Archaeopteryx
Archaeopteryx | |
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The Berlin Archaeopteryx specimen (A. siemensii) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Clade: | Paraves |
Family: | †Archaeopterygidae |
Genus: | †Archaeopteryx Meyer, 1861 (conserved name) |
Type species | |
†Archaeopteryx lithographica Meyer, 1861 (conserved name)
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Referred species | |
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Synonyms[1] | |
Genus synonymy
Species synonymy
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Archaeopteryx (
Archaeopteryx lived in the Late Jurassic around 150 million years ago, in what is now southern Germany, during a time when Europe was an archipelago of islands in a shallow warm tropical sea, much closer to the equator than it is now. Similar in size to a Eurasian magpie, with the largest individuals possibly attaining the size of a raven,[4] the largest species of Archaeopteryx could grow to about 0.5 m (1 ft 8 in) in length. Despite their small size, broad wings, and inferred ability to fly or glide, Archaeopteryx had more in common with other small Mesozoic dinosaurs than with modern birds. In particular, they shared the following features with the dromaeosaurids and troodontids: jaws with sharp teeth, three fingers with claws, a long bony tail, hyperextensible second toes ("killing claw"), feathers (which also suggest warm-bloodedness), and various features of the skeleton.[5][6]
These features make Archaeopteryx a clear candidate for a transitional fossil between non-avian dinosaurs and birds.[7][8] Thus, Archaeopteryx plays an important role, not only in the study of the origin of birds, but in the study of dinosaurs. It was named from a single feather in 1861,[9] the identity of which has been controversial.[10][11] That same year, the first complete specimen of Archaeopteryx was announced. Over the years, ten more fossils of Archaeopteryx have surfaced. Despite variation among these fossils, most experts regard all the remains that have been discovered as belonging to a single species, although this is still debated.
Archaeopteryx was long considered to be the beginning of the evolutionary tree of birds. However, in recent years, the discovery of several small, feathered dinosaurs has created a mystery for palaeontologists, raising questions about which animals are the ancestors of modern birds and which are their relatives.
History of discovery
Over the years, twelve body fossil specimens of Archaeopteryx have been found. All of the fossils come from the limestone deposits, quarried for centuries, near Solnhofen, Germany. These quarries excavate sediments from the Solnhofen Limestone formation and related units.[14][15]
The initial discovery, a single feather, was unearthed in 1860 or 1861 and described in 1861 by Hermann von Meyer.[16] It is currently located at the Natural History Museum of Berlin. Though it was the initial holotype, there were indications that it might not have been from the same animal as the body fossils.[9] In 2019 it was reported that laser imaging had revealed the structure of the quill (which had not been visible since some time after the feather was described), and that the feather was inconsistent with the morphology of all other Archaeopteryx feathers known, leading to the conclusion that it originated from another dinosaur.[10] This conclusion was challenged in 2020 as being unlikely; the feather was identified on the basis of morphology as most likely having been an upper major primary covert feather.[11]
The first skeleton, known as the London Specimen (BMNH 37001),[17] was unearthed in 1861 near Langenaltheim, Germany, and perhaps given to local physician Karl Häberlein in return for medical services. He then sold it for £700 (roughly £83,000 in 2020[18]) to the Natural History Museum in London, where it remains.[14] Missing most of its head and neck, it was described in 1863 by Richard Owen as Archaeopteryx macrura, allowing for the possibility it did not belong to the same species as the feather. In the subsequent fourth edition of his On the Origin of Species,[19] Charles Darwin described how some authors had maintained "that the whole class of birds came suddenly into existence during the eocene period; but now we know, on the authority of Professor Owen, that a bird certainly lived during the deposition of the upper greensand; and still more recently, that strange bird, the Archaeopteryx, with a long lizard-like tail, bearing a pair of feathers on each joint, and with its wings furnished with two free claws, has been discovered in the oolitic slates of Solnhofen. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world."[20]
The Greek word archaīos (ἀρχαῖος) means 'ancient, primeval'. Ptéryx primarily means 'wing', but it can also be just 'feather'. Meyer suggested this in his description. At first he referred to a single feather which appeared to resemble 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 as a "Skelett eines mit ähnlichen Federn bedeckten Tieres" ("skeleton of an animal covered in similar feathers"). In German, this ambiguity is resolved by the term Schwinge which does not necessarily mean a wing used for flying. Urschwinge was the favoured translation of Archaeopteryx among German scholars in the late nineteenth century. In English, 'ancient pinion' offers a rough approximation to this.[citation needed]
Since then, twelve specimens have been recovered:
