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121-458: One of its highlights is the specimen of the early bird Archaeopteryx discovered in 1861. The museum is also interesting because of the architecture of its building, the former urban college of arts and crafts. [REDACTED] Media related to Paläontologisches Museum München at Wikimedia Commons 48°08′51″N 11°33′50″E  /  48.14750°N 11.56389°E  / 48.14750; 11.56389 This Munich -related article

242-498: 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. 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

363-442: A clear trilling courtship call. A curve-tipped secondary on each wing is dragged against an adjacent ridged secondary at high speeds (as many as 110 times per second—slightly faster than a hummingbird's wingbeat) to create a stridulation much like that produced by some insects. Both Wilson's and common snipe have modified outer tail feathers which make noise when they are spread during the birds' roller coaster display flights; as

484-652: A complete absence of air bubbles in the rock slabs is further proof that the specimen is authentic. Most of the specimens of Archaeopteryx that have been discovered come from the Solnhofen limestone in Bavaria, southern Germany, which is a Lagerstätte , a rare and remarkable geological formation known for its superbly detailed fossils laid down during the early Tithonian stage of the Jurassic period, approximately 150.8–148.5   million years ago. Archaeopteryx

605-490: 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. 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

726-416: A dorsal 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. In

847-434: 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

968-438: A key piece of evidence for the origin of birds, the transitional fossils debate, and confirmation of evolution . 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. Over

1089-511: A large extent in size and shape (except in the case of mutation or damage), though not necessarily in the pattern. They are given different names depending on their position along the wing. Primaries are connected to the manus (the bird's "hand", composed of carpometacarpus and phalanges ); these are the longest and narrowest of the remiges (particularly those attached to the phalanges), and they can be individually rotated. These feathers are especially important for flapping flight, as they are

1210-528: A more protracted moult. In many species, there is more than one focus along the wing. Here, moult begins at all foci simultaneously, but generally proceeds only in one direction. Most grouse, for example, have two wing foci: one at the wingtip, the other between feathers P1 and S1. In this case, moult proceeds descendantly from both foci. Many large, long-winged birds have multiple wing foci. Birds that are heavily "wing-loaded"—that is, heavy-bodied birds with relatively short wings—have great difficulty flying with

1331-780: 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

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1452-473: A reversed toe—a universal feature of birds—limiting its ability to perch on branches and implying a terrestrial or trunk-climbing lifestyle. This has been interpreted as evidence of theropod ancestry. In 1988, Gregory S. Paul 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,

1573-485: A single feather which appeared to resemble a modern bird's remex (wing feather), but he had heard of and been shown a rough sketch of the London specimen, to which he referred as a " Skelett eines mit ähnlichen Federn bedeckten Tieres " ("skeleton of an animal covered in similar feathers"). In German, this ambiguity is resolved by the term Schwinge which does not necessarily mean a wing used for flying. Urschwinge

1694-593: A single species, although this is still debated. Most of these twelve fossils include impressions of feathers. Because these feathers are of an advanced form ( flight feathers ), these fossils are evidence that the evolution of feathers began before the Late Jurassic. The type specimen of Archaeopteryx was discovered just two years after Charles Darwin published On the Origin of Species . Archaeopteryx seemed to confirm Darwin's theories and has since become

1815-458: A sound during display flights. Tiny serrations on the leading edge of their remiges help owls to fly silently (and therefore hunt more successfully), while the extra-stiff rectrices of woodpeckers help them to brace against tree trunks as they hammer on them. Even flightless birds still retain flight feathers, though sometimes in radically modified forms. The remiges are divided into primary and secondary feathers based on their position along

1936-501: A sound during territorial or courtship displays. Over time, a small number of bird species have lost their ability to fly. Some of these, such as the steamer ducks , show no appreciable changes in their flight feathers. Some, such as the Titicaca grebe and a number of the flightless rails, have a reduced number of primaries. The remiges of ratites are soft and downy; they lack the interlocking hooks and barbules that help to stiffen

