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Archaeopteryx

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Archaeopteryx
Temporal range: Late Jurassic
A model of Archaeopteryx lithographica
on display at the Oxford University Museum
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Aves
Order:
Archaeopterygiformes
Family:
Archaeopterygidae

Huxley, 1871
Genus:
Archaeopteryx
Species:
A. lithographica
Binomial name
Archaeopteryx lithographica
Meyer, 1861
Synonyms

See below

Archaeopteryx (from Ancient Greek ἀρχαῖος archaios meaning 'ancient' and πτέρυξ pteryx meaning 'feather' or 'wing'; pronounced [ɑː(ɹ)kɪˈɒptəɹɪks]) is the earliest and most primitive known avian to date. It lived in the late Jurassic Period around 155-150 million years ago in what is now southern Germany. In German, Archaeopteryx is known as Urvogel, a German word meaning "protobird" or "first bird". Although the name originated in Germany, it is also used in English-speaking countries.

At the time Archaeopteryx lived, Europe was an archipelago of islands in a shallow warm tropical sea, much closer to the equator than it is now. Archaeopteryx lived during the time of the dinosaurs, yet was set apart from them because of the inclusion of both avian and theropod dinosaur features. Similar in size and shape to a European Magpie, it bore broad, rounded wings and a long tail. Archaeopteryx could grow to about half a metre, or 1.6 feet in length. Its feathers resembled those of modern birds but Archaeopteryx was rather different from any bird known today, in that it had jaws lined with sharp teeth, three 'fingers' ending in curved claws and a long bony tail. These features, which are consistent with theropod dinosaurs, have made the Archaeopteryx a hot topic in the debate on evolution. Indeed, in 1862 the description of the first intact specimen of Archaeopteryx, just two years after Charles Darwin published The Origin of Species, set off a firestorm of debate about evolution and the role of transitional fossils that endures to this day.

The eleven fossils currently classified as Archaeopteryx are the oldest evidence of feathers on the planet and the only ones dated from Jurassic times. Furthermore, their advanced nature and placement suggest their origins must have been even earlier.[1] All remains have been regarded by most as a single species, though this has been debated.

Description

Eight Archaeopteryx specimens compared to a human foot in scale.

Archaeopteryx was a primitive avian that lived during the Kimmeridgian stage of the Jurassic Period, around 155-150 million years ago.[2] The only specimens of Archaeopteryx that have been discovered come from the area that is now Germany.[2]

Archaeopteryx was roughly the size of a medium-sized, modern-day bird, with broad wings that were rounded at the ends and a long tail compared to its body length. In all, Archaeopteryx could reach up to 0.5 meters (1.6 feet) in body length. Archaeopteryx feathers, although less documented than its other features, were similar in structure and design to modern-day bird feathers.[2] However, despite the presence of numerous avian features,[3] Archaeopteryx had many theropod dinosaur characteristics. Unlike modern avians, Archaeopteryx had small teeth inside its mouth,[2] which enabled it to hunt insects and other small invertebrates of the period. Archaeopteryx also had a tail, which was long and bony, and its talons ended in three toes with long claws, more features which Archaeopteryx shared with the dinosaurs.

Because it displays a number of features common to both birds and dinosaurs, Archaeopteryx has often been considered a link between them - possibly the first avian in its change from a land dweller to a bird.[2] In the 1970s, John Ostrom argued that the birds evolved from theropod dinosaurs and Archaeopteryx was a critical piece of evidence for this argument; it preserves a number of avian features, such as a wishbone, flight feathers, wings and a partially reversed first toe, and a number of dinosaur and theropod features. For instance, it has a long ascending process of the ankle bone, interdental plates, an obturator process of the ischium, and long chevrons in the tail. In particular, Ostrom found that Archaeopteryx was remarkably similar to the theropod family Dromaeosauridae.

The first remains of Archaeopteryx were discovered just two years after Charles Darwin published The Origin of Species in 1862. Archaeopteryx seemed to confirm Darwin's theories and has since become a key piece of evidence in the dinosaur-bird connection, transitional fossils debate and the confirmation of evolution. Indeed, further research on dinosaurs from the Gobi Desert and China has since provided more evidence of a link between Archaeopteryx and the dinosaurs, such as the Chinese feathered dinosaurs. Archaeopteryx is close to the ancestry of modern birds - it shows most of the features one would expect in an ancestral bird - but it may not be the direct ancestor of living birds and it is arguable how much divergence was already present in the early birds at its time.

