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	<id>https://wiki.cas-group.net/index.php?action=history&amp;feed=atom&amp;title=Spandrel</id>
	<title>Spandrel - Revision history</title>
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	<link rel="alternate" type="text/html" href="https://wiki.cas-group.net/index.php?title=Spandrel&amp;action=history"/>
	<updated>2026-07-19T17:29:30Z</updated>
	<subtitle>Revision history for this page on the wiki</subtitle>
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	<entry>
		<id>https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=639&amp;oldid=prev</id>
		<title>Jfromm at 00:09, 11 January 2009</title>
		<link rel="alternate" type="text/html" href="https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=639&amp;oldid=prev"/>
		<updated>2009-01-11T00:09:35Z</updated>

		<summary type="html">&lt;p&gt;&lt;/p&gt;
&lt;table style=&quot;background-color: #fff; color: #202122;&quot; data-mw=&quot;interface&quot;&gt;
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				&lt;td colspan=&quot;2&quot; style=&quot;background-color: #fff; color: #202122; text-align: center;&quot;&gt;← Older revision&lt;/td&gt;
				&lt;td colspan=&quot;2&quot; style=&quot;background-color: #fff; color: #202122; text-align: center;&quot;&gt;Revision as of 19:09, 10 January 2009&lt;/td&gt;
				&lt;/tr&gt;&lt;tr&gt;&lt;td colspan=&quot;2&quot; class=&quot;diff-lineno&quot; id=&quot;mw-diff-left-l1&quot;&gt;Line 1:&lt;/td&gt;
&lt;td colspan=&quot;2&quot; class=&quot;diff-lineno&quot;&gt;Line 1:&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;−&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #ffe49c; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;&#039;&#039;&#039;Spandrel&#039;&#039;&#039; is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a [[Side-Effect|side-effect]] of an [[Adaptation|adaptation]], rather than arising from [[Natural Selection|natural selection]]. The term was coined by the Harvard paleontologist Stephen Jay Gould and population geneticist Richard Lewontin in their influential paper &quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&quot; (1979). They drew the analogy with spandrels in Renaissance architecture, which are curved areas of masonry above an arch, which they considered to be necessary side consequence arising from decisions concerned with the shape of the arch and the circumferential ring of the base of the dome, rather than being deliberately designed for direct utility in themselves. Properties that they singled out were the necessary number of four and their specific three-dimensional shape.&lt;/div&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;+&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #a3d3ff; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;&#039;&#039;&#039;Spandrel&#039;&#039;&#039; is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a [[Side-Effect|side-effect]] of an [[Adaptation|adaptation]], rather than arising from [[Natural Selection|natural selection]]&lt;ins style=&quot;font-weight: bold; text-decoration: none;&quot;&gt;. Spandrels are features of an organism which are not necessarily the product of an adaptive evolutionary process. They are traits carried along as a consequence of the architecture of organisms, like the spandrels in Gothic cathedral architecture&lt;/ins&gt;. The term was coined by the Harvard paleontologist Stephen Jay Gould and population geneticist Richard Lewontin in their influential paper &quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&quot; (1979). They drew the analogy with spandrels in Renaissance architecture, which are curved areas of masonry above an arch, which they considered to be necessary side consequence arising from decisions concerned with the shape of the arch and the circumferential ring of the base of the dome, rather than being deliberately designed for direct utility in themselves. Properties that they singled out were the necessary number of four and their specific three-dimensional shape.&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td colspan=&quot;2&quot; class=&quot;diff-side-deleted&quot;&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;+&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #a3d3ff; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt; &lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td colspan=&quot;2&quot; class=&quot;diff-side-deleted&quot;&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;+&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #a3d3ff; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt; &lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;br&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;br&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;== Links ==&lt;/div&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;== Links ==&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;/table&gt;</summary>
		<author><name>Jfromm</name></author>
	</entry>
	<entry>
		<id>https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=636&amp;oldid=prev</id>
		<title>Jfromm at 00:04, 11 January 2009</title>
		<link rel="alternate" type="text/html" href="https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=636&amp;oldid=prev"/>