The Berlin Specimen (HMN 1880/81) was discovered in 1874 or 1875 on the Blumenberg near
Composed of a torso, the
The Haarlem Specimen (TM 6428/29, also known as the Teylers Specimen) was discovered in 1855 near Riedenburg, Germany, and described as a Pterodactylus crassipes in 1857 by Meyer. It was reclassified in 1970 by John Ostrom and is currently located at the Teylers Museum in Haarlem, the Netherlands. It was the very first specimen found, but was incorrectly classified at the time. It is also one of the least complete specimens, consisting mostly of limb bones, isolated cervical vertebrae, and ribs. In 2017 it was named as a separate genus Ostromia, considered more closely related to Anchiornis from China.[24]
The Eichstätt Specimen (JM 2257) was discovered in 1951 near Workerszell, Germany, and described by Peter Wellnhofer in 1974. Currently located at the Jura Museum in Eichstätt, Germany, it is the smallest known specimen and has the second-best head. It is possibly a separate genus (Jurapteryx recurva) or species (A. recurva).[25]
The Solnhofen Specimen (unnumbered specimen) was discovered in the 1970s near Eichstätt, Germany, and described in 1988 by Wellnhofer. Currently located at the Bürgermeister-Müller-Museum in Solnhofen, it originally was classified as Compsognathus by an amateur collector, the same mayor Friedrich Müller after which the museum is named. 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.[26]
The Munich Specimen (BSP 1999 I 50, formerly known as the Solenhofer-Aktien-Verein Specimen) was discovered on 3 August 1992 near Langenaltheim and described in 1993 by Wellnhofer. It is currently located at the
An eighth, fragmentary specimen was discovered in 1990 in the younger
Another fragmentary fossil was found in 2000. It is in private possession and, since 2004, on loan to the
Long in a private collection in Switzerland, the Thermopolis Specimen (WDC CSG 100) was discovered in Bavaria 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 on 2 December 2005 Science journal article as "A well-preserved Archaeopteryx specimen with theropod features"; it shows that Archaeopteryx lacked a reversed toe—a universal feature of birds—limiting its ability to perch on branches and implying a terrestrial or trunk-climbing lifestyle.[36] This has been interpreted as evidence of theropod ancestry. In 1988, Gregory S. Paul claimed to have found evidence of a hyperextensible second toe,[37] but this was not verified and accepted by other scientists until the Thermopolis specimen was described. "Until now, the feature was thought to belong only to the species' close relatives, the deinonychosaurs."[15] The Thermopolis Specimen was assigned to Archaeopteryx siemensii in 2007.[22] The specimen is considered to represent the most complete and best-preserved Archaeopteryx remains yet.[22]
The discovery of an eleventh specimen was announced in 2011; it was described in 2014. It is one of the more complete specimens, but is missing much of the skull and one forelimb. It is privately owned and has yet to be given a name.
A twelfth specimen had been discovered by an amateur collector in 2010 at the Schamhaupten quarry, but the finding was only announced in February 2014.[42] It was scientifically described in 2018. It represents a complete and mostly articulated skeleton with skull. It is the only specimen lacking preserved feathers. It is from the Painten Formation and somewhat older than the other specimens.[43]
Authenticity
Beginning in 1985, an amateur 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 forged.[44][45][46][47] Their claims were repudiated by Alan J. Charig and others at the Natural History Museum in London.[48] Most of their supposed 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,[45] without realizing that feathers themselves would have caused a textural difference.[48] They also misinterpreted the fossils, claiming that the tail was forged as one large feather,[45] when visibly this is not the case.[48] In addition, they claimed that the other specimens of Archaeopteryx known at the time did not have feathers,[44][45] which is incorrect; the Maxberg and Eichstätt specimens have obvious feathers.[48]
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
Finally, the motives they suggested for a forgery are not strong, and are 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 wrote a detailed paper on the London specimen, so such an action would certainly backfire.[49]
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.[48] Spetner et al. then attempted to show that the cracks would have propagated naturally through their postulated cement layer,[50] 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.[49] They also attempted to show the presence of cement on the London specimen through X-ray spectroscopy, and did find something that was not rock;[50] it was not cement either, and is most probably a fragment of silicone rubber left behind when moulds were made of the specimen.[49] Their suggestions have not been taken seriously by palaeontologists, 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. Charig et al. reported a discolouration: a dark band between two layers of limestone – they say it is the product of sedimentation.[48] It is natural for limestone to take on the colour of its surroundings and most limestones are coloured (if not colour banded) to some degree, so the darkness was attributed to such impurities.[51] They also mention that a complete absence of air bubbles in the rock slabs is further proof that the specimen is authentic.[48]