Paleobiology

Plumage

A model of Archaeopteryx lithographica.

Specimens of Archaeopteryx were most notable for their well-developed flight feathers. They were markedly asymmetrical and showed the structure of flight feathers in modern birds, with vanes given stability by a barb-barbule-barbicel arrangement. The tail feathers were less asymmetrical, again in line with the situation in modern birds and also had firm vanes. The thumb however, did not bear a separately movable tuft of stiff feathers yet.

The body plumage of Archaeopteryx is less well documented and has only been properly researched in the well-preserved Berlin specimen. Thus, as more than one species seems to be involved, the research into the Berlin specimen's feathers does not necessarily hold true for the rest of the species of Archaeopteryx. In the Berlin specimen, there are "trousers" of well-developed feathers on the legs; some of these feathers seem to have a basic contour feather structure but are somewhat decomposed (i.e., lack barbicels as in ratites),[4] but in part they are firm and thus capable of supporting flight.[5]

There was a patch of pennaceous feathers running along the back which was quite similar to the contour feathers of the body plumage of modern birds in being symmetrical and firm, though not as stiff as the flight-related feathers. Apart from that, the feather traces in the Berlin specimen are limited to a sort of "proto-down" not dissimilar to that found in the dinosaur Sinosauropteryx, being decomposed and fluffy, and possibly even appeared more like fur than like feathers in life (though not in their microscopic structure). These occur on the remainder of the body, as far as such structures are both preserved and not obliterated by preparation, and the lower neck.[4]

However, there is no indication of feathering on the upper neck and head; while these may conceivably have been nude as in many closely related feathered dinosaurs for which good specimens are available, this may still be an artifact of preservation: it appears that most Archaeopteryx specimens became embedded in anoxic sediment after drifting some time on their back in the sea - the head and neck and the tail are generally bent downwards which suggests that the specimens had just started to rot when they were embedded, with tendons and muscle relaxing so that the characteristic shape of the fossil specimens was achieved. This would mean that the skin was already softened and loose, which is bolstered by the fact that in some specimens, the flight feathers were starting to detach at the point of embedding in the sediment. So it is hypothesized that the pertinent specimens moved along the sea bed in shallow water for some time before burial, the head and upper neck feathers sloughing off, while the more firmly attached tail feathers remained.[6]

Flight

As in the wings of modern birds, the flight feathers of Archaeopteryx were highly asymmetrical, and the tail feathers are rather broad. This implies that the wings and tail were used for lift generation. However, it is unclear whether Archaeopteryx was simply a glider or capable of flapping flight. The lack of a bony breastbone suggests that Archaeopteryx was not a very strong flier, but flight muscles might have attached to the thick, boomerang-shaped wishbone, the platelike coracoids, or perhaps to a cartilaginous sternum. The sideways orientation of the glenoid (shoulder) joint between scapula, coracoid and humerus - instead of the dorsally angled arrangement found in modern birds - suggests that Archaeopteryx was unable to lift its wings above its back, a requirement for the upstroke found in modern flapping flight. Thus, it seems likely that Archaeopteryx was indeed unable to use flapping flight as modern birds do, but it may well have utilized a downstroke-only flap-assisted gliding technique.[7]

Archaeopteryx wings were relatively large, which would have resulted in a low stall speed and reduced turning radius. The short and rounded shape of the wings would have increased drag, but could also have improved Archaeopteryx' ability to fly through cluttered environments such as trees and brush (similar wing shapes are seen in birds which fly through trees and brush, such as crows and pheasants). The presence of "hind wings", asymmetrical flight feathers stemming from the legs similar to those seen in dromaeosaurids such as Microraptor, would also have added to the aerial mobility of Archaeopteryx. The first detailed study of the hind wings by Longrich in 2006 suggested that the structures formed up to 12% of the total airfoil. Considering that it is not certain to what extent such feathers capable of supporting flight were present on the legs, this would have reduced stall speed by up to 6% and turning radius by up to 12%, in addition to the stall and turning radius reduction provided by the primary wing and tail feathers.[5]