2057-433: A state known as diastataxis (those that do have the fifth secondary are said to be eutaxic). In these birds, the fifth set of secondary covert feathers does not cover any remiges, possibly due to a twisting of the feather papillae during embryonic development. Loons , grebes, pelicans , hawks and eagles , cranes , sandpipers , gulls , parrots, and owls are among the families missing this feather. Tertials arise in

2178-608: A torso, the Maxberg Specimen (S5) was discovered in 1956 near Langenaltheim ; it was brought to the attention of professor Florian Heller in 1958 and described by him in 1959. The specimen is missing its head and tail, although the rest of the skeleton is mostly intact. Although it was once exhibited at the Maxberg Museum in Solnhofen , it is currently missing. It belonged to Eduard Opitsch , who loaned it to

2299-446: Is a stub . You can help Misplaced Pages by expanding it . This article about a Bavarian building or structure is a stub . You can help Misplaced Pages by expanding it . Archaeopteryx Archaeopteryx ( / ˌ ɑːr k iː ˈ ɒ p t ər ɪ k s / ; lit.   ' old-wing ' ), sometimes referred to by its German name, " Urvogel " ( lit. Primeval Bird ) is a genus of bird -like dinosaurs . The name derives from

2420-457: Is also found, though to a lesser extent, in some other species that feed along tree trunks, including treecreepers and woodcreepers . Scientists have not yet determined the function of all flight feather modifications. Male swallows in the genera Psalidoprocne and Stelgidopteryx have tiny recurved hooks on the leading edges of their outer primaries, but the function of these hooks is not yet known; some authorities suggest they may produce

2541-531: Is called saltatory or transilient wing moults. In simple forms, this involves the moulting and replacement of odd-numbered primaries and then the even-numbered primaries. There are however complex variations with differences based on life history. Arboreal woodpeckers , which depend on their tails—particularly the strong central pair of rectrices—for support while they feed, have a unique tail moult. Rather than moulting their central tail feathers first, as most birds do, they retain these feathers until last. Instead,

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2662-493: Is diminished when the outer primaries are worn, and absent when those feathers have been moulted. During the northern lapwing 's zigzagging display flight, the bird's outer primaries produce a humming sound. The outer primaries of the male American woodcock are shorter and slightly narrower than those of the female, and are likely the source of the whistling and twittering sounds made during his courtship display flights. Male club-winged manakins use modified secondaries to make

2783-473: Is known as moult (molt in the United States). The loss of wing and tail feathers can affect a bird's ability to fly (sometimes dramatically) and in certain families can impair the ability to feed or perform courtship displays . The timing and progression of flight feather moult therefore varies among families. For most birds, moult begins at a certain specific point, called a focus (plural foci), on

2904-509: Is missing much of the skull and one forelimb. It is privately owned and has yet to be given a name. Palaeontologists of the Ludwig Maximilian University of Munich studied the specimen, which revealed previously unknown features of the plumage, such as feathers on both the upper and lower legs and metatarsus , and the only preserved tail tip. A twelfth specimen had been discovered by an amateur collector in 2010 at

3025-660: Is moulted at a different time. The flight feathers of adults and juveniles can differ considerably in length, particularly among the raptors. Juveniles tend to have slightly longer rectrices and shorter, broader wings (with shorter outer primaries, and longer inner primaries and secondaries) than do adults of the same species. However, there are many exceptions. In longer-tailed species, such as swallow-tailed kite , secretary bird and European honey buzzard , for example, juveniles have shorter rectrices than adults do. Juveniles of some Buteo buzzards have narrower wings than adults do, while those of large juvenile falcons are longer. It

3146-496: 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. 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

3267-486: Is referred to as the primary extension or primary projection. As with wing formulae, this measurement is useful for distinguishing between similarly plumaged birds; however, unlike wing formulae, it is not necessary to have the bird in-hand to make the measurement. Rather, this is a useful relative measurement—some species have long primary extensions, while others have shorter ones. Among the Empidonax flycatchers of

