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class="mw-body-content"><div class="mw-content-ltr mw-parser-output" lang="en" dir="ltr"><div class="shortdescription nomobile noexcerpt noprint searchaux" style="display:none">Study of the distribution or space occupied by species</div> <style data-mw-deduplicate="TemplateStyles:r1251242444">.mw-parser-output .ambox{border:1px solid #a2a9b1;border-left:10px solid #36c;background-color:#fbfbfb;box-sizing:border-box}.mw-parser-output .ambox+link+.ambox,.mw-parser-output .ambox+link+style+.ambox,.mw-parser-output .ambox+link+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+style+.ambox,.mw-parser-output .ambox+.mw-empty-elt+link+link+.ambox{margin-top:-1px}html body.mediawiki .mw-parser-output .ambox.mbox-small-left{margin:4px 1em 4px 0;overflow:hidden;width:238px;border-collapse:collapse;font-size:88%;line-height:1.25em}.mw-parser-output .ambox-speedy{border-left:10px solid #b32424;background-color:#fee7e6}.mw-parser-output .ambox-delete{border-left:10px solid #b32424}.mw-parser-output .ambox-content{border-left:10px solid #f28500}.mw-parser-output .ambox-style{border-left:10px solid #fc3}.mw-parser-output .ambox-move{border-left:10px solid #9932cc}.mw-parser-output .ambox-protection{border-left:10px solid #a2a9b1}.mw-parser-output .ambox .mbox-text{border:none;padding:0.25em 0.5em;width:100%}.mw-parser-output .ambox .mbox-image{border:none;padding:2px 0 2px 0.5em;text-align:center}.mw-parser-output .ambox .mbox-imageright{border:none;padding:2px 0.5em 2px 0;text-align:center}.mw-parser-output .ambox .mbox-empty-cell{border:none;padding:0;width:1px}.mw-parser-output .ambox .mbox-image-div{width:52px}@media(min-width:720px){.mw-parser-output .ambox{margin:0 10%}}@media print{body.ns-0 .mw-parser-output .ambox{display:none!important}}</style><table class="box-Lead_too_short plainlinks metadata ambox ambox-content ambox-lead_too_short" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><a href="/wiki/File:Wiki_letter_w.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/40px-Wiki_letter_w.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/60px-Wiki_letter_w.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/6/6c/Wiki_letter_w.svg/80px-Wiki_letter_w.svg.png 2x" data-file-width="44" data-file-height="44" /></a></span></div></td><td class="mbox-text"><div class="mbox-text-span">This article's <a href="/wiki/Wikipedia:Manual_of_Style/Lead_section#Length" title="Wikipedia:Manual of Style/Lead section">lead section</a> <b>may be too short to adequately <a href="/wiki/Wikipedia:Summary_style" title="Wikipedia:Summary style">summarize</a> the key points</b>.<span class="hide-when-compact"> Please consider expanding the lead to <a href="/wiki/Wikipedia:Manual_of_Style/Lead_section#Provide_an_accessible_overview" title="Wikipedia:Manual of Style/Lead section">provide an accessible overview</a> of all important aspects of the article.</span> <span class="date-container"><i>(<span class="date">April 2021</span>)</i></span></div></td></tr></tbody></table> <p><b>Spatial ecology </b> studies the ultimate distributional or spatial unit occupied by a <a href="/wiki/Species" title="Species">species</a>. In a particular <a href="/wiki/Habitat" title="Habitat">habitat</a> shared by several species, each of the species is usually confined to its own <a href="/wiki/Microhabitat" class="mw-redirect" title="Microhabitat">microhabitat</a> or spatial niche because two species in the same general territory cannot usually occupy the same <a href="/wiki/Ecological_niche" title="Ecological niche">ecological niche</a> for any significant length of time. </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Overview">Overview</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=1" title="Edit section: Overview"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In nature, organisms are neither <a href="/wiki/Species_distribution#Regular_or_uniform" title="Species distribution">distributed uniformly</a> nor at <a href="/wiki/Species_distribution#Random" title="Species distribution">random</a>, forming instead some sort of <a href="/wiki/Spatial_pattern" class="mw-redirect" title="Spatial pattern">spatial pattern</a>.<sup id="cite_ref-Legendre_1-0" class="reference"><a href="#cite_note-Legendre-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This is due to various energy inputs, <a href="/wiki/Disturbance_(ecology)" title="Disturbance (ecology)">disturbances</a>, and species interactions that result in spatially patchy structures or <a href="/wiki/Gradient" title="Gradient">gradients</a>. This spatial variance in the environment creates diversity in communities of organisms, as well as in the variety of the observed biological and ecological events.<sup id="cite_ref-Legendre_1-1" class="reference"><a href="#cite_note-Legendre-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The type of spatial arrangement present may suggest certain interactions within and between species, such as <a href="/wiki/Biological_competition" class="mw-redirect" title="Biological competition">competition</a>, <a href="/wiki/Predation" title="Predation">predation</a>, and <a href="/wiki/Reproduction" title="Reproduction">reproduction</a>.<sup id="cite_ref-Perry_2-0" class="reference"><a href="#cite_note-Perry-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> On the other hand, certain spatial patterns may also rule out specific <a href="/wiki/Ecological_theory" class="mw-redirect" title="Ecological theory">ecological theories</a> previously thought to be true.<sup id="cite_ref-Liebhold_3-0" class="reference"><a href="#cite_note-Liebhold-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Although spatial ecology deals with spatial patterns, it is usually based on <a href="/wiki/Empirical" class="mw-redirect" title="Empirical">observational data</a> rather than on an <a href="/wiki/Scientific_modelling" title="Scientific modelling">existing model</a>.<sup id="cite_ref-Perry_2-1" class="reference"><a href="#cite_note-Perry-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> This is because nature rarely follows set expected order. To properly research a spatial pattern or population, the spatial extent to which it occurs must be detected. Ideally, this would be accomplished beforehand via a benchmark spatial survey, which would determine whether the pattern or process is on a local, regional, or global scale. This is rare in actual field research, however, due to the lack of time and funding, as well as the ever-changing nature of such widely-studied <a href="/wiki/Organism" title="Organism">organisms</a> such as <a href="/wiki/Insect" title="Insect">insects</a> and <a href="/wiki/Wildlife" title="Wildlife">wildlife</a>.<sup id="cite_ref-Tobin_4-0" class="reference"><a href="#cite_note-Tobin-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> With detailed information about a species' life-stages, dynamics, <a href="/wiki/Demography" title="Demography">demography</a>, movement, behavior, etc., models of spatial pattern may be developed to estimate and predict events in unsampled locations.<sup id="cite_ref-Perry_2-2" class="reference"><a href="#cite_note-Perry-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="History">History</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=2" title="Edit section: History"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Most mathematical studies in ecology in the nineteenth century assumed a uniform distribution of living organisms in their habitat.