In 2004, scientists analyzing a detailed CT scan of Archaeopteryx's braincase, concluded that its brain was significantly larger than that of most dinosaurs, indicating that it possessed the brain size necessary for flying. The overall brain anatomy was reconstructed using the scan. The reconstruction showed that the regions associated with vision took up nearly one-third of the brain. Other well-developed areas involved hearing and muscle coordination.[8] The skull scan also revealed the structure of the inner ear. The structure more closely resembles that of modern birds than the inner ear of reptiles. These characteristics taken together suggest that Archaeopteryx had the keen sense of hearing, balance, spatial perception and coordination needed to fly.[9]

Archaeopteryx continues to play an important part in scientific debates about the origin and evolution of birds. Some scientists see it as a semi-arboreal climbing animal, following the idea that birds evolved from tree-dwelling gliders (the "trees down" hypothesis for the evolution of flight proposed by O.C. Marsh). Other scientists see Archaeopteryx as running quickly along the ground, supporting the idea that birds evolved flight by running (the "ground up" hypothesis proposed by Samuel Wendell Williston). Still others suggest that Archaeopteryx might have been at home both in the trees and on the ground, like modern crows, and this latter view is what today is considered best-supported by morphological characters. Altogether, it appears that it was a species which was neither particularly specialized for running on the ground, nor for perching. Considering the current knowledge of flight-related morphology, a scenario as outlined by Elżanowski in 2002, namely that Archaeopteryx used its wings mainly to escape predators by glides punctuated with shallow downstrokes to reach successively higher perches, and alternatively to cover longer distances by (mainly) gliding down from cliffs or treetops, appears quite reasonable.[6]

Given that it is now well established that several lineages of theropods evolved feathers and flight independently, the question of how precisely the ancestors of Archaeopteryx became able to fly has lost dramatically in importance for the time being. Since it is quite likely that this species belongs to a lineage of birds unrelated to the Neornithes (the Jurassic ancestor of which remains unknown), how exactly flying ability was gained in Archaeopteryx may be a moot point, having little bearing on how this happened in the ancestors of modern birds.

Paleoecology

The richness and diversity of the Solnhofen limestones in which all specimens of Archaeopteryx have been found have shed light on an ancient Jurassic Bavaria strikingly different from the present day. The latitude was similar to Florida, though the climate was likely to have been drier as evidenced by fossils of plants with adaptations for arid conditions and lack of terrestrial sediments characteristic of rivers. Evidence of plants, though scarce, include cycads and conifers while animals found include a large number of insects, small lizards, pterosaurs and Compsognathus.[10]

The excellent preservation of Archaeopteryx fossils and other terrestrial fossils found at Solnhofen indicates that they did not travel far before becoming preserved.[11] The Archaeopteryx specimens found are likely therefore to have lived on the low islands surrounding the Solnhofen lagoon rather than been corpses that drifted in from further away. Archaeopteryx skeletons are considerably less numerous in the deposits of Solnhofen than those of pterosaurs such as Rhamphorhynchus, the group which dominated the niche currently occupied by seabirds, yet are common enough that it is unlikely that the specimens found are vagrants from the larger islands 50 km (31 miles) to the north.[12]

The islands that surrounded the Solnhofen lagoon were low lying, semi-arid and sub-tropical with a long dry season and little rain.[13] The flora of these islands was adapted to these dry conditions and consisted mostly of low (3 m [10 ft]) shrubs.[12] Contrary to reconstructions of Archaeopteryx climbing large trees, these seem to have been mostly absent from the islands; few trunks have been found in the sediments and fossilised tree pollen is also absent.

The lifestyle of Archaeopteryx is difficult to reconstruct and there are several theories regarding it. It has been suggested by some researchers that it was primarily adapted to life on the ground,[14] while other researchers suggest that it was principally arboreal. The absence of trees does not preclude Archaeopteryx from an arboreal lifestyle; several species of extant bird live exclusively in low shrubs. Various aspects of the morphology of Archaeopteryx point to either an arboreal or ground existence, the length of its legs, the elongation in its feet; and some authorities consider it likely to have been a generalist capable of feeding in both shrubs, open ground and even alongside the shores of the lagoon.[12] It most likely hunted small prey, seizing it with its jaws if it were small enough or with its claws if it were larger.

