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searchaux" style="display:none">Ecological mechanism enabling species to coexist</div> <p>The <b>storage effect</b> is a coexistence mechanism proposed in the <a href="/wiki/Ecological_theory" class="mw-redirect" title="Ecological theory">ecological theory</a> of <a href="/wiki/Species" title="Species">species</a> coexistence, which tries to explain how such a wide variety of similar species are able to coexist within the same <a href="/wiki/Ecological_community" class="mw-redirect" title="Ecological community">ecological community</a> or <a href="/wiki/Guild_(ecology)" title="Guild (ecology)">guild</a>. The storage effect was originally proposed in the 1980s<sup id="cite_ref-Chesson_&_Warner_1981_1-0" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> to explain coexistence in diverse communities of coral reef fish, however it has since been generalized to cover a variety of ecological communities.<sup id="cite_ref-Pake_&_Venable_1995_2-0" class="reference"><a href="#cite_note-Pake_&_Venable_1995-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Caceres_1997_3-0" class="reference"><a href="#cite_note-Caceres_1997-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Kelly_&_Bowler_2002_4-0" class="reference"><a href="#cite_note-Kelly_&_Bowler_2002-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> The theory proposes one way for multiple species to coexist: in a changing environment, no species can be the best under all conditions.<sup id="cite_ref-Chesson_1994_5-0" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Instead, each species must have a unique response to varying environmental conditions, and a way of buffering against the effects of bad years.<sup id="cite_ref-Chesson_1994_5-1" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The storage effect gets its name because each population "stores" the gains in good years or microhabitats (patches) to help it survive population losses in bad years or patches.<sup id="cite_ref-Chesson_&_Warner_1981_1-1" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> One strength of this theory is that, unlike most coexistence mechanisms, the storage effect can be measured and quantified, with units of per-capita growth rate (offspring per adult per generation).<sup id="cite_ref-Chesson_1994_5-2" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>The storage effect can be caused by both temporal and spatial variation. The temporal storage effect (often referred to as simply "the storage effect") occurs when species benefit from changes in year-to-year environmental patterns,<sup id="cite_ref-Chesson_&_Warner_1981_1-2" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> while the spatial storage effect occurs when species benefit from variation in microhabitats across a landscape.<sup id="cite_ref-Chesson_2000space_6-0" class="reference"><a href="#cite_note-Chesson_2000space-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="The_concept">The concept</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=1" title="Edit section: The concept"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>For the storage effect to operate, it requires variation (i.e. fluctuations) in the environment<sup id="cite_ref-Chesson_&_Warner_1981_1-3" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> and thus can be termed a "fluctuation-dependent mechanism". This variation can come from a large degree of factors, including resource availability, temperature, and predation levels. However, for the storage effect to function, this variation must change the birth, survival, or <a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">recruitment</a> rate of species from year to year (or patch to patch).<sup id="cite_ref-Chesson_&_Warner_1981_1-4" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Chesson_2000space_6-1" class="reference"><a href="#cite_note-Chesson_2000space-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p><p>For competing species within the same community to coexist, they have to meet one fundamental requirement: the impact of competition from a species on itself must exceed its competitive impact on other species. In other words, <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">intraspecific competition</a> must exceed <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a>.<sup id="cite_ref-Chesson_2000review_7-0" class="reference"><a href="#cite_note-Chesson_2000review-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> For example, jackrabbits living in the same area compete for food and nesting grounds. Such competition within the same species is called <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">intraspecific competition</a>, which limits the growth of the species itself. Members from different species can also compete. For instance, jackrabbits and cottontail rabbits also compete for food and nesting grounds. Competition between different species is called <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a>, which limits the growth of other species. Stable coexistence occurs when any one species in the community limits its own growth more strongly than the growth of others.<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (August 2023)">citation needed</span></a></i>]</sup> </p><p>The storage effect mixes three essential ingredients to assemble a community of competing species that fulfill the requirement. They are 1) correlation between the quality of an environment and the amount of competition experienced by a population in that environment (i.e. <a href="/wiki/Covariance" title="Covariance">covariance</a> between environment and competition), 2) differences in species response to the same environment (i.e. species-specific environmental responses), and 3) the ability of a population to diminish the impact of competition under worsening environment (i.e. buffered population growth).<sup id="cite_ref-Chesson_1994_5-3" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Each ingredient is described in detail below with an explanation why the combination of the three leads to species coexistence. </p> <div class="mw-heading mw-heading3"><h3 id="Covariance_between_environment_and_competition">Covariance between environment and competition</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=2" title="Edit section: Covariance between environment and competition"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The growth of a population can be strongly influenced by the environment it experiences. An environment consists of not only physical elements such as resource abundance, temperature, and level of physical disturbance, but also biological elements such as the abundance of <a href="/wiki/Predation" title="Predation">natural enemies</a> and <a href="/wiki/Mutualists" class="mw-redirect" title="Mutualists">mutualists</a>.<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> Usually organisms reproduce more in a favorable environment (i.e. either during a good year, or within a good patch), build up their population densities, and lead themselves to a high level of <a href="/wiki/Competition_(biology)" title="Competition (biology)">competition</a> due to this increasing crowding.<sup id="cite_ref-Chesson_1994_5-4" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Such a trend means that higher quality environments usually correlate with a higher strength of competition experienced by the organisms in those environments. In short, a better environment results in stronger competition.<sup id="cite_ref-Chesson_1994_5-5" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> In statistics, such <a href="/wiki/Correlation" title="Correlation">correlation</a> means that there will be a non-zero <a href="/wiki/Covariance" title="Covariance">covariance</a> between the change of population density in response to the environment and that to the competition. That is why the first ingredient is called "<a href="/wiki/Covariance" title="Covariance">covariance</a> between environment and competition".<sup class="noprint Inline-Template Template-Fact" style="white-space:nowrap;">[<i><a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed"><span title="This claim needs references to reliable sources. (August 2023)">citation needed</span></a></i>]</sup> </p> <div class="mw-heading mw-heading3"><h3 id="Species-specific_environmental_responses">Species-specific environmental responses</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=3" title="Edit section: Species-specific environmental responses"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Covariance between environment and competition suggests that organisms experience the strongest <a href="/wiki/Competition_(biology)" title="Competition (biology)">competition</a> under their optimal environmental conditions because their populations grow most rapidly in those conditions. In nature, we often find that different species from the same community respond to the same conditions in distinctive manners. For example, plant species have different preferred levels of light and water availability, which affect their germination and physical growth rates.