CINXE.COM

<!DOCTYPE html> <html class="client-nojs vector-feature-language-in-header-enabled vector-feature-language-in-main-page-header-disabled vector-feature-sticky-header-disabled vector-feature-page-tools-pinned-disabled vector-feature-toc-pinned-clientpref-1 vector-feature-main-menu-pinned-disabled vector-feature-limited-width-clientpref-1 vector-feature-limited-width-content-enabled vector-feature-custom-font-size-clientpref-1 vector-feature-appearance-pinned-clientpref-1 vector-feature-night-mode-enabled skin-theme-clientpref-day vector-toc-available" lang="en" dir="ltr"> <head> <meta charset="UTF-8"> <title>Beetle - Wikipedia</title> <script>(function(){var className="client-js vector-feature-language-in-header-enabled vector-feature-language-in-main-page-header-disabled vector-feature-sticky-header-disabled vector-feature-page-tools-pinned-disabled vector-feature-toc-pinned-clientpref-1 vector-feature-main-menu-pinned-disabled vector-feature-limited-width-clientpref-1 vector-feature-limited-width-content-enabled vector-feature-custom-font-size-clientpref-1 vector-feature-appearance-pinned-clientpref-1 vector-feature-night-mode-enabled skin-theme-clientpref-day vector-toc-available";var cookie=document.cookie.match(/(?:^|; )enwikimwclientpreferences=([^;]+)/);if(cookie){cookie[1].split('%2C').forEach(function(pref){className=className.replace(new RegExp('(^| )'+pref.replace(/-clientpref-\w+$|[^\w-]+/g,'')+'-clientpref-\\w+( |$)'),'$1'+pref+'$2');});}document.documentElement.className=className;}());RLCONF={"wgBreakFrames":false,"wgSeparatorTransformTable":["",""],"wgDigitTransformTable":["",""],"wgDefaultDateFormat":"dmy", "wgMonthNames":["","January","February","March","April","May","June","July","August","September","October","November","December"],"wgRequestId":"d948f592-1437-4eb1-9954-f34a724ced5f","wgCanonicalNamespace":"","wgCanonicalSpecialPageName":false,"wgNamespaceNumber":0,"wgPageName":"Beetle","wgTitle":"Beetle","wgCurRevisionId":1257880098,"wgRevisionId":1257880098,"wgArticleId":7044,"wgIsArticle":true,"wgIsRedirect":false,"wgAction":"view","wgUserName":null,"wgUserGroups":["*"],"wgCategories":["Pages using the WikiHiero extension","CS1 German-language sources (de)","CS1 Dutch-language sources (nl)","CS1 Russian-language sources (ru)","CS1 French-language sources (fr)","CS1 maint: numeric names: authors list","Articles with short description","Short description matches Wikidata","Good articles","Use mdy dates from September 2012","Articles with 'species' microformats","Articles containing potentially dated statements from 2015","All articles containing potentially dated statements", "All articles with unsourced statements","Articles with unsourced statements from June 2023","Articles with German-language sources (de)","Webarchive template wayback links","Taxonbars with 20–24 taxon IDs","Beetles","Extant Pennsylvanian first appearances","Insects in culture"],"wgPageViewLanguage":"en","wgPageContentLanguage":"en","wgPageContentModel":"wikitext","wgRelevantPageName":"Beetle","wgRelevantArticleId":7044,"wgIsProbablyEditable":true,"wgRelevantPageIsProbablyEditable":true,"wgRestrictionEdit":[],"wgRestrictionMove":[],"wgNoticeProject":"wikipedia","wgCiteReferencePreviewsActive":false,"wgFlaggedRevsParams":{"tags":{"status":{"levels":1}}},"wgMediaViewerOnClick":true,"wgMediaViewerEnabledByDefault":true,"wgPopupsFlags":0,"wgVisualEditor":{"pageLanguageCode":"en","pageLanguageDir":"ltr","pageVariantFallbacks":"en"},"wgMFDisplayWikibaseDescriptions":{"search":true,"watchlist":true,"tagline":false,"nearby":true},"wgWMESchemaEditAttemptStepOversample":false,"wgWMEPageLength" :200000,"wgRelatedArticlesCompat":[],"wgCentralAuthMobileDomain":false,"wgEditSubmitButtonLabelPublish":true,"wgULSPosition":"interlanguage","wgULSisCompactLinksEnabled":false,"wgVector2022LanguageInHeader":true,"wgULSisLanguageSelectorEmpty":false,"wgWikibaseItemId":"Q22671","wgCheckUserClientHintsHeadersJsApi":["brands","architecture","bitness","fullVersionList","mobile","model","platform","platformVersion"],"GEHomepageSuggestedEditsEnableTopics":true,"wgGETopicsMatchModeEnabled":false,"wgGEStructuredTaskRejectionReasonTextInputEnabled":false,"wgGELevelingUpEnabledForUser":false};RLSTATE={"ext.globalCssJs.user.styles":"ready","site.styles":"ready","user.styles":"ready","ext.globalCssJs.user":"ready","user":"ready","user.options":"loading","ext.cite.styles":"ready","mediawiki.page.gallery.styles":"ready","ext.wikihiero":"ready","skins.vector.search.codex.styles":"ready","skins.vector.styles":"ready","skins.vector.icons":"ready","jquery.makeCollapsible.styles":"ready", "ext.wikimediamessages.styles":"ready","ext.visualEditor.desktopArticleTarget.noscript":"ready","ext.uls.interlanguage":"ready","wikibase.client.init":"ready","ext.wikimediaBadges":"ready"};RLPAGEMODULES=["ext.cite.ux-enhancements","mediawiki.page.gallery","mediawiki.page.media","ext.scribunto.logs","site","mediawiki.page.ready","jquery.makeCollapsible","mediawiki.toc","skins.vector.js","ext.centralNotice.geoIP","ext.centralNotice.startUp","ext.gadget.ReferenceTooltips","ext.gadget.switcher","ext.urlShortener.toolbar","ext.centralauth.centralautologin","mmv.bootstrap","ext.popups","ext.visualEditor.desktopArticleTarget.init","ext.visualEditor.targetLoader","ext.echo.centralauth","ext.eventLogging","ext.wikimediaEvents","ext.navigationTiming","ext.uls.interface","ext.cx.eventlogging.campaigns","ext.cx.uls.quick.actions","wikibase.client.vector-2022","ext.checkUser.clientHints","ext.growthExperiments.SuggestedEditSession","wikibase.sidebar.tracking"];</script> <script>(RLQ=window.RLQ||[]).push(function(){mw.loader.impl(function(){return["user.options@12s5i",function($,jQuery,require,module){mw.user.tokens.set({"patrolToken":"+\\","watchToken":"+\\","csrfToken":"+\\"}); }];});});</script> <link rel="stylesheet" href="/w/load.php?lang=en&modules=ext.cite.styles%7Cext.uls.interlanguage%7Cext.visualEditor.desktopArticleTarget.noscript%7Cext.wikihiero%2CwikimediaBadges%7Cext.wikimediamessages.styles%7Cjquery.makeCollapsible.styles%7Cmediawiki.page.gallery.styles%7Cskins.vector.icons%2Cstyles%7Cskins.vector.search.codex.styles%7Cwikibase.client.init&only=styles&skin=vector-2022"> <script async="" src="/w/load.php?lang=en&modules=startup&only=scripts&raw=1&skin=vector-2022"></script> <meta name="ResourceLoaderDynamicStyles" content=""> <link rel="stylesheet" href="/w/load.php?lang=en&modules=site.styles&only=styles&skin=vector-2022"> <meta name="generator" content="MediaWiki 1.44.0-wmf.4"> <meta name="referrer" content="origin"> <meta name="referrer" content="origin-when-cross-origin"> <meta name="robots" content="max-image-preview:standard"> <meta name="format-detection" content="telephone=no"> <meta property="og:image" content="https://upload.wikimedia.org/wikipedia/commons/thumb/2/20/Coleoptera_collage.png/1200px-Coleoptera_collage.png"> <meta property="og:image:width" content="1200"> <meta property="og:image:height" content="1144"> <meta property="og:image" content="https://upload.wikimedia.org/wikipedia/commons/thumb/2/20/Coleoptera_collage.png/800px-Coleoptera_collage.png"> <meta property="og:image:width" content="800"> <meta property="og:image:height" content="763"> <meta property="og:image" content="https://upload.wikimedia.org/wikipedia/commons/thumb/2/20/Coleoptera_collage.png/640px-Coleoptera_collage.png"> <meta property="og:image:width" content="640"> <meta property="og:image:height" content="610"> <meta name="viewport" content="width=1120"> <meta property="og:title" content="Beetle - Wikipedia"> <meta property="og:type" content="website"> <link rel="preconnect" href="//upload.wikimedia.org"> <link rel="alternate" media="only screen and (max-width: 640px)" href="//en.m.wikipedia.org/wiki/Beetle"> <link rel="alternate" type="application/x-wiki" title="Edit this page" href="/w/index.php?title=Beetle&action=edit"> <link rel="apple-touch-icon" href="/static/apple-touch/wikipedia.png"> <link rel="icon" href="/static/favicon/wikipedia.ico"> <link rel="search" type="application/opensearchdescription+xml" href="/w/rest.php/v1/search" title="Wikipedia (en)"> <link rel="EditURI" type="application/rsd+xml" href="//en.wikipedia.org/w/api.php?action=rsd"> <link rel="canonical" href="https://en.wikipedia.org/wiki/Beetle"> <link rel="license" href="https://creativecommons.org/licenses/by-sa/4.0/deed.en"> <link rel="alternate" type="application/atom+xml" title="Wikipedia Atom feed" href="/w/index.php?title=Special:RecentChanges&feed=atom"> <link rel="dns-prefetch" href="//meta.wikimedia.org" /> <link rel="dns-prefetch" href="//login.wikimedia.org"> </head> <body class="skin--responsive skin-vector skin-vector-search-vue mediawiki ltr sitedir-ltr mw-hide-empty-elt ns-0 ns-subject mw-editable page-Beetle rootpage-Beetle skin-vector-2022 action-view"><a class="mw-jump-link" href="#bodyContent">Jump to content</a> <div class="vector-header-container"> <header class="vector-header mw-header"> <div class="vector-header-start"> <nav class="vector-main-menu-landmark" aria-label="Site"> <div id="vector-main-menu-dropdown" class="vector-dropdown vector-main-menu-dropdown vector-button-flush-left vector-button-flush-right" > <input type="checkbox" id="vector-main-menu-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-main-menu-dropdown" class="vector-dropdown-checkbox " aria-label="Main menu" > <label id="vector-main-menu-dropdown-label" for="vector-main-menu-dropdown-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" > Main menu </label> <div class="vector-dropdown-content"> <div id="vector-main-menu-unpinned-container" class="vector-unpinned-container"> <div id="vector-main-menu" class="vector-main-menu vector-pinnable-element"> <div class="vector-pinnable-header vector-main-menu-pinnable-header vector-pinnable-header-unpinned" data-feature-name="main-menu-pinned" data-pinnable-element-id="vector-main-menu" data-pinned-container-id="vector-main-menu-pinned-container" data-unpinned-container-id="vector-main-menu-unpinned-container" > <div class="vector-pinnable-header-label">Main menu</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-main-menu.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-main-menu.unpin">hide</button> </div> <div id="p-navigation" class="vector-menu mw-portlet mw-portlet-navigation" > <div class="vector-menu-heading"> Navigation </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="n-mainpage-description" class="mw-list-item"><a href="/wiki/Main_Page" title="Visit the main page [z]" accesskey="z">Main page</a></li><li id="n-contents" class="mw-list-item"><a href="/wiki/Wikipedia:Contents" title="Guides to browsing Wikipedia">Contents</a></li><li id="n-currentevents" class="mw-list-item"><a href="/wiki/Portal:Current_events" title="Articles related to current events">Current events</a></li><li id="n-randompage" class="mw-list-item"><a href="/wiki/Special:Random" title="Visit a randomly selected article [x]" accesskey="x">Random article</a></li><li id="n-aboutsite" class="mw-list-item"><a href="/wiki/Wikipedia:About" title="Learn about Wikipedia and how it works">About Wikipedia</a></li><li id="n-contactpage" class="mw-list-item"><a href="//en.wikipedia.org/wiki/Wikipedia:Contact_us" title="How to contact Wikipedia">Contact us</a></li> </ul> </div> </div> <div id="p-interaction" class="vector-menu mw-portlet mw-portlet-interaction" > <div class="vector-menu-heading"> Contribute </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="n-help" class="mw-list-item"><a href="/wiki/Help:Contents" title="Guidance on how to use and edit Wikipedia">Help</a></li><li id="n-introduction" class="mw-list-item"><a href="/wiki/Help:Introduction" title="Learn how to edit Wikipedia">Learn to edit</a></li><li id="n-portal" class="mw-list-item"><a href="/wiki/Wikipedia:Community_portal" title="The hub for editors">Community portal</a></li><li id="n-recentchanges" class="mw-list-item"><a href="/wiki/Special:RecentChanges" title="A list of recent changes to Wikipedia [r]" accesskey="r">Recent changes</a></li><li id="n-upload" class="mw-list-item"><a href="/wiki/Wikipedia:File_upload_wizard" title="Add images or other media for use on Wikipedia">Upload file</a></li> </ul> </div> </div> </div> </div> </div> </div> </nav> <a href="/wiki/Main_Page" class="mw-logo"> <img class="mw-logo-icon" src="/static/images/icons/wikipedia.png" alt="" aria-hidden="true" height="50" width="50"> <img class="mw-logo-wordmark" alt="Wikipedia" src="/static/images/mobile/copyright/wikipedia-wordmark-en.svg" style="width: 7.5em; height: 1.125em;"> <img class="mw-logo-tagline" alt="The Free Encyclopedia" src="/static/images/mobile/copyright/wikipedia-tagline-en.svg" width="117" height="13" style="width: 7.3125em; height: 0.8125em;"> </a> </div> <div class="vector-header-end"> <div id="p-search" role="search" class="vector-search-box-vue vector-search-box-collapses vector-search-box-show-thumbnail vector-search-box-auto-expand-width vector-search-box"> <a href="/wiki/Special:Search" class="cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only search-toggle" title="Search Wikipedia [f]" accesskey="f"> Search </a> <div class="vector-typeahead-search-container"> <div class="cdx-typeahead-search cdx-typeahead-search--show-thumbnail cdx-typeahead-search--auto-expand-width"> <form action="/w/index.php" id="searchform" class="cdx-search-input cdx-search-input--has-end-button"> <div id="simpleSearch" class="cdx-search-input__input-wrapper" data-search-loc="header-moved"> <div class="cdx-text-input cdx-text-input--has-start-icon"> <input class="cdx-text-input__input" type="search" name="search" placeholder="Search Wikipedia" aria-label="Search Wikipedia" autocapitalize="sentences" title="Search Wikipedia [f]" accesskey="f" id="searchInput" > </div> <input type="hidden" name="title" value="Special:Search"> </div> <button class="cdx-button cdx-search-input__end-button">Search</button> </form> </div> </div> </div> <nav class="vector-user-links vector-user-links-wide" aria-label="Personal tools"> <div class="vector-user-links-main"> <div id="p-vector-user-menu-preferences" class="vector-menu mw-portlet emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> <div id="p-vector-user-menu-userpage" class="vector-menu mw-portlet emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> <nav class="vector-appearance-landmark" aria-label="Appearance"> <div id="vector-appearance-dropdown" class="vector-dropdown " title="Change the appearance of the page's font size, width, and color" > <input type="checkbox" id="vector-appearance-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-appearance-dropdown" class="vector-dropdown-checkbox " aria-label="Appearance" > <label id="vector-appearance-dropdown-label" for="vector-appearance-dropdown-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" > Appearance </label> <div class="vector-dropdown-content"> <div id="vector-appearance-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <div id="p-vector-user-menu-notifications" class="vector-menu mw-portlet emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> <div id="p-vector-user-menu-overflow" class="vector-menu mw-portlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="pt-sitesupport-2" class="user-links-collapsible-item mw-list-item user-links-collapsible-item"><a data-mw="interface" href="https://donate.wikimedia.org/wiki/Special:FundraiserRedirector?utm_source=donate&utm_medium=sidebar&utm_campaign=C13_en.wikipedia.org&uselang=en" class="">Donate</a> </li> <li id="pt-createaccount-2" class="user-links-collapsible-item mw-list-item user-links-collapsible-item"><a data-mw="interface" href="/w/index.php?title=Special:CreateAccount&returnto=Beetle" title="You are encouraged to create an account and log in; however, it is not mandatory" class="">Create account</a> </li> <li id="pt-login-2" class="user-links-collapsible-item mw-list-item user-links-collapsible-item"><a data-mw="interface" href="/w/index.php?title=Special:UserLogin&returnto=Beetle" title="You're encouraged to log in; however, it's not mandatory. [o]" accesskey="o" class="">Log in</a> </li> </ul> </div> </div> </div> <div id="vector-user-links-dropdown" class="vector-dropdown vector-user-menu vector-button-flush-right vector-user-menu-logged-out" title="Log in and more options" > <input type="checkbox" id="vector-user-links-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-user-links-dropdown" class="vector-dropdown-checkbox " aria-label="Personal tools" > <label id="vector-user-links-dropdown-label" for="vector-user-links-dropdown-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" > Personal tools </label> <div class="vector-dropdown-content"> <div id="p-personal" class="vector-menu mw-portlet mw-portlet-personal user-links-collapsible-item" title="User menu" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="pt-sitesupport" class="user-links-collapsible-item mw-list-item"><a href="https://donate.wikimedia.org/wiki/Special:FundraiserRedirector?utm_source=donate&utm_medium=sidebar&utm_campaign=C13_en.wikipedia.org&uselang=en">Donate</a></li><li id="pt-createaccount" class="user-links-collapsible-item mw-list-item"><a href="/w/index.php?title=Special:CreateAccount&returnto=Beetle" title="You are encouraged to create an account and log in; however, it is not mandatory"> Create account</a></li><li id="pt-login" class="user-links-collapsible-item mw-list-item"><a href="/w/index.php?title=Special:UserLogin&returnto=Beetle" title="You're encouraged to log in; however, it's not mandatory. [o]" accesskey="o"> Log in</a></li> </ul> </div> </div> <div id="p-user-menu-anon-editor" class="vector-menu mw-portlet mw-portlet-user-menu-anon-editor" > <div class="vector-menu-heading"> Pages for logged out editors <a href="/wiki/Help:Introduction" aria-label="Learn more about editing">learn more</a> </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="pt-anoncontribs" class="mw-list-item"><a href="/wiki/Special:MyContributions" title="A list of edits made from this IP address [y]" accesskey="y">Contributions</a></li><li id="pt-anontalk" class="mw-list-item"><a href="/wiki/Special:MyTalk" title="Discussion about edits from this IP address [n]" accesskey="n">Talk</a></li> </ul> </div> </div> </div> </div> </nav> </div> </header> </div> <div class="mw-page-container"> <div class="mw-page-container-inner"> <div class="vector-sitenotice-container"> <div id="siteNotice"></div> </div> <div class="vector-column-start"> <div class="vector-main-menu-container"> <div id="mw-navigation"> <nav id="mw-panel" class="vector-main-menu-landmark" aria-label="Site"> <div id="vector-main-menu-pinned-container" class="vector-pinned-container"> </div> </nav> </div> </div> <div class="vector-sticky-pinned-container"> <nav id="mw-panel-toc" aria-label="Contents" data-event-name="ui.sidebar-toc" class="mw-table-of-contents-container vector-toc-landmark"> <div id="vector-toc-pinned-container" class="vector-pinned-container"> <div id="vector-toc" class="vector-toc vector-pinnable-element"> <div class="vector-pinnable-header vector-toc-pinnable-header vector-pinnable-header-pinned" data-feature-name="toc-pinned" data-pinnable-element-id="vector-toc" > <h2 class="vector-pinnable-header-label">Contents</h2> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-toc.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-toc.unpin">hide</button> </div> <ul class="vector-toc-contents" id="mw-panel-toc-list"> <li id="toc-mw-content-text" class="vector-toc-list-item vector-toc-level-1"> <a href="#" class="vector-toc-link"> <div class="vector-toc-text">(Top)</div> </a> </li> <li id="toc-Etymology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Etymology"> <div class="vector-toc-text"> 1 Etymology </div> </a> <ul id="toc-Etymology-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Distribution_and_diversity" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Distribution_and_diversity"> <div class="vector-toc-text"> 2 Distribution and diversity </div> </a> <ul id="toc-Distribution_and_diversity-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Evolution" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Evolution"> <div class="vector-toc-text"> 3 Evolution </div> </a> <button aria-controls="toc-Evolution-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Evolution subsection </button> <ul id="toc-Evolution-sublist" class="vector-toc-list"> <li id="toc-Late_Paleozoic_and_Triassic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Late_Paleozoic_and_Triassic"> <div class="vector-toc-text"> 3.1 Late Paleozoic and Triassic </div> </a> <ul id="toc-Late_Paleozoic_and_Triassic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Jurassic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Jurassic"> <div class="vector-toc-text"> 3.2 Jurassic </div> </a> <ul id="toc-Jurassic-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cretaceous" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cretaceous"> <div class="vector-toc-text"> 3.3 Cretaceous </div> </a> <ul id="toc-Cretaceous-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Cenozoic" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Cenozoic"> <div class="vector-toc-text"> 3.4 Cenozoic </div> </a> <ul id="toc-Cenozoic-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Phylogeny" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Phylogeny"> <div class="vector-toc-text"> 4 Phylogeny </div> </a> <ul id="toc-Phylogeny-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_morphology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_morphology"> <div class="vector-toc-text"> 5 External morphology </div> </a> <button aria-controls="toc-External_morphology-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle External morphology subsection </button> <ul id="toc-External_morphology-sublist" class="vector-toc-list"> <li id="toc-Head" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Head"> <div class="vector-toc-text"> 5.1 Head </div> </a> <ul id="toc-Head-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Thorax" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Thorax"> <div class="vector-toc-text"> 5.2 Thorax </div> </a> <ul id="toc-Thorax-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Legs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Legs"> <div class="vector-toc-text"> 5.3 Legs </div> </a> <ul id="toc-Legs-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Wings" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Wings"> <div class="vector-toc-text"> 5.4 Wings </div> </a> <ul id="toc-Wings-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Abdomen" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Abdomen"> <div class="vector-toc-text"> 5.5 Abdomen </div> </a> <ul id="toc-Abdomen-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Anatomy_and_physiology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Anatomy_and_physiology"> <div class="vector-toc-text"> 6 Anatomy and physiology </div> </a> <button aria-controls="toc-Anatomy_and_physiology-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Anatomy and physiology subsection </button> <ul id="toc-Anatomy_and_physiology-sublist" class="vector-toc-list"> <li id="toc-Digestive_system" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Digestive_system"> <div class="vector-toc-text"> 6.1 Digestive system </div> </a> <ul id="toc-Digestive_system-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Nervous_system" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Nervous_system"> <div class="vector-toc-text"> 6.2 Nervous system </div> </a> <ul id="toc-Nervous_system-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Respiratory_system" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Respiratory_system"> <div class="vector-toc-text"> 6.3 Respiratory system </div> </a> <ul id="toc-Respiratory_system-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Circulatory_system" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Circulatory_system"> <div class="vector-toc-text"> 6.4 Circulatory system </div> </a> <ul id="toc-Circulatory_system-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Specialized_organs" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Specialized_organs"> <div class="vector-toc-text"> 6.5 Specialized organs </div> </a> <ul id="toc-Specialized_organs-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Reproduction_and_development" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Reproduction_and_development"> <div class="vector-toc-text"> 7 Reproduction and development </div> </a> <button aria-controls="toc-Reproduction_and_development-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Reproduction and development subsection </button> <ul id="toc-Reproduction_and_development-sublist" class="vector-toc-list"> <li id="toc-Mating" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mating"> <div class="vector-toc-text"> 7.1 Mating </div> </a> <ul id="toc-Mating-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Life_cycle" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Life_cycle"> <div class="vector-toc-text"> 7.2 Life cycle </div> </a> <ul id="toc-Life_cycle-sublist" class="vector-toc-list"> <li id="toc-Egg" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Egg"> <div class="vector-toc-text"> 7.2.1 Egg </div> </a> <ul id="toc-Egg-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Larva" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Larva"> <div class="vector-toc-text"> 7.2.2 Larva </div> </a> <ul id="toc-Larva-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Pupa_and_adult" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Pupa_and_adult"> <div class="vector-toc-text"> 7.2.3 Pupa and adult </div> </a> <ul id="toc-Pupa_and_adult-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> </ul> </li> <li id="toc-Behaviour" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Behaviour"> <div class="vector-toc-text"> 8 Behaviour </div> </a> <button aria-controls="toc-Behaviour-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Behaviour subsection </button> <ul id="toc-Behaviour-sublist" class="vector-toc-list"> <li id="toc-Locomotion" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Locomotion"> <div class="vector-toc-text"> 8.1 Locomotion </div> </a> <ul id="toc-Locomotion-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Communication" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Communication"> <div class="vector-toc-text"> 8.2 Communication </div> </a> <ul id="toc-Communication-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Parental_care" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Parental_care"> <div class="vector-toc-text"> 8.3 Parental care </div> </a> <ul id="toc-Parental_care-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Eusociality" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Eusociality"> <div class="vector-toc-text"> 8.4 Eusociality </div> </a> <ul id="toc-Eusociality-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Feeding" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Feeding"> <div class="vector-toc-text"> 8.5 Feeding </div> </a> <ul id="toc-Feeding-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Ecology" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Ecology"> <div class="vector-toc-text"> 9 Ecology </div> </a> <button aria-controls="toc-Ecology-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Ecology subsection </button> <ul id="toc-Ecology-sublist" class="vector-toc-list"> <li id="toc-Anti-predator_adaptations" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Anti-predator_adaptations"> <div class="vector-toc-text"> 9.1 Anti-predator adaptations </div> </a> <ul id="toc-Anti-predator_adaptations-sublist" class="vector-toc-list"> <li id="toc-Camouflage" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Camouflage"> <div class="vector-toc-text"> 9.1.1 Camouflage </div> </a> <ul id="toc-Camouflage-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mimicry_and_aposematism" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Mimicry_and_aposematism"> <div class="vector-toc-text"> 9.1.2 Mimicry and aposematism </div> </a> <ul id="toc-Mimicry_and_aposematism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Other_defenses" class="vector-toc-list-item vector-toc-level-3"> <a class="vector-toc-link" href="#Other_defenses"> <div class="vector-toc-text"> 9.1.3 Other defenses </div> </a> <ul id="toc-Other_defenses-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Parasitism" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Parasitism"> <div class="vector-toc-text"> 9.2 Parasitism </div> </a> <ul id="toc-Parasitism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Pollination" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Pollination"> <div class="vector-toc-text"> 9.3 Pollination </div> </a> <ul id="toc-Pollination-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Mutualism" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Mutualism"> <div class="vector-toc-text"> 9.4 Mutualism </div> </a> <ul id="toc-Mutualism-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Tolerance_of_extreme_environments" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Tolerance_of_extreme_environments"> <div class="vector-toc-text"> 9.5 Tolerance of extreme environments </div> </a> <ul id="toc-Tolerance_of_extreme_environments-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-Migration" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Migration"> <div class="vector-toc-text"> 9.6 Migration </div> </a> <ul id="toc-Migration-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Relationship_to_humans" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Relationship_to_humans"> <div class="vector-toc-text"> 10 Relationship to humans </div> </a> <button aria-controls="toc-Relationship_to_humans-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle Relationship to humans subsection </button> <ul id="toc-Relationship_to_humans-sublist" class="vector-toc-list"> <li id="toc-In_ancient_cultures" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_ancient_cultures"> <div class="vector-toc-text"> 10.1 In ancient cultures </div> </a> <ul id="toc-In_ancient_cultures-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_pests" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#As_pests"> <div class="vector-toc-text"> 10.2 As pests </div> </a> <ul id="toc-As_pests-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_beneficial_resources" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#As_beneficial_resources"> <div class="vector-toc-text"> 10.3 As beneficial resources </div> </a> <ul id="toc-As_beneficial_resources-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_food_and_medicine" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#As_food_and_medicine"> <div class="vector-toc-text"> 10.4 As food and medicine </div> </a> <ul id="toc-As_food_and_medicine-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_biodiversity_indicators" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#As_biodiversity_indicators"> <div class="vector-toc-text"> 10.5 As biodiversity indicators </div> </a> <ul id="toc-As_biodiversity_indicators-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_art_and_adornment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_art_and_adornment"> <div class="vector-toc-text"> 10.6 In art and adornment </div> </a> <ul id="toc-In_art_and_adornment-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_entertainment" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_entertainment"> <div class="vector-toc-text"> 10.7 In entertainment </div> </a> <ul id="toc-In_entertainment-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_pets" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#As_pets"> <div class="vector-toc-text"> 10.8 As pets </div> </a> <ul id="toc-As_pets-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_things_to_collect" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#As_things_to_collect"> <div class="vector-toc-text"> 10.9 As things to collect </div> </a> <ul id="toc-As_things_to_collect-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-As_inspiration_for_technologies" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#As_inspiration_for_technologies"> <div class="vector-toc-text"> 10.10 As inspiration for technologies </div> </a> <ul id="toc-As_inspiration_for_technologies-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-In_conservation" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#In_conservation"> <div class="vector-toc-text"> 10.11 In conservation </div> </a> <ul id="toc-In_conservation-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Notes" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Notes"> <div class="vector-toc-text"> 11 Notes </div> </a> <ul id="toc-Notes-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-See_also" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#See_also"> <div class="vector-toc-text"> 12 See also </div> </a> <ul id="toc-See_also-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-References" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#References"> <div class="vector-toc-text"> 13 References </div> </a> <button aria-controls="toc-References-sublist" class="cdx-button cdx-button--weight-quiet cdx-button--icon-only vector-toc-toggle"> Toggle References subsection </button> <ul id="toc-References-sublist" class="vector-toc-list"> <li id="toc-Bibliography" class="vector-toc-list-item vector-toc-level-2"> <a class="vector-toc-link" href="#Bibliography"> <div class="vector-toc-text"> 13.1 Bibliography </div> </a> <ul id="toc-Bibliography-sublist" class="vector-toc-list"> </ul> </li> </ul> </li> <li id="toc-Further_reading" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#Further_reading"> <div class="vector-toc-text"> 14 Further reading </div> </a> <ul id="toc-Further_reading-sublist" class="vector-toc-list"> </ul> </li> <li id="toc-External_links" class="vector-toc-list-item vector-toc-level-1"> <a class="vector-toc-link" href="#External_links"> <div class="vector-toc-text"> 15 External links </div> </a> <ul id="toc-External_links-sublist" class="vector-toc-list"> </ul> </li> </ul> </div> </div> </nav> </div> </div> <div class="mw-content-container"> <main id="content" class="mw-body"> <header class="mw-body-header vector-page-titlebar"> <nav aria-label="Contents" class="vector-toc-landmark"> <div id="vector-page-titlebar-toc" class="vector-dropdown vector-page-titlebar-toc vector-button-flush-left" > <input type="checkbox" id="vector-page-titlebar-toc-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-titlebar-toc" class="vector-dropdown-checkbox " aria-label="Toggle the table of contents" > <label id="vector-page-titlebar-toc-label" for="vector-page-titlebar-toc-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--icon-only " aria-hidden="true" > Toggle the table of contents </label> <div class="vector-dropdown-content"> <div id="vector-page-titlebar-toc-unpinned-container" class="vector-unpinned-container"> </div> </div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading">Beetle</h1> <div id="p-lang-btn" class="vector-dropdown mw-portlet mw-portlet-lang" > <input type="checkbox" id="p-lang-btn-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-p-lang-btn" class="vector-dropdown-checkbox mw-interlanguage-selector" aria-label="Go to an article in another language. Available in 117 languages" > <label id="p-lang-btn-label" for="p-lang-btn-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet cdx-button--action-progressive mw-portlet-lang-heading-117" aria-hidden="true" > 117 languages </label> <div class="vector-dropdown-content"> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="interlanguage-link interwiki-af mw-list-item"><a href="https://af.wikipedia.org/wiki/Kewer" title="Kewer – Afrikaans" lang="af" hreflang="af" data-title="Kewer" data-language-autonym="Afrikaans" data-language-local-name="Afrikaans" class="interlanguage-link-target">Afrikaans</a></li><li class="interlanguage-link interwiki-am mw-list-item"><a href="https://am.wikipedia.org/wiki/%E1%89%A0%E1%88%AB%E1%89%A5%E1%88%AA%E1%89%B5" title="በራብሪት – Amharic" lang="am" hreflang="am" data-title="በራብሪት" data-language-autonym="አማርኛ" data-language-local-name="Amharic" class="interlanguage-link-target">አማርኛ</a></li><li class="interlanguage-link interwiki-ang mw-list-item"><a href="https://ang.wikipedia.org/wiki/Ceafer" title="Ceafer – Old English" lang="ang" hreflang="ang" data-title="Ceafer" data-language-autonym="Ænglisc" data-language-local-name="Old English" class="interlanguage-link-target">Ænglisc</a></li><li class="interlanguage-link interwiki-ar mw-list-item"><a href="https://ar.wikipedia.org/wiki/%D8%AE%D9%86%D8%A7%D9%81%D8%B3" title="خنافس – Arabic" lang="ar" hreflang="ar" data-title="خنافس" data-language-autonym="العربية" data-language-local-name="Arabic" class="interlanguage-link-target">العربية</a></li><li class="interlanguage-link interwiki-an mw-list-item"><a href="https://an.wikipedia.org/wiki/Coleoptera" title="Coleoptera – Aragonese" lang="an" hreflang="an" data-title="Coleoptera" data-language-autonym="Aragonés" data-language-local-name="Aragonese" class="interlanguage-link-target">Aragonés</a></li><li class="interlanguage-link interwiki-ast mw-list-item"><a href="https://ast.wikipedia.org/wiki/Coleoptera" title="Coleoptera – Asturian" lang="ast" hreflang="ast" data-title="Coleoptera" data-language-autonym="Asturianu" data-language-local-name="Asturian" class="interlanguage-link-target">Asturianu</a></li><li class="interlanguage-link interwiki-atj mw-list-item"><a href="https://atj.wikipedia.org/wiki/Emikwanisis" title="Emikwanisis – Atikamekw" lang="atj" hreflang="atj" data-title="Emikwanisis" data-language-autonym="Atikamekw" data-language-local-name="Atikamekw" class="interlanguage-link-target">Atikamekw</a></li><li class="interlanguage-link interwiki-gn mw-list-item"><a href="https://gn.wikipedia.org/wiki/Lembu" title="Lembu – Guarani" lang="gn" hreflang="gn" data-title="Lembu" data-language-autonym="Avañe'ẽ" data-language-local-name="Guarani" class="interlanguage-link-target">Avañe'ẽ</a></li><li class="interlanguage-link interwiki-az mw-list-item"><a href="https://az.wikipedia.org/wiki/S%C9%99rtqanadl%C4%B1lar" title="Sərtqanadlılar – Azerbaijani" lang="az" hreflang="az" data-title="Sərtqanadlılar" data-language-autonym="Azərbaycanca" data-language-local-name="Azerbaijani" class="interlanguage-link-target">Azərbaycanca</a></li><li class="interlanguage-link interwiki-bn mw-list-item"><a href="https://bn.wikipedia.org/wiki/%E0%A6%97%E0%A7%81%E0%A6%AC%E0%A6%B0%E0%A7%87_%E0%A6%AA%E0%A7%8B%E0%A6%95%E0%A6%BE" title="গুবরে পোকা – Bangla" lang="bn" hreflang="bn" data-title="গুবরে পোকা" data-language-autonym="বাংলা" data-language-local-name="Bangla" class="interlanguage-link-target">বাংলা</a></li><li class="interlanguage-link interwiki-zh-min-nan mw-list-item"><a href="https://zh-min-nan.wikipedia.org/wiki/Ku-%C3%A1_(th%C3%A2ng)" title="Ku-á (thâng) – Minnan" lang="nan" hreflang="nan" data-title="Ku-á (thâng)" data-language-autonym="閩南語 / Bân-lâm-gú" data-language-local-name="Minnan" class="interlanguage-link-target">閩南語 / Bân-lâm-gú</a></li><li class="interlanguage-link interwiki-ba mw-list-item"><a href="https://ba.wikipedia.org/wiki/%D2%A0%D0%B0%D1%82%D1%8B_%D2%A1%D0%B0%D0%BD%D0%B0%D1%82%D0%BB%D1%8B%D0%BB%D0%B0%D1%80" title="Ҡаты ҡанатлылар – Bashkir" lang="ba" hreflang="ba" data-title="Ҡаты ҡанатлылар" data-language-autonym="Башҡортса" data-language-local-name="Bashkir" class="interlanguage-link-target">Башҡортса</a></li><li class="interlanguage-link interwiki-be mw-list-item"><a href="https://be.wikipedia.org/wiki/%D0%96%D1%83%D0%BA%D1%96" title="Жукі – Belarusian" lang="be" hreflang="be" data-title="Жукі" data-language-autonym="Беларуская" data-language-local-name="Belarusian" class="interlanguage-link-target">Беларуская</a></li><li class="interlanguage-link interwiki-be-x-old mw-list-item"><a href="https://be-tarask.wikipedia.org/wiki/%D0%A6%D1%8C%D0%B2%D0%B5%D1%80%D0%B4%D0%B0%D0%BA%D1%80%D1%8B%D0%BB%D1%8B%D1%8F" title="Цьвердакрылыя – Belarusian (Taraškievica orthography)" lang="be-tarask" hreflang="be-tarask" data-title="Цьвердакрылыя" data-language-autonym="Беларуская (тарашкевіца)" data-language-local-name="Belarusian (Taraškievica orthography)" class="interlanguage-link-target">Беларуская (тарашкевіца)</a></li><li class="interlanguage-link interwiki-bcl mw-list-item"><a href="https://bcl.wikipedia.org/wiki/Amamanggi" title="Amamanggi – Central Bikol" lang="bcl" hreflang="bcl" data-title="Amamanggi" data-language-autonym="Bikol Central" data-language-local-name="Central Bikol" class="interlanguage-link-target">Bikol Central</a></li><li class="interlanguage-link interwiki-bg mw-list-item"><a href="https://bg.wikipedia.org/wiki/%D0%91%D1%80%D1%8A%D0%BC%D0%B1%D0%B0%D1%80%D0%B8" title="Бръмбари – Bulgarian" lang="bg" hreflang="bg" data-title="Бръмбари" data-language-autonym="Български" data-language-local-name="Bulgarian" class="interlanguage-link-target">Български</a></li><li class="interlanguage-link interwiki-bs mw-list-item"><a href="https://bs.wikipedia.org/wiki/Tvrdokrilci" title="Tvrdokrilci – Bosnian" lang="bs" hreflang="bs" data-title="Tvrdokrilci" data-language-autonym="Bosanski" data-language-local-name="Bosnian" class="interlanguage-link-target">Bosanski</a></li><li class="interlanguage-link interwiki-br mw-list-item"><a href="https://br.wikipedia.org/wiki/C%27hwil" title="C'hwil – Breton" lang="br" hreflang="br" data-title="C'hwil" data-language-autonym="Brezhoneg" data-language-local-name="Breton" class="interlanguage-link-target">Brezhoneg</a></li><li class="interlanguage-link interwiki-ca mw-list-item"><a href="https://ca.wikipedia.org/wiki/Cole%C3%B2pters" title="Coleòpters – Catalan" lang="ca" hreflang="ca" data-title="Coleòpters" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target">Català</a></li><li class="interlanguage-link interwiki-cv mw-list-item"><a href="https://cv.wikipedia.org/wiki/%D0%9F%D0%B8%D1%80%D1%87%D0%B5%D0%B2_%C3%A7%D1%83%D0%BD%D0%B0%D1%82%D0%BB%D0%B8%D1%81%D0%B5%D0%BC" title="Пирчев çунатлисем – Chuvash" lang="cv" hreflang="cv" data-title="Пирчев çунатлисем" data-language-autonym="Чӑвашла" data-language-local-name="Chuvash" class="interlanguage-link-target">Чӑвашла</a></li><li class="interlanguage-link interwiki-ceb mw-list-item"><a href="https://ceb.wikipedia.org/wiki/Bakukangon" title="Bakukangon – Cebuano" lang="ceb" hreflang="ceb" data-title="Bakukangon" data-language-autonym="Cebuano" data-language-local-name="Cebuano" class="interlanguage-link-target">Cebuano</a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Brouci" title="Brouci – Czech" lang="cs" hreflang="cs" data-title="Brouci" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target">Čeština</a></li><li class="interlanguage-link interwiki-cy mw-list-item"><a href="https://cy.wikipedia.org/wiki/Chwilen" title="Chwilen – Welsh" lang="cy" hreflang="cy" data-title="Chwilen" data-language-autonym="Cymraeg" data-language-local-name="Welsh" class="interlanguage-link-target">Cymraeg</a></li><li class="interlanguage-link interwiki-da mw-list-item"><a href="https://da.wikipedia.org/wiki/Biller" title="Biller – Danish" lang="da" hreflang="da" data-title="Biller" data-language-autonym="Dansk" data-language-local-name="Danish" class="interlanguage-link-target">Dansk</a></li><li class="interlanguage-link interwiki-de badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://de.wikipedia.org/wiki/K%C3%A4fer" title="Käfer – German" lang="de" hreflang="de" data-title="Käfer" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target">Deutsch</a></li><li class="interlanguage-link interwiki-nv mw-list-item"><a href="https://nv.wikipedia.org/wiki/Ch%C3%A9%C5%82chaa%CA%BC" title="Chéłchaaʼ – Navajo" lang="nv" hreflang="nv" data-title="Chéłchaaʼ" data-language-autonym="Diné bizaad" data-language-local-name="Navajo" class="interlanguage-link-target">Diné bizaad</a></li><li class="interlanguage-link interwiki-et mw-list-item"><a href="https://et.wikipedia.org/wiki/Mardikalised" title="Mardikalised – Estonian" lang="et" hreflang="et" data-title="Mardikalised" data-language-autonym="Eesti" data-language-local-name="Estonian" class="interlanguage-link-target">Eesti</a></li><li class="interlanguage-link interwiki-el mw-list-item"><a href="https://el.wikipedia.org/wiki/%CE%9A%CE%BF%CE%BB%CE%B5%CF%8C%CF%80%CF%84%CE%B5%CF%81%CE%B1" title="Κολεόπτερα – Greek" lang="el" hreflang="el" data-title="Κολεόπτερα" data-language-autonym="Ελληνικά" data-language-local-name="Greek" class="interlanguage-link-target">Ελληνικά</a></li><li class="interlanguage-link interwiki-es badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://es.wikipedia.org/wiki/Coleoptera" title="Coleoptera – Spanish" lang="es" hreflang="es" data-title="Coleoptera" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target">Español</a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Koleopteroj" title="Koleopteroj – Esperanto" lang="eo" hreflang="eo" data-title="Koleopteroj" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target">Esperanto</a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Kakalardo" title="Kakalardo – Basque" lang="eu" hreflang="eu" data-title="Kakalardo" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target">Euskara</a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D9%82%D8%A7%D8%A8%E2%80%8C%D8%A8%D8%A7%D9%84%D8%A7%D9%86" title="قاب‌بالان – Persian" lang="fa" hreflang="fa" data-title="قاب‌بالان" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target">فارسی</a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Coleoptera" title="Coleoptera – French" lang="fr" hreflang="fr" data-title="Coleoptera" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target">Français</a></li><li class="interlanguage-link interwiki-ga mw-list-item"><a href="https://ga.wikipedia.org/wiki/Coleoptera" title="Coleoptera – Irish" lang="ga" hreflang="ga" data-title="Coleoptera" data-language-autonym="Gaeilge" data-language-local-name="Irish" class="interlanguage-link-target">Gaeilge</a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Cole%C3%B3pteros" title="Coleópteros – Galician" lang="gl" hreflang="gl" data-title="Coleópteros" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target">Galego</a></li><li class="interlanguage-link interwiki-inh mw-list-item"><a href="https://inh.wikipedia.org/wiki/%D0%A7%D0%BE%D0%B0%D0%BF%D0%B8%D0%BB%D0%B3%D0%B0%D1%88" title="Чоапилгаш – Ingush" lang="inh" hreflang="inh" data-title="Чоапилгаш" data-language-autonym="ГӀалгӀай" data-language-local-name="Ingush" class="interlanguage-link-target">ГӀалгӀай</a></li><li class="interlanguage-link interwiki-hak mw-list-item"><a href="https://hak.wikipedia.org/wiki/K%C3%BB-%C3%A9" title="Kû-é – Hakka Chinese" lang="hak" hreflang="hak" data-title="Kû-é" data-language-autonym="客家語 / Hak-kâ-ngî" data-language-local-name="Hakka Chinese" class="interlanguage-link-target">客家語 / Hak-kâ-ngî</a></li><li class="interlanguage-link interwiki-ko mw-list-item"><a href="https://ko.wikipedia.org/wiki/%EB%94%B1%EC%A0%95%EB%B2%8C%EB%A0%88%EB%AA%A9" title="딱정벌레목 – Korean" lang="ko" hreflang="ko" data-title="딱정벌레목" data-language-autonym="한국어" data-language-local-name="Korean" class="interlanguage-link-target">한국어</a></li><li class="interlanguage-link interwiki-ha mw-list-item"><a href="https://ha.wikipedia.org/wiki/Buzuzu" title="Buzuzu – Hausa" lang="ha" hreflang="ha" data-title="Buzuzu" data-language-autonym="Hausa" data-language-local-name="Hausa" class="interlanguage-link-target">Hausa</a></li><li class="interlanguage-link interwiki-hy mw-list-item"><a href="https://hy.wikipedia.org/wiki/%D4%B2%D5%A6%D5%A5%D5%A6%D5%B6%D5%A5%D6%80" title="Բզեզներ – Armenian" lang="hy" hreflang="hy" data-title="Բզեզներ" data-language-autonym="Հայերեն" data-language-local-name="Armenian" class="interlanguage-link-target">Հայերեն</a></li><li class="interlanguage-link interwiki-hi mw-list-item"><a href="https://hi.wikipedia.org/wiki/%E0%A4%B5%E0%A4%B0%E0%A5%8D%E0%A4%AE%E0%A4%AA%E0%A4%82%E0%A4%96%E0%A5%80" title="वर्मपंखी – Hindi" lang="hi" hreflang="hi" data-title="वर्मपंखी" data-language-autonym="हिन्दी" data-language-local-name="Hindi" class="interlanguage-link-target">हिन्दी</a></li><li class="interlanguage-link interwiki-hsb mw-list-item"><a href="https://hsb.wikipedia.org/wiki/Bruki" title="Bruki – Upper Sorbian" lang="hsb" hreflang="hsb" data-title="Bruki" data-language-autonym="Hornjoserbsce" data-language-local-name="Upper Sorbian" class="interlanguage-link-target">Hornjoserbsce</a></li><li class="interlanguage-link interwiki-hr mw-list-item"><a href="https://hr.wikipedia.org/wiki/Kornja%C5%A1i" title="Kornjaši – Croatian" lang="hr" hreflang="hr" data-title="Kornjaši" data-language-autonym="Hrvatski" data-language-local-name="Croatian" class="interlanguage-link-target">Hrvatski</a></li><li class="interlanguage-link interwiki-io mw-list-item"><a href="https://io.wikipedia.org/wiki/Koleoptero" title="Koleoptero – Ido" lang="io" hreflang="io" data-title="Koleoptero" data-language-autonym="Ido" data-language-local-name="Ido" class="interlanguage-link-target">Ido</a></li><li class="interlanguage-link interwiki-id mw-list-item"><a href="https://id.wikipedia.org/wiki/Kumbang" title="Kumbang – Indonesian" lang="id" hreflang="id" data-title="Kumbang" data-language-autonym="Bahasa Indonesia" data-language-local-name="Indonesian" class="interlanguage-link-target">Bahasa Indonesia</a></li><li class="interlanguage-link interwiki-ia mw-list-item"><a href="https://ia.wikipedia.org/wiki/Coleoptero" title="Coleoptero – Interlingua" lang="ia" hreflang="ia" data-title="Coleoptero" data-language-autonym="Interlingua" data-language-local-name="Interlingua" class="interlanguage-link-target">Interlingua</a></li><li class="interlanguage-link interwiki-os mw-list-item"><a href="https://os.wikipedia.org/wiki/%D0%A5%D1%8A%C3%A6%D0%BD%D0%B4%D0%B8%D0%BB%D1%82%C3%A6" title="Хъæндилтæ – Ossetic" lang="os" hreflang="os" data-title="Хъæндилтæ" data-language-autonym="Ирон" data-language-local-name="Ossetic" class="interlanguage-link-target">Ирон</a></li><li class="interlanguage-link interwiki-is mw-list-item"><a href="https://is.wikipedia.org/wiki/Bj%C3%B6llur" title="Bjöllur – Icelandic" lang="is" hreflang="is" data-title="Bjöllur" data-language-autonym="Íslenska" data-language-local-name="Icelandic" class="interlanguage-link-target">Íslenska</a></li><li class="interlanguage-link interwiki-it mw-list-item"><a href="https://it.wikipedia.org/wiki/Coleoptera" title="Coleoptera – Italian" lang="it" hreflang="it" data-title="Coleoptera" data-language-autonym="Italiano" data-language-local-name="Italian" class="interlanguage-link-target">Italiano</a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%97%D7%99%D7%A4%D7%95%D7%A9%D7%99%D7%95%D7%AA" title="חיפושיות – Hebrew" lang="he" hreflang="he" data-title="חיפושיות" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target">עברית</a></li><li class="interlanguage-link interwiki-jv mw-list-item"><a href="https://jv.wikipedia.org/wiki/Kumbang" title="Kumbang – Javanese" lang="jv" hreflang="jv" data-title="Kumbang" data-language-autonym="Jawa" data-language-local-name="Javanese" class="interlanguage-link-target">Jawa</a></li><li class="interlanguage-link interwiki-kn mw-list-item"><a href="https://kn.wikipedia.org/wiki/%E0%B2%A6%E0%B3%81%E0%B2%82%E0%B2%AC%E0%B2%BF" title="ದುಂಬಿ – Kannada" lang="kn" hreflang="kn" data-title="ದುಂಬಿ" data-language-autonym="ಕನ್ನಡ" data-language-local-name="Kannada" class="interlanguage-link-target">ಕನ್ನಡ</a></li><li class="interlanguage-link interwiki-pam mw-list-item"><a href="https://pam.wikipedia.org/wiki/Coleoptera" title="Coleoptera – Pampanga" lang="pam" hreflang="pam" data-title="Coleoptera" data-language-autonym="Kapampangan" data-language-local-name="Pampanga" class="interlanguage-link-target">Kapampangan</a></li><li class="interlanguage-link interwiki-ka mw-list-item"><a href="https://ka.wikipedia.org/wiki/%E1%83%AE%E1%83%9D%E1%83%AD%E1%83%9D%E1%83%94%E1%83%91%E1%83%98" title="ხოჭოები – Georgian" lang="ka" hreflang="ka" data-title="ხოჭოები" data-language-autonym="ქართული" data-language-local-name="Georgian" class="interlanguage-link-target">ქართული</a></li><li class="interlanguage-link interwiki-kk mw-list-item"><a href="https://kk.wikipedia.org/wiki/%D2%9A%D0%B0%D1%82%D1%82%D1%8B_%D2%9B%D0%B0%D0%BD%D0%B0%D1%82%D1%82%D1%8B%D0%BB%D0%B0%D1%80" title="Қатты қанаттылар – Kazakh" lang="kk" hreflang="kk" data-title="Қатты қанаттылар" data-language-autonym="Қазақша" data-language-local-name="Kazakh" class="interlanguage-link-target">Қазақша</a></li><li class="interlanguage-link interwiki-kw mw-list-item"><a href="https://kw.wikipedia.org/wiki/Hwilen" title="Hwilen – Cornish" lang="kw" hreflang="kw" data-title="Hwilen" data-language-autonym="Kernowek" data-language-local-name="Cornish" class="interlanguage-link-target">Kernowek</a></li><li class="interlanguage-link interwiki-sw mw-list-item"><a href="https://sw.wikipedia.org/wiki/Mbawakawa" title="Mbawakawa – Swahili" lang="sw" hreflang="sw" data-title="Mbawakawa" data-language-autonym="Kiswahili" data-language-local-name="Swahili" class="interlanguage-link-target">Kiswahili</a></li><li class="interlanguage-link interwiki-ku mw-list-item"><a href="https://ku.wikipedia.org/wiki/K%C3%AAzik" title="Kêzik – Kurdish" lang="ku" hreflang="ku" data-title="Kêzik" data-language-autonym="Kurdî" data-language-local-name="Kurdish" class="interlanguage-link-target">Kurdî</a></li><li class="interlanguage-link interwiki-ky mw-list-item"><a href="https://ky.wikipedia.org/wiki/%D0%9A%D0%BE%D2%A3%D1%83%D0%B7%D0%B4%D0%B0%D1%80" title="Коңуздар – Kyrgyz" lang="ky" hreflang="ky" data-title="Коңуздар" data-language-autonym="Кыргызча" data-language-local-name="Kyrgyz" class="interlanguage-link-target">Кыргызча</a></li><li class="interlanguage-link interwiki-lld mw-list-item"><a href="https://lld.wikipedia.org/wiki/Ch%C3%ABifer" title="Chëifer – Ladin" lang="lld" hreflang="lld" data-title="Chëifer" data-language-autonym="Ladin" data-language-local-name="Ladin" class="interlanguage-link-target">Ladin</a></li><li class="interlanguage-link interwiki-la mw-list-item"><a href="https://la.wikipedia.org/wiki/Coleoptera" title="Coleoptera – Latin" lang="la" hreflang="la" data-title="Coleoptera" data-language-autonym="Latina" data-language-local-name="Latin" class="interlanguage-link-target">Latina</a></li><li class="interlanguage-link interwiki-lv badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://lv.wikipedia.org/wiki/Vaboles" title="Vaboles – Latvian" lang="lv" hreflang="lv" data-title="Vaboles" data-language-autonym="Latviešu" data-language-local-name="Latvian" class="interlanguage-link-target">Latviešu</a></li><li class="interlanguage-link interwiki-lb mw-list-item"><a href="https://lb.wikipedia.org/wiki/Kiewerleken" title="Kiewerleken – Luxembourgish" lang="lb" hreflang="lb" data-title="Kiewerleken" data-language-autonym="Lëtzebuergesch" data-language-local-name="Luxembourgish" class="interlanguage-link-target">Lëtzebuergesch</a></li><li class="interlanguage-link interwiki-lt badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://lt.wikipedia.org/wiki/Vabalai" title="Vabalai – Lithuanian" lang="lt" hreflang="lt" data-title="Vabalai" data-language-autonym="Lietuvių" data-language-local-name="Lithuanian" class="interlanguage-link-target">Lietuvių</a></li><li class="interlanguage-link interwiki-lfn mw-list-item"><a href="https://lfn.wikipedia.org/wiki/Scarabe" title="Scarabe – Lingua Franca Nova" lang="lfn" hreflang="lfn" data-title="Scarabe" data-language-autonym="Lingua Franca Nova" data-language-local-name="Lingua Franca Nova" class="interlanguage-link-target">Lingua Franca Nova</a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Bogarak" title="Bogarak – Hungarian" lang="hu" hreflang="hu" data-title="Bogarak" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target">Magyar</a></li><li class="interlanguage-link interwiki-mk mw-list-item"><a href="https://mk.wikipedia.org/wiki/%D0%A2%D0%B2%D1%80%D0%B4%D0%BE%D0%BA%D1%80%D0%B8%D0%BB%D1%86%D0%B8" title="Тврдокрилци – Macedonian" lang="mk" hreflang="mk" data-title="Тврдокрилци" data-language-autonym="Македонски" data-language-local-name="Macedonian" class="interlanguage-link-target">Македонски</a></li><li class="interlanguage-link interwiki-mg mw-list-item"><a href="https://mg.wikipedia.org/wiki/Borera" title="Borera – Malagasy" lang="mg" hreflang="mg" data-title="Borera" data-language-autonym="Malagasy" data-language-local-name="Malagasy" class="interlanguage-link-target">Malagasy</a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%B5%E0%B4%A3%E0%B5%8D%E0%B4%9F%E0%B5%8D" title="വണ്ട് – Malayalam" lang="ml" hreflang="ml" data-title="വണ്ട്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target">മലയാളം</a></li><li class="interlanguage-link interwiki-arz mw-list-item"><a href="https://arz.wikipedia.org/wiki/%D8%AE%D9%86%D8%A7%D9%81%D8%B3" title="خنافس – Egyptian Arabic" lang="arz" hreflang="arz" data-title="خنافس" data-language-autonym="مصرى" data-language-local-name="Egyptian Arabic" class="interlanguage-link-target">مصرى</a></li><li class="interlanguage-link interwiki-ms mw-list-item"><a href="https://ms.wikipedia.org/wiki/Kumbang" title="Kumbang – Malay" lang="ms" hreflang="ms" data-title="Kumbang" data-language-autonym="Bahasa Melayu" data-language-local-name="Malay" class="interlanguage-link-target">Bahasa Melayu</a></li><li class="interlanguage-link interwiki-my mw-list-item"><a href="https://my.wikipedia.org/wiki/%E1%80%81%E1%80%BB%E1%80%B1%E1%80%96%E1%80%AC%E1%80%95%E1%80%AD%E1%80%AF%E1%80%B8" title="ချေဖာပိုး – Burmese" lang="my" hreflang="my" data-title="ချေဖာပိုး" data-language-autonym="မြန်မာဘာသာ" data-language-local-name="Burmese" class="interlanguage-link-target">မြန်မာဘာသာ</a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Kevers" title="Kevers – Dutch" lang="nl" hreflang="nl" data-title="Kevers" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target">Nederlands</a></li><li class="interlanguage-link interwiki-ja mw-list-item"><a href="https://ja.wikipedia.org/wiki/%E7%94%B2%E8%99%AB%E9%A1%9E" title="甲虫類 – Japanese" lang="ja" hreflang="ja" data-title="甲虫類" data-language-autonym="日本語" data-language-local-name="Japanese" class="interlanguage-link-target">日本語</a></li><li class="interlanguage-link interwiki-frr mw-list-item"><a href="https://frr.wikipedia.org/wiki/Kraaben_(Insekten)" title="Kraaben (Insekten) – Northern Frisian" lang="frr" hreflang="frr" data-title="Kraaben (Insekten)" data-language-autonym="Nordfriisk" data-language-local-name="Northern Frisian" class="interlanguage-link-target">Nordfriisk</a></li><li class="interlanguage-link interwiki-no mw-list-item"><a href="https://no.wikipedia.org/wiki/Biller" title="Biller – Norwegian Bokmål" lang="nb" hreflang="nb" data-title="Biller" data-language-autonym="Norsk bokmål" data-language-local-name="Norwegian Bokmål" class="interlanguage-link-target">Norsk bokmål</a></li><li class="interlanguage-link interwiki-nn mw-list-item"><a href="https://nn.wikipedia.org/wiki/Biller" title="Biller – Norwegian Nynorsk" lang="nn" hreflang="nn" data-title="Biller" data-language-autonym="Norsk nynorsk" data-language-local-name="Norwegian Nynorsk" class="interlanguage-link-target">Norsk nynorsk</a></li><li class="interlanguage-link interwiki-oc mw-list-item"><a href="https://oc.wikipedia.org/wiki/Coleopt%C3%A8r" title="Coleoptèr – Occitan" lang="oc" hreflang="oc" data-title="Coleoptèr" data-language-autonym="Occitan" data-language-local-name="Occitan" class="interlanguage-link-target">Occitan</a></li><li class="interlanguage-link interwiki-uz mw-list-item"><a href="https://uz.wikipedia.org/wiki/Qo%CA%BBng%CA%BBizlar" title="Qoʻngʻizlar – Uzbek" lang="uz" hreflang="uz" data-title="Qoʻngʻizlar" data-language-autonym="Oʻzbekcha / ўзбекча" data-language-local-name="Uzbek" class="interlanguage-link-target">Oʻzbekcha / ўзбекча</a></li><li class="interlanguage-link interwiki-pa mw-list-item"><a href="https://pa.wikipedia.org/wiki/%E0%A8%AD%E0%A9%82%E0%A9%B0%E0%A8%A1" title="ਭੂੰਡ – Punjabi" lang="pa" hreflang="pa" data-title="ਭੂੰਡ" data-language-autonym="ਪੰਜਾਬੀ" data-language-local-name="Punjabi" class="interlanguage-link-target">ਪੰਜਾਬੀ</a></li><li class="interlanguage-link interwiki-pfl mw-list-item"><a href="https://pfl.wikipedia.org/wiki/K%C3%A4fer" title="Käfer – Palatine German" lang="pfl" hreflang="pfl" data-title="Käfer" data-language-autonym="Pälzisch" data-language-local-name="Palatine German" class="interlanguage-link-target">Pälzisch</a></li><li class="interlanguage-link interwiki-ps mw-list-item"><a href="https://ps.wikipedia.org/wiki/%DA%AF%D9%88%D9%86%DA%AF%D9%BC%D9%87" title="گونگټه – Pashto" lang="ps" hreflang="ps" data-title="گونگټه" data-language-autonym="پښتو" data-language-local-name="Pashto" class="interlanguage-link-target">پښتو</a></li><li class="interlanguage-link interwiki-nds mw-list-item"><a href="https://nds.wikipedia.org/wiki/K%C3%A4vers" title="Kävers – Low German" lang="nds" hreflang="nds" data-title="Kävers" data-language-autonym="Plattdüütsch" data-language-local-name="Low German" class="interlanguage-link-target">Plattdüütsch</a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Chrz%C4%85szcze" title="Chrząszcze – Polish" lang="pl" hreflang="pl" data-title="Chrząszcze" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target">Polski</a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Besouro" title="Besouro – Portuguese" lang="pt" hreflang="pt" data-title="Besouro" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target">Português</a></li><li class="interlanguage-link interwiki-ro mw-list-item"><a href="https://ro.wikipedia.org/wiki/Coleopter" title="Coleopter – Romanian" lang="ro" hreflang="ro" data-title="Coleopter" data-language-autonym="Română" data-language-local-name="Romanian" class="interlanguage-link-target">Română</a></li><li class="interlanguage-link interwiki-qu mw-list-item"><a href="https://qu.wikipedia.org/wiki/Suntu" title="Suntu – Quechua" lang="qu" hreflang="qu" data-title="Suntu" data-language-autonym="Runa Simi" data-language-local-name="Quechua" class="interlanguage-link-target">Runa Simi</a></li><li class="interlanguage-link interwiki-ru badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://ru.wikipedia.org/wiki/%D0%96%D0%B5%D1%81%D1%82%D0%BA%D0%BE%D0%BA%D1%80%D1%8B%D0%BB%D1%8B%D0%B5" title="Жесткокрылые – Russian" lang="ru" hreflang="ru" data-title="Жесткокрылые" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target">Русский</a></li><li class="interlanguage-link interwiki-stq mw-list-item"><a href="https://stq.wikipedia.org/wiki/Ruste" title="Ruste – Saterland Frisian" lang="stq" hreflang="stq" data-title="Ruste" data-language-autonym="Seeltersk" data-language-local-name="Saterland Frisian" class="interlanguage-link-target">Seeltersk</a></li><li class="interlanguage-link interwiki-si mw-list-item"><a href="https://si.wikipedia.org/wiki/%E0%B6%9A%E0%B7%94%E0%B6%BB%E0%B7%94%E0%B6%B8%E0%B7%92%E0%B6%AB%E0%B7%92%E0%B6%BA%E0%B7%8F" title="කුරුමිණියා – Sinhala" lang="si" hreflang="si" data-title="කුරුමිණියා" data-language-autonym="සිංහල" data-language-local-name="Sinhala" class="interlanguage-link-target">සිංහල</a></li><li class="interlanguage-link interwiki-simple mw-list-item"><a href="https://simple.wikipedia.org/wiki/Beetle" title="Beetle – Simple English" lang="en-simple" hreflang="en-simple" data-title="Beetle" data-language-autonym="Simple English" data-language-local-name="Simple English" class="interlanguage-link-target">Simple English</a></li><li class="interlanguage-link interwiki-sk mw-list-item"><a href="https://sk.wikipedia.org/wiki/Chrob%C3%A1ky" title="Chrobáky – Slovak" lang="sk" hreflang="sk" data-title="Chrobáky" data-language-autonym="Slovenčina" data-language-local-name="Slovak" class="interlanguage-link-target">Slovenčina</a></li><li class="interlanguage-link interwiki-sl mw-list-item"><a href="https://sl.wikipedia.org/wiki/Hro%C5%A1%C4%8Di" title="Hrošči – Slovenian" lang="sl" hreflang="sl" data-title="Hrošči" data-language-autonym="Slovenščina" data-language-local-name="Slovenian" class="interlanguage-link-target">Slovenščina</a></li><li class="interlanguage-link interwiki-ckb mw-list-item"><a href="https://ckb.wikipedia.org/wiki/%D9%82%D8%A7%D9%84%DB%86%D9%86%DA%86%DB%95" title="قالۆنچە – Central Kurdish" lang="ckb" hreflang="ckb" data-title="قالۆنچە" data-language-autonym="کوردی" data-language-local-name="Central Kurdish" class="interlanguage-link-target">کوردی</a></li><li class="interlanguage-link interwiki-sr badge-Q17437796 badge-featuredarticle mw-list-item" title="featured article badge"><a href="https://sr.wikipedia.org/wiki/%D0%A2%D0%B2%D1%80%D0%B4%D0%BE%D0%BA%D1%80%D0%B8%D0%BB%D1%86%D0%B8" title="Тврдокрилци – Serbian" lang="sr" hreflang="sr" data-title="Тврдокрилци" data-language-autonym="Српски / srpski" data-language-local-name="Serbian" class="interlanguage-link-target">Српски / srpski</a></li><li class="interlanguage-link interwiki-sh mw-list-item"><a href="https://sh.wikipedia.org/wiki/Tvrdokrilci" title="Tvrdokrilci – Serbo-Croatian" lang="sh" hreflang="sh" data-title="Tvrdokrilci" data-language-autonym="Srpskohrvatski / српскохрватски" data-language-local-name="Serbo-Croatian" class="interlanguage-link-target">Srpskohrvatski / српскохрватски</a></li><li class="interlanguage-link interwiki-su mw-list-item"><a href="https://su.wikipedia.org/wiki/Kumbang" title="Kumbang – Sundanese" lang="su" hreflang="su" data-title="Kumbang" data-language-autonym="Sunda" data-language-local-name="Sundanese" class="interlanguage-link-target">Sunda</a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Kovakuoriaiset" title="Kovakuoriaiset – Finnish" lang="fi" hreflang="fi" data-title="Kovakuoriaiset" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target">Suomi</a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Skalbaggar" title="Skalbaggar – Swedish" lang="sv" hreflang="sv" data-title="Skalbaggar" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target">Svenska</a></li><li class="interlanguage-link interwiki-tl mw-list-item"><a href="https://tl.wikipedia.org/wiki/Uwang" title="Uwang – Tagalog" lang="tl" hreflang="tl" data-title="Uwang" data-language-autonym="Tagalog" data-language-local-name="Tagalog" class="interlanguage-link-target">Tagalog</a></li><li class="interlanguage-link interwiki-ta mw-list-item"><a href="https://ta.wikipedia.org/wiki/%E0%AE%B5%E0%AE%A3%E0%AF%8D%E0%AE%9F%E0%AF%81" title="வண்டு – Tamil" lang="ta" hreflang="ta" data-title="வண்டு" data-language-autonym="தமிழ்" data-language-local-name="Tamil" class="interlanguage-link-target">தமிழ்</a></li><li class="interlanguage-link interwiki-kab mw-list-item"><a href="https://kab.wikipedia.org/wiki/Izinzer" title="Izinzer – Kabyle" lang="kab" hreflang="kab" data-title="Izinzer" data-language-autonym="Taqbaylit" data-language-local-name="Kabyle" class="interlanguage-link-target">Taqbaylit</a></li><li class="interlanguage-link interwiki-tt mw-list-item"><a href="https://tt.wikipedia.org/wiki/%D0%9A%D0%BE%D2%A3%D0%B3%D1%8B%D0%B7%D0%BB%D0%B0%D1%80" title="Коңгызлар – Tatar" lang="tt" hreflang="tt" data-title="Коңгызлар" data-language-autonym="Татарча / tatarça" data-language-local-name="Tatar" class="interlanguage-link-target">Татарча / tatarça</a></li><li class="interlanguage-link interwiki-th mw-list-item"><a href="https://th.wikipedia.org/wiki/%E0%B8%AD%E0%B8%B1%E0%B8%99%E0%B8%94%E0%B8%B1%E0%B8%9A%E0%B8%94%E0%B9%89%E0%B8%A7%E0%B8%87" title="อันดับด้วง – Thai" lang="th" hreflang="th" data-title="อันดับด้วง" data-language-autonym="ไทย" data-language-local-name="Thai" class="interlanguage-link-target">ไทย</a></li><li class="interlanguage-link interwiki-tg mw-list-item"><a href="https://tg.wikipedia.org/wiki/%D0%93%D0%B0%D0%BC%D0%B1%D1%83%D1%81%D0%BA%D2%B3%D0%BE" title="Гамбускҳо – Tajik" lang="tg" hreflang="tg" data-title="Гамбускҳо" data-language-autonym="Тоҷикӣ" data-language-local-name="Tajik" class="interlanguage-link-target">Тоҷикӣ</a></li><li class="interlanguage-link interwiki-tr mw-list-item"><a href="https://tr.wikipedia.org/wiki/K%C4%B1n_kanatl%C4%B1lar" title="Kın kanatlılar – Turkish" lang="tr" hreflang="tr" data-title="Kın kanatlılar" data-language-autonym="Türkçe" data-language-local-name="Turkish" class="interlanguage-link-target">Türkçe</a></li><li class="interlanguage-link interwiki-kcg mw-list-item"><a href="https://kcg.wikipedia.org/wiki/A%CC%B1mumu" title="A̱mumu – Tyap" lang="kcg" hreflang="kcg" data-title="A̱mumu" data-language-autonym="Tyap" data-language-local-name="Tyap" class="interlanguage-link-target">Tyap</a></li><li class="interlanguage-link interwiki-uk badge-Q17437798 badge-goodarticle mw-list-item" title="good article badge"><a href="https://uk.wikipedia.org/wiki/%D0%A2%D0%B2%D0%B5%D1%80%D0%B4%D0%BE%D0%BA%D1%80%D0%B8%D0%BB%D1%96" title="Твердокрилі – Ukrainian" lang="uk" hreflang="uk" data-title="Твердокрилі" data-language-autonym="Українська" data-language-local-name="Ukrainian" class="interlanguage-link-target">Українська</a></li><li class="interlanguage-link interwiki-ur mw-list-item"><a href="https://ur.wikipedia.org/wiki/%D8%A8%DA%BE%D9%88%D9%86%D8%B1%D8%A7" title="بھونرا – Urdu" lang="ur" hreflang="ur" data-title="بھونرا" data-language-autonym="اردو" data-language-local-name="Urdu" class="interlanguage-link-target">اردو</a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/B%E1%BB%8D_c%C3%A1nh_c%E1%BB%A9ng" title="Bọ cánh cứng – Vietnamese" lang="vi" hreflang="vi" data-title="Bọ cánh cứng" data-language-autonym="Tiếng Việt" data-language-local-name="Vietnamese" class="interlanguage-link-target">Tiếng Việt</a></li><li class="interlanguage-link interwiki-vls mw-list-item"><a href="https://vls.wikipedia.org/wiki/Keevrs" title="Keevrs – West Flemish" lang="vls" hreflang="vls" data-title="Keevrs" data-language-autonym="West-Vlams" data-language-local-name="West Flemish" class="interlanguage-link-target">West-Vlams</a></li><li class="interlanguage-link interwiki-war mw-list-item"><a href="https://war.wikipedia.org/wiki/Bagang" title="Bagang – Waray" lang="war" hreflang="war" data-title="Bagang" data-language-autonym="Winaray" data-language-local-name="Waray" class="interlanguage-link-target">Winaray</a></li><li class="interlanguage-link interwiki-wuu mw-list-item"><a href="https://wuu.wikipedia.org/wiki/%E9%9E%98%E7%BF%85%E7%9B%AE" title="鞘翅目 – Wu" lang="wuu" hreflang="wuu" data-title="鞘翅目" data-language-autonym="吴语" data-language-local-name="Wu" class="interlanguage-link-target">吴语</a></li><li class="interlanguage-link interwiki-yi mw-list-item"><a href="https://yi.wikipedia.org/wiki/%D7%96%D7%A9%D7%95%D7%A7" title="זשוק – Yiddish" lang="yi" hreflang="yi" data-title="זשוק" data-language-autonym="ייִדיש" data-language-local-name="Yiddish" class="interlanguage-link-target">ייִדיש</a></li><li class="interlanguage-link interwiki-zh-yue mw-list-item"><a href="https://zh-yue.wikipedia.org/wiki/%E7%94%B2%E8%9F%B2" title="甲蟲 – Cantonese" lang="yue" hreflang="yue" data-title="甲蟲" data-language-autonym="粵語" data-language-local-name="Cantonese" class="interlanguage-link-target">粵語</a></li><li class="interlanguage-link interwiki-bat-smg mw-list-item"><a href="https://bat-smg.wikipedia.org/wiki/Vombuol%C4%97" title="Vombuolė – Samogitian" lang="sgs" hreflang="sgs" data-title="Vombuolė" data-language-autonym="Žemaitėška" data-language-local-name="Samogitian" class="interlanguage-link-target">Žemaitėška</a></li><li class="interlanguage-link interwiki-zh mw-list-item"><a href="https://zh.wikipedia.org/wiki/%E9%9E%98%E7%BF%85%E7%9B%AE" title="鞘翅目 – Chinese" lang="zh" hreflang="zh" data-title="鞘翅目" data-language-autonym="中文" data-language-local-name="Chinese" class="interlanguage-link-target">中文</a></li> </ul> <div class="after-portlet after-portlet-lang"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q22671#sitelinks-wikipedia" title="Edit interlanguage links" class="wbc-editpage">Edit links</a></div> </div> </div> </div> </header> <div class="vector-page-toolbar"> <div class="vector-page-toolbar-container"> <div id="left-navigation"> <nav aria-label="Namespaces"> <div id="p-associated-pages" class="vector-menu vector-menu-tabs mw-portlet mw-portlet-associated-pages" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-nstab-main" class="selected vector-tab-noicon mw-list-item"><a href="/wiki/Beetle" title="View the content page [c]" accesskey="c">Article</a></li><li id="ca-talk" class="vector-tab-noicon mw-list-item"><a href="/wiki/Talk:Beetle" rel="discussion" title="Discuss improvements to the content page [t]" accesskey="t">Talk</a></li> </ul> </div> </div> <div id="vector-variants-dropdown" class="vector-dropdown emptyPortlet" > <input type="checkbox" id="vector-variants-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-variants-dropdown" class="vector-dropdown-checkbox " aria-label="Change language variant" > <label id="vector-variants-dropdown-label" for="vector-variants-dropdown-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet" aria-hidden="true" >English </label> <div class="vector-dropdown-content"> <div id="p-variants" class="vector-menu mw-portlet mw-portlet-variants emptyPortlet" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> </ul> </div> </div> </div> </div> </nav> </div> <div id="right-navigation" class="vector-collapsible"> <nav aria-label="Views"> <div id="p-views" class="vector-menu vector-menu-tabs mw-portlet mw-portlet-views" > <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-view" class="selected vector-tab-noicon mw-list-item"><a href="/wiki/Beetle">Read</a></li><li id="ca-edit" class="vector-tab-noicon mw-list-item"><a href="/w/index.php?title=Beetle&action=edit" title="Edit this page [e]" accesskey="e">Edit</a></li><li id="ca-history" class="vector-tab-noicon mw-list-item"><a href="/w/index.php?title=Beetle&action=history" title="Past revisions of this page [h]" accesskey="h">View history</a></li> </ul> </div> </div> </nav> <nav class="vector-page-tools-landmark" aria-label="Page tools"> <div id="vector-page-tools-dropdown" class="vector-dropdown vector-page-tools-dropdown" > <input type="checkbox" id="vector-page-tools-dropdown-checkbox" role="button" aria-haspopup="true" data-event-name="ui.dropdown-vector-page-tools-dropdown" class="vector-dropdown-checkbox " aria-label="Tools" > <label id="vector-page-tools-dropdown-label" for="vector-page-tools-dropdown-checkbox" class="vector-dropdown-label cdx-button cdx-button--fake-button cdx-button--fake-button--enabled cdx-button--weight-quiet" aria-hidden="true" >Tools </label> <div class="vector-dropdown-content"> <div id="vector-page-tools-unpinned-container" class="vector-unpinned-container"> <div id="vector-page-tools" class="vector-page-tools vector-pinnable-element"> <div class="vector-pinnable-header vector-page-tools-pinnable-header vector-pinnable-header-unpinned" data-feature-name="page-tools-pinned" data-pinnable-element-id="vector-page-tools" data-pinned-container-id="vector-page-tools-pinned-container" data-unpinned-container-id="vector-page-tools-unpinned-container" > <div class="vector-pinnable-header-label">Tools</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-page-tools.