Array ( [0] => {{Short description|Extinct genus of bird-like dinosaurs}} [1] => {{About| the ancient animal| the ancient plant | Archaeopteris |other uses| Archaeopteryx (disambiguation)}} [2] => {{pp-move}} [3] => {{Featured article}} [4] => {{Use dmy dates|date=June 2023}} [5] => {{Use British English|date=March 2016}} [6] => {{Use Oxford spelling|date = August 2022}} [7] => {{Automatic taxobox [8] => | fossil_range = [[Late Jurassic]] ([[Tithonian]]), {{fossil range|150.8|148.5}} [9] => | image = Archaeopteryx lithographica (Berlin specimen).jpg [10] => | image_caption = The Berlin ''Archaeopteryx'' specimen (''A. siemensii'') [11] => | image_alt = Fossil of complete Archaeopteryx, including indentations of feathers on wings and tail [12] => | taxon = Archaeopteryx [13] => | authority = [[Christian Erich Hermann von Meyer|Meyer]], 1861
([[conserved name]]) [14] => | type_species = {{extinct}}'''''Archaeopteryx lithographica''''' [15] => | type_species_authority = Meyer, 1861 (conserved name) [16] => | subdivision_ranks = Referred [[species]] [17] => | subdivision = *{{extinct}}'''''A. siemensii'''''
Dames, 1897 [18] => *{{extinct}}'''''A. albersdoerferi'''''
Kundrat ''et al.'', 2018 [19] => | display_parents = 2 [20] => | synonyms = {{collapsible list|bullets = false [21] => |title=Genus synonymy [22] => |''Griphosaurus''
Wagner, 1862 ([[rejected name]]) [23] => |''Griphornis''
Woodward, 1862 (rejected name) [24] => |''Archaeornis''
Petronievics, 1917 [25] => |''Jurapteryx''
Howgate, 1984 [26] => |''[[Wellnhoferia]]''?
Elżanowski, 2001 [27] => }} [28] => {{collapsible list|bullets = true [29] => |title=Species synonymy [30] => |''Griphosaurus problematicus''
Wagner, 1862 vide Woodward 1862 nomen rejectum [31] => |''Griphornis longicaudatus''
Owen 1862 vide Woodward 1862 nomen rejectum [32] => |''Griphosaurus longicaudatus''
(Owen 1862 vide Woodward 1862) Owen 1862 vide Brodkorb 1863 nomen rejectum [33] => |''Archaeopteryx macrura''
Owen, 1862 (rejected name) [34] => |''Archaeopteryx siemensii''?
(Dames, 1897) [35] => |''Archaeornis siemensii''
(Dames, 1897) Peteronievics vide Petroneivics & Woodward 1917 [36] => |''Archaeopteryx owenii''
Petronievics, 1917 (rejected name) [37] => |''Archaeopteryx recurva''
Howgate, 1984 [38] => |''Jurapteryx recurva''
(Howgate, 1984) Howgate 1985 [39] => |''Archaeopteryx bavarica''
Wellnhofer, 1993 [40] => |?''[[Wellnhoferia grandis]]''
Elżanowski, 2001 [41] => }} [42] => | synonyms_ref = {{Cite web |title=Troodontidae Gilmore, 1924 |url=http://theropoddatabase.com/Troodontidae.htm#Troodontidae |archive-url=https://web.archive.org/web/20190403063849/http://theropoddatabase.com/Troodontidae.htm#Troodontidae |archive-date=3 April 2019 |website=theropoddatabase.com}} [43] => }} [44] => [45] => '''''Archaeopteryx''''' ({{IPAc-en|ˌ|ɑːr|k|iː|ˈ|ɒ|p|t|ər|ᵻ|k|s}}; {{lit|old-wing}}), sometimes referred to by its German name, "'''{{lang|de|Urvogel}}'''" ({{lit}} ''Primeval Bird'') is a [[genus]] of [[bird]]-like [[dinosaur]]s. The name derives from the [[ancient Greek]] {{lang|grc|{{linktext|ἀρχαῖος}}}} (''archaīos''), meaning "ancient", and {{lang|grc|{{linktext|πτέρυξ}}}} (''ptéryx''), meaning "feather" or "wing". Between the late 19th century and the early 21st century, ''Archaeopteryx'' was generally accepted by [[paleontology|palaeontologists]] and popular reference books as the oldest-known bird (member of the group [[Avialae]]).{{Cite journal |last1=Xu |first1=X |last2=You |first2=H |last3=Du |first3=K |last4=Han |first4=F |date=28 July 2011 |title=An ''Archaeopteryx''-like theropod from China and the origin of Avialae |url=http://www.ivpp.ac.cn/qt/papers/201403/P020140314389417822583.pdf |url-status=dead |journal=Nature |volume=475 |issue=7357 |pages=465–470 |doi=10.1038/nature10288 |pmid=21796204 |archive-url=https://web.archive.org/web/20161220071919/http://www.ivpp.ac.cn/qt/papers/201403/P020140314389417822583.pdf |archive-date=20 December 2016 |access-date=5 November 2016 |s2cid=205225790}} Older potential [[avialan]]s have since been identified, including ''[[Anchiornis]]'', ''[[Xiaotingia]]'', and ''[[Aurornis]]''.{{Cite journal |last1=Godefroit |first1=Pascal |last2=Cau |first2=Andrea |last3=Hu |first3=Dong-Yu |last4=Escuillié |first4=François |last5=Wu |first5=Wenhao |last6=Dyke |first6=Gareth |year=2013 |title=A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds |journal=Nature |volume=498 |issue=7454 |pages=359–362 |bibcode=2013Natur.498..359G |doi=10.1038/nature12168 |pmid=23719374 |s2cid=4364892}} [46] => [47] => ''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 {{cvt|0.5|m}} 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 [[dromaeosauridae|dromaeosaurids]] and [[troodontidae|troodontids]]: jaws with sharp [[Tooth|teeth]], three fingers with [[claw]]s, a long bony tail, hyperextensible second toes ("killing claw"), feathers (which also suggest [[homeothermy|warm-bloodedness]]), and various features of the [[skeleton]]. [48] => [49] => 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, the identity of which has been controversial. 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. [50] => [51] => ''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.{{Cite journal|last=Kaplan |first=Matt |date=27 July 2011 |title=Archaeopteryx no longer first bird |url=http://www.nature.com/articles/news.2011.443 |journal=Nature |language=en |pages=news.2011.443 |doi=10.1038/news.2011.443 |issn=0028-0836}} Most of these eleven fossils include impressions of feathers. Because these feathers are of an advanced form ([[flight feather]]s), 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 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 of [[evolution]]. [52] => [53] => == History of discovery == [54] => {{Main|Specimens of Archaeopteryx}} [55] => [[File:Archaeopteryx (Feather).jpg|upright|thumb|left|The single feather]] [56] => 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 {{lang|de|italics=unset|[[Solnhofen]]}}, Germany. These quarries excavate sediments from the [[Solnhofen Limestone]] formation and related units. [57] => [[File:Archaeopteryx timeline.svg|thumb|Timeline of ''Archaeopteryx'' discoveries until 2007{{image reference needed|date=December 2022}}]] [58] => The initial discovery, a single feather, was unearthed in 1860 or 1861 and described in 1861 by {{lang|de|italics=unset|[[Christian Erich Hermann von Meyer|Hermann von Meyer]]}}.{{Cite journal |last=Meyer |first=Hermann von |date=15 August 1861 |title=Vogel-Federn und Palpipes priscus von Solenhofen |trans-title=Bird feathers and ''Palpipes priscus'' [a crustacean] from Solenhofen |url=https://books.google.com/books?id=6RAFAAAAQAAJ&pg=PA561 |journal=Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefaktenkunde |language=de |page=561}} ''"Aus dem lithographischen Schiefer der Brüche von ''Solenhofen'' in ''Bayern'' ist mir in den beiden Gegenplatten eine auf der Ablösungs- oder Spaltungs-Fläche des Gesteins liegende Versteinerung mitgetheilt worden, die mit grosser Deutlichkeit eine Feder erkennen lässt, welche von den Vogel-Federn nicht zu unterscheiden ist."'' (From the lithographic slates of the faults of Solenhofen in Bavaria, there has been reported to me a fossil lying on the stone's surface of detachment or cleavage, in both opposing slabs, which can be recognized with great clarity [to be] a feather, which is indistinguishable from a bird's feather.) It is currently located at the [[Natural History Museum, Berlin|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. 