		<updated>2009-01-11T00:04:22Z</updated>

		<summary type="html">&lt;p&gt;&lt;/p&gt;
&lt;table style=&quot;background-color: #fff; color: #202122;&quot; data-mw=&quot;interface&quot;&gt;
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				&lt;col class=&quot;diff-marker&quot; /&gt;
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				&lt;td colspan=&quot;2&quot; style=&quot;background-color: #fff; color: #202122; text-align: center;&quot;&gt;← Older revision&lt;/td&gt;
				&lt;td colspan=&quot;2&quot; style=&quot;background-color: #fff; color: #202122; text-align: center;&quot;&gt;Revision as of 19:04, 10 January 2009&lt;/td&gt;
				&lt;/tr&gt;&lt;tr&gt;&lt;td colspan=&quot;2&quot; class=&quot;diff-lineno&quot; id=&quot;mw-diff-left-l1&quot;&gt;Line 1:&lt;/td&gt;
&lt;td colspan=&quot;2&quot; class=&quot;diff-lineno&quot;&gt;Line 1:&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;−&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #ffe49c; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;&#039;&#039;&#039;Spandrel&#039;&#039;&#039; is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a [[Side-&lt;del style=&quot;font-weight: bold; text-decoration: none;&quot;&gt;effect&lt;/del&gt;|side-effect]] of an [[Adaptation|adaptation]], rather than arising from [[Natural Selection|natural selection]]. The term was coined by the Harvard paleontologist Stephen Jay Gould and population geneticist Richard Lewontin in their influential paper &quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&quot; (1979). They drew the analogy with spandrels in Renaissance architecture, which are curved areas of masonry above an arch, which they considered to be necessary side consequence arising from decisions concerned with the shape of the arch and the circumferential ring of the base of the dome, rather than being deliberately designed for direct utility in themselves. Properties that they singled out were the necessary number of four and their specific three-dimensional shape.&lt;/div&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;+&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #a3d3ff; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;&#039;&#039;&#039;Spandrel&#039;&#039;&#039; is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a [[Side-&lt;ins style=&quot;font-weight: bold; text-decoration: none;&quot;&gt;Effect&lt;/ins&gt;|side-effect]] of an [[Adaptation|adaptation]], rather than arising from [[Natural Selection|natural selection]]. The term was coined by the Harvard paleontologist Stephen Jay Gould and population geneticist Richard Lewontin in their influential paper &quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&quot; (1979). They drew the analogy with spandrels in Renaissance architecture, which are curved areas of masonry above an arch, which they considered to be necessary side consequence arising from decisions concerned with the shape of the arch and the circumferential ring of the base of the dome, rather than being deliberately designed for direct utility in themselves. Properties that they singled out were the necessary number of four and their specific three-dimensional shape.&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;br&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;br&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;== Links ==&lt;/div&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;== Links ==&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;/table&gt;</summary>
		<author><name>Jfromm</name></author>
	</entry>
	<entry>
		<id>https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=635&amp;oldid=prev</id>
		<title>Jfromm at 00:03, 11 January 2009</title>
		<link rel="alternate" type="text/html" href="https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=635&amp;oldid=prev"/>
		<updated>2009-01-11T00:03:17Z</updated>

		<summary type="html">&lt;p&gt;&lt;/p&gt;
&lt;table style=&quot;background-color: #fff; color: #202122;&quot; data-mw=&quot;interface&quot;&gt;
				&lt;col class=&quot;diff-marker&quot; /&gt;
				&lt;col class=&quot;diff-content&quot; /&gt;
				&lt;col class=&quot;diff-marker&quot; /&gt;
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				&lt;td colspan=&quot;2&quot; style=&quot;background-color: #fff; color: #202122; text-align: center;&quot;&gt;← Older revision&lt;/td&gt;
				&lt;td colspan=&quot;2&quot; style=&quot;background-color: #fff; color: #202122; text-align: center;&quot;&gt;Revision as of 19:03, 10 January 2009&lt;/td&gt;
				&lt;/tr&gt;&lt;tr&gt;&lt;td colspan=&quot;2&quot; class=&quot;diff-lineno&quot; id=&quot;mw-diff-left-l6&quot;&gt;Line 6:&lt;/td&gt;