Description
Most of the specimens of Archaeopteryx that have been discovered come from the Solnhofen limestone in Bavaria, southern Germany, which is a Lagerstätte, a rare and remarkable geological formation known for its superbly detailed fossils laid down during the early Tithonian stage of the Jurassic period,[52] approximately 150.8–148.5 million years ago.[53]
Archaeopteryx was roughly the size of a raven,[4] with broad wings that were rounded at the ends and a long tail compared to its body length. It could reach up to 0.5 metres (1 ft 8 in) in body length and 0.7 metres (2 ft 4 in) in wingspan, with an estimated mass of 0.5 to 1 kilogram (1.1 to 2.2 lb).[4][54] Archaeopteryx feathers, although less documented than its other features, were very similar in structure to modern-day bird feathers.[52] Despite the presence of numerous avian features,[55] Archaeopteryx had many non-avian theropod dinosaur characteristics. Unlike modern birds, Archaeopteryx had small teeth,[52] as well as a long bony tail, features which Archaeopteryx shared with other dinosaurs of the time.[56]
Because it displays features common to both birds and non-avian dinosaurs, Archaeopteryx has often been considered a link between them.
Archaeopteryx had three separate digits on each fore-leg each ending with a "claw". Few birds have such features. Some birds, such as ducks, swans, Jacanas (Jacana sp.), and the hoatzin (Opisthocomus hoazin), have them concealed beneath their leg-feathers.[61]
Plumage
Specimens of Archaeopteryx were most notable for their well-developed
The body plumage of Archaeopteryx is less well-documented and has only been properly researched in the well-preserved
A patch of pennaceous feathers is found running along its back, which was quite similar to the contour feathers of the body plumage of modern birds in being symmetrical and firm, although 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: decomposed and fluffy, and possibly even appearing more like fur than feathers in life (although not in their microscopic structure). These occur on the remainder of the body—although some feathers did not fossilize and others were obliterated during preparation, leaving bare patches on specimens—and the lower neck.[63]
There is no indication of feathering on the upper neck and head. While these conceivably may have been nude, this may still be an artefact of preservation. It appears that most Archaeopteryx specimens became embedded in
Colouration
In 2011, graduate student Ryan Carney and colleagues performed the first colour study on an Archaeopteryx specimen.
Classification
Today, fossils of the genus Archaeopteryx are usually assigned to one or two species, A. lithographica and A. siemensii, but their taxonomic history is complicated. Ten names have been published for the handful of specimens. As interpreted today, the name A. lithographica only referred to the single feather described by
It has been noted that the feather, the first specimen of Archaeopteryx described, does not correspond well with the flight-related feathers of Archaeopteryx. It certainly is a
Below is a cladogram published in 2013 by Godefroit et al.[3]
Avialae |
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Species
It has been argued that all the specimens belong to the same species, A. lithographica.[78] Differences do exist among the specimens, and while some researchers regard these as due to the different ages of the specimens, some may be related to actual species diversity. 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 the possible presence of a sternum. Due to these differences, most individual specimens have been given their own species name 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 as Wellnhoferia grandis.[21]
In 2007, a review of all well-preserved specimens including the then-newly discovered Thermopolis specimen concluded that two distinct species of Archaeopteryx could be supported: A. lithographica (consisting of at least the London and Solnhofen specimens), and A. siemensii (consisting of at least the Berlin, Munich, and Thermopolis specimens). The two species are distinguished primarily by large flexor tubercles on the foot claws in A. lithographica (the claws of A. siemensii specimens being relatively simple and straight). A. lithographica also had a constricted portion of the crown in some teeth and a stouter metatarsus. A supposed additional species, Wellnhoferia grandis (based on the Solnhofen specimen), seems to be indistinguishable from A. lithographica except in its larger size.[22]
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
- Archaeopteryx lithographica Meyer, 1861 [conserved name]
- Archaeopterix lithographica Anon., 1861 [lapsus]
- Griphosaurus problematicus Wagner, 1862 [rejected name 1961 per ICZN Opinion 607]
- Griphornis longicaudatus Owen vide Woodward, 1862 [rejected name 1961 per ICZN Opinion 607]
- Archaeopteryx macrura Owen, 1862 [rejected name 1961 per ICZN Opinion 607]
- Archaeopteryx oweni Petronievics, 1917 [rejected name 1961 per ICZN Opinion 607]
- Archaeopteryx recurva Howgate, 1984
- Jurapteryx recurva (Howgate, 1984) Howgate, 1985
- Wellnhoferia grandis Elżanowski, 2001
- Archaeopteryx siemensii Dames, 1897
- Archaeornis siemensii (Dames, 1897) Petronievics, 1917[22]
- Archaeopteryx bavarica Wellnhofer, 1993
"Archaeopteryx" vicensensis (Anon. fide Lambrecht, 1933) is a nomen nudum for what appears to be an undescribed pterosaur.
