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</div> </div> </nav> <h1 id="firstHeading" class="firstHeading mw-first-heading"><span class="mw-page-title-main">Amplexus</span></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. 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href="https://ca.wikipedia.org/wiki/Amplexus" title="Amplexus – Catalan" lang="ca" hreflang="ca" data-title="Amplexus" data-language-autonym="Català" data-language-local-name="Catalan" class="interlanguage-link-target"><span>Català</span></a></li><li class="interlanguage-link interwiki-cs mw-list-item"><a href="https://cs.wikipedia.org/wiki/Amplexus" title="Amplexus – Czech" lang="cs" hreflang="cs" data-title="Amplexus" data-language-autonym="Čeština" data-language-local-name="Czech" class="interlanguage-link-target"><span>Čeština</span></a></li><li class="interlanguage-link interwiki-de mw-list-item"><a href="https://de.wikipedia.org/wiki/Amplexus" title="Amplexus – German" lang="de" hreflang="de" data-title="Amplexus" data-language-autonym="Deutsch" data-language-local-name="German" class="interlanguage-link-target"><span>Deutsch</span></a></li><li class="interlanguage-link interwiki-es mw-list-item"><a href="https://es.wikipedia.org/wiki/Amplexo" title="Amplexo – Spanish" lang="es" hreflang="es" data-title="Amplexo" data-language-autonym="Español" data-language-local-name="Spanish" class="interlanguage-link-target"><span>Español</span></a></li><li class="interlanguage-link interwiki-eo mw-list-item"><a href="https://eo.wikipedia.org/wiki/Amplekso" title="Amplekso – Esperanto" lang="eo" hreflang="eo" data-title="Amplekso" data-language-autonym="Esperanto" data-language-local-name="Esperanto" class="interlanguage-link-target"><span>Esperanto</span></a></li><li class="interlanguage-link interwiki-eu mw-list-item"><a href="https://eu.wikipedia.org/wiki/Anplexo" title="Anplexo – Basque" lang="eu" hreflang="eu" data-title="Anplexo" data-language-autonym="Euskara" data-language-local-name="Basque" class="interlanguage-link-target"><span>Euskara</span></a></li><li class="interlanguage-link interwiki-fa mw-list-item"><a href="https://fa.wikipedia.org/wiki/%D8%A2%D8%BA%D9%88%D8%B4%E2%80%8C%DA%AF%DB%8C%D8%B1%DB%8C" title="آغوش‌گیری – Persian" lang="fa" hreflang="fa" data-title="آغوش‌گیری" data-language-autonym="فارسی" data-language-local-name="Persian" class="interlanguage-link-target"><span>فارسی</span></a></li><li class="interlanguage-link interwiki-fr mw-list-item"><a href="https://fr.wikipedia.org/wiki/Amplexus" title="Amplexus – French" lang="fr" hreflang="fr" data-title="Amplexus" data-language-autonym="Français" data-language-local-name="French" class="interlanguage-link-target"><span>Français</span></a></li><li class="interlanguage-link interwiki-gl mw-list-item"><a href="https://gl.wikipedia.org/wiki/Amplexo" title="Amplexo – Galician" lang="gl" hreflang="gl" data-title="Amplexo" data-language-autonym="Galego" data-language-local-name="Galician" class="interlanguage-link-target"><span>Galego</span></a></li><li class="interlanguage-link interwiki-he mw-list-item"><a href="https://he.wikipedia.org/wiki/%D7%97%D7%99%D7%91%D7%95%D7%A7_%D7%9B%D7%9C%D7%95%D7%9C%D7%95%D7%AA" title="חיבוק כלולות – Hebrew" lang="he" hreflang="he" data-title="חיבוק כלולות" data-language-autonym="עברית" data-language-local-name="Hebrew" class="interlanguage-link-target"><span>עברית</span></a></li><li class="interlanguage-link interwiki-hu mw-list-item"><a href="https://hu.wikipedia.org/wiki/Amplexus" title="Amplexus – Hungarian" lang="hu" hreflang="hu" data-title="Amplexus" data-language-autonym="Magyar" data-language-local-name="Hungarian" class="interlanguage-link-target"><span>Magyar</span></a></li><li class="interlanguage-link interwiki-ml mw-list-item"><a href="https://ml.wikipedia.org/wiki/%E0%B4%86%E0%B4%82%E0%B4%AA%E0%B5%8D%E0%B4%B2%E0%B4%BF%E0%B4%95%E0%B5%8D%E2%80%8C%E0%B4%B8%E0%B4%B8%E0%B5%8D" title="ആംപ്ലിക്‌സസ് – Malayalam" lang="ml" hreflang="ml" data-title="ആംപ്ലിക്‌സസ്" data-language-autonym="മലയാളം" data-language-local-name="Malayalam" class="interlanguage-link-target"><span>മലയാളം</span></a></li><li class="interlanguage-link interwiki-nl mw-list-item"><a href="https://nl.wikipedia.org/wiki/Amplexus_(dier)" title="Amplexus (dier) – Dutch" lang="nl" hreflang="nl" data-title="Amplexus (dier)" data-language-autonym="Nederlands" data-language-local-name="Dutch" class="interlanguage-link-target"><span>Nederlands</span></a></li><li class="interlanguage-link interwiki-pl mw-list-item"><a href="https://pl.wikipedia.org/wiki/Ampleksus" title="Ampleksus – Polish" lang="pl" hreflang="pl" data-title="Ampleksus" data-language-autonym="Polski" data-language-local-name="Polish" class="interlanguage-link-target"><span>Polski</span></a></li><li