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.{{Cite journal |last1=Kaye |first1=T.G. |last2=Pittman |first2=M. |last3=Mayr |first3=G. |last4=Schwarz |first4=D. |last5=Xu |first5=X. |year=2019 |title=Detection of lost calamus challenges identity of isolated ''Archaeopteryx'' feather |journal=Scientific Reports |volume=9 |issue=1 |page=1182 |bibcode=2019NatSR...9.1182K |doi=10.1038/s41598-018-37343-7 |pmc=6362147 |pmid=30718905}} 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]]. [59] => [60] => The first skeleton, known as the '''London Specimen''' (BMNH 37001), was unearthed in 1861 near {{lang|de|italics=unset|[[Langenaltheim]]}}, Germany, and perhaps given to local physician {{lang|de|italics=unset|Karl Häberlein}} in return for medical services. He then sold it for £700 (roughly £83,000 in 2020[https://www.in2013dollars.com/uk/inflation/1861?amount=700 UK inflation calculator]) to the [[Natural History Museum, London|Natural History Museum]] in London, where it remains. 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'', 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 [[Oolite|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." [61] => [62] => The [[Ancient Greek|Greek]] word {{lang|grc-Latn|archaīos}} ({{wikt-lang|grc|ἀρχαῖος}}) means 'ancient, primeval'. {{lang|grc-Latn|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 [[Flight feather|remex]] (wing feather), but he had heard of and been shown a rough sketch of the [[London]] specimen, to which he referred as a "{{lang|de|Skelett eines mit ähnlichen Federn bedeckten Tieres}}" ("skeleton of an animal covered in similar feathers"). In German, this ambiguity is resolved by the term {{lang|de|Schwinge}} which does not necessarily mean a wing used for flying. {{lang|de|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|date=March 2022}} [63] => [64] => Since then, twelve specimens have been recovered: [65] => [66] => The '''Berlin Specimen''' (HMN 1880/81) was discovered in 1874 or 1875 on the Blumenberg near {{lang|de|italics=unset|[[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 [[German gold mark|Goldmark]] by the Berlin's Natural History Museum, where it now is displayed. The transaction was financed by [[Ernst Werner von Siemens]], founder of [[Siemens|the 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. [67] => [[File:Archaeopteryx lithographica (Cast of Maxberg Specimen).jpg|upright|thumb|Cast of the Maxberg Specimen]] [68] => Composed of a torso, the '''[[Maxberg specimen|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 sold.{{Cite web |title=Lost treasures: The Maxberg Archaeopteryx |url=https://www.newscientist.com/article/mg21328501-800-lost-treasures-the-maxberg-archaeopteryx |last=Hecht |first=Jeff |date=1 February 2012 |access-date=2 June 2022 |website=New Scientist |language=en-US }} [69] => [70] => The '''Haarlem Specimen''' (TM 6428/29, also known as the ''Teylers Specimen'') was discovered in 1855 near {{lang|de|italics=unset|[[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.{{Cite journal |last1=Foth |first1=C. |last2=Rauhut |first2=O.W.M. |date=2017 |title=Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs |journal=BMC Evolutionary Biology |volume=17 |issue=1 |page=236 |doi=10.1186/s12862-017-1076-y |pmc=5712154 |pmid=29197327 |bibcode=2017BMCEE..17..236F |doi-access=free}} [71] => [[File:Archaeopteryx lithographica (Eichstätter Specimen).jpg|thumb|left|Eichstätt Specimen, once considered a distinct genus, ''Jurapteryx'']] [72] => 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'').{{Cite journal |last=Howgate |first=M. E. |date=September 1984 |title=The teeth of Archaeopteryx and a reinterpretation of the Eichstätt specimen |url= https://academic.oup.com/zoolinnean/article-lookup/doi/10.1111/j.1096-3642.1984.tb00540.x |journal=Zoological Journal of the Linnean Society |language=en |volume=82 |issue=1–2 |pages=159–175 |doi=10.1111/j.1096-3642.1984.tb00540.x}} [73] => [74] => 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.Elżanowski A 2001b. "[https://www.researchgate.net/publication/228707898_A_new_genus_and_species_for_the_largest_specimen_of_Archaeopteryx A new genus and species for the largest specimen of Archaeopteryx]". Acta Palaeontologica Polonica 46: 519–532. [75] => [76] => 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 [[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'',Wellnhofer, Peter. ''Archaeopteryx. Der Urvogel Von Solnhofen''. München: Friedrich Pfeil, 2008, p. 54. but more recent studies suggest it belongs to ''A. siemensii''. [77] => [78] => [[File:Archaeopteryx (Daiting Specimen).jpg|thumb|Daiting Specimen, the [[holotype]] of ''A. albersdoerferi'']] [79] => An eighth, fragmentary specimen was discovered in 1990 in the younger [[Mörnsheim Formation]] 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 ''Archaeopteryx'' at the Munich Mineral Show in October 2009. The Daiting Specimen was subsequently named ''Archaeopteryx albersdoerferi'' by Kundrat et al. (2018).{{Cite journal |last1=Kundrát |first1=Martin |last2=Nudds |first2=John |last3=Kear |first3=Benjamin P. |last4=Lü |first4=Junchang |last5=Ahlberg |first5=Per |year=2019 |title=The first specimen of ''Archaeopteryx'' from the Upper Jurassic Mörnsheim Formation of Germany |journal=Historical Biology |volume=31 |issue=1 |pages=3–63 |doi=10.1080/08912963.2018.1518443 |bibcode=2019HBio...31....3K |s2cid=91497638}} After a lengthy period in a closed private collection, it was moved to the Museum of Evolution at [[Knuthenborg Safaripark]] (Denmark) in 2022, where it has since been on display and also been made available for researchers.{{cite news | last=Frandsen | first=P.N. | date=16 November 2022 | title=Verdens vigtigste fossil kommer til Lolland | url=https://folketidende.dk/lolland/verdens-vigtigste-fossil-kommer-til-lolland | publisher=Folketidende | access-date=1 July 2023 | language=da }}{{cite news | date=15 November 2022 | title=Knuthenborg Safaripark åbner sin største satsning nogensinde | url=https://via.ritzau.dk/pressemeddelelse/knuthenborg-safaripark-abner-sin-storste-satsning-nogensinde?publisherId=13560971&releaseId=13664363 | publisher=Ritzau | access-date=1 July 2023 | language=da }} [80] => [[File:Archaeopteryx (Chicken Wing).jpg|thumb|left|Bürgermeister-Müller ("chicken wing") Specimen]] [81] => 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'', it is colloquially known as "chicken wing".{{Cite journal |last1=Voeten |first1=Dennis F. A. E. |last2=Cubo |first2=Jorge |last3=de Margerie |first3=Emmanuel |last4=Röper |first4=Martin |last5=Beyrand |first5=Vincent |last6=Bureš |first6=Stanislav |last7=Tafforeau |first7=Paul |last8=Sanchez |first8=Sophie |date=2018 |title=Wing bone geometry reveals active flight in Archaeopteryx |url=https://doi.org/10.1038/s41467-018-03296-8 |journal=Nature Communications |volume=9 |issue=923 |page=6 |bibcode=2018NatCo...9..923V |doi=10.1038/s41467-018-03296-8 |pmc=5849612 |pmid=29535376 |access-date=17 June 2022}} [82] => [[File:Thermopolis skeleton.jpg|thumb|Details of the Wyoming Dinosaur Center Archaeopteryx (WDC-CSG-100)]] [83] => 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. This has been interpreted as evidence of [[Theropoda|theropod]] ancestry. In 1988, [[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 and best-preserved ''Archaeopteryx'' remains yet. [84] => [[File:Eleventh Archaeopteryx specimen.jpg|alt=|thumb|The eleventh specimen]] [85] => 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.