Wagner, 1862 (rejected name)
Woodward, 1862 (rejected name)
Wagner, 1862 vide Woodward 1862 nomen rejectum
Owen 1862 vide Woodward 1862 nomen rejectum
(Owen 1862 vide Woodward 1862) Owen 1862 vide Brodkorb 1863 nomen
Owen, 1862 (rejected name)
(Dames, 1897) Peteronievics vide Petroneivics & Woodward 1917
Petronievics, 1917 (rejected name)
(Howgate, 1984) Howgate 1985
Archaeopteryx (/ˌɑːrkiːˈɒptərɪks/), meaning "old wing"
(sometimes referred to by its German name Urvogel ("original bird" or
"first bird")), is a genus of bird-like dinosaurs that is transitional
between non-avian feathered dinosaurs and modern birds. The name
derives from the ancient Greek ἀρχαῖος (archaīos) meaning
"ancient", and πτέρυξ (ptéryx), meaning "feather" or "wing".
Between the late nineteenth century and the early twenty-first
Archaeopteryx had been generally accepted by palaeontologists
and popular reference books as the oldest known bird (member of the
group Avialae). Older potential avialans have since been
identified, including Anchiornis, Xiaotingia, and Aurornis.
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, 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
These features make
Archaeopteryx a clear candidate for a transitional
fossil between non-avian dinosaurs and birds. 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. That same year, the first complete specimen
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.
Most of these eleven fossils include impressions of feathers. Because
these feathers are of an advanced form (flight feathers), these
fossils are evidence that the evolution of feathers began before the
Late Jurassic. The type specimen of
Archaeopteryx was discovered
just two years after
Charles Darwin published On the Origin of
Archaeopteryx seemed to confirm Darwin's theories and has
since become a key piece of evidence for the origin of birds, the
transitional fossils debate, and confirmation of evolution.
In March 2018, scientists reported that
Archaeopteryx was likely
capable of flight, but in a manner substantially different from that
of modern birds.
2.3 Daily activity patterns
4 History of discovery
6.2 Phylogenetic position
7 See also
9 Further reading
10 External links
Specimens compared to a human in scale
Most of the specimens of
Archaeopteryx that have been discovered come
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
Jurassic period, approximately 150.8–148.5 million years
Archaeopteryx was roughly the size of a raven, with broad wings
that were rounded at the ends and a long tail compared to its body
length. It could reach up to 500 millimetres (20 in) in body
length, with an estimated mass of 0.8 to 1 kilogram (1.8 to
Archaeopteryx feathers, although less documented than
its other features, were very similar in structure to modern-day bird
feathers. Despite the presence of numerous avian features,
Archaeopteryx had many non-avian theropod dinosaur characteristics.
Unlike modern birds,
Archaeopteryx had small teeth, as well as a
long bony tail, features which
Archaeopteryx shared with other
dinosaurs of the time.
Because it displays features common to both birds and non-avian
Archaeopteryx has often been considered a link between
them. In the 1970s, John Ostrom, following Thomas Henry Huxley's
lead in 1868, argued that birds evolved within theropod dinosaurs and
Archaeopteryx was a critical piece of evidence for this argument; it
had several avian features, such as a wishbone, flight feathers,
wings, and a partially reversed first toe along with 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
Archaeopteryx was remarkably similar to the theropod family
Anatomical illustration comparing the "frond-tail" of Archaeopteryx
with the "fan-tail" of a modern bird
Specimens of Archaeopteryx
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 did not yet bear a
separately movable tuft of stiff feathers.
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 (they lack barbicels as in ratites). In part
they are firm and thus capable of supporting flight.
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.
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 anoxic sediment after drifting some time on their backs in
the sea—the head, neck and the tail are generally bent downward,
which suggests that the specimens had just started to rot when they
were embedded, with tendons and muscle relaxing so that the
characteristic shape (death pose) of the fossil specimens was
achieved. This would mean that the skin already was 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.
