" title="Food web">Food webs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biomagnification" title="Biomagnification">Biomagnification</a></li> <li><a href="/wiki/Ecological_efficiency" title="Ecological efficiency">Ecological efficiency</a></li> <li><a href="/wiki/Ecological_pyramid" title="Ecological pyramid">Ecological pyramid</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Energy flow</a></li> <li><a href="/wiki/Food_chain" title="Food chain">Food chain</a></li> <li><a href="/wiki/Trophic_level" title="Trophic level">Trophic level</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Example webs</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lake_ecosystem#Trophic_relationships" title="Lake ecosystem">Lakes</a></li> <li><a href="/wiki/River_ecosystem#Trophic_relationships" title="River ecosystem">Rivers</a></li> <li><a href="/wiki/Soil_food_web" title="Soil food web">Soil</a></li> <li><a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></li> <li><a href="/wiki/Marine_food_web" title="Marine food web">Marine food webs</a> <ul><li><a href="/wiki/Cold_seep" title="Cold seep">cold seeps</a></li> <li><a href="/wiki/Hydrothermal_vent#Biological_communities" title="Hydrothermal vent">hydrothermal vents</a></li> <li><a href="/wiki/Intertidal_ecology" title="Intertidal ecology">intertidal</a></li> <li><a href="/wiki/Kelp_forest#Trophic_ecology" title="Kelp forest">kelp forests</a></li> <li><a href="/wiki/North_Pacific_Gyre" title="North Pacific Gyre">North Pacific Gyre</a></li> <li><a href="/wiki/Ecology_of_the_San_Francisco_Estuary#Food_web" title="Ecology of the San Francisco Estuary">San Francisco Estuary</a></li> <li><a href="/wiki/Tide_pool" title="Tide pool">tide pool</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Processes</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ascendency" title="Ascendency">Ascendency</a></li> <li><a href="/wiki/Bioaccumulation" title="Bioaccumulation">Bioaccumulation</a></li> <li><a href="/wiki/Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li> <li><a href="/wiki/Climax_community" title="Climax community">Climax community</a></li> <li><a href="/wiki/Competitive_exclusion_principle" title="Competitive exclusion principle">Competitive exclusion principle</a></li> <li><a href="/wiki/Consumer%E2%80%93resource_interactions" title="Consumer–resource interactions">Consumer–resource interactions</a></li> <li><a href="/wiki/Copiotroph" title="Copiotroph">Copiotrophs</a></li> <li><a href="/wiki/Dominance_(ecology)" title="Dominance (ecology)">Dominance</a></li> <li><a href="/wiki/Ecological_network" title="Ecological network">Ecological network</a></li> <li><a href="/wiki/Ecological_succession" title="Ecological succession">Ecological succession</a></li> <li><a href="/wiki/Energy_quality" title="Energy quality">Energy quality</a></li> <li><a href="/wiki/Energy_systems_language" title="Energy systems language">Energy systems language</a></li> <li><a href="/wiki/F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li> <li><a href="/wiki/Feed_conversion_ratio" title="Feed conversion ratio">Feed conversion ratio</a></li> <li><a href="/wiki/Feeding_frenzy" title="Feeding frenzy">Feeding frenzy</a></li> <li><a href="/wiki/Mesotrophic_soil" title="Mesotrophic soil">Mesotrophic soil</a></li> <li><a href="/wiki/Nutrient_cycle" title="Nutrient cycle">Nutrient cycle</a></li> <li><a href="/wiki/Oligotroph" title="Oligotroph">Oligotroph</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li> <li><a href="/wiki/Trophic_cascade" title="Trophic cascade">Trophic cascade</a></li> <li><a href="/wiki/Trophic_mutualism" title="Trophic mutualism">Trophic mutualism</a></li> <li><a href="/wiki/Trophic_state_index" title="Trophic state index">Trophic state index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Defense,<br />counter</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptations</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Herbivore_adaptations_to_plant_defense" title="Herbivore adaptations to plant defense">Herbivore adaptations to plant defense</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Plant_defense_against_herbivory" title="Plant defense against herbivory">Plant defense against herbivory</a></li> <li><a href="/wiki/Shoaling_and_schooling#Predator_avoidance" title="Shoaling and schooling">Predator avoidance in schooling fish</a></li></ul> </div></td></tr></tbody></table></div><div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Other_components" style="padding:3px"><table class="nowraplinks hlist mw-collapsible uncollapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Modelling_ecosystems" title="Template:Modelling