class="interlanguage-link interwiki-pt mw-list-item"><a href="https://pt.wikipedia.org/wiki/Amplexo" title="Amplexo – Portuguese" lang="pt" hreflang="pt" data-title="Amplexo" data-language-autonym="Português" data-language-local-name="Portuguese" class="interlanguage-link-target"><span>Português</span></a></li><li class="interlanguage-link interwiki-ru mw-list-item"><a href="https://ru.wikipedia.org/wiki/%D0%90%D0%BC%D0%BF%D0%BB%D0%B5%D0%BA%D1%81%D1%83%D1%81" title="Амплексус – Russian" lang="ru" hreflang="ru" data-title="Амплексус" data-language-autonym="Русский" data-language-local-name="Russian" class="interlanguage-link-target"><span>Русский</span></a></li><li class="interlanguage-link interwiki-fi mw-list-item"><a href="https://fi.wikipedia.org/wiki/Amplexus" title="Amplexus – Finnish" lang="fi" hreflang="fi" data-title="Amplexus" data-language-autonym="Suomi" data-language-local-name="Finnish" class="interlanguage-link-target"><span>Suomi</span></a></li><li class="interlanguage-link interwiki-sv mw-list-item"><a href="https://sv.wikipedia.org/wiki/Amplexus" title="Amplexus – Swedish" lang="sv" hreflang="sv" data-title="Amplexus" data-language-autonym="Svenska" data-language-local-name="Swedish" class="interlanguage-link-target"><span>Svenska</span></a></li><li class="interlanguage-link interwiki-vi mw-list-item"><a href="https://vi.wikipedia.org/wiki/C%C3%B5ng_gh%C3%A9p_%C4%91%C3%B4i" title="Cõng ghép đôi – Vietnamese" lang="vi" hreflang="vi" data-title="Cõng 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<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">Type of mating behavior exhibited by some externally fertilizing species</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 record label, see <a href="/wiki/Amplexus_(label)" title="Amplexus (label)">Amplexus (label)</a>.</div><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Strongylopus_grayii_in_amplexus.png" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Strongylopus_grayii_in_amplexus.png/220px-Strongylopus_grayii_in_amplexus.png" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/1/16/Strongylopus_grayii_in_amplexus.png/330px-Strongylopus_grayii_in_amplexus.png 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/1/16/Strongylopus_grayii_in_amplexus.png/440px-Strongylopus_grayii_in_amplexus.png 2x" data-file-width="5192" data-file-height="3472" /></a><figcaption>A pair of clicking stream frogs (<i><a href="/wiki/Gray%27s_stream_frog" title="Gray's stream frog">Strongylopus grayii</a></i>) in amplexus</figcaption></figure><figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Litoria_xanthomera_amplexus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Litoria_xanthomera_amplexus.jpg/220px-Litoria_xanthomera_amplexus.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/9/98/Litoria_xanthomera_amplexus.jpg/330px-Litoria_xanthomera_amplexus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/9/98/Litoria_xanthomera_amplexus.jpg/440px-Litoria_xanthomera_amplexus.jpg 2x" data-file-width="1600" data-file-height="1200" /></a><figcaption><a href="/wiki/Orange-thighed_frog" title="Orange-thighed frog">Orange-thighed frogs</a> (<i><a href="/wiki/Litoria_xanthomera" class="mw-redirect" title="Litoria xanthomera">Litoria xanthomera</a></i>) in amplexus</figcaption></figure> <p><b>Amplexus</b> (Latin "embrace") is a type of <a href="/wiki/Mating" title="Mating">mating behavior</a> exhibited by some <a href="/wiki/External_fertilization" title="External fertilization">externally fertilizing</a> species (chiefly <a href="/wiki/Amphibian" title="Amphibian">amphibians</a> and <a href="/wiki/Horseshoe_crabs" class="mw-redirect" title="Horseshoe crabs">horseshoe crabs</a>) in which a male grasps a female with his front legs as part of the mating process, and at the same time or with some time delay, he <a href="/wiki/Fertilize" class="mw-redirect" title="Fertilize">fertilizes</a> the eggs, as they are released from the female's body.<sup id="cite_ref-:02_1-0" class="reference"><a href="#cite_note-:02-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> In amphibians, females may be grasped by the head, waist, or armpits, and the type of amplexus is characteristic of some taxonomic groups. </p><p>Amplexus involves direct contact between male and female, distinguished from other forms of external fertilization, such as <a href="/wiki/Broadcast_spawning" class="mw-redirect" title="Broadcast spawning">broadcast spawning</a>, where sperm and eggs are freely shed into water without direct contact by individuals. In order for amplexus to be initiated, male frogs must first find a mate by attracting one through calls, typically in the evening.