Artist's restoration illustrating one interpretation of Carney's
In 2011, graduate student Ryan Carney and colleagues performed the
first colour study on an
Archaeopteryx specimen. Using scanning
electron microscopy technology and energy-dispersive X-ray analysis,
the team was able to detect the structure of melanosomes in the
isolated feather specimen described in 1861. The resultant
measurements were then compared to those of 87 modern bird
species, and the original colour was calculated with a 95% likelihood
to be black. The feather was determined to be black throughout, with
heavier pigmentation in the distal tip. The feather studied was most
probably a dorsal covert, which would have partly covered the primary
feathers on the wings. The study does not mean that
entirely black, but suggests that it had some black colouration which
included the coverts. Carney pointed out that this is consistent with
what we know of modern flight characteristics, in that black
melanosomes have structural properties that strengthen feathers for
flight. In a 2013 study published in the Journal of Analytical
Atomic Spectrometry, new analyses of Archaeopteryx's feathers
revealed that the animal may have had light-coloured inner vanes. This
interpretation was based on syncrotron imaging of organic sulfur and
trace metals, used as proxies for the presence of melanin. However,
such biomarkers have been found to be non-specific for melanin,
and thus it was suggested that the purported light-and-dark patterns
were instead due to non-pigmentary organics and secondary
precipitates. In 2014, Carney and colleagues presented additional data
that the wing feather's color was matte black (as opposed to
iridescent), due to the lower aspect ratio of the melanosomes detected
in the fossil. Here they also provided evidence that the
previous counter-interpretation of the feather having a light-coloured
inner vane was not supported, due to the abundance of matte black
melanosomes in this region.
1880 photo of the Berlin specimen, showing leg feathers that were
removed subsequently, during preparation
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
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 also could have
improved its ability to fly through cluttered environments such as
trees and brush (similar wing shapes are seen in birds that 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, also
would 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. This would have
reduced stall speed by up to 6% and turning radius by up to 12%.
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. The degree of asymmetry seen in
Archaeopteryx is more typical for slow flyers than for flightless
The Munich Specimen
In 2010, Robert L. Nudds and Gareth J. Dyke in the journal Science
published a paper in which they analysed the rachises of the primary
Confuciusornis and Archaeopteryx. The analysis suggested
that the rachises on these two genera were thinner and weaker than
those of modern birds relative to body mass. The authors determined
Archaeopteryx and Confuciusornis, were unable to use flapping
flight. This study was criticized by
Philip J. Currie
Philip J. Currie and Luis
Chiappe. Chiappe suggested that it is difficult to measure the
rachises of fossilized feathers, and Currie speculated that
Confuciusornis must have been able to fly to some
degree, as their fossils are preserved in what is believed to have
been marine or lake sediments, suggesting that they must have been
able to fly over deep water.
Gregory Paul also disagreed with the
study, arguing in a 2010 response that Nudds and Dyke had
overestimated the masses of these early birds, and that more accurate
mass estimates allowed powered flight even with relatively narrow
rachises. Nudds and Dyke had assumed a mass of 250 g
(8.8 oz) for the Munich specimen Archaeopteryx, a young juvenile,
based on published mass estimates of larger specimens. Paul argued
that a more reasonable body mass estimate for the Munich specimen is
about 140 g (4.9 oz). Paul also criticized the measurements
of the rachises themselves, noting that the feathers in the Munich
specimen are poorly preserved. Nudds and Dyke reported a diameter of
0.75 mm (0.03 in) for the longest primary feather, which
Paul could not confirm using photographs. Paul measured some of the
inner primary feathers, finding rachises 1.25–1.4 mm
(0.049–0.055 in) across. Despite these criticisms, Nudds
and Dyke stood by their original conclusions. They claimed that Paul's
statement, that an adult
Archaeopteryx would have been a better flyer
than the juvenile Munich specimen, was dubious. This, they reasoned,
would require an even thicker rachis, evidence for which has not yet
been presented. Another possibility is that they had not achieved
true flight, but instead used their wings as aids for extra lift while
running over water after the fashion of the basilisk lizard, which
could explain their presence in lake and marine deposits (see
Evolution of bird flight).
Replica of the
In 2004, scientists analysing a detailed CT scan of the braincase of
Archaeopteryx 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. The skull scan also revealed the structure of its
inner ear. The structure more closely resembles that of modern birds
than the inner ear of non-avian reptiles. These characteristics taken
together suggest that
Archaeopteryx had the keen sense of hearing,
balance, spatial perception, and coordination needed to fly.