<sup id="cite_ref-Pake_&_Venable_1995_2-1" class="reference"><a href="#cite_note-Pake_&_Venable_1995-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Such differences in their response to the environment, which is called "species-specific environmental response," means no two species from a community will have the same best environment in a given year or a given patch. As a result, when a species is under its optimal environmental conditions and thus experiencing the strongest <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">intraspecific competition</a>, other species from the same community only experience the strongest <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a> coming from that species, but not the strongest <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">intraspecific competition</a> coming from themselves. </p> <div class="mw-heading mw-heading3"><h3 id="Buffered_population_growth">Buffered population growth</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=4" title="Edit section: Buffered population growth"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A population can decline when environmental conditions worsen and when competition intensifies. If a species cannot limit the impact of competition in a hostile environment, its population will crash, and it will become locally extinct.<sup id="cite_ref-Chesson_&_Warner_1981_1-5" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Marvelously, in nature organisms are often able to slow down the rate of population decline in a hostile environment by alleviating the impact of competition. In doing so, they are able to set up a lower limit on the rate of their population decline.<sup id="cite_ref-Chesson_&_Warner_1981_1-6" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This phenomenon is called "buffered population growth", which occurs under a variety of situations. Under the temporal storage effect, it can be accomplished by the adults of a species having long life spans, which are relatively unaffected by environmental stressors. For example, an adult tree is unlikely to be killed by a few weeks of drought or a single night of freezing temperatures, whereas a seedling may not survive these conditions.<sup id="cite_ref-Kelly_&_Bowler_2002_4-1" class="reference"><a href="#cite_note-Kelly_&_Bowler_2002-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Even if all seedlings are killed by bad environmental conditions, the long-lived adults are able to keep the overall population from crashing.<sup id="cite_ref-Chesson_&_Warner_1981_1-7" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Moreover, the adults usually adopt strategies such as dormancy or hibernation under a hostile environment, which make them less sensitive to competition, and allows them to buffer against the double blades of the hostile environment and competition from their rivals. For a different example, buffered population growth is attained by annual plants with a persistent seed bank.<sup id="cite_ref-Pake_&_Venable_1995_2-2" class="reference"><a href="#cite_note-Pake_&_Venable_1995-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Thanks to these long-lived seeds, the entire population cannot be destroyed by a single bad year. Moreover, the seeds stay dormant under unfavorable environmental conditions, avoiding direct competition with rivals who are favored by the same environment, and thus diminish the impact of competition in bad years.<sup id="cite_ref-Pake_&_Venable_1995_2-3" class="reference"><a href="#cite_note-Pake_&_Venable_1995-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> There are some temporal situations in which buffered population growth is not expected to occur. Namely, when multiple generations do not overlap (such as <a href="/wiki/Furcifer_labordi" class="mw-redirect" title="Furcifer labordi">Labord's chameleon</a>) or when adults have a high mortality rate (such as many aquatic insects, or some populations of the <a href="/wiki/Eastern_Fence_Lizard" class="mw-redirect" title="Eastern Fence Lizard">Eastern Fence Lizard</a><sup id="cite_ref-Tinkle_&_Ballinger_1972_9-0" class="reference"><a href="#cite_note-Tinkle_&_Ballinger_1972-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup>), buffered growth does not occur. Under the spatial storage effect, buffered population growth is generally automatic, because the effects of a detrimental <a href="/wiki/Microhabitat" class="mw-redirect" title="Microhabitat">microhabitat</a> will only be experienced by individuals in that area, rather than the population as a whole.<sup id="cite_ref-Chesson_2000space_6-2" class="reference"><a href="#cite_note-Chesson_2000space-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Outcome">Outcome</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=5" title="Edit section: Outcome"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The combined effect of (1) <a href="/wiki/Covariance" title="Covariance">covariance</a> between environment and competition, and (2) species-specific response to the environment decouple the strongest intraspecific and interspecific competition experienced by a species.<sup id="cite_ref-Chesson_1994_5-6" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">Intraspecific competition</a> is strongest when a species is favored by the environment, whereas <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a> is strongest when its rivals are favored. After this decoupling, buffered population growth limits the impact of <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a> when a species is not favored by the environment. As a consequence, the impact of <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">intraspecific competition</a> on the species favored by a particular environment exceeds that of the <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a> on species less favored by that environment. We see that the fundamental requirement for species coexistence is fulfilled and thus storage effect is able to maintain stable coexistence in a community of competing species.<sup id="cite_ref-Chesson_1994_5-7" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p><p>For species to coexist in a community, all species must be able to recover from low density.<sup id="cite_ref-Chesson_2000review_7-1" class="reference"><a href="#cite_note-Chesson_2000review-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> Not surprisingly, being a coexistence mechanism, the storage effect helps species when they become rare. It does so by making the abundant species’ effect on itself greater than its effect on the rare species.<sup id="cite_ref-Chesson_1994_5-8" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The difference between species’ response to environmental conditions means that a rare species’ optimal environment is not the same as its competitors. Under these conditions, the rare species will experience low levels of <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a>. Because the rare species itself is rare, it will experience little impact from <a href="/wiki/Intraspecific_competition" title="Intraspecific competition">intraspecific competition</a> as well, even at its highest possible levels of intraspecific competition. Free from the impact of competition, the rare species is able to make gains in these good years or patches.<sup id="cite_ref-Chesson_&_Warner_1981_1-8" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Moreover, thanks to the buffered population growth, the rare species is able to survive the bad years or patches by "storing" the gains from the good years/patches. As a result, the population of any rare species is able to grow due to the storage effect. </p><p>One natural outcome from the <a href="/wiki/Covariance" title="Covariance">covariance</a> between environment and competition is that the species with very low densities will have more fluctuation in its <a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">recruitment</a> rates than species with normal densities.<sup id="cite_ref-Sears_&_Chesson_2007_10-0" class="reference"><a href="#cite_note-Sears_&_Chesson_2007-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> This occurs because in good environments, species with high densities will often experience large amount of crowding by members of the same species, thus limiting the benefits of good years/patches, and making good years/patches more similar to bad years/patches. Low-density species are rarely able to cause crowding, thus allowing significantly increased fitness in good years/patches.