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-page-tools.unpin">hide</button> </div> <div id="p-cactions" class="vector-menu mw-portlet mw-portlet-cactions emptyPortlet vector-has-collapsible-items" title="More options" > <div class="vector-menu-heading"> Actions </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="ca-more-view" class="selected vector-more-collapsible-item mw-list-item"><a href="/wiki/Beetle">Read</a></li><li id="ca-more-edit" class="vector-more-collapsible-item mw-list-item"><a href="/w/index.php?title=Beetle&action=edit" title="Edit this page [e]" accesskey="e">Edit</a></li><li id="ca-more-history" class="vector-more-collapsible-item mw-list-item"><a href="/w/index.php?title=Beetle&action=history">View history</a></li> </ul> </div> </div> <div id="p-tb" class="vector-menu mw-portlet mw-portlet-tb" > <div class="vector-menu-heading"> General </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="t-whatlinkshere" class="mw-list-item"><a href="/wiki/Special:WhatLinksHere/Beetle" title="List of all English Wikipedia pages containing links to this page [j]" accesskey="j">What links here</a></li><li id="t-recentchangeslinked" class="mw-list-item"><a href="/wiki/Special:RecentChangesLinked/Beetle" rel="nofollow" title="Recent changes in pages linked from this page [k]" accesskey="k">Related changes</a></li><li id="t-upload" class="mw-list-item"><a href="/wiki/Wikipedia:File_Upload_Wizard" title="Upload files [u]" accesskey="u">Upload file</a></li><li id="t-specialpages" class="mw-list-item"><a href="/wiki/Special:SpecialPages" title="A list of all special pages [q]" accesskey="q">Special pages</a></li><li id="t-permalink" class="mw-list-item"><a href="/w/index.php?title=Beetle&oldid=1257880098" title="Permanent link to this revision of this page">Permanent link</a></li><li id="t-info" class="mw-list-item"><a href="/w/index.php?title=Beetle&action=info" title="More information about this page">Page information</a></li><li id="t-cite" class="mw-list-item"><a href="/w/index.php?title=Special:CiteThisPage&page=Beetle&id=1257880098&wpFormIdentifier=titleform" title="Information on how to cite this page">Cite this page</a></li><li id="t-urlshortener" class="mw-list-item"><a href="/w/index.php?title=Special:UrlShortener&url=https%3A%2F%2Fen.wikipedia.org%2Fwiki%2FBeetle">Get shortened URL</a></li><li id="t-urlshortener-qrcode" class="mw-list-item"><a href="/w/index.php?title=Special:QrCode&url=https%3A%2F%2Fen.wikipedia.org%2Fwiki%2FBeetle">Download QR code</a></li> </ul> </div> </div> <div id="p-coll-print_export" class="vector-menu mw-portlet mw-portlet-coll-print_export" > <div class="vector-menu-heading"> Print/export </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li id="coll-download-as-rl" class="mw-list-item"><a href="/w/index.php?title=Special:DownloadAsPdf&page=Beetle&action=show-download-screen" title="Download this page as a PDF file">Download as PDF</a></li><li id="t-print" class="mw-list-item"><a href="/w/index.php?title=Beetle&printable=yes" title="Printable version of this page [p]" accesskey="p">Printable version</a></li> </ul> </div> </div> <div id="p-wikibase-otherprojects" class="vector-menu mw-portlet mw-portlet-wikibase-otherprojects" > <div class="vector-menu-heading"> In other projects </div> <div class="vector-menu-content"> <ul class="vector-menu-content-list"> <li class="wb-otherproject-link wb-otherproject-commons mw-list-item"><a href="https://commons.wikimedia.org/wiki/Category:Coleoptera" hreflang="en">Wikimedia Commons</a></li><li class="wb-otherproject-link wb-otherproject-species mw-list-item"><a href="https://species.wikimedia.org/wiki/Coleoptera" hreflang="en">Wikispecies</a></li><li class="wb-otherproject-link wb-otherproject-wikiquote mw-list-item"><a href="https://en.wikiquote.org/wiki/Beetles" hreflang="en">Wikiquote</a></li><li id="t-wikibase" class="wb-otherproject-link wb-otherproject-wikibase-dataitem mw-list-item"><a href="https://www.wikidata.org/wiki/Special:EntityPage/Q22671" title="Structured data on this page hosted by Wikidata [g]" accesskey="g">Wikidata item</a></li> </ul> </div> </div> </div> </div> </div> </div> </nav> </div> </div> </div> <div class="vector-column-end"> <div class="vector-sticky-pinned-container"> <nav class="vector-page-tools-landmark" aria-label="Page tools"> <div id="vector-page-tools-pinned-container" class="vector-pinned-container"> </div> </nav> <nav class="vector-appearance-landmark" aria-label="Appearance"> <div id="vector-appearance-pinned-container" class="vector-pinned-container"> <div id="vector-appearance" class="vector-appearance vector-pinnable-element"> <div class="vector-pinnable-header vector-appearance-pinnable-header vector-pinnable-header-pinned" data-feature-name="appearance-pinned" data-pinnable-element-id="vector-appearance" data-pinned-container-id="vector-appearance-pinned-container" data-unpinned-container-id="vector-appearance-unpinned-container" > <div class="vector-pinnable-header-label">Appearance</div> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-pin-button" data-event-name="pinnable-header.vector-appearance.pin">move to sidebar</button> <button class="vector-pinnable-header-toggle-button vector-pinnable-header-unpin-button" data-event-name="pinnable-header.vector-appearance.unpin">hide</button> </div> </div> </div> </nav> </div> </div> <div id="bodyContent" class="vector-body" aria-labelledby="firstHeading" data-mw-ve-target-container> <div class="vector-body-before-content"> <div class="mw-indicators"> <div id="mw-indicator-good-star" class="mw-indicator"><div class="mw-parser-output"><a href="/wiki/Wikipedia:Good_articles*" title="This is a good article. Click here for more information."><img alt="This is a good article. Click here for more information." src="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/19px-Symbol_support_vote.svg.png" decoding="async" width="19" height="20" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/29px-Symbol_support_vote.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/9/94/Symbol_support_vote.svg/39px-Symbol_support_vote.svg.png 2x" data-file-width="180" data-file-height="185" /></a></div></div> </div> <div id="siteSub" class="noprint">From Wikipedia, the free encyclopedia</div> </div> <div id="contentSub"><div id="mw-content-subtitle"></div></div> <div id="mw-content-text" 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">Order of insects</div> <style data-mw-deduplicate="TemplateStyles:r1236090951">.mw-parser-output .hatnote{font-style:italic}.mw-parser-output div.hatnote{padding-left:1.6em;margin-bottom:0.5em}.mw-parser-output .hatnote i{font-style:normal}.mw-parser-output .hatnote+link+.hatnote{margin-top:-0.5em}@media print{body.ns-0 .mw-parser-output .hatnote{display:none!important}}</style><div role="note" class="hatnote navigation-not-searchable">For the car, see <a href="/wiki/Volkswagen_Beetle" title="Volkswagen Beetle">Volkswagen Beetle</a>. For other uses, see <a href="/wiki/Beetle_(disambiguation)" class="mw-disambig" title="Beetle (disambiguation)">Beetle (disambiguation)</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"Beetles" redirects here. For the English rock band, see <a href="/wiki/The_Beatles" title="The Beatles">the Beatles</a>.</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">"Coleoptera" redirects here. Not to be confused with <a href="/wiki/Coeloptera" title="Coeloptera">Coeloptera</a> or <a href="/wiki/Cleopatra" title="Cleopatra">Cleopatra</a>.</div> <table class="infobox biota" style="text-align: left; width: 200px; font-size: 100%"> <tbody><tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)">Beetle <div style="font-size: 85%;">Temporal range: 299–0 <a href="/wiki/Megaannum" class="mw-redirect" title="Megaannum">Ma</a> <div id="Timeline-row" style="margin: 4px auto 0; clear:both; width:220px; padding:0px; height:18px; overflow:visible; white-space:nowrap; border:1px #666; border-style:solid none; position:relative; z-index:0; font-size:97%;"> <div style="position:absolute; height:100%; left:0px; width:207.23076923077px; padding-left:5px; text-align:left; background-color:rgb(254,217,106); background-image: linear-gradient(to right, rgba(255,255,255,1), rgba(254,217,106,1) 15%, rgba(254,217,106,1));"><a href="/wiki/Precambrian" title="Precambrian">PreꞒ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,160,86); left:37.636923076923px; width:18.073846153846px;"><a href="/wiki/Cambrian" title="Cambrian">Ꞓ</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(0,146,112); left:55.710769230769px; width:14.08px;"><a href="/wiki/Ordovician" title="Ordovician">O</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(179,225,182); left:69.790769230769px; width:8.3261538461539px;"><a href="/wiki/Silurian" title="Silurian">S</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(203,140,55); left:78.116923076923px; width:20.409230769231px;"><a href="/wiki/Devonian" title="Devonian">D</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(103,165,153); left:98.526153846154px; width:20.307692307692px;"><a href="/wiki/Carboniferous" title="Carboniferous">C</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(240,64,40); left:118.83384615385px; width:15.907015384615px;"><a href="/wiki/Permian" title="Permian">P</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(129,43,146); left:134.74086153846px; width:17.092984615385px;"><a href="/wiki/Triassic" title="Triassic">T</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(52,178,201); left:151.83384615385px; width:19.089230769231px;"><a href="/wiki/Jurassic" title="Jurassic">J</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(127,198,78); left:170.92307692308px; width:26.738461538462px;"><a href="/wiki/Cretaceous" title="Cretaceous">K</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(253,154,82); left:197.66153846154px; width:14.543692307692px;"><a href="/wiki/Paleogene" title="Paleogene">Pg</a></div> <div style="position:absolute; height:100%; text-align:center; background-color:rgb(255,230,25); left:212.20523076923px; width:6.9215384615385px;"><a href="/wiki/Neogene" title="Neogene">N</a></div> <div id="end-border" style="position:absolute; height:100%; background-color:#666; width:1px; left:219px"></div><div style="margin:0 auto; line-height:0; clear:both; width:220px; padding:0px; height:8px; overflow:visible; background-color:transparent; position:relative; top:-4px; z-index:100;"><div style="position:absolute; height:8px; left:118.8px; width:101.2px; background-color:#360; opacity:0.42;"></div> <div style="position:absolute; height:8px; left:118.8px; width:101.2px; background-color:#360; opacity:1;"></div> <div style="position:absolute; height:6px; top:1px; left:119.8px; width:99.2px; background-color:#6c3;"></div> </div> </div>Earliest <a href="/wiki/Permian" title="Permian">Permian</a>–<a href="/wiki/Holocene" title="Holocene">Present</a></div> </th></tr> <tr> <td colspan="2" style="text-align: center"><a href="/wiki/File:Coleoptera_collage.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Coleoptera_collage.png/220px-Coleoptera_collage.png" decoding="async" width="220" height="210" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/20/Coleoptera_collage.png/330px-Coleoptera_collage.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/20/Coleoptera_collage.png/440px-Coleoptera_collage.png 2x" data-file-width="1411" data-file-height="1345" /></a> </td></tr> <tr> <td colspan="2" style="text-align: center; font-size: 88%">Clockwise from top left: female golden stag beetle (<a href="/wiki/Lamprima_aurata" title="Lamprima aurata">Lamprima aurata</a>), <a href="/wiki/Rhinoceros_beetle" class="mw-redirect" title="Rhinoceros beetle">rhinoceros beetle</a> (Megasoma sp.), long nose weevil (<a href="/wiki/Rhinotia_hemistictus" class="mw-redirect" title="Rhinotia hemistictus">Rhinotia hemistictus</a>), cowboy beetle (<a href="/wiki/Chondropyga_dorsalis" title="Chondropyga dorsalis">Chondropyga dorsalis</a>), and a species of <a href="/wiki/Amblytelus" title="Amblytelus">Amblytelus</a> </td></tr> <tr> <th colspan="2" style="color:inherit; min-width:15em; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Taxonomy_(biology)" title="Taxonomy (biology)">Scientific classification</a> <a href="/wiki/Template:Taxonomy/Coleoptera" title="Edit this classification"><img alt="Edit this classification" src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/15px-OOjs_UI_icon_edit-ltr.svg.png" decoding="async" width="15" height="15" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/23px-OOjs_UI_icon_edit-ltr.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8a/OOjs_UI_icon_edit-ltr.svg/30px-OOjs_UI_icon_edit-ltr.svg.png 2x" data-file-width="20" data-file-height="20" /></a> </th></tr> <tr> <td>Domain: </td> <td><a href="/wiki/Eukaryote" title="Eukaryote">Eukaryota</a> </td></tr> <tr> <td>Kingdom: </td> <td><a href="/wiki/Animal" title="Animal">Animalia</a> </td></tr> <tr> <td>Phylum: </td> <td><a href="/wiki/Arthropod" title="Arthropod">Arthropoda</a> </td></tr> <tr> <td>Class: </td> <td><a href="/wiki/Insect" title="Insect">Insecta</a> </td></tr> <tr> <td>Clade: </td> <td><a href="/wiki/Coleopterida" title="Coleopterida">Coleopterida</a> </td></tr> <tr> <td>Order: </td> <td><a class="mw-selflink selflink">Coleoptera</a> <a href="/wiki/Linnaeus" class="mw-redirect" title="Linnaeus">Linnaeus</a>, <a href="/wiki/10th_edition_of_Systema_Naturae" title="10th edition of Systema Naturae">1758</a> </td></tr> <tr> <th colspan="2" style="color:inherit; text-align: center; background-color: rgb(235,235,210)"><a href="/wiki/Suborders" class="mw-redirect" title="Suborders">Suborders</a><a href="#cite_note-Bouchard_et_al_2011-1">[1]</a> </th></tr> <tr> <td colspan="2" style="text-align: left"> <ul><li><a href="/wiki/Adephaga" title="Adephaga">Adephaga</a></li> <li><a href="/wiki/Archostemata" title="Archostemata">Archostemata</a></li> <li><a href="/wiki/Myxophaga" title="Myxophaga">Myxophaga</a></li> <li><a href="/wiki/Polyphaga" title="Polyphaga">Polyphaga</a></li> <li>†<a href="/wiki/Protocoleoptera" title="Protocoleoptera">Protocoleoptera</a></li></ul> See <a href="/wiki/Subgroups_of_the_order_Coleoptera" class="mw-redirect" title="Subgroups of the order Coleoptera">subgroups of the order Coleoptera</a> </td></tr> </tbody></table><style data-mw-deduplicate="TemplateStyles:r1238732961">@media screen{html.skin-theme-clientpref-night .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-night .mw-parser-output .infobox.biota img{background:transparent}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .infobox.biota tr{background:transparent!important}html.skin-theme-clientpref-os .mw-parser-output .infobox.biota img{background:white}}.mw-parser-output .infobox.biota .taxobox-edit-taxonomy img{background:transparent!important}body.skin-vector .mw-parser-output table.biota.infobox{margin-top:0.5em}body.skin--responsive .mw-parser-output table.biota.infobox tr.taxonrow td{padding:2px 10px}</style> Beetles are <a href="/wiki/Insect" title="Insect">insects</a> that form the <a href="/wiki/Taxonomic_rank" title="Taxonomic rank">order</a> Coleoptera (<a href="/wiki/Help:IPA/English" title="Help:IPA/English">/koʊliːˈɒptərə/</a>), in the superorder <a href="/wiki/Holometabola" title="Holometabola">Holometabola</a>. Their front pair of wings are hardened into wing-cases, <a href="/wiki/Elytra" class="mw-redirect" title="Elytra">elytra</a>, distinguishing them from most other insects. The Coleoptera, with about 400,000 described species, is the largest of all orders, constituting almost 40% of described insects and 25% of all known animal species;<a href="#cite_note-Stork2018-2">[2]</a> new species are discovered frequently, with estimates suggesting that there are between 0.9 and 2.1 million total species. However, the number of beetle species is challenged by the number of species in <a href="/wiki/Fly" title="Fly">dipterans</a> (flies) and <a href="/wiki/Hymenoptera" title="Hymenoptera">hymenopterans</a> (wasps). Found in almost every habitat except the sea and the <a href="/wiki/Polar_region" class="mw-redirect" title="Polar region">polar regions</a>, they interact with their <a href="/wiki/Ecosystem" title="Ecosystem">ecosystems</a> in several ways: beetles often feed on plants and <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a>, break down animal and plant debris, and eat other <a href="/wiki/Invertebrate" title="Invertebrate">invertebrates</a>. Some species are serious agricultural pests, such as the <a href="/wiki/Colorado_potato_beetle" title="Colorado potato beetle">Colorado potato beetle</a>, while others such as <a href="/wiki/Coccinellidae" title="Coccinellidae">Coccinellidae</a> (ladybirds or ladybugs) eat <a href="/wiki/Aphid" title="Aphid">aphids</a>, <a href="/wiki/Scale_insect" title="Scale insect">scale insects</a>, <a href="/wiki/Thrips" title="Thrips">thrips</a>, and other plant-sucking insects that damage crops. Some others also have unusual characteristics, such as <a href="/wiki/Fireflies" class="mw-redirect" title="Fireflies">fireflies</a>, which use a light-emitting organ for mating and communication purposes. Beetles typically have a particularly hard <a href="/wiki/Exoskeleton" title="Exoskeleton">exoskeleton</a> including the <a href="/wiki/Elytra" class="mw-redirect" title="Elytra">elytra</a>, though some such as the <a href="/wiki/Rove_beetles" class="mw-redirect" title="Rove beetles">rove beetles</a> have very short elytra while <a href="/wiki/Blister_beetle" title="Blister beetle">blister beetles</a> have softer elytra. The general <a href="/wiki/Anatomy" title="Anatomy">anatomy</a> of a beetle is quite uniform and typical of insects, although there are several examples of novelty, such as adaptations in <a href="/wiki/Water_beetle" title="Water beetle">water beetles</a> which trap air bubbles under the elytra for use while diving. Beetles are <a href="/wiki/Holometabola" title="Holometabola">holometabolans</a>, which means that they undergo complete <a href="/wiki/Metamorphosis" title="Metamorphosis">metamorphosis</a>, with a series of conspicuous and relatively abrupt changes in body structure between hatching and becoming adult after a relatively immobile <a href="/wiki/Pupa" title="Pupa">pupal</a> stage. Some, such as <a href="/wiki/Stag_beetle" title="Stag beetle">stag beetles</a>, have a marked <a href="/wiki/Sexual_dimorphism" title="Sexual dimorphism">sexual dimorphism</a>, the males possessing enormously enlarged <a href="/wiki/Mandible_(insect_mouthpart)" title="Mandible (insect mouthpart)">mandibles</a> which they use to fight other males. Many beetles are <a href="/wiki/Aposematic" class="mw-redirect" title="Aposematic">aposematic</a>, with bright colors and patterns warning of their toxicity, while others are harmless <a href="/wiki/Batesian_mimics" class="mw-redirect" title="Batesian mimics">Batesian mimics</a> of such insects. Many beetles, including those that live in sandy places, have effective <a href="/wiki/Camouflage" title="Camouflage">camouflage</a>. Beetles are prominent <a href="/wiki/Insects_in_culture" class="mw-redirect" title="Insects in culture">in human culture</a>, from the <a href="/wiki/Scarab_(artifact)" title="Scarab (artifact)">sacred scarabs</a> of <a href="/wiki/Ancient_Egypt" title="Ancient Egypt">ancient Egypt</a> to <a href="/wiki/Beetlewing" title="Beetlewing">beetlewing</a> art and use as <a href="/wiki/Pet" title="Pet">pets</a> or <a href="/wiki/Insect_fighting" title="Insect fighting">fighting insects</a> for entertainment and gambling. Many beetle groups are brightly and attractively colored making them objects of collection and decorative displays. Over 300 species are <a href="/wiki/Entomophagy" title="Entomophagy">used as food</a>, mostly as <a href="/wiki/Larva" title="Larva">larvae</a>; species widely consumed include <a href="/wiki/Mealworm" title="Mealworm">mealworms</a> and <a href="/wiki/Rhinoceros_beetle" class="mw-redirect" title="Rhinoceros beetle">rhinoceros beetle</a> larvae. However, the major impact of beetles on human life is as agricultural, forestry, and horticultural <a href="/wiki/Pest_(organism)" title="Pest (organism)">pests</a>. Serious pest species include the <a href="/wiki/Boll_weevil" title="Boll weevil">boll weevil</a> of cotton, the <a href="/wiki/Colorado_potato_beetle" title="Colorado potato beetle">Colorado potato beetle</a>, the <a href="/wiki/Brontispa_longissima" title="Brontispa longissima">coconut hispine beetle</a>, the <a href="/wiki/Mountain_pine_beetle" title="Mountain pine beetle">mountain pine beetle</a>, and many others. Most beetles, however, do not cause economic damage and some, such as numerous species of <a href="/wiki/Lady_beetle" class="mw-redirect" title="Lady beetle">lady beetles</a>, are beneficial by helping to control insect pests. <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Etymology">Etymology</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=1" title="Edit section: Etymology">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Coleoptera_SMNK.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/Coleoptera_SMNK.jpg/220px-Coleoptera_SMNK.jpg" decoding="async" width="220" height="320" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/c0/Coleoptera_SMNK.jpg/330px-Coleoptera_SMNK.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/c0/Coleoptera_SMNK.jpg/440px-Coleoptera_SMNK.jpg 2x" data-file-width="3083" data-file-height="4487" /></a><figcaption>Coleoptera at the <a href="/wiki/State_Museum_of_Natural_History,_Karlsruhe" class="mw-redirect" title="State Museum of Natural History, Karlsruhe">State Museum of Natural History, Karlsruhe</a>, Germany</figcaption></figure> The name of the taxonomic order, Coleoptera, comes from the <a href="/wiki/Ancient_Greek" title="Ancient Greek">Greek</a> koleopteros (κολεόπτερος), given to the group by <a href="/wiki/Aristotle" title="Aristotle">Aristotle</a> for their <a href="/wiki/Elytra" class="mw-redirect" title="Elytra">elytra</a>, hardened shield-like forewings, from koleos, sheath, and pteron, wing. The English name beetle comes from the <a href="/wiki/Old_English" title="Old English">Old English</a> word bitela, little biter, related to bītan (to bite),<a href="#cite_note-Etymology-3">[3]</a><a href="#cite_note-Etymology_Beetle-4">[4]</a> leading to <a href="/wiki/Middle_English" title="Middle English">Middle English</a> betylle.<a href="#cite_note-Merriam_Dictionary-5">[5]</a> Another Old English name for beetle is ċeafor, chafer, used in names such as <a href="/wiki/Cockchafer" title="Cockchafer">cockchafer</a>, from the Proto-Germanic *kebrô ("beetle"; compare German Käfer, Dutch kever, Afrikaans kewer).<a href="#cite_note-Etymology_Chafer-6">[6]</a> <div class="mw-heading mw-heading2"><h2 id="Distribution_and_diversity">Distribution and diversity</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=2" title="Edit section: Distribution and diversity">edit</a>]</div> Beetles are by far the largest order of insects: the roughly 400,000 species make up about 40% of all insect species so far described, and about 25% of all animal species.<a href="#cite_note-Bouchard_et_al_2011-1">[1]</a><a href="#cite_note-Gilliott-7">[7]</a><a href="#cite_note-insenc-8">[8]</a><a href="#cite_note-Rosenzweig_1995-9">[9]</a><a href="#cite_note-Hunt_et_al_2007-10">[10]</a><a href="#cite_note-Hammond_1992-11">[11]</a> A 2015 study provided four independent estimates of the total number of beetle species, giving a mean estimate of some 1.5 million with a "surprisingly narrow range"<a href="#cite_note-Stork-12">[12]</a> spanning all four estimates from a minimum of 0.9 to a maximum of 2.1 million beetle species. The four estimates made use of host-specificity relationships (1.5 to 1.9 million), ratios with other taxa (0.9 to 1.2 million), plant:beetle ratios (1.2 to 1.3), and extrapolations based on body size by year of description (1.7 to 2.1 million).<a href="#cite_note-Stork-12">[12]</a><a href="#cite_note-13">[13]</a> This immense diversity led the evolutionary biologist <a href="/wiki/J._B._S._Haldane" title="J. B. S. Haldane">J. B. S. Haldane</a> to quip, when some <a href="/wiki/Theologians" class="mw-redirect" title="Theologians">theologians</a> asked him what could be inferred about the mind of the <a href="/wiki/Christian_God" class="mw-redirect" title="Christian God">Christian God</a> from the works of His Creation, "An inordinate fondness for beetles".<a href="#cite_note-Hutchinson_1959-14">[14]</a> However, the ranking of beetles as most diverse has been challenged. Multiple studies posit that Diptera (flies) and/or Hymenoptera (sawflies, wasps, ants and bees) may have more species.<a href="#cite_note-15">[15]</a><a href="#cite_note-16">[16]</a><a href="#cite_note-17">[17]</a> Beetles are found in nearly all habitats, including freshwater and coastal habitats, wherever vegetative foliage is found, from trees and their bark to flowers, leaves, and underground near roots - even inside plants in galls, in every plant tissue, including dead or decaying ones.<a href="#cite_note-18">[18]</a> Tropical forest canopies have a large and diverse fauna of beetles,<a href="#cite_note-19">[19]</a> including <a href="/wiki/Carabidae" class="mw-redirect" title="Carabidae">Carabidae</a>,<a href="#cite_note-20">[20]</a> <a href="/wiki/Chrysomelidae" class="mw-redirect" title="Chrysomelidae">Chrysomelidae</a>,<a href="#cite_note-21">[21]</a> and <a href="/wiki/Scarabaeidae" title="Scarabaeidae">Scarabaeidae</a>.<a href="#cite_note-22">[22]</a> The heaviest beetle, indeed the heaviest insect stage, is the <a href="/wiki/Larva" title="Larva">larva</a> of the <a href="/wiki/Goliath_beetle" class="mw-redirect" title="Goliath beetle">goliath beetle</a>, Goliathus goliatus, which can attain a mass of at least 115 g (4.1 oz) and a length of 11.5 cm (4.5 in). Adult male goliath beetles are the heaviest beetle in its adult stage, weighing 70–100 g (2.5–3.5 oz) and measuring up to 11 cm (4.3 in).<a href="#cite_note-23">[23]</a> Adult <a href="/wiki/Elephant_beetle" title="Elephant beetle">elephant beetles</a>, <a href="/wiki/Megasoma_elephas" class="mw-redirect" title="Megasoma elephas">Megasoma elephas</a> and <a href="/wiki/Megasoma_actaeon" class="mw-redirect" title="Megasoma actaeon">Megasoma actaeon</a> often reach 50 g (1.8 oz) and 10 cm (3.9 in).<a href="#cite_note-Williams2001-24">[24]</a> The longest beetle is the <a href="/wiki/Hercules_beetle" title="Hercules beetle">Hercules beetle</a> Dynastes hercules, with a maximum overall length of at least 16.7 cm (6.6 in) including the very long <a href="/wiki/Prothorax" title="Prothorax">pronotal</a> horn. The smallest recorded beetle and the smallest free-living insect (as of 2015<a class="external text" href="https://en.wikipedia.org/w/index.php?title=Beetle&action=edit">[update]</a>), is the <a href="/wiki/Featherwing_beetle" class="mw-redirect" title="Featherwing beetle">featherwing beetle</a> <a href="/wiki/Scydosella_musawasensis" class="mw-redirect" title="Scydosella musawasensis">Scydosella musawasensis</a> which may measure as little as 325 <a href="/wiki/%CE%9Cm" class="mw-redirect" title="Μm">μm</a> in length.<a href="#cite_note-25">[25]</a> <ul class="gallery mw-gallery-packed"> <li class="gallerybox" style="width: 138px"> <div class="thumb" style="width: 136px;"><a href="/wiki/File:Titanus_giganteus_MHNT_dos.jpg" class="mw-file-description" title="Titan beetle, Titanus giganteus, a tropical longhorn, is one of the largest and heaviest insects in the world."><img alt="Titan beetle, Titanus giganteus, a tropical longhorn, is one of the largest and heaviest insects in the world." src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Titanus_giganteus_MHNT_dos.jpg/204px-Titanus_giganteus_MHNT_dos.jpg" decoding="async" width="136" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Titanus_giganteus_MHNT_dos.jpg/306px-Titanus_giganteus_MHNT_dos.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5e/Titanus_giganteus_MHNT_dos.jpg/407px-Titanus_giganteus_MHNT_dos.jpg 2x" data-file-width="2809" data-file-height="3612" /></a></div> <div class="gallerytext"><a href="/wiki/Titan_beetle" title="Titan beetle">Titan beetle</a>, Titanus giganteus, a tropical <a href="/wiki/Longhorn_beetle" title="Longhorn beetle">longhorn</a>, is one of the largest and heaviest insects in the world.</div> </li> <li class="gallerybox" style="width: 134.66666666667px"> <div class="thumb" style="width: 132.66666666667px;"><a href="/wiki/File:Scydosella_musawasensis.jpg" class="mw-file-description" title="Scydosella musawasensis, the smallest known beetle: scale bar (right) is 50 μm."><img alt="Scydosella musawasensis, the smallest known beetle: scale bar (right) is 50 μm." src="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Scydosella_musawasensis.jpg/199px-Scydosella_musawasensis.jpg" decoding="async" width="133" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/86/Scydosella_musawasensis.jpg/299px-Scydosella_musawasensis.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/86/Scydosella_musawasensis.jpg/398px-Scydosella_musawasensis.jpg 2x" data-file-width="1512" data-file-height="1988" /></a></div> <div class="gallerytext"><a href="/wiki/Scydosella_musawasensis" class="mw-redirect" title="Scydosella musawasensis">Scydosella musawasensis</a>, the smallest known beetle: scale bar (right) is 50 μm.</div> </li> <li class="gallerybox" style="width: 268px"> <div class="thumb" style="width: 266px;"><a href="/wiki/File:Dynastes_hercules_ecuatorianus_MHNT.jpg" class="mw-file-description" title="Hercules beetle, Dynastes hercules ecuatorianus, the longest of all beetles."><img alt="Hercules beetle, Dynastes hercules ecuatorianus, the longest of all beetles." src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Dynastes_hercules_ecuatorianus_MHNT.jpg/399px-Dynastes_hercules_ecuatorianus_MHNT.jpg" decoding="async" width="266" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Dynastes_hercules_ecuatorianus_MHNT.jpg/598px-Dynastes_hercules_ecuatorianus_MHNT.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Dynastes_hercules_ecuatorianus_MHNT.jpg/797px-Dynastes_hercules_ecuatorianus_MHNT.jpg 2x" data-file-width="5861" data-file-height="3853" /></a></div> <div class="gallerytext"><a href="/wiki/Hercules_beetle" title="Hercules beetle">Hercules beetle</a>, Dynastes hercules ecuatorianus, the longest of all beetles.</div> </li> <li class="gallerybox" style="width: 170px"> <div class="thumb" style="width: 168px;"><a href="/wiki/File:Protaetia-cuprea-ignicollis-IZE-257.jpg" class="mw-file-description" title="Iridescent Protaetia cuprea feeding on thistle."><img alt="Iridescent Protaetia cuprea feeding on thistle." src="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Protaetia-cuprea-ignicollis-IZE-257.jpg/252px-Protaetia-cuprea-ignicollis-IZE-257.jpg" decoding="async" width="168" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/07/Protaetia-cuprea-ignicollis-IZE-257.jpg/378px-Protaetia-cuprea-ignicollis-IZE-257.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/07/Protaetia-cuprea-ignicollis-IZE-257.jpg/504px-Protaetia-cuprea-ignicollis-IZE-257.jpg 2x" data-file-width="906" data-file-height="942" /></a></div> <div class="gallerytext">Iridescent <a href="/wiki/Protaetia_cuprea" title="Protaetia cuprea">Protaetia cuprea</a> feeding on thistle.</div> </li> </ul> <div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=3" title="Edit section: Evolution">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Late_Paleozoic_and_Triassic">Late Paleozoic and Triassic</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=4" title="Edit section: Late Paleozoic and Triassic">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Moravocoleus_permianus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Moravocoleus_permianus.jpg/220px-Moravocoleus_permianus.jpg" decoding="async" width="220" height="185" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Moravocoleus_permianus.jpg/330px-Moravocoleus_permianus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2b/Moravocoleus_permianus.jpg/440px-Moravocoleus_permianus.jpg 2x" data-file-width="718" data-file-height="603" /></a><figcaption>Fossil and life restoration of Moravocoleus permianus (<a href="/wiki/Tshekardocoleidae" title="Tshekardocoleidae">Tshekardocoleidae</a>) from the Early Permian of the Czech Republic, representative of the morphology of early beetles</figcaption></figure> The oldest known beetle is <a href="/wiki/Coleopsis" title="Coleopsis">Coleopsis</a>, from the earliest Permian (<a href="/wiki/Asselian" title="Asselian">Asselian</a>) of Germany, around 295 million years ago.<a href="#cite_note-26">[26]</a> Early beetles from the Permian, which are collectively grouped into the "<a href="/wiki/Protocoleoptera" title="Protocoleoptera">Protocoleoptera</a>" are thought to have been <a href="/wiki/Xylophagous" class="mw-redirect" title="Xylophagous">xylophagous</a> (wood eating) and <a href="/wiki/Woodboring_beetle" title="Woodboring beetle">wood boring</a>. Fossils from this time have been found in Siberia and Europe, for instance in the red slate fossil beds of Niedermoschel near Mainz, Germany.<a href="#cite_note-27">[27]</a> Further fossils have been found in Obora, Czech Republic and Tshekarda in the Ural mountains, Russia.<a href="#cite_note-28">[28]</a> However, there are only a few fossils from North America before the <a href="/wiki/Middle_Permian" class="mw-redirect" title="Middle Permian">middle Permian</a>, although both Asia and North America had been united to <a href="/wiki/Euramerica" title="Euramerica">Euramerica</a>. The first discoveries from North America made in the <a href="/wiki/Wellington_Formation" title="Wellington Formation">Wellington Formation</a> of Oklahoma were published in 2005 and 2008.<a href="#cite_note-evo-29">[29]</a><a href="#cite_note-30">[30]</a> The earliest members of modern beetle lineages appeared during the <a href="/wiki/Late_Permian" class="mw-redirect" title="Late Permian">Late Permian</a>. In the <a href="/wiki/Permian%E2%80%93Triassic_extinction_event" title="Permian–Triassic extinction event">Permian–Triassic extinction event</a> at the end of the Permian, most "protocoleopteran" lineages became extinct. Beetle diversity did not recover to pre-extinction levels until the <a href="/wiki/Middle_Triassic" title="Middle Triassic">Middle Triassic</a>.<a href="#cite_note-:1-31">[31]</a> <div class="mw-heading mw-heading3"><h3 id="Jurassic">Jurassic</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=5" title="Edit section: Jurassic">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Phylogenetic_chart_of_Coleoptera.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Phylogenetic_chart_of_Coleoptera.svg/220px-Phylogenetic_chart_of_Coleoptera.svg.png" decoding="async" width="220" height="258" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Phylogenetic_chart_of_Coleoptera.svg/330px-Phylogenetic_chart_of_Coleoptera.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b5/Phylogenetic_chart_of_Coleoptera.svg/440px-Phylogenetic_chart_of_Coleoptera.svg.png 2x" data-file-width="2989" data-file-height="3505" /></a><figcaption>Beetle <a href="/wiki/Genus" title="Genus">genera</a> were mainly saprophages (<a href="/wiki/Detritivore" title="Detritivore">detritivores</a>) in the <a href="/wiki/Permian" title="Permian">Permian</a> and <a href="/wiki/Triassic" title="Triassic">Triassic</a>. During the <a href="/wiki/Jurassic" title="Jurassic">Jurassic</a>, <a href="/wiki/Herbivorous" class="mw-redirect" title="Herbivorous">herbivorous</a> and then <a href="/wiki/Carnivorous" class="mw-redirect" title="Carnivorous">carnivorous</a> genera became more common. In the <a href="/wiki/Cenozoic" title="Cenozoic">Cenozoic</a>, genera at all three <a href="/wiki/Trophic_level" title="Trophic level">trophic levels</a> became far more numerous.</figcaption></figure>During the <a href="/wiki/Jurassic" title="Jurassic">Jurassic</a> (<a rel="nofollow" class="external text" href="https://geoltime.github.io/?Ma=210–145">210 to 145</a> mya), there was a dramatic increase in the diversity of beetle families,<a href="#cite_note-evo-29">[29]</a> including the development and growth of carnivorous and herbivorous species. The <a href="/wiki/Chrysomeloidea" title="Chrysomeloidea">Chrysomeloidea</a> diversified around the same time, feeding on a wide array of plant hosts from <a href="/wiki/Cycad" title="Cycad">cycads</a> and <a href="/wiki/Conifer" title="Conifer">conifers</a> to <a href="/wiki/Angiosperm" class="mw-redirect" title="Angiosperm">angiosperms</a>.<a href="#cite_note-insenc186-32">[32]</a> Close to the Upper Jurassic, the Cupedidae decreased, but the diversity of the early plant-eating species increased. Most recent plant-eating beetles feed on flowering plants or angiosperms, whose success contributed to a doubling of plant-eating species during the <a href="/wiki/Middle_Jurassic" title="Middle Jurassic">Middle Jurassic</a>. However, the increase of the number of beetle families during the Cretaceous does not correlate with the increase of the number of angiosperm species.<a href="#cite_note-33">[33]</a> Around the same time, numerous primitive weevils (e.g. <a href="/wiki/Curculionoidea" class="mw-redirect" title="Curculionoidea">Curculionoidea</a>) and click beetles (e.g. <a href="/wiki/Elateroidea" title="Elateroidea">Elateroidea</a>) appeared. The first jewel beetles (e.g. <a href="/wiki/Buprestidae" title="Buprestidae">Buprestidae</a>) are present, but they remained rare until the Cretaceous.<a href="#cite_note-34">[34]</a><a href="#cite_note-35">[35]</a><a href="#cite_note-36">[36]</a> The first scarab beetles were not coprophagous but presumably fed on rotting wood with the help of fungus; they are an early example of a <a href="#Mutualism">mutualistic</a> relationship. There are more than 150 important fossil sites from the Jurassic, the majority in Eastern Europe and North Asia. Outstanding sites include <a href="/wiki/Solnhofen" title="Solnhofen">Solnhofen</a> in Upper <a href="/wiki/Bavaria" title="Bavaria">Bavaria</a>, Germany,<a href="#cite_note-37">[37]</a> Karatau in South <a href="/wiki/Kazakhstan" title="Kazakhstan">Kazakhstan</a>,<a href="#cite_note-38">[38]</a> the Yixian formation in <a href="/wiki/Liaoning" title="Liaoning">Liaoning</a>, North China,<a href="#cite_note-liaoning-39">[39]</a> as well as the Jiulongshan formation and further fossil sites in <a href="/wiki/Mongolia" title="Mongolia">Mongolia</a>. In North America there are only a few sites with fossil records of insects from the Jurassic, namely the shell limestone deposits in the Hartford basin, the Deerfield basin and the Newark basin.<a href="#cite_note-evo-29">[29]</a><a href="#cite_note-40">[40]</a> <div class="mw-heading mw-heading3"><h3 id="Cretaceous">Cretaceous</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=6" title="Edit section: Cretaceous">edit</a>]</div> The <a href="/wiki/Cretaceous" title="Cretaceous">Cretaceous</a> saw the fragmenting of the southern landmass, with the opening of the southern Atlantic Ocean and the isolation of New Zealand, while South America, Antarctica, and Australia grew more distant.<a href="#cite_note-insenc186-32">[32]</a> The diversity of Cupedidae and <a href="/wiki/Archostemata" title="Archostemata">Archostemata</a> decreased considerably. Predatory ground beetles (Carabidae) and rove beetles (Staphylinidae) began to distribute into different patterns; the <a href="/wiki/Carabidae" class="mw-redirect" title="Carabidae">Carabidae</a> predominantly occurred in the warm regions, while the <a href="/wiki/Staphylinidae" class="mw-redirect" title="Staphylinidae">Staphylinidae</a> and <a href="/wiki/Click_beetle" title="Click beetle">click beetles</a> (Elateridae) preferred temperate climates. Likewise, predatory species of <a href="/wiki/Cleroidea" title="Cleroidea">Cleroidea</a> and <a href="/wiki/Cucujoidea" title="Cucujoidea">Cucujoidea</a> hunted their prey under the bark of trees together with the <a href="/wiki/Jewel_beetle" class="mw-redirect" title="Jewel beetle">jewel beetles</a> (Buprestidae). The diversity of jewel beetles increased rapidly, as they were the primary consumers of wood,<a href="#cite_note-41">[41]</a> while <a href="/wiki/Longhorn_beetle" title="Longhorn beetle">longhorn beetles</a> (<a href="/wiki/Cerambycidae" class="mw-redirect" title="Cerambycidae">Cerambycidae</a>) were rather rare: their diversity increased only towards the end of the Upper Cretaceous.<a href="#cite_note-evo-29">[29]</a> The first coprophagous beetles are from the Upper Cretaceous<a href="#cite_note-42">[42]</a> and may have lived on the excrement of herbivorous dinosaurs.<a href="#cite_note-43">[43]</a> The first species where both larvae and adults are adapted to an aquatic lifestyle are found. Whirligig beetles (Gyrinidae) were moderately diverse, although other early beetles (e.g. Dytiscidae) were less, with the most widespread being the species of <a href="/wiki/Coptoclavidae" title="Coptoclavidae">Coptoclavidae</a>, which preyed on aquatic fly larvae.<a href="#cite_note-evo-29">[29]</a> A 2020 review of the palaeoecological interpretations of fossil beetles from Cretaceous ambers has suggested that <a href="/wiki/Saproxylic" class="mw-redirect" title="Saproxylic">saproxylicity</a> was the most common feeding strategy, with <a href="/wiki/Fungivorous" class="mw-redirect" title="Fungivorous">fungivorous</a> species in particular appearing to dominate.<a href="#cite_note-amber-44">[44]</a> Many fossil sites worldwide contain beetles from the Cretaceous. Most are in Europe and Asia and belong to the temperate climate zone during the Cretaceous.<a href="#cite_note-liaoning-39">[39]</a> Lower Cretaceous sites include the Crato fossil beds in the Araripe basin in the <a href="/wiki/Cear%C3%A1" title="Ceará">Ceará</a>, North Brazil, as well as overlying Santana formation; the latter was near the equator at that time. In Spain, important sites are near <a href="/wiki/Montsec" class="mw-redirect" title="Montsec">Montsec</a> and <a href="/wiki/Las_Hoyas" class="mw-redirect" title="Las Hoyas">Las Hoyas</a>. In Australia, the Koonwarra fossil beds of the Korumburra group, <a href="/wiki/South_Gippsland" title="South Gippsland">South Gippsland</a>, Victoria, are noteworthy. Major sites from the Upper Cretaceous include Kzyl-Dzhar in South Kazakhstan and Arkagala in Russia.<a href="#cite_note-evo-29">[29]</a> <div class="mw-heading mw-heading3"><h3 id="Cenozoic">Cenozoic</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=7" title="Edit section: Cenozoic">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Prachtk%C3%A4fer_aus_der_Grube_Messel.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Prachtk%C3%A4fer_aus_der_Grube_Messel.JPG/220px-Prachtk%C3%A4fer_aus_der_Grube_Messel.JPG" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Prachtk%C3%A4fer_aus_der_Grube_Messel.JPG/330px-Prachtk%C3%A4fer_aus_der_Grube_Messel.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0d/Prachtk%C3%A4fer_aus_der_Grube_Messel.JPG/440px-Prachtk%C3%A4fer_aus_der_Grube_Messel.JPG 2x" data-file-width="2480" data-file-height="2480" /></a><figcaption>Fossil <a href="/wiki/Buprestidae" title="Buprestidae">buprestid beetle</a> from the <a href="/wiki/Eocene" title="Eocene">Eocene</a> (50 mya) <a href="/wiki/Messel_pit" class="mw-redirect" title="Messel pit">Messel pit</a>, which retains its <a href="/wiki/Structural_color" class="mw-redirect" title="Structural color">structural color</a><a href="#cite_note-45">[45]</a></figcaption></figure> Beetle fossils are abundant in the Cenozoic; by the <a href="/wiki/Quaternary" title="Quaternary">Quaternary</a> (up to 1.6 mya), fossil species are identical to living ones, while from the <a href="/wiki/Late_Miocene" title="Late Miocene">Late Miocene</a> (5.7 mya) the fossils are still so close to modern forms that they are most likely the ancestors of living species. The <a href="/wiki/Quaternary_glaciation" title="Quaternary glaciation">large oscillations in climate</a> during the Quaternary caused beetles to change their geographic distributions so much that current location gives little clue to the biogeographical history of a species. It is evident that geographic isolation of populations must often have been broken as insects moved under the influence of changing climate, causing mixing of gene pools, rapid evolution, and extinctions, especially in middle latitudes.<a href="#cite_note-Coope-46">[46]</a> <div class="mw-heading mw-heading2"><h2 id="Phylogeny">Phylogeny</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=8" title="Edit section: Phylogeny">edit</a>]</div> The very large number of beetle species poses special problems for <a href="/wiki/Linnaean_taxonomy" title="Linnaean taxonomy">classification</a>. Some families contain tens of thousands of species, and need to be divided into subfamilies and tribes. <a href="/wiki/Polyphaga" title="Polyphaga">Polyphaga</a> is the largest suborder, containing more than 300,000 described species in more than 170 families, including <a href="/wiki/Rove_beetle" title="Rove beetle">rove beetles</a> (Staphylinidae), scarab beetles (<a href="/wiki/Scarabaeidae" title="Scarabaeidae">Scarabaeidae</a>), <a href="/wiki/Blister_beetle" title="Blister beetle">blister beetles</a> (Meloidae), <a href="/wiki/Stag_beetle" title="Stag beetle">stag beetles</a> (Lucanidae) and true weevils (<a href="/wiki/Curculionidae" title="Curculionidae">Curculionidae</a>).<a href="#cite_note-Hunt_et_al_2007-10">[10]</a><a href="#cite_note-TOLweb_1995-47">[47]</a> These polyphagan beetle groups can be identified by the presence of cervical <a href="/wiki/Sclerite" title="Sclerite">sclerites</a> (hardened parts of the head used as points of attachment for muscles) absent in the other suborders.<a href="#cite_note-48">[48]</a> <a href="/wiki/Adephaga" title="Adephaga">Adephaga</a> contains about 10 families of largely predatory beetles, includes <a href="/wiki/Ground_beetle" title="Ground beetle">ground beetles</a> (Carabidae), water beetles (<a href="/wiki/Dytiscidae" title="Dytiscidae">Dytiscidae</a>) and <a href="/wiki/Whirligig_beetle" title="Whirligig beetle">whirligig beetles</a> (Gyrinidae). In these insects, the <a href="/wiki/Testes" class="mw-redirect" title="Testes">testes</a> are tubular and the first abdominal sternum (a plate of the <a href="/wiki/Exoskeleton" title="Exoskeleton">exoskeleton</a>) is divided by the hind <a href="/wiki/Arthropod_leg" title="Arthropod leg">coxae</a> (the basal joints of the beetle's legs).<a href="#cite_note-49">[49]</a> <a href="/wiki/Archostemata" title="Archostemata">Archostemata</a> contains four families of mainly wood-eating beetles, including <a href="/wiki/Reticulated_beetle" class="mw-redirect" title="Reticulated beetle">reticulated beetles</a> (Cupedidae) and the <a href="/wiki/Telephone-pole_beetle" title="Telephone-pole beetle">telephone-pole beetle</a>.<a href="#cite_note-50">[50]</a> The Archostemata have an exposed plate called the metatrochantin in front of the basal segment or coxa of the hind leg.<a href="#cite_note-ausbeetle-51">[51]</a> <a href="/wiki/Myxophaga" title="Myxophaga">Myxophaga</a> contains about 65 described species in four families, mostly very small, including <a href="/wiki/Hydroscaphidae" title="Hydroscaphidae">Hydroscaphidae</a> and the genus <a href="/wiki/Sphaerius" title="Sphaerius">Sphaerius</a>.<a href="#cite_note-52">[52]</a> The myxophagan beetles are small and mostly alga-feeders. Their mouthparts are characteristic in lacking galeae and having a mobile tooth on their left mandible.<a href="#cite_note-53">[53]</a> The consistency of beetle <a href="/wiki/Morphology_(biology)" title="Morphology (biology)">morphology</a>, in particular their possession of <a href="/wiki/Elytra" class="mw-redirect" title="Elytra">elytra</a>, has long suggested that Coleoptera is <a href="/wiki/Monophyletic" class="mw-redirect" title="Monophyletic">monophyletic</a>, though there have been doubts about the arrangement of the <a href="/wiki/Taxonomic_rank" title="Taxonomic rank">suborders</a>, namely the <a href="/wiki/Adephaga" title="Adephaga">Adephaga</a>, <a href="/wiki/Archostemata" title="Archostemata">Archostemata</a>, <a href="/wiki/Myxophaga" title="Myxophaga">Myxophaga</a> and <a href="/wiki/Polyphaga" title="Polyphaga">Polyphaga</a> within that <a href="/wiki/Clade" title="Clade">clade</a>.<a href="#cite_note-whiting-54">[54]</a><a href="#cite_note-insenc186-32">[32]</a><a href="#cite_note-Beutel-55">[55]</a><a href="#cite_note-Kukalová-56">[56]</a><a href="#cite_note-57">[57]</a> The twisted-wing parasites, <a href="/wiki/Strepsiptera" title="Strepsiptera">Strepsiptera</a>, are thought to be a sister group to the beetles, having split from them in the <a href="/wiki/Early_Permian" class="mw-redirect" title="Early Permian">Early Permian</a>.<a href="#cite_note-Kukalová-56">[56]</a><a href="#cite_note-58">[58]</a><a href="#cite_note-59">[59]</a><a href="#cite_note-McKenna2015-60">[60]</a> Molecular phylogenetic analysis confirms that the Coleoptera are monophyletic. Duane McKenna et al. (2015) used eight nuclear genes for 367 species from 172 of 183 Coleopteran families. They split the Adephaga into 2 clades, Hydradephaga and Geadephaga, broke up the Cucujoidea into 3 clades, and placed the Lymexyloidea within the Tenebrionoidea. The Polyphaga appear to date from the Triassic. Most extant beetle families appear to have arisen in the Cretaceous.<a href="#cite_note-McKenna2015-60">[60]</a> The <a href="/wiki/Cladogram" title="Cladogram">cladogram</a> is based on McKenna (2015).<a href="#cite_note-McKenna2015-60">[60]</a> The number of species in each group (mainly superfamilies) is shown in parentheses, and boldface if over 10,000.<a href="#cite_note-Hunt2007-61">[61]</a> English common names are given where possible. Dates of origin of major groups are shown in italics in millions of years ago (mya).<a href="#cite_note-Hunt2007-61">[61]</a> <div class="clade"><style data-mw-deduplicate="TemplateStyles:r1258728058">body.skin-vector-2022 .mw-parser-output div.clade,body.skin-minerva .mw-parser-output div.clade{overflow-x:auto;overflow-y:hidden}body.skin-minerva .mw-parser-output div.clade p{font-size:inherit}.mw-parser-output table.clade{border-spacing:0;margin:0;font-size:100%;line-height:100%;border-collapse:separate;width:auto}.mw-parser-output table.clade table.clade{width:100%;line-height:inherit}.mw-parser-output table.clade td.clade-label{min-width:0.2em;width:0.2em;padding:0.1em 0.25em;vertical-align:bottom;text-align:center;border-left:1px solid;border-bottom:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-label::before,.mw-parser-output table.clade td.clade-slabel::before{content:"\2060 "}.mw-parser-output table.clade td.clade-fixed-width{overflow:hidden;text-overflow:ellipsis}.mw-parser-output table.clade td.clade-fixed-width:hover{overflow:visible}.mw-parser-output table.clade td.clade-label.first{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-label.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-slabel{padding:0.1em 0.25em;vertical-align:top;text-align:center;border-left:1px solid;white-space:nowrap}.mw-parser-output table.clade td.clade-slabel:hover{overflow:visible}.mw-parser-output table.clade td.clade-slabel.last{border-left:none;border-right:none}.mw-parser-output table.clade td.clade-slabel.reverse{border-left:none;border-right:1px solid}.mw-parser-output table.clade td.clade-bar{vertical-align:middle;text-align:left;padding:0 0.5em;position:relative}.mw-parser-output table.clade td.clade-bar.reverse{text-align:right;position:relative}.mw-parser-output table.clade td.clade-leaf{border:0;padding:0;text-align:left}.mw-parser-output table.clade td.clade-leafR{border:0;padding:0;text-align:right}.mw-parser-output table.clade td.clade-leaf.reverse{text-align:right}.mw-parser-output table.clade:hover span.linkA{background-color:yellow}.mw-parser-output table.clade:hover span.linkB{background-color:green}</style> <table class="clade" style="font-size:85%;line-height:85%"> <tbody><tr> <td class="clade-label first">Coleoptera </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first">240 mya </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Archostemata" title="Archostemata">Archostemata</a> 160 mya (40) <a href="/wiki/File:Archostemata_Tenomerga_mucida01_rotated.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Archostemata_Tenomerga_mucida01_rotated.jpg/80px-Archostemata_Tenomerga_mucida01_rotated.jpg" decoding="async" width="80" height="18" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Archostemata_Tenomerga_mucida01_rotated.jpg/120px-Archostemata_Tenomerga_mucida01_rotated.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Archostemata_Tenomerga_mucida01_rotated.jpg/160px-Archostemata_Tenomerga_mucida01_rotated.jpg 2x" data-file-width="784" data-file-height="177" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Myxophaga" title="Myxophaga">Myxophaga</a> 220 mya (94) <a href="/wiki/File:Sphaerius.acaroides.Reitter.tafel64_(rotated).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Sphaerius.acaroides.Reitter.tafel64_%28rotated%29.jpg/50px-Sphaerius.acaroides.Reitter.tafel64_%28rotated%29.jpg" decoding="async" width="50" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Sphaerius.acaroides.Reitter.tafel64_%28rotated%29.jpg/75px-Sphaerius.acaroides.Reitter.tafel64_%28rotated%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Sphaerius.acaroides.Reitter.tafel64_%28rotated%29.jpg/100px-Sphaerius.acaroides.Reitter.tafel64_%28rotated%29.jpg 2x" data-file-width="324" data-file-height="262" /></a> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Adephaga" title="Adephaga">Adephaga</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Hydradephaga" class="mw-redirect" title="Hydradephaga">Hydradephaga</a> (5,560) e.g. <a href="/wiki/Dytiscidae" title="Dytiscidae">Dytiscidae</a> (diving beetles) <a href="/wiki/File:Dytiscus_marginalis_Linn%C3%A9,_1758_female.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/31/Dytiscus_marginalis_Linn%C3%A9%2C_1758_female.jpg/70px-Dytiscus_marginalis_Linn%C3%A9%2C_1758_female.jpg" decoding="async" width="70" height="45" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/31/Dytiscus_marginalis_Linn%C3%A9%2C_1758_female.jpg/105px-Dytiscus_marginalis_Linn%C3%A9%2C_1758_female.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/31/Dytiscus_marginalis_Linn%C3%A9%2C_1758_female.jpg/140px-Dytiscus_marginalis_Linn%C3%A9%2C_1758_female.jpg 2x" data-file-width="2633" data-file-height="1676" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Geadephaga" class="mw-redirect" title="Geadephaga">Geadephaga</a> (35,000) e.g. <a href="/wiki/Carabidae" class="mw-redirect" title="Carabidae">Carabidae</a> (ground beetles) <a href="/wiki/File:Cicindela_sylvicola01.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Cicindela_sylvicola01.jpg/70px-Cicindela_sylvicola01.jpg" decoding="async" width="70" height="54" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/83/Cicindela_sylvicola01.jpg/105px-Cicindela_sylvicola01.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/83/Cicindela_sylvicola01.jpg/140px-Cicindela_sylvicola01.jpg 2x" data-file-width="2427" data-file-height="1859" /></a> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last">200 mya </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Polyphaga" title="Polyphaga">Polyphaga</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Scirtoidea" title="Scirtoidea">Scirtoidea</a> (800) + <a href="/wiki/Derodontoidea" class="mw-redirect" title="Derodontoidea">Derodontoidea</a> (29) 200 mya <a href="/wiki/File:Clambus_punctulum_(Beck,_1817)_(12924039694).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Clambus_punctulum_%28Beck%2C_1817%29_%2812924039694%29.png/60px-Clambus_punctulum_%28Beck%2C_1817%29_%2812924039694%29.png" decoding="async" width="60" height="46" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/49/Clambus_punctulum_%28Beck%2C_1817%29_%2812924039694%29.png/90px-Clambus_punctulum_%28Beck%2C_1817%29_%2812924039694%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/49/Clambus_punctulum_%28Beck%2C_1817%29_%2812924039694%29.png/120px-Clambus_punctulum_%28Beck%2C_1817%29_%2812924039694%29.png 2x" data-file-width="1678" data-file-height="1278" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Staphylinidae" class="mw-redirect" title="Staphylinidae">Staphylinidae</a> 195 mya (48,000, rove beetles) <a href="/wiki/File:Cordalia_tsavoana_Pace,_2008_(2912937616).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Cordalia_tsavoana_Pace%2C_2008_%282912937616%29.jpg/70px-Cordalia_tsavoana_Pace%2C_2008_%282912937616%29.jpg" decoding="async" width="70" height="39" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Cordalia_tsavoana_Pace%2C_2008_%282912937616%29.jpg/105px-Cordalia_tsavoana_Pace%2C_2008_%282912937616%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e0/Cordalia_tsavoana_Pace%2C_2008_%282912937616%29.jpg/140px-Cordalia_tsavoana_Pace%2C_2008_%282912937616%29.jpg 2x" data-file-width="2174" data-file-height="1198" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Scarabaeoidea" title="Scarabaeoidea">Scarabaeoidea</a> 145 mya (35,000, scarabs, stag beetles, etc.) <a href="/wiki/File:Mimela_splendens_left_facing.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Mimela_splendens_left_facing.jpg/70px-Mimela_splendens_left_facing.jpg" decoding="async" width="70" height="54" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Mimela_splendens_left_facing.jpg/105px-Mimela_splendens_left_facing.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f4/Mimela_splendens_left_facing.jpg/140px-Mimela_splendens_left_facing.jpg 2x" data-file-width="3559" data-file-height="2747" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Hydrophiloidea" title="Hydrophiloidea">Hydrophiloidea</a> (2,800, water scavenger beetles) <a href="/wiki/File:Hydrophilus_piceus_(Linn%C3%A9,_1758)_female_(4035156238).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Hydrophilus_piceus_%28Linn%C3%A9%2C_1758%29_female_%284035156238%29.jpg/70px-Hydrophilus_piceus_%28Linn%C3%A9%2C_1758%29_female_%284035156238%29.jpg" decoding="async" width="70" height="41" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Hydrophilus_piceus_%28Linn%C3%A9%2C_1758%29_female_%284035156238%29.jpg/105px-Hydrophilus_piceus_%28Linn%C3%A9%2C_1758%29_female_%284035156238%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a0/Hydrophilus_piceus_%28Linn%C3%A9%2C_1758%29_female_%284035156238%29.jpg/140px-Hydrophilus_piceus_%28Linn%C3%A9%2C_1758%29_female_%284035156238%29.jpg 2x" data-file-width="5676" data-file-height="3359" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Histeroidea" title="Histeroidea">Histeroidea</a> (3,800, clown beetles) <a href="/wiki/File:Hister_quadrimaculatus_MHNT_Fronton_Blanc.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Hister_quadrimaculatus_MHNT_Fronton_Blanc.jpg/60px-Hister_quadrimaculatus_MHNT_Fronton_Blanc.jpg" decoding="async" width="60" height="60" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Hister_quadrimaculatus_MHNT_Fronton_Blanc.jpg/90px-Hister_quadrimaculatus_MHNT_Fronton_Blanc.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e4/Hister_quadrimaculatus_MHNT_Fronton_Blanc.jpg/120px-Hister_quadrimaculatus_MHNT_Fronton_Blanc.jpg 2x" data-file-width="3191" data-file-height="3209" /></a> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"><a href="/wiki/Elateriformia" title="Elateriformia">Elateriformia</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Nosodendridae" title="Nosodendridae">Nosodendridae</a> (70) </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Dascilloidea" title="Dascilloidea">Dascilloidea</a> (180) <a href="/wiki/File:Dascillus_cervinus_(Linn%C3%A9,_1758)_(5598492362).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Dascillus_cervinus_%28Linn%C3%A9%2C_1758%29_%285598492362%29.jpg/60px-Dascillus_cervinus_%28Linn%C3%A9%2C_1758%29_%285598492362%29.jpg" decoding="async" width="60" height="42" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/37/Dascillus_cervinus_%28Linn%C3%A9%2C_1758%29_%285598492362%29.jpg/90px-Dascillus_cervinus_%28Linn%C3%A9%2C_1758%29_%285598492362%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/37/Dascillus_cervinus_%28Linn%C3%A9%2C_1758%29_%285598492362%29.jpg/120px-Dascillus_cervinus_%28Linn%C3%A9%2C_1758%29_%285598492362%29.jpg 2x" data-file-width="5231" data-file-height="3681" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Buprestoidea" title="Buprestoidea">Buprestoidea</a> (14,000, jewel beetles) <a href="/wiki/File:Buprestis_octoguttata_side.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/4a/Buprestis_octoguttata_side.JPG/70px-Buprestis_octoguttata_side.JPG" decoding="async" width="70" height="32" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/4a/Buprestis_octoguttata_side.JPG/105px-Buprestis_octoguttata_side.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/4a/Buprestis_octoguttata_side.JPG/140px-Buprestis_octoguttata_side.JPG 2x" data-file-width="1268" data-file-height="572" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Byrrhoidea" title="Byrrhoidea">Byrrhoidea</a> (400, pill and turtle beetles, etc.) <a href="/wiki/File:Chaetophora.spinosa.-.calwer.16.17.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Chaetophora.spinosa.-.calwer.16.17.jpg/50px-Chaetophora.spinosa.