&lt;td colspan=&quot;2&quot; class=&quot;diff-lineno&quot;&gt;Line 6:&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;* Stephen Jay Gould and Richard C. Lewontin. [http://www.aaas.org/spp/dser/03_Areas/evolution/perspectives/Gould_Lewontin_1979.shtml &amp;quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&amp;quot;] &amp;#039;&amp;#039;Proc. Roy. Soc. London B&amp;#039;&amp;#039; &amp;#039;&amp;#039;&amp;#039;205&amp;#039;&amp;#039;&amp;#039; ([http://faculty.washington.edu/lynnhank/GouldLewontin.pdf 1979]) pp. 581-598&lt;/div&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;* Stephen Jay Gould and Richard C. Lewontin. [http://www.aaas.org/spp/dser/03_Areas/evolution/perspectives/Gould_Lewontin_1979.shtml &amp;quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&amp;quot;] &amp;#039;&amp;#039;Proc. Roy. Soc. London B&amp;#039;&amp;#039; &amp;#039;&amp;#039;&amp;#039;205&amp;#039;&amp;#039;&amp;#039; ([http://faculty.washington.edu/lynnhank/GouldLewontin.pdf 1979]) pp. 581-598&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;* Stephen Jay Gould (1997). [http://www.pnas.org/cgi/content/full/94/20/10750 &amp;quot;The exaptive excellence of spandrels as a term and prototype&amp;quot;] &amp;#039;&amp;#039;Proceedings of the National Academy of Sciences USA&amp;#039;&amp;#039;. 94: 10750-10755.&lt;/div&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot;&gt;&lt;/td&gt;&lt;td style=&quot;background-color: #f8f9fa; color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #eaecf0; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;* Stephen Jay Gould (1997). [http://www.pnas.org/cgi/content/full/94/20/10750 &amp;quot;The exaptive excellence of spandrels as a term and prototype&amp;quot;] &amp;#039;&amp;#039;Proceedings of the National Academy of Sciences USA&amp;#039;&amp;#039;. 94: 10750-10755.&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td colspan=&quot;2&quot; class=&quot;diff-side-deleted&quot;&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;+&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #a3d3ff; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;&lt;ins style=&quot;font-weight: bold; text-decoration: none;&quot;&gt;&lt;/ins&gt;&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;tr&gt;&lt;td colspan=&quot;2&quot; class=&quot;diff-side-deleted&quot;&gt;&lt;/td&gt;&lt;td class=&quot;diff-marker&quot; data-marker=&quot;+&quot;&gt;&lt;/td&gt;&lt;td style=&quot;color: #202122; font-size: 88%; border-style: solid; border-width: 1px 1px 1px 4px; border-radius: 0.33em; border-color: #a3d3ff; vertical-align: top; white-space: pre-wrap;&quot;&gt;&lt;div&gt;&lt;ins style=&quot;font-weight: bold; text-decoration: none;&quot;&gt;[[Category:Evolutionary Principles]]&lt;/ins&gt;&lt;/div&gt;&lt;/td&gt;&lt;/tr&gt;
&lt;/table&gt;</summary>
		<author><name>Jfromm</name></author>
	</entry>
	<entry>
		<id>https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=634&amp;oldid=prev</id>
		<title>Jfromm: New page: &#039;&#039;&#039;Spandrel&#039;&#039;&#039; is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a side-effect of an [[...</title>
		<link rel="alternate" type="text/html" href="https://wiki.cas-group.net/index.php?title=Spandrel&amp;diff=634&amp;oldid=prev"/>
		<updated>2009-01-11T00:02:48Z</updated>

		<summary type="html">&lt;p&gt;New page: &amp;#039;&amp;#039;&amp;#039;Spandrel&amp;#039;&amp;#039;&amp;#039; is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a &lt;a href=&quot;/index.php?title=Side-effect&amp;amp;action=edit&amp;amp;redlink=1&quot; class=&quot;new&quot; title=&quot;Side-effect (page does not exist)&quot;&gt;side-effect&lt;/a&gt; of an [[...&lt;/p&gt;
&lt;p&gt;&lt;b&gt;New page&lt;/b&gt;&lt;/p&gt;&lt;div&gt;&amp;#039;&amp;#039;&amp;#039;Spandrel&amp;#039;&amp;#039;&amp;#039; is a term used in evolutionary biology describing a phenotypic characteristic that is considered to have developed during evolution as a [[Side-effect|side-effect]] of an [[Adaptation|adaptation]], rather than arising from [[Natural Selection|natural selection]]. The term was coined by the Harvard paleontologist Stephen Jay Gould and population geneticist Richard Lewontin in their influential paper &amp;quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&amp;quot; (1979). They drew the analogy with spandrels in Renaissance architecture, which are curved areas of masonry above an arch, which they considered to be necessary side consequence arising from decisions concerned with the shape of the arch and the circumferential ring of the base of the dome, rather than being deliberately designed for direct utility in themselves. Properties that they singled out were the necessary number of four and their specific three-dimensional shape.&lt;br /&gt;
&lt;br /&gt;
== Links ==&lt;br /&gt;
&lt;br /&gt;
* Wikipedia Entry for [http://en.wikipedia.org/wiki/Spandrel_(biology) Spandrel]&lt;br /&gt;
* Stephen Jay Gould and Richard C. Lewontin. [http://www.aaas.org/spp/dser/03_Areas/evolution/perspectives/Gould_Lewontin_1979.shtml &amp;quot;The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme&amp;quot;] &amp;#039;&amp;#039;Proc. Roy. Soc. London B&amp;#039;&amp;#039; &amp;#039;&amp;#039;&amp;#039;205&amp;#039;&amp;#039;&amp;#039; ([http://faculty.washington.edu/lynnhank/GouldLewontin.pdf 1979]) pp. 581-598&lt;br /&gt;
* Stephen Jay Gould (1997). [http://www.pnas.org/cgi/content/full/94/20/10750 &amp;quot;The exaptive excellence of spandrels as a term and prototype&amp;quot;] &amp;#039;&amp;#039;Proceedings of the National Academy of Sciences USA&amp;#039;&amp;#039;. 94: 10750-10755.&lt;/div&gt;</summary>
		<author><name>Jfromm</name></author>
	</entry>
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