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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_rewrite plainlinks metadata ambox ambox-style ambox-lead_rewrite" role="presentation"><tbody><tr><td class="mbox-image"><div class="mbox-image-div"><span typeof="mw:File"><span><img alt="" src="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/40px-Edit-clear.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/60px-Edit-clear.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/f/f2/Edit-clear.svg/80px-Edit-clear.svg.png 2x" data-file-width="48" data-file-height="48" /></span></span></div></td><td class="mbox-text"><div class="mbox-text-span">The article's <a href="/wiki/Wikipedia:Manual_of_Style/Lead_section" title="Wikipedia:Manual of Style/Lead section">lead section</a> <b>may need to be rewritten</b>.<span class="hide-when-compact"> Please help <a class="external text" href="https://en.wikipedia.org/w/index.php?title=Spatial_organization&action=edit">improve the lead</a> and read the <a href="/wiki/Wikipedia:Manual_of_Style/Lead_section" title="Wikipedia:Manual of Style/Lead section">lead layout guide</a>.</span> <span class="date-container"><i>(<span class="date">February 2013</span>)</i></span><span class="hide-when-compact"><i> (<small><a href="/wiki/Help:Maintenance_template_removal" title="Help:Maintenance template removal">Learn how and when to remove this message</a></small>)</i></span></div></td></tr></tbody></table> <p><b>Spatial organization</b> can be observed when components of an <a href="/wiki/Abiotic" class="mw-redirect" title="Abiotic">abiotic</a> or biological group are arranged non-randomly in space. Abiotic patterns, such as the ripple formations in <a href="/wiki/Dune" title="Dune">sand dunes</a> or the oscillating wave patterns of the <a href="/wiki/Belousov%E2%80%93Zhabotinsky_reaction" title="Belousov–Zhabotinsky reaction">Belousov–Zhabotinsky reaction</a><sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> emerge after thousands of particles interact millions of times. On the other hand, individuals in biological groups may be arranged non-randomly due to selfish behavior, <a href="/wiki/Dominance_hierarchy" title="Dominance hierarchy">dominance</a> interactions, or cooperative behavior. <a href="/wiki/W._D._Hamilton" title="W. D. Hamilton">W. D. Hamilton</a> (1971) proposed that in a non-related "<a href="/wiki/Herd" title="Herd">herd</a>" of animals, the spatial organization is likely a result of the selfish interests of individuals trying to acquire food or avoid <a href="/wiki/Predation" title="Predation">predation</a>.<sup id="cite_ref-2" class="reference"><a href="#cite_note-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> On the other hand, spatial arrangements have also been observed among highly related members of <a href="/wiki/Eusocial" class="mw-redirect" title="Eusocial">eusocial</a> groups, suggesting that the arrangement of individuals may provide advantages for the <a href="/wiki/Group_selection" title="Group selection">group</a>.<sup id="cite_ref-wilson1976_3-0" class="reference"><a href="#cite_note-wilson1976-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>Spatial organization can be defined as how something is organized internally as well as externally concerning its surroundings. It can be viewed both on a large or small scale and may also include in its discussion, the factors and conditions that lead to a specific organization pattern.<sup id="cite_ref-4" class="reference"><a href="#cite_note-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> In the frame of biology this may include the abiotic and biotic factors that a population is immersed in. In the frame of civilization, spatial organization may include if an area is rural, suburban, or urban. Spatial organization can even include how one manages and organizes their lawn to meet social norms. <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> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="In_eusocial_insects">In eusocial insects</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_organization&action=edit&section=1" title="Edit section: In eusocial insects"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Paint_marked_ants.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Paint_marked_ants.JPG/175px-Paint_marked_ants.JPG" decoding="async" width="175" height="182" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Paint_marked_ants.JPG/263px-Paint_marked_ants.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f0/Paint_marked_ants.JPG/350px-Paint_marked_ants.JPG 2x" data-file-width="1357" data-file-height="1408" /></a><figcaption> Spatial patterns exhibited by ants (<i>Temnothorax rugatulus</i>) can be determined after each individual is painted with a distinguishing mark.