-.calwer.16.17.jpg" decoding="async" width="50" height="47" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Chaetophora.spinosa.-.calwer.16.17.jpg/75px-Chaetophora.spinosa.-.calwer.16.17.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Chaetophora.spinosa.-.calwer.16.17.jpg/100px-Chaetophora.spinosa.-.calwer.16.17.jpg 2x" data-file-width="207" data-file-height="195" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Elateroidea" title="Elateroidea">Elateroidea</a> (23,000, click and soldier beetles, fireflies) <a href="/wiki/File:Agriotes_lineatus_bl2.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Agriotes_lineatus_bl2.JPG/70px-Agriotes_lineatus_bl2.JPG" decoding="async" width="70" height="48" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Agriotes_lineatus_bl2.JPG/105px-Agriotes_lineatus_bl2.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/ef/Agriotes_lineatus_bl2.JPG/140px-Agriotes_lineatus_bl2.JPG 2x" data-file-width="836" data-file-height="568" /></a> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel">190 mya </td></tr> <tr> <td class="clade-label">190 mya </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Bostrichoidea" title="Bostrichoidea">Bostrichoidea</a> (3150, deathwatch, powderpost and skin beetles) <a href="/wiki/File:Anthrenus_verbasci_MHNT_Fronton_Side_view.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/78/Anthrenus_verbasci_MHNT_Fronton_Side_view.jpg/60px-Anthrenus_verbasci_MHNT_Fronton_Side_view.jpg" decoding="async" width="60" height="41" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/78/Anthrenus_verbasci_MHNT_Fronton_Side_view.jpg/90px-Anthrenus_verbasci_MHNT_Fronton_Side_view.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/78/Anthrenus_verbasci_MHNT_Fronton_Side_view.jpg/120px-Anthrenus_verbasci_MHNT_Fronton_Side_view.jpg 2x" data-file-width="4271" data-file-height="2946" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Cucujiformia" title="Cucujiformia">Cucujiformia</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Coccinellidae" title="Coccinellidae">Coccinelloidea</a> (6,000, ladybirds or lady beetles) <a href="/wiki/File:Coccinella.quinquepunctata.jacobs24.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/87/Coccinella.quinquepunctata.jacobs24.jpg/50px-Coccinella.quinquepunctata.jacobs24.jpg" decoding="async" width="50" height="60" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/87/Coccinella.quinquepunctata.jacobs24.jpg/75px-Coccinella.quinquepunctata.jacobs24.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/87/Coccinella.quinquepunctata.jacobs24.jpg/100px-Coccinella.quinquepunctata.jacobs24.jpg 2x" data-file-width="345" data-file-height="416" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Tenebrionoidea" title="Tenebrionoidea">Tenebrionoidea</a> 180 mya (35,000, leaf/flower beetles, etc.) and <a href="/wiki/Lymexyloidea" class="mw-redirect" title="Lymexyloidea">Lymexyloidea</a> <a href="/wiki/File:Tribolium.castaneum.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fe/Tribolium.castaneum.jpg/70px-Tribolium.castaneum.jpg" decoding="async" width="70" height="53" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fe/Tribolium.castaneum.jpg/105px-Tribolium.castaneum.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fe/Tribolium.castaneum.jpg/140px-Tribolium.castaneum.jpg 2x" data-file-width="661" data-file-height="496" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Cleroidea" title="Cleroidea">Cleroidea</a> (9,900, checkered beetles and allies) <a href="/wiki/File:Trichodes_ornatus,_U,_back,_Fossil_Butte_NM,_Wyoming_2012-10-16-15.20_(8095655020).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Trichodes_ornatus%2C_U%2C_back%2C_Fossil_Butte_NM%2C_Wyoming_2012-10-16-15.20_%288095655020%29.jpg/70px-Trichodes_ornatus%2C_U%2C_back%2C_Fossil_Butte_NM%2C_Wyoming_2012-10-16-15.20_%288095655020%29.jpg" decoding="async" width="70" height="47" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Trichodes_ornatus%2C_U%2C_back%2C_Fossil_Butte_NM%2C_Wyoming_2012-10-16-15.20_%288095655020%29.jpg/105px-Trichodes_ornatus%2C_U%2C_back%2C_Fossil_Butte_NM%2C_Wyoming_2012-10-16-15.20_%288095655020%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/0e/Trichodes_ornatus%2C_U%2C_back%2C_Fossil_Butte_NM%2C_Wyoming_2012-10-16-15.20_%288095655020%29.jpg/140px-Trichodes_ornatus%2C_U%2C_back%2C_Fossil_Butte_NM%2C_Wyoming_2012-10-16-15.20_%288095655020%29.jpg 2x" data-file-width="5616" data-file-height="3744" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Cucujoidea" title="Cucujoidea">Cucujoidea</a> (8,000) <a href="/wiki/File:Cucujus_cinnaberinus_habitus_rotated.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Cucujus_cinnaberinus_habitus_rotated.jpg/70px-Cucujus_cinnaberinus_habitus_rotated.jpg" decoding="async" width="70" height="36" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Cucujus_cinnaberinus_habitus_rotated.jpg/105px-Cucujus_cinnaberinus_habitus_rotated.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/b/b9/Cucujus_cinnaberinus_habitus_rotated.jpg/140px-Cucujus_cinnaberinus_habitus_rotated.jpg 2x" data-file-width="942" data-file-height="491" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"><a href="/wiki/Phytophaga" title="Phytophaga">Phytophaga</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"><a href="/wiki/Chrysomeloidea" title="Chrysomeloidea">Chrysomeloidea</a> </td> <td rowspan="2" class="clade-leaf"> <div><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1258728058"> <table class="clade"> <tbody><tr> <td class="clade-label first"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Chrysomelidae" class="mw-redirect" title="Chrysomelidae">Chrysomelidae</a> (35,000, leaf beetles) <a href="/wiki/File:Mesoplatys_cincta_(Olivier,_1790)_(8245535128).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Mesoplatys_cincta_%28Olivier%2C_1790%29_%288245535128%29.png/70px-Mesoplatys_cincta_%28Olivier%2C_1790%29_%288245535128%29.png" decoding="async" width="70" height="49" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Mesoplatys_cincta_%28Olivier%2C_1790%29_%288245535128%29.png/105px-Mesoplatys_cincta_%28Olivier%2C_1790%29_%288245535128%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6e/Mesoplatys_cincta_%28Olivier%2C_1790%29_%288245535128%29.png/140px-Mesoplatys_cincta_%28Olivier%2C_1790%29_%288245535128%29.png 2x" data-file-width="5296" data-file-height="3674" /></a> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Cerambycidae" class="mw-redirect" title="Cerambycidae">Cerambycidae</a> (25,000, longhorn beetles) <a href="/wiki/File:Leptura_(syn._Strangalia,_Rutpela)_maculata_(Poda,_1761)_male_(3926136467).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Leptura_%28syn._Strangalia%2C_Rutpela%29_maculata_%28Poda%2C_1761%29_male_%283926136467%29.jpg/70px-Leptura_%28syn._Strangalia%2C_Rutpela%29_maculata_%28Poda%2C_1761%29_male_%283926136467%29.jpg" decoding="async" width="70" height="36" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Leptura_%28syn._Strangalia%2C_Rutpela%29_maculata_%28Poda%2C_1761%29_male_%283926136467%29.jpg/105px-Leptura_%28syn._Strangalia%2C_Rutpela%29_maculata_%28Poda%2C_1761%29_male_%283926136467%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e6/Leptura_%28syn._Strangalia%2C_Rutpela%29_maculata_%28Poda%2C_1761%29_male_%283926136467%29.jpg/140px-Leptura_%28syn._Strangalia%2C_Rutpela%29_maculata_%28Poda%2C_1761%29_male_%283926136467%29.jpg 2x" data-file-width="5267" data-file-height="2728" /></a> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel"> </td></tr> <tr> <td class="clade-label"> </td> <td rowspan="2" class="clade-leaf"> <a href="/wiki/Curculionoidea" class="mw-redirect" title="Curculionoidea">Curculionoidea</a> (97,000, weevils) <a href="/wiki/File:Curculio_glandium_Marsham,_1802_female_(8112399337).png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Curculio_glandium_Marsham%2C_1802_female_%288112399337%29.png/70px-Curculio_glandium_Marsham%2C_1802_female_%288112399337%29.png" decoding="async" width="70" height="34" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Curculio_glandium_Marsham%2C_1802_female_%288112399337%29.png/105px-Curculio_glandium_Marsham%2C_1802_female_%288112399337%29.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5a/Curculio_glandium_Marsham%2C_1802_female_%288112399337%29.png/140px-Curculio_glandium_Marsham%2C_1802_female_%288112399337%29.png 2x" data-file-width="7234" data-file-height="3485" /></a> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last">170 mya </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last"> </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last">225 mya </td></tr></tbody></table></div> </td></tr> <tr> <td class="clade-slabel last">285 mya </td></tr></tbody></table></div> <div class="mw-heading mw-heading2"><h2 id="External_morphology">External morphology</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=9" title="Edit section: External morphology">edit</a>]</div> <figure class="mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Melolontha_melolontha_insect_morphology.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Melolontha_melolontha_insect_morphology.jpg/300px-Melolontha_melolontha_insect_morphology.jpg" decoding="async" width="300" height="199" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Melolontha_melolontha_insect_morphology.jpg/450px-Melolontha_melolontha_insect_morphology.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2f/Melolontha_melolontha_insect_morphology.jpg/600px-Melolontha_melolontha_insect_morphology.jpg 2x" data-file-width="8063" data-file-height="5347" /></a><figcaption>Beetle body structure, using <a href="/wiki/Cockchafer" title="Cockchafer">cockchafer</a>. A: head, B: thorax, C: abdomen. 1: antenna, 2: compound eye, 3: femur, 4: elytron (wing cover), 5: tibia, 6: tarsus, 7: claws, 8: mouthparts, 9: prothorax, 10: mesothorax, 11: metathorax, 12: abdominal sternites, 13: pygidium.</figcaption></figure> Beetles are generally characterized by a particularly hard <a href="/wiki/Exoskeleton" title="Exoskeleton">exoskeleton</a> and hard forewings (<a href="/wiki/Elytra" class="mw-redirect" title="Elytra">elytra</a>) not usable for flying. Almost all beetles have mandibles that move in a horizontal plane. The mouthparts are rarely suctorial, though they are sometimes reduced; the maxillae always bear palps. The antennae usually have 11 or fewer segments, except in some groups like the Cerambycidae (longhorn beetles) and the Rhipiceridae (cicada parasite beetles). The coxae of the legs are usually located recessed within a coxal cavity. The genitalic structures are telescoped into the last abdominal segment in all extant beetles. Beetle larvae can often be confused with those of other holometabolan groups.<a href="#cite_note-ausbeetle-51">[51]</a> The beetle's exoskeleton is made up of numerous plates, called <a href="/wiki/Sclerite" title="Sclerite">sclerites</a>, separated by thin sutures. This design provides armored defenses while maintaining flexibility. The general <a href="/wiki/Anatomy" title="Anatomy">anatomy</a> of a beetle is quite uniform, although specific organs and <a href="/wiki/Appendage" title="Appendage">appendages</a> vary greatly in appearance and function between the many families in the order. Like all insects, beetles' bodies are divided into three sections: the head, the thorax, and the abdomen.<a href="#cite_note-insenc-8">[8]</a> Because there are so many species, identification is quite difficult, and relies on attributes including the shape of the antennae, the <a href="/wiki/Tarsal_formula" title="Tarsal formula">tarsal formulae</a><a href="#cite_note-62">[a]</a> and shapes of these small segments on the legs, the mouthparts, and the ventral plates (sterna, pleura, coxae). In many species accurate identification can only be made by examination of the unique male genitalic structures.<a href="#cite_note-63">[62]</a> <div class="mw-heading mw-heading3"><h3 id="Head">Head</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=10" title="Edit section: Head">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Head_of_the_Weaver_Beetle_(Lamia_textor).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Head_of_the_Weaver_Beetle_%28Lamia_textor%29.jpg/220px-Head_of_the_Weaver_Beetle_%28Lamia_textor%29.jpg" decoding="async" width="220" height="148" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/70/Head_of_the_Weaver_Beetle_%28Lamia_textor%29.jpg/330px-Head_of_the_Weaver_Beetle_%28Lamia_textor%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/70/Head_of_the_Weaver_Beetle_%28Lamia_textor%29.jpg/440px-Head_of_the_Weaver_Beetle_%28Lamia_textor%29.jpg 2x" data-file-width="4048" data-file-height="2729" /></a><figcaption>Front view of the head of <a href="/wiki/Lamia_textor" class="mw-redirect" title="Lamia textor">Lamia textor</a></figcaption></figure> The head, having mouthparts projecting forward or sometimes downturned, is usually heavily <a href="/wiki/Sclerotin" title="Sclerotin">sclerotized</a> and is sometimes very large.<a href="#cite_note-Gilliott-7">[7]</a> The eyes are <a href="/wiki/Compound_eye" title="Compound eye">compound</a> and may display remarkable adaptability, as in the case of the aquatic whirligig beetles (<a href="/wiki/Gyrinidae" class="mw-redirect" title="Gyrinidae">Gyrinidae</a>), where they are split to allow a view both above and below the waterline. A few <a href="/wiki/Longhorn_beetle" title="Longhorn beetle">Longhorn beetles</a> (<a href="/wiki/Cerambycidae" class="mw-redirect" title="Cerambycidae">Cerambycidae</a>) and weevils as well as some fireflies (<a href="/wiki/Rhagophthalmidae" title="Rhagophthalmidae">Rhagophthalmidae</a>) <a href="#cite_note-Lau-64">[63]</a> have divided eyes, while many have eyes that are notched, and a few have <a href="/wiki/Ocelli" class="mw-redirect" title="Ocelli">ocelli</a>, small, simple <a href="/wiki/Eye" title="Eye">eyes</a> usually farther back on the head (on the <a href="/wiki/Vertex_(anatomy)" title="Vertex (anatomy)">vertex</a>); these are more common in larvae than in adults.<a href="#cite_note-BeetlesAustralia-65">[64]</a> The anatomical organization of the compound eyes may be modified and depends on whether a species is primarily crepuscular, or diurnally or nocturnally active.<a href="#cite_note-beetles-66">[65]</a> Ocelli are found in the adult carpet beetle (as a single central ocellus in <a href="/wiki/Dermestidae" title="Dermestidae">Dermestidae</a>), some rove beetles (<a href="/wiki/Omaliinae" title="Omaliinae">Omaliinae</a>), and the <a href="/wiki/Derodontidae" title="Derodontidae">Derodontidae</a>.<a href="#cite_note-BeetlesAustralia-65">[64]</a> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Polyphylla_fullo_up.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/13/Polyphylla_fullo_up.jpg/220px-Polyphylla_fullo_up.jpg" decoding="async" width="220" height="189" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/13/Polyphylla_fullo_up.jpg/330px-Polyphylla_fullo_up.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/13/Polyphylla_fullo_up.jpg/440px-Polyphylla_fullo_up.jpg 2x" data-file-width="2722" data-file-height="2344" /></a><figcaption><a href="/wiki/Polyphylla_fullo" title="Polyphylla fullo">Polyphylla fullo</a> has distinctive fan-like <a href="/wiki/Antenna_(biology)" title="Antenna (biology)">antennae</a>, one of several distinct forms for the appendages among beetles.</figcaption></figure> Beetle <a href="/wiki/Antenna_(biology)" title="Antenna (biology)">antennae</a> are primarily organs of sensory perception and can detect motion, odor and chemical substances,<a href="#cite_note-Antennae-67">[66]</a> but may also be used to physically feel a beetle's environment. Beetle families may use antennae in different ways. For example, when moving quickly, tiger beetles may not be able to see very well and instead hold their antennae rigidly in front of them in order to avoid obstacles.<a href="#cite_note-68">[67]</a> Certain Cerambycidae use antennae to balance, and blister beetles may use them for grasping. Some aquatic beetle species may use antennae for gathering air and passing it under the body whilst submerged. Equally, some families use antennae during mating, and a few species use them for defense. In the cerambycid <a href="/wiki/Onychocerus_albitarsis" title="Onychocerus albitarsis">Onychocerus albitarsis</a>, the antennae have venom injecting structures used in defense, which is unique among <a href="/wiki/Arthropod" title="Arthropod">arthropods</a>.<a href="#cite_note-69">[68]</a> Antennae vary greatly in form, sometimes between the sexes, but are often similar within any given family. Antennae may be <a href="https://en.wiktionary.org/wiki/clavate" class="extiw" title="wiktionary:clavate">clubbed</a>, <a href="https://en.wiktionary.org/wiki/filiform" class="extiw" title="wiktionary:filiform">threadlike</a>, <a href="https://en.wiktionary.org/wiki/geniculate" class="extiw" title="wiktionary:geniculate">angled</a>, <a href="https://en.wiktionary.org/wiki/moniliform" class="extiw" title="wiktionary:moniliform">shaped like a string of beads</a>, <a href="https://en.wiktionary.org/wiki/pectinate" class="extiw" title="wiktionary:pectinate">comb-like</a> (either on one side or both, bipectinate), or <a href="https://en.wiktionary.org/wiki/serrate" class="extiw" title="wiktionary:serrate">toothed</a>. The physical variation of antennae is important for the identification of many beetle groups. The Curculionidae have elbowed or geniculate antennae. Feather like flabellate antennae are a restricted form found in the Rhipiceridae and a few other families. The Silphidae have a capitate antennae with a spherical head at the tip. The Scarabaeidae typically have lamellate antennae with the terminal segments extended into long flat structures stacked together. The Carabidae typically have thread-like antennae. The antennae arises between the eye and the mandibles and in the Tenebrionidae, the antennae rise in front of a notch that breaks the usually circular outline of the compound eye. They are segmented and usually consist of 11 parts, the first part is called the scape and the second part is the pedicel. The other segments are jointly called the flagellum.<a href="#cite_note-Antennae-67">[66]</a><a href="#cite_note-70">[69]</a><a href="#cite_note-71">[70]</a> Beetles have <a href="/wiki/Insect_mouthparts" title="Insect mouthparts">mouthparts</a> like those of <a href="/wiki/Grasshopper" title="Grasshopper">grasshoppers</a>. The <a href="/wiki/Mandible_(insect)" class="mw-redirect" title="Mandible (insect)">mandibles</a> appear as large pincers on the front of some beetles. The mandibles are a pair of hard, often tooth-like structures that move horizontally to grasp, crush, or cut food or enemies (see <a href="#Anti-predator_adaptations">defence</a>, below). Two pairs of finger-like appendages, the maxillary and labial palpi, are found around the mouth in most beetles, serving to move food into the mouth. In many species, the mandibles are sexually dimorphic, with those of the males enlarged enormously compared with those of females of the same species.<a href="#cite_note-Gilliott-7">[7]</a> <div class="mw-heading mw-heading3"><h3 id="Thorax">Thorax</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=11" title="Edit section: Thorax">edit</a>]</div> The thorax is <a href="/wiki/Segmentation_(biology)" title="Segmentation (biology)">segmented</a> into the two discernible parts, the pro- and pterothorax. The pterothorax is the fused meso- and metathorax, which are commonly separated in other insect species, although flexibly articulate from the prothorax. When viewed from below, the thorax is that part from which all three pairs of legs and both pairs of wings arise. The abdomen is everything posterior to the thorax.<a href="#cite_note-insenc-8">[8]</a> When viewed from above, most beetles appear to have three clear sections, but this is deceptive: on the beetle's upper surface, the middle section is a hard plate called the <a href="/wiki/Pronotum" class="mw-redirect" title="Pronotum">pronotum</a>, which is only the front part of the thorax; the back part of the thorax is concealed by the beetle's <a href="/wiki/Insect_wing" title="Insect wing">wings</a>. This further segmentation is usually best seen on the abdomen.[<a href="/wiki/Wikipedia:Citation_needed" title="Wikipedia:Citation needed">citation needed</a>] <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Acilius_sulcatus.gemeiner_furchenschwimmer.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/4/43/Acilius_sulcatus.gemeiner_furchenschwimmer.jpg/220px-Acilius_sulcatus.gemeiner_furchenschwimmer.jpg" decoding="async" width="220" height="211" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/4/43/Acilius_sulcatus.gemeiner_furchenschwimmer.jpg/330px-Acilius_sulcatus.gemeiner_furchenschwimmer.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/4/43/Acilius_sulcatus.gemeiner_furchenschwimmer.jpg/440px-Acilius_sulcatus.gemeiner_furchenschwimmer.jpg 2x" data-file-width="780" data-file-height="748" /></a><figcaption><a href="/wiki/Acilius_sulcatus" title="Acilius sulcatus">Acilius sulcatus</a>, a diving beetle with hind legs adapted as <a href="/wiki/Animal_locomotion#Swimming" title="Animal locomotion">swimming</a> limbs</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Legs">Legs</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=12" title="Edit section: Legs">edit</a>]</div> The multisegmented <a href="/wiki/Arthropod_leg" title="Arthropod leg">legs</a> end in two to five small segments called tarsi. Like many other insect orders, beetles have claws, usually one pair, on the end of the last tarsal segment of each leg. While most beetles use their legs for walking, legs have been variously adapted for other uses. Aquatic beetles including the <a href="/wiki/Dytiscidae" title="Dytiscidae">Dytiscidae (diving beetles)</a>, <a href="/wiki/Haliplidae" title="Haliplidae">Haliplidae</a>, and many species of <a href="/wiki/Hydrophilidae" title="Hydrophilidae">Hydrophilidae</a>, the legs, often the last pair, are modified for swimming, typically with rows of long hairs. Male diving beetles have suctorial cups on their forelegs that they use to grasp females.<a href="#cite_note-72">[71]</a> Other beetles have <a href="/wiki/Fossorial" title="Fossorial">fossorial</a> legs widened and often spined for digging. Species with such adaptations are found among the scarabs, ground beetles, and <a href="/wiki/Clown_beetle" class="mw-redirect" title="Clown beetle">clown beetles</a> (<a href="/wiki/Histeridae" title="Histeridae">Histeridae</a>). The hind legs of some beetles, such as <a href="/wiki/Flea_beetle" title="Flea beetle">flea beetles</a> (within Chrysomelidae) and flea weevils (within Curculionidae), have enlarged femurs that help them leap.<a href="#cite_note-73">[72]</a> <div class="mw-heading mw-heading3"><h3 id="Wings">Wings</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=13" title="Edit section: Wings">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Soldier_Beetle_Trichodes_alvearius_taking_off_from_Knapweed_(cropped).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Soldier_Beetle_Trichodes_alvearius_taking_off_from_Knapweed_%28cropped%29.jpg/220px-Soldier_Beetle_Trichodes_alvearius_taking_off_from_Knapweed_%28cropped%29.jpg" decoding="async" width="220" height="221" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/15/Soldier_Beetle_Trichodes_alvearius_taking_off_from_Knapweed_%28cropped%29.jpg/330px-Soldier_Beetle_Trichodes_alvearius_taking_off_from_Knapweed_%28cropped%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/15/Soldier_Beetle_Trichodes_alvearius_taking_off_from_Knapweed_%28cropped%29.jpg/440px-Soldier_Beetle_Trichodes_alvearius_taking_off_from_Knapweed_%28cropped%29.jpg 2x" data-file-width="961" data-file-height="967" /></a><figcaption>Checkered beetle <a href="/wiki/Trichodes_alvearius" title="Trichodes alvearius">Trichodes alvearius</a> taking off, showing the hard elytra (forewings adapted as wing-cases) held stiffly away from the flight wings</figcaption></figure> The forewings of beetles are not used for <a href="/wiki/Insect_flight" title="Insect flight">flight</a>, but form elytra which cover the hind part of the body and protect the hindwings. The elytra are usually hard shell-like structures which must be raised to allow the hindwings to move for flight.<a href="#cite_note-Carpenter-74">[73]</a> However, in the soldier beetles (<a href="/wiki/Cantharidae" class="mw-redirect" title="Cantharidae">Cantharidae</a>), the elytra are soft, earning this family the name of leatherwings.<a href="#cite_note-75">[74]</a> Other soft wing beetles include the <a href="/wiki/Net-winged_beetle" class="mw-redirect" title="Net-winged beetle">net-winged beetle</a> <a href="/wiki/Calopteron_discrepans" title="Calopteron discrepans">Calopteron discrepans</a>, which has brittle wings that rupture easily in order to release chemicals for defense.<a href="#cite_note-76">[75]</a> Beetles' flight wings are crossed with veins and are folded after landing, often along these veins, and stored below the elytra. A fold (jugum) of the membrane at the base of each wing is characteristic.<a href="#cite_note-Carpenter-74">[73]</a> Some beetles have lost the ability to fly. These include some ground beetles (Carabidae) and some true weevils (Curculionidae), as well as desert- and cave-dwelling species of other families. Many have the two elytra fused together, forming a solid shield over the abdomen. In a few families, both the ability to fly and the elytra have been lost, as in the <a href="/wiki/Glow-worm" class="mw-redirect" title="Glow-worm">glow-worms</a> (<a href="/wiki/Phengodidae" title="Phengodidae">Phengodidae</a>), where the females <a href="/wiki/Larviform_female" title="Larviform female">resemble larvae</a> throughout their lives.<a href="#cite_note-77">[76]</a> The presence of elytra and wings does not always indicate that the beetle will fly. For example, the <a href="/wiki/Tansy_beetle" title="Tansy beetle">tansy beetle</a> walks between habitats despite being physically capable of flight.<a href="#cite_note-78">[77]</a> <div class="mw-heading mw-heading3"><h3 id="Abdomen">Abdomen</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=14" title="Edit section: Abdomen">edit</a>]</div> The <a href="/wiki/Abdomen" title="Abdomen">abdomen</a> is the section behind the metathorax, made up of a series of rings, each with a hole for breathing and respiration, called a <a href="/wiki/Spiracle_(arthropods)" title="Spiracle (arthropods)">spiracle</a>, composing three different segmented sclerites: the tergum, pleura, and the sternum. The tergum in almost all species is membranous, or usually soft and concealed by the wings and elytra when not in flight. The pleura are usually small or hidden in some species, with each pleuron having a single spiracle. The sternum is the most widely visible part of the abdomen, being a more or less sclerotized segment. The abdomen itself does not have any appendages, but some (for example, <a href="/wiki/Mordellidae" title="Mordellidae">Mordellidae</a>) have articulating sternal lobes.<a href="#cite_note-American-79">[78]</a> <div class="mw-heading mw-heading2"><h2 id="Anatomy_and_physiology">Anatomy and physiology</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=15" title="Edit section: Anatomy and physiology">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Internal_morphology_of_Coleoptera.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/79/Internal_morphology_of_Coleoptera.svg/370px-Internal_morphology_of_Coleoptera.svg.png" decoding="async" width="370" height="184" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/79/Internal_morphology_of_Coleoptera.svg/555px-Internal_morphology_of_Coleoptera.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/79/Internal_morphology_of_Coleoptera.svg/740px-Internal_morphology_of_Coleoptera.svg.png 2x" data-file-width="485" data-file-height="241" /></a><figcaption>A beetle's body systems</figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Digestive_system">Digestive system</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=16" title="Edit section: Digestive system">edit</a>]</div> The <a href="/wiki/Digestive_system" class="mw-redirect" title="Digestive system">digestive system</a> of beetles is primarily adapted for a herbivorous diet. Digestion takes place mostly in the anterior <a href="/wiki/Midgut" title="Midgut">midgut</a>, although in predatory groups like the <a href="/wiki/Carabidae" class="mw-redirect" title="Carabidae">Carabidae</a>, most digestion occurs in the crop by means of midgut enzymes. In the <a href="/wiki/Elateridae" class="mw-redirect" title="Elateridae">Elateridae</a>, the larvae are liquid feeders that extraorally digest their food by secreting enzymes.<a href="#cite_note-insenc-8">[8]</a> The alimentary canal basically consists of a short, narrow <a href="/wiki/Pharynx" title="Pharynx">pharynx</a>, a widened expansion, the crop, and a poorly developed <a href="/wiki/Gizzard" title="Gizzard">gizzard</a>. This is followed by the midgut, that varies in dimensions between species, with a large amount of <a href="/wiki/Cecum" title="Cecum">cecum</a>, and the hindgut, with varying lengths. There are typically four to six <a href="/wiki/Malpighian_tubule" class="mw-redirect" title="Malpighian tubule">Malpighian tubules</a>.<a href="#cite_note-Gilliott-7">[7]</a> <div class="mw-heading mw-heading3"><h3 id="Nervous_system">Nervous system</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=17" title="Edit section: Nervous system">edit</a>]</div> The <a href="/wiki/Nervous_system" title="Nervous system">nervous system</a> in beetles contains all the types found in insects, varying between different species, from three thoracic and seven or eight abdominal ganglia which can be distinguished to that in which all the thoracic and abdominal ganglia are fused to form a composite structure.<a href="#cite_note-insenc-8">[8]</a> <div class="mw-heading mw-heading3"><h3 id="Respiratory_system">Respiratory system</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=18" title="Edit section: Respiratory system">edit</a>]</div> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:DytiscusSpiracles.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/DytiscusSpiracles.png/103px-DytiscusSpiracles.png" decoding="async" width="103" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/3f/DytiscusSpiracles.png/155px-DytiscusSpiracles.png 1.5x, //upload.wikimedia.org/wikipedia/commons/3/3f/DytiscusSpiracles.png 2x" data-file-width="199" data-file-height="337" /></a><figcaption><a href="/wiki/Dytiscus" title="Dytiscus">Dytiscus</a> <a href="/wiki/Spiracle_(arthropods)" title="Spiracle (arthropods)">spiracles</a> (right) on upper side of abdomen, normally covered by the elytra, are in contact with an air bubble when the beetle dives.</figcaption></figure> Like most insects, beetles inhale air, for the <a href="/wiki/Oxygen" title="Oxygen">oxygen</a> it contains, and exhale <a href="/wiki/Carbon_dioxide" title="Carbon dioxide">carbon dioxide</a>, via a <a href="/wiki/Invertebrate_trachea" class="mw-redirect" title="Invertebrate trachea">tracheal system</a>. Air enters the body through <a href="/wiki/Spiracle_(arthropods)" title="Spiracle (arthropods)">spiracles</a>, and circulates within the haemocoel in a system of <a href="/wiki/Invertebrate_trachea" class="mw-redirect" title="Invertebrate trachea">tracheae</a> and tracheoles, through whose walls the gases can diffuse.<a href="#cite_note-insenc-8">[8]</a> Diving beetles, such as the <a href="/wiki/Dytiscidae" title="Dytiscidae">Dytiscidae</a>, carry a bubble of air with them when they dive. Such a bubble may be contained under the elytra or against the body by specialized <a href="/wiki/Hydrophobic" class="mw-redirect" title="Hydrophobic">hydrophobic</a> hairs. The bubble covers at least some of the spiracles, permitting air to enter the tracheae.<a href="#cite_note-insenc-8">[8]</a> The function of the bubble is not only to contain a store of air but to act as a <a href="/wiki/Gill#Plastron" title="Gill">physical gill</a>. The air that it traps is in contact with oxygenated water, so as the animal's consumption depletes the oxygen in the bubble, more oxygen can diffuse in to replenish it.<a href="#cite_note-80">[79]</a> Carbon dioxide is more soluble in water than either oxygen or nitrogen, so it readily diffuses out faster than in. Nitrogen is the most plentiful gas in the bubble, and the least soluble, so it constitutes a relatively static component of the bubble and acts as a stable medium for respiratory gases to accumulate in and pass through. Occasional visits to the surface are sufficient for the beetle to re-establish the constitution of the bubble.<a href="#cite_note-81">[80]</a> <div class="mw-heading mw-heading3"><h3 id="Circulatory_system">Circulatory system</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=19" title="Edit section: Circulatory system">edit</a>]</div> Like other insects, beetles have <a href="/wiki/Open_circulatory_system" class="mw-redirect" title="Open circulatory system">open circulatory systems</a>, based on <a href="/wiki/Hemolymph" title="Hemolymph">hemolymph</a> rather than blood. As in other insects, a segmented tube-like heart is attached to the dorsal wall of the <a href="/wiki/Circulatory_system#Nonhuman" title="Circulatory system">hemocoel</a>. It has paired inlets or ostia at intervals down its length, and circulates the hemolymph from the main cavity of the haemocoel and out through the anterior cavity in the head.<a href="#cite_note-82">[81]</a> <div class="mw-heading mw-heading3"><h3 id="Specialized_organs">Specialized organs</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=20" title="Edit section: Specialized organs">edit</a>]</div> Different glands are specialized for different pheromones to attract mates. Pheromones from species of <a href="/wiki/Rutelinae" title="Rutelinae">Rutelinae</a> are produced from <a href="/wiki/Epithelial_cell" class="mw-redirect" title="Epithelial cell">epithelial cells</a> lining the inner surface of the apical abdominal segments; amino acid-based pheromones of <a href="/wiki/Melolonthinae" title="Melolonthinae">Melolonthinae</a> are produced from eversible glands on the abdominal apex. Other species produce different types of pheromones. <a href="/wiki/Dermestid" class="mw-redirect" title="Dermestid">Dermestids</a> produce <a href="/wiki/Ester" title="Ester">esters</a>, and species of <a href="/wiki/Elateridae" class="mw-redirect" title="Elateridae">Elateridae</a> produce <a href="/wiki/Aldehyde" title="Aldehyde">fatty acid-derived aldehydes</a> and <a href="/wiki/Acetate" title="Acetate">acetates</a>.<a href="#cite_note-insenc-8">[8]</a> To attract a mate, fireflies (<a href="/wiki/Lampyridae" class="mw-redirect" title="Lampyridae">Lampyridae</a>) use modified fat body cells with transparent surfaces backed with reflective uric acid crystals to produce light by <a href="/wiki/Bioluminescence" title="Bioluminescence">bioluminescence</a>. Light production is highly efficient, by oxidation of <a href="/wiki/Firefly_luciferin" title="Firefly luciferin">luciferin</a> catalyzed by enzymes (<a href="/wiki/Luciferase" title="Luciferase">luciferases</a>) in the presence of <a href="/wiki/Adenosine_triphosphate" title="Adenosine triphosphate">adenosine triphosphate</a> (ATP) and oxygen, producing <a href="/wiki/Oxyluciferin" class="mw-redirect" title="Oxyluciferin">oxyluciferin</a>, carbon dioxide, and light.<a href="#cite_note-insenc-8">[8]</a> <a href="/wiki/Tympanal_organ" title="Tympanal organ">Tympanal organs</a> or hearing organs consist of a membrane (tympanum) stretched across a frame backed by an air sac and associated sensory neurons, are found in two families.<a href="#cite_note-Scoble-83">[82]</a> Several species of the genus <a href="/wiki/Cicindela" title="Cicindela">Cicindela</a> (Carabidae) have hearing organs on the dorsal surfaces of their first abdominal segments beneath the wings; two tribes in the <a href="/wiki/Dynastinae" title="Dynastinae">Dynastinae</a> (within the <a href="/wiki/Scarabaeidae" title="Scarabaeidae">Scarabaeidae</a>) have hearing organs just beneath their pronotal shields or neck membranes. Both families are sensitive to ultrasonic frequencies, with strong evidence indicating they function to detect the presence of bats by their ultrasonic echolocation.<a href="#cite_note-insenc-8">[8]</a> <div class="mw-heading mw-heading2"><h2 id="Reproduction_and_development">Reproduction and development</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=21" title="Edit section: Reproduction and development">edit</a>]</div> Beetles are members of the <a href="/wiki/Order_(biology)" title="Order (biology)">superorder</a> <a href="/wiki/Holometabola" title="Holometabola">Holometabola</a>, and accordingly most of them undergo complete <a href="/wiki/Metamorphosis" title="Metamorphosis">metamorphosis</a>. The typical form of metamorphosis in beetles passes through four main stages: the <a href="/wiki/Egg" title="Egg">egg</a>, the <a href="/wiki/Larva" title="Larva">larva</a>, the <a href="/wiki/Pupa" title="Pupa">pupa</a>, and the <a href="/wiki/Imago" title="Imago">imago</a> or adult.<a href="#cite_note-Winkler-84">[83]</a> The larvae are commonly called <a href="/wiki/Larva" title="Larva">grubs</a> and the pupa sometimes is called the chrysalis. In some species, the pupa may be enclosed in a cocoon constructed by the larva towards the end of its final <a href="/wiki/Instar" title="Instar">instar</a>. Some beetles, such as typical members of the families <a href="/wiki/Meloidae" class="mw-redirect" title="Meloidae">Meloidae</a> and <a href="/wiki/Rhipiphoridae" class="mw-redirect" title="Rhipiphoridae">Rhipiphoridae</a>, go further, undergoing <a href="/wiki/Hypermetamorphosis" title="Hypermetamorphosis">hypermetamorphosis</a> in which the first instar takes the form of a <a href="/wiki/Triungulin" class="mw-redirect" title="Triungulin">triungulin</a>.<a href="#cite_note-85">[84]</a> <div class="mw-heading mw-heading3"><h3 id="Mating">Mating</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=22" title="Edit section: Mating">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Punctate_Flower_Chafer.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Punctate_Flower_Chafer.jpg/220px-Punctate_Flower_Chafer.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/66/Punctate_Flower_Chafer.jpg/330px-Punctate_Flower_Chafer.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/66/Punctate_Flower_Chafer.jpg/440px-Punctate_Flower_Chafer.jpg 2x" data-file-width="1600" data-file-height="1067" /></a><figcaption><a href="/wiki/Punctate_flower_chafer" title="Punctate flower chafer">Punctate flower chafers</a> (Neorrhina punctata, Scarabaeidae) mating</figcaption></figure> Some beetles have intricate mating behaviour. <a href="/wiki/Pheromone" title="Pheromone">Pheromone</a> communication is often important in locating a mate. Different species use different pheromones. <a href="/wiki/White_worm_beetle" title="White worm beetle">Scarab beetles</a> such as the <a href="/wiki/Rutelinae" title="Rutelinae">Rutelinae</a> use pheromones derived from <a href="/wiki/Fatty_acid_synthesis" title="Fatty acid synthesis">fatty acid synthesis</a> and others use pheromones from organic compounds, while other scarabs such as the <a href="/wiki/Melolonthinae" title="Melolonthinae">Melolonthinae</a> use <a href="/wiki/Amino_acid" title="Amino acid">amino acids</a> and terpenoids. Another way beetles find mates is seen in the <a href="/wiki/Fireflies" class="mw-redirect" title="Fireflies">fireflies</a> (Lampyridae) which are <a href="/wiki/Bioluminescent" class="mw-redirect" title="Bioluminescent">bioluminescent</a>, with abdominal light-producing organs. The males and females engage in a complex dialog before mating; each species has a unique combination of flight patterns, duration, composition, and intensity of the light produced.<a href="#cite_note-insenc-8">[8]</a> Before mating, males and females may stridulate, or vibrate the objects they are on. In the Meloidae, the male climbs onto the dorsum of the female and strokes his antennae on her head, palps, and antennae. In <a href="/wiki/Eupompha" title="Eupompha">Eupompha</a>, the male draws his antennae along his longitudinal vertex. They may not mate at all if they do not perform the precopulatory ritual.<a href="#cite_note-insenc-8">[8]</a> This mating behavior may be different amongst dispersed populations of the same species. For example, the mating of a <a href="/wiki/Russia" title="Russia">Russian</a> population of <a href="/wiki/Tansy_beetle" title="Tansy beetle">tansy beetle</a> (Chrysolina graminis) is preceded by an elaborate ritual involving the male tapping the female's eyes, pronotum and antennae with its antennae, which is not evident in the population of this species in the <a href="/wiki/United_Kingdom" title="United Kingdom">United Kingdom</a>.<a href="#cite_note-86">[85]</a> In another example, the <a href="/wiki/Intromittent_organ" title="Intromittent organ">intromittent organ</a> of male <a href="/wiki/Thistle_tortoise_beetle" title="Thistle tortoise beetle">thistle tortoise beetles</a> is a long, tube-like structure called the <a href="/wiki/Flagellum" title="Flagellum">flagellum</a> which is thin and curved. When not in use, the flagellum is stored inside the <a href="/wiki/Abdomen" title="Abdomen">abdomen</a> of the male and can extend out to be longer than the male when needed. During mating, this organ bends to the complex shape of the female <a href="/wiki/Reproductive_organ" class="mw-redirect" title="Reproductive organ">reproductive organ</a>, which includes a coiled duct that the male must penetrate with the organ. Furthermore, these physical properties of the thistle tortioise beetle have been studied because the ability of a thin, flexible structure to harden without buckling or rupturing is mechanically challenging and may have important implications for the development of microscopic <a href="/wiki/Catheter" title="Catheter">catheters</a> in modern medicine.<a href="#cite_note-87">[86]</a> <a href="/wiki/Competition_(biology)" title="Competition (biology)">Competition</a> can play a part in the mating rituals of species such as <a href="/wiki/Burying_beetle" title="Burying beetle">burying beetles</a> (Nicrophorus), the insects fighting to determine which can mate. Many male beetles are <a href="/wiki/Territory_(animal)" title="Territory (animal)">territorial</a> and fiercely defend their territories from intruding males. In such species, the male often has horns on the head or thorax, making its body length greater than that of a female. Copulation is generally quick, but in some cases lasts for several hours. During copulation, <a href="/wiki/Spermatozoon" title="Spermatozoon">sperm cells</a> are transferred to the female to <a href="/wiki/Fertilize" class="mw-redirect" title="Fertilize">fertilize</a> the egg.<a href="#cite_note-Gilliott-7">[7]</a> <div class="mw-heading mw-heading3"><h3 id="Life_cycle">Life cycle</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=23" title="Edit section: Life cycle">edit</a>]</div> <figure typeof="mw:File/Thumb"><a href="/wiki/File:Life_cycle_of_stag_beetle.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Life_cycle_of_stag_beetle.svg/300px-Life_cycle_of_stag_beetle.svg.png" decoding="async" width="300" height="301" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Life_cycle_of_stag_beetle.svg/450px-Life_cycle_of_stag_beetle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8f/Life_cycle_of_stag_beetle.svg/600px-Life_cycle_of_stag_beetle.svg.png 2x" data-file-width="472" data-file-height="474" /></a><figcaption>The life cycle of the <a href="/wiki/Stag_beetle" title="Stag beetle">stag beetle</a> includes three <a href="/wiki/Instar" title="Instar">instars</a>.</figcaption></figure> <div class="mw-heading mw-heading4"><h4 id="Egg">Egg</h4>[<a href="/w/index.php?title=Beetle&action=edit&section=24" title="Edit section: Egg">edit</a>]</div> Essentially all beetles lay eggs, though some <a href="/wiki/Myrmecophilous" class="mw-redirect" title="Myrmecophilous">myrmecophilous</a> <a href="/wiki/Aleocharinae" title="Aleocharinae">Aleocharinae</a> and some <a href="/wiki/Chrysomelinae" title="Chrysomelinae">Chrysomelinae</a> which live in mountains or the subarctic are <a href="/wiki/Ovoviviparous" class="mw-redirect" title="Ovoviviparous">ovoviviparous</a>, laying eggs which hatch almost immediately.<a href="#cite_note-Winkler-84">[83]</a> Beetle eggs generally have smooth surfaces and are soft, though the <a href="/wiki/Cupedidae" title="Cupedidae">Cupedidae</a> have hard eggs. Eggs vary widely between species: the eggs tend to be small in species with many instars (larval stages), and in those that lay large numbers of eggs. A female may lay from several dozen to several thousand eggs during her lifetime, depending on the extent of parental care. This ranges from the simple laying of eggs under a leaf, to the parental care provided by <a href="/wiki/Scarab_beetles" class="mw-redirect" title="Scarab beetles">scarab beetles</a>, which house, feed and protect their young. The <a href="/wiki/Attelabidae" title="Attelabidae">Attelabidae</a> roll leaves and lay their eggs inside the roll for protection.<a href="#cite_note-insenc-8">[8]</a><a href="#cite_note-Crowson-88">[87]</a> <div class="mw-heading mw-heading4"><h4 id="Larva">Larva</h4>[<a href="/w/index.php?title=Beetle&action=edit&section=25" title="Edit section: Larva">edit</a>]</div> The <a href="/wiki/Larva" title="Larva">larva</a> is usually the principal feeding stage of the beetle <a href="/wiki/Biological_life_cycle" title="Biological life cycle">life cycle</a>. Larvae tend to feed voraciously once they emerge from their eggs. Some feed externally on plants, such as those of certain leaf beetles, while others feed within their food sources. Examples of internal feeders are most <a href="/wiki/Buprestidae" title="Buprestidae">Buprestidae</a> and longhorn beetles. The larvae of many beetle families are predatory like the adults (ground beetles, ladybirds, rove beetles). The larval period varies between species, but can be as long as several years. The larvae of <a href="/wiki/Skin_beetle" class="mw-redirect" title="Skin beetle">skin beetles</a> undergo a degree of reversed development when starved, and later grow back to the previously attained level of maturity. The cycle can be repeated many times (see <a href="/wiki/Biological_immortality" title="Biological immortality">Biological immortality</a>).<a href="#cite_note-89">[88]</a> Larval morphology is highly varied amongst species, with well-developed and sclerotized heads, distinguishable thoracic and abdominal segments (usually the tenth, though sometimes the eighth or ninth).<a href="#cite_note-Gilliott-7">[7]</a> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Hercules_beetle_(larva).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Hercules_beetle_%28larva%29.jpg/130px-Hercules_beetle_%28larva%29.jpg" decoding="async" width="130" height="231" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Hercules_beetle_%28larva%29.jpg/195px-Hercules_beetle_%28larva%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2d/Hercules_beetle_%28larva%29.jpg/260px-Hercules_beetle_%28larva%29.jpg 2x" data-file-width="2248" data-file-height="4000" /></a><figcaption><a href="/wiki/Scarabaeoidea" title="Scarabaeoidea">Scarabaeiform</a> larva of <a href="/wiki/Hercules_beetle" title="Hercules beetle">Hercules beetle</a></figcaption></figure> Beetle larvae can be differentiated from other insect larvae by their hardened, often darkened heads, the presence of chewing mouthparts, and <a href="/wiki/Spiracle_(arthropods)" title="Spiracle (arthropods)">spiracles</a> along the sides of their bodies. Like adult beetles, the larvae are varied in appearance, particularly between beetle families. Beetles with somewhat flattened, highly mobile larvae include the ground beetles and rove beetles; their larvae are described as campodeiform. Some beetle larvae resemble hardened worms with dark head capsules and minute legs. These are elateriform larvae, and are found in the <a href="/wiki/Click_beetle" title="Click beetle">click beetle</a> (Elateridae) and <a href="/wiki/Darkling_beetle" title="Darkling beetle">darkling beetle</a> (Tenebrionidae) families. Some elateriform larvae of click beetles are known as wireworms. Beetles in the <a href="/wiki/Scarabaeoidea" title="Scarabaeoidea">Scarabaeoidea</a> have short, thick larvae described as scarabaeiform, more commonly known as grubs.<a href="#cite_note-90">[89]</a> All beetle larvae go through several <a href="/wiki/Instar" title="Instar">instars</a>, which are the developmental stages between each <a href="/wiki/Ecdysis" title="Ecdysis">moult</a>. In many species, the larvae simply increase in size with each successive instar as more food is consumed. In some cases, however, more dramatic changes occur. Among certain beetle families or genera, particularly those that exhibit parasitic lifestyles, the first instar (the <a href="/wiki/Planidium" title="Planidium">planidium</a>) is highly mobile to search out a host, while the following instars are more sedentary and remain on or within their host. This is known as <a href="/wiki/Hypermetamorphosis" title="Hypermetamorphosis">hypermetamorphosis</a>; it occurs in the <a href="/wiki/Meloidae" class="mw-redirect" title="Meloidae">Meloidae</a>, <a href="/wiki/Micromalthidae" title="Micromalthidae">Micromalthidae</a>, and <a href="/wiki/Ripiphoridae" title="Ripiphoridae">Ripiphoridae</a>.<a href="#cite_note-91">[90]</a> The blister beetle <a href="/wiki/Epicauta_vittata" title="Epicauta vittata">Epicauta vittata</a> (Meloidae), for example, has three distinct larval stages. Its first stage, the <a href="/wiki/Triungulin" class="mw-redirect" title="Triungulin">triungulin</a>, has longer legs to go in search of the eggs of grasshoppers. After feeding for a week it moults to the second stage, called the caraboid stage, which resembles the larva of a <a href="/wiki/Carabidae" class="mw-redirect" title="Carabidae">carabid beetle</a>. In another week it moults and assumes the appearance of a <a href="/wiki/Scarabaeidae" title="Scarabaeidae">scarabaeid larva</a>—the scarabaeidoid stage. Its penultimate larval stage is the pseudo-pupa or the coarcate larva, which will overwinter and pupate until the next spring.<a href="#cite_note-92">[91]</a> The larval period can vary widely. A fungus feeding staphylinid <a href="/wiki/Phanerota_fasciata" title="Phanerota fasciata">Phanerota fasciata</a> undergoes three moults in 3.2 days at room temperature while Anisotoma sp. (Leiodidae) completes its larval stage in the fruiting body of slime mold in 2 days and possibly represents the fastest growing beetles. Dermestid beetles, Trogoderma inclusum can remain in an extended larval state under unfavourable conditions, even reducing their size between moults. A larva is reported to have survived for 3.5 years in an enclosed container.<a href="#cite_note-insenc-8">[8]</a> <div class="mw-heading mw-heading4"><h4 id="Pupa_and_adult">Pupa and adult</h4>[<a href="/w/index.php?title=Beetle&action=edit&section=26" title="Edit section: Pupa and adult">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Eburia_quadrigeminata.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/Eburia_quadrigeminata.jpg/220px-Eburia_quadrigeminata.jpg" decoding="async" width="220" height="224" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/Eburia_quadrigeminata.jpg/330px-Eburia_quadrigeminata.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/57/Eburia_quadrigeminata.jpg/440px-Eburia_quadrigeminata.jpg 2x" data-file-width="549" data-file-height="560" /></a><figcaption>The ivory-marked beetle, <a href="/wiki/Eburia_quadrigeminata" title="Eburia quadrigeminata">Eburia quadrigeminata</a>, may live up to 40 years inside the <a href="/wiki/Hardwood" title="Hardwood">hardwoods</a> on which the larva feeds.</figcaption></figure> As with all holometabolans, beetle larvae pupate, and from these <a href="/wiki/Pupa" title="Pupa">pupae</a> emerge fully formed, sexually mature adult beetles, or <a href="/wiki/Imago" title="Imago">imagos</a>. Pupae never have mandibles (they are <a href="/wiki/Adecticous" class="mw-redirect" title="Adecticous">adecticous</a>). In most pupae, the appendages are not attached to the body and are said to be <a href="/wiki/Exarate" class="mw-redirect" title="Exarate">exarate</a>; in a few beetles (Staphylinidae, Ptiliidae etc.) the appendages are fused with the body (termed as <a href="/wiki/Pupa#Position_in_life_cycle" title="Pupa">obtect</a> pupae).<a href="#cite_note-Gilliott-7">[7]</a> Adults have extremely variable lifespans, from weeks to years, depending on the species.<a href="#cite_note-Gilliott-7">[7]</a><a href="#cite_note-ausbeetle-51">[51]</a> Some wood-boring beetles can have extremely long life-cycles. It is believed that when furniture or house timbers are infested by beetle larvae, the timber already contained the larvae when it was first sawn up. A <a href="/wiki/Birch" title="Birch">birch</a> bookcase 40 years old released adult <a href="/wiki/Eburia_quadrigeminata" title="Eburia quadrigeminata">Eburia quadrigeminata</a> (<a href="/wiki/Cerambycidae" class="mw-redirect" title="Cerambycidae">Cerambycidae</a>), while <a href="/wiki/Buprestis_aurulenta" title="Buprestis aurulenta">Buprestis aurulenta</a> and other <a href="/wiki/Buprestidae" title="Buprestidae">Buprestidae</a> have been documented as emerging as much as 51 years after manufacture of wooden items.<a href="#cite_note-Zeng-93">[92]</a> <div class="mw-heading mw-heading2"><h2 id="Behaviour">Behaviour</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=27" title="Edit section: Behaviour">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="Locomotion">Locomotion</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=28" title="Edit section: Locomotion">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Photinus_pyralis_Firefly_2.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Photinus_pyralis_Firefly_2.jpg/220px-Photinus_pyralis_Firefly_2.jpg" decoding="async" width="220" height="222" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/81/Photinus_pyralis_Firefly_2.jpg/330px-Photinus_pyralis_Firefly_2.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/81/Photinus_pyralis_Firefly_2.jpg/440px-Photinus_pyralis_Firefly_2.jpg 2x" data-file-width="1035" data-file-height="1046" /></a><figcaption><a href="/wiki/Photinus_pyralis" title="Photinus pyralis">Photinus pyralis</a>, firefly, in flight</figcaption></figure> The elytra allow beetles to both fly and move through confined spaces, doing so by folding the delicate wings under the elytra while not flying, and folding their wings out just before takeoff. The unfolding and folding of the wings is operated by muscles attached to the wing base; as long as the tension on the radial and cubital veins remains, the wings remain straight.<a href="#cite_note-insenc-8">[8]</a> Some beetle species (many <a href="/wiki/Cetoniinae" class="mw-redirect" title="Cetoniinae">Cetoniinae</a>; some <a href="/wiki/Scarabaeinae" title="Scarabaeinae">Scarabaeinae</a>, <a href="/wiki/Curculionidae" title="Curculionidae">Curculionidae</a> and <a href="/wiki/Buprestidae" title="Buprestidae">Buprestidae</a>) fly with the elytra closed, with the metathoracic wings extended under the lateral elytra margins.<a href="#cite_note-94">[93]</a> The altitude reached by beetles in flight varies. One study investigating the flight altitude of the ladybird species <a href="/wiki/Coccinella_septempunctata" title="Coccinella septempunctata">Coccinella septempunctata</a> and <a href="/wiki/Harmonia_axyridis" title="Harmonia axyridis">Harmonia axyridis</a> using radar showed that, whilst the majority in flight over a single location were at 150–195 m above ground level, some reached altitudes of over 1100 m.<a href="#cite_note-95">[94]</a> Many rove beetles have greatly reduced elytra, and while they are capable of flight, they most often move on the ground: their soft bodies and strong abdominal muscles make them flexible, easily able to wriggle into small cracks.<a href="#cite_note-96">[95]</a> Aquatic beetles use several techniques for retaining air beneath the water's surface. Diving beetles (Dytiscidae) hold air between the abdomen and the elytra when diving. Hydrophilidae have hairs on their under surface that retain a layer of air against their bodies. Adult crawling <a href="/wiki/Water_beetle" title="Water beetle">water beetles</a> use both their elytra and their hind <a href="/wiki/Arthropod_leg" title="Arthropod leg">coxae</a> (the basal segment of the back legs) in air retention, while <a href="/wiki/Whirligig_beetle" title="Whirligig beetle">whirligig beetles</a> simply carry an air bubble down with them whenever they dive.<a href="#cite_note-97">[96]</a> <div class="mw-heading mw-heading3"><h3 id="Communication">Communication</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=29" title="Edit section: Communication">edit</a>]</div> Beetles have a variety of ways to communicate, including the use of <a href="/wiki/Pheromone" title="Pheromone">pheromones</a>. The <a href="/wiki/Mountain_pine_beetle" title="Mountain pine beetle">mountain pine beetle</a> emits a pheromone to attract other beetles to a tree. The mass of beetles are able to overcome the chemical defenses of the tree. After the tree's defenses have been exhausted, the beetles emit an anti-aggregation pheromone. This species can <a href="/wiki/Stridulate" class="mw-redirect" title="Stridulate">stridulate</a> to communicate,<a href="#cite_note-Beetle_love-98">[97]</a> but others may use sound to defend themselves when attacked.<a href="#cite_note-stridulation-99">[98]</a> <div class="mw-heading mw-heading3"><h3 id="Parental_care">Parental care</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=30" title="Edit section: Parental care">edit</a>]</div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Scarabaeus_laticollis.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Scarabaeus_laticollis.jpg/170px-Scarabaeus_laticollis.jpg" decoding="async" width="170" height="187" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Scarabaeus_laticollis.jpg/255px-Scarabaeus_laticollis.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/e/e8/Scarabaeus_laticollis.jpg/340px-Scarabaeus_laticollis.jpg 2x" data-file-width="1000" data-file-height="1100" /></a><figcaption>A dung beetle rolling dung</figcaption></figure> Parental care is found in a few families<a href="#cite_note-100">[99]</a> of beetle, perhaps for protection against adverse conditions and predators.<a href="#cite_note-insenc-8">[8]</a> The rove beetle <a href="/wiki/Bledius_spectabilis" title="Bledius spectabilis">Bledius spectabilis</a> lives in <a href="/wiki/Salt_marsh" title="Salt marsh">salt marshes</a>, so the eggs and larvae are endangered by the rising <a href="/wiki/Tide" title="Tide">tide</a>. The maternal beetle patrols the eggs and larvae, burrowing to keep them from flooding and <a href="/wiki/Asphyxiating" class="mw-redirect" title="Asphyxiating">asphyxiating</a>, and protects them from the predatory carabid beetle <a href="/wiki/Dicheirotrichus_gustavii" title="Dicheirotrichus gustavii">Dicheirotrichus gustavii</a> and from the <a href="/wiki/Parasitism" title="Parasitism">parasitoidal</a> wasp <a href="/wiki/Barycnemis_blediator" title="Barycnemis blediator">Barycnemis blediator</a>, which kills some 15% of the larvae.<a href="#cite_note-101">[100]</a> <a href="/wiki/Burying_beetle" title="Burying beetle">Burying beetles</a> are attentive parents, and participate in cooperative care and feeding of their offspring. Both parents work to bury small animal carcass to serve as a food resource for their young and build a brood chamber around it. The parents prepare the carcass and protect it from competitors and from early decomposition. After their eggs hatch, the parents keep the larvae clean of fungus and bacteria and help the larvae feed by regurgitating food for them.<a href="#cite_note-102">[101]</a> Some <a href="/wiki/Dung_beetle" title="Dung beetle">dung beetles</a> provide parental care, collecting herbivore dung and laying eggs within that food supply, an instance of <a href="/wiki/Mass_provisioning" title="Mass provisioning">mass provisioning</a>. Some species do not leave after this stage, but remain to safeguard their offspring.<a href="#cite_note-103">[102]</a> Most species of beetles do not display parental care behaviors after the eggs have been laid.<a href="#cite_note-104">[103]</a> Subsociality, where females guard their offspring, is well-documented in two families of Chrysomelidae, Cassidinae and Chrysomelinae.<a href="#cite_note-105">[104]</a><a href="#cite_note-106">[105]</a><a href="#cite_note-107">[106]</a><a href="#cite_note-108">[107]</a><a href="#cite_note-109">[108]</a> <div class="mw-heading mw-heading3"><h3 id="Eusociality">Eusociality</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=31" title="Edit section: Eusociality">edit</a>]</div> <a href="/wiki/Eusociality" title="Eusociality">Eusociality</a> involves cooperative brood care (including brood care of offspring from other individuals), overlapping generations within a colony of adults, and a division of labor into reproductive and non-reproductive groups.<a href="#cite_note-110">[109]</a> Few organisms outside <a href="/wiki/Hymenoptera" title="Hymenoptera">Hymenoptera</a> exhibit this behavior; the only beetle to do so is the weevil <a href="/wiki/Austroplatypus_incompertus" title="Austroplatypus incompertus">Austroplatypus incompertus</a>.<a href="#cite_note-Kent1992-111">[110]</a> This <a href="/wiki/Australia" title="Australia">Australian</a> species lives in horizontal networks of tunnels, in the <a href="/wiki/Heartwood" class="mw-redirect" title="Heartwood">heartwood</a> of <a href="/wiki/Eucalyptus" title="Eucalyptus">Eucalyptus</a> trees. It is one of more than 300 species of wood-boring <a href="/wiki/Ambrosia_beetle" title="Ambrosia beetle">Ambrosia beetles</a> which distribute the spores of ambrosia fungi.<a href="#cite_note-bee-112">[111]</a> The fungi grow in the beetles' tunnels, providing food for the beetles and their larvae; female offspring remain in the tunnels and maintain the fungal growth, probably never reproducing.<a href="#cite_note-bee-112">[111]</a><a href="#cite_note-Kent1992-111">[110]</a> Cooperative brood care is also found in the bess beetles (<a href="/wiki/Passalidae" title="Passalidae">Passalidae</a>) where the larvae feed on the semi-digested faeces of the adults.<a href="#cite_note-113">[112]</a> <div class="mw-heading mw-heading3"><h3 id="Feeding">Feeding</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=32" title="Edit section: Feeding">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Orange_Blister_Beetle_(Mylabris_pustulata)_on_Ipomoea_carnea_W_IMG_0593.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Orange_Blister_Beetle_%28Mylabris_pustulata%29_on_Ipomoea_carnea_W_IMG_0593.jpg/220px-Orange_Blister_Beetle_%28Mylabris_pustulata%29_on_Ipomoea_carnea_W_IMG_0593.jpg" decoding="async" width="220" height="150" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/19/Orange_Blister_Beetle_%28Mylabris_pustulata%29_on_Ipomoea_carnea_W_IMG_0593.jpg/330px-Orange_Blister_Beetle_%28Mylabris_pustulata%29_on_Ipomoea_carnea_W_IMG_0593.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/19/Orange_Blister_Beetle_%28Mylabris_pustulata%29_on_Ipomoea_carnea_W_IMG_0593.jpg/440px-Orange_Blister_Beetle_%28Mylabris_pustulata%29_on_Ipomoea_carnea_W_IMG_0593.jpg 2x" data-file-width="882" data-file-height="600" /></a><figcaption><a href="/wiki/Hycleus" title="Hycleus">Hycleus</a> sp. (<a href="/wiki/Meloidae" class="mw-redirect" title="Meloidae">Meloidae</a>) feeding on the petals of <a href="/wiki/Ipomoea_carnea" title="Ipomoea carnea">Ipomoea carnea</a></figcaption></figure> Beetles are able to exploit a wide diversity of food sources available in their many habitats. Some are <a href="/wiki/Omnivore" title="Omnivore">omnivores</a>, eating both plants and animals. Other beetles are highly specialized in their diet. Many species of leaf beetles, longhorn beetles, and weevils are very host-specific, feeding on only a single species of plant. <a href="/wiki/Ground_beetle" title="Ground beetle">Ground beetles</a> and <a href="/wiki/Rove_beetle" title="Rove beetle">rove beetles</a> (<a href="/wiki/Staphylinidae" class="mw-redirect" title="Staphylinidae">Staphylinidae</a>), among others, are primarily carnivorous and catch and consume many other <a href="/wiki/Arthropod" title="Arthropod">arthropods</a> and small prey, such as earthworms and snails. While most predatory beetles are generalists, a few species have more specific prey requirements or preferences.<a href="#cite_note-114">[113]</a> In some species, digestive ability relies upon a <a href="/wiki/Symbiotic" class="mw-redirect" title="Symbiotic">symbiotic</a> relationship with <a href="/wiki/Fungi" class="mw-redirect" title="Fungi">fungi</a> - some beetles have yeasts living their guts, including some yeasts previously undiscovered anywhere else.<a href="#cite_note-115">[114]</a> Decaying organic matter is a primary diet for many species. This can range from <a href="/wiki/Feces" title="Feces">dung</a>, which is consumed by <a href="/wiki/Coprophagous" class="mw-redirect" title="Coprophagous">coprophagous</a> species (such as certain <a href="/wiki/Dung_beetle" title="Dung beetle">scarab beetles</a> in the <a href="/wiki/Scarabaeidae" title="Scarabaeidae">Scarabaeidae</a>), to dead animals, which are eaten by <a href="/wiki/Necrophagous" class="mw-redirect" title="Necrophagous">necrophagous</a> species (such as the <a href="/wiki/Carrion_beetle" class="mw-redirect" title="Carrion beetle">carrion beetles</a>, <a href="/wiki/Silphidae" title="Silphidae">Silphidae</a>). Some beetles found in dung and carrion are in fact predatory. These include members of the <a href="/wiki/Histeridae" title="Histeridae">Histeridae</a> and <a href="/wiki/Silphidae" title="Silphidae">Silphidae</a>, preying on the larvae of <a href="/wiki/Coprophagous" class="mw-redirect" title="Coprophagous">coprophagous</a> and <a href="/wiki/Necrophagous" class="mw-redirect" title="Necrophagous">necrophagous</a> insects.<a href="#cite_note-116">[115]</a> Many beetles feed under bark, some feed on wood while others feed on fungi growing on wood or leaf-litter. Some beetles have special <a href="/wiki/Mycangia" class="mw-redirect" title="Mycangia">mycangia</a>, structures for the transport of fungal spores.<a href="#cite_note-117">[116]</a> <div class="mw-heading mw-heading2"><h2 id="Ecology">Ecology</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=33" title="Edit section: Ecology">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Long-horn_Beetle_-_Ecyrus_dasycerus,_Leesylvania_State_Park,_Woodbridge,_Virginia.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Long-horn_Beetle_-_Ecyrus_dasycerus%2C_Leesylvania_State_Park%2C_Woodbridge%2C_Virginia.jpg/170px-Long-horn_Beetle_-_Ecyrus_dasycerus%2C_Leesylvania_State_Park%2C_Woodbridge%2C_Virginia.jpg" decoding="async" width="170" height="273" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Long-horn_Beetle_-_Ecyrus_dasycerus%2C_Leesylvania_State_Park%2C_Woodbridge%2C_Virginia.jpg/255px-Long-horn_Beetle_-_Ecyrus_dasycerus%2C_Leesylvania_State_Park%2C_Woodbridge%2C_Virginia.