<sup id="cite_ref-:1_2-0" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Once a male has successfully attracted a mate, the process of amplexus begins, while the unsuccessful males are forced to continue their search for a mate through further calls.<sup id="cite_ref-:1_2-1" class="reference"><a href="#cite_note-:1-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> </p><p>The competition for a female mate among males is considered intense, and it is not uncommon for a male amphibian to attack an already-amplexed pair of amphibians.<sup id="cite_ref-:4_3-0" class="reference"><a href="#cite_note-:4-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> When a male amphibian attacks an amplexed pair of amphibians, he is trying to force the other male to release its grasp of the female, so he can then mate with her.<sup id="cite_ref-:4_3-1" class="reference"><a href="#cite_note-:4-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> Male amphibians are also known to show mate-guarding behaviour, which is shown after amplexus, and it is the male's attempt to prevent the female amphibian from mating with other males.<sup id="cite_ref-:4_3-2" class="reference"><a href="#cite_note-:4-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup> </p><p>The duration of amplexus has been found to vary across species. In some species it may last for many days, while in others it may last a few hours.<sup id="cite_ref-:12_4-0" class="reference"><a href="#cite_note-:12-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Despite the variation in the duration of amplexus across species, typically all species that exhibit this behaviour have to use their forelimb muscles for the duration of amplexus.<sup id="cite_ref-:12_4-1" class="reference"><a href="#cite_note-:12-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> Studies have found that this reproductive behaviour of amplexus can come with different fitness costs, due to the fact that amplexus can occur for prolonged periods of time. For instance, a study found that when a male amphibian is grasping and holding onto a female amphibian, this can lead to the impairment of the female's ability to move or to feed.<sup id="cite_ref-:7_5-0" class="reference"><a href="#cite_note-:7-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> After conducting experiments, researchers confirmed that amplexus does decrease a female's locomotor performance (e.g., swimming, walking) as well as the feeding rates.<sup id="cite_ref-:7_5-1" class="reference"><a href="#cite_note-:7-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> With regard to the cost to the male engaged in amplexus, male amphibians have been found to not feed at all during amplexus.<sup id="cite_ref-:7_5-2" class="reference"><a href="#cite_note-:7-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> </p> <meta property="mw:PageProp/toc" /> <div class="mw-heading mw-heading2"><h2 id="Types">Types</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=1" title="Edit section: Types"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Many types of amplexus are identified in the literature. However, two types of amplexus are more common than others, known as inguinal and axillary amplexus.<sup id="cite_ref-:3_6-0" class="reference"><a href="#cite_note-:3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> These two types of amplexus have been classified based on the position of the male amphibian relative to the female.<sup id="cite_ref-:3_6-1" class="reference"><a href="#cite_note-:3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> When a male amphibian clasps a female around her waist (inguinal region) using his forelimbs, this is considered inguinal amplexus.<sup id="cite_ref-:3_6-2" class="reference"><a href="#cite_note-:3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> By contrast, when a male amphibian clasps behind the forelimbs (axillary region) of the female, this is considered axillary amplexus.<sup id="cite_ref-:3_6-3" class="reference"><a href="#cite_note-:3-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Physiological_and_hormonal_function">Physiological and hormonal function</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=2" title="Edit section: Physiological and hormonal function"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <div class="mw-heading mw-heading3"><h3 id="Physiology">Physiology</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=3" title="Edit section: Physiology"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Amplexus has been found to involve different muscles in both male and female amphibians. The forelimb muscles in both males and females have been identified as the key muscles for amplexus that most species use.