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<span id="openzim-page-title" class="mw-page-title-main"><span class="mw-page-title-main">Extinction debt</span></span>
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<p>In <a href="Ecology" title="Ecology">ecology</a>, <b>extinction debt</b> is the future <a href="Extinction" title="Extinction">extinction</a> of <a href="Species" title="Species">species</a> due to events in the past. The phrases <b>dead clade walking</b> and <b>survival without recovery</b> express the same idea.<sup id="cite_ref-Jablonski2001_1-0" class="reference"><a href="#cite_note-Jablonski2001-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p><p>Extinction debt occurs because of time delays between impacts on a species, such as destruction of <a href="Habitat" title="Habitat">habitat</a>, and the species' ultimate disappearance. For instance, long-lived <a href="Trees" class="mw-redirect" title="Trees">trees</a> may survive for many years even after reproduction of new trees has become impossible, and thus they may be committed to extinction. Technically, extinction debt generally refers to the <i>number of species</i> in an area likely to become extinct, rather than the prospects of any one species, but <a href="Colloquially" class="mw-redirect" title="Colloquially">colloquially</a> it refers to any occurrence of delayed extinction.
</p><p>Extinction debt may be local or global, but most examples are local as these are easier to observe and model. It is most likely to be found in long-lived species and species with very specific habitat requirements (specialists).<sup id="cite_ref-Kuussaari2009_2-0" class="reference"><a href="#cite_note-Kuussaari2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Extinction debt has important implications for conservation, as it implies that species may become extinct due to past habitat destruction, even if continued impacts cease, and that current reserves may not be sufficient to maintain the species that occupy them. Interventions such as <a href="Habitat_restoration" class="mw-redirect" title="Habitat restoration">habitat restoration</a> may reverse extinction debt.
</p><p><i>Immigration credit</i> is the corollary to extinction debt. It refers to the number of species likely to migrate to an area after an event such as the <a href="Ecosystem_restoration" class="mw-redirect" title="Ecosystem restoration">restoration of an ecosystem</a>.<sup id="cite_ref-Jackson2010_3-0" class="reference"><a href="#cite_note-Jackson2010-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>
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<div class="mw-heading mw-heading2"><h2 id="Terminology">Terminology</h2></div>
<p>The term <i>extinction debt</i> was first used in 1994 in a paper by <a href="David_Tilman" class="mw-redirect" title="David Tilman">David Tilman</a>, <a href="Robert_May%2C_Baron_May_of_Oxford" title="Robert May, Baron May of Oxford">Robert May</a>, Clarence Lehman and <a href="Martin_Nowak" title="Martin Nowak">Martin Nowak</a>,<sup id="cite_ref-Tilman1994_4-0" class="reference"><a href="#cite_note-Tilman1994-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> although <a href="Jared_Diamond" title="Jared Diamond">Jared Diamond</a> used the term "relaxation time" to describe a similar phenomenon in 1972.<sup id="cite_ref-Diamond1972_5-0" class="reference"><a href="#cite_note-Diamond1972-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>
</p><p>Extinction debt is also known by the terms <i>dead clade walking</i> and <i>survival without recovery</i><sup id="cite_ref-Jablonski2001_1-1" class="reference"><a href="#cite_note-Jablonski2001-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> when referring to the species affected. The phrase "dead clade walking" was coined by <a href="David_Jablonski" title="David Jablonski">David Jablonski</a> as early as 2001<sup id="cite_ref-Jablonski2001_1-2" class="reference"><a href="#cite_note-Jablonski2001-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> as a reference to <i><a href="Dead_Man_Walking_(film)" title="Dead Man Walking (film)">Dead Man Walking</a></i>,<sup id="cite_ref-Jablonski2002_6-0" class="reference"><a href="#cite_note-Jablonski2002-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> a film whose title is based on American <a href="Prison_slang" title="Prison slang">prison slang</a> for a condemned prisoner's last walk to the execution chamber. "Dead clade walking" has since appeared in other scientists' writings about the aftermaths of mass extinctions.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup>
</p><p>In discussions of <a href="Conservation_biology#Threats_to_biodiversity" title="Conservation biology">threats to biodiversity</a>, extinction debt is analogous to the "<a href="Climate_commitment" title="Climate commitment">climate commitment</a>" in <a href="Climate_change" title="Climate change">climate change</a>, which states that <a href="Inertia" title="Inertia">inertia</a> will cause the earth to continue to warm for centuries even if no more <a href="Greenhouse_gasses" class="mw-redirect" title="Greenhouse gasses">greenhouse gasses</a> are emitted. Similarly, the <a href="Holocene_extinction" title="Holocene extinction">current extinction</a> may continue long after <a href="Human_impacts_on_the_environment" class="mw-redirect" title="Human impacts on the environment">human impacts</a> on species halt.
</p>
<div class="mw-heading mw-heading2"><h2 id="Causes">Causes</h2></div>
<p>Extinction debt is caused by many of the same drivers as <a href="Extinction" title="Extinction">extinction</a>. The most well-known drivers of extinction debt are <a href="Habitat_fragmentation" title="Habitat fragmentation">habitat fragmentation</a> and <a href="Habitat_destruction" title="Habitat destruction">habitat destruction</a>.<sup id="cite_ref-Kuussaari2009_2-1" class="reference"><a href="#cite_note-Kuussaari2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> These cause extinction debt by reducing the ability of species to persist via immigration to new habitats. Under <a href="https://en.wiktionary.org/wiki/equilibrium" class="extiw external" title="wikt:equilibrium">equilibrium</a> conditions, a species may become extinct in one <a href="Landscape_ecology#Patch_and_mosaic" title="Landscape ecology">habitat patch</a> yet continue to survive because it can <a href="Biological_dispersal" title="Biological dispersal">disperse</a> to other patches. However, as other patches have been destroyed or rendered inaccessible due to fragmentation, this "insurance" effect is reduced and the species may ultimately become extinct.
</p><p><a href="Pollution" title="Pollution">Pollution</a> may also cause extinction debt by reducing a species' <a href="Birth_rate" title="Birth rate">birth rate</a> or increasing its <a href="Death_rate" class="mw-redirect" title="Death rate">death rate</a> so that its population slowly declines.<sup id="cite_ref-Loehle1996_9-0" class="reference"><a href="#cite_note-Loehle1996-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Extinction debts may also be caused by <a href="Invasive_species" title="Invasive species">invasive species</a><sup id="cite_ref-Sax2008_10-0" class="reference"><a href="#cite_note-Sax2008-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> or by <a href="Climate_change" title="Climate change">climate change</a>.
