The Macro Library^(beta)
A collection of macroevolutionary examples for K-12 teachers, students, and publishers

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The Origin of Birds

The origin of birds has long been considered a hallmark of the success of evolutionary theory. Just a two years after Darwin's work was published, the first Archaeopteryx was found in the Solnhofen limestone, having clear feather impressions but at the same time, many dinosaurian characteristics, seemingly an irrefutable confirmation of Darwin's theory¹. Things became even more exciting when John Ostrom found Deinonychus antirrhopus in 1962², and his 1969 anlysis prompted the 1973 landmark paper suggesting that, in fact, birds are living

dinosaurs³. However, this very significant claim needs much good evidence and careful examination, for it is easy to make several fallacious inferences and, in fact, a few paleontologists have rejected the theropod ancestry of dinosaurs in favor of as-yet undiscovered "thecodont" ancestor, which was particularly reinvigorated in the 1990's with a text by Alan Feduccia⁴. In this article we will not address this alternative theory, which to date has largely lacked a

basis and is mostly an opposition argument, and relies on several problematic (and in some cases, outdated) claims^5[1]. Indeed, the theropod descent theory is so well supported that the issue is often not just the question of how less derived dinosaurs are to birds^6[2] but in some cases the question instead becomes if some other dinosaur groups were, in fact, more derived than birds, such as the Alvarezsaurs, and finds such as those in Figure 1 only work to blur the lines further.

Figure 1. Microraptor gui. From Fastovsky et al. 2005. (GET PERMISSION) Modified with permission.

   What we would most closely like to analyze, then, is how similar are many of these dinosaurs to birds? It would appear in the fossil record that theropods converged on various birdlike characteristics several times. In fact, Tyrannosaurus rex had a

^7[3]! Furthermore, the presence of

rib segments and ossified sternum in

and oviraptorids suggest that many of the definitive skeletal features we traditionally associate with birds in fact originated seperately from flight, and was only later co-opted in the process known as

⁸. However, soft tissues would appear to be increasingly unsatisfactory in use as an evaluation of "bird-ness". Some analyses even find reason to believe that feathers can be found on all but the most basal

with even a light coat of feathers potentially covering T. rex, at least when it was young. Feathers, then, were not an adaptation for flight, with evidence for them arising very early, as shown in Figure 2.

Figure 2. Cladogram depicting how different types of feathers arose in coelurosaurs. Modified after Fastovsky et al. 2005 with permission. (GET PERMISSION)

This is even more strongly supported by finds such as that of a brooding Oviraptor⁹, in which case feathers merely for self-insulation may have been less of a requirement than the ability to thermoregulate eggs, much as modern birds today. Even some one of the most difficult features of birds to find in the fossil record, its unique air-sac system, may be a relatively basal development for saurischians in general¹⁰.

   With so many avian character scattered throughout the fossil record of saurischians, it is evident that the primary arguments against the theropod hypothesis can't be well-established in fossil characters, and must instead lie elsewhere. One common argument is that the timing of the fossil record is at odd with the earliest appearance of birds. In particular, many of the most well-known relatives of birds^[4] do not appear until the Cretaceous — for example, Velociraptor is perhaps 70 MY younger than Archaeopteryx. However, this is not the problem it would first appear to be. There are other well-known dromaeosaurids such as Utahraptor that are only 25 MY younger than Archaeopteryx, less well-known ones such as Sinovenator that are 17 MY younger, and fragmentary speciemens that may be as much as 20 MY older than Archaeopteryx — and thus this argument is rendered ineffectual with a larger sampling size¹¹. Furthermore, these gaps actually compare favorably to other gaps that are present and considered of little importance in the fossil record. Other contrary arguments focus on perhaps poor homologies of parts of the skeleton^[5]. However, most of these homologies have been well-established, such as the semilunate carpal — what was called the "half-moon shaped bone in the wrist" in the movie Jurassic Park — as shown in Figure 3

Figure 3. Semi-lunate carpal in the left hand of Ingenia, an oviraptorosaurian maniraptoriform. Modified after Fastovsky et al 2005 with permission. (GET PERMISSION)

. The other contrary arguments often levelled focus on digit homlogy (in what amounts to semantics between diciplines) and in the methods of lung ventilation, but the treatment of both can be extensive, so we refer to the cited literature for a more in-depth discussion.