</figcaption></figure> <p>Individuals in a <a href="/wiki/Eusociality" title="Eusociality">social insect</a> colony can be spatially organized, or arranged non-randomly inside the nest. These miniature territories, or spatial fidelity zones have been described in <a href="/wiki/Honey_bee" title="Honey bee">honey bees</a> (<i>Apis mellifera</i><sup id="cite_ref-seeley1982_6-0" class="reference"><a href="#cite_note-seeley1982-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup>), <a href="/wiki/Ant" title="Ant">ants</a> (<i>Odontomachus brunneus</i>;<sup id="cite_ref-powell1999_7-0" class="reference"><a href="#cite_note-powell1999-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> <i><a href="/wiki/Temnothorax_albipennis" title="Temnothorax albipennis">Temnothorax albipennis</a></i>;<sup id="cite_ref-sendova1995_8-0" class="reference"><a href="#cite_note-sendova1995-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> <i>Pheidole dentata</i><sup id="cite_ref-wilson1976_3-1" class="reference"><a href="#cite_note-wilson1976-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>), and <a href="/wiki/Paper_wasp" title="Paper wasp">paper wasps</a> (<i>Polistes dominulus</i>,<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> <i>Ropalidia revolutionalis</i><sup id="cite_ref-robson2000_10-0" class="reference"><a href="#cite_note-robson2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup>). While residing in these zones, workers perform the <a href="/wiki/Task_allocation_and_partitioning_of_social_insects" class="mw-redirect" title="Task allocation and partitioning of social insects">task</a> appropriate to the area they reside. For example, individuals that remain in the center of an ant nest are more likely to feed larvae, whereas individuals found at the periphery of the nest are more likely to forage.<sup id="cite_ref-powell1999_7-1" class="reference"><a href="#cite_note-powell1999-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-sendova1995_8-1" class="reference"><a href="#cite_note-sendova1995-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> <a href="/wiki/E._O._Wilson" title="E. O. Wilson">E. O. Wilson</a> proposed that by remaining in small, non-random areas inside the nest, the distance an individual moves between tasks may be minimized, and overall colony efficiency would increase.<sup id="cite_ref-wilson1976_3-2" class="reference"><a href="#cite_note-wilson1976-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="In_the_nest">In the nest</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_organization&action=edit&section=2" title="Edit section: In the nest"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading4"><h4 id="Foraging-for-work">Foraging-for-work</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_organization&action=edit&section=3" title="Edit section: Foraging-for-work"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There are a variety of ways in which individuals can divide space inside a nest. According to the "foraging-for-work" hypothesis, adult workers begin performing <a href="/wiki/Task_allocation_and_partitioning_of_social_insects" class="mw-redirect" title="Task allocation and partitioning of social insects">tasks</a> in the area of the nest where they emerged, and gradually move towards the periphery of the nest as demands to perform particular tasks change. This hypothesis is based on two observations: "(1) that there is spatial structure in the layout of tasks in social insect colonies and (2) that workers first become adults in or around the center of the nest".<sup id="cite_ref-franks1994_11-0" class="reference"><a href="#cite_note-franks1994-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Individuals can remain in an area for an extended period, as long as tasks need to be performed there. Over time, an individual's zone may shift as tasks are accomplished and workers search for other areas where tasks need to be performed. <a href="/wiki/Honey_bee" title="Honey bee">Honey bees</a>, for example, begin their adult life caring for brood located in the area near where they emerged (i.e. nurse bees). Eventually, workers move away from the brood rearing area and begin to perform other tasks, such as food storage, guarding, or foraging.<sup id="cite_ref-seeley1982_6-1" class="reference"><a href="#cite_note-seeley1982-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading4"><h4 id="Dominance_hierarchy">Dominance hierarchy</h4><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_organization&action=edit&section=4" title="Edit section: Dominance hierarchy"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Polistes_flavus_on_the_nest.