ecosystems"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Modelling_ecosystems" title="Template talk:Modelling ecosystems"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Modelling_ecosystems" title="Special:EditPage/Template:Modelling ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ecology:_Modelling_ecosystems:_Other_components" style="font-size:114%;margin:0 4em"><a href="/wiki/Ecology" title="Ecology">Ecology</a>: <a href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: Other components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Population_ecology" title="Population ecology">Population<br />ecology</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abundance_(ecology)" title="Abundance (ecology)">Abundance</a></li> <li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Consumer-resource_model" title="Consumer-resource model">Consumer-resource model</a></li> <li><a href="/wiki/Depensation" title="Depensation">Depensation</a></li> <li><a href="/wiki/Ecological_yield" title="Ecological yield">Ecological yield</a></li> <li><a href="/wiki/Effective_population_size" title="Effective population size">Effective population size</a></li> <li><a href="/wiki/Intraspecific_competition" title="Intraspecific competition">Intraspecific competition</a></li> <li><a href="/wiki/Logistic_function" title="Logistic function">Logistic function</a></li> <li><a href="/wiki/Malthusian_growth_model" title="Malthusian growth model">Malthusian growth model</a></li> <li><a href="/wiki/Maximum_sustainable_yield" title="Maximum sustainable yield">Maximum sustainable yield</a></li> <li><a href="/wiki/Overpopulation" title="Overpopulation">Overpopulation</a></li> <li><a href="/wiki/Overexploitation" title="Overexploitation">Overexploitation</a></li> <li><a href="/wiki/Population_cycle" title="Population cycle">Population cycle</a></li> <li><a href="/wiki/Population_dynamics" title="Population dynamics">Population dynamics</a></li> <li><a href="/wiki/Population_model" title="Population model">Population modeling</a></li> <li><a href="/wiki/Population_size" title="Population size">Population size</a></li> <li><a href="/wiki/Lotka%E2%80%93Volterra_equations" title="Lotka–Volterra equations">Predator–prey (Lotka–Volterra) equations</a></li> <li><a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">Recruitment</a></li> <li><a href="/wiki/Small_population_size" title="Small population size">Small population size</a></li> <li><a href="/wiki/Ecological_stability" title="Ecological stability">Stability</a> <ul><li><a href="/wiki/Ecological_resilience" title="Ecological resilience">Resilience</a></li> <li><a href="/wiki/Resistance_(ecology)" title="Resistance (ecology)">Resistance</a></li></ul></li> <li><a href="/wiki/Random_generalized_Lotka%E2%80%93Volterra_model" title="Random generalized Lotka–Volterra model">Random generalized Lotka–Volterra model</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Density_dependence" title="Density dependence">Density-dependent inhibition</a></li> <li><a href="/wiki/Ecological_effects_of_biodiversity" title="Ecological effects of biodiversity">Ecological effects of biodiversity</a></li> <li><a href="/wiki/Ecological_extinction" title="Ecological extinction">Ecological extinction</a></li> <li><a href="/wiki/Endemism" title="Endemism">Endemic species</a></li> <li><a href="/wiki/Flagship_species" title="Flagship species">Flagship species</a></li> <li><a href="/wiki/Ordination_(statistics)" title="Ordination (statistics)">Gradient analysis</a></li> <li><a href="/wiki/Bioindicator" title="Bioindicator">Indicator species</a></li> <li><a href="/wiki/Introduced_species" title="Introduced species">Introduced species</a></li> <li><a href="/wiki/Invasive_species" title="Invasive species">Invasive species</a> / <a href="/wiki/Native_species" title="Native species">Native species</a></li> <li><a href="/wiki/Latitudinal_gradients_in_species_diversity" title="Latitudinal gradients in species diversity">Latitudinal gradients in species diversity</a></li> <li><a href="/wiki/Minimum_viable_population" title="Minimum viable population">Minimum viable population</a></li> <li><a href="/wiki/Unified_neutral_theory_of_biodiversity" title="Unified neutral theory of biodiversity">Neutral theory</a></li> <li><a href="/wiki/Occupancy%E2%80%93abundance_relationship" title="Occupancy–abundance relationship">Occupancy–abundance relationship</a></li> <li><a href="/wiki/Population_viability_analysis" title="Population viability analysis">Population viability analysis</a></li> <li><a href="/wiki/Priority_effect" title="Priority