</p><p>Extinction debt may also occur due to the loss of <a href="Mutualism_(biology)" title="Mutualism (biology)">mutualist</a> species. In New Zealand, the local extinction of several species of pollinating birds in 1870 has caused a long-term reduction in the reproduction of the shrub species <i><a href="Rhabdothamnus_solandri" class="mw-redirect" title="Rhabdothamnus solandri">Rhabdothamnus solandri</a></i>, which requires these birds to produce seeds. However, as the plant is slow-growing and long-lived, its populations persist.<sup id="cite_ref-Anderson2010_11-0" class="reference"><a href="#cite_note-Anderson2010-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup>
</p><p>Jablonski recognized at least four patterns in the fossil record following mass extinctions:<sup id="cite_ref-Jablonski2001_1-3" class="reference"><a href="#cite_note-Jablonski2001-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>
</p>
<dl><dt>(1) survival without recovery</dt>
<dd>also called “<i>dead clade walking</i>” – a group dwindling to extinction or relegation to precarious, minor ecological niches</dd>
<dt>(2) continuity with setbacks</dt>
<dd>patterns disturbed by the extinction event but soon continuing on the previous trajectory</dd>
<dt>(3) unbroken continuity</dt>
<dd>large-scale patterns continuing with little disruption</dd>
<dt>(4) unbridled diversification</dt>
<dd>an increase in diversity and species richness, as in the mammals following the <a href="End-Cretaceous_extinction_event" class="mw-redirect" title="End-Cretaceous extinction event">end-Cretaceous extinction event</a></dd></dl>
<div class="mw-heading mw-heading2"><h2 id="Rate_of_extinction">Rate of extinction</h2></div>
<p>Jablonski found that the extinction rate of marine <a href="Invertebrate" title="Invertebrate">invertebrates</a> was significantly higher in the <a href="Geologic_timescale" class="mw-redirect" title="Geologic timescale">stage</a> (major subdivision of an <a href="Geologic_timescale" class="mw-redirect" title="Geologic timescale">epoch</a> – typically 2–10&nbsp;million years' duration) following a mass extinction than in the stages preceding the mass extinction. His analysis focused on marine <a href="Mollusca" title="Mollusca">molluscs</a> since they constitute the most abundant group of fossils and are therefore the least likely to produce <a href="Sampling_error" title="Sampling error">sampling errors</a>. Jablonski suggested that two possible explanations deserved further study:
</p>
<ul><li>Post-extinction physical environments differed from pre-extinction environments in ways which were disadvantageous to the "dead clades walking".</li>
<li><a href="Ecosystem" title="Ecosystem">Ecosystems</a> that developed after recoveries from mass extinctions may have been less favorable for the "dead clades walking".<sup id="cite_ref-Jablonski2002_6-1" class="reference"><a href="#cite_note-Jablonski2002-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup></li></ul>
<div class="mw-heading mw-heading2"><h2 id="Time_scale">Time scale</h2></div>
<p>The time to "payoff" of extinction debt can be very long. Islands that lost habitat at the end of the last <a href="Ice_age" title="Ice age">ice age</a> 10,000&nbsp;years ago still appear to be losing species as a result.<sup id="cite_ref-Diamond1972_5-1" class="reference"><a href="#cite_note-Diamond1972-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> It has been shown that some <a href="Bryozoa" title="Bryozoa">bryozoans</a>, a type of microscopic marine organism, became extinct due to the <a href="Volcano" title="Volcano">volcanic</a> rise of the <a href="Isthmus_of_Panama" title="Isthmus of Panama">Isthmus of Panama</a>. This event cut off the flow of nutrients from the <a href="Pacific_Ocean" title="Pacific Ocean">Pacific Ocean</a> to the <a href="Caribbean" title="Caribbean">Caribbean</a> 3–4.5&nbsp;million years ago. While bryozoan populations dropped severely at this time, extinction of these species took another 1–2&nbsp;million years.<sup id="cite_ref-Jackson2009_12-0" class="reference"><a href="#cite_note-Jackson2009-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup>
</p><p>Extinction debts incurred due to human actions have shorter timescales. Local extinction of birds from rainforest fragmentation occurs over years or decades,<sup id="cite_ref-Gonzalez2000_13-0" class="reference"><a href="#cite_note-Gonzalez2000-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> while plants in fragmented grasslands show debts lasting 50–100&nbsp;years.<sup id="cite_ref-Lindborg2004_14-0" class="reference"><a href="#cite_note-Lindborg2004-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Tree species in fragmented temperate forests have debts lasting 200 years or more.<sup id="cite_ref-Vellend2006_15-0" class="reference"><a href="#cite_note-Vellend2006-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Theoretical_development">Theoretical development</h2></div>
<div class="mw-heading mw-heading3"><h3 id="Origins_in_metapopulation_models">Origins in metapopulation models</h3></div>
<p>Tilman et al. demonstrated that extinction debt could occur using a mathematical <a href="Ecosystem_model" title="Ecosystem model">ecosystem model</a> of species <a href="Metapopulations" class="mw-redirect" title="Metapopulations">metapopulations</a>. <a href="Metapopulations" class="mw-redirect" title="Metapopulations">Metapopulations</a> are multiple populations of a species that live in separate <a href="Landscape_ecology#Patch_and_mosaic" title="Landscape ecology">habitat patches</a> or islands but interact via immigration between the patches. In this model, species persist via a balance between random local extinctions in patches and <a href="Colonisation_(biology)" title="Colonisation (biology)">colonization</a> of new patches. Tilman <i>et al.</i> used this model to predict that species would persist long after they no longer had sufficient habitat to support them. When used to estimate extinction debts of tropical tree species, the model predicted debts lasting 50–400&nbsp;years.<sup id="cite_ref-Tilman1994_4-1" class="reference"><a href="#cite_note-Tilman1994-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup>
</p><p>One of the assumptions underlying the original extinction debt model was a trade-off between species' <a href="Competition_(biology)" title="Competition (biology)">competitive</a> ability and colonization ability. That is, a species that competes well against other species, and is more likely to become dominant in an area, is less likely to colonize new habitats due to evolutionary trade-offs. One of the implications of this assumption is that better competitors, which may even be more common than other species, are more likely to become extinct than rarer, less competitive, better dispersing species. This has been one of the more controversial components of the model, as there is little evidence for this trade-off in many ecosystems, and in many empirical studies dominant competitors were least likely species to become extinct.<sup id="cite_ref-McCarthy1997_16-0" class="reference"><a href="#cite_note-McCarthy1997-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> A later modification of the model showed that these trade-off assumptions may be relaxed, but need to exist partially, in order for the theory to work.<sup id="cite_ref-Banks1997_17-0" class="reference"><a href="#cite_note-Banks1997-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Development_in_other_models">Development in other models</h3></div>
<p>Further theoretical work has shown that extinction debt can occur under many different circumstances, driven by different mechanisms and under different model assumptions. The original model predicted extinction debt as a result of habitat destruction in a system of small, isolated habitats such as islands. Later models showed that extinction debt could occur in systems where habitat destruction occurs in small areas within a large area of habitat, as in <a href="Slash-and-burn" class="mw-redirect" title="Slash-and-burn">slash-and-burn</a> agriculture in forests, and could also occur due to decreased growth of species from pollutants.<sup id="cite_ref-Loehle1996_9-1" class="reference"><a href="#cite_note-Loehle1996-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Predicted patterns of extinction debt differ between models, though. For instance, habitat destruction resembling slash-and-burn agriculture is thought to affect rare species rather than poor colonizers. Models that incorporate <a href="Stochastic_process" title="Stochastic process">stochasticity</a>, or random fluctuation in populations, show extinction debt occurring over different time scales than classic models.<sup id="cite_ref-Etienne2002_18-0" class="reference"><a href="#cite_note-Etienne2002-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup>
</p><p>Most recently, extinction debts have been estimated through the use models derived from <a href="Unified_neutral_theory_of_biodiversity" title="Unified neutral theory of biodiversity">neutral theory</a>. Neutral theory has very different assumptions than the metapopulation models described above. It predicts that the abundance and distribution of species can be predicted entirely through random processes, without considering the traits of individual species. As extinction debt arises in models under such different assumptions, it is robust to different kinds of models. Models derived from neutral theory have successfully predicted extinction times for a number of bird species, but perform poorly at both very small and very large spatial scales.<sup id="cite_ref-Halley2011_19-0" class="reference"><a href="#cite_note-Halley2011-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup>
</p><p><a href="Mathematical_model" title="Mathematical model">Mathematical models</a> have also shown that extinction debt will last longer if it occurs in response to large habitat impacts (as the system will move farther from equilibrium), and if species are long-lived. Also, species just below their <a href="Extinction_threshold" title="Extinction threshold">extinction threshold</a>, that is, just below the population level or habitat occupancy levels required sustain their population, will have long-term extinction debts. Finally, extinction debts are predicted to last longer in landscapes with a few large patches of habitat, rather than many small ones.<sup id="cite_ref-Hanski2002_20-0" class="reference"><a href="#cite_note-Hanski2002-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Detection">Detection</h2></div>
<p>Extinction debt is difficult to detect and measure. Processes that drive extinction debt are inherently slow and highly variable (noisy), and it is difficult to locate or count the very small populations of near-extinct species. Because of these issues, most measures of extinction debt have a great deal of uncertainty.<sup id="cite_ref-Kuussaari2009_2-2" class="reference"><a href="#cite_note-Kuussaari2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Experimental_evidence">Experimental evidence</h3></div>
<p>Due to the logistical and ethical difficulties of inciting extinction debt, there are few studies of extinction debt in controlled experiments. However, experiments <a href="Microcosm%3A_Model_/_experimental_ecosystem" class="mw-redirect" title="Microcosm: Model / experimental ecosystem">microcosms</a> of <a href="Insects" class="mw-redirect" title="Insects">insects</a> living on moss habitats demonstrated that extinction debt occurs after habitat destruction. In these experiments, it took 6–12&nbsp;months for species to die out following the destruction of habitat.<sup id="cite_ref-Gonzalez2000_13-1" class="reference"><a href="#cite_note-Gonzalez2000-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Observational_methods">Observational methods</h3></div>
<div class="mw-heading mw-heading4"><h4 id="Long-term_observation">Long-term observation</h4></div>
<p>Extinction debts that reach equilibrium in relatively short time scales (years to decades) can be observed via measuring the change in species numbers in the time following an impact on habitat. For instance, in the <a href="Amazon_rainforest" title="Amazon rainforest">Amazon rainforest</a>, researchers have measured the rate at which bird species disappear after forest is cut down.<sup id="cite_ref-Stouffer2009_21-0" class="reference"><a href="#cite_note-Stouffer2009-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> As even short-term extinction debts can take years to decades to reach equilibrium, though, such studies take many years and good data are rare.
