Archaeopteryx lived in theLate Jurassic around 150 million years ago, in what is now southern Germany, during a time when Europe was anarchipelago of islands in a shallow warm tropical sea, much closer to theequator than it is now. Similar in size to aEurasian magpie, with the largest individuals possibly attaining the size of araven,[5] the largest species ofArchaeopteryx could grow to about 50 cm (20 in) in length. Despite their small size, broad wings, and inferred ability to fly or glide,Archaeopteryx had more in common with other smallMesozoic dinosaurs than with modern birds. In particular, they shared the following features with thedromaeosaurids andtroodontids: jaws with sharpteeth, three fingers withclaws, a long bony tail, hyperextensible second toes ("killing claw"), feathers (which also suggestwarm-bloodedness), and various features of theskeleton.[6][7]
These features makeArchaeopteryx a clear candidate for atransitional fossil between non-avian dinosaurs and avian dinosaurs (birds).[8][9] Thus,Archaeopteryx plays an important role, not only in the study of theorigin of birds, but in the study of dinosaurs. It was named from a singlefeather in 1861,[10] the identity of which has been controversial.[11][12] That same year, the first complete specimen ofArchaeopteryx was announced. Over the years, twelve more fossils ofArchaeopteryx have surfaced. Despite variation among these fossils, most experts regard all the remains that have been discovered as belonging to a single species or at least genus, although this is still debated.[13]
Most of these 14 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.[14] Thetype specimen ofArchaeopteryx was discovered just two years afterCharles Darwin publishedOn the Origin of Species.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 ofevolution.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.[15]
Over the years, fourteen body fossil specimens ofArchaeopteryx have been found. All of the fossils come from thelimestone deposits, quarried for centuries, nearSolnhofen, Germany. These quarries excavate sediments from theSolnhofen Limestone formation and related units.[16][17] The initial specimen was the first dinosaur to be discovered with feathers.
The initial discovery, a single feather, was unearthed in 1860 or 1861 and described in 1861 byHermann von Meyer.[18] It is now in theNatural History Museum of Berlin. Though it was the initialholotype, there were indications that it might not have been from the same animal as the body fossils.[10] 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 otherArchaeopteryx feathers known, leading to the conclusion that it originated from another dinosaur.[11] 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 primarycovert feather.[12]
The first skeleton, known as theLondon Specimen (BMNH 37001),[19] was unearthed in 1861 nearLangenaltheim, Germany, and perhaps given to local physicianKarl Häberlein in return for medical services. He then sold it for £700 (roughly £83,000 in 2020[20]) to theNatural History Museum in London, where it remains.[16] Missing most of its head and neck, it was described in 1863 byRichard Owen asArchaeopteryx macrura, allowing for the possibility it did not belong to the same species as the feather. In the subsequent fourth edition of hisOn the Origin of Species,[21] Charles Darwin said the following about the discovery.[22]
Until quite recently these authors might have maintained, and some have maintained, that the whole class of birds came suddenly into existence during theeocene 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, theArchaeopteryx, 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 theoolitic 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.