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/4/44/Polistes_flavus_on_the_nest.JPG/175px-Polistes_flavus_on_the_nest.JPG" decoding="async" width="175" height="171" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/4/44/Polistes_flavus_on_the_nest.JPG/263px-Polistes_flavus_on_the_nest.JPG 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/4/44/Polistes_flavus_on_the_nest.JPG/350px-Polistes_flavus_on_the_nest.JPG 2x" data-file-width="1228" data-file-height="1201" /></a><figcaption> The dominant paper wasp (<i>Polistes flavus</i>) remains in the center of the nest while subordinate wasps are often at the edge or off the nest.</figcaption></figure> <p>Space inside the nest may also be divided as a result of <a href="/wiki/Dominance_(ethology)" class="mw-redirect" title="Dominance (ethology)">dominance</a> interactions. For example, in <a href="/wiki/Paper_wasp" title="Paper wasp">paper wasp</a> colonies, a single inseminated queen may find (initiate) a colony after waking up from hibernation (overwintering). However, it is common in many species that multiple inseminated females join these foundresses instead of founding their own nest.<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> When multiple inseminated females found a colony together, the colony grows quickly, yet only one individual will become the primary egg-layer.<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> Through a series of dominance interactions, the most aggressive wasp will emerge as the dominant individual and will become the primary egg-layer for the group (the prime role for ensuring your genes are passed on to subsequent generations), whereas the remaining subordinate wasps will perform other tasks, such as nest construction or foraging.<sup id="cite_ref-robson2000_10-1" class="reference"><a href="#cite_note-robson2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> There is evidence that these dominance interactions affect the spatial zones individuals occupy as well. In paper wasps (<i>Ropalidia revolutionalis</i>), as well as in the ant species <i>Odontomachus brunneus</i>,<sup id="cite_ref-powell1999_7-2" class="reference"><a href="#cite_note-powell1999-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> dominant individuals are more likely to reside in the central areas of the nest, where they take care of the brood, while the subordinate individuals are pushed towards the edge, where they are more likely to forage. It is unknown whether division of space or establishment of dominance occurs first and if the other is a result of it. </p> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Marked_Bumble_bees.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/en/thumb/5/5f/Marked_Bumble_bees.jpg/175px-Marked_Bumble_bees.jpg" decoding="async" width="175" height="164" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/5/5f/Marked_Bumble_bees.jpg/263px-Marked_Bumble_bees.jpg 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/5/5f/Marked_Bumble_bees.jpg/350px-Marked_Bumble_bees.jpg 2x" data-file-width="480" data-file-height="451" /></a><figcaption> Bumble bees, <i>Bombus impatiens</i> individually marked with plastic number tags</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Outside_the_nest">Outside the nest</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_organization&action=edit&section=5" title="Edit section: Outside the nest"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>There is also evidence that foragers, which are the insects that leave the nest to collect the valuable resources for the developing colony, can divide space outside the nest. Makino & Sakai showed that <a href="/wiki/Bumblebee" title="Bumblebee">bumble bee</a> foragers maintain foraging zones in flower patches, which means that bees consistently return to the same areas within a patch and there is little overlap between individuals.<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> These zones can expand and contract when neighboring foragers are removed or introduced, respectively.