</p>
<div class="mw-heading mw-heading4"><h4 id="Comparing_the_past_and_present">Comparing the past and present</h4></div>
<p>Most studies of extinction debt compare species numbers with habitat patterns from the past and habitat patterns in the present. If the present populations of species are more closely related to past habitat patterns than present, extinction debt is a likely explanation. The magnitude of extinction debt (i.e., number of species likely to become extinct) can not be estimated by this method.<sup id="cite_ref-Kuussaari2009_2-3" class="reference"><a href="#cite_note-Kuussaari2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p><p>If one has information on species populations from the past in addition to the present, the magnitude of extinction debt can be estimated. One can use the relationship between species and habitat from the past to predict the number of species expected in the present. The difference between this estimate and the actual number of species is the extinction debt.<sup id="cite_ref-Kuussaari2009_2-4" class="reference"><a href="#cite_note-Kuussaari2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>
</p><p>This method requires the assumption that in the past species and their habitat were in equilibrium, which is often unknown. Also, a common relationship used to equate habitat and species number is the <a href="Species-area_curve" class="mw-redirect" title="Species-area curve">species-area curve</a>, but as the species-area curve arises from very different mechanisms than those in <a href="Metapopulation" title="Metapopulation">metapopulation</a> based models, extinction debts measured in this way may not conform with metapopulation models' predictions.<sup id="cite_ref-Loehle1996_9-2" class="reference"><a href="#cite_note-Loehle1996-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> The relationship between habitat and species number can also be represented by much more complex models that simulate the behavior of many species independently.<sup id="cite_ref-Vellend2006_15-1" class="reference"><a href="#cite_note-Vellend2006-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading4"><h4 id="Comparing_impacted_and_pristine_habitats">Comparing impacted and pristine habitats</h4></div>
<p>If data on past species numbers or habitat are not available, species debt can also be estimated by comparing two different habitats: one which is mostly intact, and another which has had areas cleared and is smaller and more fragmented. One can then measure the relationship of species with the condition of habitat in the intact habitat, and, assuming this represents equilibrium, use it to predict the number of species in the cleared habitat. If this prediction is lower than the actual number of species in the cleared habitat, then the difference represents extinction debt.<sup id="cite_ref-Kuussaari2009_2-5" class="reference"><a href="#cite_note-Kuussaari2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> This method requires many of the same assumptions as methods comparing the past and present.
</p>
<div class="mw-heading mw-heading3"><h3 id="Examples">Examples</h3></div>
<div class="mw-heading mw-heading4"><h4 id="Grasslands">Grasslands</h4></div>
<p>Studies of European grasslands show evidence of extinction debt through both comparisons with the past and between present-day systems with different levels of human impacts. The species diversity of <a href="Grasslands" class="mw-redirect" title="Grasslands">grasslands</a> in <a href="Sweden" title="Sweden">Sweden</a> appears to be a remnant of more connected landscapes present 50 to 100&nbsp;years ago.<sup id="cite_ref-Lindborg2004_14-1" class="reference"><a href="#cite_note-Lindborg2004-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> In <a href="Alvar" title="Alvar">alvar grasslands</a> in <a href="Estonia" title="Estonia">Estonia</a> that have lost area since the 1930s, 17–70% of species are estimated to be committed to extinction.<sup id="cite_ref-Helm2006_22-0" class="reference"><a href="#cite_note-Helm2006-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> However, studies of similar grasslands in <a href="Belgium" title="Belgium">Belgium</a>, where similar impacts have occurred, show no evidence of extinction debt.<sup id="cite_ref-Adriaens2006_23-0" class="reference"><a href="#cite_note-Adriaens2006-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> This may be due to differences in the scale of measurement or the level of specialization of grass species.<sup id="cite_ref-Cousins2010_24-0" class="reference"><a href="#cite_note-Cousins2010-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading4"><h4 id="Forests">Forests</h4></div>
<p>Forests in <a href="Flemish_Brabant" title="Flemish Brabant">Flemish Brabant</a>, Belgium, show evidence of extinction debt remaining from <a href="Deforestation" title="Deforestation">deforestation</a> that occurred between 1775 and 1900. Detailed modeling of species behavior, based on similar forests in <a href="England" title="England">England</a> that did not experience deforestation, showed that long-lived and slow-growing species were more common than equilibrium models would predict, indicating that their presence was due to lingering extinction debt.<sup id="cite_ref-Vellend2006_15-2" class="reference"><a href="#cite_note-Vellend2006-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>
</p><p>In Sweden, some species of <a href="Lichens" class="mw-redirect" title="Lichens">lichens</a> show an extinction debt in fragments of ancient forest. However, species of lichens that are habitat <a href="Generalist_and_specialist_species" title="Generalist and specialist species">generalists, rather than specialists</a>, do not.<sup id="cite_ref-Berglund2005_25-0" class="reference"><a href="#cite_note-Berglund2005-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading4"><h4 id="Insects">Insects</h4></div>
<p>Extinction debt has been found among species of <a href="Butterflies" class="mw-redirect" title="Butterflies">butterflies</a> living in the grasslands on <a href="Saaremaa" title="Saaremaa">Saaremaa</a> and <a href="Muhu" title="Muhu">Muhu</a> – islands off the western coast of Estonia. Butterfly species distributions on these islands are better explained by the habitat in the past than current habitats.<sup id="cite_ref-Sang2010_26-0" class="reference"><a href="#cite_note-Sang2010-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup>
</p><p>On the islands of the <a href="Azores" title="Azores">Azores</a> <a href="Archipelago" title="Archipelago">Archipelago</a>, more than 95% of native forests have been destroyed in the past 600&nbsp;years. As a result, more than half of <a href="Arthropods" class="mw-redirect" title="Arthropods">arthropods</a> on these islands are believed to be committed to extinction, with many islands likely to lose more than 90% of species.<sup id="cite_ref-Kostas2010_27-0" class="reference"><a href="#cite_note-Kostas2010-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading4"><h4 id="Vertebrates">Vertebrates</h4></div>
<p>Of extinction from past <a href="Deforestation_of_the_Amazon_rainforest" title="Deforestation of the Amazon rainforest">deforestation in the Amazon</a>, 80–90% has yet to occur, based on modeling based on species-area relationships. Local extinctions of approximately 6&nbsp;species are expected in each 2500&nbsp;km<sup>2</sup> region by 2050 due to past deforestation.<sup id="cite_ref-Wearn2012_28-0" class="reference"><a href="#cite_note-Wearn2012-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> Birds in the <a href="Amazon_rainforest" title="Amazon rainforest">Amazon rainforest</a> continued to become extinct locally for 12&nbsp;years following logging that broke up <a href="Geographic_contiguity" title="Geographic contiguity">contiguous</a> forest into smaller fragments. The extinction rate slowed, however, as forest regrew in the spaces in between habitat fragments.<sup id="cite_ref-Stouffer2009_21-1" class="reference"><a href="#cite_note-Stouffer2009-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup>
</p><p>Countries in <a href="Africa" title="Africa">Africa</a> are estimated to have, on average, a local extinction debt of 30% for forest-dwelling <a href="Primates" class="mw-redirect" title="Primates">primates</a>. That is, they are expected to have 30% of their forest primate species to become extinct in the future due to loss of forest <a href="Habitat" title="Habitat">habitat</a>. The time scale for these extinctions has not been estimated.<sup id="cite_ref-29" class="reference"><a href="#cite_note-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup>
