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Foreword Fossil Birds Foreword Fossil Birds Introduction Today, there are estimated to be some 9800 species of birds on Planet Earth. This is only a window on the evolution of the class Aves. With those known in the fossil record, it can be increased to about 12,000 species, though about a third of these fossil species come from unknown lineages. Pierce Brodkorb estimated in 1959 that about 1,634,000 species of bird had existed over the 150 million years of avian evolution. We therefore know of well under 1%. However, our understanding of bird evolution is improving, particularly with the discovery of such fossil-rich sites as the Creta- ceous deposits of northern China and the early Tertiary deposits of the Messel and Quercy, in Europe, and the Green River Formation of Wyoming, North America, while fossil collecting in other areas, such as Australia, has added to our knowledge of the diversity of the birds that once existed. What follows is a very brief summary of avian diversity, starting with the basal Urvögel, moving through the early Pygostylia and other non-carinate species, before ending with a broad survey of the modern birds— the Neornithes—which include all the bird species alive today. Before beginning, special mention should perhaps be made of certain fossils that are regarded as contentious. Of particular significance is one such fossil, Protoavis texensis, as this species was hailed by its discoverer to be the primordial bird (Chatterjee 1994, 1995). Originating in the Triassic, it would have knocked the most famous of all basal birds, Archaeopteryx, off its pedestal as the oldest bird, predating it by over 100 million years. However, severe doubt has been cast as to its avian status, with indica- tions that it is actually an early coelurosaur—just as significant palaeontologically, but not as a bird (Witmer 2002). Archaeopteryx is still holding onto its title of ‘the oldest fossil bird’—for now. Time chart showing the major Eon Era Period Epoch Age (Mya) eons, eras, periods and epochs, with ages (after the UNESCO Phanerozoic Cenozoic Quaternary Holocene 000·01–000·00 International Stratigraphic Chart). Pleistocene 001·64–000·01 Added information includes the timespan of birds. Neogene Pliocene 005·20–001·64 Miocene 023·50–005·20 Oligocene 035·50–023·50 Paleogene Eocene 056·50–035·50 Tertiary Paleocene 065·00–056·50 Birds Mesozoic Cretaceous Late 097·00–065·00 Early 146·00–097·00 Jurassic Late 157·00–146·00 Middle 178·00–157·00 Early 208·00–178·00 Triassic 245·00–208·00 Paleozoic Permian 295·00–245·00 Carboniferous 355·00–295·00 Devonian 410·00–355·00 Silurian 435·00–410·00 Ordovician 500·00–435·00 Cambrian 540·00–500·00 Proterozoic 2500·00–540·00 Archean 3800·00–2500·00 Precambrian 002-foreword-HBW-12.P65 11 23/07/2007, 10:21 12 Avian Origins In 1861, a chicken-sized fossil from Bavaria turned the scientific world on its head. Archaeopteryx, Confuciusornis and relatives Named Archaeopteryx lithographica, this animal had the feathers and wings of a bird, (the ‘dawn birds’) 10–15 species, 9–10 genera but the body, tail and jaws of a reptile—even its wings had hands and claws, and so it was hailed as ‘proof’ that birds evolved from reptiles. It was not until much later that the true significance of Archaeopteryx was revealed, when comparisons were made with those of the coelurosaurian dinosaur Compsognathus. Indeed, so similar are these fossils that without its feather impressions, Archaeopteryx could have been regarded as just another compsognathid dinosaur, if a rather odd one. The discovery of Archaeopteryx spurred a debate over the origins of the class Aves that lasted until the late 1990s, when several small dinosaurs, from the primitive Sinosauropteryx to the dromaeosaurid Microraptor, were found in sites across northern China. Each dino- saur was covered by integuments that were obviously feathers (Chen et al. 1998, Xu, Compsognathus Zhou & Wang 2000, Xu, Zhou, Wang, Kuang et al. 2003). Thus, possession of feath- ers is not indicative of being a bird, but is a primitive (‘plesiomorphic’) state within the coelurosaurian theropods; even primitive relatives of Tyrannosaurus had proto- feathers (Xu et al. 2004). Rather, it is the possession of flight, contour and down feathers, and the combination of a variety of other skeletal modifications that separate birds from their closest relatives within the Dinosauria. Archaeopteryx and its kin still act as a morphological bridge, bearing a mosaic of primitive reptilian features such as teeth and the long bony tail without a pygostyle alongside more modern bird-like features, including asymmetrical flight feathers on the wings. The more we find out Sinosauropteryx about dinosaurs such as Microraptor and its relatives, the more we realise that the distinction between birds and dinosaurs is an arbitrary line based on no single, unique feature. We know that the pygostyle is