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	<title>Archetype &#187; Exoskeleton</title>
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	<description>Ant reconstruction one homology at a time</description>
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		<title>Homology Weekly: Clypeus</title>
		<link>http://roberto.kellerperez.com/2009/05/homology-weekly-clypeus/</link>
		<comments>http://roberto.kellerperez.com/2009/05/homology-weekly-clypeus/#comments</comments>
		<pubDate>Fri, 22 May 2009 17:42:05 +0000</pubDate>
		<dc:creator>Roberto Keller</dc:creator>
				<category><![CDATA[Ants]]></category>
		<category><![CDATA[Comparative Anatomy]]></category>
		<category><![CDATA[Homology Weekly]]></category>
		<category><![CDATA[Morphology]]></category>
		<category><![CDATA[Acanthoponera]]></category>
		<category><![CDATA[Cibarium]]></category>
		<category><![CDATA[Clypeus]]></category>
		<category><![CDATA[Exoskeleton]]></category>
		<category><![CDATA[Formica fusca]]></category>
		<category><![CDATA[Labrum]]></category>
		<category><![CDATA[Onychomyrmex]]></category>
		<category><![CDATA[Tetraponera aethiops]]></category>

		<guid isPermaLink="false">http://roberto.kellerperez.com/?p=984</guid>
		<description><![CDATA[When looking at an arthropod from our vertebrate perspective it is easy to forget that we are looking right at the animal&#8217;s skeleton. While our own vertebrate skeleton consists of a series of internal compact pieces with sponge-like cores that support an external layer of muscles and entrails (all nicely wrapped in skin), the reverse [...]]]></description>
			<content:encoded><![CDATA[<div id="attachment_1001" class="wp-caption aligncenter" style="width: 410px"><img class="size-full wp-image-1001" title="Tetraponera aethiops - head" src="http://roberto.kellerperez.com/wp-content/uploads/2009/05/tetraponera-aethiops-head3.jpg" alt="&lt;i&gt;Tetraponera aethiops&lt;/i&gt; worker showing the location of the clypeus in green (Scanning Electron Micrograph, Roberto Keller/AMNH)" width="400" height="459" /><p class="wp-caption-text">Tetraponera aethiops worker showing the location of the clypeus in green (Scanning Electron Micrograph, Roberto Keller/AMNH)</p></div>
<p>When looking at an <a href="http://en.wikipedia.org/wiki/Arthropod">arthropod</a> from our <a href="http://en.wikipedia.org/wiki/Vertebrate">vertebrate</a> perspective it is easy to forget that we are looking right at the animal&#8217;s skeleton. While our own vertebrate skeleton consists of a series of internal compact pieces with sponge-like cores that support an external layer of muscles and entrails (all nicely wrapped in skin), the reverse is true for arthropods. The arthropod skeleton consists of a series of external plates and hollow tubes that form enclosed spaces within which the internal musculature system attaches<sup class='footnote'><a href='#fn-984-1' id='fnref-984-1'>1</a></sup>. One consequence of this peculiar body architecture is that most of what we see on the outer surface of this <em>exoskeleton</em> is but a reflection of what is going on on the inside&#8211; minute external pits correspond to places where the cuticle folds in to form internal pillars, and innocent looking shallow furrows on the surface are large internal walls where powerful muscles originate. A simple examination of the exoskeleton, therefore, can tell us a lot about particular functions and consequently about an insect’s behavior.<span id="more-984"></span></p>
<div id="attachment_1004" class="wp-caption aligncenter" style="width: 460px"><img class="size-full wp-image-1004" title="Procryptocerus sp1 - clypeus" src="http://roberto.kellerperez.com/wp-content/uploads/2009/05/procryptocerus-sp1-clypeus.jpg" alt="The clypeus (in green) on the turtle ant &lt;i&gt;Procryptocerus&lt;/i&gt;, with a characteristic brush on its anterior border (Scanning Electron Micrograph, Roberto Keller/AMNH).  " width="450" height="337" /><p class="wp-caption-text">The clypeus (in green) on the turtle ant Procryptocerus, with a characteristic brush on its anterior border (Scanning Electron Micrograph, Roberto Keller/AMNH).  </p></div>
<p>A good example of this is provided by the <strong>clypeus</strong> in ants and its wide diversity of forms across the different species in the family. The clypeus corresponds to an unpaired skeletal plate lying right at the center of an insect face. It is normally located lower in the head just in front to where the antennae are inserted, but in many ant groups it quite commonly extends in between the antennal sockets. The anterior border of the clypeus is involved in two important articulations relating to the movement of the <a href="http://roberto.kellerperez.com/2009/04/homology-weekly-mouthparts/">mouthparts</a>. The central part forms a wide hinge with the mouth&#8217;s &#8220;lid&#8221; or labrum, allowing the latter to move forward and backwards to open and close the preoral cavity where the intricate ant tongue is stored when retracted (the actual opening of the mouth lies internally at the back-end of this preoral cavity). The sides of the clypeal border, on the other hand, form deep cavities where the anterior <a href="http://en.wiktionary.org/wiki/condyle">condyle</a> of each mandible articulate.</p>