Thegenus name derives from theAncient Greekἀρχαῖος (archaîos), meaning "ancient", andπτέρυξ (ptérux), meaning "feather, wing". Meyer suggested this in his description. At first he referred to a single feather which appeared to resemble a modern bird'sremex (wing feather), but he had heard of and been shown a rough sketch of theLondon 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 termSchwinge which does not necessarily mean a wing used for flying.Urschwinge was the favoured translation ofArchaeopteryx 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:
TheBerlin Specimen (HMN 1880/81) was discovered in 1874 or 1875 on the Blumenberg nearEichstätt, Germany, by farmer Jakob Niemeyer. He sold this preciousfossil 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 includingO. C. Marsh ofYale University's Peabody Museum, it eventually was bought for 20,000Goldmark by the Berlin's Natural History Museum, where it now is displayed. The transaction was financed byErnst Werner von Siemens, founder ofthe company that bears his name.[16] Described in 1884 byWilhelm 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 ofA. lithographica, several 21st century studies have concluded that it is a distinct species which includes the Berlin, Munich, and Thermopolis specimens.[23][24]
Cast of the Maxberg Specimen
Composed of a torso, theMaxberg Specimen (S5) was discovered in 1956 nearLangenaltheim; it was brought to the attention of professorFlorian 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 theMaxberg Museum inSolnhofen, it is currently missing. It belonged toEduard 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 sold.[25]
TheHaarlem Specimen (TM 6428/29, also known as theTeylers Specimen) was discovered in 1855 nearRiedenburg, Germany, and described as aPterodactylus crassipes in 1857 by Meyer. It was reclassified in 1970 byJohn Ostrom and is currently located at theTeylers Museum inHaarlem, 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 genusOstromia, considered more closely related toAnchiornis from China.[26]
Eichstätt Specimen, once considered a distinct genus,Jurapteryx
TheEichstätt Specimen (JM 2257) was discovered in 1951 nearWorkerszell, Germany, and described byPeter Wellnhofer in 1974. Currently located at theJura 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).[27]
TheSolnhofen Specimen (unnumbered specimen) was discovered in the 1970s near Eichstätt, Germany, and described in 1988 by Wellnhofer. Currently located at theBürgermeister-Müller-Museum in Solnhofen, it originally was classified asCompsognathus 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.[28]
TheMunich Specimen (BSP 1999 I 50, formerly known as theSolenhofer-Aktien-Verein Specimen) was discovered on 3 August 1992 near Langenaltheim and described in 1993 by Wellnhofer. It is currently located at thePaläontologisches Museum München in Munich, to which it was sold in 1999 for 1.9 millionDeutschmark. What was initially believed to be a bonysternum turned out to be part of thecoracoid,[29] but acartilaginous 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,[30] but more recent studies suggest it belongs toA. siemensii.[24]
An eighth, fragmentary specimen was discovered in 1990 in the youngerMörnsheim Formation atDaiting,Suevia. Therefore, it is known as theDaiting Specimen, and had been known since 1996 only from a cast, briefly shown at theNaturkundemuseum inBamberg. The original was purchased by palaeontologist Raimund Albertsdörfer in 2009.[31] It was on display for the first time with six other original fossils ofArchaeopteryx at the Munich Mineral Show in October 2009.[32] The Daiting Specimen was subsequently namedArchaeopteryx albersdoerferi by Kundrat et al. (2018).[33][34] After a lengthy period in a closed private collection, it was moved to the Museum of Evolution atKnuthenborg Safaripark (Denmark) in 2022, where it has since been on display and also been made available for researchers.[35][36]
Bürgermeister-Müller ("chicken wing") Specimen
Another fragmentary fossil was found in 2000. It is in private possession and, since 2004, on loan to theBürgermeister-Müller Museum in Solnhofen, so it is called theBürgermeister-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 ofArchaeopteryx, it is colloquially known as "chicken wing".[37]
Details of the Wyoming Dinosaur Center Archaeopteryx (WDC-CSG-100)
Long in a private collection in Switzerland, theThermopolis Specimen (WDC CSG 100) was discovered in Bavaria and described in 2005 by Mayr, Pohl, and Peters. Donated to theWyoming Dinosaur Center inThermopolis, 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 2005Science journal article as "A well-preservedArchaeopteryx specimen with theropod features"; it shows thatArchaeopteryx lacked a reversed toe—a universal feature of birds—limiting its ability to perch on branches and implying a terrestrial or trunk-climbing lifestyle.[38] This has been interpreted as evidence oftheropod ancestry. In 1988,Gregory S. Paul claimed to have found evidence of a hyperextensible second toe,[39] 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."[17] The Thermopolis Specimen was assigned toArchaeopteryx siemensii in 2007.[24] The specimen is considered to represent the most complete and best-preservedArchaeopteryx remains yet.[24]
The eleventh specimen
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.[40][41] Palaeontologists of theLudwig 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 andmetatarsus, and the only preserved tail tip.[42][43]
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.[44] 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 thePainten Formation and somewhat older than the other specimens.[45]