<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> By dividing foraging patches into miniature ‘foraging territories’, individuals can maximize the number of flowers visited with minimal interruptions or competition between foragers. These ‘foraging territories’ divided among individuals from the same colony are the result of <a href="/wiki/Self-organization" title="Self-organization">self-organization</a> among the foragers; that is, no lead forager is dictating where the bees will forage. Instead, the maintenance of these foraging zones is due to simple rules followed by each individual forager. Studies to determine these “rules” are an important area of research in <a href="/wiki/Computer_science" title="Computer science">computer science</a>, basic <a href="/wiki/Biology" title="Biology">biology</a>, <a href="/wiki/Behavioral_ecology" title="Behavioral ecology">behavioral ecology</a>, and <a href="/wiki/Mathematical_model" title="Mathematical model">mathematic modeling</a>. </p> <div class="mw-heading mw-heading3"><h3 id="As_an_emergent_property_of_a_self-organized_system">As an emergent property of a self-organized system</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Spatial_organization&action=edit&section=6" title="Edit section: As an emergent property of a self-organized system"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>The <a href="/wiki/Self-organization" title="Self-organization">self-organization</a> observed in foraging territories is a microcosm for the self-organization seen in the entire <a href="/wiki/Eusociality" title="Eusociality">colony</a>. Spatial organization observed across social insect colonies can be considered an <a href="/wiki/Emergence" title="Emergence">emergent</a> property of a self-organized <a href="/wiki/Complex_system" title="Complex system">complex system</a>. It is self-organized because no leader is dictating where each individual will reside, nor which task an individual will perform once they get there.<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> Instead, zones may be a by-product of division of labor, whereby individuals end up in a particular location for a period based on the task they perform,<sup id="cite_ref-franks1994_11-1" class="reference"><a href="#cite_note-franks1994-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> or dominance interactions, whereby dominant individuals are granted access to the most desirable places inside the nest.<sup id="cite_ref-powell1999_7-3" class="reference"><a href="#cite_note-powell1999-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-robson2000_10-2" class="reference"><a href="#cite_note-robson2000-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Spatial patterns exhibited by individuals of social insect colonies are not obvious, because it is difficult to observe and differentiate among individuals inside a nest cavity or flying across a foraging patch. However, when careful attention is given to the individual worker, the spatial organization of workers in the nest becomes apparent. </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_organization&action=edit&section=7" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Cellular_automaton" title="Cellular automaton">Cellular automaton</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_organization&action=edit&section=8" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles: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"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-1"><span class="mw-cite-backlink"><b><a href="#cite_ref-1">^</a></b></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="CITEREFBall" class="citation book cs1">Ball, P. <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/selfmadetapestry00ball_0"><i>The Self-Made Tapestry: Pattern formation in nature</i></a></span>. 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href="/wiki/Ant_mill" title="Ant mill">ant mill</a></li> <li><a href="/wiki/Symmetry_breaking_of_escaping_ants" title="Symmetry breaking of escaping ants">symmetry breaking of escaping ants</a></li></ul></li> <li><a href="/wiki/Shoaling_and_schooling" title="Shoaling and schooling">Shoaling and schooling</a> <ul><li><a href="/wiki/Bait_ball" title="Bait ball">bait ball</a></li></ul></li> <li><a href="/wiki/Swarm_behaviour" title="Swarm behaviour">Swarming behaviour</a></li> <li><a href="/wiki/Swarming_(honey_bee)" title="Swarming (honey bee)">Swarming (honey bee)</a></li> <li><a href="/wiki/Swarming_motility" title="Swarming motility">Swarming motility</a></li></ul> </div></td><td class="noviewer navbox-image" rowspan="6" style="width:1px;padding:0 0 0 2px"><div><span typeof="mw:File"><a href="/wiki/File:Auklet_flock_Shumagins_1986.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/140px-Auklet_flock_Shumagins_1986.jpg" decoding="async" width="140" height="95" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/210px-Auklet_flock_Shumagins_1986.