<sup id="cite_ref-Chesson_&_Warner_1981_1-9" class="reference"><a href="#cite_note-Chesson_&_Warner_1981-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> Since the fluctuation in <a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">recruitment</a> rate is an indicator of covariance between environment and competition, and since species-specific environmental response and buffered population growth can normally be assumed in nature, finding much stronger fluctuation in <a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">recruitment</a> rates in rare and low-density species provides a strong indication that the storage effect is operating within a community.<sup id="cite_ref-Kelly_&_Bowler_2002_4-2" class="reference"><a href="#cite_note-Kelly_&_Bowler_2002-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Sears_&_Chesson_2007_10-1" class="reference"><a href="#cite_note-Sears_&_Chesson_2007-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Mathematical_formulation">Mathematical formulation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=6" title="Edit section: Mathematical formulation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The storage effect is not a model for population growth (such as the <a href="/wiki/Lotka%E2%80%93Volterra_equation" class="mw-redirect" title="Lotka–Volterra equation">Lotka–Volterra equation</a>) itself, but is an effect that appears in non-additive models of population growth.<sup id="cite_ref-Chesson_1994_5-9" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> Thus, the equations shown below will work for any arbitrary model of population growth, but will only be as accurate as the original model. The derivation below is taken from Chesson 1994.<sup id="cite_ref-Chesson_1994_5-10" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> It is a derivation of the temporal storage effect, but is very similar to the spatial storage effect. </p><p>The <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> of an individual, as well as expected <a href="/wiki/Population_growth" title="Population growth">growth rate</a>, can be measured in terms of the average number of offspring it will leave during its lifetime. This parameter, <i>r</i>(<i>t</i>), is a function of both environmental factors, <i>e</i>(<i>t</i>), and how much the organism must <a href="/wiki/Competition_(biology)" title="Competition (biology)">compete</a> with other individuals (both of its own species, and different species), <i>c</i>(<i>t</i>). Thus, </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle r(t)=g(e(t),c(t))\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>r</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>=</mo> <mi>g</mi> <mo stretchy="false">(</mo> <mi>e</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>,</mo> <mi>c</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo stretchy="false">)</mo> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle r(t)=g(e(t),c(t))\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/4bbf989a9ca2c0427407edb32dbbf30b7e32f734" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:18.531ex; height:2.843ex;" alt="{\displaystyle r(t)=g(e(t),c(t))\,}"></span></dd></dl> <p>where <i>g</i> is an arbitrary function for growth rate. Throughout the article, subscripts are occasionally used to represent functions of a particular species (e.g. <i>r</i><sub> <i>j</i></sub>(<i>t</i>) is the fitness of species <i>j</i>). It is assumed that there must be some values <i>e*</i> and <i>c*</i>, such that <i>g</i>(<i>e</i>*, <i>c</i>*) = 0, representing a zero-population growth equilibrium. These values need not be unique, but for every <i>e*</i>, there is a unique <i>c*</i>. For ease of calculation, standard parameters <i>E</i>(<i>t</i>) and <i>C</i>(<i>t</i>) are defined, such that </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle E(t)=g(e(t),c^{*})\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>E</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>=</mo> <mi>g</mi> <mo stretchy="false">(</mo> <mi>e</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>,</mo> <msup> <mi>c</mi> <mrow class="MJX-TeXAtom-ORD"> <mo>∗</mo> </mrow> </msup> <mo stretchy="false">)</mo> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle E(t)=g(e(t),c^{*})\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/1bab9bfdf8682cd85e5bba3704866e216fc5d02f" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:17.663ex; height:2.843ex;" alt="{\displaystyle E(t)=g(e(t),c^{*})\,}"></span></dd></dl> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle C(t)=-g(e^{*},c(t))\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>C</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>=</mo> <mo>−</mo> <mi>g</mi> <mo stretchy="false">(</mo> <msup> <mi>e</mi> <mrow class="MJX-TeXAtom-ORD"> <mo>∗</mo> </mrow> </msup> <mo>,</mo> <mi>c</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo stretchy="false">)</mo> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle C(t)=-g(e^{*},c(t))\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/e26a7e91cc2a41e362bcb69db3eb5a28846d9ffd" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:19.462ex; height:2.843ex;" alt="{\displaystyle C(t)=-g(e^{*},c(t))\,}"></span></dd></dl> <p>Both <i>E</i> and <i>C</i> represent the effect of deviations in environmental response from equilibrium. <i>E</i> represents the effect that varying environmental conditions (e.g. rainfall patterns, temperature, food availability, etc.) have on fitness, in the absence of abnormal competitive effects. For the storage effect to occur, the environmental response for each species must be unique (i.e. <i>E</i><sub> <i>j</i></sub>(<i>t</i>) ≠ <i>E</i><sub> <i>i</i></sub>(<i>t</i>) when <i>j</i> ≠ <i>i</i>). <i>C</i>(<i>t</i>) represents how much average fitness is lowered as a result of <a href="/wiki/Competition_(biology)" title="Competition (biology)">competition</a>. For example, if there is more rain during a given year, <i>E</i>(<i>t</i>) will likely increase. If more plants begin to bloom, and thus compete for that rain, then <i>C</i>(<i>t</i>) will increase as well. Because <i>e*</i> and <i>c*</i> are not unique, <i>E</i>(<i>t</i>) and <i>C</i>(<i>t</i>) are not unique, and thus one should choose them as conveniently as possible. Under most conditions (see Chesson 1994<sup id="cite_ref-Chesson_1994_5-11" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>), <i>r</i>(<i>t</i>) can be approximated as </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle r(t)=E(t)-C(t)+\gamma E(t)C(t)\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>r</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>=</mo> <mi>E</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>−</mo> <mi>C</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>+</mo> <mi>γ</mi> <mi>E</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mi>C</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle r(t)=E(t)-C(t)+\gamma E(t)C(t)\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/66f6c65b0199ac475e2e89a55a916b1bd72bf0ae" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:31.806ex; height:2.843ex;" alt="{\displaystyle r(t)=E(t)-C(t)+\gamma E(t)C(t)\,}"></span></dd></dl> <p>where </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \gamma ={\frac {\partial ^{2}r}{\partial E\,\partial C}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>γ</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <msup> <mi mathvariant="normal">∂</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msup> <mi>r</mi> </mrow> <mrow> <mi mathvariant="normal">∂</mi> <mi>E</mi> <mspace width="thinmathspace" /> <mi mathvariant="normal">∂</mi> <mi>C</mi> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \gamma ={\frac {\partial ^{2}r}{\partial E\,\partial C}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/07c89c8bf7e6da747689f02bb0dd99aa8716f96b" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.005ex; width:11.762ex; height:5.843ex;" alt="{\displaystyle \gamma ={\frac {\partial ^{2}r}{\partial E\,\partial C}}}"></span></dd></dl> <p><i>γ</i> represents the nonadditivity of growth rates. If <i>γ</i> = 0 (known as <a href="/wiki/Additive_map" title="Additive map">additivity</a>) it means that the impact of competition on fitness does not change with the environment. If <i>γ</i> > 0 (<a href="/wiki/Superadditivity" title="Superadditivity">superadditivity</a>), it means that the adverse effects of competition during a bad year are relatively worse than during a good year. In other words, a population suffers more from competition in bad years than in good years. If <i>γ</i> < 0 (<a href="/wiki/Subadditivity" title="Subadditivity">subadditivity</a>, or buffered population growth), it means that the harm done by competition during a bad year is relatively minor when compared to a good year. In other words, the population is able to diminish the impact of competition as the environment worsens. As stated above, for the storage effect