<sup id="cite_ref-Legendre_1-2" class="reference"><a href="#cite_note-Legendre-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In the past quarter century, ecologists have begun to recognize the degree to which organisms respond to spatial patterns in their environment. Due to the rapid advances in computer technology in the same time period, more advanced methods of statistical data analysis have come into use.<sup id="cite_ref-Liebhold_3-1" class="reference"><a href="#cite_note-Liebhold-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Also, the repeated use of <a href="/wiki/Remote_sensing" title="Remote sensing">remotely sensed</a> imagery and <a href="/wiki/Geographic_information_systems" class="mw-redirect" title="Geographic information systems">geographic information systems</a> in a particular area has led to increased analysis and identification of spatial patterns over time.<sup id="cite_ref-Tobin_4-1" class="reference"><a href="#cite_note-Tobin-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> These technologies have also increased the ability to determine how human activities have impacted animal habitat and <a href="/wiki/Climate_change" title="Climate change">climate change</a>.<sup id="cite_ref-5" class="reference"><a href="#cite_note-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> The natural world has become increasingly fragmented due to human activities; anthropogenic landscape change has had a ripple-effect impacts on wildlife populations, which are now more likely to be small, restricted in distribution, and increasingly isolated from one another. In part as a reaction to this knowledge, and partially due to increasingly sophisticated theoretical developments, ecologists began stressing the importance of spatial context in research. Spatial ecology emerged from this movement toward spatial accountability; "the progressive introduction of spatial variation and complexity into ecological analysis, including changes in spatial patterns over time".<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Concepts">Concepts</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=3" title="Edit section: Concepts"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Scale">Scale</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=4" title="Edit section: Scale"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In spatial ecology, scale refers to the spatial extent of ecological processes and the spatial interpretation of the data.<sup id="cite_ref-Fortin_7-0" class="reference"><a href="#cite_note-Fortin-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The response of an organism or a species to the environment is particular to a specific scale, and may respond differently at a larger or smaller scale.<sup id="cite_ref-Bellier_8-0" class="reference"><a href="#cite_note-Bellier-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Choosing a scale that is appropriate to the ecological process in question is very important in accurately hypothesizing and determining the underlying cause.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Most often, ecological patterns are a result of multiple ecological processes, which often operate at more than one spatial scale.<sup id="cite_ref-ver_Hoef_11-0" class="reference"><a href="#cite_note-ver_Hoef-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Through the use of such spatial statistical methods such as <a href="/wiki/Geostatistics" title="Geostatistics">geostatistics</a> and principal coordinate analysis of neighbor matrices (PCNM), one can identify spatial relationships between organisms and environmental variables at multiple scales.<sup id="cite_ref-Bellier_8-1" class="reference"><a href="#cite_note-Bellier-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Spatial_autocorrelation">Spatial autocorrelation</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=5" title="Edit section: Spatial autocorrelation"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/Spatial_analysis#Spatial_dependency_or_auto-correlation" title="Spatial analysis">Spatial autocorrelation</a> refers to the value of samples taken close to each other are more likely to have similar magnitude than by chance alone.<sup id="cite_ref-Fortin_7-1" class="reference"><a href="#cite_note-Fortin-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> When a pair of values located at a certain distance apart are more similar than expected by chance, the spatial <a href="/wiki/Autocorrelation" title="Autocorrelation">autocorrelation</a> is said to be positive. When a pair of values are less similar, the spatial autocorrelation is said to be negative. It is common for values to be positively autocorrelated at shorter distances and negative autocorrelated at longer distances.<sup id="cite_ref-Legendre_1-3" class="reference"><a href="#cite_note-Legendre-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This is commonly known as <a href="/wiki/Tobler%27s_first_law_of_geography" title="Tobler's first law of geography">Tobler's first law of geography</a>, summarized as "everything is related to everything else, but nearby objects are more related than distant objects". </p><p>In ecology, there are two important sources of spatial autocorrelation, which both arise from spatial-temporal processes, such as <a href="/wiki/Biological_dispersal" title="Biological dispersal">dispersal</a> or <a href="/wiki/Human_migration" title="Human migration">migration</a>:<sup id="cite_ref-ver_Hoef_11-1" class="reference"><a href="#cite_note-ver_Hoef-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p> <ul><li>True/inherent spatial autocorrelation arises from interactions among individuals located in close proximity. This process is endogenous (internal) and results in the individuals being spatially adjacent in a <a href="/wiki/Landscape_ecology#Patch_and_mosaic" title="Landscape ecology">patchy</a> fashion.<sup id="cite_ref-Fortin_7-2" class="reference"><a href="#cite_note-Fortin-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> An example of this would be <a href="/wiki/Sexual_reproduction" title="Sexual reproduction">sexual reproduction</a>, the success of which requires the closeness of a male and female of the species.</li> <li>Induced spatial autocorrelation (or 'induced <a href="/wiki/Spatial_dependence" class="mw-redirect" title="Spatial dependence">spatial dependence</a>') arises from the species response to the spatial structure of exogenous (external) factors, which are themselves spatially autocorrelated.<sup id="cite_ref-Fortin_7-3" class="reference"><a href="#cite_note-Fortin-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> An example of this would be the winter habitat range of deer, which use conifers for heat retention and <a href="/wiki/Forage" title="Forage">forage</a>.</li></ul> <p>Most ecological data exhibit some degree of spatial autocorrelation, depending on the ecological scale (spatial resolution) of interest. As the spatial arrangement of most ecological data is not random, traditional <a href="/wiki/Random_sample" class="mw-redirect" title="Random sample">random population samples</a> tend to overestimate the true value of a variable, or infer significant <a href="/wiki/Correlation" title="Correlation">correlation</a> where there is none.<sup id="cite_ref-Legendre_1-4" class="reference"><a href="#cite_note-Legendre-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> This <a href="/wiki/Systemic_bias" title="Systemic bias">bias</a> can be corrected through the use of <a href="/wiki/Geostatistics" title="Geostatistics">geostatistics</a> and other more statistically advanced models. Regardless of method, the sample size must be appropriate to the scale and the spatial statistical method used in order to be valid.