3388-458: 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. Charig et al. pointed to the presence of hairline cracks in the slabs running through both rock and fossil impressions, and mineral growth over

3509-410: Is theorized that the differences help young birds compensate for their inexperience, weaker flight muscles and poorer flying ability. A wing formula describes the shape of distal end of a bird's wing in a mathematical way. It can be used to help distinguish between species with similar plumages, and thus is particularly useful for those who ring (band) birds. To determine a bird's wing formula,

3630-560: Is to be studied by famed paleornithologist Jingmai O'Connor . 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 in London . Most of their supposed evidence for

3751-598: The Archaeopteryx , with a long lizard-like tail, bearing a pair of feathers on each joint, and with its wings furnished with two free claws, has been discovered in the oolitic slates of Solnhofen. Hardly any recent discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world." The Greek word archaīos ( ἀρχαῖος ) means 'ancient, primeval'. Ptéryx primarily means 'wing', but it can also be just 'feather'. Meyer suggested this in his description. At first he referred to

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3872-663: The Field Museum in Chicago, US. One of two specimens in an institution outside Europe, the specimen was originally identified in a private collection in Switzerland , and had been acquired by these collectors in 1990, prior to Germany's 2015 ban on exporting Archaeopteryx specimens. The specimen was acquired by the Field Museum in 2022, and went on public display in 2024 following two years of preparation. The specimen

3993-605: 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 ). 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

4114-700: The Late Jurassic around 150 million years ago, in what is now southern Germany, during a time when Europe was an archipelago of islands in a shallow warm tropical sea, much closer to the equator than it is now. Similar in size to a Eurasian magpie , with the largest individuals possibly attaining the size of a raven , the largest species of Archaeopteryx could grow to about 0.5 m (1 ft 8 in) in length. Despite their small size, broad wings, and inferred ability to fly or glide, Archaeopteryx had more in common with other small Mesozoic dinosaurs than with modern birds. In particular, they shared

4235-476: The London Specimen (BMNH 37001), was unearthed in 1861 near Langenaltheim , Germany, and perhaps given to local physician Karl Häberlein in return for medical services. He then sold it for £700 (roughly £83,000 in 2020 ) to the 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

4356-471: The ancient Greek ἀρχαῖος ( archaīos ), meaning "ancient", and πτέρυξ ( ptéryx ), meaning "feather" or "wing". Between the late 19th century and the early 21st century, Archaeopteryx was generally accepted by palaeontologists and popular reference books as the oldest known bird (member of the group Avialae ). Older potential avialans have since been identified, including Anchiornis , Xiaotingia , and Aurornis . Archaeopteryx lived in

4477-413: The ankle bone , interdental plates , an obturator process of the ischium , and long chevrons in the tail. In particular, Ostrom found that Archaeopteryx was remarkably similar to the theropod family Dromaeosauridae . Archaeopteryx had three separate digits on each fore-leg each ending with a "claw". Few birds have such features. Some birds, such as ducks , swans , Jacanas ( Jacana sp.), and

4598-408: The hoatzin ( Opisthocomus hoazin ), have them concealed beneath their leg-feathers. Specimens of Archaeopteryx were most notable for their well-developed flight feathers . They were markedly asymmetrical and showed the structure of flight feathers in modern birds, with vanes given stability by a barb - barbule - barbicel arrangement. The tail feathers were less asymmetrical, again in line with

4719-433: The olecranon and performing the same function as true tertials) in an effort to distinguish them from other secondaries. The term humeral is sometimes used for birds such as the albatrosses and pelicans that have a long humerus. The calami of the flight feathers are protected by a layer of non-flight feathers called covert feathers or tectrices (singular tectrix ), at least one layer of them both above and beneath