<sup id="cite_ref-:12_4-2" class="reference"><a href="#cite_note-:12-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> These forelimb muscles that are used during amplexus are typically larger in males than females, and for males these muscles contain more oxidative fibers, which could mean that amplexus involves an increased rate of aerobic metabolism.<sup id="cite_ref-:12_4-3" class="reference"><a href="#cite_note-:12-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> In addition to those forelimb muscles being larger in males, male frogs also typically have keratin pads or nuptial pads, which are located on their thumbs and contribute to the success of amplexus by assisting in gripping the female for the duration of amplexus.<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> This amplexus behaviour allows the amphibians' cloacae to be in close contact, while gametes are released.<sup id="cite_ref-:2_8-0" class="reference"><a href="#cite_note-:2-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Additionally, amplexus is thought to help with the alignment of the reproductive tracts of both males and females, which furthermore contributes to successful fertilization.<sup id="cite_ref-:02_1-1" class="reference"><a href="#cite_note-:02-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> A female amphibian may not always be receptive to a male amphibian who is trying to initiate amplexus, as the female may not be ready to produce eggs.<sup id="cite_ref-:02_1-2" class="reference"><a href="#cite_note-:02-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> When the female is not ready to engage in amplexus, she will simply vibrate her body, which will then be felt by the male who is clasped to her and he will then stop the amplexus behaviour.<sup id="cite_ref-:02_1-3" class="reference"><a href="#cite_note-:02-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Hormones">Hormones</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=4" title="Edit section: Hormones"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Two major hormones have been identified to be involved in amplexus. The hormone arginine vasotocin (AVT) has been identified as having an effect on the calling behaviours produced by these male amphibians when searching for a female mate, as AVT increased the amount of calling behaviour produced in male amphibians.<sup id="cite_ref-:2_8-1" class="reference"><a href="#cite_note-:2-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Additionally, it is believed that the gonadotropin-releasing hormone (GnRH) has an influence on amplexus in amphibians, as it has been found to produce or initiate this behaviour in many anuran amphibians.<sup id="cite_ref-:2_8-2" class="reference"><a href="#cite_note-:2-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Furthermore, both arginine vasotocin and gonadotropin-releasing hormone have been found to be involved in the sexual behaviour of male amphibians.<sup id="cite_ref-:2_8-3" class="reference"><a href="#cite_note-:2-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> The stress hormone corticosterone has also been identified as associated with the sexual behaviour of amplexus. A study conducted on red-spotted newts found an acute increase in corticosterone for both males and females who were engaged in the behaviour of amplexus.<sup id="cite_ref-:5_9-0" class="reference"><a href="#cite_note-:5-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> However, overall it was found that males have a higher level of corticosterone compared to females, as amplexus is seen as more energetically costly to males.<sup id="cite_ref-:5_9-1" class="reference"><a href="#cite_note-:5-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> The increase in corticosterone found in females could be attributed to amplexus preventing them from foraging for food.<sup id="cite_ref-:5_9-2" class="reference"><a href="#cite_note-:5-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Amphibians">Amphibians</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=5" title="Edit section: Amphibians"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Mating_pair_of_common_frogs_in_amplexus.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/5/5d/Mating_pair_of_common_frogs_in_amplexus.jpg/220px-Mating_pair_of_common_frogs_in_amplexus.jpg" decoding="async" width="220" height="165" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/5/5d/Mating_pair_of_common_frogs_in_amplexus.jpg/330px-Mating_pair_of_common_frogs_in_amplexus.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/5/5d/Mating_pair_of_common_frogs_in_amplexus.jpg/440px-Mating_pair_of_common_frogs_in_amplexus.jpg 2x" data-file-width="3920" data-file-height="2940" /></a><figcaption>European <a href="/wiki/Common_frog" title="Common frog">common frog</a></figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Amplexus_Bufo_bufo_2010-03-29.