History of discovery

The London Archaeopteryx, 1863, detail, note the feathers.

Over the years, ten body fossil specimens of Archaeopteryx and a feather that may belong to it have been found. All of the fossils come from the limestone deposits, quarried for centuries, near Solnhofen, Germany.[15][10] The initial discovery, a single feather, was unearthed in 1860 and described a year later by Christian Erich Hermann von Meyer. It is currently located at the Humbolt Museum für Naturkunde in Berlin. This is generally assigned to Archaeopteryx and was the initial holotype, but whether it actually is a feather of this species or another, as yet undiscovered, proto-bird is unknown. There are some indications it is indeed not from the same animal as most of the skeletons (the "typical" A. lithographica).[16]

Soon after, the first skeleton, known as the London Specimen (BMNH 37001) was unearthed in 1861 near Langenaltheim, Germany and given to a local physician Karl Häberlein in return for medical services. He then sold it to the British Museum of Natural History in London, where it remains.[10] Missing most of its head and neck, it was described in 1863 by Richard Owen as Archaeopteryx macrura, who assumed it did not belong to the same species as the feather. In a subsequent edition of his Origin of Species (chap. 10, pp.335-336), Charles Darwin acclaimed Owen's discovery as linking lizard-like reptiles with modern birds.

It appears as if the ambiguity of the Ancient Greek πτερυξ was realized by von Meyer. At the time of the original description, he referred to a single feather which appeared like a modern bird's remex (wing feather), but he had heard of and been shown a rough sketch of the London specimen, to which he referred to as a "Skelet eines mit Federn bedeckten Thiers" ("skeleton of an animal covered in feathers"). In German, this ambiguity is amply resolved by the term Schwinge which confers exactly the same ambiguity as πτερυξ - Urschwinge was the favored translation of Archaeopteryx among German scholars in the late 19th century; in English, "ancient pinion" would be the best possible approximation.

The Berlin Archaeopteryx, 1881.

Since then nine specimens have been recovered:

The Berlin Specimen (HMN 1880) was discovered in 1876 or 1877 on the Blumenberg near Eichstätt, Germany, by Jakob Niemeyer. He exchanged this precious fossil for a cow, with Johann Dörr. Placed on sale in 1881, with potential buyers including O.C. Marsh of Yale University's Peabody Museum, it was bought by the Humbolt Museum für Naturkunde, where it is now displayed. The transaction was financed by Ernst Werner von Siemens founder of the famous company that bears his name.[10] Described in 1884 by Wilhelm Dames, it is the most complete specimen, and the first with a complete head. Once classified as a new species, A. siemensii, a recent evaluation supports the A. siemensii species definition.[6]

Composed of a torso, the Maxberg Specimen (S5) was discovered in 1956 or 1958 near Langenaltheim and described in 1959 by Heller. It is currently missing, though it was once exhibited at the Maxberg Museum in Solnhofen. It belonged to Eduard Opitsch, who loaned it to the museum. After his death in 1991, the specimen was discovered to be missing and may have been stolen or sold. The specimen is missing its head and tail, although the rest of the skeleton is mostly intact.

The Haarlem Specimen (TM 6428, also known as the Teyler Specimen) was discovered in 1855 near Riedenburg, Germany and described as a Pterodactylus crassipes in 1875 by von Meyer. It was reclassified in 1970 by John Ostrom and is currently located at the Teyler Museum in Haarlem, the Netherlands. It was the very first specimen, despite the classification error. It is also one of the least complete specimens, consisting mostly of limb bones and isolated cervical vertebrae and ribs.

The Eichstätt Specimen (JM 2257) was discovered in 1951 or 1955 near Workerszell, Germany and described by Peter Wellnhofer in 1974. Currently located at the Jura Museum in Eichstätt, Germany, it is the smallest specimen and has the second best head. It is possibly a separate genus (Jurapteryx recurva) or species (A. recurva).

The Munich Specimen.

The Solnhofen Specimen (BSP 1999) was discovered in the 1960s near Eichstätt, Germany and described in 1988 by Wellnhofer. Currently located at the Bürgermeister-Müller-Museum in Solnhofen, it was originally classified as Compsognathus by an amateur collector. It is the largest specimen known and may belong to a separate genus and species, Wellnhoferia grandis. It is missing only portions of the neck, tail, backbone, and head.