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f8/Long-horn_Beetle_-_Ecyrus_dasycerus%2C_Leesylvania_State_Park%2C_Woodbridge%2C_Virginia.jpg/340px-Long-horn_Beetle_-_Ecyrus_dasycerus%2C_Leesylvania_State_Park%2C_Woodbridge%2C_Virginia.jpg 2x" data-file-width="1704" data-file-height="2736" /></a><figcaption>A <a href="/wiki/Camouflage" title="Camouflage">camouflaged</a> <a href="/wiki/Longhorn_beetle" title="Longhorn beetle">longhorn beetle</a>, <a href="/wiki/Ecyrus_dasycerus" title="Ecyrus dasycerus">Ecyrus dasycerus</a></figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Anti-predator_adaptations">Anti-predator adaptations</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=34" title="Edit section: Anti-predator adaptations">edit</a>]</div> Beetles, both adults and larvae, are the prey of many animal <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> including <a href="/wiki/Mammals" class="mw-redirect" title="Mammals">mammals</a> from <a href="/wiki/Bat" title="Bat">bats</a> to <a href="/wiki/Rodent" title="Rodent">rodents</a>, <a href="/wiki/Birds" class="mw-redirect" title="Birds">birds</a>, <a href="/wiki/Lizard" title="Lizard">lizards</a>, <a href="/wiki/Amphibians" class="mw-redirect" title="Amphibians">amphibians</a>, <a href="/wiki/Fishes" class="mw-redirect" title="Fishes">fishes</a>, <a href="/wiki/Dragonflies" class="mw-redirect" title="Dragonflies">dragonflies</a>, <a href="/wiki/Robberflies" class="mw-redirect" title="Robberflies">robberflies</a>, <a href="/wiki/Reduviidae" title="Reduviidae">reduviid bugs</a>, <a href="/wiki/Ant" title="Ant">ants</a>, other beetles, and <a href="/wiki/Spider" title="Spider">spiders</a>.<a href="#cite_note-Evans27-118">[117]</a><a href="#cite_note-119">[118]</a> Beetles use a variety of <a href="/wiki/Anti-predator_adaptation" title="Anti-predator adaptation">anti-predator adaptations</a> to defend themselves. These include <a href="/wiki/Camouflage" title="Camouflage">camouflage</a> and <a href="/wiki/Mimicry" title="Mimicry">mimicry</a> against predators that hunt by sight, toxicity, and defensive behaviour. <div class="mw-heading mw-heading4"><h4 id="Camouflage">Camouflage</h4>[<a href="/w/index.php?title=Beetle&action=edit&section=35" title="Edit section: Camouflage">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Camouflage" title="Camouflage">Camouflage</a></div> Camouflage is common and widespread among beetle families, especially those that feed on wood or vegetation, such as <a href="/wiki/Leaf_beetle" title="Leaf beetle">leaf beetles</a> (Chrysomelidae, which are often green) and <a href="/wiki/Weevil" title="Weevil">weevils</a>. In some species, sculpturing or various colored scales or hairs cause beetles such as the <a href="/wiki/Avocado" title="Avocado">avocado</a> weevil <a href="/wiki/Heilipus_apiatus" title="Heilipus apiatus">Heilipus apiatus</a> to resemble bird dung or other inedible objects.<a href="#cite_note-Evans27-118">[117]</a> Many beetles that live in sandy environments blend in with the coloration of that substrate.<a href="#cite_note-Evans126-120">[119]</a> <div class="mw-heading mw-heading4"><h4 id="Mimicry_and_aposematism">Mimicry and aposematism</h4>[<a href="/w/index.php?title=Beetle&action=edit&section=36" title="Edit section: Mimicry and aposematism">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Clyte_b%C3%A9lier_-_MHNT_-_Vue_dorsale.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Clyte_b%C3%A9lier_-_MHNT_-_Vue_dorsale.jpg/220px-Clyte_b%C3%A9lier_-_MHNT_-_Vue_dorsale.jpg" decoding="async" width="220" height="175" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Clyte_b%C3%A9lier_-_MHNT_-_Vue_dorsale.jpg/330px-Clyte_b%C3%A9lier_-_MHNT_-_Vue_dorsale.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/c/ce/Clyte_b%C3%A9lier_-_MHNT_-_Vue_dorsale.jpg/440px-Clyte_b%C3%A9lier_-_MHNT_-_Vue_dorsale.jpg 2x" data-file-width="2891" data-file-height="2297" /></a><figcaption><a href="/wiki/Clytus_arietis" title="Clytus arietis">Clytus arietis</a> (<a href="/wiki/Cerambycidae" class="mw-redirect" title="Cerambycidae">Cerambycidae</a>), a <a href="/wiki/Batesian_mimic" class="mw-redirect" title="Batesian mimic">Batesian mimic</a> of wasps</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Mimicry" title="Mimicry">Mimicry</a> and <a href="/wiki/Aposematism" title="Aposematism">Aposematism</a></div> Some <a href="/wiki/Longhorn_beetle" title="Longhorn beetle">longhorn beetles</a> (Cerambycidae) are effective <a href="/wiki/Batesian_mimics" class="mw-redirect" title="Batesian mimics">Batesian mimics</a> of <a href="/wiki/Wasp" title="Wasp">wasps</a>. Beetles may combine coloration with behavioural mimicry, acting like the wasps they already closely resemble. Many other beetles, including <a href="/wiki/Ladybirds" class="mw-redirect" title="Ladybirds">ladybirds</a>, <a href="/wiki/Blister_beetle" title="Blister beetle">blister beetles</a>, and <a href="/wiki/Lycidae" title="Lycidae">lycid beetles</a> secrete distasteful or toxic substances to make them unpalatable or poisonous, and are often <a href="/wiki/Aposematic" class="mw-redirect" title="Aposematic">aposematic</a>, where bright or contrasting <a href="/wiki/Animal_coloration" title="Animal coloration">coloration</a> warn off predators; many beetles and other insects mimic these chemically protected species.<a href="#cite_note-Evans-121">[120]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Blister_beetle_(26390828032).jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/Blister_beetle_%2826390828032%29.jpg/220px-Blister_beetle_%2826390828032%29.jpg" decoding="async" width="220" height="176" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/f3/Blister_beetle_%2826390828032%29.jpg/330px-Blister_beetle_%2826390828032%29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/f3/Blister_beetle_%2826390828032%29.jpg/440px-Blister_beetle_%2826390828032%29.jpg 2x" data-file-width="1600" data-file-height="1280" /></a><figcaption>Blister beetles such as <a href="/wiki/Hycleus" title="Hycleus">Hycleus</a> have brilliant <a href="/wiki/Aposematic" class="mw-redirect" title="Aposematic">aposematic</a> coloration, warning of their toxicity.</figcaption></figure> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Bloody_-nosed_Beetle%5E_Timarcha_tenebricosa_-_Flickr_-_gailhampshire.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/25/Bloody_-nosed_Beetle%5E_Timarcha_tenebricosa_-_Flickr_-_gailhampshire.jpg/137px-Bloody_-nosed_Beetle%5E_Timarcha_tenebricosa_-_Flickr_-_gailhampshire.jpg" decoding="async" width="137" height="172" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/25/Bloody_-nosed_Beetle%5E_Timarcha_tenebricosa_-_Flickr_-_gailhampshire.jpg/206px-Bloody_-nosed_Beetle%5E_Timarcha_tenebricosa_-_Flickr_-_gailhampshire.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/25/Bloody_-nosed_Beetle%5E_Timarcha_tenebricosa_-_Flickr_-_gailhampshire.jpg/275px-Bloody_-nosed_Beetle%5E_Timarcha_tenebricosa_-_Flickr_-_gailhampshire.jpg 2x" data-file-width="543" data-file-height="680" /></a><figcaption>The bloody-nosed beetle, <a href="/wiki/Timarcha_tenebricosa" class="mw-redirect" title="Timarcha tenebricosa">Timarcha tenebricosa</a>, <a href="/wiki/Antipredator_adaptation" class="mw-redirect" title="Antipredator adaptation">defending itself</a> by releasing a droplet of noxious red liquid (base of leg, on right)</figcaption></figure> Chemical defense is important in some species, usually being advertised by bright aposematic colors. Some <a href="/wiki/Tenebrionidae" class="mw-redirect" title="Tenebrionidae">Tenebrionidae</a> use their posture for releasing noxious chemicals to warn off predators. Chemical defenses may serve purposes other than just protection from vertebrates, such as protection from a wide range of microbes. Some species sequester chemicals from the plants they feed on, incorporating them into their own defenses.<a href="#cite_note-Evans126-120">[119]</a> Other species have special glands to produce deterrent chemicals. The defensive glands of carabid ground beetles produce a variety of <a href="/wiki/Hydrocarbon" title="Hydrocarbon">hydrocarbons</a>, <a href="/wiki/Aldehyde" title="Aldehyde">aldehydes</a>, <a href="/wiki/Phenol" title="Phenol">phenols</a>, <a href="/wiki/Quinone" title="Quinone">quinones</a>, <a href="/wiki/Ester" title="Ester">esters</a>, and <a href="/wiki/Acid" title="Acid">acids</a> released from an opening at the end of the abdomen. African carabid beetles (for example, <a href="/wiki/Anthia" title="Anthia">Anthia</a>) employ the same chemicals as ants: <a href="/wiki/Formic_acid" title="Formic acid">formic acid</a>.<a href="#cite_note-Evans-121">[120]</a> <a href="/wiki/Bombardier_beetle" title="Bombardier beetle">Bombardier beetles</a> have well-developed pygidial glands that empty from the sides of the intersegment membranes between the seventh and eighth abdominal segments. The gland is made of two containing chambers, one for <a href="/wiki/Hydroquinone" title="Hydroquinone">hydroquinones</a> and <a href="/wiki/Hydrogen_peroxide" title="Hydrogen peroxide">hydrogen peroxide</a>, the other holding hydrogen peroxide and <a href="/wiki/Catalase" title="Catalase">catalase</a> enzymes. These chemicals mix and result in an explosive ejection, reaching a temperature of around 100 °C (212 °F), with the breakdown of hydroquinone to hydrogen, oxygen, and quinone. The oxygen propels the noxious chemical spray as a jet that can be aimed accurately at predators.<a href="#cite_note-insenc-8">[8]</a> <div class="mw-heading mw-heading4"><h4 id="Other_defenses">Other defenses</h4>[<a href="/w/index.php?title=Beetle&action=edit&section=37" title="Edit section: Other defenses">edit</a>]</div> Large ground-dwelling beetles such as <a href="/wiki/Carabidae" class="mw-redirect" title="Carabidae">Carabidae</a>, the <a href="/wiki/Rhinoceros_beetle" class="mw-redirect" title="Rhinoceros beetle">rhinoceros beetle</a> and the longhorn beetles defend themselves using strong <a href="/wiki/Mandible_(insect)" class="mw-redirect" title="Mandible (insect)">mandibles</a>, or heavily sclerotised (armored) spines or horns to deter or fight off predators.<a href="#cite_note-Evans126-120">[119]</a> Many species of weevil that feed out in the open on leaves of plants react to attack by employing a drop-off reflex. Some combine it with <a href="/wiki/Thanatosis" class="mw-redirect" title="Thanatosis">thanatosis</a>, in which they close up their appendages and "play dead".<a href="#cite_note-insenc199-122">[121]</a> The click beetles (<a href="/wiki/Elateridae" class="mw-redirect" title="Elateridae">Elateridae</a>) can suddenly catapult themselves out of danger by releasing the energy stored by a click mechanism, which consists of a stout spine on the prosternum and a matching groove in the mesosternum.<a href="#cite_note-Evans27-118">[117]</a> Some species startle an attacker by producing sounds through a process known as <a href="/wiki/Stridulation" title="Stridulation">stridulation</a>.<a href="#cite_note-stridulation-99">[98]</a> <div class="mw-heading mw-heading3"><h3 id="Parasitism">Parasitism</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=38" title="Edit section: Parasitism">edit</a>]</div> A few species of beetles are <a href="/wiki/Parasitism#Basic_concepts" title="Parasitism">ectoparasitic</a> on mammals. One such species, <a href="/wiki/Platypsyllus_castoris" class="mw-redirect" title="Platypsyllus castoris">Platypsyllus castoris</a>, parasitises beavers (<a href="/wiki/Beaver" title="Beaver">Castor</a> spp.). This beetle lives as a parasite both as a larva and as an adult, feeding on epidermal tissue and possibly on skin secretions and wound exudates. They are strikingly flattened dorsoventrally, no doubt as an adaptation for slipping between the beavers' hairs. They are wingless and eyeless, as are many other ectoparasites.<a href="#cite_note-123">[122]</a> Others are kleptoparasites of other invertebrates, such as the <a href="/wiki/Small_hive_beetle" title="Small hive beetle">small hive beetle</a> (Aethina tumida) that infests <a href="/wiki/Honey_bee" title="Honey bee">honey bee</a> nests,<a href="#cite_note-124">[123]</a> while many species are parasitic <a href="/wiki/Inquiline" title="Inquiline">inquilines</a> or <a href="/wiki/Myrmecophily_in_Staphylinidae" title="Myrmecophily in Staphylinidae">commensal in the nests of ants</a>.<a href="#cite_note-125">[124]</a> A few groups of beetles are primary <a href="/wiki/Parasitoid" title="Parasitoid">parasitoids</a> of other insects, feeding off of, and eventually killing their hosts.<a href="#cite_note-126">[125]</a> <div class="mw-heading mw-heading3"><h3 id="Pollination">Pollination</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=39" title="Edit section: Pollination">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Protaetia_cuprea_-_2019_Zachi_Evenor.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Protaetia_cuprea_-_2019_Zachi_Evenor.jpg/220px-Protaetia_cuprea_-_2019_Zachi_Evenor.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Protaetia_cuprea_-_2019_Zachi_Evenor.jpg/330px-Protaetia_cuprea_-_2019_Zachi_Evenor.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/f/fa/Protaetia_cuprea_-_2019_Zachi_Evenor.jpg/440px-Protaetia_cuprea_-_2019_Zachi_Evenor.jpg 2x" data-file-width="1210" data-file-height="1210" /></a><figcaption>An Israeli Copper Flower-Chafer (<a href="/wiki/Protaetia_cuprea" title="Protaetia cuprea">Protaetia cuprea</a> ignicollis) on a crown daisy (<a href="/wiki/Glebionis_coronaria" title="Glebionis coronaria">Glebionis coronaria</a>)</figcaption></figure> Beetle-pollinated flowers are usually large, greenish or off-white in color, and heavily scented. Scents may be spicy, fruity, or similar to decaying organic material. Beetles were most likely the first insects to pollinate flowers.<a href="#cite_note-127">[126]</a> Most beetle-pollinated flowers are flattened or dish-shaped, with pollen easily accessible, although they may include <a href="/wiki/Pollination_trap" title="Pollination trap">traps</a> to keep the beetle longer. The plants' ovaries are usually well protected from the biting mouthparts of their pollinators. The beetle families that habitually pollinate flowers are the <a href="/wiki/Buprestidae" title="Buprestidae">Buprestidae</a>, <a href="/wiki/Cantharidae" class="mw-redirect" title="Cantharidae">Cantharidae</a>, <a href="/wiki/Cerambycidae" class="mw-redirect" title="Cerambycidae">Cerambycidae</a>, <a href="/wiki/Cleridae" title="Cleridae">Cleridae</a>, <a href="/wiki/Dermestidae" title="Dermestidae">Dermestidae</a>, <a href="/wiki/Lycidae" title="Lycidae">Lycidae</a>, <a href="/wiki/Melyridae" title="Melyridae">Melyridae</a>, <a href="/wiki/Mordellidae" title="Mordellidae">Mordellidae</a>, <a href="/wiki/Nitidulidae" class="mw-redirect" title="Nitidulidae">Nitidulidae</a> and <a href="/wiki/Scarabaeidae" title="Scarabaeidae">Scarabaeidae</a>.<a href="#cite_note-Gullan-128">[127]</a> Beetles may be particularly important in some parts of the world such as semiarid areas of southern Africa and <a href="/wiki/Southern_California" title="Southern California">southern California</a><a href="#cite_note-129">[128]</a> and the montane grasslands of <a href="/wiki/KwaZulu-Natal" title="KwaZulu-Natal">KwaZulu-Natal</a> in South Africa.<a href="#cite_note-Ollerton_a-130">[129]</a> <div class="mw-heading mw-heading3"><h3 id="Mutualism">Mutualism</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=40" title="Edit section: Mutualism">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Ambrosia_beetle_life_cycle.svg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Ambrosia_beetle_life_cycle.svg/260px-Ambrosia_beetle_life_cycle.svg.png" decoding="async" width="260" height="135" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/72/Ambrosia_beetle_life_cycle.svg/390px-Ambrosia_beetle_life_cycle.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/72/Ambrosia_beetle_life_cycle.svg/520px-Ambrosia_beetle_life_cycle.svg.png 2x" data-file-width="906" data-file-height="469" /></a><figcaption> 1: Adult <a href="/wiki/Ambrosia_beetle" title="Ambrosia beetle">ambrosia beetle</a> burrows into wood and lays eggs, carrying fungal spores in its <a href="/wiki/Mycangia" class="mw-redirect" title="Mycangia">mycangia</a>. 2: Larva feeds on fungus, which digests wood, removing toxins, to mutual benefit. 3: Larva pupates.</figcaption></figure> <a href="/wiki/Mutualism_(biology)" title="Mutualism (biology)">Mutualism</a> is well known in a few beetles, such as the <a href="/wiki/Ambrosia_beetle" title="Ambrosia beetle">ambrosia beetle</a>, which partners with fungi to digest the wood of dead trees. The beetles excavate tunnels in dead trees in which they cultivate fungal gardens, their sole source of nutrition. After landing on a suitable tree, an ambrosia beetle excavates a tunnel in which it releases spores of its fungal <a href="/wiki/Symbiont" class="mw-redirect" title="Symbiont">symbiont</a>. The fungus penetrates the plant's xylem tissue, digests it, and concentrates the nutrients on and near the surface of the beetle gallery, so the weevils and the fungus both benefit. The beetles cannot eat the wood due to toxins, and uses its relationship with fungi to help overcome the defenses of its host tree in order to provide nutrition for their larvae.<a href="#cite_note-Malloch-131">[130]</a> Chemically mediated by a bacterially produced polyunsaturated peroxide,<a href="#cite_note-Scott2008-132">[131]</a> this mutualistic relationship between the beetle and the fungus is <a href="/wiki/Coevolve" class="mw-redirect" title="Coevolve">coevolved</a>.<a href="#cite_note-Malloch-131">[130]</a><a href="#cite_note-133">[132]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Bug_in_Tharparkar_District.jpg" class="mw-file-description"><img alt="Beetle found in Tharparkar District" src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Bug_in_Tharparkar_District.jpg/220px-Bug_in_Tharparkar_District.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Bug_in_Tharparkar_District.jpg/330px-Bug_in_Tharparkar_District.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Bug_in_Tharparkar_District.jpg/440px-Bug_in_Tharparkar_District.jpg 2x" data-file-width="4160" data-file-height="3120" /></a><figcaption>Tenebrionid beetle in the <a href="/wiki/Tharparkar" title="Tharparkar">Thar Desert</a></figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Tolerance_of_extreme_environments">Tolerance of extreme environments</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=41" title="Edit section: Tolerance of extreme environments">edit</a>]</div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Stenocara_gracilipes.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/Stenocara_gracilipes.jpg/220px-Stenocara_gracilipes.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/8/8e/Stenocara_gracilipes.jpg/330px-Stenocara_gracilipes.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/8/8e/Stenocara_gracilipes.jpg/440px-Stenocara_gracilipes.jpg 2x" data-file-width="2648" data-file-height="1986" /></a><figcaption>The fogstand beetle of the <a href="/wiki/Namib_Desert" class="mw-redirect" title="Namib Desert">Namib Desert</a>, <a href="/wiki/Stenocara_gracilipes" title="Stenocara gracilipes">Stenocara gracilipes</a>, is able to survive by <a href="/wiki/Fog_collection" title="Fog collection">collecting water from fog</a> on its back.</figcaption></figure> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Insect_thermoregulation" title="Insect thermoregulation">Insect thermoregulation</a> and <a href="/wiki/Insect_winter_ecology" title="Insect winter ecology">Insect winter ecology</a></div> About 90% of beetle species enter a period of adult <a href="/wiki/Diapause" title="Diapause">diapause</a>, a quiet phase with reduced metabolism to tide unfavourable environmental conditions. Adult diapause is the most common form of diapause in Coleoptera. To endure the period without food (often lasting many months) adults prepare by accumulating reserves of lipids, glycogen, proteins and other substances needed for resistance to future hazardous changes of environmental conditions. This diapause is induced by signals heralding the arrival of the unfavourable season; usually the cue is <a href="/wiki/Photoperiodic" class="mw-redirect" title="Photoperiodic">photoperiodic</a>. Short (decreasing) day length serves as a signal of approaching winter and induces winter diapause (hibernation).<a href="#cite_note-134">[133]</a> A study of hibernation in the Arctic beetle <a href="/wiki/Pterostichus_brevicornis" title="Pterostichus brevicornis">Pterostichus brevicornis</a> showed that the body fat levels of adults were highest in autumn with the <a href="/wiki/Alimentary_canal" class="mw-redirect" title="Alimentary canal">alimentary canal</a> filled with food, but empty by the end of January. This loss of body fat was a gradual process, occurring in combination with dehydration.<a href="#cite_note-135">[134]</a> All insects are <a href="/wiki/Poikilotherm" title="Poikilotherm">poikilothermic</a>,<a href="#cite_note-Outline-136">[135]</a> so the ability of a few beetles to live in extreme environments depends on their resilience to unusually high or low temperatures. The <a href="/wiki/Bark_beetle" title="Bark beetle">bark beetle</a> <a href="/w/index.php?title=Pityogenes_chalcographus&action=edit&redlink=1" class="new" title="Pityogenes chalcographus (page does not exist)">Pityogenes chalcographus</a> can survive −39°C whilst overwintering beneath tree bark;<a href="#cite_note-137">[136]</a> the Alaskan beetle <a href="/wiki/Cucujus" title="Cucujus">Cucujus</a> clavipes puniceus is able to withstand −58°C; its larvae may survive −100°C.<a href="#cite_note-138">[137]</a> At these low temperatures, the formation of ice crystals in internal fluids is the biggest threat to survival to beetles, but this is prevented through the production of antifreeze proteins that stop water molecules from grouping together. The low temperatures experienced by Cucujus clavipes can be survived through their deliberate dehydration in conjunction with the antifreeze proteins. This concentrates the antifreezes several fold.<a href="#cite_note-139">[138]</a> The <a href="/wiki/Hemolymph" title="Hemolymph">hemolymph</a> of the mealworm beetle <a href="/wiki/Tenebrio_molitor" class="mw-redirect" title="Tenebrio molitor">Tenebrio molitor</a> contains several <a href="/wiki/Antifreeze_protein" title="Antifreeze protein">antifreeze proteins</a>.<a href="#cite_note-140">[139]</a> The Alaskan beetle <a href="/wiki/Upis_ceramboides" title="Upis ceramboides">Upis ceramboides</a> can survive −60 °C: its <a href="/wiki/Cryoprotectant" title="Cryoprotectant">cryoprotectants</a> are <a href="/wiki/Xylomannan" title="Xylomannan">xylomannan</a>, a molecule consisting of a <a href="/wiki/Sugar" title="Sugar">sugar</a> bound to a <a href="/wiki/Fatty_acid" title="Fatty acid">fatty acid</a>,<a href="#cite_note-141">[140]</a> and the sugar-alcohol, <a href="/wiki/Threitol" title="Threitol">threitol</a>.<a href="#cite_note-142">[141]</a> Conversely, desert dwelling beetles are adapted to tolerate high temperatures. For example, the <a href="/wiki/Tenebrionid" class="mw-redirect" title="Tenebrionid">Tenebrionid</a> beetle <a href="/w/index.php?title=Onymacris_rugatipennis&action=edit&redlink=1" class="new" title="Onymacris rugatipennis (page does not exist)">Onymacris rugatipennis</a> can withstand 50°C.<a href="#cite_note-143">[142]</a> Tiger beetles in hot, sandy areas are often whitish (for example, <a href="/wiki/Habroscelimorpha_dorsalis" title="Habroscelimorpha dorsalis">Habroscelimorpha dorsalis</a>), to reflect more heat than a darker color would. These beetles also exhibits behavioural adaptions to tolerate the heat: they are able to stand erect on their tarsi to hold their bodies away from the hot ground, seek shade, and turn to face the sun so that only the front parts of their heads are directly exposed.<a href="#cite_note-144">[143]</a> The fogstand beetle of the <a href="/wiki/Namib_Desert" class="mw-redirect" title="Namib Desert">Namib Desert</a>, <a href="/wiki/Stenocara_gracilipes" title="Stenocara gracilipes">Stenocara gracilipes</a>, is able to <a href="/wiki/Fog_collection" title="Fog collection">collect water from fog</a>, as its elytra have a textured surface combining <a href="/wiki/Hydrophilic" class="mw-redirect" title="Hydrophilic">hydrophilic</a> (water-loving) bumps and waxy, <a href="/wiki/Hydrophobic" class="mw-redirect" title="Hydrophobic">hydrophobic</a> troughs. The beetle faces the early morning breeze, holding up its abdomen; droplets condense on the elytra and run along ridges towards their mouthparts. Similar adaptations are found in several other Namib desert beetles such as <a href="/wiki/Onymacris_unguicularis" title="Onymacris unguicularis">Onymacris unguicularis</a>.<a href="#cite_note-145">[144]</a> Some terrestrial beetles that exploit shoreline and floodplain habitats have physiological adaptations for surviving floods. In the event of flooding, adult beetles may be mobile enough to move away from flooding, but larvae and pupa often cannot. Adults of <a href="/wiki/Cicindela_togata" class="mw-redirect" title="Cicindela togata">Cicindela togata</a> are unable to survive immersion in water, but larvae are able to survive a prolonged period, up to 6 days, of <a href="/wiki/Hypoxia_(environmental)" title="Hypoxia (environmental)">anoxia</a> during floods. Anoxia tolerance in the larvae may have been sustained by switching to anaerobic metabolic pathways or by reducing metabolic rate.<a href="#cite_note-146">[145]</a> Anoxia tolerance in the adult carabid beetle <a href="/w/index.php?title=Pelophilia_borealis&action=edit&redlink=1" class="new" title="Pelophilia borealis (page does not exist)">Pelophilia borealis</a> was tested in laboratory conditions and it was found that they could survive a continuous period of up to 127 days in an atmosphere of 99.9% nitrogen at 0 °C.<a href="#cite_note-147">[146]</a> <div class="mw-heading mw-heading3"><h3 id="Migration">Migration</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=42" title="Edit section: Migration">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Insect_migration" title="Insect migration">Insect migration</a></div> Many beetle species undertake annual mass movements which are termed as migrations. These include the pollen beetle <a href="/wiki/Meligethes_aeneus" class="mw-redirect" title="Meligethes aeneus">Meligethes aeneus</a><a href="#cite_note-148">[147]</a> and many species of <a href="/wiki/Coccinellidae" title="Coccinellidae">coccinellids</a>.<a href="#cite_note-149">[148]</a> These mass movements may also be opportunistic, in search of food, rather than seasonal. A 2008 study of an unusually large outbreak of Mountain Pine Beetle (<a href="/wiki/Dendroctonus_ponderosae" class="mw-redirect" title="Dendroctonus ponderosae">Dendroctonus ponderosae</a>) in <a href="/wiki/British_Columbia" title="British Columbia">British Columbia</a> found that beetles were capable of flying 30–110 km per day in densities of up to 18,600 beetles per hectare.<a href="#cite_note-150">[149]</a> <div class="mw-heading mw-heading2"><h2 id="Relationship_to_humans">Relationship to humans</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=43" title="Edit section: Relationship to humans">edit</a>]</div> <div class="mw-heading mw-heading3"><h3 id="In_ancient_cultures">In ancient cultures</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=44" title="Edit section: In ancient cultures">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main article: <a href="/wiki/Scarab_(artifact)" title="Scarab (artifact)">Scarab (artifact)</a></div> <table style="font-size: 88%; border-collapse: collapse; border: 1px solid blue; margin: 0.5em 1em; text-align: center; clear: left; float: left; margin-left: 0;"> <tbody><tr> <td style="padding:0.75em; background-color: var(--background-color-base); color:var( --color-base ); text-align: center"><table class="mw-hiero-table mw-hiero-outer" dir="ltr"><tbody><tr><td> <table class="mw-hiero-table"><tbody><tr> <td><img class="skin-invert" style="margin: 1px;" src="/w/extensions/wikihiero/img/hiero_L1.png?8b41f" height="36" title="L1 [xpr]" alt="xpr" /></td></tr></tbody></table> </td></tr></tbody></table> </td></tr> <tr> <th style="background: navajowhite; color:black; border-bottom: 1px solid peru; padding: 0.5em">Scarabee in <a href="/wiki/Egyptian_hieroglyphs" title="Egyptian hieroglyphs">hieroglyphs</a> </th></tr> <tr> <td> </td></tr> <tr> <td><a href="/wiki/Gardiner%27s_sign_list" title="Gardiner's sign list">Gardiner</a>: L1 </td></tr> <tr> <td> </td></tr> </tbody></table> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Egypt.KV6.04.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Egypt.KV6.04.jpg/220px-Egypt.KV6.04.jpg" decoding="async" width="220" height="220" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/b/be/Egypt.KV6.04.jpg/330px-Egypt.KV6.04.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/b/be/Egypt.KV6.04.jpg 2x" data-file-width="400" data-file-height="400" /></a><figcaption>A scarab in the <a href="/wiki/Valley_of_the_Kings" title="Valley of the Kings">Valley of the Kings</a></figcaption></figure> Several species of dung beetle, especially the sacred scarab, <a href="/wiki/Scarabaeus_sacer" title="Scarabaeus sacer">Scarabaeus sacer</a>, were revered in <a href="/wiki/Ancient_Egypt" title="Ancient Egypt">Ancient Egypt</a>.<a href="#cite_note-egyptian-151">[150]</a><a href="#cite_note-152">[151]</a> The hieroglyphic image of the beetle may have had existential, fictional, or ontologic significance.<a href="#cite_note-153">[152]</a> Images of the scarab in bone, <a href="/wiki/Ivory" title="Ivory">ivory</a>, stone, <a href="/wiki/Egyptian_faience" title="Egyptian faience">Egyptian faience</a>, and precious metals are known from the Sixth Dynasty and up to the period of Roman rule. The scarab was of prime significance in the funerary cult of ancient Egypt.<a href="#cite_note-154">[153]</a> The scarab was linked to <a href="/wiki/Khepri" title="Khepri">Khepri</a>, the god of the rising <a href="/wiki/Sun" title="Sun">sun</a>, from the supposed resemblance of the rolling of the dung ball by the beetle to the rolling of the sun by the god.<a href="#cite_note-egyptian-151">[150]</a> Some of ancient Egypt's neighbors adopted the scarab motif for <a href="/wiki/Scaraboid_seal" class="mw-redirect" title="Scaraboid seal">seals</a> of varying types. The best-known of these are the Judean <a href="/wiki/LMLK_seal" title="LMLK seal">LMLK seals</a>, where eight of 21 designs contained scarab beetles, which were used exclusively to stamp impressions on storage jars during the reign of <a href="/wiki/Hezekiah" title="Hezekiah">Hezekiah</a>.<a href="#cite_note-155">[154]</a> Beetles are mentioned as a symbol of the sun, as in ancient Egypt, in <a href="/wiki/Plutarch" title="Plutarch">Plutarch</a>'s 1st century <a href="/wiki/Moralia" title="Moralia">Moralia</a>.<a href="#cite_note-156">[155]</a> The <a href="/wiki/Greek_Magical_Papyri" title="Greek Magical Papyri">Greek Magical Papyri</a> of the 2nd century BC to the 5th century AD describe scarabs as an ingredient in a spell.<a href="#cite_note-157">[156]</a> <a href="/wiki/Pliny_the_Elder" title="Pliny the Elder">Pliny the Elder</a> discusses beetles in his <a href="/wiki/Natural_History_(Pliny)" title="Natural History (Pliny)">Natural History</a>,<a href="#cite_note-PlinyXI-158">[157]</a> describing the <a href="/wiki/Lucanus_cervus" title="Lucanus cervus">stag beetle</a>: "Some insects, for the preservation of their wings, are covered with an erust (<a href="/wiki/Elytra" class="mw-redirect" title="Elytra">elytra</a>)—the beetle, for instance, the wing of which is peculiarly fine and frail. To these insects a sting has been denied by Nature; but in one large kind we find horns of a remarkable length, two-pronged at the extremities, and forming pincers, which the animal closes when it is its intention to bite."<a href="#cite_note-PlinyXI_Book34-159">[158]</a><a href="#cite_note-Beavis-160">[159]</a> The stag beetle is recorded in a Greek myth by <a href="/wiki/Nicander" title="Nicander">Nicander</a> and recalled by <a href="/wiki/Antoninus_Liberalis" title="Antoninus Liberalis">Antoninus Liberalis</a> in which <a href="/wiki/Cerambus" title="Cerambus">Cerambus</a><a href="#cite_note-161">[b]</a> is turned into a beetle: "He can be seen on trunks and has hook-teeth, ever moving his jaws together. He is black, long and has hard wings like a great dung beetle".<a href="#cite_note-162">[160]</a> The story concludes with the comment that the beetles were used as toys by young boys, and that the head was removed and worn as a pendant.<a href="#cite_note-Beavis-160">[159]</a><a href="#cite_note-163">[161]</a> <div class="mw-heading mw-heading3"><h3 id="As_pests">As pests</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=45" title="Edit section: As pests">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Boll_weevil.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/26/Boll_weevil.jpg/220px-Boll_weevil.jpg" decoding="async" width="220" height="168" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/2/26/Boll_weevil.jpg/330px-Boll_weevil.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/2/26/Boll_weevil.jpg/440px-Boll_weevil.jpg 2x" data-file-width="2800" data-file-height="2137" /></a><figcaption><a href="/wiki/Cotton_boll_weevil" class="mw-redirect" title="Cotton boll weevil">Cotton boll weevil</a></figcaption></figure> About 75% of beetle species are phytophagous in both the larval and adult stages. Many feed on economically important plants and stored plant products, including trees, cereals, tobacco, and dried fruits.<a href="#cite_note-Gilliott-7">[7]</a> Some, such as the <a href="/wiki/Boll_weevil" title="Boll weevil">boll weevil</a>, which feeds on cotton buds and flowers, can cause extremely serious damage to agriculture. The boll weevil crossed the <a href="/wiki/Rio_Grande" title="Rio Grande">Rio Grande</a> near <a href="/wiki/Brownsville,_Texas" title="Brownsville, Texas">Brownsville</a>, <a href="/wiki/Texas" title="Texas">Texas</a>, to enter the <a href="/wiki/United_States" title="United States">United States</a> from <a href="/wiki/Mexico" title="Mexico">Mexico</a> around 1892,<a href="#cite_note-msstate-164">[162]</a> and had reached southeastern <a href="/wiki/Alabama" title="Alabama">Alabama</a> by 1915. By the mid-1920s, it had entered all cotton-growing regions in the US, traveling 40 to 160 miles (60–260 km) per year. It remains the most destructive cotton pest in North America. <a href="/wiki/Mississippi_State_University" title="Mississippi State University">Mississippi State University</a> has estimated, since the boll weevil entered the United States, it has cost cotton producers about $13 billion, and in recent times about $300 million per year.<a href="#cite_note-msstate-164">[162]</a> The <a href="/wiki/Bark_beetle" title="Bark beetle">bark beetle</a>, <a href="/wiki/Elm_leaf_beetle" title="Elm leaf beetle">elm leaf beetle</a> and the Asian longhorned beetle (<a href="/wiki/Anoplophora_glabripennis" class="mw-redirect" title="Anoplophora glabripennis">Anoplophora glabripennis</a>)<a href="#cite_note-165">[163]</a> are among the species that attack <a href="/wiki/Elm" title="Elm">elm</a> trees. Bark beetles (<a href="/wiki/Scolytidae" class="mw-redirect" title="Scolytidae">Scolytidae</a>) carry <a href="/wiki/Dutch_elm_disease" title="Dutch elm disease">Dutch elm disease</a> as they move from infected breeding sites to healthy trees. The disease has devastated elm trees across Europe and North America.<a href="#cite_note-166">[164]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Potato_beetle_larvae.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/2/2e/Potato_beetle_larvae.jpg/220px-Potato_beetle_larvae.jpg" decoding="async" width="220" height="177" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/2/2e/Potato_beetle_larvae.jpg 1.5x" data-file-width="300" data-file-height="241" /></a><figcaption>Larvae of the <a href="/wiki/Colorado_potato_beetle" title="Colorado potato beetle">Colorado potato beetle</a>, Leptinotarsa decemlineata, a serious crop pest</figcaption></figure> Some species of beetle have evolved immunity to insecticides. For example, the <a href="/wiki/Colorado_potato_beetle" title="Colorado potato beetle">Colorado potato beetle</a>, Leptinotarsa decemlineata, is a destructive pest of potato plants. Its hosts include other members of the <a href="/wiki/Solanaceae" title="Solanaceae">Solanaceae</a>, such as <a href="/wiki/Nightshade" class="mw-redirect" title="Nightshade">nightshade</a>, <a href="/wiki/Tomato" title="Tomato">tomato</a>, <a href="/wiki/Eggplant" title="Eggplant">eggplant</a> and <a href="/wiki/Capsicum" title="Capsicum">capsicum</a>, as well as the potato. Different populations have between them developed resistance to all major classes of insecticide.<a href="#cite_note-167">[165]</a> The Colorado potato beetle was evaluated as a tool of <a href="/wiki/Entomological_warfare" title="Entomological warfare">entomological warfare</a> during <a href="/wiki/World_War_II" title="World War II">World War II</a>, the idea being to use the beetle and its larvae to damage the crops of enemy nations.<a href="#cite_note-lockwoodbost-168">[166]</a> Germany tested its Colorado potato beetle weaponisation program south of <a href="/wiki/Frankfurt" title="Frankfurt">Frankfurt</a>, releasing 54,000 beetles.<a href="#cite_note-heather-169">[167]</a> The <a href="/wiki/Death_watch_beetle" class="mw-redirect" title="Death watch beetle">death watch beetle</a>, Xestobium rufovillosum (<a href="/wiki/Ptinidae" title="Ptinidae">Ptinidae</a>), is a serious pest of older wooden buildings in Europe. It attacks <a href="/wiki/Hardwood" title="Hardwood">hardwoods</a> such as <a href="/wiki/Oak" title="Oak">oak</a> and <a href="/wiki/Chestnut" title="Chestnut">chestnut</a>, always where some fungal decay has taken or is taking place. The actual introduction of the pest into buildings is thought to take place at the time of construction.<a href="#cite_note-170">[168]</a> Other pests include the coconut hispine beetle, <a href="/wiki/Brontispa_longissima" title="Brontispa longissima">Brontispa longissima</a>, which feeds on young <a href="/wiki/Leaves" class="mw-redirect" title="Leaves">leaves</a>, <a href="/wiki/Seedlings" class="mw-redirect" title="Seedlings">seedlings</a> and mature <a href="/wiki/Coconut" title="Coconut">coconut</a> trees, causing serious economic damage in the <a href="/wiki/Philippines" title="Philippines">Philippines</a>.<a href="#cite_note-171">[169]</a> The <a href="/wiki/Mountain_pine_beetle" title="Mountain pine beetle">mountain pine beetle</a> is a destructive pest of mature or weakened <a href="/wiki/Lodgepole_pine" class="mw-redirect" title="Lodgepole pine">lodgepole pine</a>, sometimes affecting large areas of Canada.<a href="#cite_note-172">[170]</a> <div class="mw-heading mw-heading3"><h3 id="As_beneficial_resources">As beneficial resources</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=46" title="Edit section: As beneficial resources">edit</a>]</div> <figure class="mw-default-size mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Lady-beetle-close-up.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/Lady-beetle-close-up.jpg/220px-Lady-beetle-close-up.jpg" decoding="async" width="220" height="152" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/57/Lady-beetle-close-up.jpg/330px-Lady-beetle-close-up.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/57/Lady-beetle-close-up.jpg/440px-Lady-beetle-close-up.jpg 2x" data-file-width="1305" data-file-height="903" /></a><figcaption><a href="/wiki/Coccinella_septempunctata" title="Coccinella septempunctata">Coccinella septempunctata</a>, a predatory beetle beneficial to agriculture</figcaption></figure> Beetles can be beneficial to human economics by controlling the populations of pests. The larvae and adults of some species of <a href="/wiki/Lady_beetle" class="mw-redirect" title="Lady beetle">lady beetles</a> (<a href="/wiki/Coccinellidae" title="Coccinellidae">Coccinellidae</a>) feed on <a href="/wiki/Aphid" title="Aphid">aphids</a> that are pests. Other lady beetles feed on <a href="/wiki/Scale_insect" title="Scale insect">scale insects</a>, <a href="/wiki/Whitefly" title="Whitefly">whitefly</a> and <a href="/wiki/Mealybug" title="Mealybug">mealybugs</a>.<a href="#cite_note-173">[171]</a> If normal food sources are scarce, they may feed on small <a href="/wiki/Caterpillar" title="Caterpillar">caterpillars</a>, young <a href="/wiki/Plant_bug" class="mw-redirect" title="Plant bug">plant bugs</a>, or <a href="/wiki/Honeydew_(secretion)" title="Honeydew (secretion)">honeydew</a> and <a href="/wiki/Nectar" title="Nectar">nectar</a>.<a href="#cite_note-BBC_news-174">[172]</a> <a href="/wiki/Ground_beetle" title="Ground beetle">Ground beetles</a> (Carabidae) are common <a href="/wiki/Predator" class="mw-redirect" title="Predator">predators</a> of many insect pests, including fly eggs, caterpillars, and wireworms.<a href="#cite_note-175">[173]</a> Ground beetles can help to control <a href="/wiki/Weed" title="Weed">weeds</a> by eating their seeds in the soil, reducing the need for <a href="/wiki/Herbicide" title="Herbicide">herbicides</a> to protect crops.<a href="#cite_note-176">[174]</a> The effectiveness of some species in reducing certain plant populations has resulted in the deliberate introduction of beetles in order to control weeds. For example, the genus <a href="/wiki/Calligrapha" title="Calligrapha">Calligrapha</a> is native to North America but has been used to control <a href="/wiki/Parthenium_hysterophorus" title="Parthenium hysterophorus">Parthenium hysterophorus</a> in India and <a href="/wiki/Ambrosia_artemisiifolia" title="Ambrosia artemisiifolia">Ambrosia artemisiifolia</a> in Russia.<a href="#cite_note-ISC-177">[175]</a><a href="#cite_note-178">[176]</a> <a href="/wiki/Dung_beetles" class="mw-redirect" title="Dung beetles">Dung beetles</a> (Scarabidae) have been successfully used to reduce the populations of pestilent flies, such as <a href="/wiki/Musca_vetustissima" title="Musca vetustissima">Musca vetustissima</a> and <a href="/wiki/Haematobia_exigua" title="Haematobia exigua">Haematobia exigua</a> which are serious pests of cattle in <a href="/wiki/Australia" title="Australia">Australia</a>.<a href="#cite_note-Bornemissza-179">[177]</a> The beetles make the dung unavailable to breeding pests by quickly rolling and burying it in the soil, with the added effect of improving soil fertility, tilth, and nutrient cycling.<a href="#cite_note-180">[178]</a> The <a href="/wiki/Australian_Dung_Beetle_Project" title="Australian Dung Beetle Project">Australian Dung Beetle Project</a> (1965–1985), introduced species of dung beetle to Australia from South Africa and Europe to reduce populations of Musca vetustissima, following successful trials of this technique in <a href="/wiki/Hawaii" title="Hawaii">Hawaii</a>.<a href="#cite_note-Bornemissza-179">[177]</a> The <a href="/wiki/American_Institute_of_Biological_Sciences" title="American Institute of Biological Sciences">American Institute of Biological Sciences</a> reports that dung beetles, such as <a href="/wiki/Euoniticellus_intermedius" title="Euoniticellus intermedius">Euoniticellus intermedius</a>, save the United States cattle industry an estimated US$380 million annually through burying above-ground livestock feces.<a href="#cite_note-181">[179]</a> The <a href="/wiki/Dermestidae" title="Dermestidae">Dermestidae</a> are often used in <a href="/wiki/Taxidermy" title="Taxidermy">taxidermy</a> and in the preparation of scientific specimens, to clean soft tissue from bones.<a href="#cite_note-182">[180]</a> Larvae feed on and remove cartilage along with other soft tissue.<a href="#cite_note-183">[181]</a><a href="#cite_note-184">[182]</a> <div class="mw-heading mw-heading3"><h3 id="As_food_and_medicine">As food and medicine</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=47" title="Edit section: As food and medicine">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">See also: <a href="/wiki/Entomophagy" title="Entomophagy">Entomophagy</a> and <a href="/wiki/Insects_as_food" title="Insects as food">Insects as food</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Mealworm_01_Pengo.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Mealworm_01_Pengo.jpg/220px-Mealworm_01_Pengo.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Mealworm_01_Pengo.jpg/330px-Mealworm_01_Pengo.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/ad/Mealworm_01_Pengo.jpg/440px-Mealworm_01_Pengo.jpg 2x" data-file-width="3895" data-file-height="2608" /></a><figcaption>Mealworms in a bowl for <a href="/wiki/Entomophagy" title="Entomophagy">human consumption</a></figcaption></figure> Beetles are the most widely eaten insects, with about 344 species used as food, usually at the larval stage.<a href="#cite_note-185">[183]</a> The <a href="/wiki/Mealworm" title="Mealworm">mealworm</a> (the larva of the <a href="/wiki/Darkling_beetle" title="Darkling beetle">darkling beetle</a>) and the <a href="/wiki/Rhinoceros_beetle" class="mw-redirect" title="Rhinoceros beetle">rhinoceros beetle</a> are among the species commonly eaten.<a href="#cite_note-186">[184]</a> A wide range of species is also used in <a href="/wiki/Folk_medicine" class="mw-redirect" title="Folk medicine">folk medicine</a> to treat those suffering from a variety of disorders and illnesses, though this is done without clinical studies supporting the efficacy of such treatments.<a href="#cite_note-Meyer-Rochow-187">[185]</a> <div class="mw-heading mw-heading3"><h3 id="As_biodiversity_indicators">As biodiversity indicators</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=48" title="Edit section: As biodiversity indicators">edit</a>]</div> Due to their habitat specificity, many species of beetles have been suggested as suitable as indicators, their presence, numbers, or absence providing a measure of habitat quality. Predatory beetles such as the tiger beetles (<a href="/wiki/Cicindelidae" class="mw-redirect" title="Cicindelidae">Cicindelidae</a>) have found scientific use as an indicator taxon for measuring regional patterns of biodiversity. They are suitable for this as their taxonomy is stable; their life history is well described; they are large and simple to observe when visiting a site; they occur around the world in many habitats, with species specialised to particular habitats; and their occurrence by species accurately indicates other species, both vertebrate and invertebrate.<a href="#cite_note-188">[186]</a> According to the habitats, many other groups such as the rove beetles in human-modified habitats, dung beetles in savannas<a href="#cite_note-189">[187]</a> and saproxylic beetles in forests<a href="#cite_note-190">[188]</a> have been suggested as potential indicator species.<a href="#cite_note-191">[189]</a> <div class="mw-heading mw-heading3"><h3 id="In_art_and_adornment">In art and adornment</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=49" title="Edit section: In art and adornment">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Main articles: <a href="/wiki/Beetlewing" title="Beetlewing">Beetlewing</a> and <a href="/wiki/Live_insect_jewelry" title="Live insect jewelry">Live insect jewelry</a></div> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Zopheridae_jewelry_sjh.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Zopheridae_jewelry_sjh.jpg/220px-Zopheridae_jewelry_sjh.jpg" decoding="async" width="220" height="178" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/0/05/Zopheridae_jewelry_sjh.jpg/330px-Zopheridae_jewelry_sjh.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/0/05/Zopheridae_jewelry_sjh.jpg/440px-Zopheridae_jewelry_sjh.jpg 2x" data-file-width="1572" data-file-height="1272" /></a><figcaption><a href="/wiki/Zopheridae" title="Zopheridae">Zopheridae</a> in <a href="/wiki/Jewellery" title="Jewellery">jewellery</a> at the <a href="/wiki/Texas_A%26M_University" title="Texas A&M University">Texas A&M University</a> Insect Collection</figcaption></figure> <figure class="mw-halign-left" typeof="mw:File/Thumb"><a href="/wiki/File:Pendant_watch_in_shape_of_beetle_Switzerland_1850-1900_gold,_diamond,_enamel.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Pendant_watch_in_shape_of_beetle_Switzerland_1850-1900_gold%2C_diamond%2C_enamel.jpg/139px-Pendant_watch_in_shape_of_beetle_Switzerland_1850-1900_gold%2C_diamond%2C_enamel.jpg" decoding="async" width="139" height="225" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Pendant_watch_in_shape_of_beetle_Switzerland_1850-1900_gold%2C_diamond%2C_enamel.jpg/208px-Pendant_watch_in_shape_of_beetle_Switzerland_1850-1900_gold%2C_diamond%2C_enamel.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Pendant_watch_in_shape_of_beetle_Switzerland_1850-1900_gold%2C_diamond%2C_enamel.jpg/277px-Pendant_watch_in_shape_of_beetle_Switzerland_1850-1900_gold%2C_diamond%2C_enamel.jpg 2x" data-file-width="593" data-file-height="961" /></a><figcaption>Pendant watch in shape of beetle, Switzerland 1850–1900 gold, diamond, enamel</figcaption></figure> Many beetles have durable elytra that has been used as material in art, with <a href="/wiki/Beetlewing" title="Beetlewing">beetlewing</a> the best example.<a href="#cite_note-192">[190]</a> Sometimes, they are incorporated into ritual objects for their religious significance. Whole beetles, either as-is or encased in clear plastic, are made into objects ranging from cheap souvenirs such as key chains to expensive fine-art jewellery. In parts of Mexico, beetles of the genus <a href="/wiki/Zopherus" title="Zopherus">Zopherus</a> are made into <a href="/wiki/Living_brooch" class="mw-redirect" title="Living brooch">living brooches</a> by attaching costume jewelry and golden chains, which is made possible by the incredibly hard elytra and sedentary habits of the genus.<a href="#cite_note-Ivie-193">[191]</a> <div class="mw-heading mw-heading3"><h3 id="In_entertainment">In entertainment</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=50" title="Edit section: In entertainment">edit</a>]</div> Fighting beetles are used <a href="/wiki/Insect_fighting" title="Insect fighting">for entertainment and gambling</a>. This sport exploits the territorial behavior and mating competition of certain species of large beetles. In the <a href="/wiki/Chiang_Mai" title="Chiang Mai">Chiang Mai</a> district of northern Thailand, male <a href="/wiki/Xylotrupes" title="Xylotrupes">Xylotrupes</a> rhinoceros beetles are caught in the wild and trained for fighting. Females are held inside a log to stimulate the fighting males with their pheromones.<a href="#cite_note-194">[192]</a> These fights may be competitive and involve gambling both money and property.<a href="#cite_note-Costa-Neto-195">[193]</a> In <a href="/wiki/South_Korea" title="South Korea">South Korea</a> the <a href="/wiki/Dytiscidae" title="Dytiscidae">Dytiscidae</a> species <a href="/wiki/Cybister_tripunctatus" title="Cybister tripunctatus">Cybister tripunctatus</a> is used in a roulette-like game.<a href="#cite_note-196">[194]</a> Beetles are sometimes used as instruments: the Onabasulu of <a href="/wiki/Papua_New_Guinea" title="Papua New Guinea">Papua New Guinea</a> historically used the "<a href="/wiki/Hugu_(instrument)" title="Hugu (instrument)">hugu</a>" weevil <a href="/wiki/Rhynchophorus_ferrugineus" title="Rhynchophorus ferrugineus">Rhynchophorus ferrugineus</a> as a musical instrument by letting the human mouth serve as a variable <a href="/wiki/Resonance_chamber" title="Resonance chamber">resonance chamber</a> for the wing vibrations of the live adult beetle.<a href="#cite_note-Costa-Neto-195">[193]</a> <div class="mw-heading mw-heading3"><h3 id="As_pets">As pets</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=51" title="Edit section: As pets">edit</a>]</div> Some species of beetle are kept as <a href="/wiki/Pet" title="Pet">pets</a>, for example diving beetles (<a href="/wiki/Dytiscidae" title="Dytiscidae">Dytiscidae</a>) may be kept in a domestic fresh water tank.<a href="#cite_note-197">[195]</a> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Malay_Archipelago_Beetles.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Malay_Archipelago_Beetles.jpg/220px-Malay_Archipelago_Beetles.jpg" decoding="async" width="220" height="158" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/58/Malay_Archipelago_Beetles.jpg/330px-Malay_Archipelago_Beetles.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/58/Malay_Archipelago_Beetles.jpg/440px-Malay_Archipelago_Beetles.jpg 2x" data-file-width="1782" data-file-height="1282" /></a><figcaption>"Remarkable Beetles Found at Simunjon, Borneo".<a href="#cite_note-198">[c]</a> A few of the 2,000 species of beetle collected by <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> in <a href="/wiki/Borneo" title="Borneo">Borneo</a></figcaption></figure> In <a href="/wiki/Japan" title="Japan">Japan</a> the practice of keeping horned rhinoceros beetles (<a href="/wiki/Dynastinae" title="Dynastinae">Dynastinae</a>) and stag beetles (<a href="/wiki/Lucanidae" class="mw-redirect" title="Lucanidae">Lucanidae</a>) is particularly popular amongst young boys.<a href="#cite_note-199">[196]</a> Such is the popularity in Japan that <a href="/wiki/Vending_machines" class="mw-redirect" title="Vending machines">vending machines</a> dispensing live beetles were developed in 1999, each holding up to 100 stag beetles.<a href="#cite_note-200">[197]</a><a href="#cite_note-201">[198]</a> <div class="mw-heading mw-heading3"><h3 id="As_things_to_collect">As things to collect</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=52" title="Edit section: As things to collect">edit</a>]</div> Beetle collecting became extremely popular in the <a href="/wiki/Victorian_era" title="Victorian era">Victorian era</a>.<a href="#cite_note-202">[199]</a> The naturalist <a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a> collected (by his own count) a total of 83,200 beetles during the eight years described in his 1869 book <a href="/wiki/The_Malay_Archipelago" title="The Malay Archipelago">The Malay Archipelago</a>, including 2,000 species new to science.<a href="#cite_note-203">[200]</a> <div class="mw-heading mw-heading3"><h3 id="As_inspiration_for_technologies">As inspiration for technologies</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=53" title="Edit section: As inspiration for technologies">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1236090951"><div role="note" class="hatnote navigation-not-searchable">Further information: <a href="/wiki/Biomimetics" title="Biomimetics">Biomimetics</a></div> Several coleopteran adaptations have attracted interest in <a href="/wiki/Biomimetics" title="Biomimetics">biomimetics</a> with possible commercial applications. The <a href="/wiki/Bombardier_beetle" title="Bombardier beetle">bombardier beetle</a>'s powerful repellent spray has inspired the development of a fine mist spray technology, claimed to have a low carbon impact compared to aerosol sprays.<a href="#cite_note-204">[201]</a> Moisture harvesting behavior by the Namib desert beetle (<a href="/wiki/Stenocara_gracilipes" title="Stenocara gracilipes">Stenocara gracilipes</a>) has inspired a self-filling water bottle which utilises <a href="/wiki/Hydrophilic" class="mw-redirect" title="Hydrophilic">hydrophilic</a> and <a href="/wiki/Hydrophobic" class="mw-redirect" title="Hydrophobic">hydrophobic</a> materials to benefit people living in dry regions with no regular rainfall.<a href="#cite_note-205">[202]</a> Living beetles have been used as <a href="/wiki/Cyborg" title="Cyborg">cyborgs</a>. A <a href="/wiki/Defense_Advanced_Research_Projects_Agency" class="mw-redirect" title="Defense Advanced Research Projects Agency">Defense Advanced Research Projects Agency</a> funded project implanted electrodes into <a href="/wiki/Mecynorhina_torquata" title="Mecynorhina torquata">Mecynorhina torquata</a> beetles, allowing them to be remotely controlled via a radio receiver held on its back, as proof-of-concept for surveillance work.<a href="#cite_note-206">[203]</a> Similar technology has been applied to enable a human operator to control the free-flight steering and walking gaits of <a href="/wiki/Mecynorhina_torquata" title="Mecynorhina torquata">Mecynorhina torquata</a> as well as graded turning, backward walking and feedback control of <a href="/wiki/Zophobas_morio" class="mw-redirect" title="Zophobas morio">Zophobas morio</a>.<a href="#cite_note-207">[204]</a><a href="#cite_note-208">[205]</a><a href="#cite_note-209">[206]</a><a href="#cite_note-210">[207]</a><a href="#cite_note-211">[208]</a> Research published in 2020 sought to create a robotic camera backpack for beetles. Miniature cameras weighing 248 mg were attached to live beetles of the <a href="/wiki/Tenebrionid" class="mw-redirect" title="Tenebrionid">Tenebrionid</a> genera <a href="/wiki/Asbolus_(beetle)" title="Asbolus (beetle)">Asbolus</a> and <a href="/wiki/Eleodes" title="Eleodes">Eleodes</a>. The cameras filmed over a 60° range for up to 6 hours.<a href="#cite_note-212">[209]</a><a href="#cite_note-213">[210]</a> <div class="mw-heading mw-heading3"><h3 id="In_conservation">In conservation</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=54" title="Edit section: In conservation">edit</a>]</div> Since beetles form such a large part of the world's biodiversity, their conservation is important, and equally, loss of habitat and biodiversity is essentially certain to impact on beetles. Many species of beetles have very specific habitats and long life cycles that make them vulnerable. Some species are highly threatened while others are already feared extinct.<a href="#cite_note-214">[211]</a> Island species tend to be more susceptible as in the case of Helictopleurus undatus of Madagascar which is thought to have gone extinct during the late 20th century.<a href="#cite_note-215">[212]</a> Conservationists have attempted to arouse a liking for beetles with flagship species like the stag beetle, <a href="/wiki/Lucanus_cervus" title="Lucanus cervus">Lucanus cervus</a>,<a href="#cite_note-216">[213]</a> and tiger beetles (<a href="/wiki/Cicindelidae" class="mw-redirect" title="Cicindelidae">Cicindelidae</a>). In Japan the Genji firefly, <a href="/wiki/Luciola_cruciata" class="mw-redirect" title="Luciola cruciata">Luciola cruciata</a>, is extremely popular, and in South Africa the Addo elephant dung beetle offers promise for broadening <a href="/wiki/Ecotourism" title="Ecotourism">ecotourism</a> beyond the <a href="/wiki/Big_five_game" title="Big five game">big five tourist mammal species</a>. Popular dislike of pest beetles, too, can be turned into public interest in insects, as can unusual ecological adaptations of species like the fairy shrimp hunting beetle, <a href="/w/index.php?title=Cicinis_bruchi&action=edit&redlink=1" class="new" title="Cicinis bruchi (page does not exist)">Cicinis bruchi</a>.<a href="#cite_note-217">[214]</a> <div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=55" title="Edit section: Notes">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output .reflist-lower-alpha{list-style-type:lower-alpha}.mw-parser-output .reflist-lower-greek{list-style-type:lower-greek}.mw-parser-output .reflist-lower-roman{list-style-type:lower-roman}</style><div class="reflist reflist-lower-alpha"> <div class="mw-references-wrap"><ol class="references"> <li id="cite_note-62"><a href="#cite_ref-62">^</a> These count fore, mid, and hind leg tarsal segments, such as 5-5-4. </li> <li id="cite_note-161"><a href="#cite_ref-161">^</a> The wood-gnawing longhorn beetle genus <a href="/wiki/Cerambyx" title="Cerambyx">Cerambyx</a> is named for him. </li> <li id="cite_note-198"><a href="#cite_ref-198">^</a> The plate was labelled "Neocerambyx æneas, Cladognathus tarandus, Diurus furcellatus, Ectatorhinus Wallacei, Megacriodes Saundersii, Cyriopalpus Wallacei". </li> </ol></div></div> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=56" title="Edit section: See also">edit</a>]</div> <ul><li><a href="/wiki/Beetle_Monuments" title="Beetle Monuments">Beetle Monuments</a></li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=57" title="Edit section: References">edit</a>]</div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1239543626"><div class="reflist"> <div class="mw-references-wrap mw-references-columns"><ol class="references"> <li id="cite_note-Bouchard_et_al_2011-1">^ <a href="#cite_ref-Bouchard_et_al_2011_1-0">a</a> <a href="#cite_ref-Bouchard_et_al_2011_1-1">b</a> <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="CITEREFBouchardBousquetDaviesAlonso-Zarazaga2011" class="citation journal cs1">Bouchard, P.; Bousquet, Y.; Davies, A.; Alonso-Zarazaga, M.; Lawrence, J.; Lyal, C.; Newton, A.; Reid, C.; Schmitt, M.; Ślipiński, A.; Smith, A. (2011). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3088472">"Family-group names in Coleoptera (Insecta)"</a>. ZooKeys (88): 1–972. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2011ZooK...88....1B">2011ZooK...88....1B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.3897%2Fzookeys.88.807">10.3897/zookeys.88.807</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3088472">3088472</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/21594053">21594053</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=ZooKeys&rft.atitle=Family-group+names+in+Coleoptera+%28Insecta%29&rft.issue=88&rft.pages=1-972&rft.date=2011&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3088472%23id-name%3DPMC&rft_id=info%3Apmid%2F21594053&rft_id=info%3Adoi%2F10.3897%2Fzookeys.88.807&rft_id=info%3Abibcode%2F2011ZooK...88....1B&rft.aulast=Bouchard&rft.aufirst=P.&rft.au=Bousquet%2C+Y.&rft.au=Davies%2C+A.&rft.au=Alonso-Zarazaga%2C+M.&rft.au=Lawrence%2C+J.&rft.au=Lyal%2C+C.&rft.au=Newton%2C+A.&rft.au=Reid%2C+C.&rft.au=Schmitt%2C+M.&rft.au=%C5%9Alipi%C5%84ski%2C+A.&rft.au=Smith%2C+A.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3088472&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Stork2018-2"><a href="#cite_ref-Stork2018_2-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFStork2018" class="citation journal cs1">Stork, Nigel E. (January 7, 2018). <a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev-ento-020117-043348">"How Many Species of Insects and Other Terrestrial Arthropods Are There on Earth?"</a>. Annual Review of Entomology. 63 (1): 31–45. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev-ento-020117-043348">10.1146/annurev-ento-020117-043348</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/28938083">28938083</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:23755007">23755007</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Entomology&rft.atitle=How+Many+Species+of+Insects+and+Other+Terrestrial+Arthropods+Are+There+on+Earth%3F&rft.volume=63&rft.issue=1&rft.pages=31-45&rft.date=2018-01-07&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A23755007%23id-name%3DS2CID&rft_id=info%3Apmid%2F28938083&rft_id=info%3Adoi%2F10.1146%2Fannurev-ento-020117-043348&rft.aulast=Stork&rft.aufirst=Nigel+E.&rft_id=https%3A%2F%2Fdoi.org%2F10.1146%252Fannurev-ento-020117-043348&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Etymology-3"><a href="#cite_ref-Etymology_3-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHarper" class="citation web cs1">Harper, Douglas. <a rel="nofollow" class="external text" href="https://www.etymonline.com/?term=Coleoptera">"Coleoptera"</a>. <a href="/wiki/Online_Etymology_Dictionary" title="Online Etymology Dictionary">Online Etymology Dictionary</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Online+Etymology+Dictionary&rft.atitle=Coleoptera&rft.aulast=Harper&rft.aufirst=Douglas&rft_id=https%3A%2F%2Fwww.etymonline.com%2F%3Fterm%3DColeoptera&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Etymology_Beetle-4"><a href="#cite_ref-Etymology_Beetle_4-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHarper" class="citation web cs1">Harper, Douglas. <a rel="nofollow" class="external text" href="https://www.etymonline.com/?term=Beetle">"Beetle"</a>. <a href="/wiki/Online_Etymology_Dictionary" title="Online Etymology Dictionary">Online Etymology Dictionary</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Online+Etymology+Dictionary&rft.atitle=Beetle&rft.aulast=Harper&rft.aufirst=Douglas&rft_id=https%3A%2F%2Fwww.etymonline.com%2F%3Fterm%3DBeetle&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Merriam_Dictionary-5"><a href="#cite_ref-Merriam_Dictionary_5-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.merriam-webster.com/dictionary/Beetle">"Beetle"</a>. Merriam-Webster Online Dictionary. Retrieved February 20, 2016.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Beetle&rft.pub=Merriam-Webster+Online+Dictionary&rft_id=http%3A%2F%2Fwww.merriam-webster.com%2Fdictionary%2FBeetle&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Etymology_Chafer-6"><a href="#cite_ref-Etymology_Chafer_6-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHarper" class="citation web cs1">Harper, Douglas. <a rel="nofollow" class="external text" href="https://www.etymonline.com/?term=Chafer">"Chafer"</a>. <a href="/wiki/Online_Etymology_Dictionary" title="Online Etymology Dictionary">Online Etymology Dictionary</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Online+Etymology+Dictionary&rft.atitle=Chafer&rft.aulast=Harper&rft.aufirst=Douglas&rft_id=https%3A%2F%2Fwww.etymonline.com%2F%3Fterm%3DChafer&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Gilliott-7">^ <a href="#cite_ref-Gilliott_7-0">a</a> <a href="#cite_ref-Gilliott_7-1">b</a> <a href="#cite_ref-Gilliott_7-2">c</a> <a href="#cite_ref-Gilliott_7-3">d</a> <a href="#cite_ref-Gilliott_7-4">e</a> <a href="#cite_ref-Gilliott_7-5">f</a> <a href="#cite_ref-Gilliott_7-6">g</a> <a href="#cite_ref-Gilliott_7-7">h</a> <a href="#cite_ref-Gilliott_7-8">i</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGilliott1995" class="citation book cs1">Gilliott, Cedric (August 1995). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=DrTKxvZq_IcC&pg=PA96">Entomology</a> (2 ed.). Springer-Verlag. p. 96. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-306-44967-3" title="Special:BookSources/978-0-306-44967-3"><bdi>978-0-306-44967-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Entomology&rft.pages=96&rft.edition=2&rft.pub=Springer-Verlag&rft.date=1995-08&rft.isbn=978-0-306-44967-3&rft.aulast=Gilliott&rft.aufirst=Cedric&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DDrTKxvZq_IcC%26pg%3DPA96&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-insenc-8">^ <a href="#cite_ref-insenc_8-0">a</a> <a href="#cite_ref-insenc_8-1">b</a> <a href="#cite_ref-insenc_8-2">c</a> <a href="#cite_ref-insenc_8-3">d</a> <a href="#cite_ref-insenc_8-4">e</a> <a href="#cite_ref-insenc_8-5">f</a> <a href="#cite_ref-insenc_8-6">g</a> <a href="#cite_ref-insenc_8-7">h</a> <a href="#cite_ref-insenc_8-8">i</a> <a href="#cite_ref-insenc_8-9">j</a> <a href="#cite_ref-insenc_8-10">k</a> <a href="#cite_ref-insenc_8-11">l</a> <a href="#cite_ref-insenc_8-12">m</a> <a href="#cite_ref-insenc_8-13">n</a> <a href="#cite_ref-insenc_8-14">o</a> <a href="#cite_ref-insenc_8-15">p</a> <a href="#cite_ref-insenc_8-16">q</a> <a href="#refMcHugh">McHugh (2009)</a> </li> <li id="cite_note-Rosenzweig_1995-9"><a href="#cite_ref-Rosenzweig_1995_9-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRosenzweig1995" class="citation book cs1">Rosenzweig, M. L. (1995). Species Diversity in Space and Time. Cambridge: Cambridge University Press. p. 2. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-521-49952-1" title="Special:BookSources/978-0-521-49952-1"><bdi>978-0-521-49952-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Species+Diversity+in+Space+and+Time&rft.place=Cambridge&rft.pages=2&rft.pub=Cambridge+University+Press&rft.date=1995&rft.isbn=978-0-521-49952-1&rft.aulast=Rosenzweig&rft.aufirst=M.+L.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Hunt_et_al_2007-10">^ <a href="#cite_ref-Hunt_et_al_2007_10-0">a</a> <a href="#cite_ref-Hunt_et_al_2007_10-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHunt,_T.BergstenLevkanicovaPapadopoulou2007" class="citation journal cs1">Hunt, T.; Bergsten, J.; Levkanicova, Z.; Papadopoulou, A.; John, O. S.; Wild, R.; Hammond, P. M.; Ahrens, D.; Balke, M.; Caterino, M. S.; Gómez-Zurita, J.; Ribera, I; Barraclough, T. G.; Bocakova, M.; Bocak, L; Vogler, A. P. (2007). "A Comprehensive Phylogeny of Beetles Reveals the Evolutionary Origins of a Superradiation". <a href="/wiki/Science_(journal)" title="Science (journal)">Science</a>. 318 (5858): 1913–1916. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2007Sci...318.1913H">2007Sci...318.1913H</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.1146954">10.1126/science.1146954</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18096805">18096805</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:19392955">19392955</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=A+Comprehensive+Phylogeny+of+Beetles+Reveals+the+Evolutionary+Origins+of+a+Superradiation&rft.volume=318&rft.issue=5858&rft.pages=1913-1916&rft.date=2007&rft_id=info%3Adoi%2F10.1126%2Fscience.1146954&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A19392955%23id-name%3DS2CID&rft_id=info%3Apmid%2F18096805&rft_id=info%3Abibcode%2F2007Sci...318.1913H&rft.au=Hunt%2C+T.&rft.au=Bergsten%2C+J.&rft.au=Levkanicova%2C+Z.&rft.au=Papadopoulou%2C+A.&rft.au=John%2C+O.+S.&rft.au=Wild%2C+R.&rft.au=Hammond%2C+P.+M.&rft.au=Ahrens%2C+D.&rft.au=Balke%2C+M.&rft.au=Caterino%2C+M.+S.&rft.au=G%C3%B3mez-Zurita%2C+J.&rft.au=Ribera%2C+I&rft.au=Barraclough%2C+T.+G.&rft.au=Bocakova%2C+M.&rft.au=Bocak%2C+L&rft.au=Vogler%2C+A.+P.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Hammond_1992-11"><a href="#cite_ref-Hammond_1992_11-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHammond1992" class="citation book cs1">Hammond, Peter (1992). "Species Inventory". In Groombridge, Brian (ed.). <a rel="nofollow" class="external text" href="https://wedocs.unep.org/bitstream/handle/20.500.11822/7981/globalbiodiversi92wcmc.pdf">Global Biodiversity: Status of the Earth's Living Resources</a> (PDF). London: Chapman & Hall. pp. 17–39. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-412-47240-4" title="Special:BookSources/978-0-412-47240-4"><bdi>978-0-412-47240-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Species+Inventory&rft.btitle=Global+Biodiversity%3A+Status+of+the+Earth%27s+Living+Resources&rft.place=London&rft.pages=17-39&rft.pub=Chapman+%26+Hall&rft.date=1992&rft.isbn=978-0-412-47240-4&rft.aulast=Hammond&rft.aufirst=Peter&rft_id=https%3A%2F%2Fwedocs.unep.org%2Fbitstream%2Fhandle%2F20.500.11822%2F7981%2Fglobalbiodiversi92wcmc.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Stork-12">^ <a href="#cite_ref-Stork_12-0">a</a> <a href="#cite_ref-Stork_12-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFStorkMcBroomGelyHamilton2015" class="citation journal cs1">Stork, Nigel E.; McBroom, James; Gely, Claire; Hamilton, Andrew J. (2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4475949">"New approaches narrow global species estimates for beetles, insects, and terrestrial arthropods"</a>. PNAS. 116 (24): 7519–7523. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2015PNAS..112.7519S">2015PNAS..112.7519S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.1502408112">10.1073/pnas.1502408112</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4475949">4475949</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26034274">26034274</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PNAS&rft.atitle=New+approaches+narrow+global+species+estimates+for+beetles%2C+insects%2C+and+terrestrial+arthropods&rft.volume=116&rft.issue=24&rft.pages=7519-7523&rft.date=2015&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4475949%23id-name%3DPMC&rft_id=info%3Apmid%2F26034274&rft_id=info%3Adoi%2F10.1073%2Fpnas.1502408112&rft_id=info%3Abibcode%2F2015PNAS..112.7519S&rft.aulast=Stork&rft.aufirst=Nigel+E.&rft.au=McBroom%2C+James&rft.au=Gely%2C+Claire&rft.au=Hamilton%2C+Andrew+J.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4475949&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-13"><a href="#cite_ref-13">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGullanCranston,_P.S.2014" class="citation book cs1">Gullan, P.J.; Cranston, P.S. (2014). <a rel="nofollow" class="external text" href="https://archive.org/details/insectsoutlineen00pjgu">The Insects: An Outline of Entomology</a> (5 ed.). John Wiley & Sons. p. <a rel="nofollow" class="external text" href="https://archive.org/details/insectsoutlineen00pjgu/page/n24">6</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4443-3036-6" title="Special:BookSources/978-1-4443-3036-6"><bdi>978-1-4443-3036-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Insects%3A+An+Outline+of+Entomology&rft.pages=6&rft.edition=5&rft.pub=John+Wiley+%26+Sons&rft.date=2014&rft.isbn=978-1-4443-3036-6&rft.aulast=Gullan&rft.aufirst=P.J.&rft.au=Cranston%2C+P.S.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Finsectsoutlineen00pjgu&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Hutchinson_1959-14"><a href="#cite_ref-Hutchinson_1959_14-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHutchinson1959" class="citation journal cs1">Hutchinson, G. E. (1959). "Homage to Santa Rosalia or why are there so many kinds of animals?". The American Naturalist. 93 (870): 145–159. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1086%2F282070">10.1086/282070</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/2458768">2458768</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:26401739">26401739</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+American+Naturalist&rft.atitle=Homage+to+Santa+Rosalia+or+why+are+there+so+many+kinds+of+animals%3F&rft.volume=93&rft.issue=870&rft.pages=145-159&rft.date=1959&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A26401739%23id-name%3DS2CID&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F2458768%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.1086%2F282070&rft.aulast=Hutchinson&rft.aufirst=G.+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-15"><a href="#cite_ref-15">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHebertRatnasinghamZakharovTelfer2016" class="citation journal cs1">Hebert, Paul D. N.; Ratnasingham, Sujeevan; Zakharov, Evgeny V.; Telfer, Angela C.; Levesque-Beaudin, Valerie; Milton, Megan A.; Pedersen, Stephanie; Jannetta, Paul; deWaard, Jeremy R. (September 5, 2016). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4971185">"Counting animal species with DNA barcodes: Canadian insects"</a>. Philosophical Transactions of the Royal Society B: Biological Sciences. 371 (1702): 20150333. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frstb.2015.0333">10.1098/rstb.2015.0333</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0962-8436">0962-8436</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4971185">4971185</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/27481785">27481785</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Philosophical+Transactions+of+the+Royal+Society+B%3A+Biological+Sciences&rft.atitle=Counting+animal+species+with+DNA+barcodes%3A+Canadian+insects&rft.volume=371&rft.issue=1702&rft.pages=20150333&rft.date=2016-09-05&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4971185%23id-name%3DPMC&rft.issn=0962-8436&rft_id=info%3Apmid%2F27481785&rft_id=info%3Adoi%2F10.1098%2Frstb.2015.0333&rft.aulast=Hebert&rft.aufirst=Paul+D.+N.&rft.au=Ratnasingham%2C+Sujeevan&rft.au=Zakharov%2C+Evgeny+V.&rft.au=Telfer%2C+Angela+C.&rft.au=Levesque-Beaudin%2C+Valerie&rft.au=Milton%2C+Megan+A.&rft.au=Pedersen%2C+Stephanie&rft.au=Jannetta%2C+Paul&rft.au=deWaard%2C+Jeremy+R.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4971185&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-16"><a href="#cite_ref-16">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBorkentBrownAdlerAmorim2018" class="citation journal cs1">Borkent, Art; Brown, Brian V.; Adler, Peter H.; Amorim, Dalton De Souza; Barber, Kevin; Bickel, Daniel; Boucher, Stephanie; Brooks, Scott E.; Burger, John; Burington, Z.L.; Capellari, Renato S.; Costa, Daniel N.R.; Cumming, Jeffrey M.; Curler, Greg; Dick, Carl W. (March 27, 2018). <a rel="nofollow" class="external text" href="https://www.mapress.com/zt/article/view/zootaxa.4402.1.3">"Remarkable fly (Diptera) diversity in a patch of Costa Rican cloud forest: Why inventory is a vital science"</a>. Zootaxa. 4402 (1): 53–90. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.11646%2Fzootaxa.4402.1.3">10.11646/zootaxa.4402.1.3</a>. <a href="/wiki/Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<a rel="nofollow" class="external text" href="https://hdl.handle.net/10138%2F234433">10138/234433</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1175-5334">1175-5334</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/29690278">29690278</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:13819313">13819313</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zootaxa&rft.atitle=Remarkable+fly+%28Diptera%29+diversity+in+a+patch+of+Costa+Rican+cloud+forest%3A+Why+inventory+is+a+vital+science&rft.volume=4402&rft.issue=1&rft.pages=53-90&rft.date=2018-03-27&rft_id=info%3Ahdl%2F10138%2F234433&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A13819313%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.11646%2Fzootaxa.4402.1.3&rft.issn=1175-5334&rft_id=info%3Apmid%2F29690278&rft.aulast=Borkent&rft.aufirst=Art&rft.au=Brown%2C+Brian+V.&rft.au=Adler%2C+Peter+H.&rft.au=Amorim%2C+Dalton+De+Souza&rft.au=Barber%2C+Kevin&rft.au=Bickel%2C+Daniel&rft.au=Boucher%2C+Stephanie&rft.au=Brooks%2C+Scott+E.&rft.au=Burger%2C+John&rft.au=Burington%2C+Z.L.&rft.au=Capellari%2C+Renato+S.&rft.au=Costa%2C+Daniel+N.R.&rft.au=Cumming%2C+Jeffrey+M.&rft.au=Curler%2C+Greg&rft.au=Dick%2C+Carl+W.&rft_id=https%3A%2F%2Fwww.mapress.com%2Fzt%2Farticle%2Fview%2Fzootaxa.4402.1.3&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-17"><a href="#cite_ref-17">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFForbesBagleyBeerHippee2018" class="citation journal cs1">Forbes, Andrew A.; Bagley, Robin K.; Beer, Marc A.; Hippee, Alaine C.; Widmayer, Heather A. (July 12, 2018). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6042248">"Quantifying the unquantifiable: why Hymenoptera, not Coleoptera, is the most speciose animal order"</a>. BMC Ecology. 18 (1): 21. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2018BMCE...18...21F">2018BMCE...18...21F</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1186%2Fs12898-018-0176-x">10.1186/s12898-018-0176-x</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1472-6785">1472-6785</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6042248">6042248</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/30001194">30001194</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BMC+Ecology&rft.atitle=Quantifying+the+unquantifiable%3A+why+Hymenoptera%2C+not+Coleoptera%2C+is+the+most+speciose+animal+order&rft.volume=18&rft.issue=1&rft.pages=21&rft.date=2018-07-12&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6042248%23id-name%3DPMC&rft_id=info%3Abibcode%2F2018BMCE...18...21F&rft_id=info%3Apmid%2F30001194&rft_id=info%3Adoi%2F10.1186%2Fs12898-018-0176-x&rft.issn=1472-6785&rft.aulast=Forbes&rft.aufirst=Andrew+A.&rft.au=Bagley%2C+Robin+K.&rft.au=Beer%2C+Marc+A.&rft.au=Hippee%2C+Alaine+C.&rft.au=Widmayer%2C+Heather+A.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC6042248&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-18"><a href="#cite_ref-18">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGullanCranston,_P._S.2014" class="citation book cs1">Gullan, P. J.; Cranston, P. S. (2014). <a rel="nofollow" class="external text" href="https://archive.org/details/insectsoutlineen00pjgu">The Insects: An Outline of Entomology</a> (5 ed.). John Wiley & Sons. p. <a rel="nofollow" class="external text" href="https://archive.org/details/insectsoutlineen00pjgu/page/n541">517</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4443-3036-6" title="Special:BookSources/978-1-4443-3036-6"><bdi>978-1-4443-3036-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Insects%3A+An+Outline+of+Entomology&rft.pages=517&rft.edition=5&rft.pub=John+Wiley+%26+Sons&rft.date=2014&rft.isbn=978-1-4443-3036-6&rft.aulast=Gullan&rft.aufirst=P.+J.&rft.au=Cranston%2C+P.+S.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Finsectsoutlineen00pjgu&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-19"><a href="#cite_ref-19">^</a> Kirmse S, Adis J, Morawetz W. 2003. Flowering events and beetle diversity in Venezuela. In: Basset Y, Novotny V, Miller SE, Kitching RL, editors. Arthropods of tropical forests: Spatio-temporal dynamics and resource use in the canopy. Cambridge: Cambridge University Press; p. 256–265. </li> <li id="cite_note-20"><a href="#cite_ref-20">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArndtKirmseErwin2001" class="citation journal cs1">Arndt, Erik; Kirmse, Susan; Erwin, Terry L. (2001). "Arboreal Beetles of Neotropical Forests:Agra Fabricius, Larval Descriptions with Notes on Natural History and Behaviour (Coleoptera, Carabidae, Lebiini, Agrina)". The Coleopterists Bulletin. 55 (3): 297–310. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1649%2F0010-065x%282001%29055%5B0297%3Aabonfa%5D2.0.co%3B2">10.1649/0010-065x(2001)055[0297:abonfa]2.0.co;2</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:52065045">52065045</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Coleopterists+Bulletin&rft.atitle=Arboreal+Beetles+of+Neotropical+Forests%3AAgra+Fabricius%2C+Larval+Descriptions+with+Notes+on+Natural+History+and+Behaviour+%28Coleoptera%2C+Carabidae%2C+Lebiini%2C+Agrina%29&rft.volume=55&rft.issue=3&rft.pages=297-310&rft.date=2001&rft_id=info%3Adoi%2F10.1649%2F0010-065x%282001%29055%5B0297%3Aabonfa%5D2.0.co%3B2&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A52065045%23id-name%3DS2CID&rft.aulast=Arndt&rft.aufirst=Erik&rft.au=Kirmse%2C+Susan&rft.au=Erwin%2C+Terry+L.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-21"><a href="#cite_ref-21">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKirmseChaboo2018" class="citation journal cs1">Kirmse, Susan; Chaboo, Caroline S. (2018). "Polyphagy and florivory prevail in a leaf-beetle community (Coleoptera: Chrysomelidae) inhabiting the canopy of a tropical lowland rainforest in southern Venezuela". Journal of Natural History. 52 (41–42): 2677–2721. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2018JNatH..52.2677K">2018JNatH..52.2677K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00222933.2018.1548666">10.1080/00222933.2018.1548666</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:91732501">91732501</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Natural+History&rft.atitle=Polyphagy+and+florivory+prevail+in+a+leaf-beetle+community+%28Coleoptera%3A+Chrysomelidae%29+inhabiting+the+canopy+of+a+tropical+lowland+rainforest+in+southern+Venezuela&rft.volume=52&rft.issue=41%E2%80%9342&rft.pages=2677-2721&rft.date=2018&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A91732501%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1080%2F00222933.2018.1548666&rft_id=info%3Abibcode%2F2018JNatH..52.2677K&rft.aulast=Kirmse&rft.aufirst=Susan&rft.au=Chaboo%2C+Caroline+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-22"><a href="#cite_ref-22">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKirmseRatcliffe2019" class="citation journal cs1">Kirmse, Susan; Ratcliffe, Brett C. (2019). <a rel="nofollow" class="external text" href="https://digitalcommons.unl.edu/entomologyfacpub/799">"Composition and Host-Use Patterns of a Scarab Beetle (Coleoptera: Scarabaeidae) Community Inhabiting the Canopy of a Lowland Tropical Rainforest in Southern Venezuela"</a>. The Coleopterists Bulletin. 73: 149. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1649%2F0010-065X-73.1.149">10.1649/0010-065X-73.1.149</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:108786139">108786139</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Coleopterists+Bulletin&rft.atitle=Composition+and+Host-Use+Patterns+of+a+Scarab+Beetle+%28Coleoptera%3A+Scarabaeidae%29+Community+Inhabiting+the+Canopy+of+a+Lowland+Tropical+Rainforest+in+Southern+Venezuela&rft.volume=73&rft.pages=149&rft.date=2019&rft_id=info%3Adoi%2F10.1649%2F0010-065X-73.1.149&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A108786139%23id-name%3DS2CID&rft.aulast=Kirmse&rft.aufirst=Susan&rft.au=Ratcliffe%2C+Brett+C.&rft_id=https%3A%2F%2Fdigitalcommons.unl.edu%2Fentomologyfacpub%2F799&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-23"><a href="#cite_ref-23">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.guinnessworldrecords.com/world-records/heaviest-insect">"Heaviest insect"</a>. <a href="/wiki/Guinness_World_Records" title="Guinness World Records">Guinness World Records</a>. Retrieved February 1, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Heaviest+insect&rft.pub=Guinness+World+Records&rft_id=http%3A%2F%2Fwww.guinnessworldrecords.com%2Fworld-records%2Fheaviest-insect&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Williams2001-24"><a href="#cite_ref-Williams2001_24-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWilliams,_David_M.2001" class="citation web cs1">Williams, David M. (2001). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110718140051/http://entnemdept.ufl.edu/walker/ufbir/chapters/chapter_30.shtml">"Chapter 30 — Largest Insect"</a>. Book of Insect Records. <a href="/wiki/University_of_Florida" title="University of Florida">University of Florida</a>. Archived from <a rel="nofollow" class="external text" href="http://entnemdept.ufl.edu/walker/ufbir/chapters/chapter_30.shtml">the original</a> on July 18, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Book+of+Insect+Records&rft.atitle=Chapter+30+%E2%80%94+Largest+Insect&rft.date=2001&rft.au=Williams%2C+David+M.&rft_id=http%3A%2F%2Fentnemdept.ufl.edu%2Fwalker%2Fufbir%2Fchapters%2Fchapter_30.shtml&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-25"><a href="#cite_ref-25">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPolilov2015" class="citation journal cs1">Polilov, Alexey (2015). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4607844">"How small is the smallest? New record and remeasuring of Scydosella musawasensis Hall, 1999 (Coleoptera, Ptiliidae), the smallest known free-living insect"</a>. <a href="/wiki/ZooKeys" title="ZooKeys">ZooKeys</a> (526): 61–64. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2015ZooK..526...61P">2015ZooK..526...61P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.3897%2Fzookeys.526.6531">10.3897/zookeys.526.6531</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4607844">4607844</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/26487824">26487824</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=ZooKeys&rft.atitle=How+small+is+the+smallest%3F+New+record+and+remeasuring+of+Scydosella+musawasensis+Hall%2C+1999+%28Coleoptera%2C+Ptiliidae%29%2C+the+smallest+known+free-living+insect&rft.issue=526&rft.pages=61-64&rft.date=2015&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4607844%23id-name%3DPMC&rft_id=info%3Apmid%2F26487824&rft_id=info%3Adoi%2F10.3897%2Fzookeys.526.6531&rft_id=info%3Abibcode%2F2015ZooK..526...61P&rft.aulast=Polilov&rft.aufirst=Alexey&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4607844&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-26"><a href="#cite_ref-26">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKirejtshukPoschmannProkopGarrouste2014" class="citation journal cs1">Kirejtshuk, Alexander G.; Poschmann, Markus; Prokop, Jakub; Garrouste, Romain; Nel, André (July 4, 2014). <a rel="nofollow" class="external text" href="http://www.tandfonline.com/doi/abs/10.1080/14772019.2013.821530">"Evolution of the elytral venation and structural adaptations in the oldest Palaeozoic beetles (Insecta: Coleoptera: Tshekardocoleidae)"</a>. Journal of Systematic Palaeontology. 12 (5): 575–600. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2014JSPal..12..575K">2014JSPal..12..575K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F14772019.2013.821530">10.1080/14772019.2013.821530</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/1477-2019">1477-2019</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:85163674">85163674</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Systematic+Palaeontology&rft.atitle=Evolution+of+the+elytral+venation+and+structural+adaptations+in+the+oldest+Palaeozoic+beetles+%28Insecta%3A+Coleoptera%3A+Tshekardocoleidae%29&rft.volume=12&rft.issue=5&rft.pages=575-600&rft.date=2014-07-04&rft_id=info%3Adoi%2F10.1080%2F14772019.2013.821530&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A85163674%23id-name%3DS2CID&rft.issn=1477-2019&rft_id=info%3Abibcode%2F2014JSPal..12..575K&rft.aulast=Kirejtshuk&rft.aufirst=Alexander+G.&rft.au=Poschmann%2C+Markus&rft.au=Prokop%2C+Jakub&rft.au=Garrouste%2C+Romain&rft.au=Nel%2C+Andr%C3%A9&rft_id=http%3A%2F%2Fwww.tandfonline.com%2Fdoi%2Fabs%2F10.1080%2F14772019.2013.821530&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-27"><a href="#cite_ref-27">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHörnschemeyerStapf,_H." class="citation journal cs1 cs1-prop-foreign-lang-source">Hörnschemeyer, T.; Stapf, H. "Die Insektentaphozönose von Niedermoschel (Asselian, unt. Perm; Deutschland)". Schriften der Alfred-Wegener-Stiftung (in German) (99/8): 98.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Schriften+der+Alfred-Wegener-Stiftung&rft.atitle=Die+Insektentaphoz%C3%B6nose+von+Niedermoschel+%28Asselian%2C+unt.+Perm%3B+Deutschland%29&rft.issue=99%2F8&rft.pages=98&rft.aulast=H%C3%B6rnschemeyer&rft.aufirst=T.&rft.au=Stapf%2C+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-28"><a href="#cite_ref-28">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKukalová,_J.1969" class="citation journal cs1">Kukalová, J. (1969). "On the systematic position of the supposed Permian beetles, Tshecardocoleidae, with a description of a new collection from Moravia". Sborník Geologických Věd, Paleontologie. 11: 139–161.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Sborn%C3%ADk+Geologick%C3%BDch+V%C4%9Bd%2C+Paleontologie&rft.atitle=On+the+systematic+position+of+the+supposed+Permian+beetles%2C+Tshecardocoleidae%2C+with+a+description+of+a+new+collection+from+Moravia&rft.volume=11&rft.pages=139-161&rft.date=1969&rft.au=Kukalov%C3%A1%2C+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-evo-29">^ <a href="#cite_ref-evo_29-0">a</a> <a href="#cite_ref-evo_29-1">b</a> <a href="#cite_ref-evo_29-2">c</a> <a href="#cite_ref-evo_29-3">d</a> <a href="#cite_ref-evo_29-4">e</a> <a href="#cite_ref-evo_29-5">f</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBenisch,_Christoph2010" class="citation web cs1">Benisch, Christoph (2010). <a rel="nofollow" class="external text" href="http://www.kerbtier.de/Pages/Themenseiten/enPhylogenie.html#L9">"Phylogeny of the beetles"</a>. The beetle fauna of Germany. Kerbtier. Retrieved March 16, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=The+beetle+fauna+of+Germany&rft.atitle=Phylogeny+of+the+beetles&rft.date=2010&rft.au=Benisch%2C+Christoph&rft_id=http%3A%2F%2Fwww.kerbtier.de%2FPages%2FThemenseiten%2FenPhylogenie.html%23L9&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-30"><a href="#cite_ref-30">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeckemeyerEngel,_M._S.2008" class="citation journal cs1">Beckemeyer, R. J.; Engel, M. S. (2008). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110718202310/http://fossilinsects.net/pdfs/beckemeyer_engel_2008_JKansEntSoc_PermocoleusPermOklahoma.pdf">"A second specimen of Permocoleus (Coleoptera) from the Lower Permian Wellington Formation of Noble County, Oklahoma"</a> (PDF). <a href="/wiki/Journal_of_the_Kansas_Entomological_Society" title="Journal of the Kansas Entomological Society">Journal of the Kansas Entomological Society</a>. 81 (1): 4–7. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2317%2FJKES-708.01.1">10.2317/JKES-708.01.1</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:86835593">86835593</a>. Archived from <a rel="nofollow" class="external text" href="http://fossilinsects.net/pdfs/beckemeyer_engel_2008_JKansEntSoc_PermocoleusPermOklahoma.pdf">the original</a> (PDF) on July 18, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+Kansas+Entomological+Society&rft.atitle=A+second+specimen+of+Permocoleus+%28Coleoptera%29+from+the+Lower+Permian+Wellington+Formation+of+Noble+County%2C+Oklahoma&rft.volume=81&rft.issue=1&rft.pages=4-7&rft.date=2008&rft_id=info%3Adoi%2F10.2317%2FJKES-708.01.1&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A86835593%23id-name%3DS2CID&rft.aulast=Beckemeyer&rft.aufirst=R.+J.&rft.au=Engel%2C+M.+S.&rft_id=http%3A%2F%2Ffossilinsects.net%2Fpdfs%2Fbeckemeyer_engel_2008_JKansEntSoc_PermocoleusPermOklahoma.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-:1-31"><a href="#cite_ref-:1_31-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZhaoYuClaphamYan2021" class="citation journal cs1">Zhao, Xianye; Yu, Yilun; Clapham, Matthew E; Yan, Evgeny; Chen, Jun; Jarzembowski, Edmund A; Zhao, Xiangdong; Wang, Bo (November 8, 2021). Perry, George H; Fikacek, Martin (eds.). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8585485">"Early evolution of beetles regulated by the end-Permian deforestation"</a>. eLife. 10: e72692. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.7554%2FeLife.72692">10.7554/eLife.72692</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2050-084X">2050-084X</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8585485">8585485</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/34747694">34747694</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=eLife&rft.atitle=Early+evolution+of+beetles+regulated+by+the+end-Permian+deforestation&rft.volume=10&rft.pages=e72692&rft.date=2021-11-08&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8585485%23id-name%3DPMC&rft.issn=2050-084X&rft_id=info%3Apmid%2F34747694&rft_id=info%3Adoi%2F10.7554%2FeLife.72692&rft.aulast=Zhao&rft.aufirst=Xianye&rft.au=Yu%2C+Yilun&rft.au=Clapham%2C+Matthew+E&rft.au=Yan%2C+Evgeny&rft.au=Chen%2C+Jun&rft.au=Jarzembowski%2C+Edmund+A&rft.au=Zhao%2C+Xiangdong&rft.au=Wang%2C+Bo&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC8585485&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-insenc186-32">^ <a href="#cite_ref-insenc186_32-0">a</a> <a href="#cite_ref-insenc186_32-1">b</a> <a href="#cite_ref-insenc186_32-2">c</a> <a href="#refMcHugh">McHugh (2009)</a>, p. 186 </li> <li id="cite_note-33"><a href="#cite_ref-33">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLabandeiraSepkoski1993" class="citation journal cs1">Labandeira, C. C.; Sepkoski, J. J. (1993). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20120331061227/http://129.123.92.202/biol5530/Labandeira_Sepkoski_1993.pdf">"Insect diversity in the fossil record"</a> (PDF). <a href="/wiki/Science_(journal)" title="Science (journal)">Science</a>. 261 (5119): 310–315. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1993Sci...261..310L">1993Sci...261..310L</a>. <a href="/wiki/CiteSeerX_(identifier)" class="mw-redirect" title="CiteSeerX (identifier)">CiteSeerX</a> <a rel="nofollow" class="external text" href="https://citeseerx.ist.psu.edu/viewdoc/summary?doi=10.1.1.496.1576">10.1.1.496.1576</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.11536548">10.1126/science.11536548</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11536548">11536548</a>. Archived from <a rel="nofollow" class="external text" href="http://129.123.92.202/biol5530/Labandeira_Sepkoski_1993.pdf">the original</a> (PDF) on March 31, 2012.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Insect+diversity+in+the+fossil+record&rft.volume=261&rft.issue=5119&rft.pages=310-315&rft.date=1993&rft_id=https%3A%2F%2Fciteseerx.ist.psu.edu%2Fviewdoc%2Fsummary%3Fdoi%3D10.1.1.496.1576%23id-name%3DCiteSeerX&rft_id=info%3Apmid%2F11536548&rft_id=info%3Adoi%2F10.1126%2Fscience.11536548&rft_id=info%3Abibcode%2F1993Sci...261..310L&rft.aulast=Labandeira&rft.aufirst=C.+C.&rft.au=Sepkoski%2C+J.+J.&rft_id=http%3A%2F%2F129.123.92.202%2Fbiol5530%2FLabandeira_Sepkoski_1993.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-34"><a href="#cite_ref-34">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGratshev,_Vadim_G.Zherikhin,_Vladimir_V.2003" class="citation journal cs1">Gratshev, Vadim G.; Zherikhin, Vladimir V. (October 15, 2003). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20161229073633/http://www.isez.pan.krakow.pl/journals/azc_i/pdf/46(suppl)/15.pdf">"Insect diversity in the fossil record"</a> (PDF). Acta Zoologica Cracoviensia. 261 (5119): 129–138. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1993Sci...261..310L">1993Sci...261..310L</a>. <a href="/wiki/CiteSeerX_(identifier)" class="mw-redirect" title="CiteSeerX (identifier)">CiteSeerX</a> <a rel="nofollow" class="external text" href="https://citeseerx.ist.psu.edu/viewdoc/summary?doi=10.1.1.496.1576">10.1.1.496.1576</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.11536548">10.1126/science.11536548</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11536548">11536548</a>. Archived from <a rel="nofollow" class="external text" href="http://www.isez.pan.krakow.pl/journals/azc_i/pdf/46(suppl)/15.pdf">the original</a> (PDF) on December 29, 2016. Retrieved March 16, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Acta+Zoologica+Cracoviensia&rft.atitle=Insect+diversity+in+the+fossil+record&rft.volume=261&rft.issue=5119&rft.pages=129-138&rft.date=2003-10-15&rft_id=https%3A%2F%2Fciteseerx.ist.psu.edu%2Fviewdoc%2Fsummary%3Fdoi%3D10.1.1.496.1576%23id-name%3DCiteSeerX&rft_id=info%3Apmid%2F11536548&rft_id=info%3Adoi%2F10.1126%2Fscience.11536548&rft_id=info%3Abibcode%2F1993Sci...261..310L&rft.au=Gratshev%2C+Vadim+G.&rft.au=Zherikhin%2C+Vladimir+V.&rft_id=http%3A%2F%2Fwww.isez.pan.krakow.pl%2Fjournals%2Fazc_i%2Fpdf%2F46%28suppl%29%2F15.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-35"><a href="#cite_ref-35">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChang,_H.Zhang,_F.Ren,_D.2008" class="citation journal cs1">Chang, H.; Zhang, F.; Ren, D. (2008). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110704101413/http://202.204.209.200/upload/20080609030648.pdf">"A new genus and two new species of fossil elaterids from the Yixian Formation of Western Liaoning, China (Coleoptera: Elateridae)"</a> (PDF). <a href="/wiki/Zootaxa" title="Zootaxa">Zootaxa</a>. 1785 (1): 54–62. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.11646%2Fzootaxa.1785.1.3">10.11646/zootaxa.1785.1.3</a>. Archived from <a rel="nofollow" class="external text" href="http://202.204.209.200/upload/20080609030648.pdf">the original</a> (PDF) on July 4, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zootaxa&rft.atitle=A+new+genus+and+two+new+species+of+fossil+elaterids+from+the+Yixian+Formation+of+Western+Liaoning%2C+China+%28Coleoptera%3A+Elateridae%29&rft.volume=1785&rft.issue=1&rft.pages=54-62&rft.date=2008&rft_id=info%3Adoi%2F10.11646%2Fzootaxa.1785.1.3&rft.au=Chang%2C+H.&rft.au=Zhang%2C+F.&rft.au=Ren%2C+D.&rft_id=http%3A%2F%2F202.204.209.200%2Fupload%2F20080609030648.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-36"><a href="#cite_ref-36">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlexeev,_A._V.1993" class="citation journal cs1">Alexeev, A. V. (1993). <a rel="nofollow" class="external text" href="http://www.zin.ru/Animalia/Coleoptera/pdf/Alexeev%20-%20jurassic%20and%20lc%20buprestidae%2093.pdf">"Jurassic and Lower Cretaceous Buprestidae (Coleoptera) from Eurasia"</a> (PDF). <a href="/wiki/Paleontological_Journal" title="Paleontological Journal">Paleontological Journal</a> (1A): 9–34. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20100326031216/http://www.zin.ru/Animalia/Coleoptera/pdf/Alexeev%20-%20jurassic%20and%20lc%20buprestidae%2093.pdf">Archived</a> (PDF) from the original on March 26, 2010.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Paleontological+Journal&rft.atitle=Jurassic+and+Lower+Cretaceous+Buprestidae+%28Coleoptera%29+from+Eurasia&rft.issue=1A&rft.pages=9-34&rft.date=1993&rft.au=Alexeev%2C+A.+V.&rft_id=http%3A%2F%2Fwww.zin.ru%2FAnimalia%2FColeoptera%2Fpdf%2FAlexeev%2520-%2520jurassic%2520and%2520lc%2520buprestidae%252093.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-37"><a href="#cite_ref-37">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPonomarenko,_Alexandr_G.1985" class="citation journal cs1">Ponomarenko, Alexandr G. (1985). <a rel="nofollow" class="external text" href="http://www.landesmuseum.at/pdf_frei_remote/ANNA_87A_0135-0144.pdf">"Fossil insects from the Tithonian 'Solnhofener Plattenkalke' in the Museum of Natural History, Vienna"</a> (PDF). Annalen des Naturhistorischen Museums in Wien. 87 (1): 135–144. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110704203743/http://www.landesmuseum.at/pdf_frei_remote/ANNA_87A_0135-0144.pdf">Archived</a> (PDF) from the original on July 4, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annalen+des+Naturhistorischen+Museums+in+Wien&rft.atitle=Fossil+insects+from+the+Tithonian+%27Solnhofener+Plattenkalke%27+in+the+Museum+of+Natural+History%2C+Vienna&rft.volume=87&rft.issue=1&rft.pages=135-144&rft.date=1985&rft.au=Ponomarenko%2C+Alexandr+G.&rft_id=http%3A%2F%2Fwww.landesmuseum.at%2Fpdf_frei_remote%2FANNA_87A_0135-0144.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-38"><a href="#cite_ref-38">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFYan,_E._V.2009" class="citation journal cs1">Yan, E. V. (2009). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110718202329/http://fossilinsects.net/pdfs/Yan_2009_PalJ_ElateriformJurassicKaratau.pdf">"A new genus of elateriform beetles (Coleoptera, Polyphaga) from the Middle-Late Jurassic of Karatau"</a> (PDF). <a href="/wiki/Paleontological_Journal" title="Paleontological Journal">Paleontological Journal</a>. 43 (1): 78–82. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2009PalJ...43...78Y">2009PalJ...43...78Y</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1134%2FS0031030109010080">10.1134/S0031030109010080</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:84621777">84621777</a>. Archived from <a rel="nofollow" class="external text" href="http://fossilinsects.net/pdfs/Yan_2009_PalJ_ElateriformJurassicKaratau.pdf">the original</a> (PDF) on July 18, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Paleontological+Journal&rft.atitle=A+new+genus+of+elateriform+beetles+%28Coleoptera%2C+Polyphaga%29+from+the+Middle-Late+Jurassic+of+Karatau&rft.volume=43&rft.issue=1&rft.pages=78-82&rft.date=2009&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A84621777%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1134%2FS0031030109010080&rft_id=info%3Abibcode%2F2009PalJ...43...78Y&rft.au=Yan%2C+E.+V.&rft_id=http%3A%2F%2Ffossilinsects.net%2Fpdfs%2FYan_2009_PalJ_ElateriformJurassicKaratau.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-liaoning-39">^ <a href="#cite_ref-liaoning_39-0">a</a> <a href="#cite_ref-liaoning_39-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTan,_J.-J.Ren,_D.Liu,_M.2005" class="citation journal cs1">Tan, J.-J.; Ren, D.; Liu, M. (2005). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110718202354/http://fossilinsects.net/pdfs/tan_etal_2005.pdf">"New ommatids from the Late Jurassic of western Liaoning, China (Coleoptera: Archostemata)"</a> (PDF). Insect Science. 12 (3): 207–216. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2005InsSc..12..207T">2005InsSc..12..207T</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1005-295X.2005.00026.x">10.1111/j.1005-295X.2005.00026.x</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:83733980">83733980</a>. Archived from <a rel="nofollow" class="external text" href="http://fossilinsects.net/pdfs/tan_etal_2005.pdf">the original</a> (PDF) on July 18, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Insect+Science&rft.atitle=New+ommatids+from+the+Late+Jurassic+of+western+Liaoning%2C+China+%28Coleoptera%3A+Archostemata%29&rft.volume=12&rft.issue=3&rft.pages=207-216&rft.date=2005&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A83733980%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1111%2Fj.1005-295X.2005.00026.x&rft_id=info%3Abibcode%2F2005InsSc..12..207T&rft.au=Tan%2C+J.-J.&rft.au=Ren%2C+D.&rft.au=Liu%2C+M.&rft_id=http%3A%2F%2Ffossilinsects.net%2Fpdfs%2Ftan_etal_2005.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-40"><a href="#cite_ref-40">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPonomarenko,_A._G.1997" class="citation journal cs1">Ponomarenko, A. G. (1997). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20060925135716/http://www.palaeoentomolog.ru/Publ/PALJ389.pdf">"New beetles of the family Cupedidae from the Mesozoic of Mongolia. Ommatini, Mesocupedini, Priacmini"</a> (PDF). <a href="/wiki/Paleontological_Journal" title="Paleontological Journal">Paleontological Journal</a>. 31 (4): 389–399. Archived from <a rel="nofollow" class="external text" href="http://www.palaeoentomolog.ru/Publ/PALJ389.pdf">the original</a> (PDF) on September 25, 2006.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Paleontological+Journal&rft.atitle=New+beetles+of+the+family+Cupedidae+from+the+Mesozoic+of+Mongolia.+Ommatini%2C+Mesocupedini%2C+Priacmini&rft.volume=31&rft.issue=4&rft.pages=389-399&rft.date=1997&rft.au=Ponomarenko%2C+A.+G.&rft_id=http%3A%2F%2Fwww.palaeoentomolog.ru%2FPubl%2FPALJ389.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-41"><a href="#cite_ref-41">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlexeev2009" class="citation journal cs1">Alexeev, A. V. (2009). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110718202416/http://fossilinsects.net/pdfs/Alexeev_2009_PalJ_BuprestidaeCretaceousRussiaKazakhstanMongolia.pdf">"New Jewel Beetles (Coleoptera: Buprestidae) from the Cretaceous of Russia, Kazakhstan, and Mongolia"</a> (PDF). Paleontological Journal. 43 (3): 277–281. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2009PalJ...43..277A">2009PalJ...43..277A</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1134%2Fs0031030109030058">10.1134/s0031030109030058</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:129618839">129618839</a>. Archived from <a rel="nofollow" class="external text" href="http://fossilinsects.net/pdfs/Alexeev_2009_PalJ_BuprestidaeCretaceousRussiaKazakhstanMongolia.pdf">the original</a> (PDF) on July 18, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Paleontological+Journal&rft.atitle=New+Jewel+Beetles+%28Coleoptera%3A+Buprestidae%29+from+the+Cretaceous+of+Russia%2C+Kazakhstan%2C+and+Mongolia&rft.volume=43&rft.issue=3&rft.pages=277-281&rft.date=2009&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A129618839%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1134%2Fs0031030109030058&rft_id=info%3Abibcode%2F2009PalJ...43..277A&rft.aulast=Alexeev&rft.aufirst=A.+V.&rft_id=http%3A%2F%2Ffossilinsects.net%2Fpdfs%2FAlexeev_2009_PalJ_BuprestidaeCretaceousRussiaKazakhstanMongolia.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-42"><a href="#cite_ref-42">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChin,_K.Gill,_B._D.1996" class="citation journal cs1">Chin, K.; Gill, B. D. (1996). "Dinosaurs, dung beetles, and conifers; participants in a Cretaceous food web". PALAIOS. 11 (3): 280–285. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1996Palai..11..280C">1996Palai..11..280C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.2307%2F3515235">10.2307/3515235</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/3515235">3515235</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PALAIOS&rft.atitle=Dinosaurs%2C+dung+beetles%2C+and+conifers%3B+participants+in+a+Cretaceous+food+web&rft.volume=11&rft.issue=3&rft.pages=280-285&rft.date=1996&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F3515235%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.2307%2F3515235&rft_id=info%3Abibcode%2F1996Palai..11..280C&rft.au=Chin%2C+K.&rft.au=Gill%2C+B.+D.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-43"><a href="#cite_ref-43">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArillo,_AntonioOrtuño,_Vicente_M.2008" class="citation journal cs1">Arillo, Antonio & Ortuño, Vicente M. (2008). "Did dinosaurs have any relation with dung-beetles? (The origin of coprophagy)". <a href="/wiki/Journal_of_Natural_History" title="Journal of Natural History">Journal of Natural History</a>. 42 (19&20): 1405–1408. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2008JNatH..42.1405A">2008JNatH..42.1405A</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00222930802105130">10.1080/00222930802105130</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:83643794">83643794</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Natural+History&rft.atitle=Did+dinosaurs+have+any+relation+with+dung-beetles%3F+%28The+origin+of+coprophagy%29&rft.volume=42&rft.issue=19%2620&rft.pages=1405-1408&rft.date=2008&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A83643794%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1080%2F00222930802105130&rft_id=info%3Abibcode%2F2008JNatH..42.1405A&rft.au=Arillo%2C+Antonio&rft.au=Ortu%C3%B1o%2C+Vicente+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-amber-44"><a href="#cite_ref-amber_44-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDavid_PerisJes_Rust2020" class="citation journal cs1">David Peris; Jes Rust (2020). <a rel="nofollow" class="external text" href="https://academic.oup.com/zoolinnean/article/189/4/1085/5638889">"Cretaceous beetles (Insecta: Coleoptera) in amber: the palaeoecology of this most diverse group of insects"</a>. Zoological Journal of the Linnean Society. 189 (4): 1085–1104. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fzoolinnean%2Fzlz118">10.1093/zoolinnean/zlz118</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zoological+Journal+of+the+Linnean+Society&rft.atitle=Cretaceous+beetles+%28Insecta%3A+Coleoptera%29+in+amber%3A+the+palaeoecology+of+this+most+diverse+group+of+insects&rft.volume=189&rft.issue=4&rft.pages=1085-1104&rft.date=2020&rft_id=info%3Adoi%2F10.1093%2Fzoolinnean%2Fzlz118&rft.au=David+Peris&rft.au=Jes+Rust&rft_id=https%3A%2F%2Facademic.oup.com%2Fzoolinnean%2Farticle%2F189%2F4%2F1085%2F5638889&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-45"><a href="#cite_ref-45">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMcNamaraBriggsOrrNoh2011" class="citation journal cs1">McNamara, M. E.; Briggs, D. E. G.; Orr, P. J.; Noh, H.; Cao, H. (2011). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3267148">"The original colours of fossil beetles"</a>. Proceedings of the Royal Society B: Biological Sciences. 279 (1731): 1114–1121. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frspb.2011.1677">10.1098/rspb.2011.1677</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3267148">3267148</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/21957131">21957131</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+Royal+Society+B%3A+Biological+Sciences&rft.atitle=The+original+colours+of+fossil+beetles&rft.volume=279&rft.issue=1731&rft.pages=1114-1121&rft.date=2011&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3267148%23id-name%3DPMC&rft_id=info%3Apmid%2F21957131&rft_id=info%3Adoi%2F10.1098%2Frspb.2011.1677&rft.aulast=McNamara&rft.aufirst=M.+E.&rft.au=Briggs%2C+D.+E.+G.&rft.au=Orr%2C+P.+J.&rft.au=Noh%2C+H.&rft.au=Cao%2C+H.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3267148&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Coope-46"><a href="#cite_ref-Coope_46-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCoope1979" class="citation journal cs1">Coope, G. R. (1979). "Late Cenozoic Fossil Coleoptera: Evolution, Biogeography, and Ecology". Annual Review of Ecology and Systematics. 10: 246–267. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1146%2Fannurev.es.10.110179.001335">10.1146/annurev.es.10.110179.001335</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/2096792">2096792</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annual+Review+of+Ecology+and+Systematics&rft.atitle=Late+Cenozoic+Fossil+Coleoptera%3A+Evolution%2C+Biogeography%2C+and+Ecology&rft.volume=10&rft.pages=246-267&rft.date=1979&rft_id=info%3Adoi%2F10.1146%2Fannurev.es.10.110179.001335&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F2096792%23id-name%3DJSTOR&rft.aulast=Coope&rft.aufirst=G.+R.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-TOLweb_1995-47"><a href="#cite_ref-TOLweb_1995_47-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMaddison1995" class="citation web cs1">Maddison, D. R. (1995). <a rel="nofollow" class="external text" href="http://tolweb.org/Polyphaga/8870">"Polyphaga"</a>. Tree of Life web project. Retrieved February 27, 2016.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Tree+of+Life+web+project&rft.atitle=Polyphaga&rft.date=1995&rft.aulast=Maddison&rft.aufirst=D.+R.&rft_id=http%3A%2F%2Ftolweb.org%2FPolyphaga%2F8870&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-48"><a href="#cite_ref-48">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutel,_R._G.Lawrence,_J._F.2005" class="citation book cs1">Beutel, R. G.; Lawrence, J. F. (2005). "4. Coleoptera (Morphology)". In Kristensen, N. P.; Beutel, R. G. (eds.). Handbook of Zoology, Band 4: Arthropoda, 2: Insecta, Coleoptera, Beetles. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). p. 23. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-3-11-017130-3" title="Special:BookSources/978-3-11-017130-3"><bdi>978-3-11-017130-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=4.+Coleoptera+%28Morphology%29&rft.btitle=Handbook+of+Zoology%2C+Band+4%3A+Arthropoda%2C+2%3A+Insecta%2C+Coleoptera%2C+Beetles.+Volume+1%3A+Morphology+and+Systematics+%28Archostemata%2C+Adephaga%2C+Myxophaga%2C+Polyphaga+partim%29&rft.pages=23&rft.date=2005&rft.isbn=978-3-11-017130-3&rft.au=Beutel%2C+R.+G.&rft.au=Lawrence%2C+J.+F.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-49"><a href="#cite_ref-49">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutel,_R._G.Ribera,_I.2005" class="citation book cs1">Beutel, R. G.; Ribera, I. (2005). "7. Adephaga Schellenberg, 1806". In Kristensen, N. P.; Beutel, R. G. (eds.). Handbook of Zoology, Band 4: Arthropoda, 2: Insecta, Coleoptera, Beetles. Volume 1: Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). p. 54. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-3-11-017130-3" title="Special:BookSources/978-3-11-017130-3"><bdi>978-3-11-017130-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=7.+Adephaga+Schellenberg%2C+1806&rft.btitle=Handbook+of+Zoology%2C+Band+4%3A+Arthropoda%2C+2%3A+Insecta%2C+Coleoptera%2C+Beetles.+Volume+1%3A+Morphology+and+Systematics+%28Archostemata%2C+Adephaga%2C+Myxophaga%2C+Polyphaga+partim%29&rft.pages=54&rft.date=2005&rft.isbn=978-3-11-017130-3&rft.au=Beutel%2C+R.+G.&rft.au=Ribera%2C+I.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-50"><a href="#cite_ref-50">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://bugguide.net/node/view/40603">"Suborder Archostemata – Reticulated and Telephone-pole Beetles"</a>. BugGuide. 2006. Retrieved January 26, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Suborder+Archostemata+%E2%80%93+Reticulated+and+Telephone-pole+Beetles&rft.pub=BugGuide&rft.date=2006&rft_id=http%3A%2F%2Fbugguide.net%2Fnode%2Fview%2F40603&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-ausbeetle-51">^ <a href="#cite_ref-ausbeetle_51-0">a</a> <a href="#cite_ref-ausbeetle_51-1">b</a> <a href="#cite_ref-ausbeetle_51-2">c</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLawrenceŚlipiński2013" class="citation book cs1">Lawrence, John F.; Ślipiński, Adam (2013). Australian Beetles. Morphology, Classification and Keys. CSIRO. pp. 1–16. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-643-09728-5" title="Special:BookSources/978-0-643-09728-5"><bdi>978-0-643-09728-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Australian+Beetles.+Morphology%2C+Classification+and+Keys&rft.pages=1-16&rft.pub=CSIRO&rft.date=2013&rft.isbn=978-0-643-09728-5&rft.aulast=Lawrence&rft.aufirst=John+F.&rft.au=%C5%9Alipi%C5%84ski%2C+Adam&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-52"><a href="#cite_ref-52">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMesaros2013" class="citation journal cs1">Mesaros, Gabor (2013). <a rel="nofollow" class="external text" href="http://scindeks.ceon.rs/Article.aspx?artid=1820-95211306071M">"Sphaeriusidae (Coleoptera, Myxophaga): A new beetle family to the fauna of Serbia"</a>. Bulletin of the Natural History Museum (6): 71–74. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.5937%2Fbnhmb1306071m">10.5937/bnhmb1306071m</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Bulletin+of+the+Natural+History+Museum&rft.atitle=Sphaeriusidae+%28Coleoptera%2C+Myxophaga%29%3A+A+new+beetle+family+to+the+fauna+of+Serbia&rft.issue=6&rft.pages=71-74&rft.date=2013&rft_id=info%3Adoi%2F10.5937%2Fbnhmb1306071m&rft.aulast=Mesaros&rft.aufirst=Gabor&rft_id=http%3A%2F%2Fscindeks.ceon.rs%2FArticle.aspx%3Fartid%3D1820-95211306071M&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-53"><a href="#cite_ref-53">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutelLeschen2005" class="citation book cs1">Beutel, Rolf G.; Leschen, Richard (2005). Handbook of Zoology. Volume 4. Part 38. Arthropoda. Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). Walter de Gruyter. p. 43.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Handbook+of+Zoology.+Volume+4.+Part+38.+Arthropoda.+Morphology+and+Systematics+%28Archostemata%2C+Adephaga%2C+Myxophaga%2C+Polyphaga+partim%29&rft.pages=43&rft.pub=Walter+de+Gruyter&rft.date=2005&rft.aulast=Beutel&rft.aufirst=Rolf+G.&rft.au=Leschen%2C+Richard&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-whiting-54"><a href="#cite_ref-whiting_54-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWhiting,_Michael_F.2002" class="citation journal cs1">Whiting, Michael F. (2002). "Phylogeny of the holometabolous insect orders: molecular evidence". <a href="/wiki/Zoologica_Scripta" title="Zoologica Scripta">Zoologica Scripta</a>. 31 (1): 3–15. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1046%2Fj.0300-3256.2001.00093.x">10.1046/j.0300-3256.2001.00093.x</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:45544978">45544978</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zoologica+Scripta&rft.atitle=Phylogeny+of+the+holometabolous+insect+orders%3A+molecular+evidence&rft.volume=31&rft.issue=1&rft.pages=3-15&rft.date=2002&rft_id=info%3Adoi%2F10.1046%2Fj.0300-3256.2001.00093.x&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A45544978%23id-name%3DS2CID&rft.au=Whiting%2C+Michael+F.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Beutel-55"><a href="#cite_ref-Beutel_55-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeutelHaas,_F.2000" class="citation journal cs1">Beutel, R.; Haas, F. (2000). <a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1096-0031.2000.tb00350.x">"Phylogenetic relationships of the suborders of Coleoptera (Insecta)"</a>. <a href="/wiki/Cladistics_(journal)" title="Cladistics (journal)">Cladistics</a>. 16 (1): 103–141. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1096-0031.2000.tb00350.x">10.1111/j.1096-0031.2000.tb00350.x</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/34902922">34902922</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:56131113">56131113</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cladistics&rft.atitle=Phylogenetic+relationships+of+the+suborders+of+Coleoptera+%28Insecta%29&rft.volume=16&rft.issue=1&rft.pages=103-141&rft.date=2000&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A56131113%23id-name%3DS2CID&rft_id=info%3Apmid%2F34902922&rft_id=info%3Adoi%2F10.1111%2Fj.1096-0031.2000.tb00350.x&rft.aulast=Beutel&rft.aufirst=R.&rft.au=Haas%2C+F.&rft_id=https%3A%2F%2Fdoi.org%2F10.1111%252Fj.1096-0031.2000.tb00350.x&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Kukalová-56">^ <a href="#cite_ref-Kukalová_56-0">a</a> <a href="#cite_ref-Kukalová_56-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKukalová-PeckLawrence,_J._F.1993" class="citation journal cs1">Kukalová-Peck, J.; Lawrence, J. F. (1993). "Evolution of the hind wing in Coleoptera". <a href="/wiki/Canadian_Entomologist" class="mw-redirect" title="Canadian Entomologist">Canadian Entomologist</a>. 125 (2): 181–258. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.4039%2FEnt125181-2">10.4039/Ent125181-2</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:52888506">52888506</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Canadian+Entomologist&rft.atitle=Evolution+of+the+hind+wing+in+Coleoptera&rft.volume=125&rft.issue=2&rft.pages=181-258&rft.date=1993&rft_id=info%3Adoi%2F10.4039%2FEnt125181-2&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A52888506%23id-name%3DS2CID&rft.aulast=Kukalov%C3%A1-Peck&rft.aufirst=J.&rft.au=Lawrence%2C+J.+F.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-57"><a href="#cite_ref-57">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMaddisonMooreBakerEllis2009" class="citation journal cs1">Maddison, D. R.; Moore, W.; Baker, M. D.; Ellis, T. M.; Ober, K. A.; Cannone, J. J.; Gutell, R. R. (2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2752903">"Monophyly of terrestrial adephagan beetles as indicated by three nuclear genes (Coleoptera: Carabidae and Trachypachidae)"</a>. Zoologica Scripta. 38 (1): 43–62. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1463-6409.2008.00359.x">10.1111/j.1463-6409.2008.00359.x</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2752903">2752903</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19789725">19789725</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Zoologica+Scripta&rft.atitle=Monophyly+of+terrestrial+adephagan+beetles+as+indicated+by+three+nuclear+genes+%28Coleoptera%3A+Carabidae+and+Trachypachidae%29&rft.volume=38&rft.issue=1&rft.pages=43-62&rft.date=2009&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2752903%23id-name%3DPMC&rft_id=info%3Apmid%2F19789725&rft_id=info%3Adoi%2F10.1111%2Fj.1463-6409.2008.00359.x&rft.aulast=Maddison&rft.aufirst=D.+R.&rft.au=Moore%2C+W.&rft.au=Baker%2C+M.+D.&rft.au=Ellis%2C+T.+M.&rft.au=Ober%2C+K.+A.&rft.au=Cannone%2C+J.+J.&rft.au=Gutell%2C+R.+R.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2752903&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-58"><a href="#cite_ref-58">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMaddison2000" class="citation web cs1">Maddison, David R. (September 11, 2000). <a rel="nofollow" class="external text" href="http://tolweb.org/coleoptera">"Coleoptera. Beetle"</a>. <a href="/wiki/Tree_of_Life_Web_Project" title="Tree of Life Web Project">Tree of Life Web Project</a>. tolweb.org. Retrieved March 18, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Tree+of+Life+Web+Project&rft.atitle=Coleoptera.+Beetle&rft.date=2000-09-11&rft.aulast=Maddison&rft.aufirst=David+R.&rft_id=http%3A%2F%2Ftolweb.org%2Fcoleoptera&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-59"><a href="#cite_ref-59">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNiehuis,_OliverHartig,_GerritGrath,_Sonja2012" class="citation journal cs1">Niehuis, Oliver; Hartig, Gerrit; Grath, Sonja; et al. (2012). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2012.05.018">"Genomic and Morphological Evidence Converge to Resolve the Enigma of Strepsiptera"</a>. Current Biology. 22 (14): 1309–1313. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2012.05.018">10.1016/j.cub.2012.05.018</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/22704986">22704986</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Biology&rft.atitle=Genomic+and+Morphological+Evidence+Converge+to+Resolve+the+Enigma+of+Strepsiptera&rft.volume=22&rft.issue=14&rft.pages=1309-1313&rft.date=2012&rft_id=info%3Adoi%2F10.1016%2Fj.cub.2012.05.018&rft_id=info%3Apmid%2F22704986&rft.au=Niehuis%2C+Oliver&rft.au=Hartig%2C+Gerrit&rft.au=Grath%2C+Sonja&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.cub.2012.05.018&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-McKenna2015-60">^ <a href="#cite_ref-McKenna2015_60-0">a</a> <a href="#cite_ref-McKenna2015_60-1">b</a> <a href="#cite_ref-McKenna2015_60-2">c</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMcKennaWild2015" class="citation journal cs1">McKenna, Duane D.; Wild, Alexander L.; et al. (2015). <a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fsyen.12132">"The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution"</a>. Systematic Entomology. 40 (4): 835–880. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2015SysEn..40..835M">2015SysEn..40..835M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fsyen.12132">10.1111/syen.12132</a>. <a href="/wiki/Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<a rel="nofollow" class="external text" href="https://hdl.handle.net/10057%2F11540">10057/11540</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Systematic+Entomology&rft.atitle=The+beetle+tree+of+life+reveals+that+Coleoptera+survived+end-Permian+mass+extinction+to+diversify+during+the+Cretaceous+terrestrial+revolution&rft.volume=40&rft.issue=4&rft.pages=835-880&rft.date=2015&rft_id=info%3Ahdl%2F10057%2F11540&rft_id=info%3Adoi%2F10.1111%2Fsyen.12132&rft_id=info%3Abibcode%2F2015SysEn..40..835M&rft.aulast=McKenna&rft.aufirst=Duane+D.&rft.au=Wild%2C+Alexander+L.&rft_id=https%3A%2F%2Fdoi.org%2F10.1111%252Fsyen.12132&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Hunt2007-61">^ <a href="#cite_ref-Hunt2007_61-0">a</a> <a href="#cite_ref-Hunt2007_61-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHunt2007" class="citation journal cs1">Hunt, Toby; et al. (2007). "A Comprehensive Phylogeny of Beetles Reveals the Evolutionary Origins of a Superradiation". <a href="/wiki/Science_(journal)" title="Science (journal)">Science</a>. 318 (5858): 1913–19116. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2007Sci...318.1913H">2007Sci...318.1913H</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.1146954">10.1126/science.1146954</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18096805">18096805</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:19392955">19392955</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=A+Comprehensive+Phylogeny+of+Beetles+Reveals+the+Evolutionary+Origins+of+a+Superradiation&rft.volume=318&rft.issue=5858&rft.pages=1913-19116&rft.date=2007&rft_id=info%3Adoi%2F10.1126%2Fscience.1146954&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A19392955%23id-name%3DS2CID&rft_id=info%3Apmid%2F18096805&rft_id=info%3Abibcode%2F2007Sci...318.1913H&rft.aulast=Hunt&rft.aufirst=Toby&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-63"><a href="#cite_ref-63">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://entnemdept.ufl.edu/choate/beetles.pdf">"Introduction to the Identification of Beetles (Coleoptera)"</a> (PDF). University of Florida. Retrieved March 15, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Introduction+to+the+Identification+of+Beetles+%28Coleoptera%29&rft.pub=University+of+Florida&rft_id=http%3A%2F%2Fentnemdept.ufl.edu%2Fchoate%2Fbeetles.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Lau-64"><a href="#cite_ref-Lau_64-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLau_T.F.S.Meyer-Rochow_V.B.2006" class="citation journal cs1">Lau T.F.S.; Meyer-Rochow V.B. (2006). "Sexual dimorphism in the compound eye of Rhagophthalmus ohbai (Coleoptera; Rhagophthalmidae): I. Morphology and ultrastructure". Journal of Asia-Pacific Entomology. 9: 19–30. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS1226-8615%2808%2960271-X">10.1016/S1226-8615(08)60271-X</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Asia-Pacific+Entomology&rft.atitle=Sexual+dimorphism+in+the+compound+eye+of+Rhagophthalmus+ohbai+%28Coleoptera%3B+Rhagophthalmidae%29%3A+I.+Morphology+and+ultrastructure&rft.volume=9&rft.pages=19-30&rft.date=2006&rft_id=info%3Adoi%2F10.1016%2FS1226-8615%2808%2960271-X&rft.au=Lau+T.F.S.&rft.au=Meyer-Rochow+V.B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-BeetlesAustralia-65">^ <a href="#cite_ref-BeetlesAustralia_65-0">a</a> <a href="#cite_ref-BeetlesAustralia_65-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHangay,_G.Zborowski,_P.2010" class="citation book cs1">Hangay, G.; Zborowski, P. (2010). <a rel="nofollow" class="external text" href="https://archive.org/details/guidetobeetlesau00hang">A Guide to the Beetles of Australia</a>. CSIRO. p. <a rel="nofollow" class="external text" href="https://archive.org/details/guidetobeetlesau00hang/page/n245">10</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-643-09487-1" title="Special:BookSources/978-0-643-09487-1"><bdi>978-0-643-09487-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=A+Guide+to+the+Beetles+of+Australia&rft.pages=10&rft.pub=CSIRO&rft.date=2010&rft.isbn=978-0-643-09487-1&rft.au=Hangay%2C+G.&rft.au=Zborowski%2C+P.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fguidetobeetlesau00hang&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-beetles-66"><a href="#cite_ref-beetles_66-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGokan_N.Meyer-Rochow_V.B.2000" class="citation journal cs1">Gokan N.; Meyer-Rochow V.B. (2000). "Morphological comparisons of compound eyes in Scarabaeoidea (Coleoptera) related to the beetles' daily activity maxima and phylogenetic positions". Journal of Agricultural Science (Tokyo Nogyo Daigaku). 45 (1): 15–61.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Agricultural+Science+%28Tokyo+Nogyo+Daigaku%29&rft.atitle=Morphological+comparisons+of+compound+eyes+in+Scarabaeoidea+%28Coleoptera%29+related+to+the+beetles%27+daily+activity+maxima+and+phylogenetic+positions&rft.volume=45&rft.issue=1&rft.pages=15-61&rft.date=2000&rft.au=Gokan+N.&rft.au=Meyer-Rochow+V.B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Antennae-67">^ <a href="#cite_ref-Antennae_67-0">a</a> <a href="#cite_ref-Antennae_67-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBenisch2007" class="citation web cs1">Benisch, Christoph (2007). <a rel="nofollow" class="external text" href="http://www.kerbtier.de/Pages/Themenseiten/enKoerperbau.html#Fuehler">"3. Antennae, Beetle morphology"</a>. Kerbtier.de (Beetle fauna of Germany). Retrieved March 14, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=3.+Antennae%2C+Beetle+morphology&rft.pub=Kerbtier.de+%28Beetle+fauna+of+Germany%29&rft.date=2007&rft.aulast=Benisch&rft.aufirst=Christoph&rft_id=http%3A%2F%2Fwww.kerbtier.de%2FPages%2FThemenseiten%2FenKoerperbau.html%23Fuehler&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-68"><a href="#cite_ref-68">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZurekGilbert2014" class="citation journal cs1">Zurek, D.B.; Gilbert, C. (2014). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3924084">"Static antennae act as locomotory guides that compensate for visual motion blur in a diurnal, keen-eyed predator"</a>. Proceedings of the Royal Society B: Biological Sciences. 281 (1779): 20133072. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frspb.2013.3072">10.1098/rspb.2013.3072</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3924084">3924084</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24500171">24500171</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Proceedings+of+the+Royal+Society+B%3A+Biological+Sciences&rft.atitle=Static+antennae+act+as+locomotory+guides+that+compensate+for+visual+motion+blur+in+a+diurnal%2C+keen-eyed+predator&rft.volume=281&rft.issue=1779&rft.pages=20133072&rft.date=2014&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3924084%23id-name%3DPMC&rft_id=info%3Apmid%2F24500171&rft_id=info%3Adoi%2F10.1098%2Frspb.2013.3072&rft.aulast=Zurek&rft.aufirst=D.B.&rft.au=Gilbert%2C+C.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3924084&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-69"><a href="#cite_ref-69">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBerkovRodríguezCenteno2007" class="citation journal cs1">Berkov, Amy; Rodríguez, Nelson; Centeno, Pedro (2007). "Convergent evolution in the antennae of a cerambycid beetle, Onychocerus albitarsis, and the sting of a scorpion". Naturwissenschaften. 95 (3): 257–61. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2008NW.....95..257B">2008NW.....95..257B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00114-007-0316-1">10.1007/s00114-007-0316-1</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18004534">18004534</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:30226487">30226487</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Naturwissenschaften&rft.atitle=Convergent+evolution+in+the+antennae+of+a+cerambycid+beetle%2C+Onychocerus+albitarsis%2C+and+the+sting+of+a+scorpion&rft.volume=95&rft.issue=3&rft.pages=257-61&rft.date=2007&rft_id=info%3Adoi%2F10.1007%2Fs00114-007-0316-1&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A30226487%23id-name%3DS2CID&rft_id=info%3Apmid%2F18004534&rft_id=info%3Abibcode%2F2008NW.....95..257B&rft.aulast=Berkov&rft.aufirst=Amy&rft.au=Rodr%C3%ADguez%2C+Nelson&rft.au=Centeno%2C+Pedro&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-70"><a href="#cite_ref-70">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20180222221339/http://bugs.bio.usyd.edu.au/learning/resources/Entomology/externalMorphology/antennaTypes.html">"Antennae Types"</a>. University of Sydney. Archived from <a rel="nofollow" class="external text" href="http://bugs.bio.usyd.edu.au/learning/resources/Entomology/externalMorphology/antennaTypes.html">the original</a> on February 22, 2018. Retrieved January 26, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Antennae+Types&rft.pub=University+of+Sydney&rft_id=http%3A%2F%2Fbugs.bio.usyd.edu.au%2Flearning%2Fresources%2FEntomology%2FexternalMorphology%2FantennaTypes.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-71"><a href="#cite_ref-71">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArnett,_R._H._Jr.Thomas,_M._C.2001" class="citation book cs1">Arnett, R. H. Jr.; Thomas, M. C. (2001). American Beetles, Volume I: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia. CRC Press. pp. 3–7. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4822-7432-5" title="Special:BookSources/978-1-4822-7432-5"><bdi>978-1-4822-7432-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=American+Beetles%2C+Volume+I%3A+Archostemata%2C+Myxophaga%2C+Adephaga%2C+Polyphaga%3A+Staphyliniformia&rft.pages=3-7&rft.pub=CRC+Press&rft.date=2001&rft.isbn=978-1-4822-7432-5&rft.au=Arnett%2C+R.+H.+Jr.&rft.au=Thomas%2C+M.+C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-72"><a href="#cite_ref-72">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGreenKovalevSvenssonGorb2013" class="citation journal cs1">Green, Kristina Karlsson; Kovalev, Alexander; Svensson, Erik I.; Gorb, Stanislav N. (2013). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3730688">"Male clasping ability, female polymorphism and sexual conflict: fine-scale elytral morphology as a sexually antagonistic adaptation in female diving beetles"</a>. Journal of the Royal Society Interface. 10 (86): 20130409. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsif.2013.0409">10.1098/rsif.2013.0409</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3730688">3730688</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/23825114">23825114</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+Royal+Society+Interface&rft.atitle=Male+clasping+ability%2C+female+polymorphism+and+sexual+conflict%3A+fine-scale+elytral+morphology+as+a+sexually+antagonistic+adaptation+in+female+diving+beetles&rft.volume=10&rft.issue=86&rft.pages=20130409&rft.date=2013&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3730688%23id-name%3DPMC&rft_id=info%3Apmid%2F23825114&rft_id=info%3Adoi%2F10.1098%2Frsif.2013.0409&rft.aulast=Green&rft.aufirst=Kristina+Karlsson&rft.au=Kovalev%2C+Alexander&rft.au=Svensson%2C+Erik+I.&rft.au=Gorb%2C+Stanislav+N.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3730688&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-73"><a href="#cite_ref-73">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBurkness,_S.Hahn,_J.2007" class="citation web cs1">Burkness, S.; Hahn, J. (2007). <a rel="nofollow" class="external text" href="http://www.extension.umn.edu/garden/insects/find/flea-beetles/">"Flea beetles in home gardens"</a>. <a href="/wiki/University_of_Minnesota_Extension" class="mw-redirect" title="University of Minnesota Extension">University of Minnesota Extension</a>. Retrieved January 26, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Flea+beetles+in+home+gardens&rft.pub=University+of+Minnesota+Extension&rft.date=2007&rft.au=Burkness%2C+S.&rft.au=Hahn%2C+J.