<sup id="cite_ref-Tobin_4-2" class="reference"><a href="#cite_note-Tobin-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Pattern">Pattern</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=6" title="Edit section: Pattern"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Spatial patterns, such as the distribution of a species, are the result of either true or induced spatial autocorrelation.<sup id="cite_ref-Fortin_7-4" class="reference"><a href="#cite_note-Fortin-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> In nature, organisms are distributed neither uniformly nor at random. The environment is spatially structured by various ecological processes,<sup id="cite_ref-Legendre_1-5" class="reference"><a href="#cite_note-Legendre-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> which in combination with the behavioral response of species generally results in: </p> <ul><li>Gradients (trends): steady directional change in numbers over a specific distance</li> <li>Patches (clumps): a relatively uniform and homogenous area separated by gaps</li> <li>Noise (random fluctuations): variation not able to be explained by a model</li></ul> <p>Theoretically, any of these structures may occur at any given scale. Due to the presence of spatial autocorrelation, in nature gradients are generally found at the global level, whereas patches represent intermediate (regional) scales, and noise at local scales.<sup id="cite_ref-ver_Hoef_11-2" class="reference"><a href="#cite_note-ver_Hoef-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> </p><p>The analysis of spatial ecological patterns comprises two families of methods:<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <ul><li>Point pattern analysis deals with the distribution of individuals through space, and is used to determine whether the distribution is random.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> It also describes the type of pattern and draws conclusions on what kind of process created the observed pattern. Quadrat-density and the <a href="/wiki/Nearest_neighbor_search" title="Nearest neighbor search">nearest neighbor</a> methods are the most commonly used statistical methods.</li> <li>Surface pattern analysis deals with spatially continuous phenomena. After the spatial distribution of the variables is determined through discrete sampling, statistical methods are used to quantify the magnitude, intensity, and extent of spatial autocorrelation present in the data (such as <a href="/wiki/Correlogram" title="Correlogram">correlograms</a>, variograms, and periodograms), as well as to map the amount of spatial variation.</li></ul> <div class="mw-heading mw-heading2"><h2 id="Applications">Applications</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=7" title="Edit section: Applications"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Research">Research</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=8" title="Edit section: Research"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Analysis of spatial trends has been used to research <a href="/wiki/Wildlife_management" title="Wildlife management">wildlife management</a>, <a href="/wiki/Fire_ecology" title="Fire ecology">fire ecology</a>, <a href="/wiki/Population_ecology" title="Population ecology">population ecology</a>, <a href="/wiki/Disease_ecology" title="Disease ecology">disease ecology</a>, <a href="/wiki/Invasive_species" title="Invasive species">invasive species</a>, <a href="/wiki/Marine_ecology" class="mw-redirect" title="Marine ecology">marine ecology</a>, and <a href="/wiki/Carbon_sequestration" title="Carbon sequestration">carbon sequestration</a> modeling using the spatial relationships and patterns to determine ecological processes and their effects on the environment. Spatial patterns have different ecosystem functioning in ecology for examples enhanced productive.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Interdisciplinary">Interdisciplinary</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=9" title="Edit section: Interdisciplinary"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The concepts of spatial ecology are fundamental to understanding the spatial dynamics of <a href="/wiki/Population_ecology" title="Population ecology">population</a> and <a href="/wiki/Community_ecology" class="mw-redirect" title="Community ecology">community ecology</a>. The <a href="/wiki/Spatial_heterogeneity" title="Spatial heterogeneity">spatial heterogeneity</a> of populations and communities plays a central role in such ecological theories as <a href="/wiki/Ecological_succession" title="Ecological succession">succession</a>, <a href="/wiki/Adaptation" title="Adaptation">adaptation</a>, community stability, <a href="/wiki/Competition_(biology)" title="Competition (biology)">competition</a>, <a href="/wiki/Predator-prey_interaction" class="mw-redirect" title="Predator-prey interaction">predator-prey interactions</a>, <a href="/wiki/Parasitism" title="Parasitism">parasitism</a>, and <a href="/wiki/Epidemic" title="Epidemic">epidemics</a>.<sup id="cite_ref-Legendre_1-6" class="reference"><a href="#cite_note-Legendre-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> The rapidly expanding field of landscape ecology utilizes the basic aspects of spatial ecology in its research.<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 practical use of spatial ecology concepts is essential to understanding the consequences of <a href="/wiki/Habitat_fragmentation" title="Habitat fragmentation">fragmentation</a> and habitat loss for wildlife. Understanding the response of a species to a spatial structure provides useful information in regards to <a href="/wiki/Biodiversity" title="Biodiversity">biodiversity conservation</a> and habitat restoration.<sup id="cite_ref-15" class="reference"><a href="#cite_note-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p><p>Spatial ecology modeling uses components of remote sensing and <a href="/wiki/Geographical_information_system" class="mw-redirect" title="Geographical information system">geographical information systems</a> (GIS).<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-heading2"><h2 id="Statistical_tests">Statistical tests</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=10" title="Edit section: Statistical tests"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>A number of statistical tests have been developed to study such relations. </p> <div class="mw-heading mw-heading3"><h3 id="Tests_based_on_distance">Tests based on distance</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=11" title="Edit section: Tests based on distance"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Clark_and_Evans'_R"><span id="Clark_and_Evans.27_R"></span>Clark and Evans' R</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=12" title="Edit section: Clark and Evans' R"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Clark and Evans in 1954<sup id="cite_ref-Clark1954_16-0" class="reference"><a href="#cite_note-Clark1954-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> proposed a test based on the density and distance between organisms. Under the <a href="/wiki/Null_hypothesis" title="Null hypothesis">null hypothesis</a> the expected distance ( <i>r</i><sub>e</sub> ) between the organisms (measured as the nearest neighbor's distance) with a known constant density ( <i>ρ</i> ) 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 