</p><p>Based on historical species-area relationships, <a href="Hungary" title="Hungary">Hungary</a> currently has approximately nine more species of <a href="Bird_of_prey" title="Bird of prey">raptors</a> than are thought to be able to be supported by current nature reserves.<sup id="cite_ref-Baldi2006_30-0" class="reference"><a href="#cite_note-Baldi2006-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Applications_to_conservation">Applications to conservation</h2></div>
<p>The existence of extinction debt in many different ecosystems has important implications for <a href="Conservation_biology" title="Conservation biology">conservation</a>. It implies that in the absence of further <a href="Habitat_destruction" title="Habitat destruction">habitat destruction</a> or other environmental impacts, many species are still likely to become extinct. Protection of existing <a href="Habitats" class="mw-redirect" title="Habitats">habitats</a> may not be sufficient to protect species from extinction.<sup id="cite_ref-Baldi2006_30-1" class="reference"><a href="#cite_note-Baldi2006-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup> However, the long time scales of extinction debt may allow for habitat <a href="Restoration_ecology" class="mw-redirect" title="Restoration ecology">restoration</a> in order to prevent extinction,<sup id="cite_ref-Kuussaari2009_2-6" class="reference"><a href="#cite_note-Kuussaari2009-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> as occurred in the slowing of extinction in Amazon forest birds above.<sup id="cite_ref-Stouffer2009_21-2" class="reference"><a href="#cite_note-Stouffer2009-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup> In another example, it has been found that <a href="Grizzly_bears" class="mw-redirect" title="Grizzly bears">grizzly bears</a> in very small reserves in the <a href="Rocky_Mountains" title="Rocky Mountains">Rocky Mountains</a> are likely to become extinct, but this finding allows the modification of reserve networks to better support their populations.<sup id="cite_ref-Carroll2004_31-0" class="reference"><a href="#cite_note-Carroll2004-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup>
</p><p>The extinction debt concept may require revision of the value of land for species conservation, as the number of species currently present in a habitat may not be a good measure of the habitat's ability to support species (see <a href="Carrying_capacity" title="Carrying capacity">carrying capacity</a>) in the future.<sup id="cite_ref-Berglund2005_25-1" class="reference"><a href="#cite_note-Berglund2005-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup> As extinction debt may last longest near extinction thresholds, it may be hardest to detect the threat of extinction for species that conservation could benefit the most.<sup id="cite_ref-Hanski2002_20-1" class="reference"><a href="#cite_note-Hanski2002-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup>
</p><p>Economic analyses have shown that including extinction in management decision-making process changes decision outcomes, as the decision to destroy habitat changes conservation value in the future as well as the present. It is estimated that in <a href="Costa_Rica" title="Costa Rica">Costa Rica</a>, ongoing extinction debt may cost between $88 million and $467 million.<sup id="cite_ref-Leroux2009_32-0" class="reference"><a href="#cite_note-Leroux2009-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="In_popular_culture">In popular culture</h2></div>
<ul><li>An episode of the <a href="CBS" title="CBS">CBS</a> series <i><a href="Elementary_(TV_series)" title="Elementary (TV series)">Elementary</a></i> was named "<a href="List_of_Elementary_episodes" title="List of Elementary episodes">Dead Clade Walking</a>".<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup></li></ul>
<div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2></div>
<ul><li><span class="nowrap"><span class="mw-image-border noviewer" typeof="mw:File"><span></span></span> </span><a href="Portal%3AEcology" title="Portal:Ecology">Ecology portal</a></li></ul>
<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>
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<li id="cite_note-Jablonski2001-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Jablonski2001_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Jablonski2001_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Jablonski2001_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Jablonski2001_1-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><style data-mw-deduplicate="TemplateStyles:r1238218222">
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<li id="cite_note-Kuussaari2009-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Kuussaari2009_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Kuussaari2009_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Kuussaari2009_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Kuussaari2009_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Kuussaari2009_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Kuussaari2009_2-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Kuussaari2009_2-6"><sup><i><b>g</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFKuussaariBommarcoHeikkinenHelm2009" class="citation journal cs1">Kuussaari, M.; Bommarco, R.; Heikkinen, R. K.; Helm, A.; Krauss, J.; Lindborg, R.; Öckinger, E.; Pärtel, M.; Pino, J.; Rodà, F.; Stefanescu, C.; Teder, T.; Zobel, M.; Steffan-Dewenter, I. (2009). "Extinction debt: a challenge for biodiversity conservation". <i>Trends in Ecology &amp; Evolution</i>. <b>24</b> (10): <span class="nowrap">564–</span>71. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.tree.2009.04.011">10.1016/j.tree.2009.04.011</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19665254">19665254</a>.</cite></span>
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<li id="cite_note-Jackson2010-3"><span class="mw-cite-backlink"><b><a href="#cite_ref-Jackson2010_3-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFJacksonSax2010" class="citation journal cs1">Jackson, S. T.; Sax, D. F. (2010). "Balancing biodiversity in a changing environment: extinction debt, immigration credit and species turnover". <i>Trends in Ecology &amp; Evolution</i>. <b>25</b> (3): <span class="nowrap">153–</span>60. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.tree.2009.10.001">10.1016/j.tree.2009.10.001</a>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19879014">19879014</a>.</cite></span>
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<li id="cite_note-Tilman1994-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Tilman1994_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Tilman1994_4-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFTilmanMayLehmanNowak1994" class="citation journal cs1">Tilman, D.; May, R. M.; Lehman, C. L.; Nowak, M. A. (1994). "Habitat destruction and the extinction debt". <i>Nature</i>. <b>371</b> (6492): 65. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1994Natur.371...65T">1994Natur.371...65T</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F371065a0">10.1038/371065a0</a>. <a href="S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a>&nbsp;<a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4308409">4308409</a>.</cite></span>
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<li id="cite_note-Diamond1972-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-Diamond1972_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Diamond1972_5-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFDiamond1972" class="citation journal cs1">Diamond, JM (1972). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC389735">"Biogeographic kinetics: estimation of relaxation times for avifaunas of southwest pacific islands"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>69</b> (11): <span class="nowrap">3199–</span>203. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1972PNAS...69.3199D">1972PNAS...69.3199D</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.69.11.3199">10.1073/pnas.69.11.3199</a></span>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&nbsp;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC389735">389735</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/16592024">16592024</a>.</cite></span>
