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A field guide to mesozoic birds and other winged dinosaurs

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feather fossils have been reported by Vinther & al. (2008), <strong>and</strong> it is sometimes<br />

apparent even <strong>to</strong> the naked eye. Structurally colored feathers have<br />

been recognized by a distinct arrangement where a thin layer of densely<br />

aligned melanin granules overlies a looser conglomerate of melanin. This<br />

can be seen even if the overlying keratin scattering layer has degraded (Vinther<br />

& al. 2008). This arrangement where structure is produced by the arrangement<br />

of melanin as well as ‘bubbles’ in the keratin layer is notably<br />

found in the dazzling iridescent plumage of humming<strong>birds</strong> (Prum, 2006).<br />

The mechanics of structural color in feathers have implications for<br />

how extinct bird species are res<strong>to</strong>red in art. Blue, green, jet black <strong>and</strong> bright<br />

white can’t be present in <strong>birds</strong> that lack structural color in their feathers.<br />

Structural colors may or may not have been possible in the monofilament<br />

feathers of some primitive coelurosaurs <strong>and</strong> ornithischians. Note that<br />

structural coloration is never observed in the monofilament hair of modern<br />

mammals. The primary difference between hair <strong>and</strong> simple feathers, however,<br />

isn’t the macrostructure of the filaments, but the microstructure of the<br />

underlying molecules. Hair is composed of alpha-keratin, a helix-shaped<br />

molecule like DNA. Beta-keratin, which makes up feathers, has a layered<br />

<strong>and</strong> pleated underlying molecular structure more conducive <strong>to</strong> scattering<br />

light. On the <strong>other</strong> h<strong>and</strong>, in all of the iridescent fossil feathers studied by<br />

Vinther & al. (2008), the structural color was restricted <strong>to</strong> the barbules,<br />

which are not present in many primitive feathered <strong>dinosaurs</strong>. Additionally,<br />

structural colors are not observed in modern plumulaceous feathers (down)<br />

or in the downy after-feathers of <strong>other</strong>wise structurally colored pennaceous<br />

feathers. It is therefore likely that blue, green, iridescent or vivid downy <strong>and</strong><br />

monofilament feathers were extremely rare, if they existed at all in Mesozoic<br />

<strong>birds</strong> <strong>and</strong> more primitive feathered <strong>dinosaurs</strong>.<br />

The vast majority of bird colors are due in whole or in part <strong>to</strong> pigmentation,<br />

or lack thereof (Stettenheim, 2000). There are several different kinds<br />

of pigments, with the two most common being melanins <strong>and</strong> carotenoids.<br />

Melanins are easily identified in fossil feathers, <strong>and</strong> their shape <strong>and</strong><br />

concentration can indicate what color they produced. Melanins are responsible<br />

for black (though not deep, solid black, which requires the addition<br />

of structural color), gray, <strong>and</strong> a wide variety of browns <strong>to</strong> rufous orange<br />

or rusty red colors. A lack of melanin will produce white, as evidenced by<br />

albino specimens. Note that some albino <strong>birds</strong> <strong>to</strong>day are not completely<br />

white, but retain some darker coloration due <strong>to</strong> the structural colors of the<br />

feathers which are not erased by an absence of melanin in all cases.<br />

Carotenoids are, by <strong>and</strong> large, what give <strong>birds</strong> their characteristically<br />

bright colors. Carotenoids cannot be directly synthesized by the body<br />

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