to contribute to species coexistence, we must have buffered population growth (i.e. it must be the case that <i>γ</i> < 0). </p><p>The long-term average of the above equation is </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\bar {r}}={\bar {E}}-{\bar {C}}+\gamma ({\bar {E}}{\bar {C}}+{\text{Cov}}(E(t),C(t)))\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>r</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>E</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>−</mo> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>C</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>+</mo> <mi>γ</mi> <mo stretchy="false">(</mo> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>E</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>C</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>+</mo> <mrow class="MJX-TeXAtom-ORD"> <mtext>Cov</mtext> </mrow> <mo stretchy="false">(</mo> <mi>E</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>,</mo> <mi>C</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo stretchy="false">)</mo> <mo stretchy="false">)</mo> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\bar {r}}={\bar {E}}-{\bar {C}}+\gamma ({\bar {E}}{\bar {C}}+{\text{Cov}}(E(t),C(t)))\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/80d57bca9db951a805f50c695b729f6306713c85" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:39.205ex; height:3.176ex;" alt="{\displaystyle {\bar {r}}={\bar {E}}-{\bar {C}}+\gamma ({\bar {E}}{\bar {C}}+{\text{Cov}}(E(t),C(t)))\,}"></span></dd></dl> <p>which, under environments with sufficient variation relative to mean effects, can be approximated as </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\bar {r}}\approx {\bar {E}}-{\bar {C}}+\gamma {\text{Cov}}(E(t),C(t))}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>r</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>≈</mo> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>E</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>−</mo> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <mi>C</mi> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>+</mo> <mi>γ</mi> <mrow class="MJX-TeXAtom-ORD"> <mtext>Cov</mtext> </mrow> <mo stretchy="false">(</mo> <mi>E</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo>,</mo> <mi>C</mi> <mo stretchy="false">(</mo> <mi>t</mi> <mo stretchy="false">)</mo> <mo stretchy="false">)</mo> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\bar {r}}\approx {\bar {E}}-{\bar {C}}+\gamma {\text{Cov}}(E(t),C(t))}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/2a38255532b3a0ae09597325940941126090cfbf" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.838ex; width:30.626ex; height:3.176ex;" alt="{\displaystyle {\bar {r}}\approx {\bar {E}}-{\bar {C}}+\gamma {\text{Cov}}(E(t),C(t))}"></span></dd></dl> <p>For any effect to act as a coexistence mechanism, it must boost the average fitness of an individual when they are at below-normal population density. Otherwise, a species at low density (known as an `invader') will continue to dwindle, and this negative feedback will cause its extinction. When a species is at equilibrium (known as a `resident'), its average long-term fitness must be 0. For a species to recover from low density, its average fitness must be greater than 0. For the remainder of the text, we refer to functions of the invader with the subscript <i>i</i>, and to the resident with the subscript <i>r</i>. </p><p>A long-term average growth rate of an invader is often written as </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle {\bar {r_{i}}}=\Delta E-\Delta C+\Delta I\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <msub> <mi>r</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>=</mo> <mi mathvariant="normal">Δ</mi> <mi>E</mi> <mo>−</mo> <mi mathvariant="normal">Δ</mi> <mi>C</mi> <mo>+</mo> <mi mathvariant="normal">Δ</mi> <mi>I</mi> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle {\bar {r_{i}}}=\Delta E-\Delta C+\Delta I\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/40b9f1364a648343fd8b4d2676466b88664347a1" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.671ex; width:21.536ex; height:2.509ex;" alt="{\displaystyle {\bar {r_{i}}}=\Delta E-\Delta C+\Delta I\,}"></span></dd></dl> <p>where, </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Delta E={\bar {E_{i}}}-\sum _{i\neq r}q_{ir}{\bar {E_{r}}}\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi mathvariant="normal">Δ</mi> <mi>E</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <msub> <mi>E</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>−</mo> <munder> <mo>∑</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mo>≠</mo> <mi>r</mi> </mrow> </munder> <msub> <mi>q</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>r</mi> </mrow> </msub> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <msub> <mi>E</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Delta E={\bar {E_{i}}}-\sum _{i\neq r}q_{ir}{\bar {E_{r}}}\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/8eedb2b3669bcda996119c0a06638ce1c2028e69" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.505ex; width:21.562ex; height:6.009ex;" alt="{\displaystyle \Delta E={\bar {E_{i}}}-\sum _{i\neq r}q_{ir}{\bar {E_{r}}}\,}"></span></dd></dl> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Delta C={\bar {C_{i}}}-\sum _{i\neq r}q_{ir}{\bar {C_{r}}}\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi mathvariant="normal">Δ</mi> <mi>C</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mo>−</mo> <munder> <mo>∑</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mo>≠</mo> <mi>r</mi> </mrow> </munder> <msub> <mi>q</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>r</mi> </mrow> </msub> <mrow class="MJX-TeXAtom-ORD"> <mrow class="MJX-TeXAtom-ORD"> <mover> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <mo stretchy="false">¯</mo> </mover> </mrow> </mrow> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Delta C={\bar {C_{i}}}-\sum _{i\neq r}q_{ir}{\bar {C_{r}}}\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/f634246743782d051f319cec148affa5a4743317" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.505ex; width:21.446ex; height:6.009ex;" alt="{\displaystyle \Delta C={\bar {C_{i}}}-\sum _{i\neq r}q_{ir}{\bar {C_{r}}}\,}"></span></dd></dl> <p>and, ΔI, the storage effect, </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle \Delta I=\gamma _{i}{\text{Cov}}(E_{i},C_{i})-\sum _{i\neq r}q_{ir}\gamma _{r}{\text{Cov}}(E_{r},C_{r})\,}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi mathvariant="normal">Δ</mi> <mi>I</mi> <mo>=</mo> <msub> <mi>γ</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mrow class="MJX-TeXAtom-ORD"> <mtext>Cov</mtext> </mrow> <mo stretchy="false">(</mo> <msub> <mi>E</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo>,</mo> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> <mo stretchy="false">)</mo> <mo>−</mo> <munder> <mo>∑</mo> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mo>≠</mo> <mi>r</mi> </mrow> </munder> <msub> <mi>q</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>r</mi> </mrow> </msub> <msub> <mi>γ</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <mrow class="MJX-TeXAtom-ORD"> <mtext>Cov</mtext> </mrow> <mo stretchy="false">(</mo> <msub> <mi>E</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <mo>,</mo> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> <mo stretchy="false">)</mo> <mspace width="thinmathspace" /> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \Delta I=\gamma _{i}{\text{Cov}}(E_{i},C_{i})-\sum _{i\neq r}q_{ir}\gamma _{r}{\text{Cov}}(E_{r},C_{r})\,}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/f18dca4d9434a781582b61cdfba2644d1f4993a3" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.505ex; width:44.059ex; height:6.009ex;" alt="{\displaystyle \Delta I=\gamma _{i}{\text{Cov}}(E_{i},C_{i})-\sum _{i\neq r}q_{ir}\gamma _{r}{\text{Cov}}(E_{r},C_{r})\,}"></span></dd></dl> <p>where </p> <dl><dd><span class="mwe-math-element"><span class="mwe-math-mathml-inline mwe-math-mathml-a11y" style="display: none;"><math xmlns="http://www.w3.org/1998/Math/MathML" alttext="{\displaystyle q_{ir}={\frac {\partial C_{i}}{\partial C_{r}}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>q</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> <mi>r</mi> </mrow> </msub> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <mi mathvariant="normal">∂</mi> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>i</mi> </mrow> </msub> </mrow> <mrow> <mi mathvariant="normal">∂</mi> <msub> <mi>C</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>r</mi> </mrow> </msub> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle q_{ir}={\frac {\partial C_{i}}{\partial C_{r}}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/9204124c4db4f2b05b24a6c9fd7d26d8f3abb3c6" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.338ex; width:10.466ex; height:5.843ex;" alt="{\displaystyle q_{ir}={\frac {\partial C_{i}}{\partial C_{r}}}}"></span></dd></dl> <p>In this equation, <i>q</i><sub><i>ir</i></sub> tells us how much the competition experienced by <i>r</i> affects the competition experienced by <i>i</i>. </p><p>The biological meaning of the storage effect is expressed in the mathematical form of ΔI. The first term of the expression is <a href="/wiki/Covariance" title="Covariance">covariance</a> between environment and competition (Cov(<i>E</i> <i>C</i>)), scaled by a factor representing buffered population growth (<i>γ</i>). The difference between the first term and the second term represents the difference in species responses to the environment between the invader and the sum of the residents, scaled by the effect each resident has on the invader (<i>q</i><sub><i>ir</i></sub>).