Archaeopteryx had a cerebrum-to-brain-volume ratio 78% of the way to
modern birds from the condition of non-coelurosaurian dinosaurs such
Carcharodontosaurus or Allosaurus, which had a crocodile-like
anatomy of the brain and inner ear. Newer research shows that
Archaeopteryx brain was more complex than that of more
primitive theropods, it had a more generalized brain volume among
maniraptoran dinosaurs, even smaller than that of other non-avian
dinosaurs in several instances, which indicates the neurological
development required for flight was already a common trait in the
Recent studies of flight feather barb geometry reveal that modern
birds possess a larger barb angle in the trailing vane of the feather,
Archaeopteryx lacks this large barb angle, indicating
potentially weak flight abilities.
Reconstructed skeleton, Academy of Natural Sciences of Drexel
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 currently is
considered best-supported by morphological characters. Altogether, it
appears that the species was not particularly specialized for running
on the ground or for perching. A scenario outlined by Elżanowski in
2002 suggested 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 (mainly) by gliding down from cliffs or treetops.
Growth trends compared with other dinosaurs and birds
A histological study by Erickson, Norell, Zhongue, and others in 2009
Archaeopteryx grew relatively slowly compared to modern
birds, presumably because the outermost portions of Archaeopteryx
bones appear poorly vascularized; in living vertebrates, poorly
vascularized bone is correlated with slow growth rate. They also
assume that all known skeletons of
Archaeopteryx come from juvenile
specimens. Because the bones of
Archaeopteryx could not be
histologically sectioned in a formal skeletochronological (growth
ring) analysis, Erickson and colleagues used bone vascularity
(porosity) to estimate bone growth rate. They assumed that poorly
vascularized bone grows at similar rates in all birds and in
Archaeopteryx. The poorly vascularized bone of
have grown as slowly as that in a mallard (2.5 micrometres per day) or
as fast as that in an ostrich (4.2 micrometres per day). Using this
range of bone growth rates, they calculated how long it would take to
"grow" each specimen of
Archaeopteryx to the observed size; it may
have taken at least 970 days (there were 375 days in a Late Jurassic
year) to reach an adult size of 0.8–1 kg (1.8–2.2 lb).
The study also found that the avialans
relatively slowly, as did the dromaeosaurid Mahakala. The avialans
Ichthyornis grew relatively quickly, following a
growth trend similar to that of modern birds. One of the few
modern birds that exhibit slow growth is the flightless kiwi, and the
authors speculated that
Archaeopteryx and the kiwi had similar basal
Daily activity patterns
Comparisons between the scleral rings of
Archaeopteryx and modern
birds and reptiles indicate that it may have been diurnal, similar to
most modern birds.
Archaeopteryx chasing a juvenile Compsognathus
The richness and diversity of the
Solnhofen limestones in which all
Archaeopteryx have been found have shed light on an
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 a lack of terrestrial sediments characteristic
of rivers. Evidence of plants, although scarce, include cycads and
conifers while animals found include a large number of insects, small
lizards, pterosaurs, and Compsognathus.
The excellent preservation of
Archaeopteryx fossils and other
terrestrial fossils found at
Solnhofen indicates that they did not
travel far before becoming preserved. The
found were therefore likely to have lived on the low islands
Solnhofen lagoon rather than to have been corpses that
drifted in from farther away.
Archaeopteryx skeletons are considerably
less numerous in the deposits of
Solnhofen than those of pterosaurs,
of which seven genera have been found. The pterosaurs included
species such as
Rhamphorhynchus belonging to the Rhamphorhynchidae,
the group which dominated the niche currently occupied by seabirds,
and which became extinct at the end of the Jurassic. The pterosaurs,
which also included Pterodactylus, were common enough that it is
unlikely that the specimens found are vagrants from the larger islands
50 km (31 mi) to the north.