effect">Priority effect</a></li> <li><a href="/wiki/Rapoport%27s_rule" title="Rapoport's rule">Rapoport's rule</a></li> <li><a href="/wiki/Relative_abundance_distribution" title="Relative abundance distribution">Relative abundance distribution</a></li> <li><a href="/wiki/Relative_species_abundance" title="Relative species abundance">Relative species abundance</a></li> <li><a href="/wiki/Species_diversity" title="Species diversity">Species diversity</a></li> <li><a href="/wiki/Species_homogeneity" title="Species homogeneity">Species homogeneity</a></li> <li><a href="/wiki/Species_richness" title="Species richness">Species richness</a></li> <li><a href="/wiki/Species_distribution" title="Species distribution">Species distribution</a></li> <li><a href="/wiki/Species%E2%80%93area_relationship" title="Species–area relationship">Species–area curve</a></li> <li><a href="/wiki/Umbrella_species" title="Umbrella species">Umbrella species</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species<br />interaction</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antibiosis" title="Antibiosis">Antibiosis</a></li> <li><a href="/wiki/Biological_interaction" title="Biological interaction">Biological interaction</a></li> <li><a href="/wiki/Commensalism" title="Commensalism">Commensalism</a></li> <li><a href="/wiki/Community_(ecology)" title="Community (ecology)">Community ecology</a></li> <li><a href="/wiki/Ecological_facilitation" title="Ecological facilitation">Ecological facilitation</a></li> <li><a href="/wiki/Interspecific_competition" title="Interspecific competition">Interspecific competition</a></li> <li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li> <li><a href="/wiki/Parasitism" title="Parasitism">Parasitism</a></li> <li><a href="/wiki/Storage_effect" title="Storage effect">Storage effect</a></li> <li><a href="/wiki/Symbiosis" title="Symbiosis">Symbiosis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Spatial_ecology" title="Spatial ecology">Spatial<br />ecology</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</a></li> <li><a href="/wiki/Cross-boundary_subsidy" title="Cross-boundary subsidy">Cross-boundary subsidy</a></li> <li><a href="/wiki/Cline_(biology)" title="Cline (biology)">Ecocline</a></li> <li><a href="/wiki/Ecotone" title="Ecotone">Ecotone</a></li> <li><a href="/wiki/Ecotype" title="Ecotype">Ecotype</a></li> <li><a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">Disturbance</a></li> <li><a href="/wiki/Edge_effects" title="Edge effects">Edge effects</a></li> <li><a href="/wiki/Foster%27s_rule" title="Foster's rule">Foster's rule</a></li> <li><a href="/wiki/Habitat_fragmentation" title="Habitat fragmentation">Habitat fragmentation</a></li> <li><a href="/wiki/Ideal_free_distribution" title="Ideal free distribution">Ideal free distribution</a></li> <li><a href="/wiki/Intermediate_disturbance_hypothesis" title="Intermediate disturbance hypothesis">Intermediate disturbance hypothesis</a></li> <li><a href="/wiki/Insular_biogeography" title="Insular biogeography">Insular biogeography</a></li> <li><a href="/wiki/Land_change_modeling" title="Land change modeling">Land change modeling</a></li> <li><a href="/wiki/Landscape_ecology" title="Landscape ecology">Landscape ecology</a></li> <li><a href="/wiki/Landscape_epidemiology" title="Landscape epidemiology">Landscape epidemiology</a></li> <li><a href="/wiki/Landscape_limnology" title="Landscape limnology">Landscape limnology</a></li> <li><a href="/wiki/Metapopulation" title="Metapopulation">Metapopulation</a></li> <li><a href="/wiki/Patch_dynamics" title="Patch dynamics">Patch dynamics</a></li> <li><a href="/wiki/R/K_selection_theory" title="R/K selection theory"><i>r</i>/<i>K</i> selection theory</a></li> <li><a href="/wiki/Resource_selection_function" title="Resource selection function">Resource selection function</a></li> <li><a href="/wiki/Source%E2%80%93sink_dynamics" title="Source–sink dynamics">Source–sink dynamics</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Ecological_niche" title="Ecological niche">Niche</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ecological_niche" title="Ecological niche">Ecological niche</a></li> <li><a href="/wiki/Ecological_trap" title="Ecological trap">Ecological trap</a></li> <li><a href="/wiki/Ecosystem_engineer" title="Ecosystem engineer">Ecosystem engineer</a></li> <li><a href="/wiki/Species_distribution_modelling" title="Species distribution modelling">Environmental niche modelling</a></li> <li><a href="/wiki/Guild_(ecology)" title="Guild (ecology)">Guild</a></li> <li><a href="/wiki/Habitat" title="Habitat">Habitat</a> <ul><li><a