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Amplexus_Bufo_bufo_2010-03-29.jpg/220px-Amplexus_Bufo_bufo_2010-03-29.jpg" decoding="async" width="220" height="143" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/33/Amplexus_Bufo_bufo_2010-03-29.jpg/330px-Amplexus_Bufo_bufo_2010-03-29.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/33/Amplexus_Bufo_bufo_2010-03-29.jpg/440px-Amplexus_Bufo_bufo_2010-03-29.jpg 2x" data-file-width="3747" data-file-height="2429" /></a><figcaption>European <a href="/wiki/Common_toad" title="Common toad">common toad</a> (<i><a href="/wiki/Bufo_bufo" class="mw-redirect" title="Bufo bufo">Bufo bufo</a></i>).</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Common_toad_Bufo_bufo_multiple_amplexus_mating_ball.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Common_toad_Bufo_bufo_multiple_amplexus_mating_ball.jpg/220px-Common_toad_Bufo_bufo_multiple_amplexus_mating_ball.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Common_toad_Bufo_bufo_multiple_amplexus_mating_ball.jpg/330px-Common_toad_Bufo_bufo_multiple_amplexus_mating_ball.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/d/d8/Common_toad_Bufo_bufo_multiple_amplexus_mating_ball.jpg/440px-Common_toad_Bufo_bufo_multiple_amplexus_mating_ball.jpg 2x" data-file-width="1620" data-file-height="1080" /></a><figcaption><a href="/wiki/Common_toad" title="Common toad">Common toad</a> <a href="/wiki/Mating_ball" title="Mating ball">mating ball</a> (multiple amplexus).</figcaption></figure> <figure class="mw-default-size" typeof="mw:File/Thumb"><a href="/wiki/File:Red-eyed_tree_frogs_mating.JPG" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/6/68/Red-eyed_tree_frogs_mating.JPG/220px-Red-eyed_tree_frogs_mating.JPG" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/6/68/Red-eyed_tree_frogs_mating.JPG/330px-Red-eyed_tree_frogs_mating.JPG 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/6/68/Red-eyed_tree_frogs_mating.JPG/440px-Red-eyed_tree_frogs_mating.JPG 2x" data-file-width="3312" data-file-height="2208" /></a><figcaption><a href="/wiki/Agalychnis_callidryas" title="Agalychnis callidryas">Red-eyed tree frog</a> (<i><a href="/wiki/Agalychnis_callidryas" title="Agalychnis callidryas">Agalychnis callidryas</a></i>), <a href="/wiki/Costa_Rica" title="Costa Rica">Costa Rica</a></figcaption></figure> <div class="mw-heading mw-heading3"><h3 id="Anurans">Anurans</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=6" title="Edit section: Anurans"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Amplexus chiefly occurs aquatically, but some terrestrial <a href="/wiki/Frog" title="Frog">anurans</a> (frogs and toads) like the <a href="/wiki/Disc-tongued_frog" class="mw-redirect" title="Disc-tongued frog">disc-tongued frogs</a> (<a href="/wiki/Discoglossidae" class="mw-redirect" title="Discoglossidae">Discoglossidae</a>) perform amplexus on land. In crown-group anurans, like the true frogs (<a href="/wiki/Ranidae" class="mw-redirect" title="Ranidae">Ranidae</a>), the <a href="/wiki/Tree_frog" title="Tree frog">tree frogs</a> (Hylidae), and the true toads (<a href="/wiki/Bufonidae" class="mw-redirect" title="Bufonidae">Bufonidae</a>), amplexus is axillary (in the armpits). Other anurans (the <a href="/wiki/Archaeobatrachia" title="Archaeobatrachia">Archaeobatrachia</a>, <a href="/wiki/Sooglossidae" title="Sooglossidae">Sooglossidae</a> and <a href="/wiki/Myobatrachidae" title="Myobatrachidae">Myobatrachidae</a>), show the ancestral state which is inguinal or lumbar amplexus (abdominal, in front of the hindlegs). Some species show cephalic amplexus where the head of the female is held while others show complete lack of amplexus.<sup id="cite_ref-Duellman&Trueb_10-0" class="reference"><a href="#cite_note-Duellman&Trueb-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> Additionally, anurans species have been observed to engage in multiple amplexus, which can also be referred to as a mating ball, as many toads attach themselves to a female trying to initiate amplexus.<sup id="cite_ref-:8_11-0" class="reference"><a href="#cite_note-:8-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> However, multiple amplexus is not common among anurans, which could indicate that the costs associated with multiple amplexus are higher than the advantages associated with it.<sup id="cite_ref-:8_11-1" class="reference"><a href="#cite_note-:8-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> For female anurans, the idea of multiple amplexus would probably be more advantageous, because mating with more than one male would increase fertilization chances or increase offspring genetic diversity.