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A field guide to mesozoic birds and other winged dinosaurs

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fined Aves the way Mammalia came <strong>to</strong> be defined--as a “crown group”, that<br />

is, the group containing all modern bird groups <strong>and</strong> any species closer <strong>to</strong><br />

them than <strong>to</strong> more distantly related prehis<strong>to</strong>ric lineages. This use of Aves<br />

preserves Linnaeus’ usage (he didn’t know of any prehis<strong>to</strong>ric <strong>birds</strong>), but<br />

also excludes fossil bird groups such as Archaeopteryx <strong>and</strong> even those very<br />

similar <strong>to</strong> modern <strong>birds</strong> like Ichthyornis. Subsequently, attempts were made<br />

<strong>to</strong> re-define Aves <strong>to</strong> include Archaeopteryx for his<strong>to</strong>rical reasons; after all,<br />

it has always been considered the “first bird”. Critics called these attempts<br />

arbitrary: why include Archaeopteryx <strong>and</strong> not their closest relatives (such<br />

as Microrap<strong>to</strong>r), which were more like modern <strong>birds</strong> in some respects? As<br />

of this writing, a body known as the International Society for Phylogenetic<br />

Nomenclature is being organized <strong>to</strong> govern the naming of clades, <strong>and</strong> it<br />

seems likely that it will adopt the crown group usage as official.<br />

When this usage is officially adopted, it will leave most Mesozoic<br />

<strong>birds</strong> outside of Aves. This will only require a semantic change; true “<strong>birds</strong>”<br />

will be restricted <strong>to</strong> the group of modern-style <strong>birds</strong> only. Non-avian members<br />

of the lineage leading <strong>to</strong> modern <strong>birds</strong> such as Ichthyornis will be considered<br />

“stem-<strong>birds</strong>” (<strong>and</strong>, somewhat ironically, also a “non-avian dinosaur”<br />

genus, as all <strong>birds</strong> are included in the clade Dinosauria).<br />

So what is a “bird”? As a vernacular rather than scientific term, this is<br />

a matter of loose convention rather than scientific precision. Most common<br />

definitions of the word in English hinge on certain key aspects: egg-laying,<br />

feathers, <strong>and</strong> flight. If this usage is adopted, Archaeopteryx may still be considered<br />

<strong>birds</strong>, depending on their controversial flight abilities (though they<br />

were almost certainly at least glissant, i.e. capable of passive gliding). Microrap<strong>to</strong>r<br />

are probably “<strong>birds</strong>” as well under this definition. Of course, not<br />

all <strong>birds</strong> fly, <strong>and</strong> just as there are ground <strong>birds</strong> <strong>and</strong> flightless <strong>birds</strong> <strong>to</strong>day,<br />

many Mesozoic lineages lost flight, or evolved before flight. Therefore, any<br />

<strong>dinosaurs</strong> which would now be included by one of the first definitions proposed<br />

for the clade Aves, by Allan Charig in 1985, are featured in this book.<br />

Charig’s definition of Aves linked it <strong>to</strong> the presence of feathers. While he did<br />

not specify what he meant by feathers, for the purposes of this book I am<br />

using a conservative interpretation: feathers with a fully modern ana<strong>to</strong>my,<br />

consisting of a rachis (central “quill”), <strong>and</strong> a vane comprised of barbs linked<br />

<strong>to</strong>gether by barbules (see diagram on p. 39). This is also the same characteristic<br />

that was used <strong>to</strong> classify Archaeopteryx as <strong>birds</strong> in the first place, so it<br />

seems most appropriate for use in this work. This definition is equivalent <strong>to</strong><br />

the modern clade Aviremigia, that is, all <strong>winged</strong> <strong>dinosaurs</strong>.<br />

By their nature, feathers do not fossilize well, nor often, so there<br />

may certainly be some species or even entire lineages which were “<strong>birds</strong>”<br />

under the above definition, but are excluded here because we do not have<br />

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