The Munich Specimen (S6, formerly known as the Solnhofen-Aktien-Verein Specimen) was discovered in 1991 near Langenaltheim and described in 1993 by Wellnhofer. It is currently located at the Paläontologische Museum München in Munich. What was initially believed to be a bony sternum turned out to be part of the coracoid,[17] but a cartilaginous sternum may have been present. Only the front of its face is missing. It may be a new species, A. bavarica.

An eighth, fragmentary specimen, the Bürgermeister-Müller Specimen was discovered in 1997 and it is currently kept at the Bürgermeister-Müller Museum. Other than the above remains discovered, a further fragmentary fossil was found in 2004.

Long in a private collection, the Thermopolis Specimen (WDC CSG 100) was discovered in Germany and described in 2005 by Mayr, Pohl, and Peters. Donated to the Wyoming Dinosaur Center in Thermopolis, Wyoming, it has the best-preserved head and feet; most of the neck and the lower jaw have not been preserved. The "Thermopolis" specimen was described in the December 2, 2005 Science journal article as "A well-preserved Archaeopteryx specimen with theropod features", shows that the Archaeopteryx lacked a reversed toe—a universal feature of birds—limiting its ability to perch in trees and implying a terrestrial lifestyle.[18] This has been interpreted as evidence of theropod ancestry. The specimen also has a hyperextendible second toe. "Until now, the feature was thought to belong only to the species' close relatives, the deinonychosaurs."[19] This tenth and latest specimen was assigned to Archaeopteryx siemensii in 2007.[20] The specimen itself, currently on loan to the Forschungsinstitut Senckenberg in Frankfurt, is considered the most complete and well preserved Archaeopteryx remains yet.[20]

Taxonomy

Today, there is only one recognized species of Archaeopteryx, A. lithographica, but its taxonomic history is extremely confused. Dozens of names have been published for the handful of specimens, most of which are simply spelling errors (lapsus). Originally, the name A. lithographica only referred to the single feather described by von Meyer. In 1960, Swinton proposed that the name Archaeopteryx lithographica be officially transferred from the feather to the London specimen.[21] The ICZN did suppress the plethora of alternative names initially proposed for the first skeleton specimens,[22] which mainly resulted from the acrimonious dispute between von Meyer and his opponent Johann Andreas Wagner (whose Griphosaurus problematicus - "enigmatic lizard" - was a vitriolic sneer at von Meyer's Archaeopteryx).[23] In addition, descriptions of Archaeopteryx fossils as pterosaurs before their true nature was realized were also suppressed.[24]

The relationships of the specimens are problematic. Most subsequent specimens have been given their own species at one point or another. The Berlin specimen has been designated as Archaeornis siemensii, the Eichstätt specimen as Jurapteryx recurva, the Munich specimen as Archaeopteryx bavarica and the Solnhofen specimen was designated as Wellnhoferia grandis.

Recently, it has been argued that all the specimens belong to the same species.[25] However, significant differences exist among the specimens. In particular, the Munich, Eichstätt, Solnhofen and Thermopolis specimens differ from the London, Berlin, and Haarlem specimens in being smaller or much larger, having different finger proportions, having more slender snouts, lined with forward-pointing teeth and possible presence of a sternum. These differences are as large as or larger than the differences seen today between adults of different bird species. However, it is also possible that these differences could be explained by different ages of the living birds.

Finally, it is worth noting that the feather, the first specimen of Archaeopteryx described, does not agree too well with the flight-related feathers of Archaeopteryx. It certainly is a remix of a contemporary species, but its size and proportions indicate that it probably belongs to an as of yet undiscovered species of primitive bird or possibly bird-like dinosaur. As the feather was the original type specimen, this has created quite some nomenclatorial confusion.

Synonyms

If two names are given, the first denotes the original describer of the "species", the second the author on whom the given name combination is based. As always in zoological nomenclature, putting an author's name in parentheses denotes that the taxon was originally described in a different genus.