{{Cite web |last=Switek |first=Brian |date=19 October 2011 |title=Paleontologists Unveil the 11th Archaeopteryx |url=http://blogs.smithsonianmag.com/dinosaur/2011/10/paleontologists-unveil-the-11th-archaeopteryx/ |website=Dinosaur Tracking Blog |publisher=Smithsonian Institution}}{{Cite journal |last=Hecht |first=Jeff |date=20 October 2011 |title=Another stunning Archaeopteryx fossil found in Germany |url=https://www.newscientist.com/blogs/shortsharpscience/2011/10/stunning-new-archaeopteryx-fos.html |journal=New Scientist}} 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 tip.{{Cite journal |last1=Foth |first1=C |last2=Tischlinger |first2=H |last3=Rauhut |first3=O. W |date=3 July 2014 |title=New specimen of ''Archaeopteryx'' provides insights into the evolution of pennaceous feathers |journal=Nature |volume=511 |issue=7507 |pages=79–82 |bibcode=2014Natur.511...79F |doi=10.1038/nature13467 |pmid=24990749 |s2cid=4464659}}{{Cite web |title=First show off, then take off |date=3 July 2014 |url=http://www.en.uni-muenchen.de/news/press-services/press-releases/2014/archaeopteryx.html |publisher=Ludwig-Maximilians-Universität |archive-url=https://web.archive.org/web/20140707074102/http://www.en.uni-muenchen.de/news/press-services/press-releases/2014/archaeopteryx.html |archive-date=7 July 2014}} [86] => [87] => 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.{{Cite web |title=Schamhaupten: Fossil des Archaeopteryx entdeckt |url=http://www.donaukurier.de/lokales/kurzmeldungen/riedenburg/Fossil-des-Archaeopteryx-entdeckt;art74375,2880005#plx982820499 |website=donaukurier.de |access-date=18 February 2014 |archive-date=19 October 2014 |archive-url=https://web.archive.org/web/20141019150803/http://www.donaukurier.de/lokales/kurzmeldungen/riedenburg/Fossil-des-Archaeopteryx-entdeckt;art74375,2880005#plx982820499 |url-status=dead }} 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.{{Cite journal |last1=Rauhut |first1=Oliver W.M. |last2=Foth |first2=Christian |last3=Tischlinger |first3=Helmut |year=2018 |title=The oldest ''Archaeopteryx'' (Theropoda: Avialiae): a new specimen from the Kimmeridgian/Tithonian boundary of Schamhaupten, Bavaria |journal=PeerJ |volume=6 |pages=e4191 |doi=10.7717/peerj.4191 |pmc=5788062 |pmid=29383285 |doi-access=free }} [88] => [89] => === Authenticity === [90] => 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. Their claims were repudiated by [[Alan J. Charig]] and others at the [[Natural History Museum, London|Natural History Museum in London]]. 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]], 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 feathers. [91] => [92] => 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 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. [93] => [94] => 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. [95] => [96] => 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 authentic. [97] => [98] => == Description == [99] => [[File:Archiesizeall1.svg|thumb|400px|alt=Archaeopteryx sizes ranging between about 25 and 50 cm long and between 25 and 60 cm in wingspan|Specimens compared to a [[human]] in scale]] [100] => Most of the specimens of ''Archaeopteryx'' that have been discovered come from the Solnhofen limestone in Bavaria, southern Germany, which is a {{lang|de|italics=unset|[[Lagerstätte]]}}, a rare and remarkable geological formation known for its superbly detailed fossils laid down during the early Tithonian stage of the Jurassic period, approximately 150.8–148.5{{nbsp}}million years ago.{{Cite journal |last=Schweigert |first=G. |year=2007 |title=Ammonite biostratigraphy as a tool for dating Upper Jurassic lithographic limestones from South Germany – first results and open questions |journal=Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen |volume=245 |issue=1 |pages=117–125 |doi=10.1127/0077-7749/2007/0245-0117|s2cid=140597349 |url=http://doc.rero.ch/record/13825/files/PAL_E722.pdf }} [101] => [102] => ''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 {{convert|0.5|m|ftin}} in body length and {{convert|0.7|m|ftin}} in [[wingspan]], with an estimated mass of {{convert|0.5|to|1|kg}}.{{Cite book |last=Paul |first=Gregory S. |url=http://worldcat.org/oclc/985402380 |title=The Princeton Field Guide to Dinosaurs |publisher=Princeton University Press |year=2016 |isbn=978-1-78684-190-2 |pages=144 |oclc=985402380}} ''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 [[Theropoda|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. [103] => [104] => Because it displays features common to both birds and non-avian dinosaurs, ''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 plate]]s, an [[obturator foramen|obturator]] process of the [[ischium]], and long chevrons in the tail. In particular, Ostrom found that ''Archaeopteryx'' was remarkably similar to the theropod family [[Dromaeosauridae]]. [105] => [106] => Archaeopteryx had three separate digits on each fore-leg each ending with a "claw". Few birds have such features. Some birds, such as [[duck]]s, [[swan]]s, [[Jacana (genus)|Jacanas]] (''Jacana'' sp.), and the [[hoatzin]] (''Opisthocomus hoazin''), have them concealed beneath their leg-feathers.{{cite book |last=Attenborough |first=David |title=The Life of Birds |publisher=BBC Books |year=1998 |isbn=0563387920 |author-link=David Attenborough}} [107] => [108] => === Plumage === [109] => [[File:Archaeopteryx-modern bird tails.png|thumb|left|Anatomical illustration comparing the "frond-tail" of ''Archaeopteryx'' with the "fan-tail" of a modern bird]] [110] => Specimens of ''Archaeopteryx'' were most notable for their well-developed [[flight feather]]s. They were markedly asymmetrical and showed the structure of flight feathers in modern birds, with vanes given stability by a [[barb (feather)|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 [[alula|stiff feathers]]. [111] => [112] => The body plumage of ''Archaeopteryx'' is less well-documented and has only been properly researched in the well-preserved [[The Berlin specimen|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 [[ratite]]s). In part they are firm and thus capable of supporting flight. [113] => [114] => A patch of [[pennaceous feather]]s 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 feather|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. [115] => [116] => 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 [[Oxygen saturation|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.