Phylogenetic position
Modern palaeontology has often classified Archaeopteryx as the most primitive bird. However, it is not thought to be a true ancestor of modern birds, but rather a close relative of that ancestor.[79] Nonetheless, Archaeopteryx was often used as a model of the true ancestral bird. Several authors have done so.[80] Lowe (1935)[81] and Thulborn (1984)[82] questioned whether Archaeopteryx truly was the first bird. They suggested that Archaeopteryx was a dinosaur that was no more closely related to birds than were other dinosaur groups. Kurzanov (1987) suggested that Avimimus was more likely to be the ancestor of all birds than Archaeopteryx.[83] Barsbold (1983)[84] and Zweers and Van den Berge (1997)[85] noted that many maniraptoran lineages are extremely birdlike, and they suggested that different groups of birds may have descended from different dinosaur ancestors.
The discovery of the closely related
Phylogenetic studies conducted by Senter, et al. (2012) and Turner, Makovicky, and Norell (2012) also found Archaeopteryx to be more closely related to living birds than to dromaeosaurids and troodontids.[87][88] On the other hand, Godefroit et al. (2013) recovered Archaeopteryx as more closely related to dromaeosaurids and troodontids in the analysis included in their description of Eosinopteryx brevipenna. The authors used a modified version of the matrix from the study describing Xiaotingia, adding Jinfengopteryx elegans and Eosinopteryx brevipenna to it, as well as adding four additional characters related to the development of the plumage. Unlike the analysis from the description of Xiaotingia, the analysis conducted by Godefroit, et al. did not find Archaeopteryx to be related particularly closely to Anchiornis and Xiaotingia, which were recovered as basal troodontids instead.[89]
Agnolín and Novas (2013) found Archaeopteryx and (possibly synonymous) Wellnhoferia to be form a clade sister to the lineage including Jeholornis and Pygostylia, with Microraptoria, Unenlagiinae, and the clade containing Anchiornis and Xiaotingia being successively closer outgroups to the Avialae (defined by the authors as the clade stemming from the last common ancestor of Archaeopteryx and Aves).[90] Another phylogenetic study by Godefroit, et al., using a more inclusive matrix than the one from the analysis in the description of Eosinopteryx brevipenna, also found Archaeopteryx to be a member of Avialae (defined by the authors as the most inclusive clade containing Passer domesticus, but not Dromaeosaurus albertensis or Troodon formosus). Archaeopteryx was found to form a grade at the base of Avialae with Xiaotingia, Anchiornis, and Aurornis. Compared to Archaeopteryx, Xiaotingia was found to be more closely related to extant birds, while both Anchiornis and Aurornis were found to be more distantly so.[3]
Hu et al. (2018),[91] Wang et al. (2018)[92] and Hartman et al. (2019)[93] found Archaeopteryx to have been a deinonychosaur instead of an avialan. More specifically, it and closely related taxa were considered basal deinonychosaurs, with dromaeosaurids and troodontids forming together a parallel lineage within the group. Because Hartman et al. found Archaeopteryx isolated in a group of flightless deinonychosaurs (otherwise considered "anchiornithids"), they considered it highly probable that this animal evolved flight independently from bird ancestors (and from Microraptor and Yi). The following cladogram illustrates their hypothesis regarding the position of Archaeopteryx:
Paraves |
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The authors, however, found that the Archaeopteryx being an avialan was only slightly less likely than this hypothesis, and as likely as Archaeopterygidae and Troodontidae being sister clades.[93]
Palaeobiology
Flight
As in the wings of modern birds, the flight feathers of Archaeopteryx were somewhat asymmetrical and the tail feathers were rather broad. This implies that the wings and tail were used for lift generation, but it is unclear whether Archaeopteryx was capable of flapping flight or simply a glider. 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—may indicate that Archaeopteryx was unable to lift its wings above its back, a requirement for the upstroke found in modern flapping flight. According to a study by Philip Senter in 2006, Archaeopteryx was indeed unable to use flapping flight as modern birds do, but it may well have used a downstroke-only flap-assisted gliding technique.[94] However, a more recent study solves this issue by suggesting a different flight stroke configuration for non-avian flying theropods.[95]