href="/wiki/Marine_habitat" title="Marine habitat">marine habitats</a></li></ul></li> <li><a href="/wiki/Limiting_similarity" title="Limiting similarity">Limiting similarity</a></li> <li><a href="/wiki/Niche_apportionment_models" title="Niche apportionment models">Niche apportionment models</a></li> <li><a href="/wiki/Niche_construction" title="Niche construction">Niche construction</a></li> <li><a href="/wiki/Niche_differentiation" class="mw-redirect" title="Niche differentiation">Niche differentiation</a></li> <li><a href="/wiki/Ontogenetic_niche_shift" title="Ontogenetic niche shift">Ontogenetic niche shift</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Non-trophic_networks" title="Non-trophic networks">Other<br />networks</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Assembly_rules" title="Assembly rules">Assembly rules</a></li> <li><a href="/wiki/Bateman%27s_principle" title="Bateman's principle">Bateman's principle</a></li> <li><a href="/wiki/Bioluminescence" title="Bioluminescence">Bioluminescence</a></li> <li><a href="/wiki/Ecological_collapse" class="mw-redirect" title="Ecological collapse">Ecological collapse</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological debt</a></li> <li><a href="/wiki/Ecological_debt" title="Ecological debt">Ecological deficit</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Ecological energetics</a></li> <li><a href="/wiki/Ecological_indicator" title="Ecological indicator">Ecological indicator</a></li> <li><a href="/wiki/Ecological_threshold" title="Ecological threshold">Ecological threshold</a></li> <li><a href="/wiki/Ecosystem_diversity" title="Ecosystem diversity">Ecosystem diversity</a></li> <li><a href="/wiki/Emergence" title="Emergence">Emergence</a></li> <li><a href="/wiki/Extinction_debt" title="Extinction debt">Extinction debt</a></li> <li><a href="/wiki/Kleiber%27s_law" title="Kleiber's law">Kleiber's law</a></li> <li><a href="/wiki/Liebig%27s_law_of_the_minimum" title="Liebig's law of the minimum">Liebig's law of the minimum</a></li> <li><a href="/wiki/Marginal_value_theorem" title="Marginal value theorem">Marginal value theorem</a></li> <li><a href="/wiki/Thorson%27s_rule" title="Thorson's rule">Thorson's rule</a></li> <li><a href="/wiki/Xerosere" title="Xerosere">Xerosere</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Allometry" title="Allometry">Allometry</a></li> <li><a class="mw-selflink selflink">Alternative stable state</a></li> <li><a href="/wiki/Balance_of_nature" title="Balance of nature">Balance of nature</a></li> <li><a href="/wiki/Biological_data_visualization" title="Biological data visualization">Biological data visualization</a></li> <li><a href="/wiki/Ecological_economics" title="Ecological economics">Ecological economics</a></li> <li><a href="/wiki/Ecological_footprint" title="Ecological footprint">Ecological footprint</a></li> <li><a href="/wiki/Ecological_forecasting" title="Ecological forecasting">Ecological forecasting</a></li> <li><a href="/wiki/Environmental_humanities" title="Environmental humanities">Ecological humanities</a></li> <li><a href="/wiki/Ecological_stoichiometry" title="Ecological stoichiometry">Ecological stoichiometry</a></li> <li><a href="/wiki/Ecopath" title="Ecopath">Ecopath</a></li> <li><a href="/wiki/Ecosystem_based_fisheries" class="mw-redirect" title="Ecosystem based fisheries">Ecosystem based fisheries</a></li> <li><a href="/wiki/Endolith" title="Endolith">Endolith</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Functional_ecology" title="Functional ecology">Functional ecology</a></li> <li><a href="/wiki/Industrial_ecology" title="Industrial ecology">Industrial ecology</a></li> <li><a href="/wiki/Macroecology" title="Macroecology">Macroecology</a></li> <li><a href="/wiki/Microecosystem" title="Microecosystem">Microecosystem</a></li> <li><a href="/wiki/Natural_environment" title="Natural environment">Natural environment</a></li> <li><a href="/wiki/Regime_shift" title="Regime shift">Regime shift</a></li> <li><a href="/wiki/Sexecology" title="Sexecology">Sexecology</a></li> <li><a href="/wiki/Systems_ecology" title="Systems ecology">Systems ecology</a></li> <li><a href="/wiki/Urban_ecology" title="Urban ecology">Urban ecology</a></li> <li><a href="/wiki/Theoretical_ecology" title="Theoretical ecology">Theoretical ecology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Outline_of_ecology" title="Outline of ecology">Outline of ecology</a></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1&useformat=desktop" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" 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