not a uniquely avian feature either and is found in a number of oviraptorosaurs as well (e.g. Barsbold et al. 2000). We know that the wishbone, or ‘furcula’, is found in many theropods, including Tyrannosaurus, one of the most advanced members of that group. There is also evidence that pneumaticisation of the bone—the evolution of hollow bones supported by internal struts—is a theropod characteristic, not a purely avian one. However, no non-avian theropod found so far has all these traits in one skeleton, while the fusion of the trunk vertebrae and the The primitive Tyrannosaurid supporting struts of the ribs are uniquely avian. The closest relatives of the birds, the Dilong Dromaeosauridae, possess a suite of other features that can shed light on how birds themselves came into being. If we take a closer look at the arboreal Microraptor gui, for example, we find that not only does this tiny dinosaur have flight feathers, but it has them on all four legs. These feathers are asymmetrical in form, like those of mod- ern birds. Yet, this is a bona fide dinosaur, not a bird. Microraptor has given credence to the ‘tree-down’ hypothesis of bird origins, which had floundered through the lack of examples of arboreal dinosaurs. Until Microraptor was discovered, the alternative theory, the ‘ground-up’ concept, was gaining favour, despite there being so many Tyrannosaurus problems as to how this could be the accurate scenario, given that all modern species of flying and gliding organisms are plesiomorphically arboreal. It has since been sug- gested that the initiation to flying came through these tiny animals jumping about in the branches of the trees of their forest home, with the hindwings forming feathered ‘edges’, which aided lift. The four-winged approach to gliding in this dromaeosaurid came as a complete surprise and led to the re-examination of Archaeopteryx and later forms (Christiansen & Bonde 2004, Zhang & Zhou 2004), which revealed that their The primitive Oviraptorosaur leg feathers still bore some similarities with those of M. gui rather than with the more Caudipteryx typical contour feathers that we see today on the legs of birds. Although there are still several opponents to the ‘birds-as-dinosaurs’ theory, general opinion is that the evi- dence for birds being descendents of theropod dinosaurs is overwhelming. Despite all of the recent discoveries in China, the Archaeopterygidae are still re- garded as the oldest known group of birds, although they are not regarded as directly ancestral. There are still a large suite of reptilian features in the skeleton, including a reptilian physiology, a ‘reversed’ pelvis more similar to their Saurischian ancestors, Velociraptor, an unfused rib cage, and a long, bony tail. Also unlike modern birds, the most notable a typical Dromaeosaur feature is the lack of a horny beak. The recent description of a tenth specimen has revealed more about Archaeopteryx and may throw our interpretation of the evolution of birds through a new twist: this example reveals features not shown in the others, clarifying the position of the four toes in relation to each other and, more amazingly, the presence of a ‘switch-claw’, otherwise found only in the dromaeosaurs and the fossil bird Rahonavis. Cladistic study also indicates that this discussion on fossil birds should include the dromaeosaurs and the related troodontids as both would be classed as members of the Aves if this is defined as the clade including the ancestor of Archaeopteryx and the birds (Mayr, Pohl & Peters 2005, Mayr, Pohl, Hartman & Peters 2007), rather than lying outside the group phylogenetically (as per Sereno 1999), Troodon 002-foreword-HBW-12.P65 12 23/07/2007, 10:21 13 although there is still debate as to whether this is an accurate interpretation (Corfe & Butler 2006, Mayr & Peters 2006). Archaeopteryx itself had poorly-developed flight muscles and could not fly as well as modern birds. Its long tail shifted its centre of balance behind the wings, again unlike modern species. Its claws tell us two different tales. Those on the wings were adapted primarily for climbing, while those on the hindlimbs suggest a ground-based animal. Current consensus is that Archaeopteryx was primarily terrestrial, but may also have been a facultative percher, if it perched at all (Mayr et al. 2005). Archaeopteryx inhabited a coastal scrubland of low-growing conifers, cycads and similar species, intermingled with more tree-like species such as firs and ginkgos. This vegetation was concentrated around shallow coastal lagoons that abounded in Europe in the Late Jurassic. It may be that, at the coastal edge, as around Solnhofen, the habitat was more open than that known to exist further ‘inland’ 10 cm during the period. Archaeopteryx may have utilised its limited flight capabilities in the open spaces, adopting a ground-hugging flight pattern (O’Farrell et al.
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