&rft_id=http%3A%2F%2Fwww.extension.umn.edu%2Fgarden%2Finsects%2Ffind%2Fflea-beetles%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Carpenter-74">^ <a href="#cite_ref-Carpenter_74-0">a</a> <a href="#cite_ref-Carpenter_74-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCarpenter1899" class="citation book cs1">Carpenter, George Herbert (1899). Insects, their structure and life.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Insects%2C+their+structure+and+life&rft.date=1899&rft.aulast=Carpenter&rft.aufirst=George+Herbert&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-75"><a href="#cite_ref-75">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPhilips,_ChrisFread,_ElizabethKuhar,_Tom" class="citation cs2">Philips, Chris; Fread, Elizabeth; Kuhar, Tom, <a rel="nofollow" class="external text" href="https://web.archive.org/web/20161130002157/http://www.pubs.ext.vt.edu/ENTO/ENTO-53/ENTO-53-pdf.pdf">Leatherwing (Soldier) Beetles</a> (PDF), archived from <a rel="nofollow" class="external text" href="http://pubs.ext.vt.edu/ENTO/ENTO-53/ENTO-53-pdf.pdf">the original</a> (PDF) on November 30, 2016, retrieved March 14, 2017</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Leatherwing+%28Soldier%29+Beetles&rft.au=Philips%2C+Chris&rft.au=Fread%2C+Elizabeth&rft.au=Kuhar%2C+Tom&rft_id=http%3A%2F%2Fpubs.ext.vt.edu%2FENTO%2FENTO-53%2FENTO-53-pdf.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-76"><a href="#cite_ref-76">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDonald_W._HallMarc_A._Branham2016" class="citation web cs1">Donald W. Hall; Marc A. Branham (2016). <a rel="nofollow" class="external text" href="http://entomology.ifas.ufl.edu/creatures/misc/beetles/banded_net-winged_beetle.htm">"Calopteron discrepans (Newman) (Insecta: Coleoptera: Lycidae)"</a>. <a href="/wiki/University_of_Florida" title="University of Florida">University of Florida</a>. Retrieved March 15, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Calopteron+discrepans+%28Newman%29+%28Insecta%3A+Coleoptera%3A+Lycidae%29&rft.pub=University+of+Florida&rft.date=2016&rft.au=Donald+W.+Hall&rft.au=Marc+A.+Branham&rft_id=http%3A%2F%2Fentomology.ifas.ufl.edu%2Fcreatures%2Fmisc%2Fbeetles%2Fbanded_net-winged_beetle.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-77"><a href="#cite_ref-77">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLawrence,_J._F.Hastings,_A._M.Dallwitz,_M._J.Paine,_T._A.2005" class="citation web cs1">Lawrence, J. F.; Hastings, A. M.; Dallwitz, M. J.; Paine, T. A.; Zurcher, E. J. (2005). <a rel="nofollow" class="external text" href="http://delta-intkey.com/elateria/www/phenmplf.htm">"Elateriformia (Coleoptera): descriptions, illustrations, identification, and information retrieval for families and subfamilies"</a>. Retrieved January 26, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Elateriformia+%28Coleoptera%29%3A+descriptions%2C+illustrations%2C+identification%2C+and+information+retrieval+for+families+and+subfamilies&rft.date=2005&rft.au=Lawrence%2C+J.+F.&rft.au=Hastings%2C+A.+M.&rft.au=Dallwitz%2C+M.+J.&rft.au=Paine%2C+T.+A.&rft.au=Zurcher%2C+E.+J.&rft_id=http%3A%2F%2Fdelta-intkey.com%2Felateria%2Fwww%2Fphenmplf.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-78"><a href="#cite_ref-78">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeenen,_RWinkelman,_J._K.2001" class="citation journal cs1 cs1-prop-foreign-lang-source">Beenen, R; Winkelman, J. K. (2001). "Aantekeningen over Chrysomelidae in Nederland 5". Entomologische Berichten (in Dutch). 61: 63–67.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Entomologische+Berichten&rft.atitle=Aantekeningen+over+Chrysomelidae+in+Nederland+5&rft.volume=61&rft.pages=63-67&rft.date=2001&rft.au=Beenen%2C+R&rft.au=Winkelman%2C+J.+K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-American-79"><a href="#cite_ref-American_79-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArnettThomas2000" class="citation book cs1">Arnett, Ross H. Jr; Thomas, Michael C. (2000). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=bJjSBQAAQBAJ&pg=PA8">American Beetles, Volume I: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia</a>. CRC Press. p. 8. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4822-7432-5" title="Special:BookSources/978-1-4822-7432-5"><bdi>978-1-4822-7432-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=American+Beetles%2C+Volume+I%3A+Archostemata%2C+Myxophaga%2C+Adephaga%2C+Polyphaga%3A+Staphyliniformia&rft.pages=8&rft.pub=CRC+Press&rft.date=2000&rft.isbn=978-1-4822-7432-5&rft.aulast=Arnett&rft.aufirst=Ross+H.+Jr&rft.au=Thomas%2C+Michael+C.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DbJjSBQAAQBAJ%26pg%3DPA8&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-80"><a href="#cite_ref-80">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSeymourMatthews2012" class="citation journal cs1">Seymour, Roger S.; Matthews, Philip G. D. (2012). <a rel="nofollow" class="external text" href="https://doi.org/10.1242%2Fjeb.070276">"Physical gills in diving insects and spiders: Theory and experiment"</a>. The Journal of Experimental Biology. 216 (2): 164–70. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1242%2Fjeb.070276">10.1242/jeb.070276</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/23255190">23255190</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Journal+of+Experimental+Biology&rft.atitle=Physical+gills+in+diving+insects+and+spiders%3A+Theory+and+experiment&rft.volume=216&rft.issue=2&rft.pages=164-70&rft.date=2012&rft_id=info%3Adoi%2F10.1242%2Fjeb.070276&rft_id=info%3Apmid%2F23255190&rft.aulast=Seymour&rft.aufirst=Roger+S.&rft.au=Matthews%2C+Philip+G.+D.&rft_id=https%3A%2F%2Fdoi.org%2F10.1242%252Fjeb.070276&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-81"><a href="#cite_ref-81">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSchmidt-Nielsen1997" class="citation book cs1">Schmidt-Nielsen, Knut (January 15, 1997). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=Af7IwQWJoCMC&pg=PA55">"Insect Respiration"</a>. Animal Physiology: Adaptation and Environment (5th ed.). <a href="/wiki/Cambridge_University_Press" title="Cambridge University Press">Cambridge University Press</a>. p. 55. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-521-57098-5" title="Special:BookSources/978-0-521-57098-5"><bdi>978-0-521-57098-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Insect+Respiration&rft.btitle=Animal+Physiology%3A+Adaptation+and+Environment&rft.pages=55&rft.edition=5th&rft.pub=Cambridge+University+Press&rft.date=1997-01-15&rft.isbn=978-0-521-57098-5&rft.aulast=Schmidt-Nielsen&rft.aufirst=Knut&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DAf7IwQWJoCMC%26pg%3DPA55&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-82"><a href="#cite_ref-82">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMiller1985" class="citation book cs1">Miller, T. A. (1985). "Chapter 8: Structure and Physiology of the Circulatory System". In Kerkut, G. A.; Gilbert, L. I. (eds.). Comprehensive Insect Physiology, Biochemistry and Pharmacology. Volume 3: Integument, Respiration and Circulation. Pergamom Press. pp. 289–355. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-08-030804-3" title="Special:BookSources/978-0-08-030804-3"><bdi>978-0-08-030804-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Chapter+8%3A+Structure+and+Physiology+of+the+Circulatory+System&rft.btitle=Comprehensive+Insect+Physiology%2C+Biochemistry+and+Pharmacology.+Volume+3%3A+Integument%2C+Respiration+and+Circulation&rft.pages=289-355&rft.pub=Pergamom+Press&rft.date=1985&rft.isbn=978-0-08-030804-3&rft.aulast=Miller&rft.aufirst=T.+A.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Scoble-83"><a href="#cite_ref-Scoble_83-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFScoble,_M._J.1992" class="citation book cs1">Scoble, M. J. (1992). <a rel="nofollow" class="external text" href="https://archive.org/details/lepidopteraformf0000scob">The Lepidoptera: Form, function, and diversity</a>. Oxford University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4020-6242-1" title="Special:BookSources/978-1-4020-6242-1"><bdi>978-1-4020-6242-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Lepidoptera%3A+Form%2C+function%2C+and+diversity&rft.pub=Oxford+University+Press&rft.date=1992&rft.isbn=978-1-4020-6242-1&rft.au=Scoble%2C+M.+J.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Flepidopteraformf0000scob&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Winkler-84">^ <a href="#cite_ref-Winkler_84-0">a</a> <a href="#cite_ref-Winkler_84-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWinkler1964" class="citation book cs1">Winkler, Josef Rudolf (1964). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=xP5CAAAAYAAJ">A Book of Beetles</a>. <a href="/wiki/Spring_Books" class="mw-redirect" title="Spring Books">Spring Books</a> Science. pp. 30–32.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=A+Book+of+Beetles&rft.pages=30-32&rft.pub=Spring+Books+Science&rft.date=1964&rft.aulast=Winkler&rft.aufirst=Josef+Rudolf&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DxP5CAAAAYAAJ&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-85"><a href="#cite_ref-85">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOzbek,_H.Szalokia,_D.1998" class="citation journal cs1">Ozbek, H.; Szalokia, D. (1998). <a rel="nofollow" class="external text" href="https://www.researchgate.net/publication/242221698">"A contribution to the knowledge of the Meloidae (Coleoptera) fauna of Turkey along with new record"</a>. Turkish Journal of Zoology. 22: 23–40.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Turkish+Journal+of+Zoology&rft.atitle=A+contribution+to+the+knowledge+of+the+Meloidae+%28Coleoptera%29+fauna+of+Turkey+along+with+new+record&rft.volume=22&rft.pages=23-40&rft.date=1998&rft.au=Ozbek%2C+H.&rft.au=Szalokia%2C+D.&rft_id=https%3A%2F%2Fwww.researchgate.net%2Fpublication%2F242221698&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-86"><a href="#cite_ref-86">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMedvedev,_L._N.Pavlov,_S._I.1988" class="citation journal cs1">Medvedev, L. N.; Pavlov, S. I. (1988). "Mating behavior of the Chrysomelidae (Coleoptera)". Entomological Review. 67: 100–108.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Entomological+Review&rft.atitle=Mating+behavior+of+the+Chrysomelidae+%28Coleoptera%29&rft.volume=67&rft.pages=100-108&rft.date=1988&rft.au=Medvedev%2C+L.+N.&rft.au=Pavlov%2C+S.+I.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-87"><a href="#cite_ref-87">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMatsumuraKovalevGorb2017" class="citation journal cs1">Matsumura, Yoko; Kovalev, Alexander E.; Gorb, Stanislav N. (December 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5738233">"Penetration mechanics of a beetle intromittent organ with bending stiffness gradient and a soft tip"</a>. Science Advances. 3 (12): eaao5469. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fsciadv.aao5469">10.1126/sciadv.aao5469</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2375-2548">2375-2548</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5738233">5738233</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/29279866">29279866</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science+Advances&rft.atitle=Penetration+mechanics+of+a+beetle+intromittent+organ+with+bending+stiffness+gradient+and+a+soft+tip&rft.volume=3&rft.issue=12&rft.pages=eaao5469&rft.date=2017-12&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5738233%23id-name%3DPMC&rft.issn=2375-2548&rft_id=info%3Apmid%2F29279866&rft_id=info%3Adoi%2F10.1126%2Fsciadv.aao5469&rft.aulast=Matsumura&rft.aufirst=Yoko&rft.au=Kovalev%2C+Alexander+E.&rft.au=Gorb%2C+Stanislav+N.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5738233&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Crowson-88"><a href="#cite_ref-Crowson_88-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCrowson2013" class="citation book cs1">Crowson, R. A. (2013). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=WMXYBAAAQBAJ&pg=PA360">The Biology of the Coleoptera</a>. <a href="/wiki/Elsevier" title="Elsevier">Elsevier</a> Science. pp. 358–370. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4832-1760-4" title="Special:BookSources/978-1-4832-1760-4"><bdi>978-1-4832-1760-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Biology+of+the+Coleoptera&rft.pages=358-370&rft.pub=Elsevier+Science&rft.date=2013&rft.isbn=978-1-4832-1760-4&rft.aulast=Crowson&rft.aufirst=R.+A.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DWMXYBAAAQBAJ%26pg%3DPA360&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-89"><a href="#cite_ref-89">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeckBharadwaj1972" class="citation journal cs1">Beck, S. D.; Bharadwaj, R. K. (1972). "Reversed development and cellular ageing in an insect". <a href="/wiki/Science_(journal)" title="Science (journal)">Science</a>. 178 (4066): 1210–1211. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1972Sci...178.1210B">1972Sci...178.1210B</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.178.4066.1210">10.1126/science.178.4066.1210</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/4637808">4637808</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:34101370">34101370</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Reversed+development+and+cellular+ageing+in+an+insect&rft.volume=178&rft.issue=4066&rft.pages=1210-1211&rft.date=1972&rft_id=info%3Adoi%2F10.1126%2Fscience.178.4066.1210&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A34101370%23id-name%3DS2CID&rft_id=info%3Apmid%2F4637808&rft_id=info%3Abibcode%2F1972Sci...178.1210B&rft.aulast=Beck&rft.aufirst=S.+D.&rft.au=Bharadwaj%2C+R.+K.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-90"><a href="#cite_ref-90">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://www.amentsoc.org/insects/glossary/terms/scarabaeiform">"Definition of 'Scarabaeiform'"</a>. <a href="/wiki/Amateur_Entomologists%27_Society" title="Amateur Entomologists' Society">Amateur Entomologists' Society</a>. Retrieved January 27, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Definition+of+%27Scarabaeiform%27&rft.pub=Amateur+Entomologists%27+Society&rft_id=https%3A%2F%2Fwww.amentsoc.org%2Finsects%2Fglossary%2Fterms%2Fscarabaeiform&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-91"><a href="#cite_ref-91">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKrinsky,_W._L.2009" class="citation book cs1">Krinsky, W. L. (2009). "8 Beetles (Coleoptera)". In Mullen, G. R.; Durden, L. A. (eds.). Medical and Veterinary Entomology (2nd ed.). <a href="/wiki/Elsevier" title="Elsevier">Elsevier</a>. pp. 101–115. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-372500-4" title="Special:BookSources/978-0-12-372500-4"><bdi>978-0-12-372500-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=8+Beetles+%28Coleoptera%29&rft.btitle=Medical+and+Veterinary+Entomology&rft.pages=101-115&rft.edition=2nd&rft.pub=Elsevier&rft.date=2009&rft.isbn=978-0-12-372500-4&rft.au=Krinsky%2C+W.+L.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-92"><a href="#cite_ref-92">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://bugguide.net/node/view/149686">"Hypermetamorphosis of Striped Blister Beetle – Epicauta vittata"</a>. BugGuide. 2007. Retrieved January 27, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Hypermetamorphosis+of+Striped+Blister+Beetle+%E2%80%93+Epicauta+vittata&rft.pub=BugGuide&rft.date=2007&rft_id=http%3A%2F%2Fbugguide.net%2Fnode%2Fview%2F149686&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Zeng-93"><a href="#cite_ref-Zeng_93-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZeng1995" class="citation web cs1">Zeng, Yong (1995). <a rel="nofollow" class="external text" href="http://entnemdept.ifas.ufl.edu/walker/ufbir/chapters/chapter_12.shtml">"Longest Life Cycle"</a>. University of Florida. Retrieved March 17, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Longest+Life+Cycle&rft.pub=University+of+Florida&rft.date=1995&rft.aulast=Zeng&rft.aufirst=Yong&rft_id=http%3A%2F%2Fentnemdept.ifas.ufl.edu%2Fwalker%2Fufbir%2Fchapters%2Fchapter_12.shtml&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-94"><a href="#cite_ref-94">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFŠípekFabriziEberleAhrens2016" class="citation journal cs1">Šípek, Petr; Fabrizi, Silvia; Eberle, Jonas; Ahrens, Dirk (2016). <a rel="nofollow" class="external text" href="https://linkinghub.elsevier.com/retrieve/pii/S1055790316300938">"A molecular phylogeny of rose chafers (Coleoptera: Scarabaeidae: Cetoniinae) reveals a complex and concerted morphological evolution related to their flight mode"</a>. Molecular Phylogenetics and Evolution. 101: 163–175. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.ympev.2016.05.012">10.1016/j.ympev.2016.05.012</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/27165937">27165937</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Molecular+Phylogenetics+and+Evolution&rft.atitle=A+molecular+phylogeny+of+rose+chafers+%28Coleoptera%3A+Scarabaeidae%3A+Cetoniinae%29+reveals+a+complex+and+concerted+morphological+evolution+related+to+their+flight+mode&rft.volume=101&rft.pages=163-175&rft.date=2016&rft_id=info%3Adoi%2F10.1016%2Fj.ympev.2016.05.012&rft_id=info%3Apmid%2F27165937&rft.aulast=%C5%A0%C3%ADpek&rft.aufirst=Petr&rft.au=Fabrizi%2C+Silvia&rft.au=Eberle%2C+Jonas&rft.au=Ahrens%2C+Dirk&rft_id=https%3A%2F%2Flinkinghub.elsevier.com%2Fretrieve%2Fpii%2FS1055790316300938&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-95"><a href="#cite_ref-95">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFJeffries,_Daniel_L.Chapman,_JasonRoy,_Helen_E.Humphries,_Stuart2013" class="citation journal cs1">Jeffries, Daniel L.; Chapman, Jason; <a href="/wiki/Helen_Roy" title="Helen Roy">Roy, Helen E.</a>; Humphries, Stuart; Harrington, Richard; Brown, Peter M. J.; Handley, Lori-J. Lawson (2013). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3867359">"Characteristics and Drivers of High-Altitude Ladybird Flight: Insights from Vertical-Looking Entomological Radar"</a>. <a href="/wiki/PLOS_ONE" class="mw-redirect" title="PLOS ONE">PLOS ONE</a>. 8 (2): e82278. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2013PLoSO...882278J">2013PLoSO...882278J</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1371%2Fjournal.pone.0082278">10.1371/journal.pone.0082278</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3867359">3867359</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/24367512">24367512</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PLOS+ONE&rft.atitle=Characteristics+and+Drivers+of+High-Altitude+Ladybird+Flight%3A+Insights+from+Vertical-Looking+Entomological+Radar&rft.volume=8&rft.issue=2&rft.pages=e82278&rft.date=2013&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3867359%23id-name%3DPMC&rft_id=info%3Apmid%2F24367512&rft_id=info%3Adoi%2F10.1371%2Fjournal.pone.0082278&rft_id=info%3Abibcode%2F2013PLoSO...882278J&rft.au=Jeffries%2C+Daniel+L.&rft.au=Chapman%2C+Jason&rft.au=Roy%2C+Helen+E.&rft.au=Humphries%2C+Stuart&rft.au=Harrington%2C+Richard&rft.au=Brown%2C+Peter+M.+J.&rft.au=Handley%2C+Lori-J.+Lawson&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC3867359&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-96"><a href="#cite_ref-96">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCapinera2008" class="citation book cs1">Capinera, John L. (2008). <a rel="nofollow" class="external text" href="https://archive.org/details/encyclopediaento00capi_189">Encyclopedia of Entomology</a> (2 ed.). Springer Science & Business Media. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/encyclopediaento00capi_189/page/n3281">3218</a>–3219. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4020-6242-1" title="Special:BookSources/978-1-4020-6242-1"><bdi>978-1-4020-6242-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Encyclopedia+of+Entomology&rft.pages=3218-3219&rft.edition=2&rft.pub=Springer+Science+%26+Business+Media&rft.date=2008&rft.isbn=978-1-4020-6242-1&rft.aulast=Capinera&rft.aufirst=John+L.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fencyclopediaento00capi_189&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-97"><a href="#cite_ref-97">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFArnett,_R._H._Jr.Thomas,_M._C.2001" class="citation book cs1">Arnett, R. H. Jr.; Thomas, M. C. (2001). "Haliplidae". American Beetles, Volume 1. CRC Press. pp. 138–143. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8493-1925-9" title="Special:BookSources/978-0-8493-1925-9"><bdi>978-0-8493-1925-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Haliplidae&rft.btitle=American+Beetles%2C+Volume+1&rft.pages=138-143&rft.pub=CRC+Press&rft.date=2001&rft.isbn=978-0-8493-1925-9&rft.au=Arnett%2C+R.+H.+Jr.&rft.au=Thomas%2C+M.+C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Beetle_love-98"><a href="#cite_ref-Beetle_love_98-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.pc.gc.ca/eng/docs/v-g/dpp-mpb/sec2/dpp-mpb2b.aspx">"Mountain Pine Beetle – Beetle Love"</a>. Parks Canada. Retrieved March 13, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Mountain+Pine+Beetle+%E2%80%93+Beetle+Love&rft.pub=Parks+Canada&rft_id=http%3A%2F%2Fwww.pc.gc.ca%2Feng%2Fdocs%2Fv-g%2Fdpp-mpb%2Fsec2%2Fdpp-mpb2b.aspx&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-stridulation-99">^ <a href="#cite_ref-stridulation_99-0">a</a> <a href="#cite_ref-stridulation_99-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMeyer-Rochow1971" class="citation journal cs1">Meyer-Rochow, V.B. (1971). "Observations on stridulating Australian beetles (Hydrophilidae, Cerambycidae, Passalidae, Dynastinae) using scanning electron microscopical and electrophysiological techniques". Forma et Functio. 4: 326–339.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Forma+et+Functio&rft.atitle=Observations+on+stridulating+Australian+beetles+%28Hydrophilidae%2C+Cerambycidae%2C+Passalidae%2C+Dynastinae%29+using+scanning+electron+microscopical+and+electrophysiological+techniques&rft.volume=4&rft.pages=326-339&rft.date=1971&rft.aulast=Meyer-Rochow&rft.aufirst=V.B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-100"><a href="#cite_ref-100">^</a> Brandmayr P. 1992. Short review of the presocial evolution in Coleoptera. Ethol Ecol Evol. 4:7–16. </li> <li id="cite_note-101"><a href="#cite_ref-101">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWyatt,_T._D.Foster,_W._A.1989" class="citation journal cs1">Wyatt, T. D. & Foster, W. A. (1989). "Parental care in the subsocial intertidal beetle, Bledius spectabilis, in relation to parasitism by the ichneumonid wasp, Barycnemis blediator". <a href="/wiki/Behaviour_(journal)" title="Behaviour (journal)">Behaviour</a>. 110 (1–4): 76–92. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1163%2F156853989X00394">10.1163/156853989X00394</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/4534785">4534785</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Behaviour&rft.atitle=Parental+care+in+the+subsocial+intertidal+beetle%2C+Bledius+spectabilis%2C+in+relation+to+parasitism+by+the+ichneumonid+wasp%2C+Barycnemis+blediator&rft.volume=110&rft.issue=1%E2%80%934&rft.pages=76-92&rft.date=1989&rft_id=info%3Adoi%2F10.1163%2F156853989X00394&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F4534785%23id-name%3DJSTOR&rft.au=Wyatt%2C+T.+D.&rft.au=Foster%2C+W.+A.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-102"><a href="#cite_ref-102">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMilneMilne1944" class="citation journal cs1">Milne, Lorus J.; Milne, Margery J. (1944). "Notes on the Behavior of Burying Beetles (Nicrophorus spp.)". Journal of the New York Entomological Society. 52 (4): 311–327. <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/25005075">25005075</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+New+York+Entomological+Society&rft.atitle=Notes+on+the+Behavior+of+Burying+Beetles+%28Nicrophorus+spp.%29&rft.volume=52&rft.issue=4&rft.pages=311-327&rft.date=1944&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F25005075%23id-name%3DJSTOR&rft.aulast=Milne&rft.aufirst=Lorus+J.&rft.au=Milne%2C+Margery+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-103"><a href="#cite_ref-103">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHanskiYves1991" class="citation book cs1">Hanski, Ilkka; Yves, Cambefort (1991). Dung Beetle Ecology. Princeton University Press. pp. 626–672. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-691-08739-9" title="Special:BookSources/978-0-691-08739-9"><bdi>978-0-691-08739-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Dung+Beetle+Ecology&rft.pages=626-672&rft.pub=Princeton+University+Press&rft.date=1991&rft.isbn=978-0-691-08739-9&rft.aulast=Hanski&rft.aufirst=Ilkka&rft.au=Yves%2C+Cambefort&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-104"><a href="#cite_ref-104">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://animals.sandiegozoo.org/animals/beetle">"Beetle | San Diego Zoo Animals & Plants"</a>. animals.sandiegozoo.org. Retrieved July 18, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=animals.sandiegozoo.org&rft.atitle=Beetle+%7C+San+Diego+Zoo+Animals+%26+Plants&rft_id=http%3A%2F%2Fanimals.sandiegozoo.org%2Fanimals%2Fbeetle&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-105"><a href="#cite_ref-105">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChabooFrieiro-CostaGómez-ZuritaWesterduijn2014" class="citation journal cs1">Chaboo, C.S.; Frieiro-Costa, F.A.; Gómez-Zurita, J.; Westerduijn, R. (2014). "Subsociality in leaf beetles (Coleoptera: Chrysomelidae: Cassidinae, Chrysomelinae)". Journal of Natural History. 48: 1–44. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00222933.2014.909060">10.1080/00222933.2014.909060</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:84683405">84683405</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Natural+History&rft.atitle=Subsociality+in+leaf+beetles+%28Coleoptera%3A+Chrysomelidae%3A+Cassidinae%2C+Chrysomelinae%29&rft.volume=48&rft.pages=1-44&rft.date=2014&rft_id=info%3Adoi%2F10.1080%2F00222933.2014.909060&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A84683405%23id-name%3DS2CID&rft.aulast=Chaboo&rft.aufirst=C.S.&rft.au=Frieiro-Costa%2C+F.A.&rft.au=G%C3%B3mez-Zurita%2C+J.&rft.au=Westerduijn%2C+R.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-106"><a href="#cite_ref-106">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFChaboo2002" class="citation journal cs1">Chaboo, CS (2002). "First report of immatures, genitalia and maternal care in Eugenysa columbiana (Boheman) (Coleoptera: Chrysomelidae: Cassidinae: Eugenysini)". The Coleopterists Bulletin. 56: 50–67. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1649%2F0010-065x%282002%29056%5B0050%3Afroiga%5D2.0.co%3B2">10.1649/0010-065x(2002)056[0050:froiga]2.0.co;2</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:85885981">85885981</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Coleopterists+Bulletin&rft.atitle=First+report+of+immatures%2C+genitalia+and+maternal+care+in+Eugenysa+columbiana+%28Boheman%29+%28Coleoptera%3A+Chrysomelidae%3A+Cassidinae%3A+Eugenysini%29&rft.volume=56&rft.pages=50-67&rft.date=2002&rft_id=info%3Adoi%2F10.1649%2F0010-065x%282002%29056%5B0050%3Afroiga%5D2.0.co%3B2&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A85885981%23id-name%3DS2CID&rft.aulast=Chaboo&rft.aufirst=CS&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-107"><a href="#cite_ref-107">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWindsor1987" class="citation journal cs1">Windsor, DM (1987). <a rel="nofollow" class="external text" href="https://doi.org/10.1155%2F1987%2F19861">"Natural History of a Subsocial Tortoise Beetle, Acromis sparsa Boheman (Chrysomelidae, Cassidinae) in Panama"</a>. Psyche: A Journal of Entomology. 94 (1–2): 127–150. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1155%2F1987%2F19861">10.1155/1987/19861</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Psyche%3A+A+Journal+of+Entomology&rft.atitle=Natural+History+of+a+Subsocial+Tortoise+Beetle%2C+Acromis+sparsa+Boheman+%28Chrysomelidae%2C+Cassidinae%29+in+Panama&rft.volume=94&rft.issue=1%E2%80%932&rft.pages=127-150&rft.date=1987&rft_id=info%3Adoi%2F10.1155%2F1987%2F19861&rft.aulast=Windsor&rft.aufirst=DM&rft_id=https%3A%2F%2Fdoi.org%2F10.1155%252F1987%252F19861&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-108"><a href="#cite_ref-108">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFReidBeatsonHasenpusch2009" class="citation journal cs1">Reid, CAM; Beatson, M; Hasenpusch, J (2009). "The morphology and biology of Pterodunga mirabile Daccordi, an unusual subsocial chrysomeline (Coleoptera: Chrysomelidae)". J. Nat. Hist. 43 (7–8): 373–398. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2009JNatH..43..373R">2009JNatH..43..373R</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F00222930802586016">10.1080/00222930802586016</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:84744056">84744056</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=J.+Nat.+Hist.&rft.atitle=The+morphology+and+biology+of+Pterodunga+mirabile+Daccordi%2C+an+unusual+subsocial+chrysomeline+%28Coleoptera%3A+Chrysomelidae%29&rft.volume=43&rft.issue=7%E2%80%938&rft.pages=373-398&rft.date=2009&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A84744056%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1080%2F00222930802586016&rft_id=info%3Abibcode%2F2009JNatH..43..373R&rft.aulast=Reid&rft.aufirst=CAM&rft.au=Beatson%2C+M&rft.au=Hasenpusch%2C+J&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-109"><a href="#cite_ref-109">^</a> Windsor DM, Choe JC. 1994. Origins of parental care in chrysomelid beetles. In: Jolivet PH, Cox ML, Petitipierre E, editors. Novel aspects of the biology of Chrysomelidae. Series Entomologica 50. Dordrecht: Kluwer Academic Publishers; p. 111–117. </li> <li id="cite_note-110"><a href="#cite_ref-110">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCrespi,_B._J.Yanega,_D.1995" class="citation journal cs1">Crespi, B. J.; Yanega, D. (1995). "The definition of eusociality". <a href="/wiki/Behavioral_Ecology" class="mw-redirect" title="Behavioral Ecology">Behavioral Ecology</a>. 6 (1): 109–115. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fbeheco%2F6.1.109">10.1093/beheco/6.1.109</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Behavioral+Ecology&rft.atitle=The+definition+of+eusociality&rft.volume=6&rft.issue=1&rft.pages=109-115&rft.date=1995&rft_id=info%3Adoi%2F10.1093%2Fbeheco%2F6.1.109&rft.au=Crespi%2C+B.+J.&rft.au=Yanega%2C+D.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Kent1992-111">^ <a href="#cite_ref-Kent1992_111-0">a</a> <a href="#cite_ref-Kent1992_111-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKent,_D._S.Simpson,_J._A.1992" class="citation journal cs1">Kent, D. S. & Simpson, J. A. (1992). "Eusociality in the beetle Austroplatypus incompertus (Coleoptera: Curculionidae)". <a href="/wiki/Naturwissenschaften" class="mw-redirect" title="Naturwissenschaften">Naturwissenschaften</a>. 79 (2): 86–87. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1992NW.....79...86K">1992NW.....79...86K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2FBF01131810">10.1007/BF01131810</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:35534268">35534268</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Naturwissenschaften&rft.atitle=Eusociality+in+the+beetle+Austroplatypus+incompertus+%28Coleoptera%3A+Curculionidae%29&rft.volume=79&rft.issue=2&rft.pages=86-87&rft.date=1992&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A35534268%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1007%2FBF01131810&rft_id=info%3Abibcode%2F1992NW.....79...86K&rft.au=Kent%2C+D.+S.&rft.au=Simpson%2C+J.+A.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-bee-112">^ <a href="#cite_ref-bee_112-0">a</a> <a href="#cite_ref-bee_112-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation magazine cs1"><a rel="nofollow" class="external text" href="https://www.newscientist.com/article/mg13418203.100-science-the-australian-beetle-that-behaves-like-a-bee.html">"Science: The Australian beetle that behaves like a bee"</a>. New Scientist. May 9, 1992. Retrieved October 31, 2010.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=New+Scientist&rft.atitle=Science%3A+The+Australian+beetle+that+behaves+like+a+bee&rft.date=1992-05-09&rft_id=https%3A%2F%2Fwww.newscientist.com%2Farticle%2Fmg13418203.100-science-the-australian-beetle-that-behaves-like-a-bee.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-113"><a href="#cite_ref-113">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSchusterSchuster1985" class="citation journal cs1">Schuster, Jack C.; Schuster, Laura B. (1985). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20160302014348/http://journals.fcla.edu/flaent/article/viewFile/58011/55690">"Social behavior in Passalid beetles (Coleoptera: Passalidae): Cooperative brood care"</a>. Florida Entomologist. 68 (2): 266–272. <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%2F3494359">10.2307/3494359</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/3494359">3494359</a>. Archived from <a rel="nofollow" class="external text" href="http://journals.fcla.edu/flaent/article/viewFile/58011/55690">the original</a> on March 2, 2016. Retrieved March 17, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Florida+Entomologist&rft.atitle=Social+behavior+in+Passalid+beetles+%28Coleoptera%3A+Passalidae%29%3A+Cooperative+brood+care&rft.volume=68&rft.issue=2&rft.pages=266-272&rft.date=1985&rft_id=info%3Adoi%2F10.2307%2F3494359&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F3494359%23id-name%3DJSTOR&rft.aulast=Schuster&rft.aufirst=Jack+C.&rft.au=Schuster%2C+Laura+B.&rft_id=http%3A%2F%2Fjournals.fcla.edu%2Fflaent%2Farticle%2FviewFile%2F58011%2F55690&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-114"><a href="#cite_ref-114">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLobanov,_A.L.2002" class="citation web cs1">Lobanov, A.L. (2002). <a rel="nofollow" class="external text" href="http://www.zin.ru/animalia/coleoptera/eng/biol3.htm">"feeding"</a>. Beetle Biology And Ecology. Beetles (Coleoptera) and Coleopterologist. Retrieved March 13, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Beetle+Biology+And+Ecology&rft.atitle=feeding&rft.date=2002&rft.au=Lobanov%2C+A.L.&rft_id=http%3A%2F%2Fwww.zin.ru%2Fanimalia%2Fcoleoptera%2Feng%2Fbiol3.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-115"><a href="#cite_ref-115">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMcCoy2016" class="citation book cs1">McCoy, Peter (2016). <a rel="nofollow" class="external text" href="https://archive.org/stream/radical-mycology-a-treatise-on-seeing-and-working-with-fungi/Radical_Mycology_A_treatise_on_seeing_and_working_with_fungi">Radical Mycology: A Treatise on Seeing & Working with Fungi</a>. Chthaeus Press. p. 187. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-9863996-0-2" title="Special:BookSources/978-0-9863996-0-2"><bdi>978-0-9863996-0-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Radical+Mycology%3A+A+Treatise+on+Seeing+%26+Working+with+Fungi&rft.pages=187&rft.pub=Chthaeus+Press&rft.date=2016&rft.isbn=978-0-9863996-0-2&rft.aulast=McCoy&rft.aufirst=Peter&rft_id=https%3A%2F%2Farchive.org%2Fstream%2Fradical-mycology-a-treatise-on-seeing-and-working-with-fungi%2FRadical_Mycology_A_treatise_on_seeing_and_working_with_fungi&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-116"><a href="#cite_ref-116">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFIslam,_M.Hossain,_A.Mostafa,_M._G.Hossain,_M._M.2016" class="citation journal cs1">Islam, M.; Hossain, A.; Mostafa, M. G.; Hossain, M. M. (2016). <a rel="nofollow" class="external text" href="http://banglajol.info/index.php/JUJBS/article/view/29739/19949">"Forensically important insects associated with the decomposition of mice carrion in Bangladesh"</a>. Jahangirnagar University Journal of Biological Science. 5 (1): 11–20. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.3329%2Fjujbs.v5i1.29739">10.3329/jujbs.v5i1.29739</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Jahangirnagar+University+Journal+of+Biological+Science&rft.atitle=Forensically+important+insects+associated+with+the+decomposition+of+mice+carrion+in+Bangladesh&rft.volume=5&rft.issue=1&rft.pages=11-20&rft.date=2016&rft_id=info%3Adoi%2F10.3329%2Fjujbs.v5i1.29739&rft.au=Islam%2C+M.&rft.au=Hossain%2C+A.&rft.au=Mostafa%2C+M.+G.&rft.au=Hossain%2C+M.+M.&rft_id=http%3A%2F%2Fbanglajol.info%2Findex.php%2FJUJBS%2Farticle%2Fview%2F29739%2F19949&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-117"><a href="#cite_ref-117">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGrebennikovLeschen2010" class="citation journal cs1">Grebennikov, Vasily V.; Leschen, Richard A. B. (2010). "External exoskeletal cavities in Coleoptera and their possible mycangial functions". Entomological Science. 13 (1): 81–98. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1479-8298.2009.00351.x">10.1111/j.1479-8298.2009.00351.x</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:84593757">84593757</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Entomological+Science&rft.atitle=External+exoskeletal+cavities+in+Coleoptera+and+their+possible+mycangial+functions&rft.volume=13&rft.issue=1&rft.pages=81-98&rft.date=2010&rft_id=info%3Adoi%2F10.1111%2Fj.1479-8298.2009.00351.x&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A84593757%23id-name%3DS2CID&rft.aulast=Grebennikov&rft.aufirst=Vasily+V.&rft.au=Leschen%2C+Richard+A.+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Evans27-118">^ <a href="#cite_ref-Evans27_118-0">a</a> <a href="#cite_ref-Evans27_118-1">b</a> <a href="#cite_ref-Evans27_118-2">c</a> <a href="#refEvans">Evans & Bellamy (2000)</a>, pp. 27–28 </li> <li id="cite_note-119"><a href="#cite_ref-119">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCott,_H._B.1940" class="citation book cs1"><a href="/wiki/Hugh_B._Cott" title="Hugh B. Cott">Cott, H. B.</a> (1940). <a href="/wiki/Adaptive_Coloration_in_Animals" title="Adaptive Coloration in Animals">Adaptive Coloration in Animals</a>. Methuen. p. <a rel="nofollow" class="external text" href="https://archive.org/details/adaptivecolorati00cott/page/414">414</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Adaptive+Coloration+in+Animals&rft.pages=414&rft.pub=Methuen&rft.date=1940&rft.au=Cott%2C+H.+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Evans126-120">^ <a href="#cite_ref-Evans126_120-0">a</a> <a href="#cite_ref-Evans126_120-1">b</a> <a href="#cite_ref-Evans126_120-2">c</a> <a href="#refEvans">Evans & Bellamy (2000)</a>, p. 126 </li> <li id="cite_note-Evans-121">^ <a href="#cite_ref-Evans_121-0">a</a> <a href="#cite_ref-Evans_121-1">b</a> <a href="#refEvans">Evans & Bellamy (2000)</a> </li> <li id="cite_note-insenc199-122"><a href="#cite_ref-insenc199_122-0">^</a> <a href="#refMcHugh">McHugh (2009)</a>, p. 199 </li> <li id="cite_note-123"><a href="#cite_ref-123">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPeck,_Stewart_B.2006" class="citation journal cs1">Peck, Stewart B. (2006). <a rel="nofollow" class="external text" href="http://digitalcommons.unl.edu/cgi/viewcontent.cgi?article=1107&context=insectamundi">"Distribution and biology of the ectoparasitic beaver beetle Platypsyllus castoris Ritsema in North America (Coleoptera: Leiodidae: Platypsyllinae)"</a>. Insecta Mundi. 20 (1–2): 85–94.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Insecta+Mundi&rft.atitle=Distribution+and+biology+of+the+ectoparasitic+beaver+beetle+Platypsyllus+castoris+Ritsema+in+North+America+%28Coleoptera%3A+Leiodidae%3A+Platypsyllinae%29&rft.volume=20&rft.issue=1%E2%80%932&rft.pages=85-94&rft.date=2006&rft.au=Peck%2C+Stewart+B.&rft_id=http%3A%2F%2Fdigitalcommons.unl.edu%2Fcgi%2Fviewcontent.cgi%3Farticle%3D1107%26context%3Dinsectamundi&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-124"><a href="#cite_ref-124">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNeumann,_P.Elzen,_P._J.2004" class="citation journal cs1">Neumann, P. & Elzen, P. J. (2004). <a rel="nofollow" class="external text" href="https://doi.org/10.1051%2Fapido%3A2004010">"The biology of the small hive beetle (Aethina tumida, Coleoptera: Nitidulidae): Gaps in our knowledge of an invasive species"</a>. Apidologie. 35 (3): 229–247. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1051%2Fapido%3A2004010">10.1051/apido:2004010</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Apidologie&rft.atitle=The+biology+of+the+small+hive+beetle+%28Aethina+tumida%2C+Coleoptera%3A+Nitidulidae%29%3A+Gaps+in+our+knowledge+of+an+invasive+species&rft.volume=35&rft.issue=3&rft.pages=229-247&rft.date=2004&rft_id=info%3Adoi%2F10.1051%2Fapido%3A2004010&rft.au=Neumann%2C+P.&rft.au=Elzen%2C+P.+J.&rft_id=https%3A%2F%2Fdoi.org%2F10.1051%252Fapido%253A2004010&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-125"><a href="#cite_ref-125">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMeyer,_John_R.2005" class="citation web cs1">Meyer, John R. (March 8, 2005). <a rel="nofollow" class="external text" href="https://wayback.archive-it.org/all/20000524004320/http://www.cals.ncsu.edu/course/ent425/compendium/coleop~1.html">"Coleoptera"</a>. <a href="/wiki/North_Carolina_State_University" title="North Carolina State University">North Carolina State University</a>. Archived from <a rel="nofollow" class="external text" href="http://www.cals.ncsu.edu/course/ent425/compendium/coleop~1.html">the original</a> on May 24, 2000. Retrieved March 13, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Coleoptera&rft.pub=North+Carolina+State+University&rft.date=2005-03-08&rft.au=Meyer%2C+John+R.&rft_id=http%3A%2F%2Fwww.cals.ncsu.edu%2Fcourse%2Fent425%2Fcompendium%2Fcoleop~1.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-126"><a href="#cite_ref-126">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWeberSaskaChaboo2008" class="citation book cs1">Weber, Donald C.; Saska, Pavel; Chaboo, Caroline S. (2008). "Carabid Beetles (Coleoptera: Carabidae) as Parasitoids". Encyclopedia of Entomology. Springer Netherlands. pp. 719–721. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-1-4020-6359-6_492">10.1007/978-1-4020-6359-6_492</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4020-6242-1" title="Special:BookSources/978-1-4020-6242-1"><bdi>978-1-4020-6242-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Carabid+Beetles+%28Coleoptera%3A+Carabidae%29+as+Parasitoids&rft.btitle=Encyclopedia+of+Entomology&rft.pages=719-721&rft.pub=Springer+Netherlands&rft.date=2008&rft_id=info%3Adoi%2F10.1007%2F978-1-4020-6359-6_492&rft.isbn=978-1-4020-6242-1&rft.aulast=Weber&rft.aufirst=Donald+C.&rft.au=Saska%2C+Pavel&rft.au=Chaboo%2C+Caroline+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-127"><a href="#cite_ref-127">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLivingston2019" class="citation journal cs1">Livingston, Stephanie (November 2019). <a rel="nofollow" class="external text" href="https://www.science.org/content/article/amber-encased-beetle-may-have-been-one-first-insects-pollinate-flowers">"This amber-encased beetle may have been one of the first insects to pollinate flowers"</a>. Science. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.aba1758">10.1126/science.aba1758</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:213876270">213876270</a>. Retrieved January 10, 2021.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=This+amber-encased+beetle+may+have+been+one+of+the+first+insects+to+pollinate+flowers&rft.date=2019-11&rft_id=info%3Adoi%2F10.1126%2Fscience.aba1758&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A213876270%23id-name%3DS2CID&rft.aulast=Livingston&rft.aufirst=Stephanie&rft_id=https%3A%2F%2Fwww.science.org%2Fcontent%2Farticle%2Famber-encased-beetle-may-have-been-one-first-insects-pollinate-flowers&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Gullan-128"><a href="#cite_ref-Gullan_128-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGullanCranston,_P._S.2014" class="citation book cs1">Gullan, P. J.; Cranston, P. S. (2014). The Insects: An Outline of Entomology (5 ed.). Wiley, John & Sons. p. 314. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4443-3036-6" title="Special:BookSources/978-1-4443-3036-6"><bdi>978-1-4443-3036-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Insects%3A+An+Outline+of+Entomology&rft.pages=314&rft.edition=5&rft.pub=Wiley%2C+John+%26+Sons&rft.date=2014&rft.isbn=978-1-4443-3036-6&rft.aulast=Gullan&rft.aufirst=P.+J.&rft.au=Cranston%2C+P.+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-129"><a href="#cite_ref-129">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFJones,_G._D.Jones,_S._D.2001" class="citation journal cs1">Jones, G. D. & Jones, S. D. (2001). <a rel="nofollow" class="external text" href="https://doi.org/10.1590%2FS1519-566X2001000300001">"The uses of pollen and its implication for entomology"</a>. Neotropical Entomology. 30 (3): 314–349. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1590%2FS1519-566X2001000300001">10.1590/S1519-566X2001000300001</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Neotropical+Entomology&rft.atitle=The+uses+of+pollen+and+its+implication+for+entomology&rft.volume=30&rft.issue=3&rft.pages=314-349&rft.date=2001&rft_id=info%3Adoi%2F10.1590%2FS1519-566X2001000300001&rft.au=Jones%2C+G.+D.&rft.au=Jones%2C+S.+D.&rft_id=https%3A%2F%2Fdoi.org%2F10.1590%252FS1519-566X2001000300001&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Ollerton_a-130"><a href="#cite_ref-Ollerton_a_130-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFOllerton,_J.Johnson,_S._D.Cranmer,_L.Kellie,_S.2003" class="citation journal cs1">Ollerton, J.; Johnson, S. D.; Cranmer, L. & Kellie, S. (2003). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4243623">"The pollination ecology of an assemblage of grassland asclepiads in South Africa"</a>. <a href="/wiki/Annals_of_Botany" title="Annals of Botany">Annals of Botany</a>. 92 (6): 807–834. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Faob%2Fmcg206">10.1093/aob/mcg206</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4243623">4243623</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/14612378">14612378</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annals+of+Botany&rft.atitle=The+pollination+ecology+of+an+assemblage+of+grassland+asclepiads+in+South+Africa&rft.volume=92&rft.issue=6&rft.pages=807-834&rft.date=2003&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4243623%23id-name%3DPMC&rft_id=info%3Apmid%2F14612378&rft_id=info%3Adoi%2F10.1093%2Faob%2Fmcg206&rft.au=Ollerton%2C+J.&rft.au=Johnson%2C+S.+D.&rft.au=Cranmer%2C+L.&rft.au=Kellie%2C+S.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4243623&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Malloch-131">^ <a href="#cite_ref-Malloch_131-0">a</a> <a href="#cite_ref-Malloch_131-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMallochBlackwell1993" class="citation book cs1">Malloch, D.; Blackwell, M. (1993). "Dispersal biology of ophiostomatoid fungi". In Wingfield, M. J.; K. A. Seifert; J. F. Webber (eds.). Ceratocystis and Ophiostoma: Taxonomy, Ecology and Pathology. St. Paul: APS. pp. 195–206. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-89054-156-2" title="Special:BookSources/978-0-89054-156-2"><bdi>978-0-89054-156-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Dispersal+biology+of+ophiostomatoid+fungi&rft.btitle=Ceratocystis+and+Ophiostoma%3A+Taxonomy%2C+Ecology+and+Pathology&rft.place=St.+Paul&rft.pages=195-206&rft.pub=APS&rft.date=1993&rft.isbn=978-0-89054-156-2&rft.aulast=Malloch&rft.aufirst=D.&rft.au=Blackwell%2C+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Scott2008-132"><a href="#cite_ref-Scott2008_132-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFScottOh,_D._C.Yuceer,_M._C.Klepzig,_K._D.2008" class="citation journal cs1">Scott, J. J.; Oh, D. C.; Yuceer, M. C.; Klepzig, K. D.; Clardy, J.; Currie, C. R. (2008). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2761720">"Bacterial protection of beetle-fungus mutualism"</a>. <a href="/wiki/Science_(journal)" title="Science (journal)">Science</a>. 322 (5898): 63. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2008Sci...322...63S">2008Sci...322...63S</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscience.1160423">10.1126/science.1160423</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2761720">2761720</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/18832638">18832638</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science&rft.atitle=Bacterial+protection+of+beetle-fungus+mutualism&rft.volume=322&rft.issue=5898&rft.pages=63&rft.date=2008&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2761720%23id-name%3DPMC&rft_id=info%3Apmid%2F18832638&rft_id=info%3Adoi%2F10.1126%2Fscience.1160423&rft_id=info%3Abibcode%2F2008Sci...322...63S&rft.aulast=Scott&rft.aufirst=J.+J.&rft.au=Oh%2C+D.+C.&rft.au=Yuceer%2C+M.+C.&rft.au=Klepzig%2C+K.+D.&rft.au=Clardy%2C+J.&rft.au=Currie%2C+C.+R.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2761720&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-133"><a href="#cite_ref-133">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFrancke-Grossmann,_H.1967" class="citation book cs1">Francke-Grossmann, H. (1967). "Ectosymbiosis in wood inhabiting insects". In M. Henry (ed.). Symbiosis. Vol. 2. New York: <a href="/wiki/Academic_Press" title="Academic Press">Academic Press</a>. pp. 141–205.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Ectosymbiosis+in+wood+inhabiting+insects&rft.btitle=Symbiosis&rft.place=New+York&rft.pages=141-205&rft.pub=Academic+Press&rft.date=1967&rft.au=Francke-Grossmann%2C+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-134"><a href="#cite_ref-134">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHodek2012" class="citation journal cs1">Hodek, Ivo (2012). <a rel="nofollow" class="external text" href="https://doi.org/10.1155%2F2012%2F249081">"Review Article: Adult diapause in Coleoptera"</a>. Psyche: A Journal of Entomology. 2012: 1–10. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1155%2F2012%2F249081">10.1155/2012/249081</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Psyche%3A+A+Journal+of+Entomology&rft.atitle=Review+Article%3A+Adult+diapause+in+Coleoptera&rft.volume=2012&rft.pages=1-10&rft.date=2012&rft_id=info%3Adoi%2F10.1155%2F2012%2F249081&rft.aulast=Hodek&rft.aufirst=Ivo&rft_id=https%3A%2F%2Fdoi.org%2F10.1155%252F2012%252F249081&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-135"><a href="#cite_ref-135">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKaufmann,_T.1971" class="citation journal cs1">Kaufmann, T. (1971). "Hibernation in the Arctic beetle, Pterostichus brevicornis, in Alaska". Journal of the Kansas Entomological Society. 44 (1): 81–92.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+Kansas+Entomological+Society&rft.atitle=Hibernation+in+the+Arctic+beetle%2C+Pterostichus+brevicornis%2C+in+Alaska&rft.volume=44&rft.issue=1&rft.pages=81-92&rft.date=1971&rft.au=Kaufmann%2C+T.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Outline-136"><a href="#cite_ref-Outline_136-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGullan,_P._J.Cranston,_P._S.1994" class="citation book cs1">Gullan, P. J.; Cranston, P. S. (1994). The Insects: An Outline of Entomology. Chapman and Hall. pp. 103–104. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-412-49360-7" title="Special:BookSources/978-0-412-49360-7"><bdi>978-0-412-49360-7</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Insects%3A+An+Outline+of+Entomology&rft.pages=103-104&rft.pub=Chapman+and+Hall&rft.date=1994&rft.isbn=978-0-412-49360-7&rft.au=Gullan%2C+P.+J.&rft.au=Cranston%2C+P.+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-137"><a href="#cite_ref-137">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLombadero,_Maria_J.Ayres,_Matthew_P.Ayres,_Bruce_D.Reeve,_John_D.2000" class="citation journal cs1">Lombadero, Maria J.; Ayres, Matthew P.; Ayres, Bruce D.; Reeve, John D. (2000). <a rel="nofollow" class="external text" href="http://www.dartmouth.edu/~mpayres/pubs/Fina.Cold.pdf">"Cold tolerance of four species of bark beetle (Coleoptera: Scolytidae) in North America"</a> (PDF). Environmental Ecology. 29 (3): 421–432. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20070417144045/http://www.dartmouth.edu/~mpayres/pubs/Fina.Cold.pdf">Archived</a> (PDF) from the original on April 17, 2007.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Environmental+Ecology&rft.atitle=Cold+tolerance+of+four+species+of+bark+beetle+%28Coleoptera%3A+Scolytidae%29+in+North+America&rft.volume=29&rft.issue=3&rft.pages=421-432&rft.date=2000&rft.au=Lombadero%2C+Maria+J.&rft.au=Ayres%2C+Matthew+P.&rft.au=Ayres%2C+Bruce+D.&rft.au=Reeve%2C+John+D.&rft_id=http%3A%2F%2Fwww.dartmouth.edu%2F~mpayres%2Fpubs%2FFina.Cold.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-138"><a href="#cite_ref-138">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSformo,_T.Walters,_K.Jeannet,_K.Wowk,_B.2010" class="citation journal cs1">Sformo, T.; Walters, K.; Jeannet, K.; Wowk, B.; Fahy, G. M.; Barnes, B. M.; Duman, J. G. (2010). <a rel="nofollow" class="external text" href="https://doi.org/10.1242%2Fjeb.035758">"Deep supercooling, vitrification and limited survival to −100°C in the Alaskan beetle Cucujus clavipes puniceus (Coleoptera: Cucujidae) larvae"</a>. <a href="/wiki/Journal_of_Experimental_Biology" class="mw-redirect" title="Journal of Experimental Biology">Journal of Experimental Biology</a>. 213 (3): 502–509. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1242%2Fjeb.035758">10.1242/jeb.035758</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/20086136">20086136</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Experimental+Biology&rft.atitle=Deep+supercooling%2C+vitrification+and+limited+survival+to+%E2%88%92100%C2%B0C+in+the+Alaskan+beetle+Cucujus+clavipes+puniceus+%28Coleoptera%3A+Cucujidae%29+larvae&rft.volume=213&rft.issue=3&rft.pages=502-509&rft.date=2010&rft_id=info%3Adoi%2F10.1242%2Fjeb.035758&rft_id=info%3Apmid%2F20086136&rft.au=Sformo%2C+T.&rft.au=Walters%2C+K.&rft.au=Jeannet%2C+K.&rft.au=Wowk%2C+B.&rft.au=Fahy%2C+G.+M.&rft.au=Barnes%2C+B.+M.&rft.au=Duman%2C+J.+G.&rft_id=https%3A%2F%2Fdoi.org%2F10.1242%252Fjeb.035758&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-139"><a href="#cite_ref-139">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrooks2013" class="citation news cs1">Brooks, Christopher (March 26, 2013). <a rel="nofollow" class="external text" href="https://www.bbc.co.uk/nature/21923937">"The life of extremophiles: Surviving in hostile habitats"</a>. BBC Nature. Retrieved March 16, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.atitle=The+life+of+extremophiles%3A+Surviving+in+hostile+habitats&rft.date=2013-03-26&rft.aulast=Brooks&rft.aufirst=Christopher&rft_id=https%3A%2F%2Fwww.bbc.co.uk%2Fnature%2F21923937&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-140"><a href="#cite_ref-140">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGraham,_L._ALiou,_Y._C.Walker,_V._K.Davies,_P._L.1997" class="citation journal cs1">Graham, L. A; Liou, Y. C.; Walker, V. K.; Davies, P. L. (August 1997). <a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F41908">"Hyperactive antifreeze protein from beetles"</a>. <a href="/wiki/Nature_(journal)" title="Nature (journal)">Nature</a>. 388 (6644): 727–728. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1997Natur.388..727G">1997Natur.388..727G</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F41908">10.1038/41908</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/9285581">9285581</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:205029622">205029622</a>. <q>The yellow mealworm beetle, Tenebrio molitor, contains a family of small Cys-rich and Thr-rich thermal hysteresis proteins that depress the hemolymph freezing point below the melting point by as much as 5.58°C(ΔT=thermal hysteresis). Thermal hysteresis protein expression was evaluated throughout development and after exposure to altered environmental conditions.</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Hyperactive+antifreeze+protein+from+beetles&rft.volume=388&rft.issue=6644&rft.pages=727-728&rft.date=1997-08&rft_id=info%3Adoi%2F10.1038%2F41908&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A205029622%23id-name%3DS2CID&rft_id=info%3Apmid%2F9285581&rft_id=info%3Abibcode%2F1997Natur.388..727G&rft.au=Graham%2C+L.+A&rft.au=Liou%2C+Y.+C.&rft.au=Walker%2C+V.+K.&rft.au=Davies%2C+P.+L.&rft_id=https%3A%2F%2Fdoi.org%2F10.1038%252F41908&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-141"><a href="#cite_ref-141">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWalters,_K._R._JrSerianni,_A._S.Sformo,_T.Barnes,_B._M.2009" class="citation journal cs1">Walters, K. R. Jr; Serianni, A. S.; Sformo, T.; Barnes, B. M.; Duman, J. G. (2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2787118">"A nonprotein thermal hysteresis-producing xylomannan antifreeze in the freeze-tolerant Alaskan beetle Upis ceramboides"</a>. <a href="/wiki/PNAS" class="mw-redirect" title="PNAS">PNAS</a>. 106 (48): 20210–20215. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2009PNAS..10620210W">2009PNAS..10620210W</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1073%2Fpnas.0909872106">10.1073/pnas.0909872106</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2787118">2787118</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19934038">19934038</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=PNAS&rft.atitle=A+nonprotein+thermal+hysteresis-producing+xylomannan+antifreeze+in+the+freeze-tolerant+Alaskan+beetle+Upis+ceramboides&rft.volume=106&rft.issue=48&rft.pages=20210-20215&rft.date=2009&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2787118%23id-name%3DPMC&rft_id=info%3Apmid%2F19934038&rft_id=info%3Adoi%2F10.1073%2Fpnas.0909872106&rft_id=info%3Abibcode%2F2009PNAS..10620210W&rft.au=Walters%2C+K.+R.+Jr&rft.au=Serianni%2C+A.+S.&rft.au=Sformo%2C+T.&rft.au=Barnes%2C+B.+M.&rft.au=Duman%2C+J.+G.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2787118&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-142"><a href="#cite_ref-142">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWalters,_K._R._Jr.Pan,_Q.Serianni,_A._S.Duman,_J._G.2009" class="citation journal cs1">Walters, K. R. Jr.; Pan, Q.; Serianni, A. S.; Duman, J. G. (2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2719318">"Cryoprotectant biosynthesis and the selective accumulation of threitol in the freeze-tolerant Alaskan beetle, Upis ceramboides"</a>. <a href="/wiki/Journal_of_Biological_Chemistry" title="Journal of Biological Chemistry">Journal of Biological Chemistry</a>. 284 (25): 16822–16831. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1074%2Fjbc.M109.013870">10.1074/jbc.M109.013870</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2719318">2719318</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19403530">19403530</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Biological+Chemistry&rft.atitle=Cryoprotectant+biosynthesis+and+the+selective+accumulation+of+threitol+in+the+freeze-tolerant+Alaskan+beetle%2C+Upis+ceramboides&rft.volume=284&rft.issue=25&rft.pages=16822-16831&rft.date=2009&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2719318%23id-name%3DPMC&rft_id=info%3Apmid%2F19403530&rft_id=info%3Adoi%2F10.1074%2Fjbc.M109.013870&rft.au=Walters%2C+K.+R.+Jr.&rft.au=Pan%2C+Q.&rft.au=Serianni%2C+A.+S.&rft.au=Duman%2C+J.+G.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC2719318&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-143"><a href="#cite_ref-143">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEdney,_E._B.1971" class="citation journal cs1">Edney, E. B. (1971). <a rel="nofollow" class="external text" href="http://jeb.biologists.org/content/jexbio/55/1/253.full.pdf">"The body temperature of tenebrionid beetles in the Namib desert of southern Africa"</a> (PDF). <a href="/wiki/Journal_of_Experimental_Biology" class="mw-redirect" title="Journal of Experimental Biology">Journal of Experimental Biology</a>. 55: 253–272. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1242%2Fjeb.55.1.253">10.1242/jeb.55.1.253</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170211080953/http://jeb.biologists.org/content/jexbio/55/1/253.full.pdf">Archived</a> (PDF) from the original on February 11, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Experimental+Biology&rft.atitle=The+body+temperature+of+tenebrionid+beetles+in+the+Namib+desert+of+southern+Africa&rft.volume=55&rft.pages=253-272&rft.date=1971&rft_id=info%3Adoi%2F10.1242%2Fjeb.55.1.253&rft.au=Edney%2C+E.+B.&rft_id=http%3A%2F%2Fjeb.biologists.org%2Fcontent%2Fjexbio%2F55%2F1%2F253.full.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-144"><a href="#cite_ref-144">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKnisleySchultzHasewinkel1990" class="citation journal cs1">Knisley, C. B.; Schultz, T. D.; Hasewinkel, T. H. (1990). "Seasonal activity and thermoregulatory behavior of Cicindela patruela (Coleoptera: Cicindelidae)". Annals of the Entomological Society of America. 83 (5): 911–915. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Faesa%2F83.5.911">10.1093/aesa/83.5.911</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Annals+of+the+Entomological+Society+of+America&rft.atitle=Seasonal+activity+and+thermoregulatory+behavior+of+Cicindela+patruela+%28Coleoptera%3A+Cicindelidae%29&rft.volume=83&rft.issue=5&rft.pages=911-915&rft.date=1990&rft_id=info%3Adoi%2F10.1093%2Faesa%2F83.5.911&rft.aulast=Knisley&rft.aufirst=C.+B.&rft.au=Schultz%2C+T.+D.&rft.au=Hasewinkel%2C+T.+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-145"><a href="#cite_ref-145">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFParkerLawrence2001" class="citation journal cs1">Parker, Andrew R.; Lawrence, Chris R. (November 1, 2001). "Water capture by a desert beetle". Nature. 414 (6859): 33–34. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2001Natur.414...33P">2001Natur.414...33P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F35102108">10.1038/35102108</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/11689930">11689930</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:34785113">34785113</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Water+capture+by+a+desert+beetle&rft.volume=414&rft.issue=6859&rft.pages=33-34&rft.date=2001-11-01&rft_id=info%3Adoi%2F10.1038%2F35102108&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A34785113%23id-name%3DS2CID&rft_id=info%3Apmid%2F11689930&rft_id=info%3Abibcode%2F2001Natur.414...33P&rft.aulast=Parker&rft.aufirst=Andrew+R.&rft.au=Lawrence%2C+Chris+R.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-146"><a href="#cite_ref-146">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHoback,_W._WyattStanley,_David_W.Higley,_Leon_G.Barnhart,_M._Christopher1998" class="citation journal cs1">Hoback, W. Wyatt; Stanley, David W.; Higley, Leon G.; Barnhart, M. Christopher (1998). "Survival of immersion and anoxia by larval tiger beetles, Cicindela togata". The American Midland Naturalist. 140 (1): 27–33. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1674%2F0003-0031%281998%29140%5B0027%3ASOIAAB%5D2.0.CO%3B2">10.1674/0003-0031(1998)140[0027:SOIAAB]2.0.CO;2</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:86163282">86163282</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+American+Midland+Naturalist&rft.atitle=Survival+of+immersion+and+anoxia+by+larval+tiger+beetles%2C+Cicindela+togata&rft.volume=140&rft.issue=1&rft.pages=27-33&rft.date=1998&rft_id=info%3Adoi%2F10.1674%2F0003-0031%281998%29140%5B0027%3ASOIAAB%5D2.0.CO%3B2&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A86163282%23id-name%3DS2CID&rft.au=Hoback%2C+W.+Wyatt&rft.au=Stanley%2C+David+W.&rft.au=Higley%2C+Leon+G.&rft.au=Barnhart%2C+M.+Christopher&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-147"><a href="#cite_ref-147">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFConradi-Larsen,_Else-MargreteSømme,_Lauritz1973" class="citation journal cs1">Conradi-Larsen, Else-Margrete; Sømme, Lauritz (1973). "Anaerobiosis in the overwintering beetle Pelophila borealis". Nature. 245 (5425): 388–390. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1973Natur.245..388C">1973Natur.245..388C</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1038%2F245388a0">10.1038/245388a0</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:4288059">4288059</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Nature&rft.atitle=Anaerobiosis+in+the+overwintering+beetle+Pelophila+borealis&rft.volume=245&rft.issue=5425&rft.pages=388-390&rft.date=1973&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A4288059%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1038%2F245388a0&rft_id=info%3Abibcode%2F1973Natur.245..388C&rft.au=Conradi-Larsen%2C+Else-Margrete&rft.au=S%C3%B8mme%2C+Lauritz&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-148"><a href="#cite_ref-148">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAllison,_R.2015" class="citation web cs1">Allison, R. (March 17, 2015). <a rel="nofollow" class="external text" href="http://www.fwi.co.uk/arable/half-of-the-uk-sees-start-of-pollen-beetle-migration.htm">"Half of the UK sees start of pollen beetle migration"</a>. Retrieved March 16, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Half+of+the+UK+sees+start+of+pollen+beetle+migration&rft.date=2015-03-17&rft.au=Allison%2C+R.&rft_id=http%3A%2F%2Fwww.fwi.co.uk%2Farable%2Fhalf-of-the-uk-sees-start-of-pollen-beetle-migration.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-149"><a href="#cite_ref-149">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSouthwood1962" class="citation journal cs1">Southwood, T. R. E. (1962). "Migration of Terrestrial Arthropods in Relation to Habitat". Biological Reviews. 37 (2): 171–211. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1469-185X.1962.tb01609.x">10.1111/j.1469-185X.1962.tb01609.x</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:84711127">84711127</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biological+Reviews&rft.atitle=Migration+of+Terrestrial+Arthropods+in+Relation+to+Habitat&rft.volume=37&rft.issue=2&rft.pages=171-211&rft.date=1962&rft_id=info%3Adoi%2F10.1111%2Fj.1469-185X.1962.tb01609.x&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A84711127%23id-name%3DS2CID&rft.aulast=Southwood&rft.aufirst=T.+R.+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-150"><a href="#cite_ref-150">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDingle,_H.2014" class="citation book cs1">Dingle, H. (2014). Migration: The Biology of Life on the Move. Oxford University Press.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Migration%3A+The+Biology+of+Life+on+the+Move&rft.pub=Oxford+University+Press&rft.date=2014&rft.au=Dingle%2C+H.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-egyptian-151">^ <a href="#cite_ref-egyptian_151-0">a</a> <a href="#cite_ref-egyptian_151-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFZabludoff,_Marc2008" class="citation book cs1">Zabludoff, Marc (2008). Beetles. Malaysia: Michelle Bison. pp. 14–17. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7614-2532-8" title="Special:BookSources/978-0-7614-2532-8"><bdi>978-0-7614-2532-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Beetles&rft.place=Malaysia&rft.pages=14-17&rft.pub=Michelle+Bison&rft.date=2008&rft.isbn=978-0-7614-2532-8&rft.au=Zabludoff%2C+Marc&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-152"><a href="#cite_ref-152">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCambefort,_Yves2011" class="citation web cs1">Cambefort, Yves (2011). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20161018110844/http://www.insects.org/ced1/beetles_rel_sym.html">"Beetles as religious symbols"</a>. Insects.org. Archived from <a rel="nofollow" class="external text" href="http://www.insects.org/ced1/beetles_rel_sym.html">the original</a> on October 18, 2016. Retrieved February 10, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Beetles+as+religious+symbols&rft.pub=Insects.org&rft.date=2011&rft.au=Cambefort%2C+Yves&rft_id=http%3A%2F%2Fwww.insects.org%2Fced1%2Fbeetles_rel_sym.