<sup id="cite_ref-Chesson_1994_5-12" class="reference"><a href="#cite_note-Chesson_1994-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Predation">Predation</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=7" title="Edit section: Predation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Recent work has extended what is known about the storage effect to include <a href="/wiki/Apparent_competition" class="mw-redirect" title="Apparent competition">apparent competition</a> (i.e., competition mediated through a shared predator). </p><p>These models showed that generalist predators can undermine the benefits of the storage effect that from competition.<sup id="cite_ref-Kuang_&_Chesson_2009_11-0" class="reference"><a href="#cite_note-Kuang_&_Chesson_2009-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> This occurs because generalist predators depress population levels by eating individuals. When this happens, there are fewer individuals competing for resources. As a result, relatively abundant species are less constrained by competition for resource in favorable years (i.e., the covariance between environment and competition is weakened), and therefore the storage effect from competition is weakened. This conclusion follows the general trend that the introduction of a generalist predator will often weaken other competition-based coexistence mechanisms, and which result in <a href="/wiki/Competitive_exclusion" class="mw-redirect" title="Competitive exclusion">competitive exclusion</a>.<sup id="cite_ref-Chesson_&_Kuang_2008_nature_12-0" class="reference"><a href="#cite_note-Chesson_&_Kuang_2008_nature-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Holt_1977_13-0" class="reference"><a href="#cite_note-Holt_1977-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> </p><p>Additionally, certain types of predators can produce a storage effect from predation. This effect has been shown for <a href="/wiki/Prey_switching" title="Prey switching">frequency-dependent predators</a>, who are more likely to attack prey that are abundant,<sup id="cite_ref-Chesson_&_Kuang_2010_multi_14-0" class="reference"><a href="#cite_note-Chesson_&_Kuang_2010_multi-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> and for generalist pathogens, who cause outbreaks when prey are abundant.<sup id="cite_ref-Mordecai_2011_15-0" class="reference"><a href="#cite_note-Mordecai_2011-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> When prey species are especially numerous and active, frequency-dependent predators become more active, and pathogens outbreaks become more severe (i.e., there was a positive covariance between the environment and predation, analogous to the covariance between the environment and competition). As a result, abundant species are limited during their best years by high predation – an effect that is analogous to the storage effect from competition. </p> <div class="mw-heading mw-heading2"><h2 id="Empirical_studies">Empirical studies</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Storage_effect&action=edit&section=8" title="Edit section: Empirical studies"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The first empirical study that tested the requirements of the storage effect was done by Pake and Venable,<sup id="cite_ref-Pake_&_Venable_1995_2-4" class="reference"><a href="#cite_note-Pake_&_Venable_1995-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> who looked at three desert <a href="/wiki/Annual_plant" title="Annual plant">annual plants</a>. They experimentally manipulated density and water availability over a two-year period, and found that <a href="/wiki/Fitness_(biology)" title="Fitness (biology)">fitness</a> and <a href="/wiki/Germination" title="Germination">germination</a> rates varied greatly from year to year, and over different environmental conditions. This shows that each species has a unique environmental response, and implied that likely there is a <a href="/wiki/Covariance" title="Covariance">covariance</a> between environment and competition. This, combined with the buffered population growth that is a product of a long-lived <a href="/wiki/Soil_seed_bank" title="Soil seed bank">seed bank</a>, showed that a temporal storage effect was probably an important factor in mediating coexistence. This study was also important, because it showed that variation in <a href="/wiki/Germination" title="Germination">germination</a> conditions could be a major factor promoting species coexistence.<sup id="cite_ref-Pake_&_Venable_1995_2-5" class="reference"><a href="#cite_note-Pake_&_Venable_1995-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>The first attempt made at quantifying the temporal storage effect was by <a href="/wiki/Carla_C%C3%A1ceres" title="Carla Cáceres">Carla Cáceres</a> in 1997.<sup id="cite_ref-Caceres_1997_3-1" class="reference"><a href="#cite_note-Caceres_1997-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Using 30 years of water-column data from <a href="/wiki/Oneida_Lake" title="Oneida Lake">Oneida Lake</a>, New York, she studied the effect the storage effect had on two species of <a href="/wiki/Daphnia" title="Daphnia">plankton</a> (<a href="/wiki/Daphnia_galeata_mendotae" class="mw-redirect" title="Daphnia galeata mendotae">Daphnia galeata mendotae</a> and <i>D. pulicaria</i>). These species of plankton lay <a href="/wiki/Diapause" title="Diapause">diapausing</a> eggs which, much like the seeds of annual plants, lay dormant in the sediment for many years before hatching. Cáceres found that the size of reproductive bouts were fairly uncorrelated between the two species. She also found, in the absence of the storage effect, D. galeata mendotae would have gone extinct. She was unable to measure certain important parameters (such as the rate of egg predation), but found that her results were robust to a wide range of estimates.<sup id="cite_ref-Caceres_1997_3-2" class="reference"><a href="#cite_note-Caceres_1997-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>The first test of the spatial storage effect was done by Sears and Chesson <sup id="cite_ref-Sears_&_Chesson_2007_10-2" class="reference"><a href="#cite_note-Sears_&_Chesson_2007-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> in the desert area east of Portal, Arizona. Using a common neighbor-removal experiment, they examined whether coexistence between two annual plants, <a href="/wiki/Erodium_cicutarium" title="Erodium cicutarium">Erodium cicutarium</a> and Phacelia popeii, was due to the spatial storage effect or resource partitioning. The storage effect was quantified in terms of number of inflorescences (a proxy for fitness) instead of actual population growth rate. They found that E. cicutarium was able to outcompete P. popeii in many situations, and in the absence of the storage effect, would likely competitively exclude P. popeii. However, they found a very strong difference in the <a href="/wiki/Covariance" title="Covariance">covariance</a> between environment and competition, which showed that some of the most favorable areas for P. popeii (the rare species), were unfavorable to E. cicutarium (the common species). This suggests that P. popeii is able to avoid strong <a href="/wiki/Interspecific_competition" title="Interspecific competition">interspecific competition</a> in some good patches, and that this may be enough to compensate for losses in areas favorable to E. cicutarium.<sup id="cite_ref-Sears_&_Chesson_2007_10-3" class="reference"><a href="#cite_note-Sears_&_Chesson_2007-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> </p><p>Colleen Kelly and colleagues have used congeneric species pairs to examine storage dynamics where species similarity is a natural outcome of relatedness and not dependent on researcher-based estimates. Initial studies were of 12 species of trees coexisting in a tropical deciduous forest at the Chamela Biological Station in <a href="/wiki/Jalisco" title="Jalisco">Jalisco</a>, Mexico.