   The final aspect of bird evolution to consider is, of course, the matter of flight. The advantages of flight are readily apparent^[6], but the evolution of an evidently terrestrial animal such as more basal theropods into an animal capable of flight is a question that has been actively debated within paleontology. While clear and persuasive arguments have been made against using Archaeopteryx too much and relying on it as the model for the transition to flight and treating it as the de-facto direct ancestor of birds, it is nevertheless a useful model organism so long as one is aware that it is indeed just a model that came from a yet more primitive proto-bird, and almost certainly was not the direct ancestor of modern birds¹². Furthermore, it is important to note that the "origin of flight" and the "origin of birds" may be linked, but they did not necessarily evolve concurrently and treating it as such could lead to many potentially fallacious inferences. Thus, for the two competing theories of

and

the fact that Archaeopteryx was apparently a terrestrial forager and escape climber (as shown in Figure 4)

Figure 4. The escape behavior of Archaeopteryx as reconstructed from the locomotor adaptations and universal functional interdependencies. The launching "perch" is assumed not to be predator-safe. Image modified after Elzanowski 2002, with permission.

does not necessarily impose either/both conditions on the direct ancestor of birds or on the origin of flight itself¹³. There are relative merits to both positions. The cursorial approach is perhaps easier to reconcile with what we currently have of the fossil record, which indicates that as a trend, dromaeosaurids shrank as they approached the K-Pg^[7] boundary. However, as the fossil record is very incomplete, and only fragmentary Jurassic dromaeosaurid fossils have been found, using fossils to justify this approach is somewhat fallacious, and the theory has to stand on its merit in a purely theoretical regime. The cursorial approach hypotheses that the origin of flight originated with the flight-stroke of birds, but highly modified. The arm structures of dromaeosaurids and related dinosaurs are very good for jackknifing forward and grasping. Since it is known that dromaeosaurids had flight feathers¹⁴, this action could have provided some degree of lift during predatory strikes, enabling long leaps; been used to capture insects; been used for display purposes, etc.

   In contrast, while arboreality may be a bit more difficult to reconcile with the fossil record at first glance, there are reasons to support this view. Some studies suggest, for example, that dromaeosaurid "killer claws" were in fact best suited as climbing instruments, and that their shape and curvature were nonideal for cutting and tearing flesh. This is further supported by the short but sharp claws that many early birds still retained, which would also assist in climbing. However, selecting either arboreality or cursoriality as the "true" and final answer may be a false dichotomy. There is no reason to suppose that flight did not originate in some hybrid of these two approaches, much like the proposed model of Archaeopteryx in Figure 4. Also, while it goes against the generally-accepted axiom of

there is no reason why flight could not have independently evolved in the bird line more than once. While the origin of birds is fairly uncontroversial, the associated but independent question of the origin of bird flight has yet to be resolved, and may never be definitively answered.

Philip Kahn
University of California, Berkeley
Submitted August 29 2008

• [1] For example, the second edition of Feduccia's text claims "Many authors do not distinguish between carnosaurs and coelurosaurs, attributing the differences to size alone rather than to a phyletic split." However, as early as 1984-86 in studies by Gauthier, it was known that large forms such as T. rex could be coelurosaurs.
• [2] Most cladograms will have birds as a terminal branch of theropods, with no "more derived" theropods existing outside of Avialae
• [3] In many ways, the furcula of T. rex was more derived than that of Velociraptor. For more information, see the cited paper.
• [4] Here, we are referring to "maniraptoriformes".
• [5] By this, we mean the equivalent of questioning if what we call the humerus in a whale fin is the equivalent bone to a human humerus
• [6] Predator evasion, new predatory niches, etc.
• [7] K-Pg is now preferentially used to K-T to describe the extinction event approximately 66 million years ago, since the "Tertiary" is not a proper geological time. "K" is used to express Cretaceous to avoid confusion with other "C" periods, such as the Cambrian or Carboniferous.

1. Fastovsky et. al. The Evolution and Extinction of the Dinosaurs. 2005. ISBN 0-521-81172-4
2. Glut. Deinonychus. Dinosaurs: The Encyclopedia. 1997. ISBN: 0-89950-917-7
3. Witmer. The Debate on Avian Ancestry in Mesozoic Birds: Above the Heads of Dinosaurs. 2002. ISBN 0-520-20094-2
4. Feduccia. The Origin and Evolution of Birds. 1999. ISBN 0-300-07861-7.
5. Witmer. The Debate on Avian Ancestry in Mesozoic Birds: Above the Heads of Dinosaurs. 2002. ISBN 0-520-20094-2
6. Benton. Vertebrate Paleontology. 2005. ISBN 0-632-05637-1
7. Larson et al. Furcula of Tyrannosaurus rex.. In The Carnivorous Dinosaurs. 2005. ISBN 0-253-34539-1.
8. Clark et al. Cladistic Approaches to Bird Relationships, in Mesozoic Birds: Above the Heads of Dinosaurs. 2002. ISBN 0-520-20094-2
9. Norell et al. A nesting dinosaur. Nature, 1995.
10. Wedel. Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs. Paleobiology 2003.
11. Witmer. The Debate on Avian Ancestry in Mesozoic Birds: Above the Heads of Dinosaurs. 2002. ISBN 0-520-20094-2
12. Clark et al. Cladistic Approaches to Bird Relationships. In Mesozoic Birds: Above the Heads of Dinosaurs. 2002. ISBN 0-520-20094-2
13. Elzanowski. Archaeopterygidae. In Mesozoic Birds: Above the Heads of Dinosaurs. 2002. ISBN 0-520-20094-2
14. Turner et al. Feather Quill Knobs in the Dinosaur Velociraptor. Science 2007.