<sup id="cite_ref-:8_11-2" class="reference"><a href="#cite_note-:8-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup> Multiple amplexus would typically be common in explosive breeding amphibians, when there is a larger number of adults ready to breed at a breeding site in a short period of time.<sup id="cite_ref-:02_1-4" class="reference"><a href="#cite_note-:02-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> When this occurs female amphibians are viewed as a very important resource for males, as there are typically more males present than females, thus leading to more chances for multiple amplexus to occur.<sup id="cite_ref-:02_1-5" class="reference"><a href="#cite_note-:02-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> </p><p>In most anurans, the males deposit sperm onto the eggs as they are being laid, however males of the genus <i><a href="/wiki/Ascaphus" class="mw-redirect" title="Ascaphus">Ascaphus</a></i> possess an <a href="/wiki/Intromittent_organ" title="Intromittent organ">intromittent organ</a>, unique among anurans, for internal fertilization. Internal fertilization does occur in a few other genera, including <i><a href="/wiki/Nectophrynoides" title="Nectophrynoides">Nectophrynoides</a></i>, <i><a href="/wiki/Mertensophryne" title="Mertensophryne">Mertensophryne</a></i>, and <i><a href="/wiki/Eleutherodactylus" title="Eleutherodactylus">Eleutherodactylus</a></i>.<sup id="cite_ref-Duellman&Trueb_10-1" class="reference"><a href="#cite_note-Duellman&Trueb-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading3"><h3 id="Newts">Newts</h3><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=7" title="Edit section: Newts"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>In the case of <a href="/wiki/Newt" title="Newt">newts</a>, the process of amplexus is often observed soon after the newts become seasonally active. In the Western USA, for example, this time is typically soon after the onset of the winter rainy season, when <a href="/wiki/Intermittent_stream" class="mw-redirect" title="Intermittent stream">intermittent streams</a> and <a href="/wiki/Vernal_pools" class="mw-redirect" title="Vernal pools">vernal pools</a> become available as a breeding habitat. The <a href="/wiki/Rough-skinned_newt" title="Rough-skinned newt">rough-skinned newt</a> is a specific widespread example of a newt in the western USA that can be observed in quiet stream pools and shallow ponds engaging in amplexus.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> During amplexus in newts, males will typically show the behaviour of tail fanning and chin rubbing which is thought to prompt the mating receptivity of the female newt.<sup id="cite_ref-:11_14-0" class="reference"><a href="#cite_note-:11-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Studies have shown that male newts who have deeper tail-fins have better control of females during amplexus and are also more successful in catching the females for amplexus.<sup id="cite_ref-:11_14-1" class="reference"><a href="#cite_note-:11-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Additionally, it has been found that the probability of a male newt who has a deeper tail-fin to achieve amplexus is greater than those newts who do not contain a deeper tail-fin, as male newts tend to use their tails during male-male competition.<sup id="cite_ref-:11_14-2" class="reference"><a href="#cite_note-:11-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> When a male newt, who is unpaired, encounters a female and male newt engaged in amplexus, the unpaired newt will try to displace the paired male newt by using wrestling tactics.<sup id="cite_ref-:9_15-0" class="reference"><a href="#cite_note-:9-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> A study examining the wrestling behaviour of newts found that of the observed wrestling encounters, 90% were "won" by the paired male, meaning he would retain the female newt.<sup id="cite_ref-:9_15-1" class="reference"><a href="#cite_note-:9-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> The study found that the invading unpaired newt rarely successfully displaces the paired male newt, engaged in amplexus.<sup id="cite_ref-:9_15-2" class="reference"><a href="#cite_note-:9-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Horseshoe_crabs">Horseshoe crabs</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=8" title="Edit section: Horseshoe crabs"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <figure class="mw-default-size mw-halign-right" typeof="mw:File/Thumb"><a href="/wiki/File:Limulus_polyphemus1.jpg" class="mw-file-description"><img src="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Limulus_polyphemus1.jpg/220px-Limulus_polyphemus1.jpg" decoding="async" width="220" height="147" class="mw-file-element" srcset="//upload.wikimedia.org/wikipedia/commons/thumb/3/35/Limulus_polyphemus1.jpg/330px-Limulus_polyphemus1.jpg 1.5x, //upload.wikimedia.org/wikipedia/commons/thumb/3/35/Limulus_polyphemus1.jpg/440px-Limulus_polyphemus1.jpg 2x" data-file-width="3072" data-file-height="2048" /></a><figcaption>Amplectant pair of <i><a href="/wiki/Limulus_polyphemus" class="mw-redirect" title="Limulus polyphemus">Limulus polyphemus</a></i>. The male is the smaller individual.