{{Cite journal |last1=Reisdorf |first1=A. G. |last2=Wuttke |first2=M. |year=2012 |title=Re-evaluating Moodie's Opisthotonic-Posture Hypothesis in fossil vertebrates. Part I: Reptiles – The taphonomy of the bipedal dinosaurs ''Compsognathus longipes'' and ''Juravenator starki'' from the Solnhofen Archipelago (Jurassic, Germany) |journal=Palaeobiodiversity and Palaeoenvironments |volume=92 |issue=1 |pages=119–168 |doi=10.1007/s12549-011-0068-y |bibcode=2012PdPe...92..119R |s2cid=129785393}} 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. [117] => [118] => ==== Colouration ==== [119] => [[File:Archaeopteryx NT.jpg|thumb|right|Artist's restoration illustrating one interpretation of Carney's study{{Cite journal |last1=Carney |first1=R. |last2=Vinther |first2=Jakob |last3=Shawkey |first3=Matthew D. |last4=d'Alba |first4=Liliana |last5=Ackermann |first5=Jörg |year=2012 |title=New evidence on the colour and nature of the isolated Archaeopteryx feather |journal=Nature Communications |volume=3 |page=637 |bibcode=2012NatCo...3..637C |doi=10.1038/ncomms1642 |pmid=22273675 |doi-access=free}}]] [120] => [121] => In 2011, graduate student Ryan Carney and colleagues performed the first colour study on an ''Archaeopteryx'' specimen. Using [[Scanning electron microscope|scanning electron microscopy]] technology and [[Energy-dispersive X-ray spectroscopy|energy-dispersive X-ray]] analysis, the team was able to detect the structure of [[melanosome]]s in the isolated feather specimen described in 1861. The resultant measurements were then compared to those of 87{{nbsp}}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 (feather)|covert]], which would have partly covered the primary feathers on the wings. The study does not mean that ''Archaeopteryx'' 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.{{Cite news |last=Switek |first=Brian |date=9 November 2011 |title=Archaeopteryx was robed in black |work=New Scientist |location=Las Vegas |url=https://www.newscientist.com/article/dn21146-archaeopteryx-was-robed-in-black.html}} 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 complex light- and dark-coloured plumage, with heavier pigmentation in the distal tips and outer vanes.{{Cite journal |last1=Manning, Phillip. L. |last2=Edwards |first2=Nicholas P. |last3=Wogelius |first3=Roy A. |last4=Bergmann |first4=Uwe |last5=Barden |first5=Holly E. |last6=Larson |first6=Peter L. |last7=Schwarz-Wings |first7=Daniela |last8=Egerton |first8=Victoria M. |last9=Sokaras |first9=Dimosthenis |display-authors=etal |year=2013 |title=Synchrotron-based chemical imaging reveals plumage patterns in a 150 million year old early bird |journal=Journal of Analytical Atomic Spectrometry |volume=28 |issue=7 |page=1024 |doi=10.1039/c3ja50077b |doi-access=free}} This analysis of colour distribution was based primarily on the distribution of sulphate within the fossil. An author on the previous ''Archaeopteryx'' colour study argued against the interpretation of such biomarkers as an indicator of eumelanin in the full ''Archaeopteryx'' specimen.{{Cite journal |last=Vinther |first=J |year=2015 |title=A guide to the field of palaeo colour: melanin and other pigments can fossilise: reconstructing colour patterns from ancient organisms can give new insights to ecology and behaviour |journal=BioEssays |volume=6 |issue=37 |pages=643–656 |doi=10.1002/bies.201500018 |pmid=25854512 |s2cid=24966334}} Carney and other colleagues also argued against the 2013 study's interpretation of the sulphate and trace metals,{{Cite journal |last1=Carney |first1=Ryan |last2=Molnar |first2=Julia |last3=Updike |first3=Earl |last4=Brown |first4=William |last5=Jackson |first5=Jessie |last6=Shawkey |first6=Matthew |last7=Lindgren |first7=Johan |last8=Sjövall |first8=Peter |last9=Falkingham |first9=Peter |last10=Gauthier |first10=Jacques |date=2014 |title=Archaeopteryx in 4D |url=http://vertpaleo.org/GlobalPDFS/SVP-2014-Program-and-Abstract-Book-9-18-2014.aspx |journal=Journal of Vertebrate Paleontology |language=en |volume=103 |access-date=15 January 2018 |archive-date=25 April 2018 |archive-url=https://web.archive.org/web/20180425135859/http://vertpaleo.org/GlobalPDFS/SVP-2014-Program-and-Abstract-Book-9-18-2014.aspx |url-status=dead }}{{Cite web |title=SVP – Press Release – Taking a Deeper Look at "Ancient Wing" |url=http://vertpaleo.org/Society-News/SVP-Paleo-News/Society-News,-Press-Releases/Press-Release-Taking-a-Deeper-Look-at-Ancient-Wing.aspx |access-date=3 April 2016 |website=vertpaleo.org |archive-date=19 April 2016 |archive-url=https://web.archive.org/web/20160419054434/http://vertpaleo.org/Society-News/SVP-Paleo-News/Society-News,-Press-Releases/Press-Release-Taking-a-Deeper-Look-at-Ancient-Wing.aspx |url-status=dead }} and in a 2020 study published in ''Scientific 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 of ''Archaeopteryx'' remain unknown". [122] => [123] => == Classification == [124] => [[File:Thermopolis Archaeopteryx.jpg|thumb|left|The Thermopolis Specimen]] [125] => 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 [[Christian Erich Hermann von Meyer|Meyer]]. In 1954 [[Gavin de Beer]] concluded that the London specimen was the [[holotype]]. In 1960, Swinton accordingly proposed that the name ''Archaeopteryx lithographica'' 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]] (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 species. [126] => [127] => 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 dinosaur|feathered theropod]], of which only this feather is known so far. As the feather had been designated the [[Biological type|type specimen]], the name ''Archaeopteryx'' should then no longer be applied to the skeletons, thus creating significant [[International Code of Zoological Nomenclature|nomenclatorial]] 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, or [[neotype]].{{Cite journal |last1=Bock |first1=W. J. |last2=Bühler |first2=P. |year=2007 |title=''Archaeopteryx lithographica'' von Meyer, 1861 (Aves): proposed conservation of usage by designation of a neotype |url=http://iczn.org/content/archaeopteryx-lithographica-von-meyer-1861-aves-proposed-conservation-usage-designation-neot |journal=Bulletin of Zoological Nomenclature |volume=64 |issue=4 |pages=261–262}} This suggestion was upheld by the ICZN after four years of debate, and the London specimen was designated the neotype on 3 October 2011.{{Cite journal |last=ICZN |year=2011 |title=OPINION 2283 (Case 3390) Archaeopteryx lithographica von Meyer, 1861 (Aves): conservation of usage by designation of a neotype |url=http://iczn.org/content/opinion-2283-case-3390-archaeopteryx-lithographica-von-meyer-1861-aves-conservation-usage-de |journal=Bulletin of Zoological Nomenclature |volume=68 |issue=3 |pages=230–233 |doi=10.21805/bzn.v68i3.a16 |s2cid=160784119}} [128] => [[File:Twelfth Archaeopteryx specimen.jpg|thumb|The twelfth specimen]] [129] => Below is a [[cladogram]] published in 2013 by [[Pascal Godefroit|Godefroit]] ''et al.'' [130] => [131] => {{clade| style=font-size:100%; line-height:100% [132] => |label1=[[Avialae]] [133] => |1={{clade [134] => |1=''[[Aurornis]]'' [[File:Aurornis.jpg|50 px]] [135] => |2={{clade [136] => |1=''[[Anchiornis]]'' [[File:Anchiornis_martyniuk.png|50 px]] [137] => |2={{clade [138] => |1='''''Archaeopteryx''''' [[File:Archaeopteryx.png|50 px]] [139] => |2={{clade [140] => |1=''[[Xiaotingia]]'' [[File:Xiaotingia .jpg|50 px]] [141] => |2={{clade [142] => |1={{clade [143] => |1=''[[Jeholornis]]'' [[File:Jeholornis_mmartyniuk_wiki.jpg|50 px]] [144] => |2=''[[Rahonavis]]'' [[File:Rahonavis NT.jpg|50 px]]}} [145] => |2={{clade [146] => |1=''[[Balaur (dinosaur)|Balaur]]'' [[File:Balaur by Mark P. Witton.png|50 px]] [147] => |2={{clade [148] => |1=[[Avebrevicauda]] (includes modern