4840-527: The taxon was originally described in a different genus. "Archaeopteryx" vicensensis (Anon. fide Lambrecht, 1933) is a nomen nudum for what appears to be an undescribed pterosaur. Flight feather Flight feathers ( Pennae volatus ) are the long, stiff, asymmetrically shaped, but symmetrically paired pennaceous feathers on the wings or tail of a bird; those on the wings are called remiges ( / ˈ r ɛ m ɪ dʒ iː z / ), singular remex ( / ˈ r iː m ɛ k s / ), while those on

4961-399: 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

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5082-557: The ICZN after four years of debate, and the London specimen was designated the neotype on 3 October 2011. Below is a cladogram published in 2013 by Godefroit et al. Aurornis [REDACTED] Anchiornis [REDACTED] Archaeopteryx [REDACTED] Xiaotingia [REDACTED] Jeholornis [REDACTED] Rahonavis [REDACTED] Balaur [REDACTED] Avebrevicauda (includes modern birds) [REDACTED] It has been argued that all

5203-716: The Latin word for "helmsman", help the bird to brake and steer in flight. These feathers lie in a single horizontal row on the rear margin of the anatomic tail. Only the central pair are attached (via ligaments ) to the tail bones; the remaining rectrices are embedded into the rectricial bulbs , complex structures of fat and muscle that surround those bones. Rectrices are always paired, with a vast majority of species having six pairs. They are absent in grebes and some ratites , and greatly reduced in size in penguins. Many grouse species have more than 12 rectrices. In some species (including ruffed grouse , hazel grouse and common snipe ),

5324-420: 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

5445-551: 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. The Eichstätt Specimen (JM 2257) was discovered in 1951 near Workerszell , Germany, and described by Peter Wellnhofer in 1974. Currently located at

5566-650: The Schamhaupten quarry, but the finding was only announced in February 2014. It was scientifically described in 2018. It represents a complete and mostly articulated skeleton with skull. It is the only specimen lacking preserved feathers. It is from the Painten Formation and somewhat older than the other specimens. The existence of a thirteenth specimen (the Chicago specimen ) was announced in 2024 by

5687-603: 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

5808-418: The airfoil shape of the bird's wing. Secondaries tend to be shorter and broader than primaries, with blunter ends (see illustration). They vary in number from 6 in hummingbirds to as many as 40 in some species of albatross . In general, larger and longer-winged species have a larger number of secondaries. Birds in more than 40 non-passerine families seem to be missing the fifth secondary feather on each wing,

5929-649: The basis for a distinct species, A. bavarica , but more recent studies suggest it belongs to A. siemensii . An eighth, fragmentary specimen was discovered in 1990 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

6050-682: The bird dives, wind flows through the modified feathers and creates a series of rising and falling notes, which is known as "winnowing". Differences between the sounds produced by these two former conspecific subspecies—and the fact that the outer two pairs of rectrices in Wilson's snipe are modified, while only the single outermost pair are modified in common snipe—were among the characteristics used to justify their splitting into two distinct and separate species. Flight feathers are also used by some species in visual displays. Male standard-winged and pennant-winged nightjars have modified P2 primaries (using

6171-405: The bird is in flight, especially in the case of raptors. The trailing edge of the wing of a juvenile bird can appear almost serrated, due to the feathers' sharp tips, while that of an older bird will be straighter-edged. The flight feathers of a juvenile bird will also be uniform in length, since they all grew at the same time. Those of adults will be of various lengths and levels of wear, since each

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6292-444: The bird's "thumb" and normally lie flush against the anterior edge of the wing—function in the same way as the slats on an airplane wing, allowing the wing to achieve a higher than normal angle of attack  – and thus lift  – without resulting in a stall . By manipulating its thumb to create a gap between the alula and the rest of the wing, a bird can avoid stalling when flying at low speeds or landing. The development of

6413-404: The bird's newly strengthened tail is best able to cope with the loss of the crucial central rectrices. Ground-feeding woodpeckers, such as the wrynecks , do not have this modified moult strategy; in fact, wrynecks moult their outer tail feathers first, with moult proceeding proximally from there. There are often substantial differences between the remiges and rectrices of adults and juveniles of