<p>Those articulations occur externally. But what is going on the inside of the clypeus? The inner surface on the clypeus provides attachment to a set of muscles that originate right at the anterodorsal section of a special elastic chamber located just before the mouth known as <em>cibarium</em>. When these muscles contract the cibarium expands producing a suction action. It is basically the sucking pump of the insect, and the bigger the clypeus the bigger the pump muscles and the larger the sucking force. You may have probably noticed the big goofy snout in <a href="http://en.wikipedia.org/wiki/Cicada">cicadas</a>; well it is nothing but the hypertrophied clypeus attesting to the large sucking pump of these dedicated suckers. Ants never reach such extremes, but the clypeus can be quite large in some groups.</p>
<div id="attachment_1005" class="wp-caption aligncenter" style="width: 460px"><img class="size-full wp-image-1005" title="Onychomyrmex doddi - clypeus" src="http://roberto.kellerperez.com/wp-content/uploads/2009/05/onychomyrmex-doddi-head2.jpg" alt="Clypeus (in green) on a &lt;i&gt;Onychomyrmex doddi&lt;/i&gt; worker. Species in this genus display a convergent army ant like behavior (Scanning Electron Micrograph, Roberto Keller/AMNH)" width="450" height="337" /><p class="wp-caption-text">Clypeus (in green) on a Onychomyrmex doddi worker. Species in this genus display a convergent army ant like behavior (Scanning Electron Micrograph, Roberto Keller/AMNH)</p></div>
<p>In clades of chiefly predatory ants, like amblyoponines, the clypeus is never large and has become rather reduced in the more specialized genera like <em>Apomyrma</em> and <em>Onychomyrmex</em>. The same pattern occurs more or less among ponerines.</p>
<div id="attachment_1007" class="wp-caption aligncenter" style="width: 410px"><img class="size-full wp-image-1007" title="Formica fusca - clypeus" src="http://roberto.kellerperez.com/wp-content/uploads/2009/05/formica-fusca-head.jpg" alt="The large clypeus (in green) on a &lt;i&gt;Formica fusca&lt;/i&gt; worker (Scanning Electron Micrograph, Roberto Keller/AMNH)." width="400" height="533" /><p class="wp-caption-text">The large clypeus (in green) on a Formica fusca worker (left antenna removed. Scanning Electron Micrograph, Roberto Keller/AMNH).</p></div>
<p>Oh, but formicines and dolichoderines, those ants are such big suckers. Those are the ants you will most commonly see wandering between flowers looking for nectar and <a href="http://www.myrmecos.net/formicinae/ForInt11.html">tending aphids</a> for honeydew (that is, they suck ass big time<sup class='footnote'><a href='#fn-984-2' id='fnref-984-2'>2</a></sup>). Myrmecines ants have large clypeus in general, and it is not surprising to see a correlation between tending other insects and having a well developed clypeus in genera like <a href="http://www.myrmecos.net/myrmicinae/CreCer1.html"><em>Crematogaster</em></a>.</p>
<div id="attachment_1010" class="wp-caption aligncenter" style="width: 460px"><img class="size-full wp-image-1010" title="Labidus coecus - clypeus" src="http://roberto.kellerperez.com/wp-content/uploads/2009/05/labidus-coecus-clypeus.jpg" alt="The clypeus (in green, maybe) in a Neotropical army ant &lt;i&gt;Labidus coecus&lt;/i&gt; worker (is there, I swear. Scanning Electron Micrograph, Roberto Keller/AMNH)" width="450" height="337" /><p class="wp-caption-text">The clypeus (in green, maybe) in a Neotropical army ant Labidus coecus worker (the clypeus is there, I swear. Left antenna removed. Scanning Electron Micrograph, Roberto Keller/AMNH)</p></div>
<p>Now army ants, the ultimate specialized predators of the insect world, they are not suckers at all. All clades can be easily characterized by having almost no clypeus, so that the antennal sockets seem to fall off their heads forward.</p>
<div id="attachment_1013" class="wp-caption aligncenter" style="width: 460px"><img class="size-full wp-image-1013" title="Acanthoponera minor - clypeus" src="http://roberto.kellerperez.com/wp-content/uploads/2009/05/acanthoponera-minor-clypeus.jpg" alt="The swollen clypeus (in green) on an &lt;i&gt;Acanthoponera minor&lt;/i&gt; worker (Scanning Electron Micrograph, Roberto Keller/AMNH)" width="450" height="337" /><p class="wp-caption-text">The swollen clypeus (in green) on an Acanthoponera minor worker (left antenna removed. Scanning Electron Micrograph, Roberto Keller/AMNH)</p></div>
<p>One genus that intriges me is <em>Acanthoponera</em>. Species in this genus have a very large and swollen clypeus for what you will expect given the group&#8217;s phylogenetic position in between other major clades of ants. I don&#8217;t think much is known about the biology of this genus other than it is a nocturnal ant. But I bet you this ant is sucking around something.</p>
<p><strong>Notes</strong></p>
<div class='footnotes'>
<div class='footnotedivider'></div>
<ol>
<li id='fn-984-1'>The only enclosed cavity  formed by the skeleton in vertebrates is the cranium, but there are no muscles inside it. <span class='footnotereverse'><a href='#fnref-984-1'>&#8617;</a></span></li>
<li id='fn-984-2'>Sorry, just couldn&#8217;t resist. <span class='footnotereverse'><a href='#fnref-984-2'>&#8617;</a></span></li>
</ol>
</div>
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