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-153"><a href="#cite_ref-153">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFDollinger,_André2002" class="citation web cs1">Dollinger, André (January 2002). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20150401025144/http://www.reshafim.org.il/ad/egypt/bestiary/insects.htm">"Ancient Egyptian bestiary: Insects"</a>. Archived from <a rel="nofollow" class="external text" href="http://www.reshafim.org.il/ad/egypt/bestiary/insects.htm">the original</a> on April 1, 2015. Retrieved July 19, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Ancient+Egyptian+bestiary%3A+Insects&rft.date=2002-01&rft.au=Dollinger%2C+Andr%C3%A9&rft_id=http%3A%2F%2Fwww.reshafim.org.il%2Fad%2Fegypt%2Fbestiary%2Finsects.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-154"><a href="#cite_ref-154">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMorales-Correa,_Ben2006" class="citation web cs1">Morales-Correa, Ben (2006). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110820175032/http://www.all-about-egypt.com/egyptian-symbols.html">"Egyptian Symbols"</a>. All-About-Egypt. Archived from <a rel="nofollow" class="external text" href="http://www.all-about-egypt.com/egyptian-symbols.html">the original</a> on August 20, 2011. Retrieved July 19, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Egyptian+Symbols&rft.pub=All-About-Egypt&rft.date=2006&rft.au=Morales-Correa%2C+Ben&rft_id=http%3A%2F%2Fwww.all-about-egypt.com%2Fegyptian-symbols.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-155"><a href="#cite_ref-155">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFUssishkin2004" class="citation book cs1">Ussishkin, David (2004). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20180703050411/http://archaeology.tau.ac.il/?page_id=2059">The New Archaeological Excavations at Lachish (1973–1994)</a>. Tel Aviv: Institute of Archaeology of <a href="/wiki/Tel_Aviv_University" title="Tel Aviv University">Tel Aviv University</a>. Archived from <a rel="nofollow" class="external text" href="http://archaeology.tau.ac.il/?page_id=2059">the original</a> on July 3, 2018. Retrieved December 28, 2015.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+New+Archaeological+Excavations+at+Lachish+%281973%E2%80%931994%29&rft.place=Tel+Aviv&rft.pub=Institute+of+Archaeology+of+Tel+Aviv+University&rft.date=2004&rft.aulast=Ussishkin&rft.aufirst=David&rft_id=http%3A%2F%2Farchaeology.tau.ac.il%2F%3Fpage_id%3D2059&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-156"><a href="#cite_ref-156">^</a> <a rel="nofollow" class="external text" href="https://penelope.uchicago.edu/Thayer/E/Roman/Texts/Plutarch/Moralia/Isis_and_Osiris*/E.html#ref373">"Isis and Osiris"</a>, Moralia, in volume V of the <a href="/wiki/Loeb_Classical_Library" title="Loeb Classical Library">Loeb Classical Library</a>, 1936. "The race of beetles has no female, but all the males eject their sperm into a round pellet of material which they roll up by pushing it from the opposite side, just as the sun seems to turn the heavens in the direction opposite to its own course, which is from west to east. </li> <li id="cite_note-157"><a href="#cite_ref-157">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBetz,_H._D.1992" class="citation book cs1"><a href="/wiki/Hans_Dieter_Betz" title="Hans Dieter Betz">Betz, H. D.</a>, ed. (1992). The Greek Magical Papyri in Translation (Including the Demotic Spells) (2nd ed.). <a href="/wiki/University_of_Chicago_Press" title="University of Chicago Press">University of Chicago Press</a>. pp. IV.52–85, VII.520, XII.101, XIII.1065–1070, XXXVI.170.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Greek+Magical+Papyri+in+Translation+%28Including+the+Demotic+Spells%29&rft.pages=IV.52-85%2C+VII.520%2C+XII.101%2C+XIII.1065-1070%2C+XXXVI.170&rft.edition=2nd&rft.pub=University+of+Chicago+Press&rft.date=1992&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-PlinyXI-158"><a href="#cite_ref-PlinyXI_158-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPliny_the_Elder" class="citation web cs1"><a href="/wiki/Pliny_the_Elder" title="Pliny the Elder">Pliny the Elder</a>. <a rel="nofollow" class="external text" href="https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.02.0137%3Abook%3D11%3Achapter%3D1">"Natural History Book 11"</a>. Perseus.tufts.edu. Retrieved January 25, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Natural+History+Book+11&rft.pub=Perseus.tufts.edu&rft.au=Pliny+the+Elder&rft_id=https%3A%2F%2Fwww.perseus.tufts.edu%2Fhopper%2Ftext%3Fdoc%3DPerseus%253Atext%253A1999.02.0137%253Abook%253D11%253Achapter%253D1&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-PlinyXI_Book34-159"><a href="#cite_ref-PlinyXI_Book34_159-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPliny_the_Elder" class="citation web cs1"><a href="/wiki/Pliny_the_Elder" title="Pliny the Elder">Pliny the Elder</a>. <a rel="nofollow" class="external text" href="https://www.perseus.tufts.edu/hopper/text?doc=Perseus%3Atext%3A1999.02.0137%3Abook%3D11%3Achapter%3D34">"Natural History Book 11, Chapter 34"</a>. Perseus.tufts.edu. Retrieved January 25, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Natural+History+Book+11%2C+Chapter+34&rft.pub=Perseus.tufts.edu&rft.au=Pliny+the+Elder&rft_id=https%3A%2F%2Fwww.perseus.tufts.edu%2Fhopper%2Ftext%3Fdoc%3DPerseus%253Atext%253A1999.02.0137%253Abook%253D11%253Achapter%253D34&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Beavis-160">^ <a href="#cite_ref-Beavis_160-0">a</a> <a href="#cite_ref-Beavis_160-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeavis,_I._C.1988" class="citation book cs1">Beavis, I. C. (1988). Insects and other Invertebrates in Classical Antiquity. <a href="/wiki/University_of_Exeter" title="University of Exeter">University of Exeter</a>. pp. 153–154.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Insects+and+other+Invertebrates+in+Classical+Antiquity&rft.pages=153-154&rft.pub=University+of+Exeter&rft.date=1988&rft.au=Beavis%2C+I.+C.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-162"><a href="#cite_ref-162">^</a> <a href="/wiki/Antoninus_Liberalis" title="Antoninus Liberalis">Antoninus Liberalis</a>. Metamorphoses. Trans. Celoria, F. 1992. The Metamorphoses of Antoninus Liberalis: A translation with a commentary. London and New York, Routledge. </li> <li id="cite_note-163"><a href="#cite_ref-163">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSprecher-Uebersax2008" class="citation journal cs1">Sprecher-Uebersax, E. (2008). "The Stag Beetle Lucanus Cervus (Coleoptera, Lucanidae) in Art and Mythology". Revue d'Écologie. 63: 145–151.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Revue+d%27%C3%89cologie&rft.atitle=The+Stag+Beetle+Lucanus+Cervus+%28Coleoptera%2C+Lucanidae%29+in+Art+and+Mythology&rft.volume=63&rft.pages=145-151&rft.date=2008&rft.aulast=Sprecher-Uebersax&rft.aufirst=E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-msstate-164">^ <a href="#cite_ref-msstate_164-0">a</a> <a href="#cite_ref-msstate_164-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMississippi_State_University" class="citation web cs1">Mississippi State University. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20080512023346/http://www.bollweevil.ext.msstate.edu/webpage_history.htm">"History of the Boll Weevil in the United States"</a>. Economic impacts of the boll weevil. Archived from <a rel="nofollow" class="external text" href="http://www.bollweevil.ext.msstate.edu/webpage_history.htm">the original</a> on May 12, 2008.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Economic+impacts+of+the+boll+weevil&rft.atitle=History+of+the+Boll+Weevil+in+the+United+States&rft.au=Mississippi+State+University&rft_id=http%3A%2F%2Fwww.bollweevil.ext.msstate.edu%2Fwebpage_history.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-165"><a href="#cite_ref-165">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAllenHumble2002" class="citation journal cs1">Allen, E. A.; Humble, L. M. (2002). "Nonindigenous species introductions: A threat to Canada's forests and forest economy". Canadian Journal of Plant Pathology. 24 (2): 103–110. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2002CaJPP..24..103A">2002CaJPP..24..103A</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1080%2F07060660309506983">10.1080/07060660309506983</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:85073955">85073955</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Canadian+Journal+of+Plant+Pathology&rft.atitle=Nonindigenous+species+introductions%3A+A+threat+to+Canada%27s+forests+and+forest+economy&rft.volume=24&rft.issue=2&rft.pages=103-110&rft.date=2002&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A85073955%23id-name%3DS2CID&rft_id=info%3Adoi%2F10.1080%2F07060660309506983&rft_id=info%3Abibcode%2F2002CaJPP..24..103A&rft.aulast=Allen&rft.aufirst=E.+A.&rft.au=Humble%2C+L.+M.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-166"><a href="#cite_ref-166">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWebber2000" class="citation book cs1">Webber, Joan F. (2000). "Insect Vector Behavior and the Evolution of Dutch Elm Disease". The Elms. p. 47. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2F978-1-4615-4507-1_3">10.1007/978-1-4615-4507-1_3</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4613-7032-1" title="Special:BookSources/978-1-4613-7032-1"><bdi>978-1-4613-7032-1</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Insect+Vector+Behavior+and+the+Evolution+of+Dutch+Elm+Disease&rft.btitle=The+Elms&rft.pages=47&rft.date=2000&rft_id=info%3Adoi%2F10.1007%2F978-1-4615-4507-1_3&rft.isbn=978-1-4613-7032-1&rft.aulast=Webber&rft.aufirst=Joan+F.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-167"><a href="#cite_ref-167">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAlyokhin,_A.Baker,_M.Mota-Sanchez,_D.Dively,_G.2008" class="citation journal cs1">Alyokhin, A.; Baker, M.; Mota-Sanchez, D.; Dively, G.; Grafius, E. (2008). "Colorado potato beetle resistance to insecticides". American Journal of Potato Research. 85 (6): 395–413. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs12230-008-9052-0">10.1007/s12230-008-9052-0</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:41206911">41206911</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Journal+of+Potato+Research&rft.atitle=Colorado+potato+beetle+resistance+to+insecticides&rft.volume=85&rft.issue=6&rft.pages=395-413&rft.date=2008&rft_id=info%3Adoi%2F10.1007%2Fs12230-008-9052-0&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A41206911%23id-name%3DS2CID&rft.au=Alyokhin%2C+A.&rft.au=Baker%2C+M.&rft.au=Mota-Sanchez%2C+D.&rft.au=Dively%2C+G.&rft.au=Grafius%2C+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-lockwoodbost-168"><a href="#cite_ref-lockwoodbost_168-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLockwood,_Jeffrey_A.2007" class="citation news cs1">Lockwood, Jeffrey A. (October 21, 2007). <a rel="nofollow" class="external text" href="http://www.boston.com/news/globe/ideas/articles/2007/10/21/bug_bomb">"Bug bomb:Why our next terrorist attack could come on six legs"</a>. Boston Globe. Retrieved February 13, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Boston+Globe&rft.atitle=Bug+bomb%3AWhy+our+next+terrorist+attack+could+come+on+six+legs&rft.date=2007-10-21&rft.au=Lockwood%2C+Jeffrey+A.&rft_id=http%3A%2F%2Fwww.boston.com%2Fnews%2Fglobe%2Fideas%2Farticles%2F2007%2F10%2F21%2Fbug_bomb&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-heather-169"><a href="#cite_ref-heather_169-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHeather,_Neil_W.Hallman,_Guy_J.2008" class="citation book cs1">Heather, Neil W.; Hallman, Guy J. (2008). <a rel="nofollow" class="external text" href="https://archive.org/details/pestmanagementph00hall">Pest Management and Phytosanitary Trade Barriers</a>. CABI. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/pestmanagementph00hall/page/n27">17</a>–18. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1079%2F9781845933432.0000">10.1079/9781845933432.0000</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-84593-343-2" title="Special:BookSources/978-1-84593-343-2"><bdi>978-1-84593-343-2</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Pest+Management+and+Phytosanitary+Trade+Barriers&rft.pages=17-18&rft.pub=CABI&rft.date=2008&rft_id=info%3Adoi%2F10.1079%2F9781845933432.0000&rft.isbn=978-1-84593-343-2&rft.au=Heather%2C+Neil+W.&rft.au=Hallman%2C+Guy+J.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fpestmanagementph00hall&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-170"><a href="#cite_ref-170">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFAdcock,_Edward2005" class="citation web cs1">Adcock, Edward (2005). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20110710144935/http://www.bodley.ox.ac.uk/dept/preservation/training/pests/watch.htm">"Pests – Death watch beetle"</a>. Conservation and collective care. <a href="/wiki/University_of_Oxford" title="University of Oxford">University of Oxford</a>. Archived from <a rel="nofollow" class="external text" href="http://www.bodley.ox.ac.uk/dept/preservation/training/pests/watch.htm">the original</a> on July 10, 2011. Retrieved July 17, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=unknown&rft.jtitle=Conservation+and+collective+care&rft.atitle=Pests+%E2%80%93+Death+watch+beetle&rft.date=2005&rft.au=Adcock%2C+Edward&rft_id=http%3A%2F%2Fwww.bodley.ox.ac.uk%2Fdept%2Fpreservation%2Ftraining%2Fpests%2Fwatch.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-171"><a href="#cite_ref-171">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTakanoTakasuFushimiIchiki2014" class="citation journal cs1">Takano, Shun-Ichiro; Takasu, Keiji; Fushimi, Tsutomu; Ichiki, Ryoko T.; Nakamura, Satoshi (2014). "Suitability of four palm species for the development of the invasive pest Brontispa longissima(Coleoptera: Chrysomelidae) in the field". Entomological Science. 17 (2): 265–268. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fens.12048">10.1111/ens.12048</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:85910791">85910791</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Entomological+Science&rft.atitle=Suitability+of+four+palm+species+for+the+development+of+the+invasive+pest+Brontispa+longissima%28Coleoptera%3A+Chrysomelidae%29+in+the+field&rft.volume=17&rft.issue=2&rft.pages=265-268&rft.date=2014&rft_id=info%3Adoi%2F10.1111%2Fens.12048&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A85910791%23id-name%3DS2CID&rft.aulast=Takano&rft.aufirst=Shun-Ichiro&rft.au=Takasu%2C+Keiji&rft.au=Fushimi%2C+Tsutomu&rft.au=Ichiki%2C+Ryoko+T.&rft.au=Nakamura%2C+Satoshi&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-172"><a href="#cite_ref-172">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20100419214859/http://mpb.cfs.nrcan.gc.ca/biology/introduction_e.html">"The Mountain Pine Beetle in British Columbia"</a>. <a href="/wiki/Natural_Resources_Canada" title="Natural Resources Canada">Natural Resources Canada</a>. August 19, 2008. Archived from <a rel="nofollow" class="external text" href="http://mpb.cfs.nrcan.gc.ca/biology/introduction_e.html">the original</a> on April 19, 2010. Retrieved June 24, 2010.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=The+Mountain+Pine+Beetle+in+British+Columbia&rft.pub=Natural+Resources+Canada&rft.date=2008-08-19&rft_id=http%3A%2F%2Fmpb.cfs.nrcan.gc.ca%2Fbiology%2Fintroduction_e.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-173"><a href="#cite_ref-173">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://entnemdept.ufl.edu/creatures/beneficial/lady_beetles.htm">"Insecta: Coleoptera: Coccinellidae"</a>. Institute of Food and Agricultural Services, <a href="/wiki/University_of_Florida" title="University of Florida">University of Florida</a>. 2014. Retrieved February 12, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Insecta%3A+Coleoptera%3A+Coccinellidae&rft.pub=Institute+of+Food+and+Agricultural+Services%2C+University+of+Florida&rft.date=2014&rft_id=http%3A%2F%2Fentnemdept.ufl.edu%2Fcreatures%2Fbeneficial%2Flady_beetles.htm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-BBC_news-174"><a href="#cite_ref-BBC_news_174-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation news cs1"><a rel="nofollow" class="external text" href="http://news.bbc.co.uk/1/hi/england/essex/3715120.stm">"'Deadly ladybird' sighted in UK"</a>. BBC News. October 5, 2004. Retrieved June 17, 2010.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BBC+News&rft.atitle=%27Deadly+ladybird%27+sighted+in+UK&rft.date=2004-10-05&rft_id=http%3A%2F%2Fnews.bbc.co.uk%2F1%2Fhi%2Fengland%2Fessex%2F3715120.stm&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-175"><a href="#cite_ref-175">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKromp,_B.1999" class="citation journal cs1">Kromp, B. (1999). "Carabid beetles in sustainable agriculture: a review on pest control efficacy, cultivation aspects and enhancement". <a href="/wiki/Agriculture,_Ecosystems_and_Environment" class="mw-redirect" title="Agriculture, Ecosystems and Environment">Agriculture, Ecosystems and Environment</a>. 74 (1–3): 187–228. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1999AgEE...74..187K">1999AgEE...74..187K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0167-8809%2899%2900037-7">10.1016/S0167-8809(99)00037-7</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Agriculture%2C+Ecosystems+and+Environment&rft.atitle=Carabid+beetles+in+sustainable+agriculture%3A+a+review+on+pest+control+efficacy%2C+cultivation+aspects+and+enhancement&rft.volume=74&rft.issue=1%E2%80%933&rft.pages=187-228&rft.date=1999&rft_id=info%3Adoi%2F10.1016%2FS0167-8809%2899%2900037-7&rft_id=info%3Abibcode%2F1999AgEE...74..187K&rft.au=Kromp%2C+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-176"><a href="#cite_ref-176">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="https://web.archive.org/web/20180703050543/https://www.rothamsted.ac.uk/news/beetles-play-important-role-reducing-weeds">"Beetles play an important role in reducing weeds"</a>. Rothamsted Research. Archived from <a rel="nofollow" class="external text" href="http://www.rothamsted.ac.uk/news/beetles-play-important-role-reducing-weeds">the original</a> on July 3, 2018. Retrieved March 14, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Beetles+play+an+important+role+in+reducing+weeds&rft.pub=Rothamsted+Research&rft_id=http%3A%2F%2Fwww.rothamsted.ac.uk%2Fnews%2Fbeetles-play-important-role-reducing-weeds&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-ISC-177"><a href="#cite_ref-ISC_177-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation web cs1"><a rel="nofollow" class="external text" href="http://www.cabi.org/isc/datasheet/57506">"Zygogramma bicolorata (Mexican beetle)"</a>. Invasive Species Compendium. 2012. Retrieved February 12, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Zygogramma+bicolorata+%28Mexican+beetle%29&rft.pub=Invasive+Species+Compendium&rft.date=2012&rft_id=http%3A%2F%2Fwww.cabi.org%2Fisc%2Fdatasheet%2F57506&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-178"><a href="#cite_ref-178">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKovalev,_O._V.Reznik,_S._Y.Cherkashin,_V._N.1983" class="citation journal cs1 cs1-prop-foreign-lang-source">Kovalev, O. V.; Reznik, S. Y.; Cherkashin, V. N. (1983). "Specific features of the methods of using Zygogramma Chevr. (Coleoptera, Chrysomelidae) in biological control of ragweeds (Ambrosia artemisiifolia L., A. psilostachya D.C.)". Entomologicheskoe Obozrenije (in Russian). 62: 402–408.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Entomologicheskoe+Obozrenije&rft.atitle=Specific+features+of+the+methods+of+using+Zygogramma+Chevr.+%28Coleoptera%2C+Chrysomelidae%29+in+biological+control+of+ragweeds+%28Ambrosia+artemisiifolia+L.%2C+A.+psilostachya+D.C.%29&rft.volume=62&rft.pages=402-408&rft.date=1983&rft.au=Kovalev%2C+O.+V.&rft.au=Reznik%2C+S.+Y.&rft.au=Cherkashin%2C+V.+N.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Bornemissza-179">^ <a href="#cite_ref-Bornemissza_179-0">a</a> <a href="#cite_ref-Bornemissza_179-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBornemissza,_George1970" class="citation journal cs1"><a href="/wiki/George_Bornemissza" title="George Bornemissza">Bornemissza, George</a> (1970). <a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1440-6055.1970.tb00767.x">"Insectary Studies on the Control of Dung Breeding Flies by the Activity of the Dung Beetle, Onthophagus Gazella F. (Coleoptera: Scarabaeinae)"</a>. <a href="/wiki/Australian_Journal_of_Entomology" class="mw-redirect" title="Australian Journal of Entomology">Australian Journal of Entomology</a>. 9: 31–41. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Fj.1440-6055.1970.tb00767.x">10.1111/j.1440-6055.1970.tb00767.x</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Australian+Journal+of+Entomology&rft.atitle=Insectary+Studies+on+the+Control+of+Dung+Breeding+Flies+by+the+Activity+of+the+Dung+Beetle%2C+Onthophagus+Gazella+F.+%28Coleoptera%3A+Scarabaeinae%29&rft.volume=9&rft.pages=31-41&rft.date=1970&rft_id=info%3Adoi%2F10.1111%2Fj.1440-6055.1970.tb00767.x&rft.au=Bornemissza%2C+George&rft_id=https%3A%2F%2Fdoi.org%2F10.1111%252Fj.1440-6055.1970.tb00767.x&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-180"><a href="#cite_ref-180">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBrown,_JacquelineScholtz,_Clarke_H.Janeau,_Jean-LouisGrellier,_Seraphine2010" class="citation journal cs1">Brown, Jacqueline; Scholtz, Clarke H.; Janeau, Jean-Louis; Grellier, Seraphine & Podwojewski, Pascal (2010). "Dung beetles (Coleoptera: Scarabaeidae) can improve soil hydrological properties". Applied Soil Ecology. 46 (1): 9–16. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.apsoil.2010.05.010">10.1016/j.apsoil.2010.05.010</a>. <a href="/wiki/Hdl_(identifier)" class="mw-redirect" title="Hdl (identifier)">hdl</a>:<a rel="nofollow" class="external text" href="https://hdl.handle.net/2263%2F14419">2263/14419</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Applied+Soil+Ecology&rft.atitle=Dung+beetles+%28Coleoptera%3A+Scarabaeidae%29+can+improve+soil+hydrological+properties&rft.volume=46&rft.issue=1&rft.pages=9-16&rft.date=2010&rft_id=info%3Ahdl%2F2263%2F14419&rft_id=info%3Adoi%2F10.1016%2Fj.apsoil.2010.05.010&rft.au=Brown%2C+Jacqueline&rft.au=Scholtz%2C+Clarke+H.&rft.au=Janeau%2C+Jean-Louis&rft.au=Grellier%2C+Seraphine&rft.au=Podwojewski%2C+Pascal&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-181"><a href="#cite_ref-181">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLosey,_John_E.Vaughan,_Mace2006" class="citation journal cs1">Losey, John E.; Vaughan, Mace (2006). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20180619202116/http://www.xerces.org/wp-content/uploads/2008/09/economic_value_insects.pdf">"The economic value of ecological services provided by insects"</a> (PDF). BioScience. 56 (4): 311–323. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1641%2F0006-3568%282006%2956%5B311%3ATEVOES%5D2.0.CO%3B2">10.1641/0006-3568(2006)56[311:TEVOES]2.0.CO;2</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:2970747">2970747</a>. Archived from <a rel="nofollow" class="external text" href="http://www.xerces.org/wp-content/uploads/2008/09/economic_value_insects.pdf">the original</a> (PDF) on June 19, 2018. Retrieved July 17, 2011.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BioScience&rft.atitle=The+economic+value+of+ecological+services+provided+by+insects&rft.volume=56&rft.issue=4&rft.pages=311-323&rft.date=2006&rft_id=info%3Adoi%2F10.1641%2F0006-3568%282006%2956%5B311%3ATEVOES%5D2.0.CO%3B2&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A2970747%23id-name%3DS2CID&rft.au=Losey%2C+John+E.&rft.au=Vaughan%2C+Mace&rft_id=http%3A%2F%2Fwww.xerces.org%2Fwp-content%2Fuploads%2F2008%2F09%2Feconomic_value_insects.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-182"><a href="#cite_ref-182">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFTomberlin,_Jeffery_K.Sanford,_Michelle_R.2012" class="citation book cs1">Tomberlin, Jeffery K.; Sanford, Michelle R. (2012). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=EstjD_GgFvQC&pg=PT123">"Forensic entomology and wildlife"</a>. In Huffman, Jane E.; Wallace, John R. (eds.). Wildlife Forensics: Methods and Applications. Developments in Forensic Science. Vol. 6 (2nd ed.). <a href="/wiki/John_Wiley_%26_Sons" class="mw-redirect" title="John Wiley & Sons">John Wiley & Sons</a>. pp. 81–107. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-119-95429-3" title="Special:BookSources/978-1-119-95429-3"><bdi>978-1-119-95429-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Forensic+entomology+and+wildlife&rft.btitle=Wildlife+Forensics%3A+Methods+and+Applications&rft.series=Developments+in+Forensic+Science&rft.pages=81-107&rft.edition=2nd&rft.pub=John+Wiley+%26+Sons&rft.date=2012&rft.isbn=978-1-119-95429-3&rft.au=Tomberlin%2C+Jeffery+K.&rft.au=Sanford%2C+Michelle+R.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DEstjD_GgFvQC%26pg%3DPT123&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-183"><a href="#cite_ref-183">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation audio-visual cs1"><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?v=VxYrVEJp3kY">Dermestid Beetles working on a female Black Bear skull</a>. 2014. <a rel="nofollow" class="external text" href="https://ghostarchive.org/varchive/youtube/20211116/VxYrVEJp3kY">Archived</a> from the original on November 16, 2021. Retrieved January 26, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Dermestid+Beetles+working+on+a+female+Black+Bear+skull&rft.date=2014&rft_id=https%3A%2F%2Fwww.youtube.com%2Fwatch%3Fv%3DVxYrVEJp3kY&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-184"><a href="#cite_ref-184">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFFernández-JalvoMonfort2008" class="citation journal cs1">Fernández-Jalvo, Yolanda; Monfort, Maria Dolores Marín (2008). "Experimental taphonomy in museums: Preparation protocols for skeletons and fossil vertebrates under the scanning electron microscopy". Geobios. 41 (1): 157–181. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2008Geobi..41..157F">2008Geobi..41..157F</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.geobios.2006.06.006">10.1016/j.geobios.2006.06.006</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Geobios&rft.atitle=Experimental+taphonomy+in+museums%3A+Preparation+protocols+for+skeletons+and+fossil+vertebrates+under+the+scanning+electron+microscopy&rft.volume=41&rft.issue=1&rft.pages=157-181&rft.date=2008&rft_id=info%3Adoi%2F10.1016%2Fj.geobios.2006.06.006&rft_id=info%3Abibcode%2F2008Geobi..41..157F&rft.aulast=Fern%C3%A1ndez-Jalvo&rft.aufirst=Yolanda&rft.au=Monfort%2C+Maria+Dolores+Mar%C3%ADn&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-185"><a href="#cite_ref-185">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRamos-ElorduyMenzel,_Peter1998" class="citation book cs1">Ramos-Elorduy, Julieta; Menzel, Peter (1998). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=Q7f1LkFz11gC">Creepy crawly cuisine: the gourmet guide to edible insects</a>. Inner Traditions / Bear & Company. p. 5. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-89281-747-4" title="Special:BookSources/978-0-89281-747-4"><bdi>978-0-89281-747-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Creepy+crawly+cuisine%3A+the+gourmet+guide+to+edible+insects&rft.pages=5&rft.pub=Inner+Traditions+%2F+Bear+%26+Company&rft.date=1998&rft.isbn=978-0-89281-747-4&rft.aulast=Ramos-Elorduy&rft.aufirst=Julieta&rft.au=Menzel%2C+Peter&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DQ7f1LkFz11gC&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-186"><a href="#cite_ref-186">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHolland2013" class="citation magazine cs1">Holland, Jennifer S. (May 14, 2013). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20130606223425/http://news.nationalgeographic.com/news/2013/13/130514-edible-insects-entomophagy-science-food-bugs-beetles">"U.N. Urges Eating Insects; 8 Popular Bugs to Try"</a>. National Geographic. Archived from <a rel="nofollow" class="external text" href="http://news.nationalgeographic.com/news/2013/13/130514-edible-insects-entomophagy-science-food-bugs-beetles/">the original</a> on June 6, 2013. Retrieved January 26, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=National+Geographic&rft.atitle=U.N.+Urges+Eating+Insects%3B+8+Popular+Bugs+to+Try&rft.date=2013-05-14&rft.aulast=Holland&rft.aufirst=Jennifer+S.&rft_id=http%3A%2F%2Fnews.nationalgeographic.com%2Fnews%2F2013%2F13%2F130514-edible-insects-entomophagy-science-food-bugs-beetles%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Meyer-Rochow-187"><a href="#cite_ref-Meyer-Rochow_187-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMeyer-Rochow2017" class="citation journal cs1">Meyer-Rochow, V. B. (January 2017). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5296966">"Therapeutic arthropods and other, largely terrestrial, folk-medicinally important invertebrates: a comparative survey and review"</a>. Journal of Ethnobiology and Ethnomedicine. 13 (9): 9. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1186%2Fs13002-017-0136-0">10.1186/s13002-017-0136-0</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5296966">5296966</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/28173820">28173820</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Ethnobiology+and+Ethnomedicine&rft.atitle=Therapeutic+arthropods+and+other%2C+largely+terrestrial%2C+folk-medicinally+important+invertebrates%3A+a+comparative+survey+and+review.&rft.volume=13&rft.issue=9&rft.pages=9&rft.date=2017-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5296966%23id-name%3DPMC&rft_id=info%3Apmid%2F28173820&rft_id=info%3Adoi%2F10.1186%2Fs13002-017-0136-0&rft.aulast=Meyer-Rochow&rft.aufirst=V.+B.&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC5296966&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-188"><a href="#cite_ref-188">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPearsonCassola1992" class="citation journal cs1">Pearson, David L.; Cassola, Fabio (1992). "World-Wide Species Richness Patterns of Tiger Beetles (Coleoptera: Cicindelidae): Indicator Taxon for Biodiversity and Conservation Studies". Conservation Biology. 6 (3): 376–391. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1992ConBi...6..376P">1992ConBi...6..376P</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1046%2Fj.1523-1739.1992.06030376.x">10.1046/j.1523-1739.1992.06030376.x</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/2386038">2386038</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Conservation+Biology&rft.atitle=World-Wide+Species+Richness+Patterns+of+Tiger+Beetles+%28Coleoptera%3A+Cicindelidae%29%3A+Indicator+Taxon+for+Biodiversity+and+Conservation+Studies&rft.volume=6&rft.issue=3&rft.pages=376-391&rft.date=1992&rft_id=https%3A%2F%2Fwww.jstor.org%2Fstable%2F2386038%23id-name%3DJSTOR&rft_id=info%3Adoi%2F10.1046%2Fj.1523-1739.1992.06030376.x&rft_id=info%3Abibcode%2F1992ConBi...6..376P&rft.aulast=Pearson&rft.aufirst=David+L.&rft.au=Cassola%2C+Fabio&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-189"><a href="#cite_ref-189">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMcGeochVan_RensburgBotes2002" class="citation journal cs1">McGeoch, Melodie A.; Van Rensburg, Berndt J.; Botes, Antoinette (2002). "The verification and application of bioindicators: A case study of dung beetles in a savanna ecosystem". Journal of Applied Ecology. 39 (4): 661–672. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2002JApEc..39..661M">2002JApEc..39..661M</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1046%2Fj.1365-2664.2002.00743.x">10.1046/j.1365-2664.2002.00743.x</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+Applied+Ecology&rft.atitle=The+verification+and+application+of+bioindicators%3A+A+case+study+of+dung+beetles+in+a+savanna+ecosystem&rft.volume=39&rft.issue=4&rft.pages=661-672&rft.date=2002&rft_id=info%3Adoi%2F10.1046%2Fj.1365-2664.2002.00743.x&rft_id=info%3Abibcode%2F2002JApEc..39..661M&rft.aulast=McGeoch&rft.aufirst=Melodie+A.&rft.au=Van+Rensburg%2C+Berndt+J.&rft.au=Botes%2C+Antoinette&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-190"><a href="#cite_ref-190">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFLachatWermelingerGossnerBussler2012" class="citation journal cs1">Lachat, Thibault; Wermelinger, Beat; Gossner, Martin M.; Bussler, Heinz; Isacsson, Gunnar; Müller, Jörg (2012). "Saproxylic beetles as indicator species for dead-wood amount and temperature in European beech forests". Ecological Indicators. 23: 323–331. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.ecolind.2012.04.013">10.1016/j.ecolind.2012.04.013</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Ecological+Indicators&rft.atitle=Saproxylic+beetles+as+indicator+species+for+dead-wood+amount+and+temperature+in+European+beech+forests&rft.volume=23&rft.pages=323-331&rft.date=2012&rft_id=info%3Adoi%2F10.1016%2Fj.ecolind.2012.04.013&rft.aulast=Lachat&rft.aufirst=Thibault&rft.au=Wermelinger%2C+Beat&rft.au=Gossner%2C+Martin+M.&rft.au=Bussler%2C+Heinz&rft.au=Isacsson%2C+Gunnar&rft.au=M%C3%BCller%2C+J%C3%B6rg&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-191"><a href="#cite_ref-191">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBohac1999" class="citation journal cs1">Bohac, Jaroslav (1999). <a rel="nofollow" class="external text" href="http://www.zin.ru/animalia/coleoptera/pdf/bohac_1999_357-372.pdf">"Staphylinid beetles as bioindicators"</a> (PDF). Agriculture, Ecosystems & Environment. 74 (1–3): 357–372. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/1999AgEE...74..357B">1999AgEE...74..357B</a>. <a href="/wiki/CiteSeerX_(identifier)" class="mw-redirect" title="CiteSeerX (identifier)">CiteSeerX</a> <a rel="nofollow" class="external text" href="https://citeseerx.ist.psu.edu/viewdoc/summary?doi=10.1.1.496.4273">10.1.1.496.4273</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2FS0167-8809%2899%2900043-2">10.1016/S0167-8809(99)00043-2</a>. <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170811032854/http://www.zin.ru/Animalia/Coleoptera/pdf/bohac_1999_357-372.pdf">Archived</a> (PDF) from the original on August 11, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Agriculture%2C+Ecosystems+%26+Environment&rft.atitle=Staphylinid+beetles+as+bioindicators&rft.volume=74&rft.issue=1%E2%80%933&rft.pages=357-372&rft.date=1999&rft_id=https%3A%2F%2Fciteseerx.ist.psu.edu%2Fviewdoc%2Fsummary%3Fdoi%3D10.1.1.496.4273%23id-name%3DCiteSeerX&rft_id=info%3Adoi%2F10.1016%2FS0167-8809%2899%2900043-2&rft_id=info%3Abibcode%2F1999AgEE...74..357B&rft.aulast=Bohac&rft.aufirst=Jaroslav&rft_id=http%3A%2F%2Fwww.zin.ru%2Fanimalia%2Fcoleoptera%2Fpdf%2Fbohac_1999_357-372.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-192"><a href="#cite_ref-192">^</a> <a rel="nofollow" class="external text" href="http://www.malaeng.com/blog/?p=4262">Life cycle of the rounded jewel beetles, Sternocera spp. วงจรชีวิตของแมลงทับกลมใช้เวลานานถึง 2 ปี – Siam Insect Zoo-Museum</a>. Malaeng.com (2008-10-20). Retrieved on 2013-04-04. </li> <li id="cite_note-Ivie-193"><a href="#cite_ref-Ivie_193-0">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFIvie,_Michael_A.2002" class="citation book cs1">Ivie, Michael A. (2002). "105. Zopheridae". In Ross H. Arnett; Michael Charles Thomas (eds.). <a rel="nofollow" class="external text" href="https://archive.org/details/americanbeetlesv00jr">American Beetles: Polyphaga: Scarabaeoidea through Curculionoidea</a>. Volume 2 of American Beetles. <a href="/wiki/CRC_Press" title="CRC Press">CRC Press</a>. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/americanbeetlesv00jr/page/n471">457</a>–462. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-8493-0954-0" title="Special:BookSources/978-0-8493-0954-0"><bdi>978-0-8493-0954-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=105.+Zopheridae&rft.btitle=American+Beetles%3A+Polyphaga%3A+Scarabaeoidea+through+Curculionoidea&rft.series=Volume+2+of+American+Beetles&rft.pages=457-462&rft.pub=CRC+Press&rft.date=2002&rft.isbn=978-0-8493-0954-0&rft.au=Ivie%2C+Michael+A.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Famericanbeetlesv00jr&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-194"><a href="#cite_ref-194">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRennessonCesardGrimaud2008" class="citation journal cs1 cs1-prop-foreign-lang-source">Rennesson, Stephane; Cesard, Nicolas; Grimaud, Emmanuel (2008). <a rel="nofollow" class="external text" href="http://www7.inra.fr/opie-insectes/pdf/i151-renesson-et-al.pdf">"Duels en miniature: la délicate mise en scène des combats de scarabées au nord de la Thaïlande"</a> (PDF). Insectes (in French). 3 (151). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20100821174942/http://www.inra.fr/opie-insectes/pdf/i151-renesson-et-al.pdf">Archived</a> (PDF) from the original on August 21, 2010.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Insectes&rft.atitle=Duels+en+miniature%3A+la+d%C3%A9licate+mise+en+sc%C3%A8ne+des+combats+de+scarab%C3%A9es+au+nord+de+la+Tha%C3%AFlande&rft.volume=3&rft.issue=151&rft.date=2008&rft.aulast=Rennesson&rft.aufirst=Stephane&rft.au=Cesard%2C+Nicolas&rft.au=Grimaud%2C+Emmanuel&rft_id=http%3A%2F%2Fwww7.inra.fr%2Fopie-insectes%2Fpdf%2Fi151-renesson-et-al.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-Costa-Neto-195">^ <a href="#cite_ref-Costa-Neto_195-0">a</a> <a href="#cite_ref-Costa-Neto_195-1">b</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFEraldo_Medeiros_Costa-Neto2003" class="citation journal cs1">Eraldo Medeiros Costa-Neto (2003). <a rel="nofollow" class="external text" href="https://dialnet.unirioja.es/descarga/articulo/5294425.pdf">"Entertainment with insects: singing and fighting insects around the world. A brief review"</a> (PDF). Etnobiología. 3: 21–29. Retrieved January 30, 2021.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Etnobiolog%C3%ADa&rft.atitle=Entertainment+with+insects%3A+singing+and+fighting+insects+around+the+world.+A+brief+review&rft.volume=3&rft.pages=21-29&rft.date=2003&rft.au=Eraldo+Medeiros+Costa-Neto&rft_id=https%3A%2F%2Fdialnet.unirioja.es%2Fdescarga%2Farticulo%2F5294425.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-196"><a href="#cite_ref-196">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFPemberton,_R._W.1990" class="citation journal cs1">Pemberton, R. W. (1990). "The Korean water beetle game". Pan-Pacific Entomologist. 66 (2): 172–174.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Pan-Pacific+Entomologist&rft.atitle=The+Korean+water+beetle+game&rft.volume=66&rft.issue=2&rft.pages=172-174&rft.date=1990&rft.au=Pemberton%2C+R.+W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-197"><a href="#cite_ref-197">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBateman,_C.Hulcr,_J.2016" class="citation web cs1">Bateman, C.; Hulcr, J. (2016). <a rel="nofollow" class="external text" href="https://edis.ifas.ufl.edu/pdffiles/FR/FR39800.pdf">"Predaceous Diving Beetles as Pets and the Self-Cleaning Aquarium"</a> (PDF). <a href="/wiki/University_of_Florida" title="University of Florida">University of Florida</a> (IFAS Extension). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20170202054428/https://edis.ifas.ufl.edu/pdffiles/FR/FR39800.pdf">Archived</a> (PDF) from the original on February 2, 2017. Retrieved January 27, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=unknown&rft.btitle=Predaceous+Diving+Beetles+as+Pets+and+the+Self-Cleaning+Aquarium&rft.pub=University+of+Florida+%28IFAS+Extension%29&rft.date=2016&rft.au=Bateman%2C+C.&rft.au=Hulcr%2C+J.&rft_id=https%3A%2F%2Fedis.ifas.ufl.edu%2Fpdffiles%2FFR%2FFR39800.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-199"><a href="#cite_ref-199">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKawahara,_A._Y.2007" class="citation journal cs1">Kawahara, A. Y. (2007). <a rel="nofollow" class="external text" href="https://web.archive.org/web/20160412042904/http://www.flmnh.ufl.edu/mcguire/kawahara/wp-content/uploads/2012/02/Kawahara_2007_Am_Ent_sm.pdf">"Thirty-foot telescopic nets, bug-collecting video games, and beetle pets: Entomology in modern Japan"</a> (PDF). American Entomologist. 53 (3): 160–172. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fae%2F53.3.160">10.1093/ae/53.3.160</a>. Archived from <a rel="nofollow" class="external text" href="http://www.flmnh.ufl.edu/mcguire/kawahara/wp-content/uploads/2012/02/Kawahara_2007_Am_Ent_sm.pdf">the original</a> (PDF) on April 12, 2016. Retrieved January 27, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=American+Entomologist&rft.atitle=Thirty-foot+telescopic+nets%2C+bug-collecting+video+games%2C+and+beetle+pets%3A+Entomology+in+modern+Japan&rft.volume=53&rft.issue=3&rft.pages=160-172&rft.date=2007&rft_id=info%3Adoi%2F10.1093%2Fae%2F53.3.160&rft.au=Kawahara%2C+A.+Y.&rft_id=http%3A%2F%2Fwww.flmnh.ufl.edu%2Fmcguire%2Fkawahara%2Fwp-content%2Fuploads%2F2012%2F02%2FKawahara_2007_Am_Ent_sm.pdf&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-200"><a href="#cite_ref-200">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMay1999" class="citation news cs1">May, Mitchell (July 11, 1999). <a rel="nofollow" class="external text" href="https://www.chicagotribune.com/1999/07/11/yen-for-bugs/">"Yen For Bugs"</a>. <a href="/wiki/Chicago_Tribune" title="Chicago Tribune">Chicago Tribune</a>. Retrieved January 27, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Chicago+Tribune&rft.atitle=Yen+For+Bugs&rft.date=1999-07-11&rft.aulast=May&rft.aufirst=Mitchell&rft_id=https%3A%2F%2Fwww.chicagotribune.com%2F1999%2F07%2F11%2Fyen-for-bugs%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-201"><a href="#cite_ref-201">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWatts1999" class="citation news cs1">Watts, Jonathan (August 11, 1999). <a rel="nofollow" class="external text" href="https://www.theguardian.com/uk/1999/aug/11/jonathanwatts">"Vending machine beetles"</a>. The Guardian. Retrieved January 27, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=The+Guardian&rft.atitle=Vending+machine+beetles&rft.date=1999-08-11&rft.aulast=Watts&rft.aufirst=Jonathan&rft_id=https%3A%2F%2Fwww.theguardian.com%2Fuk%2F1999%2Faug%2F11%2Fjonathanwatts&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-202"><a href="#cite_ref-202">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMorseDanahay2007" class="citation book cs1">Morse, Deborah Denenholz; Danahay, Martin A. (2007). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=2eRn_GqPbBoC&pg=PA5">Victorian Animal Dreams: Representations of Animals in Victorian Literature and Culture</a>. Ashgate Publishing. p. 5. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7546-5511-4" title="Special:BookSources/978-0-7546-5511-4"><bdi>978-0-7546-5511-4</bdi></a>. <q>the Victorian mania for beetle collecting</q></cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Victorian+Animal+Dreams%3A+Representations+of+Animals+in+Victorian+Literature+and+Culture&rft.pages=5&rft.pub=Ashgate+Publishing&rft.date=2007&rft.isbn=978-0-7546-5511-4&rft.aulast=Morse&rft.aufirst=Deborah+Denenholz&rft.au=Danahay%2C+Martin+A.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D2eRn_GqPbBoC%26pg%3DPA5&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-203"><a href="#cite_ref-203">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWallace,_Alfred_Russel1869" class="citation book cs1"><a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Wallace, Alfred Russel</a> (1869). The Malay Archipelago: The land of the orang–utan, and the bird of paradise. A narrative of travel, with sketches of man and nature (1 ed.). Macmillan. pp. vii–xiv.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=The+Malay+Archipelago%3A+The+land+of+the+orang%E2%80%93utan%2C+and+the+bird+of+paradise.+A+narrative+of+travel%2C+with+sketches+of+man+and+nature&rft.pages=vii-xiv&rft.edition=1&rft.pub=Macmillan&rft.date=1869&rft.au=Wallace%2C+Alfred+Russel&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-204"><a href="#cite_ref-204">^</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20131213022851/http://www.swedishbiomimetics.com/biomimetics_folder.pdf">Swedish Biomimetics: The μMist Platform Technology</a> (original URL = <a rel="nofollow" class="external free" href="http://www.swedishbiomimetics.com/biomimetics_folder.pdf">http://www.swedishbiomimetics.com/biomimetics_folder.pdf</a>) (archive date = December 13, 2013) </li> <li id="cite_note-205"><a href="#cite_ref-205">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation news cs1"><a rel="nofollow" class="external text" href="https://www.bbc.co.uk/news/technology-20465982">"Namib Desert beetle inspires self-filling water bottle"</a>. BBC News. November 23, 2012.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=BBC+News&rft.atitle=Namib+Desert+beetle+inspires+self-filling+water+bottle&rft.date=2012-11-23&rft_id=https%3A%2F%2Fwww.bbc.co.uk%2Fnews%2Ftechnology-20465982&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-206"><a href="#cite_ref-206">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSinger2009" class="citation magazine cs1">Singer, Emily (January 29, 2009). <a rel="nofollow" class="external text" href="https://www.technologyreview.com/s/411814/the-armys-remote-controlled-beetle/">"The Army's Remote-Controlled Beetle"</a>. MIT Technology Review. Retrieved March 16, 2017.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=MIT+Technology+Review&rft.atitle=The+Army%27s+Remote-Controlled+Beetle&rft.date=2009-01-29&rft.aulast=Singer&rft.aufirst=Emily&rft_id=https%3A%2F%2Fwww.technologyreview.com%2Fs%2F411814%2Fthe-armys-remote-controlled-beetle%2F&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-207"><a href="#cite_ref-207">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCaoZhangHao_Yu_ChooSato2016" class="citation journal cs1">Cao, Feng; Zhang, Chao; Hao Yu Choo; Sato, Hirotaka (2016). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4843679">"Insect–computer hybrid legged robot with user-adjustable speed, step length and walking gait"</a>. Journal of the Royal Society Interface. 13 (116): 20160060. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsif.2016.0060">10.1098/rsif.2016.0060</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4843679">4843679</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/27030043">27030043</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Journal+of+the+Royal+Society+Interface&rft.atitle=Insect%E2%80%93computer+hybrid+legged+robot+with+user-adjustable+speed%2C+step+length+and+walking+gait&rft.volume=13&rft.issue=116&rft.pages=20160060&rft.date=2016&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4843679%23id-name%3DPMC&rft_id=info%3Apmid%2F27030043&rft_id=info%3Adoi%2F10.1098%2Frsif.2016.0060&rft.aulast=Cao&rft.aufirst=Feng&rft.au=Zhang%2C+Chao&rft.au=Hao+Yu+Choo&rft.au=Sato%2C+Hirotaka&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC4843679&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-208"><a href="#cite_ref-208">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFSatoDoanKolevHuynh2015" class="citation journal cs1">Sato, Hirotaka; Doan, Tat Thang Vo; Kolev, Svetoslav; Huynh, Ngoc Anh; Zhang, Chao; Massey, Travis L.; Kleef, Joshua van; Ikeda, Kazuo; <a href="/wiki/Pieter_Abbeel" title="Pieter Abbeel">Abbeel, Pieter</a> (March 16, 2015). <a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2015.01.051">"Deciphering the Role of a Coleopteran Steering Muscle via Free Flight Stimulation"</a>. Current Biology. 25 (6): 798–803. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.cub.2015.01.051">10.1016/j.cub.2015.01.051</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/25784033">25784033</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Current+Biology&rft.atitle=Deciphering+the+Role+of+a+Coleopteran+Steering+Muscle+via+Free+Flight+Stimulation&rft.volume=25&rft.issue=6&rft.pages=798-803&rft.date=2015-03-16&rft_id=info%3Adoi%2F10.1016%2Fj.cub.2015.01.051&rft_id=info%3Apmid%2F25784033&rft.aulast=Sato&rft.aufirst=Hirotaka&rft.au=Doan%2C+Tat+Thang+Vo&rft.au=Kolev%2C+Svetoslav&rft.au=Huynh%2C+Ngoc+Anh&rft.au=Zhang%2C+Chao&rft.au=Massey%2C+Travis+L.&rft.au=Kleef%2C+Joshua+van&rft.au=Ikeda%2C+Kazuo&rft.au=Abbeel%2C+Pieter&rft_id=https%3A%2F%2Fdoi.org%2F10.1016%252Fj.cub.2015.01.051&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-209"><a href="#cite_ref-209">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVo_DoanTanBuiSato2017" class="citation journal cs1">Vo Doan, Tat Thang; Tan, Melvin Y.W.; Bui, Xuan Hien; Sato, Hirotaka (November 3, 2017). "An Ultralightweight and Living Legged Robot". Soft Robotics. 5 (1): 17–23. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1089%2Fsoro.2017.0038">10.1089/soro.2017.0038</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/29412086">29412086</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Soft+Robotics&rft.atitle=An+Ultralightweight+and+Living+Legged+Robot&rft.volume=5&rft.issue=1&rft.pages=17-23&rft.date=2017-11-03&rft_id=info%3Adoi%2F10.1089%2Fsoro.2017.0038&rft_id=info%3Apmid%2F29412086&rft.aulast=Vo+Doan&rft.aufirst=Tat+Thang&rft.au=Tan%2C+Melvin+Y.W.&rft.au=Bui%2C+Xuan+Hien&rft.au=Sato%2C+Hirotaka&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-210"><a href="#cite_ref-210">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNguyenDungSatoVo-Doan2023" class="citation journal cs1">Nguyen, H. Duoc; Dung, V. Than; Sato, Hirotaka; Vo-Doan, T. Thang (February 1, 2023). <a rel="nofollow" class="external text" href="https://www.sciencedirect.com/science/article/abs/pii/S0925400522016318">"Efficient autonomous navigation for terrestrial insect-machine hybrid systems"</a>. Sensors and Actuators B: Chemical. 376: 132988. <a href="/wiki/ArXiv_(identifier)" class="mw-redirect" title="ArXiv (identifier)">arXiv</a>:<a rel="nofollow" class="external text" href="https://arxiv.org/abs/2204.13281">2204.13281</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1016%2Fj.snb.2022.132988">10.1016/j.snb.2022.132988</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/0925-4005">0925-4005</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Sensors+and+Actuators+B%3A+Chemical&rft.atitle=Efficient+autonomous+navigation+for+terrestrial+insect-machine+hybrid+systems&rft.volume=376&rft.pages=132988&rft.date=2023-02-01&rft_id=info%3Aarxiv%2F2204.13281&rft.issn=0925-4005&rft_id=info%3Adoi%2F10.1016%2Fj.snb.2022.132988&rft.aulast=Nguyen&rft.aufirst=H.+Duoc&rft.au=Dung%2C+V.+Than&rft.au=Sato%2C+Hirotaka&rft.au=Vo-Doan%2C+T.+Thang&rft_id=https%3A%2F%2Fwww.sciencedirect.com%2Fscience%2Farticle%2Fabs%2Fpii%2FS0925400522016318&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-211"><a href="#cite_ref-211">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVo-DoanDungSato2022" class="citation journal cs1">Vo-Doan, T. Thang; Dung, V. Than; Sato, Hirotaka (January 2022). <a rel="nofollow" class="external text" href="https://spj.science.org/doi/10.34133/2022/9780504">"A Cyborg Insect Reveals a Function of a Muscle in Free Flight"</a>. Cyborg and Bionic Systems. 2022. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.34133%2F2022%2F9780504">10.34133/2022/9780504</a>. <a href="/wiki/ISSN_(identifier)" class="mw-redirect" title="ISSN (identifier)">ISSN</a> <a rel="nofollow" class="external text" href="https://search.worldcat.org/issn/2692-7632">2692-7632</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC9494732">9494732</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/36285304">36285304</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Cyborg+and+Bionic+Systems&rft.atitle=A+Cyborg+Insect+Reveals+a+Function+of+a+Muscle+in+Free+Flight&rft.volume=2022&rft.date=2022-01&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC9494732%23id-name%3DPMC&rft.issn=2692-7632&rft_id=info%3Apmid%2F36285304&rft_id=info%3Adoi%2F10.34133%2F2022%2F9780504&rft.aulast=Vo-Doan&rft.aufirst=T.+Thang&rft.au=Dung%2C+V.+Than&rft.au=Sato%2C+Hirotaka&rft_id=https%3A%2F%2Fspj.science.org%2Fdoi%2F10.34133%2F2022%2F9780504&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-212"><a href="#cite_ref-212">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVikram_Iyer1Ali_NajafiJohannes_JamesSawyer_Fuller2020" class="citation journal cs1">Vikram Iyer1; Ali Najafi; Johannes James; Sawyer Fuller; Shyamnath Gollakota (July 2020). <a rel="nofollow" class="external text" href="https://www.science.org/doi/10.1126/scirobotics.abb0839">"Wireless steerable vision for live insects and insect-scale robots"</a>. Science Robotics. 5 (44): eabb0839. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1126%2Fscirobotics.abb0839">10.1126/scirobotics.abb0839</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/33022605">33022605</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:220688078">220688078</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Science+Robotics&rft.atitle=Wireless+steerable+vision+for+live+insects+and+insect-scale+robots&rft.volume=5&rft.issue=44&rft.pages=eabb0839&rft.date=2020-07&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A220688078%23id-name%3DS2CID&rft_id=info%3Apmid%2F33022605&rft_id=info%3Adoi%2F10.1126%2Fscirobotics.abb0839&rft.au=Vikram+Iyer1&rft.au=Ali+Najafi&rft.au=Johannes+James&rft.au=Sawyer+Fuller&rft.au=Shyamnath+Gollakota&rft_id=https%3A%2F%2Fwww.science.org%2Fdoi%2F10.1126%2Fscirobotics.abb0839&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"><code class="cs1-code">{{<a href="/wiki/Template:Cite_journal" title="Template:Cite journal">cite journal</a>}}</code>: CS1 maint: numeric names: authors list (<a href="/wiki/Category:CS1_maint:_numeric_names:_authors_list" title="Category:CS1 maint: numeric names: authors list">link</a>) </li> <li id="cite_note-213"><a href="#cite_ref-213">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFRob_Picheta2020" class="citation news cs1">Rob Picheta (July 15, 2020). <a rel="nofollow" class="external text" href="https://edition.cnn.com/2020/07/15/us/beetle-tiny-cameras-scli-scn-intl/index.html">"Scientists strapped a tiny camera to a beetle to test just how small video technology can get"</a>. CNN.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=CNN&rft.atitle=Scientists+strapped+a+tiny+camera+to+a+beetle+to+test+just+how+small+video+technology+can+get&rft.date=2020-07-15&rft.au=Rob+Picheta&rft_id=https%3A%2F%2Fedition.cnn.com%2F2020%2F07%2F15%2Fus%2Fbeetle-tiny-cameras-scli-scn-intl%2Findex.html&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-214"><a href="#cite_ref-214">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFKotzeO'Hara2003" class="citation journal cs1">Kotze, D. Johan; O'Hara, Robert B. (2003). "Species decline—but why? Explanations of carabid beetle (Coleoptera, Carabidae) declines in Europe". Oecologia. 135 (1): 138–148. <a href="/wiki/Bibcode_(identifier)" class="mw-redirect" title="Bibcode (identifier)">Bibcode</a>:<a rel="nofollow" class="external text" href="https://ui.adsabs.harvard.edu/abs/2003Oecol.135..138K">2003Oecol.135..138K</a>. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs00442-002-1174-3">10.1007/s00442-002-1174-3</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/12647113">12647113</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:11692514">11692514</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Oecologia&rft.atitle=Species+decline%E2%80%94but+why%3F+Explanations+of+carabid+beetle+%28Coleoptera%2C+Carabidae%29+declines+in+Europe&rft.volume=135&rft.issue=1&rft.pages=138-148&rft.date=2003&rft_id=info%3Adoi%2F10.1007%2Fs00442-002-1174-3&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A11692514%23id-name%3DS2CID&rft_id=info%3Apmid%2F12647113&rft_id=info%3Abibcode%2F2003Oecol.135..138K&rft.aulast=Kotze&rft.aufirst=D.+Johan&rft.au=O%27Hara%2C+Robert+B.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-215"><a href="#cite_ref-215">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHanskiKoivulehtoCameronRahagalala2007" class="citation journal cs1">Hanski, Ilkka; Koivulehto, Helena; Cameron, Alison; Rahagalala, Pierre (2007). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1995085">"Deforestation and apparent extinctions of endemic forest beetles in Madagascar"</a>. Biology Letters. 3 (3): 344–347. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1098%2Frsbl.2007.0043">10.1098/rsbl.2007.0043</a>. <a href="/wiki/PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1995085">1995085</a>. <a href="/wiki/PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/17341451">17341451</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biology+Letters&rft.atitle=Deforestation+and+apparent+extinctions+of+endemic+forest+beetles+in+Madagascar&rft.volume=3&rft.issue=3&rft.pages=344-347&rft.date=2007&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1995085%23id-name%3DPMC&rft_id=info%3Apmid%2F17341451&rft_id=info%3Adoi%2F10.1098%2Frsbl.2007.0043&rft.aulast=Hanski&rft.aufirst=Ilkka&rft.au=Koivulehto%2C+Helena&rft.au=Cameron%2C+Alison&rft.au=Rahagalala%2C+Pierre&rft_id=https%3A%2F%2Fwww.ncbi.nlm.nih.gov%2Fpmc%2Farticles%2FPMC1995085&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-216"><a href="#cite_ref-216">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCampanaro,_A.Zapponi,_L.Hardersen,_S.Méndez,_M.2016" class="citation journal cs1">Campanaro, A.; Zapponi, L.; Hardersen, S.; Méndez, M.; Al Fulaij, N.; Audisio, P.; Bardiani, M.; Carpaneto, G. M.; Corezzola, S.; Della Rocca, F.; Harvey, D.; Hawes, C.; Kadej, M.; Karg, J.; Rink, M.; Smolis, A.; Sprecher, E.; Thomaes, A.; Toni, I.; Vrezec, A.; Zauli, A.; Zilioli, M.; Chiari, S. (2016). "A European monitoring protocol for the stag beetle, a saproxylic flagship species". Insect Conservation and Diversity. 9 (6): 574–584. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1111%2Ficad.12194">10.1111/icad.12194</a>. <a href="/wiki/S2CID_(identifier)" class="mw-redirect" title="S2CID (identifier)">S2CID</a> <a rel="nofollow" class="external text" href="https://api.semanticscholar.org/CorpusID:88754595">88754595</a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Insect+Conservation+and+Diversity&rft.atitle=A+European+monitoring+protocol+for+the+stag+beetle%2C+a+saproxylic+flagship+species&rft.volume=9&rft.issue=6&rft.pages=574-584&rft.date=2016&rft_id=info%3Adoi%2F10.1111%2Ficad.12194&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A88754595%23id-name%3DS2CID&rft.au=Campanaro%2C+A.&rft.au=Zapponi%2C+L.&rft.au=Hardersen%2C+S.&rft.au=M%C3%A9ndez%2C+M.&rft.au=Al+Fulaij%2C+N.&rft.au=Audisio%2C+P.&rft.au=Bardiani%2C+M.&rft.au=Carpaneto%2C+G.+M.&rft.au=Corezzola%2C+S.&rft.au=Della+Rocca%2C+F.&rft.au=Harvey%2C+D.&rft.au=Hawes%2C+C.&rft.au=Kadej%2C+M.&rft.au=Karg%2C+J.&rft.au=Rink%2C+M.&rft.au=Smolis%2C+A.&rft.au=Sprecher%2C+E.&rft.au=Thomaes%2C+A.&rft.au=Toni%2C+I.&rft.au=Vrezec%2C+A.&rft.au=Zauli%2C+A.&rft.au=Zilioli%2C+M.&rft.au=Chiari%2C+S.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> <li id="cite_note-217"><a href="#cite_ref-217">^</a> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFNew2009" class="citation book cs1">New, T. R. (2009). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=44uv6YiZH4YC&pg=PR4">Beetles in Conservation</a>. John Wiley & Sons. pp. ix, 1–26, passim. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-1-4443-1863-0" title="Special:BookSources/978-1-4443-1863-0"><bdi>978-1-4443-1863-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Beetles+in+Conservation&rft.pages=ix%2C+1-26%2C+passim&rft.pub=John+Wiley+%26+Sons&rft.date=2009&rft.isbn=978-1-4443-1863-0&rft.aulast=New&rft.aufirst=T.+R.&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3D44uv6YiZH4YC%26pg%3DPR4&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"> </li> </ol></div></div> <div class="mw-heading mw-heading3"><h3 id="Bibliography">Bibliography</h3>[<a href="/w/index.php?title=Beetle&action=edit&section=58" title="Edit section: Bibliography">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1239549316">.mw-parser-output .refbegin{margin-bottom:0.5em}.mw-parser-output .refbegin-hanging-indents>ul{margin-left:0}.mw-parser-output .refbegin-hanging-indents>ul>li{margin-left:0;padding-left:3.2em;text-indent:-3.2em}.mw-parser-output .refbegin-hanging-indents ul,.mw-parser-output .refbegin-hanging-indents ul li{list-style:none}@media(max-width:720px){.mw-parser-output .refbegin-hanging-indents>ul>li{padding-left:1.6em;text-indent:-1.6em}}.mw-parser-output .refbegin-columns{margin-top:0.3em}.mw-parser-output .refbegin-columns ul{margin-top:0}.mw-parser-output .refbegin-columns li{page-break-inside:avoid;break-inside:avoid-column}@media screen{.mw-parser-output .refbegin{font-size:90%}}</style><div class="refbegin" style=""> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="refEvans" class="citation book cs1">Evans, Arthur V.; Bellamy, Charles (2000). <a rel="nofollow" class="external text" href="https://books.google.com/books?id=ZZ_hfpMo8oAC&pg=PA31">An Inordinate Fondness for Beetles</a>. <a href="/wiki/University_of_California_Press" title="University of California Press">University of California Press</a>. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-520-22323-3" title="Special:BookSources/978-0-520-22323-3"><bdi>978-0-520-22323-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=An+Inordinate+Fondness+for+Beetles&rft.pub=University+of+California+Press&rft.date=2000&rft.isbn=978-0-520-22323-3&rft.aulast=Evans&rft.aufirst=Arthur+V.&rft.au=Bellamy%2C+Charles&rft_id=https%3A%2F%2Fbooks.google.com%2Fbooks%3Fid%3DZZ_hfpMo8oAC%26pg%3DPA31&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="refMcHugh" class="citation book cs1">McHugh, Joseph V.; Liebherr, James K. (2009). "Coleoptera". In Vincent H. Resh; Ring T. Cardé (eds.). <a rel="nofollow" class="external text" href="https://archive.org/details/encyclopediainse00resh_059">Encyclopedia of Insects</a> (2nd ed.). Academic Press. pp. <a rel="nofollow" class="external text" href="https://archive.org/details/encyclopediainse00resh_059/page/n219">183</a>–201. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-12-374144-8" title="Special:BookSources/978-0-12-374144-8"><bdi>978-0-12-374144-8</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=bookitem&rft.atitle=Coleoptera&rft.btitle=Encyclopedia+of+Insects&rft.pages=183-201&rft.edition=2nd&rft.pub=Academic+Press&rft.date=2009&rft.isbn=978-0-12-374144-8&rft.aulast=McHugh&rft.aufirst=Joseph+V.&rft.au=Liebherr%2C+James+K.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fencyclopediainse00resh_059&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li></ul> </div> <div class="mw-heading mw-heading2"><h2 id="Further_reading">Further reading</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=59" title="Edit section: Further reading">edit</a>]</div> <ul><li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFBeckmann,_Poul2001" class="citation book cs1">Beckmann, Poul (2001). Living Jewels: The Natural Design of Beetles. Prestel. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-3-7913-2528-6" title="Special:BookSources/978-3-7913-2528-6"><bdi>978-3-7913-2528-6</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Living+Jewels%3A+The+Natural+Design+of+Beetles&rft.pub=Prestel&rft.date=2001&rft.isbn=978-3-7913-2528-6&rft.au=Beckmann%2C+Poul&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFCooter,_J.M._V._L._Barclay2006" class="citation book cs1">Cooter, J.; <a href="/wiki/Max_Barclay" title="Max Barclay">M. V. L. Barclay</a>, eds. (2006). A Coleopterist's Handbook. Amateur Entomological Society. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-900054-70-9" title="Special:BookSources/978-0-900054-70-9"><bdi>978-0-900054-70-9</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=A+Coleopterist%27s+Handbook&rft.pub=Amateur+Entomological+Society&rft.date=2006&rft.isbn=978-0-900054-70-9&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite class="citation book cs1">Beetle Larvae of the World. <a href="/wiki/Entomological_Society_of_America" title="Entomological Society of America">Entomological Society of America</a>. December 1994. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-643-05506-3" title="Special:BookSources/978-0-643-05506-3"><bdi>978-0-643-05506-3</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Beetle+Larvae+of+the+World&rft.pub=Entomological+Society+of+America&rft.date=1994-12&rft.isbn=978-0-643-05506-3&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFGrimaldi,_DavidMichael_S._Engel2005" class="citation book cs1"><a href="/wiki/David_Grimaldi_(entomologist)" title="David Grimaldi (entomologist)">Grimaldi, David</a>; <a href="/wiki/Michael_S._Engel" title="Michael S. Engel">Michael S. Engel</a> (May 16, 2005). Evolution of the Insects. Cambridge University Press. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-521-82149-0" title="Special:BookSources/978-0-521-82149-0"><bdi>978-0-521-82149-0</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Evolution+of+the+Insects&rft.pub=Cambridge+University+Press&rft.date=2005-05-16&rft.isbn=978-0-521-82149-0&rft.au=Grimaldi%2C+David&rft.au=Michael+S.+Engel&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFHarde,_K._W.1984" class="citation book cs1">Harde, K. W. (1984). A Field Guide in Color to Beetles. Octopus Books. pp. 7–24. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-7064-1937-5" title="Special:BookSources/978-0-7064-1937-5"><bdi>978-0-7064-1937-5</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=A+Field+Guide+in+Color+to+Beetles&rft.pages=7-24&rft.pub=Octopus+Books&rft.date=1984&rft.isbn=978-0-7064-1937-5&rft.au=Harde%2C+K.+W.&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li> <li><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFWhite,_R._E.1983" class="citation book cs1">White, R. E. (1983). <a rel="nofollow" class="external text" href="https://archive.org/details/isbn_9780395910894">Beetles</a>. Houghton Mifflin. <a href="/wiki/ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <a href="/wiki/Special:BookSources/978-0-395-91089-4" title="Special:BookSources/978-0-395-91089-4"><bdi>978-0-395-91089-4</bdi></a>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Abook&rft.genre=book&rft.btitle=Beetles&rft.pub=Houghton+Mifflin&rft.date=1983&rft.isbn=978-0-395-91089-4&rft.au=White%2C+R.+E.&rft_id=https%3A%2F%2Farchive.org%2Fdetails%2Fisbn_9780395910894&rfr_id=info%3Asid%2Fen.wikipedia.org%3ABeetle" class="Z3988"></li></ul> <div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2>[<a href="/w/index.php?title=Beetle&action=edit&section=60" title="Edit section: External links">edit</a>]</div> <style data-mw-deduplicate="TemplateStyles:r1235681985">.mw-parser-output .side-box{margin:4px 0;box-sizing:border-box;border:1px solid #aaa;font-size:88%;line-height:1.25em;background-color:var(--background-color-interactive-subtle,#f8f9fa);display:flow-root}.mw-parser-output .side-box-abovebelow,.mw-parser-output .side-box-text{padding:0.25em 0.9em}.mw-parser-output .side-box-image{padding:2px 0 2px 0.9em;text-align:center}.mw-parser-output .side-box-imageright{padding:2px 0.9em 2px 0;text-align:center}@media(min-width:500px){.mw-parser-output .side-box-flex{display:flex;align-items:center}.mw-parser-output .side-box-text{flex:1;min-width:0}}@media(min-width:720px){.mw-parser-output .side-box{width:238px}.mw-parser-output .side-box-right{clear:right;float:right;margin-left:1em}.mw-parser-output .side-box-left{margin-right:1em}}</style><style data-mw-deduplicate="TemplateStyles:r1237033735">@media print{body.ns-0 .mw-parser-output .sistersitebox{display:none!important}}@media screen{html.skin-theme-clientpref-night .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}@media screen and (prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .sistersitebox img[src*="Wiktionary-logo-en-v2.svg"]{background-color:white}}</style><div class="side-box side-box-right plainlinks sistersitebox"><style data-mw-deduplicate="TemplateStyles:r1126788409">.mw-parser-output .plainlist ol,.mw-parser-output .plainlist ul{line-height:inherit;list-style:none;margin:0;padding:0}.mw-parser-output .plainlist ol li,.mw-parser-output .plainlist ul li{margin-bottom:0}</style> <div class="side-box-flex"> <div class="side-box-image"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/34px-Wikispecies-logo.svg.png" decoding="async" width="34" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/51px-Wikispecies-logo.