6534-454: The bird's wing closed, so as to maintain the relative positions of the feathers. While there can be considerable variation across members of a species—and while the results are obviously impacted by the effects of moult and feather regeneration—even very closely related species show clear differences in their wing formulas. The distance that a bird's longest primaries extend beyond its longest secondaries (or tertials) when its wings are folded

6655-451: The brachial region and are not considered true remiges as they are not supported by attachment to the corresponding bone, in this case the humerus. These elongated "true" tertials act as a protective cover for all or part of the folded primaries and secondaries, and do not qualify as flight feathers as such. However, many authorities use the term tertials to refer to the shorter, more symmetrical innermost secondaries of passerines (arising from

6776-609: The carpal joint) was formerly thought to be absent in some species, but the modern view of this diastataxy is that there is a gap between the fourth and fifth secondaries. Tertiary feathers growing upon the adjoining portion of the brachium are not considered true remiges. The moult of their flight feathers can cause serious problems for birds, as it can impair their ability to fly. Different species have evolved different strategies for coping with this, ranging from dropping all their flight feathers at once (and thus becoming flightless for some relatively short period of time) to extending

6897-463: The centermost pair outwards in both directions. The flight feathers of some species provide additional functionality. In some species, for example, either remiges or rectrices make a sound during flight. These sounds are most often associated with courtship or territorial displays. The outer primaries of male broad-tailed hummingbirds produce a distinctive high-pitched trill, both in direct flight and in power-dives during courtship displays; this trill

7018-415: 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. Composed of

7139-467: The dead animals would fall onto hardened surfaces, which would form a natural plane for the future slabs to split along and would leave the bulk of the fossil on one side and little on the other. Finally, the motives they suggested for a forgery are not strong, and are contradictory; one is that Richard Owen wanted to forge evidence in support of Charles Darwin's theory of evolution, which is unlikely given Owen's views toward Darwin and his theory. The other

7260-456: The descendant numbering scheme explained above) which are displayed during their courtship rituals. In the standard-winged nightjar, this modified primary consists of an extremely long shaft with a small "pennant" (actually a large web of barbules) at the tip. In the pennant-winged nightjar, the P2 primary is an extremely long (but otherwise normal) feather, while P3, P4 and P5 are successively shorter;

7381-453: The distance between the tip of the most distal primary and the tip of its greater covert (the longest of the feathers that cover and protect the shaft of that primary) is measured in millimeters. In some cases, this results in a positive number (e.g., the primary extends beyond its greater covert), while in other cases it is a negative number (e.g. the primary is completely covered by the greater covert, as happens in some passerine species). Next,

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7502-504: The dramatically coiled twin plumes of the magnificent bird-of-paradise . Owls have remiges which are serrated rather than smooth on the leading edge. This adaptation disrupts the flow of air over the wings, eliminating the noise that airflow over a smooth surface normally creates, and allowing the birds to fly and hunt silently. The rectrices of woodpeckers are proportionately short and very stiff, allowing them to better brace themselves against tree trunks while feeding. This adaptation

7623-412: The ends. These plumes are raised up over the bird's head (along with a fine spray of modified uppertail coverts) during his extraordinary display. Rectrix modification reaches its pinnacle among the birds of paradise , which display an assortment of often bizarrely modified feathers, ranging from the extremely long plumes of the ribbon-tailed astrapia (nearly three times the length of the bird itself) to

7744-443: The event that a species is missing the small distal 10th primary, as some passerines are, its lack does not impact the numbering of the remaining primaries. Ascendant numbering, on the other hand, allows for uniformity in the numbering of non-passerine primaries, as they almost invariably have four attached to the manus regardless of how many primaries they have overall. This method is particularly useful for indicating wing formulae, as

7865-413: The feather edges. These narrowings are called either notches or emarginations depending on the degree of their slope. An emargination is a gradual change, and can be found on either side of the feather. A notch is an abrupt change, and is only found on the wider trailing edge of the remex. (Both are visible on the primary in the photo showing the feathers; they can be found about halfway along both sides of