birds) [[File:Confuciusornis sanctus mmartyniuk.png|50 px]]}} }} }} }} }} }} }} }} [149] => [150] => === Species === [151] => [[File:Archaeopteryx skeletals.jpg|thumb|upright=2|Skeletal restorations of various specimens]] [152] => 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 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''. [153] => [154] => 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 [[Tubercle (bone)|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. [155] => [156] => === Synonyms === [157] => [[File:Eichstätt Wellnhoferia grandis.jpg|thumb|upright|The Solnhofen Specimen, by some considered as belonging to the genus ''[[Wellnhoferia]]'']] [158] => 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. [159] => * '''''Archaeopteryx lithographica''''' Meyer, 1861 [conserved name] [160] => **''Archaeopterix lithographica'' Anon., 1861 [''lapsus''] [161] => ** ''Griphosaurus problematicus'' Wagner, 1862 [rejected name 1961 per ICZN Opinion 607] [162] => ** ''Griphornis longicaudatus'' Owen ''vide'' Woodward, 1862 [rejected name 1961 per ICZN Opinion 607] [163] => ** ''Archaeopteryx macrura'' Owen, 1862 [rejected name 1961 per ICZN Opinion 607] [164] => ** ''Archaeopteryx oweni'' Petronievics, 1917 [rejected name 1961 per ICZN Opinion 607] [165] => ** ''Archaeopteryx recurva'' Howgate, 1984 [166] => ** ''Jurapteryx recurva'' (Howgate, 1984) Howgate, 1985 [167] => ** ''Wellnhoferia grandis'' Elżanowski, 2001 [168] => * '''''Archaeopteryx siemensii''''' Dames, 1897 [169] => **''Archaeornis siemensii'' (Dames, 1897) Petronievics, 1917 [170] => ** ''Archaeopteryx bavarica'' Wellnhofer, 1993 [171] => [172] => ''"Archaeopteryx" vicensensis'' (Anon. ''fide'' Lambrecht, 1933) is a ''[[nomen nudum]]'' for what appears to be an undescribed pterosaur. [173] => [174] => === Phylogenetic position === [175] => [[File:Archaeo-deinony hands.svg|thumb|upright|alt=Outline of bones in forelimbs of Deinonychus and Archaeopteryx; both have two fingers and an opposed claw with very similar layout, although Archaeopteryx has thinner bones|Comparison of the forelimb of ''Archaeopteryx'' (right) with that of ''[[Deinonychus]]'' (left)]] [176] => [177] => 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. 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 [[maniraptora]]n lineages are extremely birdlike, and they suggested that different groups of birds may have descended from different dinosaur ancestors. [178] => [179] => 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.{{Cite journal |last1=Lee |first1=M. S. |last2=Worthy |first2=T. H. |year=2012 |title=Likelihood reinstates ''Archaeopteryx'' as a primitive bird |journal=Biology Letters |volume=8 |issue=2 |pages=299–303 |doi=10.1098/rsbl.2011.0884 |pmc=3297401 |pmid=22031726}} [180] => [181] => 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.{{Cite journal |last1=Senter |first1=Phil |last2=Kirkland, James I. |last3=DeBlieux, Donald D. |last4=Madsen, Scott |last5=Toth, Natalie |year=2012 |editor-last=Dodson |editor-first=Peter |title=New Dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the Evolution of the Dromaeosaurid Tail |journal=PLOS ONE |volume=7 |issue=5 |pages=e36790 |bibcode=2012PLoSO...736790S |doi=10.1371/journal.pone.0036790 |pmc=3352940 |pmid=22615813 |doi-access=free}}{{Cite journal |last1=Turner |first1=Alan H. |last2=Makovicky |first2=Peter J. |last3=Norell |first3=Mark A. |year=2012 |title=A review of dromaeosaurid systematics and paravian phylogeny |url=http://digitallibrary.amnh.org/bitstream/2246/6352/5/B371-cover.pdf |archive-url= https://ghostarchive.org/archive/20221009/http://digitallibrary.amnh.org/bitstream/2246/6352/5/B371-cover.pdf |archive-date=9 October 2022 |url-status=live |journal=Bulletin of the American Museum of Natural History |volume=371 |pages=1–206 |doi=10.1206/748.1 |hdl=2246/6352 |s2cid=83572446}} 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 |Eosinopteryx brevipenna]]''. The authors used a modified version of the matrix from the study describing ''Xiaotingia'', adding ''[[Jinfengopteryx |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.{{Cite journal |last1=Godefroit |first1=P |last2=Demuynck |first2=H |last3=Dyke |first3=G |last4=Hu |first4=D |last5=Escuillié |first5=F |last6=Claeys |first6=P |year=2013 |title=Reduced plumage and flight ability of a new Jurassic paravian theropod from China |journal=Nature Communications |volume=4 |pages=Article number 1394 |bibcode=2013NatCo...4.1394G |doi=10.1038/ncomms2389 |pmid=23340434 |doi-access=free}} [182] => [183] => 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).{{Cite book |last1=Agnolin |first1=Federico |title=Avian ancestors. A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, ''Anchiornis'', and Scansoriopterygidae |last2=Novas |first2=Fernando E |year=2013 |isbn=978-94-007-5636-6 |series=SpringerBriefs in Earth System Sciences |pages=1–96 |doi=10.1007/978-94-007-5637-3 |s2cid=199493087}} 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 ''[[house sparrow|Passer domesticus]]'', but not ''[[Dromaeosaurus |Dromaeosaurus albertensis]]'' or ''[[Troodon |Troodon formosus]]''). ''Archaeopteryx'' was found to form a [[Evolutionary grade|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. [184] => [185] => Hu ''et al''. (2018),{{Cite journal |last1=Hu |first1=Dongyu |last2=Clarke |first2=Julia A. |last3=Eliason |first3=Chad M. |last4=Qiu |first4=Rui |last5=Li |first5=Quanguo |last6=Shawkey |first6=Matthew D. |last7=Zhao |first7=Cuilin |last8=D’Alba |first8=Liliana |last9=Jiang |first9=Jinkai |last10=Xu |first10=Xing |date=15 January 2018 |title=A bony-crested Jurassic dinosaur with evidence of iridescent plumage highlights complexity in early paravian evolution |journal=Nature Communications |language=en |volume=9 |issue=1 |pages=217 |bibcode=2018NatCo...9..217H |doi=10.1038/s41467-017-02515-y |issn=2041-1723 |pmc=5768872 |pmid=29335537 |doi-access=free}} Wang ''et al''. (2018){{Cite journal |last1=Wang |first1=Min |last2=O’Connor |first2=Jingmai K. |last3=Xu |first3=Xing |last4=Zhou |first4=Zhonghe |date=May 2019 |title=A new Jurassic scansoriopterygid and the loss of membranous wings in theropod dinosaurs |url=http://www.nature.com/articles/s41586-019-1137-z |journal=Nature |language=en |volume=569 |issue=7755 |pages=256–259 |bibcode=2019Natur.569..256W |doi=10.1038/s41586-019-1137-z |issn=0028-0836 |pmid=31068719 |s2cid=148571099}} and Hartman ''et al''. (2019){{Cite journal|last1=Hartman |first1=Scott |last2=Mortimer |first2=Mickey |last3=Wahl |first3=William R. |last4=Lomax |first4=Dean R. |last5=Lippincott |first5=Jessica |last6=Lovelace |first6=David M. |date=10 July 2019 |title=A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight |journal=PeerJ |volume=7 |pages=e7247 |doi=10.7717/peerj.7247 |issn=2167-8359 |pmc=6626525 |pmid=31333906 |doi-access=free }} 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 "[[Anchiornithidae|anchiornithids]]"), they considered it highly probable that this animal [[Convergent evolution|evolved flight independently]] from bird ancestors (and from ''Microraptor'' and ''[[Yi (dinosaur)|Yi]]''). The following cladogram illustrates their hypothesis regarding the position of ''Archaeopteryx'': [186] => [187] => {{clade [188] => |1=[[Oviraptorosauria]] [189] => |label2=[[Paraves]] [190] => |2={{clade [191] => |1=[[Avialae]] [192] => |label2=[[Deinonychosauria]] [193] => |2={{clade [194] => |1={{clade [195] => |1=[[Unenlagiidae]] [196] => |2={{clade [197] => |1=[[Dromaeosauridae]] [198] => |2=[[Troodontidae]] }} }} [199] => |label2='''Archaeopterygidae''' (=[[Anchiornithidae]]) [200] => |2={{clade [201] => |1=''[[Serikornis]]'' [202] => |2={{clade [203] => |1=''[[Caihong]]'' [204] => |2={{clade [205] => |1=''[[Anchiornis]]'' [206] => |2={{clade [207] => |1='''''Archaeopteryx''''' [208] => |2={{clade [209] => |1=''[[Eosinopteryx]]'' [210] => |2=''[[Aurornis]]'' [211] => }} }} }} }} }} }} }} }} [212] => 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. [213] => [214] => == Palaeobiology == [215] => [216] => === Flight === [217] => [[File:Vog1h.jpg|thumb|upright|left|1880 photo of the Berlin Specimen, showing leg feathers that were removed subsequently, during preparation]] [218] => 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 [[Sternum|breastbone]] suggests that ''Archaeopteryx'' was not a very strong flier, but flight muscles might have attached to the thick, boomerang-shaped wishbone, the platelike [[coracoid]]s, or perhaps, to a [[cartilage|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. However, a more recent study solves this issue by suggesting a different flight stroke configuration for non-avian flying theropods.{{Cite journal |last=Voeten |first=Dennis F.A.E. |display-authors=etal |year=2018 |title=Wing bone geometry reveals active flight in ''Archaeopteryx'' |journal=Nature Communications |volume=9 |issue=1 |page=923 |bibcode=2018NatCo...9..923V |doi=10.1038/s41467-018-03296-8 |pmc=5849612 |pmid=29535376}} [219] => [220] => ''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 [[pheasant]]s). 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%. [221] => [222] => 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.{{Cite journal |last1=Speakman |first1=J. R. |last2=Thomson |first2=S. C. |year=1994 |title=Flight capabilities of ''Archaeopteryx'' |journal=Nature |volume=370 |issue=6490 |pages=336–340 |bibcode=1994Natur.370..514S |doi=10.1038/370514a0 |pmid=28568098 |s2cid=4248184|doi-access=free }} The degree of asymmetry seen in ''Archaeopteryx'' is more typical for slow flyers than for flightless birds.{{Cite journal |last=Norberg |first=R. A. |year=1995 |title=Feather asymmetry in ''Archaeopteryx'' |journal=Nature |volume=374 |issue=6519 |page=211 |bibcode=1995Natur.374..211M |doi=10.1038/374211a0 |doi-access=free |s2cid=4352260}} [223] => [224] => [[File:Archaeopteryx bavarica Detail.jpg|thumb|The Munich Specimen]] [225] => [226] => In 2010, Robert L. Nudds and Gareth J. Dyke in the journal ''Science'' published a paper in which they analysed the [[rachis]]es of the primary feathers of ''[[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 that ''Archaeopteryx'' and ''Confuciusornis'', were unable to use flapping flight.{{Cite journal |last1=Nudds |first1=Robert L. |last2=Dyke |first2=Gareth J. |date=14 May 2010 |title=Narrow Primary Feather Rachises in ''Confuciusornis'' and ''Archaeopteryx'' Suggest Poor Flight Ability |journal=Science |volume=328 |issue=5980 |pages=887–889 |bibcode=2010Sci...328..887N |doi=10.1126/science.1188895 |pmid=20466930 |s2cid=12340187}} This study was criticized by [[Philip J. Currie]] and Luis Chiappe. Chiappe suggested that it is difficult to measure the rachises of fossilized feathers, and Currie speculated that ''Archaeopteryx'' and ''[[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.Balter, M. (2010). [http://news.sciencemag.org/sciencenow/2010/05/did-first-feathers-prevent-early.html "Did First Feathers Prevent Early Flight?"] ''Science Now'', 13 May 2010. [[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 {{cvt|250|g}} 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 {{cvt|140|g}}. 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 {{cvt|0.75|mm|+2}} for the longest primary feather, which Paul could not confirm using photographs. Paul measured some of the inner primary feathers, finding rachises {{cvt|1.25–1.4|mm}} across.{{Cite journal |last=Paul |first=G. S. |date=15 October 2010 |title=Comment on 'Narrow Primary Feather Rachises in ''Confuciusornis'' and ''Archaeopteryx'' Suggest Poor Flight Ability.' |journal=Science |volume=330 |issue=6002 |page=320 |bibcode=2010Sci...330..320P |doi=10.1126/science.1192963 |pmid=20947747 |doi-access=free}} 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.{{Cite journal |last1=Dyke |first1=G. J. |last2=Nudds |first2=R. L. |date=15 October 2010 |title=Response to Comments on "Narrow Primary Feather Rachises in Confuciusornis and Archaeopteryx Suggest Poor Flight Ability" |url=http://www.sciencemag.org/cgi/reprint/330/6002/320-d.pdf |archive-url=https://ghostarchive.org/archive/20221009/http://www.sciencemag.org/cgi/reprint/330/6002/320-d.pdf |archive-date=9 October 2022 |url-status=live |journal=Science |volume=330 |issue=6002 |page=320 |bibcode=2010Sci...330..320N |doi=10.1126/science.1193474 |doi-access=free |s2cid=85044108}} 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 [[Origin of avian flight]]).Videler, JJ (2005) Avian Flight. Oxford University Press. {{ISBN|0-19-856603-4}} pages 98–117{{Cite web |last=Videler, John J |date=January 2005 |title=How Archaeopteryx could run over water |url=https://www.researchgate.net/publication/38957284 |website=ResearchGate}} [227] => [228] => [[File:Archaeopteryx lithographica, replica of London specimen, Staatliches Museum für Naturkunde Karlsruhe, Germany - 20100925.jpg|left|upright|thumb|Replica of the London Specimen]] [229] => [230] => In 2004, scientists analysing a detailed [[CT scan]] of the [[braincase]] of the London ''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 as ''[[Carcharodontosaurus]]'' or ''[[Allosaurus]]'', which had a crocodile-like anatomy of the brain and inner ear.Larsson, H. C. E. (2001). "Endocranial anatomy of ''Carcharodontosaurus saharicus'' (Theropoda: Allosauroidea) and its implications for theropod brain evolution". In: Tanke, D. H.; Carpenter, K.; Skrepnick, M. W. (eds.) ''Mesozoic Vertebrate Life''. Indiana University Press. pp. 19–33. Newer research shows that while the ''Archaeopteryx'' brain was more complex than that of more primitive theropods, it had a more generalized brain volume among [[maniraptoran|Maniraptora]] 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.{{Cite journal |last1=Balanoff |first1=Amy M. |last2=Bever |first2=Gabe S. |last3=Rowe |first3=Timothy B. |last4=Norell |first4=Mark A. |year=2013 |title=Evolutionary origins of the avian brain |journal=Nature |volume=501 |issue=7465 |pages=93–6 |bibcode=2013Natur.501...93B |doi=10.1038/nature12424 |pmid=23903660 |s2cid=4451895}} [231] => [232] => 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.{{Cite journal |last1=Feo |first1=Teresa J. |last2=Field |first2=Daniel J. |last3=Prum |first3=Richard O. |date=22 March 2015 |title=Barb geometry of asymmetrical feathers reveals a transitional morphology in the evolution of avian flight |journal=Proceedings of the Royal Society of London B: Biological Sciences |volume=282 |issue=1803 |page=20142864 |doi=10.1098/rspb.2014.2864 |issn=0962-8452 |pmc=4345455 |pmid=25673687}} [233] => [[File:Archaeopteryx Philadelphia.jpg|thumb|Reconstructed skeleton, [[Academy of Natural Sciences of Drexel University]]]] [234] => ''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 [[Predation|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. [235] => [236] => In March 2018, scientists reported that ''Archaeopteryx'' was likely capable of a flight stroke cycle morphologically closer to the grabbing motion of [[maniraptorans]] and distinct from that of [[Birds|modern birds]].