A thirteenth specimen,SMNK-PAL 10,000, was published in January 2025, this one from theMörnsheim Formation. It preserves the right forelimb, shoulder, and fragments of the other limbs, with various features of the shoulder and forelimb resemblingArchaeopteryx more than any other avialan within the Mörnsheim Formation. However, due to the fragmentary nature of this specimen, it cannot be assigned to a specific species withinArchaeopteryx.[46]
The Chicago archaeopteryx
The existence of a fourteenth specimen (theChicago specimen) was first informally announced in 2024 by theField Museum in Chicago, US. One of two specimens in an institution outside Europe, the specimen was originally identified in a private collection inSwitzerland, and had been acquired by these collectors in 1990, prior to Germany's 2015 ban on exportingArchaeopteryx specimens. The specimen was acquired by the Field Museum in 2022, and went on public display in 2024 following two years of preparation.[47][48] In 2025, the paleornithologistJingmai O'Connor and colleagues officially published a study describing this fourteenth specimen, reporting the first knowntertials (specialized inner secondary flight feathers) and other novel features inArchaeopteryx.[13]
Beginning in 1985, an amateur group including astronomerFred Hoyle and physicistLee Spetner, published a series of papers claiming that the feathers on the Berlin and London specimens ofArchaeopteryx were forged.[49][50][51][52] Their claims were repudiated byAlan J. Charig and others at theNatural History Museum in London.[53] Most of their supposed evidence for a forgery was based on unfamiliarity with the processes oflithification; for example, they proposed that, based on the difference in texture associated with the feathers, feather impressions were applied to a thin layer ofcement,[50] without realizing that feathers themselves would have caused a textural difference.[53] They also misinterpreted the fossils, claiming that the tail was forged as one large feather,[50] when visibly this is not the case.[53] In addition, they claimed that the other specimens ofArchaeopteryx known at the time did not have feathers,[49][50] which is incorrect; the Maxberg and Eichstätt specimens have obvious feathers.[53]
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 thecounterslab.[49][51] These 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 and would leave the bulk of the fossil on one side and little on the other.[53]
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.[54]
Chariget 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.[53] Spetneret al. then attempted to show that the cracks would have propagated naturally through their postulated cement layer,[55] but neglected to account for the fact that the cracks were old and had been filled withcalcite, and thus were not able to propagate.[54] They also attempted to show the presence of cement on the London specimen throughX-ray spectroscopy, and did find something that was not rock;[55] it was not cement either, and is most probably a fragment of silicone rubber left behind when moulds were made of the specimen.[54] 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. Chariget al. reported a discolouration: a dark band between two layers of limestone – they say it is the product of sedimentation.[53] 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.[56] They also mention that a complete absence of air bubbles in the rock slabs is further proof that the specimen is authentic.[53]
Most of the specimens ofArchaeopteryx that have been discovered come from the Solnhofen limestone in Bavaria, southern Germany, which is aLagerstätte, a rare and remarkable geological formation known for its superbly detailed fossils laid down during the early Tithonian stage of the Jurassic period,[57] approximately 150.8–148.5million years ago.[58]
Archaeopteryx was roughly the size of araven,[5] with broad wings that were rounded at the ends and a long tail compared to its body length. It could reach up to 50 centimetres (1 ft 8 in) in body length and 70 centimetres (2 ft 4 in) inwingspan, with an estimated mass of 500 to 1,000 grams (18 to 35 oz).[5][59]Archaeopteryx feathers, although less documented than its other features, were very similar in structure to modern-day bird feathers.[57] Despite the presence of numerous avian features,[60]Archaeopteryx had many non-aviantheropod dinosaur characteristics. Unlike modern birds,Archaeopteryx had small teeth,[57] as well as a long bony tail, features whichArchaeopteryx shared with other dinosaurs of the time.[61]
Because it displays features common to both birds and non-avian dinosaurs,Archaeopteryx has often been considered a link between them.[57] In the 1970s,John Ostrom, followingThomas Henry Huxley's lead in 1868, argued that birds evolved within theropod dinosaurs andArchaeopteryx 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 theankle bone,interdental plates, anobturator process of theischium, and long chevrons in the tail. In particular, Ostrom found thatArchaeopteryx was remarkably similar to the theropod familyDromaeosauridae.[62][63][64][65]
Archaeopteryx had three separate digits on each fore-leg each ending with a "claw". Few birds have such features. Some birds, such asducks,swans,Jacanas (Jacana sp.), and thehoatzin (Opisthocomus hoazin), have them concealed beneath their leg-feathers.[66]
Anatomical illustration comparing the "frond-tail" ofArchaeopteryx with the "fan-tail" of a modern bird
Specimens ofArchaeopteryx were most notable for their well-developedflight feathers. They were markedly asymmetrical and showed the structure of flight feathers in modern birds, with vanes given stability by abarb-barbule-barbicel arrangement.[67] The tail feathers were less asymmetrical, again in line with the situation in modern birds and also had firm vanes. Thethumb did not yet bear a separately movable tuft ofstiff feathers.