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<li id="cite_note-Jablonski2002-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Jablonski2002_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Jablonski2002_6-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFJablonski2002" class="citation journal cs1">Jablonski, D (2002). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC123034">"Survival without recovery after mass extinctions"</a>. <i><a href="Proceedings_of_the_National_Academy_of_Sciences" class="mw-redirect" title="Proceedings of the National Academy of Sciences">PNAS</a></i>. <b>99</b> (12): <span class="nowrap">8139–</span>8144. <a href="Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2002PNAS...99.8139J">2002PNAS...99.8139J</a>. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.102163299">10.1073/pnas.102163299</a></span>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a>&nbsp;<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC123034">123034</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a>&nbsp;<a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12060760">12060760</a>.</cite></span>
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<li id="cite_note-7"><span class="mw-cite-backlink"><b><a href="#cite_ref-7">^</a></b></span> <span class="reference-text"><cite id="CITEREFKorn,_D.Belka,_Z.Fröhlich,_S.Rücklin,_M.2007" class="citation journal cs1">Korn, D.; Belka, Z.; Fröhlich, S.; Rücklin, M. &amp; Wendt, J. (Jan 2007). "The youngest African clymeniids (Ammonoidea, Late Devonian) – failed survivors of the Hangenberg Event". <i>Lethaia</i>. <b>37</b> (3): <span class="nowrap">307–</span>315. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00241160410002054">10.1080/00241160410002054</a>.</cite></span>
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<li id="cite_note-Carroll2004-31"><span class="mw-cite-backlink"><b><a href="#cite_ref-Carroll2004_31-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFCarrollNossPaquetSchumaker2004" class="citation journal cs1">Carroll, C.; Noss, R. F.; Paquet, P. C.; Schumaker, N. H. (2004). "Extinction Debt of Protected Areas in Developing Landscapes". <i>Conservation Biology</i>. <b>18</b> (4): 1110. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1523-1739.2004.00083.x">10.1111/j.1523-1739.2004.00083.x</a>.</cite></span>
</li>
<li id="cite_note-Leroux2009-32"><span class="mw-cite-backlink"><b><a href="#cite_ref-Leroux2009_32-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFLerouxMartinGoeschl2009" class="citation journal cs1">Leroux, A. D.; Martin, V. L.; Goeschl, T. (2009). "Optimal conservation, extinction debt, and the augmented quasi-option value☆". <i>Journal of Environmental Economics and Management</i>. <b>58</b>: <span class="nowrap">43–</span>57. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.jeem.2008.10.002">10.1016/j.jeem.2008.10.002</a>.</cite></span>
</li>
<li id="cite_note-33"><span class="mw-cite-backlink"><b><a href="#cite_ref-33">^</a></b></span> <span class="reference-text"><cite id="CITEREFMoore" class="citation web cs1">Moore, A. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20140202161949/http://www.tv.com/shows/elementary/dead-clade-walking-3002035/">"'Elementary' Season 2, Episode 15: 'Dead Clade Walking'"</a>. <i>Atlanta Blackstar</i>. Archived from <a rel="nofollow" class="external text" href="http://www.tv.com/shows/elementary/dead-clade-walking-3002035/">the original</a> on 2 February 2014<span class="reference-accessdate">. Retrieved <span class="nowrap">31 January</span> 2014</span>.</cite></span>
</li>
</ol></div>
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</style><div id="Extinction374" style="font-size:114%;margin:0 4em"><a href="Extinction" title="Extinction">Extinction</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Phenomena</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Background_extinction_rate" title="Background extinction rate">Background extinction rate</a></li>
<li><a href="Coextinction" title="Coextinction">Coextinction</a></li>
<li><a href="De-extinction" title="De-extinction">De-extinction</a></li>
<li><a href="Ecological_extinction" title="Ecological extinction">Ecological extinction</a></li>
<li><a href="Extinct_in_the_wild" title="Extinct in the wild">Extinct in the wild</a></li>
<li><a href="Functional_extinction" title="Functional extinction">Functional extinction</a></li>
<li><a href="Genetic_pollution" title="Genetic pollution">Genetic pollution</a></li>
<li><a href="Lazarus_taxon" title="Lazarus taxon">Lazarus taxon</a></li>
<li><a href="Local_extinction" title="Local extinction">Local extinction</a></li>
<li><a href="Pseudoextinction" title="Pseudoextinction">Pseudoextinction</a></li></ul>
</div></td><td class="noviewer navbox-image" rowspan="8" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="Golden_toad" title="Golden toad"></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Models</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Extinction_vortex" title="Extinction vortex">Extinction vortex</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Causes</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Climate_variability_and_change" title="Climate variability and change">Climate variability and change</a></li>
<li><a href="Genetic_erosion" title="Genetic erosion">Genetic erosion</a></li>
<li><a href="Invasive_species" title="Invasive species">Invasive species</a></li>
<li><a href="Habitat_destruction" title="Habitat destruction">Habitat destruction</a></li>
<li><a href="Human_impact_on_the_environment" title="Human impact on the environment">Human impact on the environment</a></li>
<li><a href="Muller's_ratchet" title="Muller's ratchet">Muller's ratchet</a></li>
<li><a href="Mutational_meltdown" title="Mutational meltdown">Mutational meltdown</a></li>
<li><a href="Overabundant_species" title="Overabundant species">Overabundant species</a></li>
<li><a href="Overexploitation" title="Overexploitation">Overexploitation</a></li>
<li><a href="Overshoot_(population)" title="Overshoot (population)">Overshoot</a></li>
<li><a href="Paradox_of_enrichment" title="Paradox of enrichment">Paradox of enrichment</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Theories<br>and concepts</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul>
<li><a href="Extinction_risk_from_climate_change" title="Extinction risk from climate change">Extinction risk from climate change</a></li>
<li><a href="Extinction_threshold" title="Extinction threshold">Extinction threshold</a></li>
<li><a href="Quasi-extinction" title="Quasi-extinction">Quasi-extinction</a></li>
<li><a href="Field_of_bullets" title="Field of bullets">Field of bullets</a></li>
<li><a href="Hypothetical_species" title="Hypothetical species">Hypothetical species</a></li>
<li><a href="Latent_extinction_risk" title="Latent extinction risk">Latent extinction risk</a></li>
<li><a href="Living_fossil" title="Living fossil">Living fossil</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Extinction_event" title="Extinction event">Extinction events</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Major</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Late_Ordovician_mass_extinction" title="Late Ordovician mass extinction">Ordovician–Silurian</a></li>
<li><a href="Late_Devonian_extinction" class="mw-redirect" title="Late Devonian extinction">Late Devonian</a></li>
<li><a href="Permian%E2%80%93Triassic_extinction_event" title="Permian–Triassic extinction event">Permian–Triassic</a></li>
<li><a href="Triassic%E2%80%93Jurassic_extinction_event" title="Triassic–Jurassic extinction event">Triassic–Jurassic</a></li>
<li><a href="Cretaceous%E2%80%93Paleogene_extinction_event" title="Cretaceous–Paleogene extinction event">Cretaceous–Paleogene</a></li>
<li><a href="Holocene_extinction" title="Holocene extinction">Holocene</a>
<ul><li><a href="Timeline_of_extinctions_in_the_Holocene" title="Timeline of extinctions in the Holocene">Timeline</a></li></ul></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Great_Oxidation_Event" title="Great Oxidation Event">Great Oxidation</a></li>
<li><a href="End-Ediacaran_extinction" title="End-Ediacaran extinction">End-Ediacaran</a></li>
<li><a href="End-Botomian_mass_extinction" title="End-Botomian mass extinction">End-Botomian</a></li>
<li><a href="Dresbachian" title="Dresbachian">Dresbachian</a></li>
<li><a href="Cambrian%E2%80%93Ordovician_extinction_event" title="Cambrian–Ordovician extinction event">Cambrian–Ordovician</a></li>
<li><a href="Ireviken_event" title="Ireviken event">Ireviken</a></li>
<li><a href="Mulde_event" title="Mulde event">Mulde</a></li>
<li><a href="Lau_event" title="Lau event">Lau</a></li>