</figcaption></figure> <p>Amplexus occurs in all four species of <a href="/wiki/Horseshoe_crab" title="Horseshoe crab">horseshoe crab</a>. Horseshoe crabs typically go ashore for amplexus in high tide, and end up on beaches where the eggs are more protected.<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> The first pair of walking legs is used to tightly clasp the female in all species, and the second pair is also employed in all but <i><a href="/wiki/Limulus_polyphemus" class="mw-redirect" title="Limulus polyphemus">Limulus polyphemus</a></i>.<sup id="cite_ref-Bottonetal1996_17-0" class="reference"><a href="#cite_note-Bottonetal1996-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> A male horseshoe crab develops modified <a href="/wiki/Clasper" title="Clasper">claspers</a> during sexual maturity when the male moults; these modified claspers can then help during the process of amplexus.<sup id="cite_ref-:6_18-0" class="reference"><a href="#cite_note-:6-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> The male's pair of posterior claspers are known for having the ability to maintain long-term amplexus which have been found to always attach to the female's <a href="/wiki/Opisthosoma" title="Opisthosoma">opisthosoma</a> during amplexus.<sup id="cite_ref-:6_18-1" class="reference"><a href="#cite_note-:6-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> In contrast, the male's anterior claspers have been found to attach to the female's opisthosoma as well, but on the lateral edges of the opisthosoma and function to resist displacement from environmental factors.<sup id="cite_ref-:6_18-2" class="reference"><a href="#cite_note-:6-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Uniquely, amplexus is most likely to occur between horseshoe crabs when the female horseshoe crab has a hard shell.<sup id="cite_ref-:6_18-3" class="reference"><a href="#cite_note-:6-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> Additionally, males who inhabit a clean shell are more likely to enter amplexus, compared to males who contain a dirty shell, as it seems that females have a preference for clean shells on males.<sup id="cite_ref-:6_18-4" class="reference"><a href="#cite_note-:6-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup> A male horseshoe crab's claspers may also be an important factor of consideration for the initiation of amplexus. Since claspers are used for the attachment to the female, claspers that are in good condition are more successful for the initiation of amplexus.<sup id="cite_ref-:10_19-0" class="reference"><a href="#cite_note-:10-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> If a male horseshoe crab has a damaged or missing clasper, then that puts the male at a disadvantage and increases the probability of being displaced by other competing male crabs.<sup id="cite_ref-:10_19-1" class="reference"><a href="#cite_note-:10-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="Other_animals">Other animals</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=9" title="Edit section: Other animals"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <p>Fossil evidence suggests that a certain <a href="/wiki/Euthycarcinoid" class="mw-redirect" title="Euthycarcinoid">euthycarcinoid</a> (an extinct arthropod) from the <a href="/wiki/Cambrian" title="Cambrian">Cambrian</a> may also have mated by amplexus.<sup id="cite_ref-Collette2012_20-0" class="reference"><a href="#cite_note-Collette2012-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> </p> <div class="mw-heading mw-heading2"><h2 id="See_also">See also</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=10" title="Edit section: See also"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <ul><li><a href="/wiki/Nuptial_pad" title="Nuptial pad">Nuptial pad</a> – Secondary sex characteristic on some mature male frogs and salamanders</li></ul> <div class="mw-heading mw-heading2"><h2 id="References">References</h2><span class="mw-editsection"><span class="mw-editsection-bracket">[</span><a href="/w/index.php?title=Amplexus&action=edit&section=11" title="Edit section: References"><span>edit</span></a><span class="mw-editsection-bracket">]</span></span></div> <style data-mw-deduplicate="TemplateStyles:r1239543626">.mw-parser-output .reflist{margin-bottom:0.5em;list-style-type:decimal}@media screen{.mw-parser-output .reflist{font-size:90%}}.mw-parser-output .reflist .references{font-size:100%;margin-bottom:0;list-style-type:inherit}.mw-parser-output .reflist-columns-2{column-width:30em}.mw-parser-output .reflist-columns-3{column-width:25em}.mw-parser-output .reflist-columns{margin-top:0.3em}.mw-parser-output .reflist-columns ol{margin-top:0}.mw-parser-output .reflist-columns li{page-break-inside:avoid;break-inside:avoid-column}.mw-parser-output .reflist-upper-alpha{list-style-type:upper-alpha}.mw-parser-output .reflist-upper-roman{list-style-type:upper-roman}.mw-parser-output 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.