<sup id="cite_ref-Kelly_&_Bowler_2002_4-3" class="reference"><a href="#cite_note-Kelly_&_Bowler_2002-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> For each of the 12 species they examined age structure (calculated from size and species-specific growth rate), and found that <a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">recruitment</a> of young trees varies from year to year. Grouping the species into 6 congeneric pairs, the locally rarer species of each pair unanimously had a more irregular <a href="/wiki/Age_distribution" class="mw-redirect" title="Age distribution">age distributions</a> than the more common species. This finding strongly suggests that between closely competing tree species, the rarer species experiences stronger <a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">recruitment</a> fluctuation than the commoner species. Such difference in <a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">recruitment</a> fluctuation, combined with evidence of greater competitive ability in the rarer species of each pair, indicates a difference in <a href="/wiki/Covariance" title="Covariance">covariance</a> between the environment and competition between rare and common species. Since species-specific environmental response and buffered population growth can be naturally assumed, their finding strongly suggests that the storage effect operates in this tropical deciduous forest so as to maintain the coexistence between different tree species. Further work with these species has shown that the storage dynamic is a pairwise, competitive relationship, between congeneric species pairs, and possibly extending as successively nested pairs within a genus.<sup id="cite_ref-Kelly_&_Bowler_2005_17-0" class="reference"><a href="#cite_note-Kelly_&_Bowler_2005-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> </p><p>Angert and colleagues demonstrated the temporal storage effect occurring in the desert annual plant community on <a href="/wiki/Tumamoc_Hill" title="Tumamoc Hill">Tumamoc Hill</a>, Arizona.<sup id="cite_ref-Angert_etal_2009_18-0" class="reference"><a href="#cite_note-Angert_etal_2009-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Previous studies<sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> had shown the <a href="/wiki/Annual_plant" title="Annual plant">annual plants</a> in that community exhibited a trade-off between growth rate (a proxy for competitive ability) and water use efficiency (a proxy for drought tolerance). As a result, some plants grew better during wet years, while others grew better during dry years. This, combined with variation in <a href="/wiki/Germination" title="Germination">germination</a> rates, produced an overall community average storage effect of 0.103. In other words, the storage effect is expected to help the population of any species at low density to increase, on average, by 10.3% each generation, until it recovers from low density.<sup id="cite_ref-Angert_etal_2009_18-1" class="reference"><a href="#cite_note-Angert_etal_2009-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <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=Storage_effect&action=edit&section=9" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-columns references-column-width reflist-columns-2"> <ol class="references"> <li id="cite_note-Chesson_&_Warner_1981-1"><span class="mw-cite-backlink">^ <a href="#cite_ref-Chesson_&_Warner_1981_1-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Chesson_&_Warner_1981_1-9"><sup><i><b>j</b></i></sup></a></span> <span class="reference-text"><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="CITEREFChessonWarner,_Robert1981" class="citation journal cs1">Chesson, Peter; Warner, Robert (1981). "Environmental variability promotes coexistence in lottery competitive systems". <i>The American Naturalist</i>. <b>117</b> (6): 923–943. <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%2F283778">10.1086/283778</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:84164336">84164336</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=Environmental+variability+promotes+coexistence+in+lottery+competitive+systems&rft.volume=117&rft.issue=6&rft.pages=923-943&rft.date=1981&rft_id=info%3Adoi%2F10.1086%2F283778&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A84164336%23id-name%3DS2CID&rft.aulast=Chesson&rft.aufirst=Peter&rft.au=Warner%2C+Robert&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Pake_&_Venable_1995-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-Pake_&_Venable_1995_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Pake_&_Venable_1995_2-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Pake_&_Venable_1995_2-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Pake_&_Venable_1995_2-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Pake_&_Venable_1995_2-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Pake_&_Venable_1995_2-5"><sup><i><b>f</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPakeVenable,_D._Lawerance1995" class="citation journal cs1">Pake, Catherine; Venable, D. Lawerance (1995). "Is coexistence of Sonoran desert annual plants mediated by temporal variability reproductive success". <i>Ecology</i>. <b>76</b> (1): 246–261. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2307%2F1940646">10.2307/1940646</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/1940646">1940646</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecology&rft.atitle=Is+coexistence+of+Sonoran+desert+annual+plants+mediated+by+temporal+variability+reproductive+success&rft.volume=76&rft.issue=1&rft.pages=246-261&rft.date=1995&rft_id=info%3Adoi%2F10.2307%2F1940646&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F1940646%23id-name%3DJSTOR&rft.aulast=Pake&rft.aufirst=Catherine&rft.au=Venable%2C+D.+Lawerance&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Caceres_1997-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-Caceres_1997_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Caceres_1997_3-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Caceres_1997_3-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCáceres1997" class="citation journal cs1"><a href="/wiki/Carla_C%C3%A1ceres" title="Carla Cáceres">Cáceres, Carla</a> (1997). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC23092">"Temoral variation, dormancy, and coexistence: a field test of the storage effect"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>94</b> (17): 9171–9175. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1997PNAS...94.9171C">1997PNAS...94.9171C</a>. <a href="/wiki/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.94.17.9171">10.1073/pnas.94.17.9171</a></span>. <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/PMC23092">23092</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/11038565">11038565</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Temoral+variation%2C+dormancy%2C+and+coexistence%3A+a+field+test+of+the+storage+effect&rft.volume=94&rft.issue=17&rft.pages=9171-9175&rft.date=1997&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC23092%23id-name%3DPMC&rft_id=info%3Apmid%2F11038565&rft_id=info%3Adoi%2F10.1073%2Fpnas.94.17.9171&rft_id=info%3Abibcode%2F1997PNAS...94.9171C&rft.aulast=C%C3%A1ceres&rft.aufirst=Carla&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC23092&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Kelly_&_Bowler_2002-4"><span class="mw-cite-backlink">^ <a href="#cite_ref-Kelly_&_Bowler_2002_4-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Kelly_&_Bowler_2002_4-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Kelly_&_Bowler_2002_4-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Kelly_&_Bowler_2002_4-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKellyBowler,_Michael2002" class="citation journal cs1">Kelly, Colleen; Bowler, Michael (2002). "Coexistence and relative abundance in forest trees". <i>Nature</i>. <b>417</b> (6887): 437–440. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2002Natur.417..437K">2002Natur.417..437K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F417437a">10.1038/417437a</a>. <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/12024212">12024212</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:4318307">4318307</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Coexistence+and+relative+abundance+in+forest+trees&rft.volume=417&rft.issue=6887&rft.pages=437-440&rft.date=2002&rft_id=info%3Adoi%2F10.1038%2F417437a&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4318307%23id-name%3DS2CID&rft_id=info%3Apmid%2F12024212&rft_id=info%3Abibcode%2F2002Natur.417..437K&rft.aulast=Kelly&rft.aufirst=Colleen&rft.au=Bowler%2C+Michael&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Chesson_1994-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-Chesson_1994_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-3"><sup><i><b>d</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-4"><sup><i><b>e</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-5"><sup><i><b>f</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-6"><sup><i><b>g</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-7"><sup><i><b>h</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-8"><sup><i><b>i</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-9"><sup><i><b>j</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-10"><sup><i><b>k</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-11"><sup><i><b>l</b></i></sup></a> <a href="#cite_ref-Chesson_1994_5-12"><sup><i><b>m</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChesson1994" class="citation journal cs1">Chesson, Peter (1994). "Multispecies competition in variable environments". <i>Theoretical Population Biology</i>. <b>45</b> (3): 227–276. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1006%2Ftpbi.1994.1013">10.1006/tpbi.1994.1013</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Theoretical+Population+Biology&rft.atitle=Multispecies+competition+in+variable+environments&rft.volume=45&rft.issue=3&rft.pages=227-276&rft.date=1994&rft_id=info%3Adoi%2F10.1006%2Ftpbi.1994.1013&rft.aulast=Chesson&rft.aufirst=Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Chesson_2000space-6"><span class="mw-cite-backlink">^ <a href="#cite_ref-Chesson_2000space_6-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Chesson_2000space_6-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Chesson_2000space_6-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChesson2000" class="citation journal cs1">Chesson, Peter (2000). "General theory of competitive coexistence in spatially varying environments". <i>Theoretical Population Biology</i>. <b>58</b> (3): 211–237. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1006%2Ftpbi.2000.1486">10.1006/tpbi.2000.1486</a>. <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/11120650">11120650</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Theoretical+Population+Biology&rft.atitle=General+theory+of+competitive+coexistence+in+spatially+varying+environments&rft.volume=58&rft.issue=3&rft.pages=211-237&rft.date=2000&rft_id=info%3Adoi%2F10.1006%2Ftpbi.2000.1486&rft_id=info%3Apmid%2F11120650&rft.aulast=Chesson&rft.aufirst=Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Chesson_2000review-7"><span class="mw-cite-backlink">^ <a href="#cite_ref-Chesson_2000review_7-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Chesson_2000review_7-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChesson2000" class="citation journal cs1">Chesson, Peter (2000). "Mechanisms of maintenance of species diversity". <i>Annual Review of Ecology and Systematics</i>. <b>31</b>: 343–366. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.ecolsys.31.1.343">10.1146/annurev.ecolsys.31.1.343</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:403954">403954</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Ecology+and+Systematics&rft.atitle=Mechanisms+of+maintenance+of+species+diversity&rft.volume=31&rft.pages=343-366&rft.date=2000&rft_id=info%3Adoi%2F10.1146%2Fannurev.ecolsys.31.1.343&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A403954%23id-name%3DS2CID&rft.aulast=Chesson&rft.aufirst=Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-8"><span class="mw-cite-backlink"><b><a href="#cite_ref-8">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAndrewarthaBirch,_Charles1954" class="citation book cs1">Andrewartha, Herbert George; Birch, Charles (1954). <i>The distribution and abundance of animals</i>. University of Chicago Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/0-226-02026-6" title="Special:BookSources/0-226-02026-6"><bdi>0-226-02026-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+distribution+and+abundance+of+animals&rft.pub=University+of+Chicago+Press&rft.date=1954&rft.isbn=0-226-02026-6&rft.aulast=Andrewartha&rft.aufirst=Herbert+George&rft.au=Birch%2C+Charles&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Tinkle_&_Ballinger_1972-9"><span class="mw-cite-backlink"><b><a href="#cite_ref-Tinkle_&_Ballinger_1972_9-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTinkleBallinger,_Royce1972" class="citation journal cs1">Tinkle, Donald; Ballinger, Royce (1972). "Sceloporus undulatus: a study of intraspecific comparative demography of a lizard". <i>Ecology</i>. <b>53</b> (4): 570–584. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1972Ecol...53..570T">1972Ecol...53..570T</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2307%2F1934772">10.2307/1934772</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/1934772">1934772</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecology&rft.atitle=Sceloporus+undulatus%3A+a+study+of+intraspecific+comparative+demography+of+a+lizard&rft.volume=53&rft.issue=4&rft.pages=570-584&rft.date=1972&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F1934772%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.2307%2F1934772&rft_id=info%3Abibcode%2F1972Ecol...53..570T&rft.aulast=Tinkle&rft.aufirst=Donald&rft.au=Ballinger%2C+Royce&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Sears_&_Chesson_2007-10"><span class="mw-cite-backlink">^ <a href="#cite_ref-Sears_&_Chesson_2007_10-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Sears_&_Chesson_2007_10-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-Sears_&_Chesson_2007_10-2"><sup><i><b>c</b></i></sup></a> <a href="#cite_ref-Sears_&_Chesson_2007_10-3"><sup><i><b>d</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSearsChesson,_Peter2007" class="citation journal cs1">Sears, Anna; Chesson, Peter (2007). "New methods for quantifying the spatial storage effect: an illustration with desert annuals". <i>Ecology</i>. <b>88</b> (9): 2240–2247. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2007Ecol...88.2240S">2007Ecol...88.2240S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1890%2F06-0645.1">10.1890/06-0645.1</a>. <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/17918402">17918402</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:18486888">18486888</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecology&rft.atitle=New+methods+for+quantifying+the+spatial+storage+effect%3A+an+illustration+with+desert+annuals&rft.volume=88&rft.issue=9&rft.pages=2240-2247&rft.date=2007&rft_id=info%3Adoi%2F10.1890%2F06-0645.1&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A18486888%23id-name%3DS2CID&rft_id=info%3Apmid%2F17918402&rft_id=info%3Abibcode%2F2007Ecol...88.2240S&rft.aulast=Sears&rft.aufirst=Anna&rft.au=Chesson%2C+Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Kuang_&_Chesson_2009-11"><span class="mw-cite-backlink"><b><a href="#cite_ref-Kuang_&_Chesson_2009_11-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKuangChesson2009" class="citation journal cs1">Kuang, Jessica; Chesson, Peter (2009). "Coexistence of annual plants: Generalist seed predation weakens the storage effect". <i>Ecology</i>. <b>90</b> (1): 170–182. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2009Ecol...90..170K">2009Ecol...90..170K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1890%2F08-0207.1">10.1890/08-0207.1</a>. <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/19294923">19294923</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:17363406">17363406</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecology&rft.atitle=Coexistence+of+annual+plants%3A+Generalist+seed+predation+weakens+the+storage+effect&rft.volume=90&rft.issue=1&rft.pages=170-182&rft.date=2009&rft_id=info%3Adoi%2F10.1890%2F08-0207.1&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A17363406%23id-name%3DS2CID&rft_id=info%3Apmid%2F19294923&rft_id=info%3Abibcode%2F2009Ecol...90..170K&rft.aulast=Kuang&rft.aufirst=Jessica&rft.au=Chesson%2C+Peter&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Chesson_&_Kuang_2008_nature-12"><span class="mw-cite-backlink"><b><a href="#cite_ref-Chesson_&_Kuang_2008_nature_12-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChessonKuang2008" class="citation journal cs1">Chesson, Peter; Kuang, Jessica (2008). "The interaction between predation and competition". <i>Nature</i>. <b>456</b> (7219): 235–238. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2008Natur.456..235C">2008Natur.456..235C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2Fnature07248">10.1038/nature07248</a>. <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/19005554">19005554</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:4342701">4342701</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=The+interaction+between+predation+and+competition&rft.volume=456&rft.issue=7219&rft.pages=235-238&rft.date=2008&rft_id=info%3Adoi%2F10.1038%2Fnature07248&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4342701%23id-name%3DS2CID&rft_id=info%3Apmid%2F19005554&rft_id=info%3Abibcode%2F2008Natur.456..235C&rft.aulast=Chesson&rft.aufirst=Peter&rft.au=Kuang%2C+Jessica&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Holt_1977-13"><span class="mw-cite-backlink"><b><a href="#cite_ref-Holt_1977_13-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHolt1977" class="citation journal cs1">Holt, Robert (1977). "Predation, apparent competition, and structure of prey communities". <i>Theoretical Population Biology</i>. <b>12</b> (2): 197–229. <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%2F0040-5809%2877%2990042-9">10.1016/0040-5809(77)90042-9</a>. <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/929457">929457</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Theoretical+Population+Biology&rft.atitle=Predation%2C+apparent+competition%2C+and+structure+of+prey+communities&rft.volume=12&rft.issue=2&rft.pages=197-229&rft.date=1977&rft_id=info%3Adoi%2F10.1016%2F0040-5809%2877%2990042-9&rft_id=info%3Apmid%2F929457&rft.aulast=Holt&rft.aufirst=Robert&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Chesson_&_Kuang_2010_multi-14"><span class="mw-cite-backlink"><b><a href="#cite_ref-Chesson_&_Kuang_2010_multi_14-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChessonKuang2010" class="citation journal cs1">Chesson, Peter; Kuang, Jessica (2010). "The storage effect due to frequency-dependent predation in multispecies plant communities". <i>Theoretical Population Biology</i>. <b>78</b> (2): 148–164. <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%2Fj.tpb.2010.06.003">10.1016/j.tpb.2010.06.003</a>. <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/20600208">20600208</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Theoretical+Population+Biology&rft.atitle=The+storage+effect+due+to+frequency-dependent+predation+in+multispecies+plant+communities&rft.volume=78&rft.issue=2&rft.pages=148-164&rft.date=2010&rft_id=info%3Adoi%2F10.1016%2Fj.tpb.2010.06.003&rft_id=info%3Apmid%2F20600208&rft.aulast=Chesson&rft.aufirst=Peter&rft.au=Kuang%2C+Jessica&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Mordecai_2011-15"><span class="mw-cite-backlink"><b><a href="#cite_ref-Mordecai_2011_15-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMordecai2011" class="citation journal cs1">Mordecai, Erin (2011). "Pathogen impacts on plant communities: Unifying theory, concepts, and empirical work". <i>Ecological Monographs</i>. <b>81</b> (3): 429–441. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2011EcoM...81..429M">2011EcoM...81..429M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1890%2F10-2241.1">10.1890/10-2241.1</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecological+Monographs&rft.atitle=Pathogen+impacts+on+plant+communities%3A+Unifying+theory%2C+concepts%2C+and+empirical+work&rft.volume=81&rft.issue=3&rft.pages=429-441&rft.date=2011&rft_id=info%3Adoi%2F10.1890%2F10-2241.1&rft_id=info%3Abibcode%2F2011EcoM...81..429M&rft.aulast=Mordecai&rft.aufirst=Erin&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-16"><span class="mw-cite-backlink"><b><a href="#cite_ref-16">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKellyBowler,_Michael2002" class="citation journal cs1">Kelly, Colleen; Bowler, Michael (2002). "Coexistence and relative abundance in forest tree species". <i>Nature</i>. <b>417</b> (6887): 437–440. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2002Natur.417..437K">2002Natur.417..437K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F417437a">10.1038/417437a</a>. <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/12024212">12024212</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:4318307">4318307</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Coexistence+and+relative+abundance+in+forest+tree+species&rft.volume=417&rft.issue=6887&rft.pages=437-440&rft.date=2002&rft_id=info%3Adoi%2F10.1038%2F417437a&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4318307%23id-name%3DS2CID&rft_id=info%3Apmid%2F12024212&rft_id=info%3Abibcode%2F2002Natur.417..437K&rft.aulast=Kelly&rft.aufirst=Colleen&rft.au=Bowler%2C+Michael&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Kelly_&_Bowler_2005-17"><span class="mw-cite-backlink"><b><a href="#cite_ref-Kelly_&_Bowler_2005_17-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKellyBowler,_Michael2005" class="citation journal cs1">Kelly, Colleen; Bowler, Michael (2005). "A new application of storage dynamics: differential sensitivity, diffuse competition and temporal niches". <i>Ecology</i>. <b>86</b> (4): 1012–1022. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2005Ecol...86.1012K">2005Ecol...86.1012K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1890%2F04-0091">10.1890/04-0091</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecology&rft.atitle=A+new+application+of+storage+dynamics%3A+differential+sensitivity%2C+diffuse+competition+and+temporal+niches&rft.volume=86&rft.issue=4&rft.pages=1012-1022&rft.date=2005&rft_id=info%3Adoi%2F10.1890%2F04-0091&rft_id=info%3Abibcode%2F2005Ecol...86.1012K&rft.aulast=Kelly&rft.aufirst=Colleen&rft.au=Bowler%2C+Michael&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-Angert_etal_2009-18"><span class="mw-cite-backlink">^ <a href="#cite_ref-Angert_etal_2009_18-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Angert_etal_2009_18-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAngertHuxman,_TravisChesson,_PeterVenable,_D._Lawerance2009" class="citation journal cs1">Angert, Amy; Huxman, Travis; Chesson, Peter; Venable, D. Lawerance (2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2710622">"Functional tradeoffs determine species coexistence via the storage effect"</a>. <i>Proceedings of the National Academy of Sciences of the United States of America</i>. <b>106</b> (28): 11641–11645. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2009PNAS..10611641A">2009PNAS..10611641A</a>. <a href="/wiki/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.0904512106">10.1073/pnas.0904512106</a></span>. <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/PMC2710622">2710622</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/19571002">19571002</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+National+Academy+of+Sciences+of+the+United+States+of+America&rft.atitle=Functional+tradeoffs+determine+species+coexistence+via+the+storage+effect&rft.volume=106&rft.issue=28&rft.pages=11641-11645&rft.date=2009&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2710622%23id-name%3DPMC&rft_id=info%3Apmid%2F19571002&rft_id=info%3Adoi%2F10.1073%2Fpnas.0904512106&rft_id=info%3Abibcode%2F2009PNAS..10611641A&rft.aulast=Angert&rft.aufirst=Amy&rft.au=Huxman%2C+Travis&rft.au=Chesson%2C+Peter&rft.au=Venable%2C+D.+Lawerance&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2710622&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-19"><span class="mw-cite-backlink"><b><a href="#cite_ref-19">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAngertHuxman,_TravisBarron-Gafford,_GregGerst,_Katherine2009" class="citation journal cs1">Angert, Amy; Huxman, Travis; Barron-Gafford, Greg; Gerst, Katherine; Venable, D. Lawerance (2009). <a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1365-2745.2006.01203.x">"Linking growth strategies to long-term population dynamics in a guild of desert annuals"</a>. <i>Journal of Ecology</i>. <b>95</b> (2): 321–331. <a href="/wiki/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/10.1111%2Fj.1365-2745.2006.01203.x">10.1111/j.1365-2745.2006.01203.x</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Ecology&rft.atitle=Linking+growth+strategies+to+long-term+population+dynamics+in+a+guild+of+desert+annuals&rft.volume=95&rft.issue=2&rft.pages=321-331&rft.date=2009&rft_id=info%3Adoi%2F10.1111%2Fj.1365-2745.2006.01203.x&rft.aulast=Angert&rft.aufirst=Amy&rft.au=Huxman%2C+Travis&rft.au=Barron-Gafford%2C+Greg&rft.au=Gerst%2C+Katherine&rft.au=Venable%2C+D.+Lawerance&rft_id=https%3A%2F%2Fdoi.org%2F10.1111%252Fj.1365-2745.2006.01203.x&rfr_id=info%3Asid%2Fen.wikipedia.org%3AStorage+effect" class="Z3988"></span></span> </li> <li id="cite_note-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-20">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHuxmanBarron-Gafford,_GregGerst,_KatherineAngert,_Amy2009" class="citation journal cs1">Huxman, Travis; Barron-Gafford, Greg; Gerst, Katherine; Angert, Amy; Tyler, Anna; Venable, D. Lawerance (2009). "Photosynthetic resource-use efficiency and demographic variability in desert winter annual plants". <i>Ecology</i>. <b>89</b> (6): 1554–1563. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1890%2F06-2080.1">10.1890/06-2080.1</a>. <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/18589520">18589520</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:34025348">34025348</a>.</cite><span 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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 class="mw-selflink selflink">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 href="/wiki/Alternative_stable_state" title="Alternative stable 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