<li><a href="Carboniferous_rainforest_collapse" title="Carboniferous rainforest collapse">Carboniferous</a></li>
<li><a href="Olson's_Extinction" title="Olson's Extinction">Olson's</a></li>
<li><a href="Emeishan_Traps" title="Emeishan Traps">End-Capitanian</a></li>
<li><a href="Carnian_pluvial_episode" title="Carnian pluvial episode">Carnian Pluvial</a></li>
<li><a href="Toarcian_turnover" class="mw-redirect" title="Toarcian turnover">Toarcian</a></li>
<li><a href="Jurassic" title="Jurassic">End-Jurassic or Tithonian</a></li>
<li><a href="Aptian_extinction" title="Aptian extinction">Aptian</a></li>
<li><a href="Cenomanian-Turonian_boundary_event" title="Cenomanian-Turonian boundary event">Cenomanian-Turonian</a></li>
<li><a href="Eocene%E2%80%93Oligocene_extinction_event" title="Eocene–Oligocene extinction event">Eocene–Oligocene</a></li>
<li><a href="Middle_Miocene_disruption" title="Middle Miocene disruption">Middle Miocene</a></li>
<li><a href="Pliocene#Supernovae" title="Pliocene">Pliocene–Pleistocene</a></li>
<li><a href="Quaternary_extinction_event" class="mw-redirect" title="Quaternary extinction event">Quaternary</a></li></ul>
</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Extinct species</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Lists_of_extinct_species" title="Lists of extinct species">Lists of extinct species</a></li>
<li>IUCN Red List extinct species</li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Organizations</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="International_Union_for_Conservation_of_Nature" title="International Union for Conservation of Nature">International Union for Conservation of Nature</a></li>
<li><a href="IUCN_Species_Survival_Commission" title="IUCN Species Survival Commission">IUCN Species Survival Commission</a></li>
<li><a href="Extinction_Rebellion" title="Extinction Rebellion">Extinction Rebellion</a></li>
<li><a href="Voluntary_Human_Extinction_Movement" title="Voluntary Human Extinction Movement">Voluntary Human Extinction Movement</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">
<ul><li><a href="Anthropocene" title="Anthropocene">Anthropocene</a></li>
<li><a href="Decline_in_amphibian_populations" title="Decline in amphibian populations">Decline in amphibian populations</a></li>
<li><a href="Decline_in_insect_populations" title="Decline in insect populations">Decline in insect populations</a></li>
<li><a href="Endling" title="Endling">Endling</a></li>
<li><a href="Extinction_symbol" title="Extinction symbol">Extinction symbol</a></li>
<li><a href="Human_extinction" title="Human extinction">Human extinction</a></li>
<li><i><a href="The_Sixth_Extinction%3A_An_Unnatural_History" title="The Sixth Extinction: An Unnatural History">The Sixth Extinction: An Unnatural History</a></i></li></ul>
</div></td></tr><tr><td class="navbox-abovebelow hlist" colspan="3"><div>
<ul><li><span class="noviewer" typeof="mw:File"><span title="Category"></span></span> Category</li>
<li><span class="noviewer" typeof="mw:File"><span title="Commons page"></span></span> <a href="https://commons.wikimedia.org/wiki/Category:Extinction" class="extiw external" title="commons:Category:Extinction">Commons</a></li></ul>
</div></td></tr></tbody></table></div>
<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Trophic_components279" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Ecology:_Modelling_ecosystems:_Trophic_components279" style="font-size:114%;margin:0 4em"><a href="Ecology" title="Ecology">Ecology</a>: <a href="Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: <a href="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="Abiotic_component" title="Abiotic component">Abiotic component</a></li>
<li><a href="Abiotic_stress" title="Abiotic stress">Abiotic stress</a></li>
<li><a href="Behavioral_ecology" title="Behavioral ecology">Behaviour</a></li>
<li><a href="Biogeochemical_cycle" title="Biogeochemical cycle">Biogeochemical cycle</a></li>
<li><a href="Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li>
<li><a href="Biotic_component" class="mw-redirect" title="Biotic component">Biotic component</a></li>
<li><a href="Biotic_stress" title="Biotic stress">Biotic stress</a></li>
<li><a href="Carrying_capacity" title="Carrying capacity">Carrying capacity</a></li>
<li><a href="Competition_(biology)" title="Competition (biology)">Competition</a></li>
<li><a href="Ecosystem" title="Ecosystem">Ecosystem</a></li>
<li><a href="Ecosystem_ecology" title="Ecosystem ecology">Ecosystem ecology</a></li>
<li><a href="Ecosystem_model" title="Ecosystem model">Ecosystem model</a></li>
<li><a href="Green_world_hypothesis" title="Green world hypothesis">Green world hypothesis</a></li>
<li><a href="Keystone_species" title="Keystone species">Keystone species</a></li>
<li><a href="List_of_feeding_behaviours" title="List of feeding behaviours">List of feeding behaviours</a></li>
<li><a href="Metabolic_theory_of_ecology" title="Metabolic theory of ecology">Metabolic theory of ecology</a></li>
<li><a href="Productivity_(ecology)" title="Productivity (ecology)">Productivity</a></li>
<li><a href="Resource_(biology)" title="Resource (biology)">Resource</a></li>
<li><a href="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="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="Autotroph" title="Autotroph">Autotrophs</a></li>
<li><a href="Chemosynthesis" title="Chemosynthesis">Chemosynthesis</a></li>
<li><a href="Chemotroph" title="Chemotroph">Chemotrophs</a></li>
<li><a href="Foundation_species" title="Foundation species">Foundation species</a></li>
<li><a href="Kinetotroph" title="Kinetotroph">Kinetotrophs</a></li>
<li><a href="Mixotroph" title="Mixotroph">Mixotrophs</a></li>
<li><a href="Myco-heterotrophy" title="Myco-heterotrophy">Myco-heterotrophy</a></li>
<li><a href="Mycotroph" title="Mycotroph">Mycotroph</a></li>
<li><a href="Organotroph" title="Organotroph">Organotrophs</a></li>
<li><a href="Photoheterotroph" title="Photoheterotroph">Photoheterotrophs</a></li>
<li><a href="Photosynthesis" title="Photosynthesis">Photosynthesis</a></li>
<li><a href="Photosynthetic_efficiency" title="Photosynthetic efficiency">Photosynthetic efficiency</a></li>
<li><a href="Phototroph" title="Phototroph">Phototrophs</a></li>
<li><a href="Primary_nutritional_groups" title="Primary nutritional groups">Primary nutritional groups</a></li>
<li><a href="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="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="Apex_predator" title="Apex predator">Apex predator</a></li>
<li><a href="Bacterivore" title="Bacterivore">Bacterivore</a></li>
<li><a href="Carnivore" title="Carnivore">Carnivores</a></li>
<li><a href="Chemoorganotroph" class="mw-redirect" title="Chemoorganotroph">Chemoorganotroph</a></li>
<li><a href="Foraging" title="Foraging">Foraging</a></li>
<li><a href="Generalist_and_specialist_species" title="Generalist and specialist species">Generalist and specialist species</a></li>
<li><a href="Intraguild_predation" title="Intraguild predation">Intraguild predation</a></li>
<li><a href="Herbivore" title="Herbivore">Herbivores</a></li>
<li><a href="Heterotroph" title="Heterotroph">Heterotroph</a></li>
<li><a href="Heterotrophic_nutrition" title="Heterotrophic nutrition">Heterotrophic nutrition</a></li>
<li><a href="Insectivore" title="Insectivore">Insectivore</a></li>
<li><a href="Mesopredator" title="Mesopredator">Mesopredators</a></li>
<li><a href="Mesopredator_release_hypothesis" title="Mesopredator release hypothesis">Mesopredator release hypothesis</a></li>
<li><a href="Omnivore" title="Omnivore">Omnivores</a></li>
<li><a href="Optimal_foraging_theory" title="Optimal foraging theory">Optimal foraging theory</a></li>
<li><a href="Planktivore" title="Planktivore">Planktivore</a></li>
<li><a href="Predation" title="Predation">Predation</a></li>
<li><a href="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="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="Chemoorganoheterotrophy" class="mw-redirect" title="Chemoorganoheterotrophy">Chemoorganoheterotrophy</a></li>
<li><a href="Decomposition" title="Decomposition">Decomposition</a></li>
<li><a href="Detritivore" title="Detritivore">Detritivores</a></li>
<li><a href="Detritus" title="Detritus">Detritus</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="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="Archaea" title="Archaea">Archaea</a></li>
<li><a href="Bacteriophage" title="Bacteriophage">Bacteriophage</a></li>
<li><a href="Lithoautotroph" title="Lithoautotroph">Lithoautotroph</a></li>
<li><a href="Lithotroph" title="Lithotroph">Lithotrophy</a></li>