  • Pterodactylus crassipes Meyer, 1857 [suppressed in favor of A. lithographica 1977 per ICZN Opinion 1070]
  • Rhamphorhynchus crassipes (Meyer, 1857) (as Pterodactylus (Rhamphorhynchus) crassipes) [suppressed in favor of A. lithographica 1977 per ICZN Opinion 1070]
  • Archaeopteryx lithographica Meyer, 1861 [nomen conservandum]
  • Scaphognathus crassipes (Meyer, 1857) Wagner, 1861 [suppressed in favor of A. lithographica 1977 per ICZN Opinion 1070]
  • Archaeopterix lithographica Anon., 1861 [lapsus]
  • Griphosaurus problematicus Wagner, 1861 [nomen oblitum 1961 per ICZN Opinion 607]
  • Griphornis longicaudatus Woodward, 1862 [nomen oblitum 1961 per ICZN Opinion 607]
  • Griphosaurus longicaudatum (Woodward, 1862) [lapsus]
  • Griphosaurus longicaudatus (Owen, 1862) [nomen oblitum 1961 per ICZN Opinion 607]
  • Archaeopteryx macrura Owen, 1862 [nomen oblitum 1961 per ICZN Opinion 607]
  • Archaeopterix macrura Owen, 1862 [lapsus]
  • Archaeopterix macrurus Egerton, 1862 [lapsus]
  • Archeopteryx macrurus Owen, 1863 [unjustified emendation]
  • Archaeopteryx macroura Vogt, 1879 [lapsus]
  • Archaeopteryx siemensii Dames, 1897
  • Archaeopteryx siemensi Dames, 1897 [lapsus]
  • Archaeornis siemensii (Dames, 1897) Petronievics, 1917[20]
  • Archaeopteryx oweni Petronievics, 1917 [nomen oblitum 1961 per ICZN Opinion 607]
  • Gryphornis longicaudatus Lambrecht, 1933 [lapsus]
  • Gryphosaurus problematicus Lambrecht, 1933 [lapsus]
  • Archaeopteryx macrourus Owen, 1862 fide Lambrecht, 1933 [lapsus]
  • Archaeornis siemensi (Dames, 1897) fide Lambrecht, 1933? [lapsus]
  • Archeopteryx macrura Ostrom, 1970 [lapsus]
  • Archaeopteryx crassipes (Meyer, 1857) Ostrom, 1972 [suppressed in favor of A. lithographica 1977 per ICZN Opinion 1070]
  • Archaeopterix lithographica di Gregorio, 1984 [lapsus]
  • Archaeopteryx recurva Howgate, 1984
  • Jurapteryx recurva (Howgate, 1984) Howgate, 1985
  • Archaeopteryx bavarica Wellnhofer, 1993
  • Wellnhoferia grandis Elżanowski, 2001

The last 4 taxa may be valid genera and species.

"Archaeopteryx" vicensensis (Anon. fide Lambrecht, 1933) is a nomen nudum for what appears to be an undescribed pterosaur.

Controversies

Authenticity

Beginning in 1985, a group including astronomer Fred Hoyle and physicist Lee Spetner published a series of papers claiming that the feathers on the Berlin and London specimens of Archaeopteryx were actually forged.[26][27][28][29] Their claims were repudiated by Alan J. Charig and others at the British Museum of Natural History.[30] Most of their evidence for a forgery was based on unfamiliarity with the processes of lithification; for example, they proposed that based on the difference in texture associated with the feathers, feather impressions were applied to a thin layer of cement,[27] without realizing that feathers themselves would have caused a textural difference.[30] They also expressed disbelief that slabs would split so smoothly, or that one half of a slab containing fossils would have good preservation, but not the counterslab.[28][26] These, though, are common properties of Solnhofen fossils because the dead animals would fall onto hardened surfaces which would form a natural plane for the future slabs to split along, leaving the bulk of the fossil on one side and little on the other.[30] They also misinterpreted the fossils, claiming that the tail was forged as one large feather,[27] when this is visibly not the case.[30] In addition, they claimed that the other specimens of Archaeopteryx known at the time did not have feathers,[27][26] which is untrue; the Maxberg and Eichstätt specimens have obvious feathers.[30] Finally, the motives they suggested for a forgery are not strong, and contradictory; one is that Richard Owen wanted to forge evidence in support of Charles Darwin's theory of evolution, which is unlikely given Owen's views toward Darwin and his theory. The other is that Owen wanted to set a trap for Darwin, hoping the latter would support the fossils so Owen could discredit him with the forgery; this is unlikely because Owen himself wrote a detailed paper on the London specimen, so such an action would certainly backfire.[31]