The islands that surrounded the
Solnhofen lagoon were low lying,
semi-arid, and sub-tropical with a long dry season and little
rain. The closest modern analogue for the
Solnhofen conditions is
said to be
Orca Basin in the northern Gulf of Mexico, although it is
much deeper than the
Solnhofen lagoons. The flora of these islands
was adapted to these dry conditions and consisted mostly of low
(3 m (10 ft)) shrubs. 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 fossilized tree pollen also is absent.
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, while other
researchers suggest that it was principally arboreal. 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. It most likely hunted
small prey, seizing it with its jaws if it was small enough, or with
its claws if it was larger.
History of discovery
Main article: Specimens of Archaeopteryx
Archaeopteryx discoveries until 2007
Over the years, twelve 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,
The single feather
The initial discovery, a single feather, was unearthed in 1860 or 1861
and described in 1861 by Hermann von Meyer. It is currently
located at the Natural History Museum of Berlin. This is generally
Archaeopteryx and was the initial holotype, but whether it
is a feather of this species, or another, as yet undiscovered,
proto-bird is unknown. There are some indications it is not from the
same animal as most of the skeletons (the "typical" A.
The first skeleton, known as the
London Specimen (BMNH 37001), 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 to the Natural History Museum in London, where
it remains. Missing most of its head and neck, it was described in
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,
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."
The Greek term "pteryx" (πτέρυξ) primarily means "wing", but can
also designate merely "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
Archaeopteryx among German scholars in the late nineteenth century.
In English, "ancient pinion" offers a rough approximation.
Since then twelve specimens have been recovered:
The Berlin Specimen (HMN 1880/81) was discovered in 1874 or 1875 on
the Blumenberg near Eichstätt, Germany, by farmer Jakob Niemeyer. He
sold this precious fossil for the money to buy a cow in 1876, to
innkeeper Johann Dörr, who again sold it to Ernst Otto Häberlein,
the son of K. Häberlein. Placed on sale between 1877 and 1881, with
potential buyers including O. C. Marsh of Yale University's Peabody
Museum, it eventually was bought for 20,000 Goldmark by the Berlin's
Natural History Museum, where it now is displayed. The transaction was
financed by Ernst Werner von Siemens, founder of the famous company
that bears his name. Described in 1884 by Wilhelm Dames, it is the
most complete specimen, and the first with a complete head. In 1897 it
was named by Dames as a new species, A. siemensii; though often
considered a synonym of A. lithographica, several 21st century studies
have concluded that it is a distinct species which includes the
Berlin, Munich, and Thermopolis specimens.
Cast of the Maxberg Specimen
Composed of a torso, the Maxberg Specimen (S5) was discovered in 1956
near Langenaltheim; it was brought to the attention of professor
Florian Heller in 1958 and described by him in 1959. The specimen is
missing its head and tail, although the rest of the skeleton is mostly
intact. Although it was once exhibited at the
Maxberg Museum in
Solnhofen, it is currently missing. It belonged to Eduard Opitsch, who
loaned it to the museum until 1974. After his death in 1991, it was
discovered that the specimen was missing and may have been stolen or
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
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
Eichstätt Specimen, once considered a distinct genus, Jurapteryx
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).
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
Wellnhoferia grandis. It is missing only portions of the
neck, tail, backbone, and head.
The Munich Specimen (BSP 1999 I 50, formerly known as the
Solenhofer-Aktien-Verein Specimen) was discovered on 3 August 1992
Langenaltheim and described in 1993 by Wellnhofer. It is
currently located at the
Paläontologisches Museum München
Paläontologisches Museum München in Munich,
to which it was sold in 1999 for 1.9 million Deutschmark. What was
initially believed to be a bony sternum turned out to be part of the
coracoid, but a cartilaginous sternum may have been present. Only
the front of its face is missing. It has been used as the basis for a
distinct species, A. bavarica, but more recent studies suggest it
belongs to A. siemensii.
An eighth, fragmentary specimen was discovered in 1990, not in
Solnhofen limestone, but in somewhat younger sediments at Daiting,
Suevia. Therefore, it is known as the
Daiting Specimen, and had been
known since 1996 only from a cast, briefly shown at the
Naturkundemuseum in Bamberg. The original was purchased by
palaeontologist Raimund Albertsdörfer in 2009. It was on display
for the first time with six other original fossils of
the Munich Mineral Show in October 2009. A first, quick look by
scientists indicates that this specimen might represent a new species
of Archaeopteryx. It was found in a limestone bed that was a few
hundred thousand years younger than the other finds.