<li><a href="Marine_microorganisms" title="Marine microorganisms">Marine</a></li>
<li><a href="Microbial_cooperation" title="Microbial cooperation">Microbial cooperation</a></li>
<li><a href="Microbial_ecology" title="Microbial ecology">Microbial ecology</a></li>
<li><a href="Microbial_food_web" title="Microbial food web">Microbial food web</a></li>
<li><a href="Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li>
<li><a href="Microbial_loop" title="Microbial loop">Microbial loop</a></li>
<li><a href="Microbial_mat" title="Microbial mat">Microbial mat</a></li>
<li><a href="Microbial_metabolism" title="Microbial metabolism">Microbial metabolism</a></li>
<li><a href="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="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="Biomagnification" title="Biomagnification">Biomagnification</a></li>
<li><a href="Ecological_efficiency" title="Ecological efficiency">Ecological efficiency</a></li>
<li><a href="Ecological_pyramid" title="Ecological pyramid">Ecological pyramid</a></li>
<li><a href="Energy_flow_(ecology)" title="Energy flow (ecology)">Energy flow</a></li>
<li><a href="Food_chain" title="Food chain">Food chain</a></li>
<li><a href="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="Lake_ecosystem#Trophic_relationships" title="Lake ecosystem">Lakes</a></li>
<li><a href="River_ecosystem#Trophic_relationships" title="River ecosystem">Rivers</a></li>
<li><a href="Soil_food_web" title="Soil food web">Soil</a></li>
<li><a href="Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></li>
<li><a href="Marine_food_web" title="Marine food web">Marine food webs</a>
<ul><li><a href="Cold_seep" title="Cold seep">cold seeps</a></li>
<li><a href="Hydrothermal_vent#Biological_communities" title="Hydrothermal vent">hydrothermal vents</a></li>
<li><a href="Intertidal_ecology" title="Intertidal ecology">intertidal</a></li>
<li><a href="Kelp_forest#Trophic_ecology" title="Kelp forest">kelp forests</a></li>
<li><a href="North_Pacific_Gyre" title="North Pacific Gyre">North Pacific Gyre</a></li>
<li><a href="Ecology_of_the_San_Francisco_Estuary#Food_web" title="Ecology of the San Francisco Estuary">San Francisco Estuary</a></li>
<li><a href="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="Ascendency" title="Ascendency">Ascendency</a></li>
<li><a href="Bioaccumulation" title="Bioaccumulation">Bioaccumulation</a></li>
<li><a href="Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li>
<li><a href="Climax_community" title="Climax community">Climax community</a></li>
<li><a href="Competitive_exclusion_principle" title="Competitive exclusion principle">Competitive exclusion principle</a></li>
<li><a href="Consumer%E2%80%93resource_interactions" title="Consumer–resource interactions">Consumer–resource interactions</a></li>
<li><a href="Copiotroph" title="Copiotroph">Copiotrophs</a></li>
<li><a href="Dominance_(ecology)" title="Dominance (ecology)">Dominance</a></li>
<li><a href="Ecological_network" title="Ecological network">Ecological network</a></li>
<li><a href="Ecological_succession" title="Ecological succession">Ecological succession</a></li>
<li><a href="Energy_quality" title="Energy quality">Energy quality</a></li>
<li><a href="Energy_systems_language" title="Energy systems language">Energy systems language</a></li>
<li><a href="F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li>
<li><a href="Feed_conversion_ratio" title="Feed conversion ratio">Feed conversion ratio</a></li>
<li><a href="Feeding_frenzy" title="Feeding frenzy">Feeding frenzy</a></li>
<li><a href="Mesotrophic_soil" title="Mesotrophic soil">Mesotrophic soil</a></li>
<li><a href="Nutrient_cycle" title="Nutrient cycle">Nutrient cycle</a></li>
<li><a href="Oligotroph" title="Oligotroph">Oligotroph</a></li>
<li><a href="Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li>
<li><a href="Trophic_cascade" title="Trophic cascade">Trophic cascade</a></li>
<li><a href="Trophic_mutualism" title="Trophic mutualism">Trophic mutualism</a></li>
<li><a href="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="Animal_coloration" title="Animal coloration">Animal coloration</a></li>
<li><a href="Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptations</a></li>
<li><a href="Camouflage" title="Camouflage">Camouflage</a></li>
<li><a href="Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li>
<li><a href="Herbivore_adaptations_to_plant_defense" title="Herbivore adaptations to plant defense">Herbivore adaptations to plant defense</a></li>
<li><a href="Mimicry" title="Mimicry">Mimicry</a></li>
<li><a href="Plant_defense_against_herbivory" title="Plant defense against herbivory">Plant defense against herbivory</a></li>
<li><a href="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"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Other_components285" 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"><div id="Ecology:_Modelling_ecosystems:_Other_components285" style="font-size:114%;margin:0 4em"><a href="Ecology" title="Ecology">Ecology</a>: <a href="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="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="Abundance_(ecology)" title="Abundance (ecology)">Abundance</a></li>
<li><a href="Allee_effect" title="Allee effect">Allee effect</a></li>
<li><a href="Consumer-resource_model" title="Consumer-resource model">Consumer-resource model</a></li>
<li><a href="Depensation" title="Depensation">Depensation</a></li>
<li><a href="Ecological_yield" title="Ecological yield">Ecological yield</a></li>
<li><a href="Effective_population_size" title="Effective population size">Effective population size</a></li>
<li><a href="Intraspecific_competition" title="Intraspecific competition">Intraspecific competition</a></li>
<li><a href="Logistic_function" title="Logistic function">Logistic function</a></li>
<li><a href="Malthusian_growth_model" title="Malthusian growth model">Malthusian growth model</a></li>
<li><a href="Maximum_sustainable_yield" title="Maximum sustainable yield">Maximum sustainable yield</a></li>
<li><a href="Overpopulation" title="Overpopulation">Overpopulation</a></li>
<li><a href="Overexploitation" title="Overexploitation">Overexploitation</a></li>
<li><a href="Population_cycle" title="Population cycle">Population cycle</a></li>
<li><a href="Population_dynamics" title="Population dynamics">Population dynamics</a></li>
<li><a href="Population_model" title="Population model">Population modeling</a></li>
<li><a href="Population_size" title="Population size">Population size</a></li>
<li><a href="Lotka%E2%80%93Volterra_equations" title="Lotka–Volterra equations">Predator–prey (Lotka–Volterra) equations</a></li>
<li><a href="Recruitment_(biology)" title="Recruitment (biology)">Recruitment</a></li>
<li><a href="Small_population_size" title="Small population size">Small population size</a></li>
<li><a href="Ecological_stability" title="Ecological stability">Stability</a>
<ul><li><a href="Ecological_resilience" title="Ecological resilience">Resilience</a></li>
<li><a href="Resistance_(ecology)" title="Resistance (ecology)">Resistance</a></li></ul></li>
<li><a href="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="Biodiversity" title="Biodiversity">Biodiversity</a></li>
<li><a href="Density_dependence" title="Density dependence">Density-dependent inhibition</a></li>
<li><a href="Ecological_effects_of_biodiversity" title="Ecological effects of biodiversity">Ecological effects of biodiversity</a></li>
<li><a href="Ecological_extinction" title="Ecological extinction">Ecological extinction</a></li>
<li><a href="Endemism" title="Endemism">Endemic species</a></li>
<li><a href="Flagship_species" title="Flagship species">Flagship species</a></li>
<li><a href="Ordination_(statistics)" title="Ordination (statistics)">Gradient analysis</a></li>
<li><a href="Bioindicator" title="Bioindicator">Indicator species</a></li>
<li><a href="Introduced_species" title="Introduced species">Introduced species</a></li>
<li><a href="Invasive_species" title="Invasive species">Invasive species</a> / <a href="Native_species" title="Native species">Native species</a></li>
<li><a href="Latitudinal_gradients_in_species_diversity" title="Latitudinal gradients in species diversity">Latitudinal gradients in species diversity</a></li>
<li><a href="Minimum_viable_population" title="Minimum viable population">Minimum viable population</a></li>
<li><a href="Unified_neutral_theory_of_biodiversity" title="Unified neutral theory of biodiversity">Neutral theory</a></li>
<li><a href="Occupancy%E2%80%93abundance_relationship" title="Occupancy–abundance relationship">Occupancy–abundance relationship</a></li>