r_{e}={\frac {1}{2{\sqrt {\rho }}}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msub> <mi>r</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>e</mi> </mrow> </msub> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mn>1</mn> <mrow> <mn>2</mn> <mrow class="MJX-TeXAtom-ORD"> <msqrt> <mi>ρ</mi> </msqrt> </mrow> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle r_{e}={\frac {1}{2{\sqrt {\rho }}}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/4f52ceadbf356912843e87a90aace2e97034f362" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.838ex; width:10.282ex; height:6.176ex;" alt="{\displaystyle r_{e}={\frac {1}{2{\sqrt {\rho }}}}}"></span></dd></dl> <p>The difference between the observed ( <i>r</i><sub>o</sub> ) and the expected ( <i>r</i><sub>e</sub> ) can be tested with a Z test </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 Z={\frac {r_{o}-r_{e}}{SE}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>Z</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mrow> <msub> <mi>r</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>o</mi> </mrow> </msub> <mo>−</mo> <msub> <mi>r</mi> <mrow class="MJX-TeXAtom-ORD"> <mi>e</mi> </mrow> </msub> </mrow> <mrow> <mi>S</mi> <mi>E</mi> </mrow> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle Z={\frac {r_{o}-r_{e}}{SE}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/8b91f7ba017813b9816f4657d82e91760f3d1130" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -2.005ex; width:12.581ex; height:5.176ex;" alt="{\displaystyle Z={\frac {r_{o}-r_{e}}{SE}}}"></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 SE={\frac {0.26136}{\sqrt {N\rho }}}}"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>S</mi> <mi>E</mi> <mo>=</mo> <mrow class="MJX-TeXAtom-ORD"> <mfrac> <mn>0.26136</mn> <msqrt> <mi>N</mi> <mi>ρ</mi> </msqrt> </mfrac> </mrow> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle SE={\frac {0.26136}{\sqrt {N\rho }}}}</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/b618be7d05ced6a4a0fff5e733641708dfdaf572" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -3.171ex; width:14.831ex; height:6.509ex;" alt="{\displaystyle SE={\frac {0.26136}{\sqrt {N\rho }}}}"></span></dd></dl> <p>where <i>N</i> is the number of nearest neighbor measurements. For large samples <i>Z</i> is distributed normally. The results are usually reported in the form of a ratio: <i>R</i> = ( <i>r</i><sub>o</sub> ) / ( <i>r</i><sub>e</sub> ) </p> <div class="mw-heading mw-heading4"><h4 id="Pielou's_α"><span id="Pielou.27s_.CE.B1"></span>Pielou's α</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=13" title="Edit section: Pielou's α"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p><a href="/wiki/E._C._Pielou" class="mw-redirect" title="E. C. Pielou">Pielou</a> in 1959 devised a different statistic.<sup id="cite_ref-Pielou1959_17-0" class="reference"><a href="#cite_note-Pielou1959-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> She considered instead of the nearest neighbors the distance between an organism and a set of pre-chosen random points within the sampling area, again assuming a constant density. If the population is randomly dispersed in the area these distances will equal the nearest neighbor distances. Let <i>ω</i> be the ratio between the distances from the random points and the distances calculated from the nearest neighbor calculations. The <i>α</i> is<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> <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 \alpha =\pi d\omega }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <mi>α</mi> <mo>=</mo> <mi>π</mi> <mi>d</mi> <mi>ω</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \alpha =\pi d\omega }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/d158d9409687b27ba205f147b64be8f1ce4c64f0" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -0.338ex; width:8.58ex; height:2.176ex;" alt="{\displaystyle \alpha =\pi d\omega }"></span></dd></dl> <p>where <i>d</i> is the constant common density and π has its usual numerical value. Values of α less than, equal to or greater than 1 indicate uniformity, randomness (a <a href="/wiki/Poisson_distribution" title="Poisson distribution">Poisson distribution</a>) or aggregation respectively. Alpha may be tested for a significant deviation from 1 by computing the test statistic </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 \chi _{2n}^{2}=2n\alpha }"> <semantics> <mrow class="MJX-TeXAtom-ORD"> <mstyle displaystyle="true" scriptlevel="0"> <msubsup> <mi>χ</mi> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> <mi>n</mi> </mrow> <mrow class="MJX-TeXAtom-ORD"> <mn>2</mn> </mrow> </msubsup> <mo>=</mo> <mn>2</mn> <mi>n</mi> <mi>α</mi> </mstyle> </mrow> <annotation encoding="application/x-tex">{\displaystyle \chi _{2n}^{2}=2n\alpha }</annotation> </semantics> </math></span><img src="https://wikimedia.org/api/rest_v1/media/math/render/svg/53792d17049d5ce23b6a535077049bda0a1a3c3a" class="mwe-math-fallback-image-inline mw-invert skin-invert" aria-hidden="true" style="vertical-align: -1.005ex; width:10.639ex; height:3.176ex;" alt="{\displaystyle \chi _{2n}^{2}=2n\alpha }"></span></dd></dl> <p>where <i>χ</i><sup>2</sup> is distributed with 2<i>n</i> degrees of freedom. <i>n</i> here is the number of organisms sampled. </p><p>Montford in 1961 showed that when the density is estimated rather than a known constant, this version of alpha tended to overestimate the actual degree of aggregation. He provided a revised formulation which corrects this error. There is a wide range of mathematical problems related to spatial ecological models, relating to spatial patterns and processes associated with chaotic phenomena, bifurcations and instability.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=14" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Edge_effects" title="Edge effects">Edge effects</a></li> <li><a href="/wiki/Spatial_analysis" title="Spatial analysis">Spatial analysis</a></li> <li><a href="/wiki/Taylor%27s_law" title="Taylor's law">Taylor's law</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=15" 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 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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="CITEREFLegendreFortin,_M.-J.1989" class="citation journal cs1">Legendre, P.; Fortin, M.-J. 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Liebhold; M.S. Rosenberg; J. Dungan; M. Miriti; A. Jakomulska; S. Citron-Pousty (2002). <a rel="nofollow" class="external text" href="https://repository.rothamsted.ac.uk/download/948e96ce68bb1377dac3ef63f2b787b2da16fcee643c33a151c552daaf320a04/3932523/Perry-2002-Illustrations-and-guidelines-for-se.pdf">"Illustrations and guidelines for selecting statistical methods for quantifying spatial pattern in ecological data"</a> <span class="cs1-format">(PDF)</span>. <i>Ecography</i>. <b>25</b> (5): 578–600. <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/10.1034%2Fj.1600-0587.2002.250507.x">10.1034/j.1600-0587.2002.250507.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=Ecography&rft.atitle=Illustrations+and+guidelines+for+selecting+statistical+methods+for+quantifying+spatial+pattern+in+ecological+data&rft.volume=25&rft.issue=5&rft.pages=578-600&rft.date=2002&rft_id=info%3Adoi%2F10.1034%2Fj.1600-0587.2002.250507.x&rft.aulast=Perry&rft.aufirst=J.N.&rft.au=A.M.+Liebhold&rft.au=M.S.+Rosenberg&rft.au=J.+Dungan&rft.au=M.+Miriti&rft.au=A.+Jakomulska&rft.au=S.+Citron-Pousty&rft_id=https%3A%2F%2Frepository.rothamsted.ac.uk%2Fdownload%2F948e96ce68bb1377dac3ef63f2b787b2da16fcee643c33a151c552daaf320a04%2F3932523%2FPerry-2002-Illustrations-and-guidelines-for-se.