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/68px-Wikispecies-logo.svg.png 2x" data-file-width="941" data-file-height="1103" /></div> <div class="side-box-text plainlist"><a href="/wiki/Wikispecies" title="Wikispecies">Wikispecies</a> has information related to <a href="https://species.wikimedia.org/wiki/Coleoptera" class="extiw" title="wikispecies:Coleoptera">Coleoptera</a>.</div></div> </div> <link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1235681985"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1237033735"><div class="side-box side-box-right plainlinks sistersitebox"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1126788409"> <div class="side-box-flex"> <div class="side-box-image"><img alt="" src="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikibooks-logo-en-noslogan.svg/40px-Wikibooks-logo-en-noslogan.svg.png" decoding="async" width="40" height="40" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikibooks-logo-en-noslogan.svg/60px-Wikibooks-logo-en-noslogan.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikibooks-logo-en-noslogan.svg/80px-Wikibooks-logo-en-noslogan.svg.png 2x" data-file-width="400" data-file-height="400" /></div> <div class="side-box-text plainlist">The Wikibook <a href="https://en.wikibooks.org/wiki/Dichotomous_Key" class="extiw" title="wikibooks:Dichotomous Key">Dichotomous Key</a> has a page on the topic of: <a href="https://en.wikibooks.org/wiki/Dichotomous_Key/Coleoptera" class="extiw" title="wikibooks:Dichotomous Key/Coleoptera">Coleoptera</a></div></div> </div> <ul><li><a rel="nofollow" class="external text" href="http://tolweb.org/Coleoptera/8221">Coleoptera</a> from the <a href="/wiki/Tree_of_Life_Web_Project" title="Tree of Life Web Project">Tree of Life Web Project</a></li> <li>(in German) <a rel="nofollow" class="external text" href="https://www.kaefer-der-welt.de/index.htm">Käfer der Welt</a></li> <li><a rel="nofollow" class="external text" href="http://www.coleoptera-atlas.com/">Coleoptera Atlas</a> <a rel="nofollow" class="external text" href="https://web.archive.org/web/20140106113024/http://www.coleoptera-atlas.com/">Archived</a> January 6, 2014, at the <a href="/wiki/Wayback_Machine" title="Wayback Machine">Wayback Machine</a></li> <li><a rel="nofollow" class="external text" href="https://web.archive.org/web/20120313193822/http://www.insectlifeforms.com/Orders736/BEETLES_COLEOPTERA_60502060_736.aspx">Beetles – Coleoptera</a></li></ul> <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 ul{display:inline}.mw-parser-output .hlist .mw-empty-li{display:none}.mw-parser-output .hlist dt::after{content:": "}.mw-parser-output .hlist dd::after,.mw-parser-output .hlist li::after{content:" · ";font-weight:bold}.mw-parser-output .hlist dd:last-child::after,.mw-parser-output .hlist dt:last-child::after,.mw-parser-output .hlist li:last-child::after{content:none}.mw-parser-output .hlist dd dd:first-child::before,.mw-parser-output .hlist dd dt:first-child::before,.mw-parser-output .hlist dd li:first-child::before,.mw-parser-output .hlist dt dd:first-child::before,.mw-parser-output .hlist dt dt:first-child::before,.mw-parser-output .hlist dt li:first-child::before,.mw-parser-output .hlist li dd:first-child::before,.mw-parser-output .hlist li dt:first-child::before,.mw-parser-output .hlist li li:first-child::before{content:" (";font-weight:normal}.mw-parser-output .hlist dd dd:last-child::after,.mw-parser-output .hlist dd dt:last-child::after,.mw-parser-output .hlist dd li:last-child::after,.mw-parser-output .hlist dt dd:last-child::after,.mw-parser-output .hlist dt dt:last-child::after,.mw-parser-output .hlist dt li:last-child::after,.mw-parser-output .hlist li dd:last-child::after,.mw-parser-output .hlist li dt:last-child::after,.mw-parser-output .hlist li li:last-child::after{content:")";font-weight:normal}.mw-parser-output .hlist ol{counter-reset:listitem}.mw-parser-output .hlist ol>li{counter-increment:listitem}.mw-parser-output .hlist ol>li::before{content:" "counter(listitem)"\a0 "}.mw-parser-output .hlist dd ol>li:first-child::before,.mw-parser-output .hlist dt ol>li:first-child::before,.mw-parser-output .hlist li ol>li:first-child::before{content:" ("counter(listitem)"\a0 "}</style><style data-mw-deduplicate="TemplateStyles:r1236075235">.mw-parser-output .navbox{box-sizing:border-box;border:1px solid #a2a9b1;width:100%;clear:both;font-size:88%;text-align:center;padding:1px;margin:1em auto 0}.mw-parser-output .navbox .navbox{margin-top:0}.mw-parser-output .navbox+.navbox,.mw-parser-output .navbox+.navbox-styles+.navbox{margin-top:-1px}.mw-parser-output .navbox-inner,.mw-parser-output .navbox-subgroup{width:100%}.mw-parser-output .navbox-group,.mw-parser-output .navbox-title,.mw-parser-output .navbox-abovebelow{padding:0.25em 1em;line-height:1.5em;text-align:center}.mw-parser-output .navbox-group{white-space:nowrap;text-align:right}.mw-parser-output .navbox,.mw-parser-output .navbox-subgroup{background-color:#fdfdfd}.mw-parser-output .navbox-list{line-height:1.5em;border-color:#fdfdfd}.mw-parser-output .navbox-list-with-group{text-align:left;border-left-width:2px;border-left-style:solid}.mw-parser-output tr+tr>.navbox-abovebelow,.mw-parser-output tr+tr>.navbox-group,.mw-parser-output tr+tr>.navbox-image,.mw-parser-output tr+tr>.navbox-list{border-top:2px solid #fdfdfd}.mw-parser-output .navbox-title{background-color:#ccf}.mw-parser-output .navbox-abovebelow,.mw-parser-output .navbox-group,.mw-parser-output .navbox-subgroup .navbox-title{background-color:#ddf}.mw-parser-output .navbox-subgroup .navbox-group,.mw-parser-output .navbox-subgroup .navbox-abovebelow{background-color:#e6e6ff}.mw-parser-output .navbox-even{background-color:#f7f7f7}.mw-parser-output .navbox-odd{background-color:transparent}.mw-parser-output .navbox .hlist td dl,.mw-parser-output .navbox .hlist td ol,.mw-parser-output .navbox .hlist td ul,.mw-parser-output .navbox td.hlist dl,.mw-parser-output .navbox td.hlist ol,.mw-parser-output .navbox td.hlist ul{padding:0.125em 0}.mw-parser-output .navbox .navbar{display:block;font-size:100%}.mw-parser-output .navbox-title .navbar{float:left;text-align:left;margin-right:0.5em}body.skin--responsive .mw-parser-output .navbox-image img{max-width:none!important}@media print{body.ns-0 .mw-parser-output .navbox{display:none!important}}</style></div><div role="navigation" class="navbox" aria-labelledby="Insect_orders" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed 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"><style data-mw-deduplicate="TemplateStyles:r1239400231">.mw-parser-output .navbar{display:inline;font-size:88%;font-weight:normal}.mw-parser-output .navbar-collapse{float:left;text-align:left}.mw-parser-output .navbar-boxtext{word-spacing:0}.mw-parser-output .navbar ul{display:inline-block;white-space:nowrap;line-height:inherit}.mw-parser-output .navbar-brackets::before{margin-right:-0.125em;content:"[ "}.mw-parser-output .navbar-brackets::after{margin-left:-0.125em;content:" ]"}.mw-parser-output .navbar li{word-spacing:-0.125em}.mw-parser-output .navbar a>span,.mw-parser-output .navbar a>abbr{text-decoration:inherit}.mw-parser-output .navbar-mini abbr{font-variant:small-caps;border-bottom:none;text-decoration:none;cursor:inherit}.mw-parser-output .navbar-ct-full{font-size:114%;margin:0 7em}.mw-parser-output .navbar-ct-mini{font-size:114%;margin:0 4em}html.skin-theme-clientpref-night .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}@media(prefers-color-scheme:dark){html.skin-theme-clientpref-os .mw-parser-output .navbar li a abbr{color:var(--color-base)!important}}@media print{.mw-parser-output .navbar{display:none!important}}</style><div class="navbar plainlinks hlist navbar-mini"><ul><li class="nv-view"><a href="/wiki/Template:Orders_of_Insects" title="Template:Orders of Insects"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Orders_of_Insects" title="Template talk:Orders of Insects"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Orders_of_Insects" title="Special:EditPage/Template:Orders of Insects"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Insect_orders" style="font-size:114%;margin:0 4em"><a href="/wiki/Insect" title="Insect">Insect</a> <a href="/wiki/Category:Insect_orders" title="Category:Insect orders">orders</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li>Kingdom: <a href="/wiki/Animal" title="Animal">Animalia</a></li> <li>Phylum: <a href="/wiki/Arthropod" title="Arthropod">Arthropoda</a></li> <li>(unranked): <a href="/wiki/Pancrustacea" title="Pancrustacea">Pancrustacea</a></li> <li>Subphylum: <a href="/wiki/Hexapoda" title="Hexapoda">Hexapoda</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible uncollapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Extant" style="font-size:114%;margin:0 4em">Extant</div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Monocondylia" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Archaeognatha" title="Archaeognatha">Monocondylia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Archaeognatha" title="Archaeognatha">Archaeognatha (jumping bristletails)</a></li></ul> </div></td></tr></tbody></table><div></div></td><td class="noviewer navbox-image" rowspan="2" style="width:1px;padding:0 0 0 2px"><div><a href="/wiki/File:Insect_collage.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/7/74/Insect_collage.png/75px-Insect_collage.png" decoding="async" width="75" height="73" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/7/74/Insect_collage.png/113px-Insect_collage.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/7/74/Insect_collage.png/150px-Insect_collage.png 2x" data-file-width="2967" data-file-height="2887" /></a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1em;text-align:center;"><a href="/wiki/Dicondylia" title="Dicondylia">D i c o n d y l i a</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Zygentoma" title="Zygentoma">Zygentoma (silverfish, firebrats)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1em;text-align:center;"><a href="/wiki/Pterygota" title="Pterygota">P t e r y g o t a</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Palaeoptera" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Palaeoptera" title="Palaeoptera">Palaeoptera</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Mayfly" title="Mayfly">Ephemeropteroidea</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Mayfly" title="Mayfly">Ephemeroptera (mayflies)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Odonatoptera" title="Odonatoptera">Odonatoptera</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Odonata" title="Odonata">Odonata (dragonflies, damselflies)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1em;text-align:center;"><a href="/wiki/Neoptera" title="Neoptera">N e o p t e r a</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Polyneoptera" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Polyneoptera" title="Polyneoptera">Polyneoptera</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Plecoptera" title="Plecoptera">Plecoptera (stoneflies)</a></li> <li><a href="/wiki/Earwig" title="Earwig">Dermaptera (earwigs)</a></li> <li><a href="/wiki/Embioptera" title="Embioptera">Embioptera (webspinners)</a></li> <li><a href="/wiki/Phasmatodea" title="Phasmatodea">Phasmatodea (stick and leaf insects)</a></li> <li><a href="/wiki/Orthoptera" title="Orthoptera">Orthoptera (crickets, wetas, grasshoppers, locusts)</a></li> <li><a href="/wiki/Zoraptera" title="Zoraptera">Zoraptera (angel insects)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Notoptera" title="Notoptera">Notoptera (Xenonomia)</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Grylloblattidae" title="Grylloblattidae">Grylloblattodea (ice-crawlers)</a></li> <li><a href="/wiki/Mantophasmatidae" title="Mantophasmatidae">Mantophasmatodea (gladiators)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Dictyoptera" title="Dictyoptera">Dictyoptera</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Blattodea" title="Blattodea">Blattodea (cockroaches, termites)</a></li> <li><a href="/wiki/Mantis" title="Mantis">Mantodea (mantises)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.0em;text-align:center;"><a href="/wiki/Eumetabola" title="Eumetabola">E u m e t a b o l a</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Paraneoptera_*" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Paraneoptera" title="Paraneoptera">Paraneoptera</a><abbr title="Eumetabola = (Psocodea + (rest of Paraneoptera + Holometabola))"> *</abbr></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Psocodea" title="Psocodea">Psocodea (barklice, lice)</a></li> <li><a href="/wiki/Thrips" title="Thrips">Thysanoptera (thrips)</a></li> <li><a href="/wiki/Hemiptera" title="Hemiptera">Hemiptera (cicadas, aphids, true bugs)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;line-height:1.0em;text-align:center;font-size:85%;"><a href="/wiki/Holometabola" title="Holometabola">H o l o m e t a b o l a</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hymenoptera" title="Hymenoptera">Hymenoptera (sawflies, wasps, ants, bees)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Neuropteroidea</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Coleopterida" title="Coleopterida">Coleopterida</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/Strepsiptera" title="Strepsiptera">Strepsiptera (twisted-winged parasites)</a></li> <li><a class="mw-selflink selflink">Coleoptera (beetles)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Neuropterida" title="Neuropterida">Neuropterida</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/Snakefly" title="Snakefly">Raphidioptera (snakeflies)</a></li> <li><a href="/wiki/Megaloptera" title="Megaloptera">Megaloptera (alderflies, dobsonflies, fishflies)</a></li> <li><a href="/wiki/Neuroptera" title="Neuroptera">Neuroptera (net-winged insects: lacewings, mantidflies, antlions)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Panorpida" title="Panorpida">Panorpida (Mecopterida)</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:10em;line-height:2em;">Antliophora</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em"> <ul><li><abbr title="A clade comprising paraphyletic order Mecoptera and monophyletic order Siphonaptera."><a href="/wiki/Mecoptera" title="Mecoptera">Mecoptera (scorpionflies)</a> + <a href="/wiki/Flea" title="Flea">Siphonaptera (fleas)</a></abbr></li> <li><a href="/wiki/Fly" title="Fly">Diptera (gnats, mosquitoes, flies)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em;line-height:2em;"><a href="/wiki/Amphiesmenoptera" title="Amphiesmenoptera">Amphiesmenoptera</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/Caddisfly" title="Caddisfly">Trichoptera (caddisflies)</a></li> <li><a href="/wiki/Lepidoptera" title="Lepidoptera">Lepidoptera (moths, butterflies)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="3" style="font-size:90%;"><div> <ul><li>Four most speciose orders are marked in bold</li> <li>Italic are paraphyletic groups</li> <li>Based on <a rel="nofollow" class="external text" href="http://www.biomedcentral.com/1471-2148/13/236">Sasaki et al. (2013)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Extinct" style="font-size:114%;margin:0 4em">Extinct</div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Aethiocarenodea" class="mw-redirect" title="Aethiocarenodea">Aethiocarenodea</a></li> <li><a href="/wiki/Roachoid" title="Roachoid">Blattoptera (roachoids)</a></li> <li><a href="/wiki/Caloneurodea" title="Caloneurodea">Caloneurodea</a></li> <li><a href="/wiki/Campylopteridae" class="mw-redirect" title="Campylopteridae">Campylopteridae</a></li> <li><a href="/w/index.php?title=Carbotriplurida&action=edit&redlink=1" class="new" title="Carbotriplurida (page does not exist)">Carbotriplurida</a></li> <li><a href="/wiki/Coxoplectoptera" title="Coxoplectoptera">Coxoplectoptera</a></li> <li><a href="/wiki/Diaphanopterodea" title="Diaphanopterodea">Diaphanopterodea</a></li> <li><a href="/w/index.php?title=Eoblattodea&action=edit&redlink=1" class="new" title="Eoblattodea (page does not exist)">Eoblattodea</a></li> <li><a href="/w/index.php?title=Eudiaphanoptera&action=edit&redlink=1" class="new" title="Eudiaphanoptera (page does not exist)">Eudiaphanoptera</a></li> <li><a href="/wiki/Geroptera" class="mw-redirect" title="Geroptera">Geroptera</a></li> <li><a href="/wiki/Glosselytrodea" title="Glosselytrodea">Glosselytrodea</a></li> <li><a href="/w/index.php?title=Heraridea&action=edit&redlink=1" class="new" title="Heraridea (page does not exist)">Heraridea</a></li> <li><a href="/w/index.php?title=Hypoperlida&action=edit&redlink=1" class="new" title="Hypoperlida (page does not exist)">Hypoperlida</a></li> <li><a href="/wiki/Lapeyriidae" class="mw-redirect" title="Lapeyriidae">Lapeyriidae</a></li> <li><a href="/wiki/Meganisoptera" title="Meganisoptera">Meganisoptera (griffinflies)</a></li> <li><a href="/wiki/Megasecoptera" title="Megasecoptera">Megasecoptera</a></li> <li><a href="/wiki/Miomoptera" title="Miomoptera">Miomoptera</a></li> <li><a href="/wiki/Palaeodictyoptera" title="Palaeodictyoptera">Palaeodictyoptera</a></li> <li><a href="/wiki/Paoliida" title="Paoliida">Paoliida</a></li> <li><a href="/w/index.php?title=Permoplectoptera&action=edit&redlink=1" class="new" title="Permoplectoptera (page does not exist)">Permoplectoptera</a></li> <li><a href="/wiki/Permothemistida" title="Permothemistida">Permothemistida</a></li> <li><a href="/wiki/Protanisoptera" class="mw-redirect" title="Protanisoptera">Protanisoptera</a></li> <li><a href="/wiki/Protelytroptera" title="Protelytroptera">Protelytroptera</a></li> <li><a href="/w/index.php?title=Protephemerida&action=edit&redlink=1" class="new" title="Protephemerida (page does not exist)">Protephemerida</a></li> <li><a href="/wiki/Protorthoptera" title="Protorthoptera">Protorthoptera</a></li> <li><a href="/wiki/Protozygoptera" class="mw-redirect" title="Protozygoptera">Protozygoptera</a></li> <li><a href="/w/index.php?title=Syntonoptera&action=edit&redlink=1" class="new" title="Syntonoptera (page does not exist)">Syntonoptera</a></li> <li><a href="/wiki/Titanoptera" title="Titanoptera">Titanoptera</a></li> <li><a href="/wiki/Triadophlebioptera" class="mw-redirect" title="Triadophlebioptera">Triadophlebioptera</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li><a href="/wiki/File:Beetle_icon.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Beetle_icon.svg/25px-Beetle_icon.svg.png" decoding="async" width="25" height="28" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Beetle_icon.svg/38px-Beetle_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Beetle_icon.svg/50px-Beetle_icon.svg.png 2x" data-file-width="143" data-file-height="159" /></a> <a href="/wiki/Portal:Insects" title="Portal:Insects">Insects portal</a></li> <li><a href="https://species.wikimedia.org/wiki/Insecta" class="extiw" title="species:Insecta">Wikispecies</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="Extant_Coleoptera_families" style="padding:3px"><table class="nowraplinks hlist mw-collapsible mw-collapsed 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:Coleoptera" title="Template:Coleoptera"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Coleoptera" title="Template talk:Coleoptera"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Coleoptera" title="Special:EditPage/Template:Coleoptera"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Extant_Coleoptera_families" style="font-size:114%;margin:0 4em">Extant <a class="mw-selflink selflink">Coleoptera</a> families</div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div> <ul><li>Kingdom: <a href="/wiki/Animal" title="Animal">Animalia</a></li> <li>Phylum: <a href="/wiki/Arthropod" title="Arthropod">Arthropoda</a></li> <li>Class: <a href="/wiki/Insect" title="Insect">Insecta</a></li> <li>Subclass: <a href="/wiki/Pterygota" title="Pterygota">Pterygota</a></li> <li>Infraclass: <a href="/wiki/Neoptera" title="Neoptera">Neoptera</a></li> <li>Superorder: <a href="/wiki/Holometabola" title="Holometabola">Holometabola</a></li></ul> </div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Suborder_Archostemata" style="font-size:114%;margin:0 4em">Suborder <a href="/wiki/Archostemata" title="Archostemata">Archostemata</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Crowsoniella" title="Crowsoniella">Crowsoniellidae (Crowsoniella relicta)</a></li> <li><a href="/wiki/Cupedidae" title="Cupedidae">Cupedidae (reticulated beetles)</a></li> <li><a href="/wiki/Jurodidae" title="Jurodidae">Jurodidae (Sikhotealinia zhiltzovae)</a></li> <li><a href="/wiki/Micromalthidae" title="Micromalthidae">Micromalthidae</a></li> <li><a href="/wiki/Ommatidae" title="Ommatidae">Ommatidae</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Suborder_Adephaga" style="font-size:114%;margin:0 4em">Suborder <a href="/wiki/Adephaga" title="Adephaga">Adephaga</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th id="Extant_families" scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Extant_taxon" class="mw-redirect" title="Extant taxon">Extant</a> families</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Amphizoa" title="Amphizoa">Amphizoidae (trout-stream beetles)</a></li> <li><a href="/wiki/Aspidytidae" title="Aspidytidae">Aspidytidae</a></li> <li><a href="/wiki/Ground_beetle" title="Ground beetle">Carabidae (ground beetles)</a></li> <li><a href="/wiki/Tiger_beetle" title="Tiger beetle">Cicindelidae (tiger beetles)</a></li> <li><a href="/wiki/Dytiscidae" title="Dytiscidae">Dytiscidae (predaceous diving beetles)</a></li> <li><a href="/wiki/Whirligig_beetle" title="Whirligig beetle">Gyrinidae (whirligig beetles)</a></li> <li><a href="/wiki/Haliplidae" title="Haliplidae">Haliplidae (crawling water beetles)</a></li> <li><a href="/wiki/Hygrobia" title="Hygrobia">Hygrobiidae</a></li> <li><a href="/wiki/Meru_phyllisae" title="Meru phyllisae">Meruidae (Meru phyllisae)</a></li> <li><a href="/wiki/Noteridae" title="Noteridae">Noteridae (burrowing water beetles)</a></li> <li><a href="/wiki/Trachypachidae" title="Trachypachidae">Trachypachidae (false ground beetles)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Suborder_Myxophaga" style="font-size:114%;margin:0 4em">Suborder <a href="/wiki/Myxophaga" title="Myxophaga">Myxophaga</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Hydroscaphidae" title="Hydroscaphidae">Hydroscaphidae (skiff beetles)</a></li> <li><a href="/wiki/Lepicerus" title="Lepicerus">Lepiceridae</a></li> <li><a href="/wiki/Sphaerius" title="Sphaerius">Sphaeriusidae</a></li> <li><a href="/wiki/Torridincolidae" title="Torridincolidae">Torridincolidae</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks mw-collapsible mw-collapsed navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Suborder_Polyphaga" style="font-size:114%;margin:0 4em">Suborder <a href="/wiki/Polyphaga" title="Polyphaga">Polyphaga</a></div></th></tr><tr><td colspan="2" class="navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Bostrichiformia" title="Bostrichiformia">Bostrichiformia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Bostrichoidea" title="Bostrichoidea">Bostrichoidea</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/Bostrichidae" title="Bostrichidae">Bostrichidae (auger beetles)</a></li> <li><a href="/wiki/Dermestidae" title="Dermestidae">Dermestidae (skin beetles)</a></li> <li><a href="/wiki/Endecatomus" title="Endecatomus">Endecatomidae</a></li> <li><a href="/wiki/Jacobsoniidae" title="Jacobsoniidae">Jacobsoniidae (Jacobson's beetles)</a></li> <li><a href="/wiki/Nosodendridae" title="Nosodendridae">Nosodendridae (wounded-tree beetles)</a></li> <li><a href="/wiki/Ptinidae" title="Ptinidae">Ptiniidae (furniture beetles, death watch beetles, spider beetles)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Derodontidae" title="Derodontidae">Derodontoidea</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/Derodontidae" title="Derodontidae">Derodontidae (tooth-necked fungus beetles)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Cucujiformia" title="Cucujiformia">Cucujiformia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Chrysomeloidea" title="Chrysomeloidea">Chrysomeloidea</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/Longhorn_beetle" title="Longhorn beetle">Cerambycidae (longhorn beetles)</a></li> <li><a href="/wiki/Leaf_beetle" title="Leaf beetle">Chrysomelidae (leaf beetles)</a></li> <li><a href="/wiki/Disteniidae" title="Disteniidae">Disteniidae</a></li> <li><a href="/wiki/Megalopodidae" title="Megalopodidae">Megalopodidae</a></li> <li><a href="/wiki/Orsodacnidae" title="Orsodacnidae">Orsodacnidae</a></li> <li><a href="/wiki/Oxypeltidae" title="Oxypeltidae">Oxypeltidae</a></li> <li><a href="/wiki/Vesperidae" title="Vesperidae">Vesperidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Cleroidea" title="Cleroidea">Cleroidea</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/Acanthocnemus" title="Acanthocnemus">Acanthocnemidae (Acanthocnemus nigricans)</a></li> <li><a href="/wiki/Biphyllidae" title="Biphyllidae">Biphyllidae (false skin beetles)</a></li> <li><a href="/wiki/Byturidae" title="Byturidae">Byturidae (fruitworm beetles)</a></li> <li><a href="/wiki/Chaetosomatidae" title="Chaetosomatidae">Chaetosomatidae</a></li> <li><a href="/wiki/Cleridae" title="Cleridae">Cleridae (checkered beetles)</a></li> <li><a href="/wiki/Lophocateridae" title="Lophocateridae">Lophocateridae</a></li> <li><a href="/wiki/Mauroniscidae" title="Mauroniscidae">Mauroniscidae</a></li> <li><a href="/wiki/Melyridae" title="Melyridae">Melyridae (soft-wing flower beetles)</a></li> <li><a href="/wiki/Metaxina" title="Metaxina">Metaxinidae (Metaxina ornata)</a></li> <li><a href="/wiki/Peltis" title="Peltis">Peltidae</a></li> <li><a href="/wiki/Phloiophilus" title="Phloiophilus">Phloiophilidae (Phloiophilus edwardsi)</a></li> <li><a href="/wiki/Phycosecidae" title="Phycosecidae">Phycosecidae</a></li> <li><a href="/wiki/Prionoceridae" title="Prionoceridae">Prionoceridae</a></li> <li><a href="/wiki/Protopeltis" title="Protopeltis">Protopeltidae</a></li> <li><a href="/wiki/Rentoniidae" title="Rentoniidae">Rentoniidae</a></li> <li><a href="/wiki/Rhadalidae" title="Rhadalidae">Rhadalidae</a></li> <li><a href="/wiki/Thanerocleridae" title="Thanerocleridae">Thanerocleridae</a></li> <li><a href="/wiki/Thymalidae" title="Thymalidae">Thymalidae</a></li> <li><a href="/wiki/Trogossitidae" title="Trogossitidae">Trogossitidae (bark-gnawing beetles)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Coccinelloidea" title="Coccinelloidea">Coccinelloidea</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/Akalyptoischion" title="Akalyptoischion">Akalyptoischiidae</a></li> <li><a href="/wiki/Alexiidae" title="Alexiidae">Alexiidae</a></li> <li><a href="/wiki/Anamorphidae" title="Anamorphidae">Anamorphidae</a></li> <li><a href="/wiki/Bothrideridae" title="Bothrideridae">Bothrideridae (dry bark beetles)</a></li> <li><a href="/wiki/Cerylonidae" title="Cerylonidae">Cerylonidae (minute bark beetles)</a></li> <li><a href="/wiki/Coccinellidae" title="Coccinellidae">Coccinellidae (lady beetles, or God's cows)</a></li> <li><a href="/wiki/Corylophidae" title="Corylophidae">Corylophidae (minute fungus beetles)</a></li> <li><a href="/wiki/Discolomatidae" title="Discolomatidae">Discolomatidae</a></li> <li><a href="/wiki/Endomychidae" title="Endomychidae">Endomychidae (handsome fungus beetles)</a></li> <li><a href="/wiki/Eupsilobiidae" title="Eupsilobiidae">Eupsilobiidae</a></li> <li><a href="/wiki/Euxestidae" title="Euxestidae">Euxestidae (well polished beetles)</a></li> <li><a href="/wiki/Latridiidae" title="Latridiidae">Latridiidae (minute brown scavenger beetles)</a></li> <li><a href="/wiki/Murmidiidae" title="Murmidiidae">Murmidiidae</a></li> <li><a href="/wiki/Mycetaeidae" title="Mycetaeidae">Mycetaeidae</a></li> <li><a href="/wiki/Teredidae" title="Teredidae">Teredidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Cucujoidea" title="Cucujoidea">Cucujoidea</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/Agapytho" title="Agapytho">Agapythidae (Agapytho foveicollis)</a></li> <li><a href="/wiki/Boganiidae" title="Boganiidae">Boganiidae</a></li> <li><a href="/wiki/Cavognathidae" title="Cavognathidae">Cavognathidae</a></li> <li><a href="/wiki/Cryptophagidae" title="Cryptophagidae">Cryptophagidae (silken fungus beetles)</a></li> <li><a href="/wiki/Cucujidae" title="Cucujidae">Cucujidae (flat bark beetles)</a></li> <li><a href="/wiki/Cybocephalidae" title="Cybocephalidae">Cybocephalidae</a></li> <li><a href="/wiki/Cyclaxyridae" title="Cyclaxyridae">Cyclaxyridae</a></li> <li><a href="/wiki/Erotylidae" title="Erotylidae">Erotylidae (pleasing fungus beetles)</a></li> <li><a href="/wiki/Helotidae" title="Helotidae">Helotidae</a></li> <li><a href="/wiki/Hobartiidae" title="Hobartiidae">Hobartiidae</a></li> <li><a href="/wiki/Kateretidae" title="Kateretidae">Kateretidae (short-winged flower beetles)</a></li> <li><a href="/wiki/Laemophloeidae" title="Laemophloeidae">Laemophloeidae (lined flat bark beetles)</a></li> <li><a href="/wiki/Lamingtonium" title="Lamingtonium">Lamingtoniidae (Lamingtonium binnaberrense)</a></li> <li><a href="/wiki/Monotomidae" title="Monotomidae">Monotomidae (root-eating beetles)</a></li> <li><a href="/wiki/Myraboliidae" class="mw-redirect" title="Myraboliidae">Myraboliidae</a></li> <li><a href="/wiki/Sap_beetle" title="Sap beetle">Nitidulidae (sap beetles)</a></li> <li><a href="/wiki/Passandridae" title="Passandridae">Passandridae (parasitic flat bark beetles)</a></li> <li><a href="/wiki/Phalacridae" title="Phalacridae">Phalacridae (shining flower beetles)</a></li> <li><a href="/wiki/Phloeostichidae" title="Phloeostichidae">Phloeostichidae</a></li> <li><a href="/wiki/Priasilphidae" title="Priasilphidae">Priasilphidae</a></li> <li><a href="/wiki/Propalticidae" title="Propalticidae">Propalticidae</a></li> <li><a href="/wiki/Protocucujidae" title="Protocucujidae">Protocucujidae</a></li> <li><a href="/wiki/Silvanidae" title="Silvanidae">Silvanidae (silvanid flat bark beetles)</a></li> <li><a href="/wiki/Smicripidae" title="Smicripidae">Smicripidae (palmetto beetles)</a></li> <li><a href="/wiki/Sphindidae" title="Sphindidae">Sphindidae (dry-fungus beetles)</a></li> <li><a href="/wiki/Tasmosalpingus" title="Tasmosalpingus">Tasmosalpingidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Weevil" title="Weevil">Curculionoidea</a> (weevils)</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/Anthribidae" title="Anthribidae">Anthribidae (fungus weevils)</a></li> <li><a href="/wiki/Attelabidae" title="Attelabidae">Attelabidae (leaf-rolling weevils)</a></li> <li><a href="/wiki/Belidae" title="Belidae">Belidae (primitive weevils)</a></li> <li><a href="/wiki/Brentidae" title="Brentidae">Brentidae (straight snout weevils, New York weevil)</a></li> <li><a href="/wiki/Caridae" title="Caridae">Caridae</a></li> <li><a href="/wiki/Curculionidae" title="Curculionidae">Curculionidae (true weevils, bark beetles, ambrosia beetles)</a></li> <li><a href="/wiki/Nemonychidae" title="Nemonychidae">Nemonychidae (pine flower weevils)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Lymexylidae" title="Lymexylidae">Lymexyloidea</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/Lymexylidae" title="Lymexylidae">Lymexylidae (ship-timber beetles)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Tenebrionoidea" title="Tenebrionoidea">Tenebrionoidea</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/Aderidae" title="Aderidae">Aderidae (ant-like leaf beetles)</a></li> <li><a href="/wiki/Anthicidae" title="Anthicidae">Anthicidae (ant-like flower beetles)</a></li> <li><a href="/wiki/Archeocrypticidae" title="Archeocrypticidae">Archeocrypticidae (cryptic fungus beetles)</a></li> <li><a href="/wiki/Boridae" title="Boridae">Boridae (conifer bark beetles)</a></li> <li><a href="/wiki/Chalcodryidae" title="Chalcodryidae">Chalcodryidae</a></li> <li><a href="/wiki/Ciidae" title="Ciidae">Ciidae (minute tree-fungus beetles)</a></li> <li><a href="/wiki/Melandryidae" title="Melandryidae">Melandryidae (false darkling beetles)</a></li> <li><a href="/wiki/Blister_beetle" title="Blister beetle">Meloidae (blister beetles)</a></li> <li><a href="/wiki/Mordellidae" title="Mordellidae">Mordellidae (tumbling flower beetles)</a></li> <li><a href="/wiki/Mycetophagidae" title="Mycetophagidae">Mycetophagidae (hairy fungus beetles)</a></li> <li><a href="/wiki/Mycteridae" title="Mycteridae">Mycteridae (palm and flower beetles)</a></li> <li><a href="/wiki/Oedemeridae" title="Oedemeridae">Oedemeridae (false blister beetle)</a></li> <li><a href="/wiki/Perimylopidae" class="mw-redirect" title="Perimylopidae">Perimylopidae, or Promecheilidae</a></li> <li><a href="/wiki/Prostomidae" title="Prostomidae">Prostomidae (jugular-horned beetles)</a></li> <li><a href="/wiki/Pterogeniidae" title="Pterogeniidae">Pterogeniidae</a></li> <li><a href="/wiki/Fire-coloured_beetle" title="Fire-coloured beetle">Pyrochroidae (fire-coloured beetles)</a></li> <li><a href="/wiki/Pythidae" title="Pythidae">Pythidae (dead log bark beetles)</a></li> <li><a href="/wiki/Ripiphoridae" title="Ripiphoridae">Ripiphoridae (wedge-shaped beetles)</a></li> <li><a href="/wiki/Salpingidae" title="Salpingidae">Salpingidae (narrow-waisted bark beetles)</a></li> <li><a href="/wiki/Scraptiidae" title="Scraptiidae">Scraptiidae (false flower beetles)</a></li> <li><a href="/wiki/Stenotrachelidae" title="Stenotrachelidae">Stenotrachelidae (false longhorn beetles)</a></li> <li><a href="/wiki/Synchroidae" title="Synchroidae">Synchroidae (synchroa bark beetles)</a></li> <li><a href="/wiki/Darkling_beetle" title="Darkling beetle">Tenebrionidae (darkling beetles)</a></li> <li><a href="/wiki/Tetratomidae" title="Tetratomidae">Tetratomidae (polypore fungus beetles)</a></li> <li><a href="/wiki/Trictenotomidae" title="Trictenotomidae">Trictenotomidae</a></li> <li><a href="/wiki/Ulodidae" title="Ulodidae">Ulodidae</a></li> <li><a href="/wiki/Zopheridae" title="Zopheridae">Zopheridae (ironclad beetles, cylindrical bark beetles)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Elateriformia" title="Elateriformia">Elateriformia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Buprestoidea" title="Buprestoidea">Buprestoidea</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/Buprestidae" title="Buprestidae">Buprestidae (jewel beetles, or metallic wood-boring beetles)</a></li> <li><a href="/wiki/Schizopodidae" title="Schizopodidae">Schizopodidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Byrrhoidea" title="Byrrhoidea">Byrrhoidea</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/Byrrhidae" title="Byrrhidae">Byrrhidae (pill beetles)</a></li> <li><a href="/wiki/Callirhipidae" title="Callirhipidae">Callirhipidae (cedar beetles)</a></li> <li><a href="/wiki/Chelonariidae" title="Chelonariidae">Chelonariidae (turtle beetles)</a></li> <li><a href="/wiki/Cneoglossidae" title="Cneoglossidae">Cneoglossidae</a></li> <li><a href="/wiki/Dryopidae" title="Dryopidae">Dryopidae (long-toed water beetles)</a></li> <li><a href="/wiki/Elmidae" title="Elmidae">Elmidae (riffle beetles)</a></li> <li><a href="/wiki/Eulichadidae" title="Eulichadidae">Eulichadidae (forest stream beetles)</a></li> <li><a href="/wiki/Heteroceridae" title="Heteroceridae">Heteroceridae (variegated mud-loving beetles)</a></li> <li><a href="/wiki/Limnichidae" title="Limnichidae">Limnichidae (minute mud beetles)</a></li> <li><a href="/wiki/Lutrochidae" class="mw-redirect" title="Lutrochidae">Lutrochidae (travertine beetles)</a></li> <li><a href="/wiki/Water-penny_beetles" class="mw-redirect" title="Water-penny beetles">Psephenidae (water-penny beetles)</a></li> <li><a href="/wiki/Ptilodactylidae" title="Ptilodactylidae">Ptilodactylidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Dascilloidea" title="Dascilloidea">Dascilloidea</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/Dascillidae" title="Dascillidae">Dascillidae (soft bodied plant beetles)</a></li> <li><a href="/wiki/Rhipiceridae" title="Rhipiceridae">Rhipiceridae (cicada beetle, cicada parasite beetles)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Elateroidea" title="Elateroidea">Elateroidea</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/Artematopodidae" title="Artematopodidae">Artematopodidae (soft-bodied plant beetles)</a></li> <li><a href="/wiki/Brachypsectridae" title="Brachypsectridae">Brachypsectridae (Texas beetles)</a></li> <li><a href="/wiki/Soldier_beetle" title="Soldier beetle">Cantharidae (soldier beetles)</a></li> <li><a href="/wiki/Cerophytidae" title="Cerophytidae">Cerophytidae (rare click beetles)</a></li> <li><a href="/wiki/Click_beetle" title="Click beetle">Elateridae (click beetles)</a></li> <li><a href="/wiki/Eucnemidae" title="Eucnemidae">Eucnemidae (false click beetles)</a></li> <li><a href="/wiki/Jurasaidae" title="Jurasaidae">Jurasaidae</a></li> <li><a href="/wiki/Firefly" title="Firefly">Lampyridae (fireflies)</a></li> <li><a href="/wiki/Lycidae" title="Lycidae">Lycidae (net-winged beetles)</a></li> <li><a href="/wiki/Omethidae" title="Omethidae">Omethidae (false fireflies, long-lipped beetles)</a></li> <li><a href="/wiki/Phengodidae" title="Phengodidae">Phengodidae (glowworm beetles)</a></li> <li><a href="/wiki/Rhagophthalmidae" title="Rhagophthalmidae">Rhagophthalmidae</a></li> <li><a href="/wiki/Sinopyrophoridae" class="mw-redirect" title="Sinopyrophoridae">Sinopyrophoridae</a></li> <li><a href="/wiki/Throscidae" title="Throscidae">Throscidae (false metallic wood-boring beetles)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Rhinorhipus" title="Rhinorhipus">Rhinorhipoidea</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/Rhinorhipus" title="Rhinorhipus">Rhinorhipidae (Rhinorhipus tamborinensis)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Scirtoidea" title="Scirtoidea">Scirtoidea</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/Clambidae" title="Clambidae">Clambidae</a></li> <li><a href="/wiki/Declinia" title="Declinia">Decliniidae (Declinia relicta)</a></li> <li><a href="/wiki/Eucinetidae" title="Eucinetidae">Eucinetidae (plate-thigh beetles)</a></li> <li><a href="/wiki/Scirtidae" title="Scirtidae">Scirtidae</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Scarabaeoidea" title="Scarabaeoidea">Scarabaeiformia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Scarabaeoidea" title="Scarabaeoidea">Scarabaeoidea</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/Belohina" title="Belohina">Belohinidae (Belohina inexpectata)</a></li> <li><a href="/wiki/Bolboceratidae" title="Bolboceratidae">Bolboceratidae</a></li> <li><a href="/wiki/False_stag_beetle" title="False stag beetle">Diphyllostomatidae (false stag beetles)</a></li> <li><a href="/wiki/Geotrupidae" title="Geotrupidae">Geotrupidae (dor beetles)</a></li> <li><a href="/wiki/Glaphyridae" title="Glaphyridae">Glaphyridae (bumble bee scarab beetles)</a></li> <li><a href="/wiki/Glaresis" title="Glaresis">Glaresidae (enigmatic scarab beetles)</a></li> <li><a href="/wiki/Hybosoridae" title="Hybosoridae">Hybosoridae (scavenger scarab beetles)</a></li> <li><a href="/wiki/Stag_beetle" title="Stag beetle">Lucanidae (stag beetles)</a></li> <li><a href="/wiki/Ochodaeidae" title="Ochodaeidae">Ochodaeidae (sand-loving scarab beetles)</a></li> <li><a href="/wiki/Passalidae" title="Passalidae">Passalidae (betsy beetles)</a></li> <li><a href="/wiki/Rain_beetle" title="Rain beetle">Pleocomidae (rain beetles)</a></li> <li><a href="/wiki/Scarabaeidae" title="Scarabaeidae">Scarabaeidae (scarabs)</a></li> <li><a href="/wiki/Trogidae" title="Trogidae">Trogidae (hide beetles)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Staphyliniformia" title="Staphyliniformia">Staphyliniformia</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Histeroidea" title="Histeroidea">Histeroidea</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/Histeridae" title="Histeridae">Histeridae (clown beetles)</a></li> <li><a href="/wiki/Sphaerites" title="Sphaerites">Sphaeritidae (false clown beetles)</a></li> <li><a href="/wiki/Syntelia" title="Syntelia">Synteliidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Hydrophiloidea" title="Hydrophiloidea">Hydrophiloidea</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/Epimetopidae" title="Epimetopidae">Epimetopidae</a></li> <li><a href="/wiki/Georissus" title="Georissus">Georissidae (minute mud-loving beetles)</a></li> <li><a href="/wiki/Helophorus" title="Helophorus">Helophoridae</a></li> <li><a href="/wiki/Hydrochidae" class="mw-redirect" title="Hydrochidae">Hydrochidae</a></li> <li><a href="/wiki/Hydrophilidae" title="Hydrophilidae">Hydrophilidae (water scavenger beetles)</a></li> <li><a href="/wiki/Spercheidae" class="mw-redirect" title="Spercheidae">Spercheidae</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:10em"><a href="/wiki/Staphylinoidea" title="Staphylinoidea">Staphylinoidea</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/Agyrtidae" title="Agyrtidae">Agyrtidae (primitive carrion beetles)</a></li> <li><a href="/wiki/Hydraenidae" title="Hydraenidae">Hydraenidae</a></li> <li><a href="/wiki/Leiodidae" title="Leiodidae">Leiodidae (round fungus beetles)</a></li> <li><a href="/wiki/Ptiliidae" title="Ptiliidae">Ptiliidae (feather-winged beetles)</a></li> <li><a href="/wiki/Silphidae" title="Silphidae">Silphidae (carrion beetles)</a></li> <li><a href="/wiki/Rove_beetle" title="Rove beetle">Staphylinidae (rove beetles)</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="/wiki/List_of_subgroups_of_the_order_Coleoptera" title="List of subgroups of the order Coleoptera">List of subgroups of the order Coleoptera</a></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="Human_interactions_with_insects" 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:Insects_in_culture" title="Template:Insects in culture"><abbr title="View this template">v</abbr></a></li><li class="nv-talk"><a href="/wiki/Template_talk:Insects_in_culture" title="Template talk:Insects in culture"><abbr title="Discuss this template">t</abbr></a></li><li class="nv-edit"><a href="/wiki/Special:EditPage/Template:Insects_in_culture" title="Special:EditPage/Template:Insects in culture"><abbr title="Edit this template">e</abbr></a></li></ul></div><div id="Human_interactions_with_insects" style="font-size:114%;margin:0 4em"><a href="/wiki/Human_interactions_with_insects" title="Human interactions with insects">Human interactions with insects</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Aspects of <a href="/wiki/Insect" title="Insect">insects</a> in culture</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">In the arts</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Insects_in_art" title="Insects in art">Insects in art</a> <ul><li><a href="/wiki/Beetlewing" title="Beetlewing">Beetlewing</a></li> <li><a href="/wiki/Musca_depicta" title="Musca depicta">Musca depicta</a></li></ul></li> <li><a href="/wiki/Arthropods_in_film" title="Arthropods in film">Arthropods in film</a></li> <li><a href="/wiki/Insects_in_literature" title="Insects in literature">Insects in literature</a></li> <li><a href="/wiki/Insects_in_music" title="Insects in music">Insects in music</a> <ul><li><a href="/wiki/List_of_insect-inspired_songs" title="List of insect-inspired songs">List of insect-inspired songs</a></li></ul></li> <li><a href="/wiki/Insects_on_stamps" title="Insects on stamps">Insects on stamps</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">In fishing</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Fishing_bait" title="Fishing bait">Fishing bait</a></li> <li><a href="/wiki/Fly_fishing" title="Fly fishing">Fly fishing</a></li> <li><a href="/wiki/Artificial_fly" title="Artificial fly">Artificial fly</a></li> <li><a href="/wiki/Fly_tying" title="Fly tying">Fly tying</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Insects_in_medicine" title="Insects in medicine">In medicine</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Apitherapy" title="Apitherapy">Apitherapy</a> <ul><li><a href="/wiki/Apitoxin" title="Apitoxin">Apitoxin</a></li> <li><a href="/wiki/Melittin" title="Melittin">Melittin</a></li></ul></li> <li><a href="/wiki/Lytta_vesicatoria" title="Lytta vesicatoria">Spanish fly</a> <ul><li><a href="/wiki/Cantharidin" title="Cantharidin">Cantharidin</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Insects_in_mythology" title="Insects in mythology">In mythology</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bees_in_mythology" title="Bees in mythology">Bees in mythology</a></li> <li><a href="/wiki/Cicadas_in_mythology" title="Cicadas in mythology">Cicadas in mythology</a></li> <li><a href="/wiki/Scarab_(artifact)" title="Scarab (artifact)">Scarab (artifact)</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Biomimetics" title="Biomimetics">Biomimicry</a></li> <li><a href="/wiki/Cockroach_racing" title="Cockroach racing">Cockroach racing</a></li> <li><a href="/wiki/Cockroaches_in_popular_culture" title="Cockroaches in popular culture">Cockroaches in popular culture</a></li> <li><a href="/wiki/Cricket_fighting" title="Cricket fighting">Cricket fighting</a></li> <li><a href="/wiki/Entomological_warfare" title="Entomological warfare">Entomological warfare</a></li> <li><a href="/wiki/Entomophagy_in_humans" title="Entomophagy in humans">Entomophagy</a></li> <li><a href="/wiki/Insect_farming" title="Insect farming">Insect farming</a></li> <li><a href="/wiki/Flea_circus" title="Flea circus">Flea circus</a></li> <li><a href="/wiki/Insects_in_ethics" title="Insects in ethics">Insects in ethics</a></li> <li><a href="/wiki/Insects_in_religion" title="Insects in religion">Insects in religion</a></li> <li><a href="/wiki/Jingzhe" title="Jingzhe">Jingzhe</a></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Economic_entomology" title="Economic entomology">Economic entomology</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Beneficial_insect" title="Beneficial insect">Beneficial insect</a></li> <li><a href="/wiki/Biological_pest_control" title="Biological pest control">Biological pest control</a></li> <li><a href="/wiki/Beekeeping" title="Beekeeping">Beekeeping</a> <ul><li><a href="/wiki/Bee_pollen" title="Bee pollen">Bee pollen</a></li> <li><a href="/wiki/List_of_crop_plants_pollinated_by_bees" title="List of crop plants pollinated by bees">List of crop plants pollinated by bees</a></li> <li><a href="/wiki/Beeswax" title="Beeswax">Beeswax</a></li> <li><a href="/wiki/Honey" title="Honey">Honey</a></li> <li><a href="/wiki/Propolis" title="Propolis">Propolis</a></li> <li><a href="/wiki/Royal_jelly" title="Royal jelly">Royal jelly</a></li></ul></li> <li><a href="/wiki/Carmine" title="Carmine">Carmine/Cochineal</a> <ul><li><a href="/wiki/Polish_cochineal" title="Polish cochineal">Polish</a></li></ul></li> <li><a href="/wiki/Chitin" title="Chitin">Chitin</a></li> <li><a href="/wiki/Kermes_(dye)" title="Kermes (dye)">Kermes</a></li> <li><a href="/wiki/Sericulture" title="Sericulture">Sericulture</a> <ul><li><a href="/wiki/Silk" title="Silk">Silk</a></li></ul></li> <li><a href="/wiki/Shellac" title="Shellac">Shellac</a></li> <li><a href="/wiki/Model_organism" title="Model organism">Model organism</a> <ul><li><a href="/wiki/Drosophila_melanogaster" title="Drosophila melanogaster">Drosophila melanogaster</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="/wiki/Pest_(organism)#Insects" title="Pest (organism)">Harmful insects</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Insect_bites_and_stings" class="mw-redirect" title="Insect bites and stings">Insect bites and stings</a></li> <li><a href="/wiki/Insect_sting_allergy" title="Insect sting allergy">Insect sting allergy</a></li> <li><a href="/wiki/Bed_bug" title="Bed bug">Bed bug</a></li> <li><a href="/wiki/Woodworm" title="Woodworm">Woodworm</a></li> <li><a href="/wiki/Home-stored_product_entomology" title="Home-stored product entomology">Home-stored product entomology</a></li> <li><a href="/wiki/Clothes_moth" title="Clothes moth">Clothes moth</a></li></ul> </div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Pioneers</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Jan_Swammerdam" title="Jan Swammerdam">Jan Swammerdam</a></li> <li><a href="/wiki/Alfred_Russel_Wallace" title="Alfred Russel Wallace">Alfred Russel Wallace</a></li> <li><a href="/wiki/Jean-Henri_Fabre" title="Jean-Henri Fabre">Jean-Henri Fabre</a></li> <li><a href="/wiki/Hans_Zinsser" title="Hans Zinsser">Hans Zinsser</a> (<a href="/wiki/Rats,_Lice_and_History" title="Rats, Lice and History">Rats, Lice and History</a>)</li> <li><a href="/wiki/Lafcadio_Hearn" title="Lafcadio Hearn">Lafcadio Hearn</a> (Insect Literature)</li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Concerns</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Bees_and_toxic_chemicals" title="Bees and toxic chemicals">Bees and toxic chemicals</a></li> <li><a href="/wiki/Colony_collapse_disorder" title="Colony collapse disorder">Colony collapse disorder</a></li> <li><a href="/wiki/Decline_in_insect_populations" title="Decline in insect populations">Decline in insect populations</a></li> <li><a href="/wiki/Habitat_destruction" title="Habitat destruction">Habitat destruction</a></li> <li><a href="/wiki/List_of_endangered_insects" title="List of endangered insects">List of endangered insects</a></li> <li><a href="/wiki/Pesticide" title="Pesticide">Pesticide</a> <ul><li><a href="/wiki/Insecticide" title="Insecticide">Insecticide</a></li> <li><a href="/wiki/Neonicotinoid" title="Neonicotinoid">Neonicotinoid</a></li> <li><a href="/wiki/Pesticide_toxicity_to_bees" title="Pesticide toxicity to bees">Pesticide toxicity to bees</a></li></ul></li></ul> </div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Categories, templates</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Category:Insects_and_humans" title="Category:Insects and humans">Insects and humans</a></li> <li><a href="/wiki/Template:Insecticides" title="Template:Insecticides">Insecticides</a></li> <li><a href="/wiki/Template:Pesticides" title="Template:Pesticides">Pesticides</a></li></ul> </div></td></tr><tr><td class="navbox-abovebelow" colspan="2" style="font-weight:bold;"><div><a href="/wiki/File:Beetle_icon.svg" class="mw-file-description"><img alt="icon" src="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Beetle_icon.svg/25px-Beetle_icon.svg.png" decoding="async" width="25" height="28" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Beetle_icon.svg/38px-Beetle_icon.svg.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/6c/Beetle_icon.svg/50px-Beetle_icon.svg.png 2x" data-file-width="143" data-file-height="159" /></a> <a href="/wiki/Portal:Insects" title="Portal:Insects">Insects portal</a></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="Taxon_identifiers" style="padding:3px"><table class="nowraplinks hlist navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Taxon_identifiers" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Taxon_identifiers" title="Help:Taxon identifiers">Taxon identifiers</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: left;">Coleoptera</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"> <ul><li><a href="/wiki/Wikidata" title="Wikidata">Wikidata</a>: <a href="https://www.wikidata.org/wiki/Q22671" class="extiw" title="wikidata:Q22671">Q22671</a></li> <li><a href="/wiki/Wikispecies" title="Wikispecies">Wikispecies</a>: <a href="https://species.wikimedia.org/wiki/Coleoptera" class="extiw" title="wikispecies:Coleoptera">Coleoptera</a></li> <li><a href="/wiki/Animal_Diversity_Web" title="Animal Diversity Web">ADW</a>: <a rel="nofollow" class="external text" href="https://animaldiversity.org/accounts/Coleoptera/">Coleoptera</a></li> <li><a href="/wiki/Australian_Faunal_Directory" title="Australian Faunal Directory">AFD</a>: <a rel="nofollow" class="external text" href="https://biodiversity.org.au/afd/taxa/Coleoptera">Coleoptera</a></li> <li><a href="/wiki/Barcode_of_Life_Data_System" title="Barcode of Life Data System">BOLD</a>: <a rel="nofollow" class="external text" href="http://www.boldsystems.org/index.php/TaxBrowser_TaxonPage?taxid=413">413</a></li> <li><a href="/wiki/BugGuide" title="BugGuide">BugGuide</a>: <a rel="nofollow" class="external text" href="https://bugguide.net/node/view/60">60</a></li> <li><a href="/wiki/Catalogue_of_Life" title="Catalogue of Life">CoL</a>: <a rel="nofollow" class="external text" href="https://www.catalogueoflife.org/data/taxon/C2L">C2L</a></li> <li><a href="/wiki/Encyclopedia_of_Life" title="Encyclopedia of Life">EoL</a>: <a rel="nofollow" class="external text" href="https://eol.org/pages/345">345</a></li> <li><a href="/wiki/EPPO_Code" title="EPPO Code">EPPO</a>: <a rel="nofollow" class="external text" href="https://gd.eppo.int/taxon/1COLEO">1COLEO</a></li> <li><a href="/wiki/Fauna_Europaea" title="Fauna Europaea">Fauna Europaea</a>: <a rel="nofollow" class="external text" href="http://www.eu-nomen.eu/portal/taxon.php?GUID=urn:lsid:faunaeur.org:taxname:10993">10993</a></li> <li><a href="/wiki/Fauna_Europaea" title="Fauna Europaea">Fauna Europaea (new)</a>: <a rel="nofollow" class="external text" href="https://fauna-eu.org/cdm_dataportal/taxon/3cf53a59-439c-4e58-90e0-88fd23e92e49">3cf53a59-439c-4e58-90e0-88fd23e92e49</a></li> <li><a href="/wiki/Global_Biodiversity_Information_Facility" title="Global Biodiversity Information Facility">GBIF</a>: <a rel="nofollow" class="external text" href="https://www.gbif.org/species/1470">1470</a></li> <li><a href="/wiki/INaturalist" title="INaturalist">iNaturalist</a>: <a rel="nofollow" class="external text" href="https://inaturalist.org/taxa/47208">47208</a></li> <li><a href="/wiki/Interim_Register_of_Marine_and_Nonmarine_Genera" title="Interim Register of Marine and Nonmarine Genera">IRMNG</a>: <a rel="nofollow" class="external text" href="https://www.irmng.org/aphia.php?p=taxdetails&id=10279">10279</a></li> <li><a href="/wiki/Integrated_Taxonomic_Information_System" title="Integrated Taxonomic Information System">ITIS</a>: <a rel="nofollow" class="external text" href="https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=109216">109216</a></li> <li><a href="/wiki/National_Biodiversity_Network" title="National Biodiversity Network">NBN</a>: <a rel="nofollow" class="external text" href="https://data.nbn.org.uk/Taxa/NHMSYS0001717710">NHMSYS0001717710</a></li> <li><a href="/wiki/National_Center_for_Biotechnology_Information" title="National Center for Biotechnology Information">NCBI</a>: <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Info&id=7041">7041</a></li> <li>NZOR: <a rel="nofollow" class="external text" href="https://www.nzor.org.nz/names/ccd627b3-0250-4bcc-b61d-ed319c395a16">ccd627b3-0250-4bcc-b61d-ed319c395a16</a></li> <li><a href="/wiki/Open_Tree_of_Life" title="Open Tree of Life">Open Tree of Life</a>: <a rel="nofollow" class="external text" href="https://tree.opentreeoflife.org/taxonomy/browse?id=865243">865243</a></li> <li><a href="/wiki/Paleobiology_Database" title="Paleobiology Database">Paleobiology Database</a>: <a rel="nofollow" class="external text" href="https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=69148">69148</a></li> <li>PPE: <a rel="nofollow" class="external text" href="https://bladmineerders.nl/parasites/animalia/arthropoda/insecta/coleoptera/">coleoptera</a></li> <li><a href="/wiki/Plazi" title="Plazi">Plazi</a>: <a rel="nofollow" class="external text" href="https://treatment.plazi.org/id/F97A87E2-7400-E660-6638-FEB4FC50FECB">F97A87E2-7400-E660-6638-FEB4FC50FECB</a></li> <li><a href="/wiki/World_Register_of_Marine_Species" title="World Register of Marine Species">WoRMS</a>: <a rel="nofollow" class="external text" href="https://www.marinespecies.org/aphia.php?p=taxdetails&id=118085">118085</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"><style data-mw-deduplicate="TemplateStyles:r1038841319">.mw-parser-output .tooltip-dotted{border-bottom:1px dotted;cursor:help}</style><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1038841319"></div><div role="navigation" class="navbox authority-control" aria-labelledby="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q22671#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="padding:3px"><table class="nowraplinks hlist mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Authority_control_databases_frameless&#124;text-top&#124;10px&#124;alt=Edit_this_at_Wikidata&#124;link=https&#58;//www.wikidata.org/wiki/Q22671#identifiers&#124;class=noprint&#124;Edit_this_at_Wikidata" style="font-size:114%;margin:0 4em"><a href="/wiki/Help:Authority_control" title="Help:Authority control">Authority control databases</a> <a href="https://www.wikidata.org/wiki/Q22671#identifiers" title="Edit this at Wikidata"><img alt="Edit this at Wikidata" src="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/10px-OOjs_UI_icon_edit-ltr-progressive.svg.png" decoding="async" width="10" height="10" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/15px-OOjs_UI_icon_edit-ltr-progressive.svg.png 1.5x, //upload.wikimedia.org/wikipedia/en/thumb/8/8a/OOjs_UI_icon_edit-ltr-progressive.svg/20px-OOjs_UI_icon_edit-ltr-progressive.svg.png 2x" data-file-width="20" data-file-height="20" /></a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">National</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><a rel="nofollow" class="external text" href="https://d-nb.info/gnd/4130263-1">Germany</a></li><li><a rel="nofollow" class="external text" href="https://id.loc.gov/authorities/sh85012867">United States</a></li><li><a rel="nofollow" class="external text" href="https://catalogue.bnf.fr/ark:/12148/cb11937784z">France</a></li><li><a rel="nofollow" class="external text" href="https://data.bnf.fr/ark:/12148/cb11937784z">BnF data</a></li><li><a rel="nofollow" class="external text" href="https://id.ndl.go.jp/auth/ndlna/00566843">Japan</a></li><li><a rel="nofollow" class="external text" href="https://aleph.nkp.cz/F/?func=find-c&local_base=aut&ccl_term=ica=ph114202&CON_LNG=ENG">Czech Republic</a></li><li><a rel="nofollow" class="external text" href="http://catalogo.bne.es/uhtbin/authoritybrowse.cgi?action=display&authority_id=XX526431">Spain</a></li><li><a rel="nofollow" class="external text" href="http://olduli.nli.org.il/F/?func=find-b&local_base=NLX10&find_code=UID&request=987007283145705171">Israel</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" style="width:100%;padding:0"><div style="padding:0 0.25em"><ul><li><a rel="nofollow" class="external text" href="https://catalog.archives.gov/id/10637432">NARA</a></li></ul></div></td></tr></tbody></table></div>    </div><noscript><img src="https://login.wikimedia.org/wiki/Special:CentralAutoLogin/start?type=1x1" alt="" width="1" height="1" style="border: none; position: absolute;"></noscript> <div class="printfooter" data-nosnippet="">Retrieved from "<a dir="ltr" href="https://en.wikipedia.org/w/index.php?title=Beetle&oldid=1257880098">https://en.wikipedia.org/w/index.php?title=Beetle&oldid=1257880098</a>"</div></div> <div id="catlinks" class="catlinks" data-mw="interface"><div id="mw-normal-catlinks" class="mw-normal-catlinks"><a href="/wiki/Help:Category" title="Help:Category">Categories</a>: <ul><li><a href="/wiki/Category:Beetles" title="Category:Beetles">Beetles</a></li><li><a href="/wiki/Category:Extant_Pennsylvanian_first_appearances" title="Category:Extant Pennsylvanian first appearances">Extant Pennsylvanian first appearances</a></li><li><a href="/wiki/Category:Insects_in_culture" title="Category:Insects in culture">Insects in culture</a></li></ul></div><div id="mw-hidden-catlinks" class="mw-hidden-catlinks mw-hidden-cats-hidden">Hidden categories: <ul><li><a href="/wiki/Category:Pages_using_the_WikiHiero_extension" title="Category:Pages using the WikiHiero extension">Pages using the WikiHiero extension</a></li><li><a href="/wiki/Category:CS1_German-language_sources_(de)" title="Category:CS1 German-language sources (de)">CS1 German-language sources (de)</a></li><li><a href="/wiki/Category:CS1_Dutch-language_sources_(nl)" title="Category:CS1 Dutch-language sources (nl)">CS1 Dutch-language sources (nl)</a></li><li><a href="/wiki/Category:CS1_Russian-language_sources_(ru)" title="Category:CS1 Russian-language sources (ru)">CS1 Russian-language sources (ru)</a></li><li><a href="/wiki/Category:CS1_French-language_sources_(fr)" title="Category:CS1 French-language sources (fr)">CS1 French-language sources (fr)</a></li><li><a href="/wiki/Category:CS1_maint:_numeric_names:_authors_list" title="Category:CS1 maint: numeric names: authors list">CS1 maint: numeric names: authors list</a></li><li><a href="/wiki/Category:Articles_with_short_description" title="Category:Articles with short description">Articles with short description</a></li><li><a href="/wiki/Category:Short_description_matches_Wikidata" title="Category:Short description matches Wikidata">Short description matches Wikidata</a></li><li><a href="/wiki/Category:Good_articles" title="Category:Good articles">Good articles</a></li><li><a href="/wiki/Category:Use_mdy_dates_from_September_2012" title="Category:Use mdy dates from September 2012">Use mdy dates from September 2012</a></li><li><a href="/wiki/Category:Articles_with_%27species%27_microformats" title="Category:Articles with 'species' microformats">Articles with 'species' microformats</a></li><li><a href="/wiki/Category:Articles_containing_potentially_dated_statements_from_2015" title="Category:Articles containing potentially dated statements from 2015">Articles containing potentially dated statements from 2015</a></li><li><a href="/wiki/Category:All_articles_containing_potentially_dated_statements" title="Category:All articles containing potentially dated statements">All articles containing potentially dated statements</a></li><li><a href="/wiki/Category:All_articles_with_unsourced_statements" title="Category:All articles with unsourced statements">All articles with unsourced statements</a></li><li><a href="/wiki/Category:Articles_with_unsourced_statements_from_June_2023" title="Category:Articles with unsourced statements from June 2023">Articles with unsourced statements from June 2023</a></li><li><a href="/wiki/Category:Articles_with_German-language_sources_(de)" title="Category:Articles with German-language sources (de)">Articles with German-language sources (de)</a></li><li><a href="/wiki/Category:Webarchive_template_wayback_links" title="Category:Webarchive template wayback links">Webarchive template wayback links</a></li><li><a href="/wiki/Category:Taxonbars_with_20%E2%80%9324_taxon_IDs" title="Category:Taxonbars with 20–24 taxon IDs">Taxonbars with 20–24 taxon IDs</a></li></ul></div></div> </div> </main> </div> <div class="mw-footer-container"> <footer id="footer" class="mw-footer" > <ul id="footer-info"> <li id="footer-info-lastmod"> This page was last edited on 17 November 2024, at 02:21 (UTC).</li> <li id="footer-info-copyright">Text is available under the <a href="/wiki/Wikipedia:Text_of_the_Creative_Commons_Attribution-ShareAlike_4.0_International_License" title="Wikipedia:Text of the Creative Commons Attribution-ShareAlike 4.0 International License">Creative Commons Attribution-ShareAlike 4.0 License</a>; additional terms may apply. By using this site, you agree to the <a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Terms_of_Use" class="extiw" title="foundation:Special:MyLanguage/Policy:Terms of Use">Terms of Use</a> and <a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Privacy_policy" class="extiw" title="foundation:Special:MyLanguage/Policy:Privacy policy">Privacy Policy</a>. Wikipedia® is a registered trademark of the <a rel="nofollow" class="external text" href="https://wikimediafoundation.org/">Wikimedia Foundation, Inc.</a>, a non-profit organization.</li> </ul> <ul id="footer-places"> <li id="footer-places-privacy"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Privacy_policy">Privacy policy</a></li> <li id="footer-places-about"><a href="/wiki/Wikipedia:About">About Wikipedia</a></li> <li id="footer-places-disclaimers"><a href="/wiki/Wikipedia:General_disclaimer">Disclaimers</a></li> <li id="footer-places-contact"><a href="//en.wikipedia.org/wiki/Wikipedia:Contact_us">Contact Wikipedia</a></li> <li id="footer-places-wm-codeofconduct"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Universal_Code_of_Conduct">Code of Conduct</a></li> <li id="footer-places-developers"><a href="https://developer.wikimedia.org">Developers</a></li> <li id="footer-places-statslink"><a href="https://stats.wikimedia.org/#/en.wikipedia.org">Statistics</a></li> <li id="footer-places-cookiestatement"><a href="https://foundation.wikimedia.org/wiki/Special:MyLanguage/Policy:Cookie_statement">Cookie statement</a></li> <li id="footer-places-mobileview"><a href="//en.m.wikipedia.org/w/index.php?title=Beetle&mobileaction=toggle_view_mobile" class="noprint stopMobileRedirectToggle">Mobile view</a></li> </ul> <ul id="footer-icons" class="noprint"> <li id="footer-copyrightico"><a href="https://wikimediafoundation.org/" class="cdx-button cdx-button--fake-button cdx-button--size-large cdx-button--fake-button--enabled"><img src="/static/images/footer/wikimedia-button.svg" width="84" height="29" alt="Wikimedia Foundation" loading="lazy"></a></li> <li id="footer-poweredbyico"><a href="https://www.mediawiki.org/" class="cdx-button cdx-button--fake-button cdx-button--size-large cdx-button--fake-button--enabled"><img src="/w/resources/assets/poweredby_mediawiki.svg" alt="Powered by MediaWiki" width="88" height="31" loading="lazy"></a></li> </ul> </footer> </div> </div> </div> <div class="vector-settings" id="p-dock-bottom"> <ul></ul> </div><script>(RLQ=window.RLQ||[]).push(function(){mw.config.set({"wgHostname":"mw-web.codfw.main-f69cdc8f6-fn5x6","wgBackendResponseTime":162,"wgPageParseReport":{"limitreport":{"cputime":"2.455","walltime":"2.849","ppvisitednodes":{"value":29243,"limit":1000000},"postexpandincludesize":{"value":914484,"limit":2097152},"templateargumentsize":{"value":26109,"limit":2097152},"expansiondepth":{"value":28,"limit":100},"expensivefunctioncount":{"value":49,"limit":500},"unstrip-depth":{"value":1,"limit":20},"unstrip-size":{"value":887119,"limit":5000000},"entityaccesscount":{"value":23,"limit":400},"timingprofile":["100.00% 2387.275 1 -total"," 45.88% 1095.260 2 Template:Reflist"," 25.93% 619.029 111 Template:Cite_journal"," 13.51% 322.550 23 Template:Clade"," 12.38% 295.429 1 Template:Automatic_taxobox"," 8.03% 191.723 50 Template:Cite_book"," 7.24% 172.833 1 Template:Taxonbar"," 6.05% 144.446 24 Template:Navbox"," 5.33% 127.316 36 Template:Cite_web"," 5.27% 125.916 2 Template:Navbox_with_collapsible_groups"]},"scribunto":{"limitreport-timeusage":{"value":"1.576","limit":"10.000"},"limitreport-memusage":{"value":16788070,"limit":52428800},"limitreport-logs":"table#1 {\n}\ntable#1 {\n [\"size\"] = \"small\",\n}\n","limitreport-profile":[["?","400","24.4"],["dataWrapper \u003Cmw.lua:672\u003E","140","8.5"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::getEntity","100","6.1"],["tostring","100","6.1"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::gsub","100","6.1"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::callParserFunction","80","4.9"],["recursiveClone \u003CmwInit.lua:45\u003E","80","4.9"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::match","60","3.7"],["init \u003CModule:Citation/CS1/Date_validation\u003E","60","3.7"],["MediaWiki\\Extension\\Scribunto\\Engines\\LuaSandbox\\LuaSandboxCallback::len","60","3.7"],["[others]","460","28.0"]]},"cachereport":{"origin":"mw-web.codfw.main-f69cdc8f6-69xkf","timestamp":"20241124113041","ttl":2592000,"transientcontent":false}}});});</script> <script type="application/ld+json">{"@context":"https:\/\/schema.org","@type":"Article","name":"Beetle","url":"https:\/\/en.wikipedia.org\/wiki\/Beetle","sameAs":"http:\/\/www.wikidata.org\/entity\/Q22671","mainEntity":"http:\/\/www.wikidata.org\/entity\/Q22671","author":{"@type":"Organization","name":"Contributors to Wikimedia projects"},"publisher":{"@type":"Organization","name":"Wikimedia Foundation, Inc.","logo":{"@type":"ImageObject","url":"https:\/\/www.wikimedia.org\/static\/images\/wmf-hor-googpub.png"}},"datePublished":"2001-11-08T14:06:00Z","dateModified":"2024-11-17T02:21:09Z","image":"https:\/\/upload.wikimedia.org\/wikipedia\/commons\/2\/20\/Coleoptera_collage.png","headline":"order of insects"}</script> </body> </html>