7986-401: The feather had been designated the type specimen , the name Archaeopteryx should then no longer be applied to the skeletons, thus creating significant nomenclatorial confusion. In 2007, two sets of scientists therefore petitioned the ICZN requesting that the London specimen explicitly be made the type by designating it as the new holotype specimen, or neotype . This suggestion was upheld by

8107-411: 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. 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 . The first skeleton, known as

8228-413: 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. The discovery of an eleventh specimen was announced in 2011; it was described in 2014. It is one of the more complete specimens, but

8349-415: The flight feathers of other birds. In addition, the emu 's remiges are proportionately much reduced in size, while those of the cassowaries are reduced both in number and structure, consisting merely of 5–6 bare quills. Most ratites have completely lost their rectrices; only the ostrich still has them. Penguins have lost their differentiated flight feathers. As adults, their wings and tail are covered with

8470-407: The flight feathers of the wings as well as above and below the rectrices of the tail. These feathers may vary widely in size – in fact, the upper tail tectrices of the male peafowl , rather than its rectrices, are what constitute its elaborate and colorful "train". The outermost primaries of large soaring birds, particularly raptors, often show a pronounced narrowing at some variable distance along

8591-478: The following features with the dromaeosaurids and troodontids : jaws with sharp teeth , three fingers with claws , a long bony tail, hyperextensible second toes ("killing claw"), feathers (which also suggest warm-bloodedness ), and various features of the skeleton . These features make Archaeopteryx a clear candidate for a transitional fossil between non-avian dinosaurs and avian dinosaurs (birds). Thus, Archaeopteryx plays an important role, not only in

8712-418: The fully formed feather. These growth bars and their widths have been used to determine the daily nutritional status of birds. Each light and dark bar correspond to around 24 hours and the use of this technique has been called ptilochronology (analogous to dendrochronology ). In general, juveniles have feathers which are narrower and more sharply pointed at the tip. This can be particularly visible when

8833-470: The interpretation of such biomarkers as an indicator of eumelanin in the full Archaeopteryx specimen. Carney and other colleagues also argued against the 2013 study's interpretation of the sulphate and trace metals, 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

8954-541: The left hand feather—a shallow notch on the left, and a gradual emargination on the right.) The presence of notches and emarginations creates gaps at the wingtip; air is forced through these gaps, increasing the generation of lift. Feathers on the alula or bastard wing are not generally considered to be flight feathers in the strict sense; though they are asymmetrical, they lack the length and stiffness of most true flight feathers. However, alula feathers are definitely an aid to slow flight. These feathers—which are attached to

9075-414: The letter S, those of tertials with T and those of rectrices with R. Most authorities number the primaries descendantly, starting from the innermost primary (the one closest to the secondaries) and working outwards; others number them ascendantly, from the most distal primary inwards. There are some advantages to each method. Descendant numbering follows the normal sequence of most birds' primary moult. In

9196-404: The longest primary feather is identified, and the differences between the length of that primary and that of all remaining primaries and of the longest secondary are also measured, again in millimeters. If any primary shows a notch or emargination, this is noted, and the distance between the feather's tip and any notch is measured, as is the depth of the notch. All distance measurements are made with

9317-528: The loss of even a few flight feathers. A protracted moult like the one described above would leave them vulnerable to predators for a sizeable portion of the year. Instead, these birds lose all their flight feathers at once. This leaves them completely flightless for a period of three to four weeks, but means their overall period of vulnerability is significantly shorter than it would otherwise be. Eleven families of birds, including loons , grebes and most waterfowl , have this moult strategy. The cuckoos show what

9438-433: The money to buy a cow in 1876, to innkeeper Johann Dörr, who again sold it to Ernst Otto Häberlein, the son of K. Häberlein. Placed on sale between 1877 and 1881, with potential buyers including O. C. Marsh of Yale University 's Peabody Museum, it eventually was bought for 20,000 Goldmark by the Berlin's Natural History Museum, where it now is displayed. The transaction was financed by Ernst Werner von Siemens , founder of