The body plumage ofArchaeopteryx is less well-documented and has only been properly researched in the well-preservedBerlin 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 ofArchaeopteryx. 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 inratites).[68] In part they are firm and thus capable of supporting flight.[69]
A patch ofpennaceous 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 dinosaurSinosauropteryx: 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.[68]
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 mostArchaeopteryx specimens became embedded inanoxic 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.[70] 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.[23]
Artist's restoration illustrating one interpretation of Carney's study[71]
In 2011, graduate student Ryan Carney and colleagues performed the first colour study on anArchaeopteryx specimen.[71] Usingscanning electron microscopy technology andenergy-dispersive X-ray analysis, the team was able to detect the structure ofmelanosomes in the isolated feather specimen described in 1861. The resultant measurements were then compared to those of 87modern 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 dorsalcovert, which would have partly covered the primary feathers on the wings. The study does not mean thatArchaeopteryx was entirely black, but suggests that it had some black colouration which included the coverts. Carney pointed out that this is consistent with what is known of modern flight characteristics, in that black melanosomes have structural properties that strengthen feathers for flight.[72] In a 2013 study published in theJournal of Analytical Atomic Spectrometry, new analyses ofArchaeopteryx's feathers revealed that the animal may have had complex light- and dark-coloured plumage, with heavier pigmentation in the distal tips and outer vanes.[73] This analysis of colour distribution was based primarily on the distribution of sulphate within the fossil. An author on the previousArchaeopteryx colour study argued against the interpretation of such biomarkers as an indicator of eumelanin in the fullArchaeopteryx specimen.[74] Carney and other colleagues also argued against the 2013 study's interpretation of the sulphate and trace metals,[75][76] and in a 2020 study published inScientific Reports demonstrated that the isolated covert feather was entirely matte black (as opposed to black and white, or iridescent) and that the remaining "plumage patterns ofArchaeopteryx remain unknown".[12]
Today, fossils of the genusArchaeopteryx are usually assigned to one or two species,A. lithographica andA. siemensii, but their taxonomic history is complicated. Ten names have been published for the handful of specimens. As interpreted today, the nameA. lithographica only referred to the single feather described byMeyer. In 1954Gavin de Beer concluded that the London specimen was theholotype. In 1960, Swinton accordingly proposed that the nameArchaeopteryx lithographica be placed on the official genera list making the alternative namesGriphosaurus andGriphornis invalid.[77] TheICZN, implicitly accepting De Beer's standpoint, did indeed suppress the plethora of alternative names initially proposed for the first skeleton specimens,[78] which mainly resulted from the acrimonious dispute between Meyer and his opponentJohann Andreas Wagner (whoseGriphosaurus problematicus—'problematicriddle-lizard'—was a vitriolic sneer at Meyer'sArchaeopteryx).[79] In addition, in 1977, the Commission ruled that the first species name of the Haarlem specimen,crassipes, described by Meyer as apterosaur before its true nature was realized, was not to be given preference overlithographica in instances where scientists considered them to represent the same species.[8][80]
It has been noted that the feather, the first specimen ofArchaeopteryx described, does not correspond well with the flight-related feathers ofArchaeopteryx. It certainly is aflight feather of a contemporary species, but its size and proportions indicate that it may belong to another, smaller species offeathered theropod, of which only this feather is known so far.[10] As the feather had been designated thetype specimen, the nameArchaeopteryx should then no longer be applied to the skeletons, thus creating significantnomenclatorial confusion. In 2007, two sets of scientists therefore petitioned the ICZN requesting that the London specimen explicitly be made the type by designating it as the new holotype specimen, orneotype.[81] This suggestion was upheld by the ICZN after four years of debate, and the London specimen was designated the neotype on 3 October 2011.[82]
It has been argued that all the specimens belong to the same species,A. lithographica.[83] 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 asternum. 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 asArchaeornis siemensii, the Eichstätt specimen asJurapteryx recurva, the Munich specimen asArchaeopteryx bavarica, and the Solnhofen specimen asWellnhoferia grandis.[23]
In 2007, a review of all well-preserved specimens including the then-newly discovered Thermopolis specimen concluded that two distinct species ofArchaeopteryx could be supported:A. lithographica (consisting of at least the London and Solnhofen specimens), andA. siemensii (consisting of at least the Berlin, Munich, and Thermopolis specimens). The two species are distinguished primarily by large flexortubercles on the foot claws inA. lithographica (the claws ofA. 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 fromA. lithographica except in its larger size.[24]
The Solnhofen Specimen, by some considered as belonging to the genusWellnhoferia
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 inzoological nomenclature, putting an author's name in parentheses denotes that thetaxon was originally described in a different genus.