{{Cite journal |last=Voeten, Dennis F.A.E. |display-authors=et al |date=13 March 2018 |title=Wing bone geometry reveals active flight in Archaeopteryx |journal=[[Nature Communications]] |volume=9 |pages=923 |bibcode=2018NatCo...9..923V |doi=10.1038/s41467-018-03296-8 |pmc=5849612 |pmid=29535376 |number=923}}{{Cite news |last=Guarino |first=Ben |date=13 March 2018 |title=This feathery dinosaur probably flew, but not like any bird you know |newspaper=[[The Washington Post]] |url=https://www.washingtonpost.com/news/speaking-of-science/wp/2018/03/13/this-feathery-dinosaur-probably-flew-but-not-like-any-bird-you-know/ |access-date=13 March 2018}} This study on ''Archaeopteryx''{{'}}s bone histology identified biomechanical and physiological adaptations exhibited by modern volant birds that perform intermittent flapping, such as [[pheasant]]s and other burst flyers. [237] => [238] => 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.Kaye, T.G. ''et al''. (2020) [https://www.nature.com/articles/s42003-020-01467-2#disqus_thread ''Archaeopteryx'' feather sheaths reveal sequential center-out flight-related molting strategy]. ''Communications Biology'' 3:745. [239] => [240] => === Growth === [241] => [[File:Archaeopteryx growth curve.jpg|thumb|upright=1.75|Growth trends compared with other dinosaurs and birds]] [242] => [243] => An [[histological]] study by Erickson, Norell, Zhongue, and others in 2009 estimated that ''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 ''Archaeopteryx'' might have grown as slowly as that in a mallard (2.5{{nbsp}}micrometres per day) or as fast as that in an ostrich (4.2{{nbsp}}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 {{cvt|0.8–1|kg}}. The study also found that the avialans ''[[Jeholornis]]'' and ''[[Sapeornis]]'' grew relatively slowly, as did the dromaeosaurid ''[[Mahakala (dinosaur)|Mahakala]]''. The avialans ''[[Confuciusornis]]'' and ''[[Ichthyornis]]'' grew relatively quickly, following a growth trend similar to that of modern birds.''EurekAlert!'' (8 October 2009), [http://www.eurekalert.org/pub_releases/2009-10/amon-wn100809.php "''Archaeopteryx'' was not very bird-like"]. One of the few modern birds that exhibit slow growth is the flightless [[Kiwi (bird)|kiwi]], and the authors speculated that ''Archaeopteryx'' and the kiwi had similar [[basal metabolic rate]]. [244] => [245] => === Daily activity patterns === [246] => Comparisons between the [[sclerotic ring|scleral ring]]s of ''Archaeopteryx'' and modern birds and reptiles indicate that it may have been [[Diurnality|diurnal]], similar to most modern birds.{{Cite journal |last1=Schmitz, L. |last2=Motani, R. |year=2011 |title=Nocturnality in Dinosaurs Inferred from Scleral Ring and Orbit Morphology |journal=Science |volume=332 |issue=6030 |pages=705–8 |bibcode=2011Sci...332..705S |doi=10.1126/science.1200043 |pmid=21493820 |s2cid=33253407}} [247] => [248] => == Palaeoecology == [249] => [[File:Archaeopteryx lithographica by durbed.jpg|thumb|Restoration of ''Archaeopteryx'' chasing a juvenile ''[[Compsognathus]]'']] [250] => The richness and diversity of the [[Solnhofen limestone]]s in which all specimens of ''Archaeopteryx'' have been found have shed light on an ancient Jurassic Bavaria strikingly different from the present day. The latitude was similar to [[Florida]], though the climate was likely to have been drier, as evidenced by fossils of plants with adaptations for arid conditions and a lack of terrestrial sediments characteristic of rivers. Evidence of plants, although scarce, include [[cycad]]s and conifers while animals found include a large number of insects, small lizards, [[pterosaur]]s, and ''[[Compsognathus]]''. [251] => [252] => The excellent preservation of ''Archaeopteryx'' fossils and other terrestrial fossils found at [[Solnhofen]] indicates that they did not travel far before becoming preserved. The ''Archaeopteryx'' 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. The pterosaurs included species such as ''[[Rhamphorhynchus (pterosaur)|Rhamphorhynchus]]'' belonging to the [[Rhamphorhynchidae]], the group which dominated the [[ecological niche]] currently occupied by [[seabird]]s, 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 [[Vagrancy (biology)|vagrants]] from the larger islands {{cvt|50|km}} to the north. [253] => [254] => 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 ({{cvt|3|m|disp=sqbr|0}}) 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. [255] => [[File:Archaeopteryx in flight at AMNH.jpg|thumb|Skeletal reconstruction of ''Archaeopteryx'' in gliding posture, [[American Museum of Natural History]]]] [256] => 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 on the basis of the curvature of the claws which has since been questioned.{{Cite journal |last1=Pike |first1=A. V. L. |last2=Maitland |first2=D. P. |date=2004 |title=Scaling of bird claws |journal=Journal of Zoology |language=en |volume=262 |issue=1 |pages=73–81 |doi=10.1017/s0952836903004382 |issn=0952-8369}} 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 and specialist species|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. [257] => [258] => == See also == [259] => {{Portal|Paleontology|Dinosaurs|Birds|Evolutionary biology}} [260] => * [[Dinosaur coloration]] [261] => * [[Evolution of birds]] [262] => * [[Feathered dinosaur]] [263] => * [[Origin of birds]] [264] => * ''[[Ostromia]]'' [265] => * ''[[Rhamphorhynchus]]'' [266] => * [[Temporal paradox (paleontology)]] [267] => * ''[[Xiaotingia]]'' [268] => [269] => == References == [270] => {{Reflist|30em|refs= [271] => {{Cite journal |last1=Mayr |first1=Gerald |last2=Pohl |first2=Burkhard |last3=Hartman |first3=Scott |last4=Peters |first4=D. Stefan |date=January 2007 |title=The tenth skeletal specimen of Archaeopteryx |journal=Zoological Journal of the Linnean Society |volume=149 |issue=1 |pages=97–116 |doi=10.1111/j.1096-3642.2006.00245.x |doi-access=free}} [272] => [273] => {{Cite journal |last1=Charig |first1=A. J. |author-link1=Alan J. Charig |last2=Greenaway |first2=F. |last3=Milner |first3=A. C. |last4=Walker |first4=C. A. |last5=Whybrow |first5=P. 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E. |year=1960 |title=Opinion 1084, Proposed addition of the generic name ''Archaeopteryx'' VON MEYER, 1861 and the specific name ''Lithographica'', VON MEYER, 1861, as published in the binomen ''Archaeopteryx Lithographica'' to the official lists (Class Aves) |url=https://www.biodiversitylibrary.org/page/12220197 |journal=Bulletin of Zoological Nomenclature |volume=17 |issue=6–8 |pages=224–226}} [376] => [377] => {{Cite journal |last=Thulborn |first=R. A. |year=1984 |title=The avian relationships of ''Archaeopteryx'', and the origin of birds |journal=Zoological Journal of the Linnean Society |volume=82 |issue=1–2 |pages=119–158 |doi=10.1111/j.1096-3642.1984.tb00539.x}} [378] => [379] => {{Cite web |last=Nedin |first=Chris |date=15 December 2007 |title=On ''Archaeopteryx'', Astronomers, and Forgery |url=http://www.talkorigins.org/faqs/archaeopteryx/forgery.html |url-status=live |archive-url=https://web.archive.org/web/20070315180204/http://www.talkorigins.org/faqs/archaeopteryx/forgery.html |archive-date=15 March 2007 |access-date=17 March 2007}} [380] => [381] => [http://www.ucmp.berkeley.edu/diapsids/birds/archaeopteryx.html ''Archaeopteryx'': An Early Bird] – [[University of California, Berkeley]], Museum of Paleontology. Retrieved 18 October 2006. 