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASpatial+ecology" class="Z3988"></span></span> </li> <li id="cite_note-Liebhold-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-Liebhold_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-Liebhold_3-1"><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="CITEREFLiebholdJ._Gurevitch2002" class="citation journal cs1">Liebhold, A.M.; J. 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"Distance to nearest neighbor as a measure of spatial relationships in populations". <i>Ecology</i>. <b>35</b> (4): 445–453. <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%2F1931034">10.2307/1931034</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/1931034">1931034</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=Distance+to+nearest+neighbor+as+a+measure+of+spatial+relationships+in+populations&rft.volume=35&rft.issue=4&rft.pages=445-453&rft.date=1954&rft_id=info%3Adoi%2F10.2307%2F1931034&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F1931034%23id-name%3DJSTOR&rft.aulast=Clark&rft.aufirst=PJ&rft.au=Evans%2C+FC&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASpatial+ecology" class="Z3988"></span></span> </li> <li id="cite_note-Pielou1959-17"><span class="mw-cite-backlink"><b><a href="#cite_ref-Pielou1959_17-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPielou1959" class="citation journal cs1">Pielou, E C (1959). "The use of point-to-plant distances in the study of patterns in plant populations". <i>J Ecol</i>. <b>47</b>: 607–613. <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%2F2257293">10.2307/2257293</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/2257293">2257293</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=J+Ecol&rft.atitle=The+use+of+point-to-plant+distances+in+the+study+of+patterns+in+plant+populations&rft.volume=47&rft.pages=607-613&rft.date=1959&rft_id=info%3Adoi%2F10.2307%2F2257293&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F2257293%23id-name%3DJSTOR&rft.aulast=Pielou&rft.aufirst=E+C&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASpatial+ecology" class="Z3988"></span></span> </li> <li id="cite_note-18"><span class="mw-cite-backlink"><b><a href="#cite_ref-18">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPapadimitriou2010" class="citation journal cs1">Papadimitriou, Fivos (2010). <a rel="nofollow" class="external text" href="https://ges.rgo.ru/jour/article/view/266">"Mathematical Modelling of Spatial-Ecological Complex Systems: An evaluation"</a>. <i>Geography, Environment, Sustainability</i>. <b>1</b> (3): 67–80. <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/10.24057%2F2071-9388-2010-3-1-67-80">10.24057/2071-9388-2010-3-1-67-80</a></span>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geography%2C+Environment%2C+Sustainability&rft.atitle=Mathematical+Modelling+of+Spatial-Ecological+Complex+Systems%3A+An+evaluation&rft.volume=1&rft.issue=3&rft.pages=67-80&rft.date=2010&rft_id=info%3Adoi%2F10.24057%2F2071-9388-2010-3-1-67-80&rft.aulast=Papadimitriou&rft.aufirst=Fivos&rft_id=https%3A%2F%2Fges.rgo.ru%2Fjour%2Farticle%2Fview%2F266&rfr_id=info%3Asid%2Fen.wikipedia.org%3ASpatial+ecology" class="Z3988"></span></span> </li> </ol></div> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_ecology&action=edit&section=16" title="Edit section: External links"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a rel="nofollow" class="external text" href="http://www.spatialecology.com/">Spatial Ecology</a>, hosts software for use in spatial ecological analysis.</li> <li><a rel="nofollow" class="external text" href="http://www.helsinki.fi/bioscience/spatialecology/">Spatial Ecology Research Programme at the University of Helsinki</a></li> <li><a rel="nofollow" class="external text" href="http://www.uq.edu.au/spatialecology/">Spatial Ecology Lab at the University of Queensland</a></li> <li><a rel="nofollow" class="external text" href="http://www.blackwellpublishing.com/eco">Ecography</a> publishes <a href="/wiki/Peer-reviewed" class="mw-redirect" title="Peer-reviewed">peer-reviewed</a> articles on spatial ecology.</li> <li><a rel="nofollow" class="external text" href="http://www.nceas.ucsb.edu/">National Center for Ecological Analysis and Synthesis at the University of California, Santa Barbara</a></li> <li><a rel="nofollow" class="external text" href="https://archive.today/20011229073059/http://picea.sel.uaf.edu/">Spatial Ecology Lab at the University of Alaska, Fairbanks</a></li> <li><a rel="nofollow" class="external text" href="http://www.spatial-ecology.net/">Spatial Ecology wikipedia</a>, online resources for learning spatial ecological analysis and data processing using Open source software.</li></ul> <div style="clear:both;" class=""></div> <div class="navbox-styles"><style data-mw-deduplicate="TemplateStyles:r1129693374">.mw-parser-output .hlist dl,.mw-parser-output .hlist ol,.mw-parser-output .hlist ul{margin:0;padding:0}.mw-parser-output .hlist dd,.mw-parser-output .hlist dt,.mw-parser-output .hlist li{margin:0;display:inline}.mw-parser-output .hlist.inline,.mw-parser-output .hlist.inline dl,.mw-parser-output .hlist.inline ol,.mw-parser-output .hlist.inline ul,.mw-parser-output .hlist dl dl,.mw-parser-output .hlist dl ol,.mw-parser-output .hlist dl ul,.mw-parser-output .hlist ol dl,.mw-parser-output .hlist ol ol,.mw-parser-output .hlist ol ul,.mw-parser-output .hlist ul dl,.mw-parser-output .hlist ul ol,.mw-parser-output .hlist ul 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class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Polar_ecology" title="Polar ecology">Polar ecology</a> <ul><li><a href="/wiki/Arctic_ecology" title="Arctic ecology">Arctic ecology</a></li></ul></li> <li><a href="/wiki/Desert_ecology" title="Desert ecology">Desert ecology</a></li> <li><a href="/wiki/Forest_ecology" title="Forest ecology">Forest ecology</a></li> <li><a href="/wiki/Soil_ecology" title="Soil ecology">Soil ecology</a></li> <li><a href="/wiki/Tropical_ecology" title="Tropical ecology">Tropical ecology</a></li> <li><a href="/wiki/Urban_ecology" title="Urban ecology">Urban ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Phenomena#Science" class="mw-redirect" title="Phenomena">Ecological phenomena</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behavioral ecology</a></li> <li><a href="/wiki/Chemical_ecology" title="Chemical ecology">Chemical ecology</a></li> <li><a href="/wiki/Disease_ecology" title="Disease ecology">Disease ecology</a></li> <li><a href="/wiki/Ecophysiology" title="Ecophysiology">Ecophysiology</a></li> <li><a href="/wiki/Ecotoxicology" title="Ecotoxicology">Ecotoxicology</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Fire_ecology" title="Fire ecology">Fire ecology</a></li> <li><a href="/wiki/Functional_ecology" title="Functional ecology">Functional ecology</a></li> <li><a href="/wiki/Genetic_ecology" title="Genetic ecology">Genetic ecology</a></li> <li><a href="/wiki/Landscape_ecology" title="Landscape ecology">Landscape ecology</a></li> <li><a href="/wiki/Molecular_ecology" title="Molecular ecology">Molecular ecology</a></li> <li><a href="/wiki/Paleoecology" title="Paleoecology">Paleoecology</a></li> <li><a href="/wiki/Social_ecology_(academic_field)" title="Social ecology (academic field)">Social ecology</a></li> <li><a class="mw-selflink selflink">Spatial ecology</a></li> <li><a href="/wiki/Thermal_ecology" title="Thermal ecology">Thermal ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Interdisciplinarity" title="Interdisciplinarity">Interdisciplinarity</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Agroecology" title="Agroecology">Agroecology</a></li> <li><a href="/wiki/Ecological_anthropology" title="Ecological anthropology">Ecological anthropology</a></li> <li><a href="/wiki/Ecological_economics" title="Ecological economics">Ecological economics</a></li> <li><a href="/wiki/Ecological_engineering" title="Ecological engineering">Ecological engineering</a></li> <li><a href="/wiki/Political_ecology" title="Political ecology">Political ecology</a></li> <li><a href="/wiki/Systems_ecology" title="Systems ecology">Systems ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/History_of_ecology" title="History of ecology">History of ecology</a></li> <li><a href="/wiki/Glossary_of_ecology" title="Glossary of ecology">Glossary of ecology</a></li> <li><a href="/wiki/Applied_ecology" title="Applied ecology">Applied ecology</a> <ul><li><a href="/wiki/Restoration_ecology" class="mw-redirect" title="Restoration ecology">Restoration ecology</a></li></ul></li> <li><a href="/wiki/Ecosophy" title="Ecosophy">Ecosophy</a></li> <li><a href="/wiki/Natural_history" title="Natural history">Natural history</a></li> <li><a href="/wiki/Biogeography" title="Biogeography">Biogeography</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:_Trophic_components" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><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:_Trophic_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>: <a href="/wiki/Trophic_level" title="Trophic level">Trophic</a> components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">General</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abiotic_component" title="Abiotic component">Abiotic component</a></li> <li><a href="/wiki/Abiotic_stress" title="Abiotic stress">Abiotic stress</a></li> <li><a href="/wiki/Behavioral_ecology" title="Behavioral ecology">Behaviour</a></li> <li><a href="/wiki/Biogeochemical_cycle" title="Biogeochemical cycle">Biogeochemical cycle</a></li> <li><a href="/wiki/Biomass_(ecology)" title="Biomass (ecology)">Biomass</a></li> <li><a href="/wiki/Biotic_component" class="mw-redirect" title="Biotic component">Biotic component</a></li> <li><a href="/wiki/Biotic_stress" title="Biotic stress">Biotic stress</a></li> <li><a href="/wiki/Carrying_capacity" title="Carrying capacity">Carrying capacity</a></li> <li><a href="/wiki/Competition_(biology)" title="Competition (biology)">Competition</a></li> <li><a href="/wiki/Ecosystem" title="Ecosystem">Ecosystem</a></li> <li><a href="/wiki/Ecosystem_ecology" title="Ecosystem ecology">Ecosystem ecology</a></li> <li><a href="/wiki/Ecosystem_model" title="Ecosystem model">Ecosystem model</a></li> <li><a href="/wiki/Green_world_hypothesis" title="Green world hypothesis">Green world hypothesis</a></li> <li><a href="/wiki/Keystone_species" title="Keystone species">Keystone species</a></li> <li><a href="/wiki/List_of_feeding_behaviours" title="List of feeding behaviours">List of feeding behaviours</a></li> <li><a href="/wiki/Metabolic_theory_of_ecology" title="Metabolic theory of ecology">Metabolic theory of ecology</a></li> <li><a href="/wiki/Productivity_(ecology)" title="Productivity (ecology)">Productivity</a></li> <li><a href="/wiki/Resource_(biology)" title="Resource (biology)">Resource</a></li> <li><a href="/wiki/Restoration_ecology" class="mw-redirect" title="Restoration ecology">Restoration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Autotroph" title="Autotroph">Producers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Autotroph" title="Autotroph">Autotrophs</a></li> <li><a href="/wiki/Chemosynthesis" title="Chemosynthesis">Chemosynthesis</a></li> <li><a href="/wiki/Chemotroph" title="Chemotroph">Chemotrophs</a></li> <li><a href="/wiki/Foundation_species" title="Foundation species">Foundation species</a></li> <li><a href="/wiki/Kinetotroph" title="Kinetotroph">Kinetotrophs</a></li> <li><a href="/wiki/Mixotroph" title="Mixotroph">Mixotrophs</a></li> <li><a href="/wiki/Myco-heterotrophy" title="Myco-heterotrophy">Myco-heterotrophy</a></li> <li><a href="/wiki/Mycotroph" title="Mycotroph">Mycotroph</a></li> <li><a href="/wiki/Organotroph" title="Organotroph">Organotrophs</a></li> <li><a href="/wiki/Photoheterotroph" title="Photoheterotroph">Photoheterotrophs</a></li> <li><a href="/wiki/Photosynthesis" title="Photosynthesis">Photosynthesis</a></li> <li><a href="/wiki/Photosynthetic_efficiency" title="Photosynthetic efficiency">Photosynthetic efficiency</a></li> <li><a href="/wiki/Phototroph" title="Phototroph">Phototrophs</a></li> <li><a href="/wiki/Primary_nutritional_groups" title="Primary nutritional groups">Primary nutritional groups</a></li> <li><a href="/wiki/Primary_production" title="Primary production">Primary production</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Consumer_(food_chain)" title="Consumer (food chain)">Consumers</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Apex_predator" title="Apex predator">Apex predator</a></li> <li><a href="/wiki/Bacterivore" title="Bacterivore">Bacterivore</a></li> <li><a href="/wiki/Carnivore" title="Carnivore">Carnivores</a></li> <li><a href="/wiki/Chemoorganotroph" class="mw-redirect" title="Chemoorganotroph">Chemoorganotroph</a></li> <li><a href="/wiki/Foraging" title="Foraging">Foraging</a></li> <li><a href="/wiki/Generalist_and_specialist_species" title="Generalist and specialist species">Generalist and specialist species</a></li> <li><a href="/wiki/Intraguild_predation" title="Intraguild predation">Intraguild predation</a></li> <li><a href="/wiki/Herbivore" title="Herbivore">Herbivores</a></li> <li><a href="/wiki/Heterotroph" title="Heterotroph">Heterotroph</a></li> <li><a href="/wiki/Heterotrophic_nutrition" title="Heterotrophic nutrition">Heterotrophic nutrition</a></li> <li><a href="/wiki/Insectivore" title="Insectivore">Insectivore</a></li> <li><a href="/wiki/Mesopredator" title="Mesopredator">Mesopredators</a></li> <li><a href="/wiki/Mesopredator_release_hypothesis" title="Mesopredator release hypothesis">Mesopredator release hypothesis</a></li> <li><a href="/wiki/Omnivore" title="Omnivore">Omnivores</a></li> <li><a href="/wiki/Optimal_foraging_theory" title="Optimal foraging theory">Optimal foraging theory</a></li> <li><a href="/wiki/Planktivore" title="Planktivore">Planktivore</a></li> <li><a href="/wiki/Predation" title="Predation">Predation</a></li> <li><a href="/wiki/Prey_switching" title="Prey switching">Prey switching</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Decomposer" title="Decomposer">Decomposers</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Chemoorganoheterotrophy" class="mw-redirect" title="Chemoorganoheterotrophy">Chemoorganoheterotrophy</a></li> <li><a href="/wiki/Decomposition" title="Decomposition">Decomposition</a></li> <li><a href="/wiki/Detritivore" title="Detritivore">Detritivores</a></li> <li><a href="/wiki/Detritus" title="Detritus">Detritus</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Microorganism#Habitats_and_ecology" title="Microorganism">Microorganisms</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaea" title="Archaea">Archaea</a></li> <li><a href="/wiki/Bacteriophage" title="Bacteriophage">Bacteriophage</a></li> <li><a href="/wiki/Lithoautotroph" title="Lithoautotroph">Lithoautotroph</a></li> <li><a href="/wiki/Lithotroph" title="Lithotroph">Lithotrophy</a></li> <li><a href="/wiki/Marine_microorganisms" title="Marine microorganisms">Marine</a></li> <li><a href="/wiki/Microbial_cooperation" title="Microbial