<li><a href="Population_viability_analysis" title="Population viability analysis">Population viability analysis</a></li>
<li><a href="Priority_effect" title="Priority effect">Priority effect</a></li>
<li><a href="Rapoport's_rule" title="Rapoport's rule">Rapoport's rule</a></li>
<li><a href="Relative_abundance_distribution" title="Relative abundance distribution">Relative abundance distribution</a></li>
<li><a href="Relative_species_abundance" title="Relative species abundance">Relative species abundance</a></li>
<li><a href="Species_diversity" title="Species diversity">Species diversity</a></li>
<li><a href="Species_homogeneity" title="Species homogeneity">Species homogeneity</a></li>
<li><a href="Species_richness" title="Species richness">Species richness</a></li>
<li><a href="Species_distribution" title="Species distribution">Species distribution</a></li>
<li><a href="Species%E2%80%93area_relationship" title="Species–area relationship">Species–area curve</a></li>
<li><a href="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="Antibiosis" title="Antibiosis">Antibiosis</a></li>
<li><a href="Biological_interaction" title="Biological interaction">Biological interaction</a></li>
<li><a href="Commensalism" title="Commensalism">Commensalism</a></li>
<li><a href="Community_(ecology)" title="Community (ecology)">Community ecology</a></li>
<li><a href="Ecological_facilitation" title="Ecological facilitation">Ecological facilitation</a></li>
<li><a href="Interspecific_competition" title="Interspecific competition">Interspecific competition</a></li>
<li><a href="Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li>
<li><a href="Parasitism" title="Parasitism">Parasitism</a></li>
<li><a href="Storage_effect" title="Storage effect">Storage effect</a></li>
<li><a href="Symbiosis" title="Symbiosis">Symbiosis</a></li></ul>
</div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="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="Biogeography" title="Biogeography">Biogeography</a></li>
<li><a href="Cross-boundary_subsidy" title="Cross-boundary subsidy">Cross-boundary subsidy</a></li>
<li><a href="Cline_(biology)" title="Cline (biology)">Ecocline</a></li>
<li><a href="Ecotone" title="Ecotone">Ecotone</a></li>
<li><a href="Ecotype" title="Ecotype">Ecotype</a></li>
<li><a href="Disturbance_(ecology)" title="Disturbance (ecology)">Disturbance</a></li>
<li><a href="Edge_effects" title="Edge effects">Edge effects</a></li>
<li><a href="Foster's_rule" title="Foster's rule">Foster's rule</a></li>
<li><a href="Habitat_fragmentation" title="Habitat fragmentation">Habitat fragmentation</a></li>
<li><a href="Ideal_free_distribution" title="Ideal free distribution">Ideal free distribution</a></li>
<li><a href="Intermediate_disturbance_hypothesis" title="Intermediate disturbance hypothesis">Intermediate disturbance hypothesis</a></li>
<li><a href="Insular_biogeography" title="Insular biogeography">Insular biogeography</a></li>
<li><a href="Land_change_modeling" title="Land change modeling">Land change modeling</a></li>
<li><a href="Landscape_ecology" title="Landscape ecology">Landscape ecology</a></li>
<li><a href="Landscape_epidemiology" title="Landscape epidemiology">Landscape epidemiology</a></li>
<li><a href="Landscape_limnology" title="Landscape limnology">Landscape limnology</a></li>
<li><a href="Metapopulation" title="Metapopulation">Metapopulation</a></li>
<li><a href="Patch_dynamics" title="Patch dynamics">Patch dynamics</a></li>
<li><a href="R/K_selection_theory" title="R/K selection theory"><i>r</i>/<i>K</i> selection theory</a></li>
<li><a href="Resource_selection_function" title="Resource selection function">Resource selection function</a></li>
<li><a href="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="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="Ecological_trap" title="Ecological trap">Ecological trap</a></li>
<li><a href="Ecosystem_engineer" title="Ecosystem engineer">Ecosystem engineer</a></li>
<li><a href="Species_distribution_modelling" title="Species distribution modelling">Environmental niche modelling</a></li>
<li><a href="Guild_(ecology)" title="Guild (ecology)">Guild</a></li>
<li><a href="Habitat" title="Habitat">Habitat</a>
<ul><li><a href="Marine_habitat" title="Marine habitat">Marine</a></li>
<li><a href="Semiaquatic" title="Semiaquatic">Semiaquatic</a></li>
<li><a href="Terrestrial_animal" title="Terrestrial animal">Terrestrial</a></li></ul></li>
<li><a href="Limiting_similarity" title="Limiting similarity">Limiting similarity</a></li>
<li><a href="Niche_apportionment_models" title="Niche apportionment models">Niche apportionment models</a></li>
<li><a href="Niche_construction" title="Niche construction">Niche construction</a></li>
<li><a href="Niche_differentiation" class="mw-redirect" title="Niche differentiation">Niche differentiation</a></li>
<li><a href="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="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="Assembly_rules" title="Assembly rules">Assembly rules</a></li>
<li><a href="Bateman's_principle" title="Bateman's principle">Bateman's principle</a></li>
<li><a href="Bioluminescence" title="Bioluminescence">Bioluminescence</a></li>
<li><a href="Ecological_collapse" class="mw-redirect" title="Ecological collapse">Ecological collapse</a></li>
<li><a href="Ecological_debt" title="Ecological debt">Ecological debt</a></li>
<li><a href="Ecological_debt" title="Ecological debt">Ecological deficit</a></li>
<li><a href="Energy_flow_(ecology)" title="Energy flow (ecology)">Ecological energetics</a></li>
<li><a href="Ecological_indicator" title="Ecological indicator">Ecological indicator</a></li>
<li><a href="Ecological_threshold" title="Ecological threshold">Ecological threshold</a></li>
<li><a href="Ecosystem_diversity" title="Ecosystem diversity">Ecosystem diversity</a></li>
<li><a href="Emergence" title="Emergence">Emergence</a></li>

<li><a href="Kleiber's_law" title="Kleiber's law">Kleiber's law</a></li>
<li><a href="Liebig's_law_of_the_minimum" title="Liebig's law of the minimum">Liebig's law of the minimum</a></li>
<li><a href="Marginal_value_theorem" title="Marginal value theorem">Marginal value theorem</a></li>
<li><a href="Thorson's_rule" title="Thorson's rule">Thorson's rule</a></li>
<li><a href="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="Allometry" title="Allometry">Allometry</a></li>
<li><a href="Alternative_stable_state" title="Alternative stable state">Alternative stable state</a></li>
<li><a href="Balance_of_nature" title="Balance of nature">Balance of nature</a></li>
<li><a href="Biological_data_visualization" title="Biological data visualization">Biological data visualization</a></li>
<li><a href="Ecological_economics" title="Ecological economics">Ecological economics</a></li>
<li><a href="Ecological_footprint" title="Ecological footprint">Ecological footprint</a></li>
<li><a href="Ecological_forecasting" title="Ecological forecasting">Ecological forecasting</a></li>
<li><a href="Environmental_humanities" title="Environmental humanities">Ecological humanities</a></li>
<li><a href="Ecological_stoichiometry" title="Ecological stoichiometry">Ecological stoichiometry</a></li>
<li><a href="Ecopath" title="Ecopath">Ecopath</a></li>
<li><a href="Ecosystem_based_fisheries" class="mw-redirect" title="Ecosystem based fisheries">Ecosystem based fisheries</a></li>
<li><a href="Endolith" title="Endolith">Endolith</a></li>
<li><a href="Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li>
<li><a href="Functional_ecology" title="Functional ecology">Functional ecology</a></li>
<li><a href="Industrial_ecology" title="Industrial ecology">Industrial ecology</a></li>
<li><a href="Macroecology" title="Macroecology">Macroecology</a></li>
<li><a href="Microecosystem" title="Microecosystem">Microecosystem</a></li>
<li><a href="Natural_environment" title="Natural environment">Natural environment</a></li>
<li><a href="Regime_shift" title="Regime shift">Regime shift</a></li>
<li><a href="Sexecology" title="Sexecology">Sexecology</a></li>
<li><a href="Systems_ecology" title="Systems ecology">Systems ecology</a></li>
<li><a href="Urban_ecology" title="Urban ecology">Urban ecology</a></li>
<li><a href="Theoretical_ecology" title="Theoretical ecology">Theoretical ecology</a></li></ul>
</div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="Outline_of_ecology" title="Outline of ecology">Outline of ecology</a></div></td></tr></tbody></table></div></div><!--htdig_noindex--><div><div class="zim-footer">
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