Comparison of the forelimb ofArchaeopteryx (right) with that ofDeinonychus (left)
Modern palaeontology has often classifiedArchaeopteryx 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.[84] Nonetheless,Archaeopteryx was often used as a model of the true ancestral bird. Several authors have done so.[85] Lowe (1935)[86] and Thulborn (1984)[87] questioned whetherArchaeopteryx truly was the first bird. They suggested thatArchaeopteryx was a dinosaur that was no more closely related to birds than were other dinosaur groups. Kurzanov (1987) suggested thatAvimimus was more likely to be the ancestor of all birds thanArchaeopteryx.[88] Barsbold (1983)[89] and Zweers and Van den Berge (1997)[90] noted that manymaniraptoran lineages are extremely birdlike, and they suggested that different groups of birds may have descended from different dinosaur ancestors.
The discovery of the closely relatedXiaotingia in 2011 led to new phylogenetic analyses that suggested thatArchaeopteryx is adeinonychosaur rather than an avialan, and therefore, not a "bird" under most common uses of that term.[2] A more thorough analysis was published soon after to test this hypothesis, and failed to arrive at the same result; it foundArchaeopteryx in its traditional position at the base ofAvialae, whileXiaotingia 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 notArchaeopteryx is a member of Avialae or not, barring new and better specimens of relevant species.[91]
Phylogenetic studies conducted by Senter,et al. (2012) and Turner, Makovicky, and Norell (2012) also foundArchaeopteryx to be more closely related to living birds than to dromaeosaurids and troodontids.[92][93] On the other hand, Godefroitet al. (2013) recoveredArchaeopteryx as more closely related to dromaeosaurids and troodontids in the analysis included in their description ofEosinopteryx brevipenna. The authors used a modified version of the matrix from the study describingXiaotingia, addingJinfengopteryx elegans andEosinopteryx brevipenna to it, as well as adding four additional characters related to the development of the plumage. Unlike the analysis from the description ofXiaotingia, the analysis conducted by Godefroit,et al. did not findArchaeopteryx to be related particularly closely toAnchiornis andXiaotingia, which were recovered as basal troodontids instead.[94]
Agnolín and Novas (2013) foundArchaeopteryx and (possibly synonymous)Wellnhoferia to form a clade sister to the lineage includingJeholornis and Pygostylia, withMicroraptoria,Unenlagiinae, and the clade containingAnchiornis andXiaotingia being successively closer outgroups to the Avialae (defined by the authors as the clade stemming from the last common ancestor ofArchaeopteryx and Aves).[95] Another phylogenetic study by Godefroit,et al., using a more inclusive matrix than the one from the analysis in the description ofEosinopteryx brevipenna, also foundArchaeopteryx to be a member of Avialae (defined by the authors as the most inclusive clade containingPasser domesticus, but notDromaeosaurus albertensis orTroodon formosus).Archaeopteryx was found to form agrade at the base of Avialae withXiaotingia,Anchiornis, andAurornis. Compared toArchaeopteryx,Xiaotingia was found to be more closely related to extant birds, while bothAnchiornis andAurornis were found to be more distantly so.[3]
Huet al. (2018),[96] Wanget al. (2018)[97] and Hartmanet al. (2019)[98] foundArchaeopteryx 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 Hartmanet al. foundArchaeopteryx isolated in a group of flightless deinonychosaurs (otherwise considered "anchiornithids"), they considered it highly probable that this animalevolved flight independently from bird ancestors (and fromMicroraptor andYi). The following cladogram illustrates their hypothesis regarding the position ofArchaeopteryx:
The authors, however, found that theArchaeopteryx being an avialan was only slightly less likely than this hypothesis, and as likely as Archaeopterygidae and Troodontidae being sister clades.[98]