Charig et al. pointed to the presence of hairline cracks in the slabs running through both rock and fossil impressions, and mineral growth over the slabs that had occurred before discovery and preparation, as evidence that the feathers were original.[30] Spetner et al. then attempted to show that the cracks would have naturally propagated through their postulated cement layer,[32] but neglected to account for the fact that the cracks were old and had been filled with calcite, and thus were not able to propagate.[31] They also attempted to show the presence of cement on the London specimen through X-ray spectrometry, and did find something that was not rock.[32] However, it was not cement, either, and is most probably from a fragment of silicone rubber left behind when molds were made of the specimen.[31] Their suggestions have not been taken seriously by paleontologists, as their evidence was largely based on misunderstandings of geology, and they never discussed the other feather-bearing specimens, which have increased in number since then.

Archaeopteryx and Protoavis

In 1984, Sankar Chatterjee discovered fossils which he claimed in 1991 belonged to a fossil avian far older than Archaeopteryx. These fossils, believed to be around 210 to 225 million years old, have been assigned the name Protoavis.[33] The fossils are too badly preserved to allow an estimate of flying ability; although Chatterjee's reconstructions usually show feathers, many paleontologists, including Paul (2002) and Witmer (2002) have rejected the claims that Protoavis was an earlier bird (or, alternately, that it existed at all).[12][34] The fossils were found disarticulated, and were collected from different locations. Because the fossils are in poor condition, Archaeopteryx remains the earliest universally recognized avian.[35]

Archaeopteryx is the best known early avian, and has thus received widespread attention. Its easily recognizable appearance and the intense public interest in dinosaurs have caused Archaeopteryx to become a feature of worldwide popular culture. A main belt asteroid discovered in 1991, 9860 Archaeopteryx, was named in honor of the genus.[36][37]

Track 11 on the Lemon Demon album Dinosaurchestra is entitled "Archaeopteryx". The song is about a person with an intense dislike of birdkind based on a jealousy of their flight ability. The person then travels back in time and kills the original Archaeopteryx, thus killing off all future birds.

See also

Footnotes

  1. ^ Wellnhofer P (2004). "The Plumage of Archaeopteryx". In Currie PJ, Koppelhus EB, Shugar MA, Wright JL (ed.). Feathered Dragons. Indiana University Press. pp. 282–300. ISBN 0-253-34373-9.{{cite book}}: CS1 maint: multiple names: editors list (link)
  2. ^ a b c d e Lambert, David (1993). The Ultimate Dinosaur Book. New York: Dorling Kindersley. pp. 38–81. ISBN 1-56458-304-X. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  3. ^ Holtz, Thomas, Jr. (1995). "Archaeopteryxs Relationship With Modern Birds". Journal of Dinosaur Paleontology. Retrieved 2007-03-01. {{cite web}}: Cite has empty unknown parameter: |coauthors= (help)CS1 maint: multiple names: authors list (link)
  4. ^ a b Christensen P, Bonde N. (2004). Body plumage in Archaeopteryx: a review, and new evidence from the Berlin specimen. Comptes Rendus Palevol. 3: 99–118. PDF fulltext
  5. ^ a b Longrich N. (2006): Structure and function of hindlimb feathers in Archaeopteryx lithographica. Paleobiology. 32(3): 417–431. doi:10.1666/04014.1 (HTML abstract) Cite error: The named reference "Longrich" was defined multiple times with different content (see the help page).
  6. ^ a b c Elżanowski A. (2002): Archaeopterygidae (Upper Jurassic of Germany). In: Chiappe, L. M. & Witmer, L. M (eds.), Mesozoic Birds: Above the Heads of Dinosaurs: 129–159. University of California Press, Berkeley.
  7. ^ Senter, P. (2006). Scapular orientation in theropods and basal birds and the origin of flapping flight. Acta Palaeontologica Polonica. 51(2): 305–313. PDF fulltext
  8. ^ Witmer, L. M. (2004). Palaeontology: Inside the oldest bird brain. Nature. 430(7000): 619-620. PMID 15295579 doi:10.1038/430619a
  9. ^ Alonso, P. D., Milner, A. C., Ketcham, R. A., Cookson, M. J. & Rowe, T. B. (2004). The avian nature of the brain and inner ear of Archaeopteryx. Nature. 430(7000): 666-669. PMID 15295597. doi:10.1038/nature02706. PDF fulltext Supplementary info
  10. ^ a b c d Chiappe, Luis M. (2007). Glorified Dinosaurs. Sydney: UNSW Press. pp. 118–146. ISBN 0-471-24723-5. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)
  11. ^ Davis, P. (1998). "The impact of decay and disarticulation on the preservation of fossil birds". Palaios. 13 (1): 3–13. Retrieved 2007-03-25. {{cite journal}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
  12. ^ a b c d Paul, Gregory S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press. ISBN 0-8018-6763-0. Cite error: The named reference "Paul" was defined multiple times with different content (see the help page).
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  21. ^ Swinton, W. E. (1960). Opinion 1084, Proposed addition of the generic name Archaeopteryx VON MEYER, 1861 and the specific name Lithographica, VON MEYER, 1861, as published in the binomen Archaeopteryx Lithographica to the official lists (Class Aves). Bulletin of Zoological Nomenclature. 17(6-8): 224-226.
  22. ^ ICZN. (1961). Opinion 607, Archaeopteryx VON MEYER, 1861 (Aves); Addition to the Official list. Bulletin of Zoological Nomenclature. 18(4): 260-261.
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  33. ^ Chatterjee, S. (1991). Cranial anatomy and relationships of a new Triassic bird from Texas. Philosophical Transactions of the Royal Society of London, B 332: 277-342.
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References