Bürgermeister-Müller ("chicken wing") Specimen
Another fragmentary fossil was found in 2000. It is in private
possession and, since 2004, on loan to the Bürgermeister-Müller
Museum in Solnhofen, so it is called the Bürgermeister-Müller
Specimen; the institute itself officially refers to it as the
"Exemplar of the families Ottman & Steil, Solnhofen". As the
fragment represents the remains of a single wing of Archaeopteryx, the
popular name of this fossil is "chicken wing".
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 in 2 December 2005 Science journal article as
Archaeopteryx specimen with theropod features"; it
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. This has been interpreted
as evidence of theropod ancestry. In 1988,
Gregory S. Paul
Gregory S. Paul claimed to
have found evidence of a hyperextensible second toe, 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."
The Thermopolis Specimen was assigned to
Archaeopteryx siemensii in
2007. The specimen is considered to represent the most complete
Archaeopteryx remains yet.
The eleventh specimen
The discovery of an eleventh specimen was announced in 2011, and 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. Palaeontologists of the Ludwig
Maximilian University of Munich studied the specimen, which revealed
previously unknown features of the plumage, such as feathers on both
the upper and lower legs and metatarsus, and the only preserved tail
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. 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.
The Thermopolis Specimen
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 Meyer. In 1954 Gavin de
Beer concluded that the
London specimen was the holotype. In 1960,
Swinton accordingly proposed that the name
be placed on the official genera list making the alternative names
Griphosaurus and Griphornis invalid. The ICZN, implicitly
accepting de Beer's standpoint, did indeed suppress the plethora of
alternative names initially proposed for the first skeleton
specimens, which mainly resulted from the acrimonious dispute
between Meyer and his opponent
Johann Andreas Wagner
Johann Andreas Wagner (whose
Griphosaurus problematicus – "problematic riddle-lizard" – was a
vitriolic sneer at Meyer's Archaeopteryx). In addition, in 1977,
the Commission ruled that the first species name of the Haarlem
specimen, crassipes, described by Meyer as a pterosaur before its true
nature was realized, was not to be given preference over lithographica
in instances where scientists considered them to represent the same
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 flight
feather of a contemporary species, but its size and proportions
indicate that it may belong to another, smaller species of feathered
theropod, of which only this feather is known so far. As the
feather had been designated the type specimen, the name Archaeopteryx
should then no longer be applied to the skeletons, thus creating
significant nomenclatorial confusion. In 2007, two sets of scientists
therefore petitioned the ICZN requesting that the
explicitly be made the type by designating it as the new holotype
specimen, or neotype. This suggestion was upheld by the ICZN after
four years of debate, and the
London specimen was designated the
neotype on 3 October 2011.
The twelfth specimen
Below is a cladogram published in 2013 by Godefroit et al.
Avebrevicauda (includes modern birds)
Skeletal restorations of various specimens
It has been argued that all the specimens belong to the same species,
A. lithographica. 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
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
The Berlin specimen has been designated as
Archaeornis siemensii, the
Eichstätt specimen as Jurapteryx recurva,
the Munich specimen as
Archaeopteryx bavarica, and the Solnhofen
In 2007, a review of all well-preserved specimens including the
then-newly discovered Thermopolis specimen concluded that two distinct
Archaeopteryx could be supported: A. lithographica
(consisting of at least the
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.
Haarlem specimen, which is now considered Ostromia
Solnhofen Specimen, by some considered as belonging to the genus
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.