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 ofArchaeopteryx 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 whetherArchaeopteryx was capable of flapping flight or simply a glider. The lack of a bonybreastbone suggests thatArchaeopteryx was not a very strong flier, but flight muscles might have attached to the thick, boomerang-shaped wishbone, the platelikecoracoids, or perhaps, to acartilaginoussternum. The sideways orientation of the glenoid (shoulder) joint betweenscapula, coracoid, andhumerus—instead of the dorsally angled arrangement found in modern birds—may indicate thatArchaeopteryx was unable to lift its wings above its back, a requirement for the upstroke found in modern flapping flight. According to a study byPhilip 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.[99] However, a more recent study solves this issue by suggesting a different flight stroke configuration for non-avian flying theropods.[100]
Archaeopteryx wings were relatively large, which would have resulted in a low stall speed and reducedturning 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 ascrows andpheasants). The presence of "hind wings", asymmetrical flight feathers stemming from the legs similar to those seen in dromaeosaurids such asMicroraptor, also would have added to the aerial mobility ofArchaeopteryx. The first detailed study of the hind wings by Longrich in 2006, suggested that the structures formed up to 12% of the totalairfoil. This would have reduced stall speed by up to 6% and turning radius by up to 12%.[69]
The feathers ofArchaeopteryx 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 ofArchaeopteryx fall into this range.[101] The degree of asymmetry seen inArchaeopteryx is more typical for slow flyers than for flightless birds.[102]
The Munich Specimen
In 2010, Robert L. Nudds and Gareth J. Dyke in the journalScience published a paper in which they analysed therachises of the primary feathers ofConfuciusornis andArchaeopteryx. 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 thatArchaeopteryx andConfuciusornis, were unable to use flapping flight.[103] This study was criticized byPhilip J. Currie and Luis Chiappe. Chiappe suggested that it is difficult to measure the rachises of fossilized feathers, and Currie speculated thatArchaeopteryx andConfuciusornis 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.[104]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 specimenArchaeopteryx, 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.[105] Despite these criticisms, Nudds and Dyke stood by their original conclusions. They claimed that Paul's statement, that an adultArchaeopteryx 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.[106] 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 thebasilisk lizard, which could explain their presence in lake and marine deposits (seeOrigin of avian flight).[107][108]
Replica of the London Specimen
In 2004, scientists analysing a detailedCT scan of thebraincase of the LondonArchaeopteryx 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.[109] 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 thatArchaeopteryx had the keen sense of hearing, balance, spatial perception, and coordination needed to fly.[110]Archaeopteryx had a cerebrum-to-brain-volume ratio 78% of the way to modern birds from the condition of non-coelurosaurian dinosaurs such asCarcharodontosaurus orAllosaurus, which had a crocodile-like anatomy of the brain and inner ear.[111] Newer research shows that while theArchaeopteryx brain was more complex than that of more primitive theropods, it had a more generalized brain volume amongManiraptora 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 maniraptoran clade.[112]
Recent studies of flight feather barb geometry reveal that modern birds possess a larger barb angle in the trailing vane of the feather, whereasArchaeopteryx lacks this large barb angle, indicating potentially weak flight abilities.[113]
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 byO. C. Marsh). Other scientists seeArchaeopteryx as running quickly along the ground, supporting the idea that birds evolved flight by running (the "ground up" hypothesis proposed bySamuel Wendell Williston). Still others suggest thatArchaeopteryx 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 thatArchaeopteryx used its wings mainly to escapepredators 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.[23]
In March 2018, scientists reported thatArchaeopteryx was likely capable of a flight stroke cycle morphologically closer to the grabbing motion ofmaniraptorans and distinct from that ofmodern birds.[114][115] This study onArchaeopteryx's bone histology identified biomechanical and physiological adaptations exhibited by modern volant birds that perform intermittent flapping, such aspheasants and other burst flyers.