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Auklet_flock_Shumagins_1986.jpg/280px-Auklet_flock_Shumagins_1986.jpg 2x" data-file-width="3756" data-file-height="2561" /></a></span></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Animal_migration" title="Animal migration">Animal migration</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/Animal_migration" title="Animal migration">Animal migration</a> <ul><li><a href="/wiki/Altitudinal_migration" title="Altitudinal migration">altitudinal</a></li> <li><a href="/wiki/Animal_migration_tracking" title="Animal migration tracking">tracking</a> <ul><li><a href="/wiki/History_of_wildlife_tracking_technology" title="History of wildlife tracking technology">history</a></li></ul></li> <li><a href="/wiki/Coded_wire_tag" title="Coded wire tag">coded wire tag</a></li></ul></li> <li><a href="/wiki/Bird_migration" title="Bird migration">Bird migration</a> <ul><li><a href="/wiki/Category:Bird_migration_flyways" title="Category:Bird migration flyways">flyways</a></li> <li><a href="/wiki/Reverse_migration_(birds)" title="Reverse migration (birds)">reverse migration</a></li></ul></li> <li><a href="/wiki/Cell_migration" title="Cell migration">Cell migration</a></li> <li><a href="/wiki/Fish_migration" title="Fish migration">Fish migration</a> <ul><li><a href="/wiki/Diel_vertical_migration" title="Diel vertical migration">diel vertical</a></li> <li><a href="/wiki/Lessepsian_migration" title="Lessepsian migration">Lessepsian</a></li> <li><a href="/wiki/Salmon_run" title="Salmon run">salmon run</a></li> <li><a href="/wiki/Sardine_run" title="Sardine run">sardine run</a></li></ul></li> <li><a href="/wiki/Homing_(biology)" title="Homing (biology)">Homing</a> <ul><li><a href="/wiki/Natal_homing" title="Natal homing">natal</a></li> <li><a href="/wiki/Philopatry" title="Philopatry">philopatry</a></li></ul></li> <li><a href="/wiki/Insect_migration" title="Insect migration">Insect migration</a> <ul><li><a href="/wiki/Lepidoptera_migration" title="Lepidoptera migration">butterflies</a> <ul><li><a href="/wiki/Monarch_butterfly_migration" title="Monarch butterfly migration">monarch</a></li></ul></li></ul></li> <li><a href="/wiki/Sea_turtle_migration" title="Sea turtle migration">Sea turtle migration</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_behaviour#Models" title="Swarm behaviour">Swarm algorithms</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/Agent-based_model" title="Agent-based model">Agent-based models</a></li> <li><a href="/wiki/Ant_colony_optimization_algorithm" class="mw-redirect" title="Ant colony optimization algorithm">Ant colony optimization</a></li> <li><a href="/wiki/Boids" title="Boids">Boids</a></li> <li><a href="/wiki/Crowd_simulation" title="Crowd simulation">Crowd simulation</a></li> <li><a href="/wiki/Particle_swarm_optimization" title="Particle swarm optimization">Particle swarm optimization</a></li> <li><a href="/wiki/Swarm_intelligence" title="Swarm intelligence">Swarm intelligence</a></li> <li><a href="/wiki/Swarm_(simulation)" title="Swarm (simulation)">Swarm (simulation)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Collective_motion" title="Collective motion">Collective motion</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/Active_matter" title="Active matter">Active matter</a></li> <li><a href="/wiki/Collective_motion" title="Collective motion">Collective motion</a></li> <li><a href="/wiki/Self-propelled_particles" title="Self-propelled particles">Self-propelled particles</a> <ul><li><a href="/wiki/Clustering_of_self-propelled_particles" title="Clustering of self-propelled particles">clustering</a></li></ul></li> <li><a href="/wiki/Vicsek_model" title="Vicsek model">Vicsek model</a></li> <li><a href="/wiki/BIO-LGCA" title="BIO-LGCA">BIO-LGCA</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Swarm_robotics" title="Swarm robotics">Swarm robotics</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/Ant_robotics" title="Ant robotics">Ant robotics</a></li> <li><a href="/wiki/Microbotics" title="Microbotics">Microbotics</a></li> <li><a href="/wiki/Nanorobotics" title="Nanorobotics">Nanorobotics</a></li> <li><a href="/wiki/Swarm_robotics" title="Swarm robotics">Swarm robotics</a></li> <li><a href="/wiki/Symbrion" title="Symbrion">Symbrion</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Related topics</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/Allee_effect" title="Allee effect">Allee effect</a></li> <li><a href="/wiki/Animal_navigation" title="Animal navigation">Animal navigation</a></li> <li><a href="/wiki/Collective_intelligence" title="Collective intelligence">Collective intelligence</a></li> <li><a href="/wiki/Decentralised_system" title="Decentralised system">Decentralised system</a></li> <li><a href="/wiki/Eusociality" title="Eusociality">Eusociality</a></li> <li><a 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