cooperation">Microbial cooperation</a></li> <li><a href="/wiki/Microbial_ecology" title="Microbial ecology">Microbial ecology</a></li> <li><a href="/wiki/Microbial_food_web" title="Microbial food web">Microbial food web</a></li> <li><a href="/wiki/Microbial_intelligence" title="Microbial intelligence">Microbial intelligence</a></li> <li><a href="/wiki/Microbial_loop" title="Microbial loop">Microbial loop</a></li> <li><a href="/wiki/Microbial_mat" title="Microbial mat">Microbial mat</a></li> <li><a href="/wiki/Microbial_metabolism" title="Microbial metabolism">Microbial metabolism</a></li> <li><a href="/wiki/Phage_ecology" title="Phage ecology">Phage ecology</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Food_web" title="Food web">Food webs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biomagnification" title="Biomagnification">Biomagnification</a></li> <li><a href="/wiki/Ecological_efficiency" title="Ecological efficiency">Ecological efficiency</a></li> <li><a href="/wiki/Ecological_pyramid" title="Ecological pyramid">Ecological pyramid</a></li> <li><a href="/wiki/Energy_flow_(ecology)" title="Energy flow (ecology)">Energy flow</a></li> <li><a href="/wiki/Food_chain" title="Food chain">Food chain</a></li> <li><a href="/wiki/Trophic_level" title="Trophic level">Trophic level</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Example webs</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Lake_ecosystem#Trophic_relationships" title="Lake ecosystem">Lakes</a></li> <li><a href="/wiki/River_ecosystem#Trophic_relationships" title="River ecosystem">Rivers</a></li> <li><a href="/wiki/Soil_food_web" title="Soil food web">Soil</a></li> <li><a href="/wiki/Tritrophic_interactions_in_plant_defense" title="Tritrophic interactions in plant defense">Tritrophic interactions in plant defense</a></li> <li><a href="/wiki/Marine_food_web" title="Marine food web">Marine food webs</a> <ul><li><a href="/wiki/Cold_seep" title="Cold seep">cold seeps</a></li> <li><a href="/wiki/Hydrothermal_vent#Biological_communities" title="Hydrothermal vent">hydrothermal vents</a></li> <li><a href="/wiki/Intertidal_ecology" title="Intertidal ecology">intertidal</a></li> <li><a href="/wiki/Kelp_forest#Trophic_ecology" title="Kelp forest">kelp forests</a></li> <li><a href="/wiki/North_Pacific_Gyre" title="North Pacific Gyre">North Pacific Gyre</a></li> <li><a href="/wiki/Ecology_of_the_San_Francisco_Estuary#Food_web" title="Ecology of the San Francisco Estuary">San Francisco Estuary</a></li> <li><a href="/wiki/Tide_pool" title="Tide pool">tide pool</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Processes</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Ascendency" title="Ascendency">Ascendency</a></li> <li><a href="/wiki/Bioaccumulation" title="Bioaccumulation">Bioaccumulation</a></li> <li><a href="/wiki/Cascade_effect_(ecology)" title="Cascade effect (ecology)">Cascade effect</a></li> <li><a href="/wiki/Climax_community" title="Climax community">Climax community</a></li> <li><a href="/wiki/Competitive_exclusion_principle" title="Competitive exclusion principle">Competitive exclusion principle</a></li> <li><a href="/wiki/Consumer%E2%80%93resource_interactions" title="Consumer–resource interactions">Consumer–resource interactions</a></li> <li><a href="/wiki/Copiotroph" title="Copiotroph">Copiotrophs</a></li> <li><a href="/wiki/Dominance_(ecology)" title="Dominance (ecology)">Dominance</a></li> <li><a href="/wiki/Ecological_network" title="Ecological network">Ecological network</a></li> <li><a href="/wiki/Ecological_succession" title="Ecological succession">Ecological succession</a></li> <li><a href="/wiki/Energy_quality" title="Energy quality">Energy quality</a></li> <li><a href="/wiki/Energy_systems_language" title="Energy systems language">Energy systems language</a></li> <li><a href="/wiki/F-ratio_(oceanography)" title="F-ratio (oceanography)">f-ratio</a></li> <li><a href="/wiki/Feed_conversion_ratio" title="Feed conversion ratio">Feed conversion ratio</a></li> <li><a href="/wiki/Feeding_frenzy" title="Feeding frenzy">Feeding frenzy</a></li> <li><a href="/wiki/Mesotrophic_soil" title="Mesotrophic soil">Mesotrophic soil</a></li> <li><a href="/wiki/Nutrient_cycle" title="Nutrient cycle">Nutrient cycle</a></li> <li><a href="/wiki/Oligotroph" title="Oligotroph">Oligotroph</a></li> <li><a href="/wiki/Paradox_of_the_plankton" title="Paradox of the plankton">Paradox of the plankton</a></li> <li><a href="/wiki/Trophic_cascade" title="Trophic cascade">Trophic cascade</a></li> <li><a href="/wiki/Trophic_mutualism" title="Trophic mutualism">Trophic mutualism</a></li> <li><a href="/wiki/Trophic_state_index" title="Trophic state index">Trophic state index</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Defense,<br />counter</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Animal_coloration" title="Animal coloration">Animal coloration</a></li> <li><a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">Anti-predator adaptations</a></li> <li><a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></li> <li><a href="/wiki/Deimatic_behaviour" title="Deimatic behaviour">Deimatic behaviour</a></li> <li><a href="/wiki/Herbivore_adaptations_to_plant_defense" title="Herbivore adaptations to plant defense">Herbivore adaptations to plant defense</a></li> <li><a href="/wiki/Mimicry" title="Mimicry">Mimicry</a></li> <li><a href="/wiki/Plant_defense_against_herbivory" title="Plant defense against herbivory">Plant defense against herbivory</a></li> <li><a href="/wiki/Shoaling_and_schooling#Predator_avoidance" title="Shoaling and schooling">Predator avoidance in schooling fish</a></li></ul> </div></td></tr></tbody></table></div><div class="navbox-styles"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236075235"></div><div role="navigation" class="navbox" aria-labelledby="Ecology:_Modelling_ecosystems:_Other_components" style="padding:3px"><table class="nowraplinks hlist mw-collapsible uncollapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1129693374"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239400231"><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Modelling_ecosystems" title="Template:Modelling ecosystems"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Modelling_ecosystems" title="Template talk:Modelling ecosystems"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Modelling_ecosystems" title="Special:EditPage/Template:Modelling ecosystems"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Ecology:_Modelling_ecosystems:_Other_components" style="font-size:114%;margin:0 4em"><a href="/wiki/Ecology" title="Ecology">Ecology</a>: <a href="/wiki/Ecosystem_model" title="Ecosystem model">Modelling ecosystems</a>: Other components</div></th></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a href="/wiki/Population_ecology" title="Population ecology">Population<br />ecology</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Abundance_(ecology)" title="Abundance (ecology)">Abundance</a></li> <li><a href="/wiki/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Consumer-resource_model" title="Consumer-resource model">Consumer-resource model</a></li> <li><a href="/wiki/Depensation" title="Depensation">Depensation</a></li> <li><a href="/wiki/Ecological_yield" title="Ecological yield">Ecological yield</a></li> <li><a href="/wiki/Effective_population_size" title="Effective population size">Effective population size</a></li> <li><a href="/wiki/Intraspecific_competition" title="Intraspecific competition">Intraspecific competition</a></li> <li><a href="/wiki/Logistic_function" title="Logistic