Some researchers suggested that the feather sheaths ofArchaeopteryx shows a center-out, flight relatedmoulting strategy like modern birds. As it was a weak flier, this would have been extremely advantageous in preserving its maximum flight performance.[116] Kiat and colleagues reinterpreted this purported moulting evidence to be problematic and equivocal at best, and considered that these structures more likely represents the calami traces of the fully grown feathers,[117] though the original authors still remained by their conclusion.[118]
Growth trends compared with other dinosaurs and birds
Anhistological study by Erickson, Norell, Zhongue, and others in 2009 estimated thatArchaeopteryx grew relatively slowly compared to modern birds, presumably because the outermost portions ofArchaeopteryx bones appear poorly vascularized;[5] in living vertebrates, poorly vascularized bone is correlated with slow growth rate. They also assume that all known skeletons ofArchaeopteryx come from juvenile specimens. Because the bones ofArchaeopteryx 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 inArchaeopteryx. The poorly vascularized bone ofArchaeopteryx might have grown as slowly as that in a mallard (2.5micrometres per day) or as fast as that in an ostrich (4.2micrometres per day). Using this range of bone growth rates, they calculated how long it would take to "grow" each specimen ofArchaeopteryx 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 avialansJeholornis andSapeornis grew relatively slowly, as did the dromaeosauridMahakala. The avialansConfuciusornis andIchthyornis grew relatively quickly, following a growth trend similar to that of modern birds.[119] One of the few modern birds that exhibit slow growth is the flightlesskiwi, and the authors speculated thatArchaeopteryx and the kiwi had similarbasal metabolic rate.[5]
Comparisons between thescleral rings ofArchaeopteryx and modern birds and reptiles indicate that it may have beendiurnal, similar to most modern birds.[120]
Restoration ofArchaeopteryx chasing a juvenileCompsognathus
The richness and diversity of theSolnhofen limestones in which all specimens ofArchaeopteryx have been found have shed light on an ancient Jurassic Bavaria strikingly different from the present day. The latitude was similar toFlorida, 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, includecycads and conifers while animals found include a large number of insects, small lizards,pterosaurs, andCompsognathus.[16]
The excellent preservation ofArchaeopteryx fossils and other terrestrial fossils found atSolnhofen indicates that they did not travel far before becoming preserved.[121] TheArchaeopteryx specimens found were therefore likely to have lived on the low islands surrounding the 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.[122] The pterosaurs included species such asRhamphorhynchus belonging to theRhamphorhynchidae, the group which dominated theecological niche currently occupied byseabirds, and which became extinct at the end of the Jurassic. The pterosaurs, which also includedPterodactylus, were common enough that it is unlikely that the specimens found arevagrants from the larger islands 50 km (31 mi) to the north.[123]
The islands that surrounded the Solnhofen lagoon were low lying,semi-arid, and sub-tropical with a longdry season and little rain.[124] The closest modern analogue for the Solnhofen conditions is said to beOrca Basin in the northernGulf of Mexico, although it is much deeper than the Solnhofen lagoons.[122] Theflora of these islands was adapted to these dry conditions and consisted mostly of low (3 m [10 ft]) shrubs.[123] Contrary to reconstructions ofArchaeopteryx climbing large trees, these seem to have been mostly absent from the islands; few trunks have been found in the sediments and fossilized treepollen also is absent.
The lifestyle ofArchaeopteryx is difficult to reconstruct and there are several theories regarding it. Some researchers suggest that it was primarily adapted to life on the ground,[125] while other researchers suggest that it was principally arboreal on the basis of the curvature of the claws[126] which has since been questioned.[127] The absence of trees does not precludeArchaeopteryx from an arboreal lifestyle, as several species of bird live exclusively in low shrubs. Various aspects of the morphology ofArchaeopteryx 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 ageneralist capable of feeding in both shrubs and open ground, as well as along the shores of the lagoon.[123] It most likely hunted small prey, seizing it with its jaws if it was small enough, or with its claws if it was larger.[citation needed]
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^British Museum of Natural History – 'BMNH 37001' – thetype specimen
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