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="CITEREFMollovPopgeorgievNaumovTzankov2010" class="citation journal cs1">Mollov, Ivelin; Popgeorgiev, Georgi; Naumov, Borislav; Tzankov, Nikolay; Stoyanov, Andrei (2010). "Cases of abnormal amplexus in anurans (Amphibia: Anura) from Bulgaria and Greece". <i>Biharean Biologist</i>.</cite><span title="ctx_ver=Z39.88-2004&rft_val_fmt=info%3Aofi%2Ffmt%3Akev%3Amtx%3Ajournal&rft.genre=article&rft.jtitle=Biharean+Biologist&rft.atitle=Cases+of+abnormal+amplexus+in+anurans+%28Amphibia%3A+Anura%29+from+Bulgaria+and+Greece&rft.date=2010&rft.aulast=Mollov&rft.aufirst=Ivelin&rft.au=Popgeorgiev%2C+Georgi&rft.au=Naumov%2C+Borislav&rft.au=Tzankov%2C+Nikolay&rft.au=Stoyanov%2C+Andrei&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAmplexus" class="Z3988"></span></span> </li> <li id="cite_note-:1-2"><span class="mw-cite-backlink">^ <a href="#cite_ref-:1_2-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:1_2-1"><sup><i><b>b</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFMcLister2003" class="citation journal cs1">McLister, James (2003). 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"Scramble competition selects for greater tailfin size in male red-spotted newts (Amphibia: Salamandridae)". <i>Behavioral Ecology and Sociobiology</i>. <b>46</b> (6): 423–428. <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/1999BEcoS..46..423A">1999BEcoS..46..423A</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%2Fs002650050638">10.1007/s002650050638</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/0340-5443">0340-5443</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:34638425">34638425</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+and+Sociobiology&rft.atitle=Scramble+competition+selects+for+greater+tailfin+size+in+male+red-spotted+newts+%28Amphibia%3A+Salamandridae%29&rft.volume=46&rft.issue=6&rft.pages=423-428&rft.date=1999-11-04&rft_id=info%3Adoi%2F10.1007%2Fs002650050638&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A34638425%23id-name%3DS2CID&rft.issn=0340-5443&rft_id=info%3Abibcode%2F1999BEcoS..46..423A&rft.aulast=Able&rft.aufirst=David+J.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAmplexus" class="Z3988"></span></span> </li> <li id="cite_note-:9-15"><span class="mw-cite-backlink">^ <a href="#cite_ref-:9_15-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-:9_15-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-:9_15-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFVerrell1986" class="citation journal cs1">Verrell, Paul A. 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"Eye and clasper damage influence male mating tactics in the horseshoe crab, Limulus polyphemus". <i>Journal of Ethology</i>. <b>24</b> (1): 67–74. <a href="/wiki/Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1007%2Fs10164-005-0163-5">10.1007/s10164-005-0163-5</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/0289-0771">0289-0771</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:1398829">1398829</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+Ethology&rft.atitle=Eye+and+clasper+damage+influence+male+mating+tactics+in+the+horseshoe+crab%2C+Limulus+polyphemus&rft.volume=24&rft.issue=1&rft.pages=67-74&rft.date=2005-08-11&rft_id=https%3A%2F%2Fapi.semanticscholar.org%2FCorpusID%3A1398829%23id-name%3DS2CID&rft.issn=0289-0771&rft_id=info%3Adoi%2F10.1007%2Fs10164-005-0163-5&rft.aulast=Duffy&rft.aufirst=Erin+E.&rft.au=Penn%2C+Dustin+J.&rft.au=Botton%2C+Mark+L.&rft.au=Brockmann%2C+H.+Jane&rft.au=Loveland%2C+Robert+E.&rfr_id=info%3Asid%2Fen.wikipedia.org%3AAmplexus" class="Z3988"></span></span> </li> <li id="cite_note-Collette2012-20"><span class="mw-cite-backlink"><b><a href="#cite_ref-Collette2012_20-0">^</a></b></span> <span class="reference-text"><link rel="mw-deduplicated-inline-style" href="mw-data:TemplateStyles:r1238218222"><cite id="CITEREFColletteGass,_Kenneth_C.Hagadorn,_James_W.2012" class="citation journal cs1">Collette, Joseph H.; Gass, Kenneth C.; Hagadorn, James W. (May 2012). "<i>Protichnites eremita</i> Unshelled? 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