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[388] => [389] => Wellnhofer, P., & Tischlinger, H. (2004). ''{{lang|de|Das "Brustbein" von Archaeopteryx bavarica Wellnhofer 1993 – eine Revision}}''. Archaeopteryx. 22: 3–15. [Article in German]. [390] => [391] => {{Cite book |last=P. Wellnhofer |title=Feathered Dragons |publisher=Indiana University Press |year=2004 |isbn=978-0-253-34373-4 |veditors=Currie PJ, Koppelhus EB, Shugar MA, Wright JL |pages=282–300 |chapter=The Plumage of ''Archaeopteryx''}} [392] => [393] => [http://www.focus.de/wissen/natur/messen-wiedergefundener-archaeopteryx-ist-wohl-neue-art_aid_448677.html ''{{lang|de|Wiedergefundener Archaeopteryx ist wohl neue Art}}''] {{in lang|de}}. ''[[Die Zeit]]''. Accessed: 17 July 2012. [394] => [395] => {{Cite journal |last=Witmer |first=Lawrence M. |date=4 August 2004 |title=Inside the oldest bird brain |journal=Nature |volume=430 |issue=7000 |pages=619–620 |doi=10.1038/430619a |pmid=15295579 |s2cid=1317955}} [396] => [397] => {{Cite book |last=Witmer |first=Lawrence M. |url=https://archive.org/details/mesozoicbirdsabo00chia_070 |title=Mesozoic Birds: Above the Heads of Dinosaurs |publisher=University of California Press |year=2002 |isbn=978-0-520-20094-4 |editor-last=Witmer, L. |location=Berkeley |pages=[https://archive.org/details/mesozoicbirdsabo00chia_070/page/n13 3]–30 |chapter=The debate on avian ancestry |editor-last2=Chiappe, L. |url-access=limited}} [398] => [399] => {{Cite journal |last=Yalden D. W. |year=1984 |title=What size was ''Archaeopteryx''? |journal=Zoological Journal of the Linnean Society |volume=82 |issue=1–2 |pages=177–188 |doi=10.1111/j.1096-3642.1984.tb00541.x}} [400] => [401] => {{Cite journal |last1=Zweers |first1=G. A. |last2=Van den Berge, J. C. |year=1997 |title=Evolutionary patterns of avian trophic diversification |journal=Zoology: Analysis of Complex Systems |volume=100 |pages=25–57 |issn=0944-2006}} [402] => [403] => }} [404] => [405] => == Further reading == [406] => * G. R. de Beer (1954). ''Archaeopteryx lithographica: a study based upon the British Museum specimen''. Trustees of the British Museum, London. [407] => * P. Chambers (2002). ''Bones of Contention: The Fossil that Shook Science.'' John Murray, London. {{ISBN|0-7195-6059-4}}. [408] => * A. Feduccia (1996). ''The Origin and Evolution of Birds''. Yale University Press, New Haven. {{ISBN|0-300-06460-8}}. [409] => * [[Gerhard Heilmann|Heilmann, G.]] (1926). ''[[The Origin of Birds (book)|The Origin of Birds]]''. Witherby, London. [410] => * T. H. Huxley. (1871). ''Manual of the anatomy of vertebrate animals''. London. [411] => * H. von Meyer (1861). ''Archaeopterix lithographica (Vogel-Feder) und Pterodactylus von Solenhofen''. {{lang|de|Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde}}. ''1861'': 678–679, plate V. [Article in German]. [https://books.google.com/books?id=6RAFAAAAQAAJ&pg=PA678 Full text, Google Books]. [412] => * P. Shipman (1998). ''Taking Wing: Archaeopteryx and the Evolution of Bird Flight''. Weidenfeld & Nicolson, London. {{ISBN|0-297-84156-4}}. [413] => * P. Wellnhofer (2008). ''Archaeopteryx – Der Urvogel von Solnhofen'' (in German). Verlag Friedrich Pfeil, Munich. {{ISBN|978-3-89937-076-8}}. [414] => [415] => == External links == [416] => {{Wikispecies|Archaeopteryx}} [417] => {{commons category|Archaeopteryx}} [418] => * [http://www.talkorigins.org/faqs/archaeopteryx/info.html All About ''Archaeopteryx''], from [[Talk.Origins]]. [419] => * [https://www.sciencedaily.com/releases/2010/05/100510151348.htm Use of SSRL X-ray takes 'transformative glimpse'] – A look at chemicals linking birds and dinosaurs [420] => * [http://www.ucmp.berkeley.edu/diapsids/birds/archaeopteryx.html ''Archaeopteryx'': An Early Bird] – University of California Museum of Paleontology [421] => * [http://www.ucmp.berkeley.edu/diapsids/avians.html Are Birds Really Dinosaurs?] – University of California Museum of Paleontology [422] => [423] => {{Jurassic Footer}} [424] => {{Paraves|A.|state=autocollapse}} [425] => {{Birds}} [426] => {{Taxonbar|from=Q100196}} [427] => {{Authority control}} [428] => [429] => [[Category:Paravians]] [430] => [[Category:Feathered dinosaurs]] [431] => [[Category:Late Jurassic dinosaurs of Europe]] [432] => [[Category:Jurassic Germany]] [433] => [[Category:Solnhofen fauna]] [434] => [[Category:Fossils of Germany]] [435] => [[Category:Transitional fossils]] [436] => [[Category:Fossil taxa described in 1861]] [437] => [[Category:Taxa named by Christian Erich Hermann von Meyer]] [438] => [[Category:Multispecific non-avian dinosaur genera]] [] => )
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Archaeopteryx

Archaeopteryx is a well-known fossil bird from the Late Jurassic period, approximately 150 million years ago. It is considered to be a transitional species between non-avian dinosaurs and modern birds, possessing both reptilian and avian characteristics.

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It is considered to be a transitional species between non-avian dinosaurs and modern birds, possessing both reptilian and avian characteristics. The fossil was first discovered in Germany in the early 1860s and has since been extensively studied, making it one of the most important fossils in the field of paleontology. Archaeopteryx is notable for its feathered wings and skeletal features that resemble both reptiles and birds. It had teeth, claws, and a long bony tail, which are typical of non-avian dinosaurs, while also possessing feathers, wings, and a wishbone, characteristic of modern birds. These anatomical features provide strong evidence for the evolutionary connection between dinosaurs and birds. The discovery of Archaeopteryx fueled the debate about the origin of birds and their relationship to dinosaurs. Its unique combination of avian and reptilian traits supported the hypothesis that birds are descended from theropod dinosaurs. The fossils of Archaeopteryx have been crucial in shaping our understanding of the evolution of flight and the development of bird-like features. Several species of Archaeopteryx have been identified, all known from fossil specimens found in Germany. They were likely small, carnivorous creatures that lived in a forested environment. The detailed preservation of their feathers has provided valuable insights into the evolutionary development and structure of feathers. Due to its significant role in understanding avian evolution, Archaeopteryx has become an iconic fossil and has had a major impact on paleontological discoveries and theories. It is often considered the missing link between dinosaurs and birds, and its fossils have greatly contributed to our knowledge of the origins and evolution of birds.

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