Archaeopteryx wings were relatively large, which would have resulted in a low stall speed and reduced
The feathers of Archaeopteryx were asymmetrical. This has been interpreted as evidence that it was a flyer, because flightless birds tend to have symmetrical feathers. Some scientists, including Thomson and Speakman, have questioned this. They studied more than 70 families of living birds, and found that some flightless types do have a range of asymmetry in their feathers, and that the feathers of Archaeopteryx fall into this range.[96] The degree of asymmetry seen in Archaeopteryx is more typical for slow flyers than for flightless birds.[97]
In 2010, Robert L. Nudds and Gareth J. Dyke in the journal Science published a paper in which they analysed the
In 2004, scientists analysing a detailed
Recent studies of flight feather barb geometry reveal that modern birds possess a larger barb angle in the trailing vane of the feather, whereas Archaeopteryx lacks this large barb angle, indicating potentially weak flight abilities.[108]
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
In March 2018, scientists reported that Archaeopteryx was likely capable of a flight stroke cycle morphologically closer to the grabbing motion of
Studies of Archaeopteryx's feather sheaths revealed that like modern birds, it had a center-out, flight related molting strategy. As it was a weak flier, this was extremely advantageous in preserving its maximum flight performance.[111]
Growth
An
Daily activity patterns
Comparisons between the
Palaeoecology
The richness and diversity of the
The excellent preservation of Archaeopteryx fossils and other terrestrial fossils found at
The islands that surrounded the Solnhofen lagoon were low lying,
The lifestyle of Archaeopteryx is difficult to reconstruct and there are several theories regarding it. Some researchers suggest that it was primarily adapted to life on the ground,[118] while other researchers suggest that it was principally arboreal on the basis of the curvature of the claws[119] which has since been questioned.[120] The absence of trees does not preclude Archaeopteryx from an arboreal lifestyle, as several species of bird live exclusively in low shrubs. Various aspects of the morphology of Archaeopteryx point to either an arboreal or ground existence, including the length of its legs and the elongation in its feet; some authorities consider it likely to have been a generalist capable of feeding in both shrubs and open ground, as well as along the shores of the lagoon.[116] It most likely hunted small prey, seizing it with its jaws if it was small enough, or with its claws if it was larger.
See also
- Dinosaur coloration
- Evolution of birds
- Feathered dinosaur
- Origin of birds
- Ostromia
- Rhamphorhynchus
- Temporal paradox (paleontology)
- Xiaotingia
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Further reading
- G. R. de Beer (1954). Archaeopteryx lithographica: a study based upon the British Museum specimen. Trustees of the British Museum, London.
- P. Chambers (2002). Bones of Contention: The Fossil that Shook Science. John Murray, London. ISBN 0-7195-6059-4.
- A. Feduccia (1996). The Origin and Evolution of Birds. Yale University Press, New Haven. ISBN 0-300-06460-8.
- The Origin of Birds. Witherby, London.
- T. H. Huxley. (1871). Manual of the anatomy of vertebrate animals. London.
- H. von Meyer (1861). Archaeopterix lithographica (Vogel-Feder) und Pterodactylus von Solenhofen. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde. 1861: 678–679, plate V. [Article in German]. Full text, Google Books.
- P. Shipman (1998). Taking Wing: Archaeopteryx and the Evolution of Bird Flight. Weidenfeld & Nicolson, London. ISBN 0-297-84156-4.
- P. Wellnhofer (2008). Archaeopteryx – Der Urvogel von Solnhofen (in German). Verlag Friedrich Pfeil, Munich. ISBN 978-3-89937-076-8.
External links
- All About Archaeopteryx, from Talk.Origins.
- Use of SSRL X-ray takes 'transformative glimpse' – A look at chemicals linking birds and dinosaurs
- Archaeopteryx: An Early Bird – University of California Museum of Paleontology
- Are Birds Really Dinosaurs? – University of California Museum of Paleontology