Archaeopteryx lithographica Meyer, 1861 [conserved name]
Archaeopterix lithographica Anon., 1861 [lapsus]
Griphosaurus problematicus Wagner, 1862 [rejected name 1961 per ICZN
Griphornis longicaudatus Owen vide Woodward, 1862 [rejected name 1961
per ICZN Opinion 607]
Archaeopteryx macrura Owen, 1862 [rejected name 1961 per ICZN Opinion
Archaeopteryx oweni Petronievics, 1917 [rejected name 1961 per ICZN
Archaeopteryx recurva Howgate, 1984
Jurapteryx recurva (Howgate, 1984) Howgate, 1985
Archaeopteryx siemensii Dames, 1897
Archaeornis siemensii (Dames, 1897) Petronievics, 1917
Archaeopteryx bavarica Wellnhofer, 1993
Wellnhoferia grandis Elżanowski, 2001
"Archaeopteryx" vicensensis (Anon. fide Lambrecht, 1933) is a nomen
nudum for what appears to be an undescribed pterosaur.
Beginning in 1985, a group including astronomer
Fred Hoyle and
Lee Spetner published a series of papers claiming that the
feathers on the Berlin and
London specimens of
forged. Their claims were repudiated by Alan J. Charig
and others at the Natural History Museum in London. 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, without
realizing that feathers themselves would have caused a textural
difference. They also misinterpreted the fossils, claiming that
the tail was forged as one large feather, when visibly this is not
the case. In addition, they claimed that the other specimens of
Archaeopteryx known at the time did not have feathers, which
is incorrect; the Maxberg and
Eichstätt specimens have obvious
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. These are common
Solnhofen fossils, because the dead animals would fall
onto hardened surfaces, which would form a natural plane for the
future slabs to split along and would leave the bulk of the fossil on
one side and little on the other.
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
so such an action would certainly backfire.
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. Spetner et al. then
attempted to show that the cracks would have propagated naturally
through their postulated cement layer, 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. 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; it was not
cement either, and is most probably a fragment of silicone rubber left
behind when moulds were made of the specimen. 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. 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. They also mention that a complete absence of air
bubbles in the rock slabs is further proof that the specimen is
Comparison of the forelimb of
Archaeopteryx (right) with that of
Modern paleontology has often classified
Archaeopteryx as the most
primitive bird. It is not thought to be a true ancestor of modern
birds, but rather, a close relative of that ancestor. Nonetheless,
Archaeopteryx was often used as a model of the true ancestral bird.
Several authors have done so. Lowe (1935) and Thulborn
(1984) 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. Barsbold (1983) and Zweers and
Van den Berge (1997) 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
Xiaotingia in 2011 led to new
phylogenetic analyses that suggested that
Archaeopteryx is a
deinonychosaur rather than an avialan, and therefore, not a "bird"
under most common uses of that term. A more thorough analysis was
published soon after to test this hypothesis, and failed to arrive at
the same result; it found
Archaeopteryx in its traditional position at
the base of Avialae, while
Xiaotingia was recovered as a basal
dromaeosaurid or troodontid. The authors of the follow-up study noted
that uncertainties still exist, and that it may not be possible to
state confidently whether or not
Archaeopteryx is a member of Avialae
or not, barring new and better specimens of relevant species.
Phylogenetic studies conducted by Senter, et al. (2012) and Turner,
Makovicky, and Norell (2012) confirmed that
Archaeopteryx was more
closely related to living birds than to dromaeosaurids and
troodontids. On the other hand, Godefroit, et al. (2013)
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
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.
Agnolín and Novas (2013) found
Archaeopteryx and (possibly
Wellnhoferia to be the basalmost avialans (
defined by the authors as including
Archaeopteryx lithographica and
Passer, their most recent common ancestor and all of its descendants),
with Microraptoria, Unenlagiinae, and the clade containing Anchiornis
Xiaotingia being successively closer outgroups to the
Avialae. Another phylogenetic study by Godefroit, et al., using a
more inclusive matrix than the one from the analysis in the
Eosinopteryx brevipenna, also found
Archaeopteryx to be
a member of
Avialae (defined by the authors as the most inclusive
Passer domesticus, but not
Archaeopteryx was found to form a grade at the
Avialae with Xiaotingia, Anchiornis, and Aurornis. Compared to
Xiaotingia was found to be more closely related to
extant birds, while both
Aurornis were found to be more
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Evolution of birds
Origin of birds
Temporal paradox (paleontology)
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Wikispecies has information related to Archaeopteryx
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
Birds Really Dinosaurs? — University of California Museum of
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