function">Logistic function</a></li> <li><a href="/wiki/Malthusian_growth_model" title="Malthusian growth model">Malthusian growth model</a></li> <li><a href="/wiki/Maximum_sustainable_yield" title="Maximum sustainable yield">Maximum sustainable yield</a></li> <li><a href="/wiki/Overpopulation" title="Overpopulation">Overpopulation</a></li> <li><a href="/wiki/Overexploitation" title="Overexploitation">Overexploitation</a></li> <li><a href="/wiki/Population_cycle" title="Population cycle">Population cycle</a></li> <li><a href="/wiki/Population_dynamics" title="Population dynamics">Population dynamics</a></li> <li><a href="/wiki/Population_model" title="Population model">Population modeling</a></li> <li><a href="/wiki/Population_size" title="Population size">Population size</a></li> <li><a href="/wiki/Lotka%E2%80%93Volterra_equations" title="Lotka–Volterra equations">Predator–prey (Lotka–Volterra) equations</a></li> <li><a href="/wiki/Recruitment_(biology)" title="Recruitment (biology)">Recruitment</a></li> <li><a href="/wiki/Small_population_size" title="Small population size">Small population size</a></li> <li><a href="/wiki/Ecological_stability" title="Ecological stability">Stability</a> <ul><li><a href="/wiki/Ecological_resilience" title="Ecological resilience">Resilience</a></li> <li><a href="/wiki/Resistance_(ecology)" title="Resistance (ecology)">Resistance</a></li></ul></li> <li><a href="/wiki/Random_generalized_Lotka%E2%80%93Volterra_model" title="Random generalized Lotka–Volterra model">Random generalized Lotka–Volterra model</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biodiversity" title="Biodiversity">Biodiversity</a></li> <li><a href="/wiki/Density_dependence" title="Density dependence">Density-dependent inhibition</a></li> <li><a href="/wiki/Ecological_effects_of_biodiversity" title="Ecological effects of biodiversity">Ecological effects of biodiversity</a></li> <li><a href="/wiki/Ecological_extinction" title="Ecological extinction">Ecological extinction</a></li> <li><a href="/wiki/Endemism" title="Endemism">Endemic species</a></li> <li><a href="/wiki/Flagship_species" title="Flagship species">Flagship species</a></li> <li><a href="/wiki/Ordination_(statistics)" title="Ordination (statistics)">Gradient analysis</a></li> <li><a href="/wiki/Bioindicator" title="Bioindicator">Indicator species</a></li> <li><a href="/wiki/Introduced_species" title="Introduced species">Introduced species</a></li> <li><a href="/wiki/Invasive_species" title="Invasive species">Invasive species</a> / <a href="/wiki/Native_species" title="Native species">Native species</a></li> <li><a href="/wiki/Latitudinal_gradients_in_species_diversity" title="Latitudinal gradients in species diversity">Latitudinal gradients in species diversity</a></li> <li><a href="/wiki/Minimum_viable_population" title="Minimum viable population">Minimum viable population</a></li> <li><a href="/wiki/Unified_neutral_theory_of_biodiversity" title="Unified neutral theory of biodiversity">Neutral theory</a></li> <li><a href="/wiki/Occupancy%E2%80%93abundance_relationship" title="Occupancy–abundance relationship">Occupancy–abundance relationship</a></li> <li><a href="/wiki/Population_viability_analysis" title="Population viability analysis">Population viability analysis</a></li> <li><a href="/wiki/Priority_effect" title="Priority effect">Priority effect</a></li> <li><a href="/wiki/Rapoport%27s_rule" title="Rapoport's rule">Rapoport's rule</a></li> <li><a href="/wiki/Relative_abundance_distribution" title="Relative abundance distribution">Relative abundance distribution</a></li> <li><a href="/wiki/Relative_species_abundance" title="Relative species abundance">Relative species abundance</a></li> <li><a href="/wiki/Species_diversity" title="Species diversity">Species diversity</a></li> <li><a href="/wiki/Species_homogeneity" title="Species homogeneity">Species homogeneity</a></li> <li><a href="/wiki/Species_richness" title="Species richness">Species richness</a></li> <li><a href="/wiki/Species_distribution" title="Species distribution">Species distribution</a></li> <li><a href="/wiki/Species%E2%80%93area_relationship" title="Species–area relationship">Species–area curve</a></li> <li><a href="/wiki/Umbrella_species" title="Umbrella species">Umbrella species</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em">Species<br />interaction</th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Antibiosis" title="Antibiosis">Antibiosis</a></li> <li><a href="/wiki/Biological_interaction" title="Biological interaction">Biological interaction</a></li> <li><a href="/wiki/Commensalism" title="Commensalism">Commensalism</a></li> <li><a href="/wiki/Community_(ecology)" title="Community (ecology)">Community ecology</a></li> <li><a href="/wiki/Ecological_facilitation" title="Ecological facilitation">Ecological facilitation</a></li> <li><a href="/wiki/Interspecific_competition" title="Interspecific competition">Interspecific competition</a></li> <li><a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a></li> <li><a href="/wiki/Parasitism" title="Parasitism">Parasitism</a></li> <li><a href="/wiki/Storage_effect" title="Storage effect">Storage effect</a></li> <li><a href="/wiki/Symbiosis" title="Symbiosis">Symbiosis</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:7.5em"><a class="mw-selflink selflink">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 state">Alternative stable state</a></li> <li><a href="/wiki/Balance_of_nature" title="Balance of nature">Balance of nature</a></li> <li><a href="/wiki/Biological_data_visualization" title="Biological data visualization">Biological data visualization</a></li> <li><a href="/wiki/Ecological_economics" title="Ecological economics">Ecological economics</a></li> <li><a href="/wiki/Ecological_footprint" title="Ecological footprint">Ecological footprint</a></li> <li><a href="/wiki/Ecological_forecasting" title="Ecological forecasting">Ecological forecasting</a></li> <li><a href="/wiki/Environmental_humanities" title="Environmental humanities">Ecological humanities</a></li> <li><a href="/wiki/Ecological_stoichiometry" title="Ecological stoichiometry">Ecological stoichiometry</a></li> <li><a href="/wiki/Ecopath" title="Ecopath">Ecopath</a></li> <li><a href="/wiki/Ecosystem_based_fisheries" class="mw-redirect" title="Ecosystem based fisheries">Ecosystem based fisheries</a></li> <li><a href="/wiki/Endolith" title="Endolith">Endolith</a></li> <li><a href="/wiki/Evolutionary_ecology" title="Evolutionary ecology">Evolutionary ecology</a></li> <li><a href="/wiki/Functional_ecology" title="Functional ecology">Functional ecology</a></li> <li><a href="/wiki/Industrial_ecology" title="Industrial ecology">Industrial ecology</a></li> <li><a href="/wiki/Macroecology" title="Macroecology">Macroecology</a></li> <li><a href="/wiki/Microecosystem" title="Microecosystem">Microecosystem</a></li> <li><a href="/wiki/Natural_environment" title="Natural environment">Natural environment</a></li> <li><a href="/wiki/Regime_shift" title="Regime shift">Regime shift</a></li> <li><a href="/wiki/Sexecology" title="Sexecology">Sexecology</a></li> <li><a href="/wiki/Systems_ecology" title="Systems ecology">Systems ecology</a></li> <li><a href="/wiki/Urban_ecology" title="Urban ecology">Urban ecology</a></li> <li><a href="/wiki/Theoretical_ecology" title="Theoretical ecology">Theoretical ecology</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/Outline_of_ecology" title="Outline of ecology">Outline of ecology</a></div></td></tr></tbody></table></div>    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