  • de Beer, G. R. (1954). Archaeopteryx lithographica: A Study Based upon the British Museum Specimen. Trustees of the British Museum, London.
  • Bühler, P. & Bock, W. J. (2002). Zur Archaeopteryx-Nomenklatur: Missverständnisse und Lösung. Journal of Ornithology. 143(3): 269-286. [Article in German, English abstract] doi:10.1046/j.1439-0361.2002.02006.x (HTML abstract)
  • Chambers, P. (2002). Bones of Contention: The Fossil that Shook Science. John Murray, London. ISBN 0-7195-6059-4.
  • Feduccia, A. (1996). The Origin and Evolution of Birds. Yale University Press, New Haven. ISBN 0-300-06460-8.
  • Feduccia, A. (1993). Evidence from claw geometry indicating arboreal habits of Archaeopteryx. Science. 259(5096): 790-793. HTML abstract
  • Feduccia, A. & Tordoff, H. B. (1979). Feathers of Archaeopteryx: Asymmetric vanes indicate aerodynamic function. Science. 203(4384): 1021-1022. HTML abstract
  • Heilmann, G. (1926). The Origin of Birds. Witherby, London.
  • Huxley, T. H. (1871). Manual of the anatomy of vertebrate animals. London.
  • Kennedy, Elaine (2000). Solnhofen Limestone: Home of Archaeopteryx. Geoscience Reports. 30: 1-4. Retrieved 2006-10-18.
  • Nedin, C. (1999). All About Archaeopteryx. talk.origins archive. Version of June 10, 2002; retrieved 2006-10-18.
  • Olson, S. L. & Feduccia, A. (1979). Flight capability and the pectoral girdle of Archaeopteryx. Nature. 278(5701). 247-248. doi:10.1038/278247a0 (HTML abstract)
  • Ostrom, J. H. (1976). Archaeopteryx and the origin of birds. Biol. J. Linn. Soc.. 8: 91-182.
  • Ostrom, J. H. (1985). Introduction to Archaeopteryx. In: Hecht, M. K. O.; Ostrom, J.H.; Viohl, G. & Wellnhofer, P. (eds.) The Beginnings of Birds: Proceedings of the International Archaeopteryx Conference: 9-20. Eichstätt, Freunde des Jura-Museums Eichstätt.
  • Owen, R. (1863). On the Archaeopteryx of Von Meyer, with a description of the fossil remains of a long-tailed species from the lithographic stone of Solnhofen. Phil. Trans. Roy. Soc. London. 153: 33-47. First page image
  • von Meyer, H. (1861). Archaeopteryx litographica (Vogel-Feder) und Pterodactylus von Solenhofen. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde. 1861: 678-679, plate V [Article in German].
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