9559-422: The moult over a period of several years. Remiges (from the Latin for "oarsman") are located on the posterior side of the wing. Ligaments attach the long calami (quills) firmly to the wing bones, and a thick, strong band of tendinous tissue known as the postpatagium helps to hold and support the remiges in place. Corresponding remiges on individual birds are symmetrical between the two wings, matching to

9680-616: The museum until 1974. After his death in 1991, it was discovered that the specimen was missing and may have been stolen or sold. The Haarlem Specimen (TM 6428/29, also known as the Teylers Specimen ) was discovered in 1855 near Riedenburg , Germany, and described as a Pterodactylus crassipes in 1857 by Meyer. It was reclassified in 1970 by John Ostrom and is currently located at the Teylers Museum in Haarlem ,

9801-570: The next feathers in line (P2 and S2 on the wing, and both R2s on the tail) are dropped. This pattern of drop and replacement continues until moult reaches either end of the wing or tail. The speed of the moult can vary somewhat within a species. Some passerines that breed in the Arctic , for example, drop many more flight feathers at once (sometimes becoming briefly flightless) in order to complete their entire wing moult prior to migrating south, while those same species breeding at lower latitudes undergo

9922-450: The number of primaries they possess. The number in non-passerines generally varies between 9 and 11, but grebes , storks and flamingos have 12, and ostriches have 16. While most modern passerines have ten primaries, some have only nine. Those with nine are missing the most distal primary (sometimes called the remicle) which is typically very small and sometimes rudimentary in passerines. The outermost primaries—those connected to

10043-452: The number varies among individuals. Domestic pigeons have a highly variable number as a result of changes brought about over centuries of selective breeding. In order to make the discussion of such topics as moult processes or body structure easier, ornithologists assign a number to each flight feather. By convention, the numbers assigned to primary feathers always start with the letter P (P1, P2, P3, etc.) , those of secondaries with

10164-431: 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

10285-418: The outermost primary is the one with which the measurements begin. Secondaries are always numbered ascendantly, starting with the outermost secondary (the one closest to the primaries) and working inwards. Tertials are also numbered ascendantly, but in this case, the numbers continue on consecutively from that given to the last secondary (e.g. ... S5, S6, T7, T8, ... etc.). Rectrices are always numbered from

10406-418: The overall effect is a broadly forked wingtip with a very long plume beyond the lower half of the fork. Males of many species, ranging from the widely introduced ring-necked pheasant to Africa's many whydahs , have one or more elongated pairs of rectrices, which play an often-critical role in their courtship rituals. The outermost pair of rectrices in male lyrebirds are extremely long and strongly curved at

10527-423: The phalanges—are sometimes known as pinions . Secondaries are connected to the ulna . In some species, the ligaments that bind these remiges to the bone connect to small, rounded projections, known as quill knobs , on the ulna; in other species, no such knobs exist. Secondary feathers remain close together in flight (they cannot be individually separated like the primaries can) and help to provide lift by creating

10648-473: 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. In 2011, graduate student Ryan Carney and colleagues performed the first colour study on an Archaeopteryx specimen. Using scanning electron microscopy technology and energy-dispersive X-ray analysis,

10769-454: 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,

10890-488: 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 . In 2007, a review of all well-preserved specimens including

11011-410: 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

11132-414: The presence of numerous avian features, Archaeopteryx had many non-avian theropod dinosaur characteristics. Unlike modern birds, Archaeopteryx had small teeth, as well as a long bony tail, features which Archaeopteryx shared with other dinosaurs of the time. Because it displays features common to both birds and non-avian dinosaurs, Archaeopteryx has often been considered a link between them. In

11253-426: The principal source of thrust , moving the bird forward through the air. The mechanical properties of primaries are important in supporting flight. Most thrust is generated on the downstroke of flapping flight. However, on the upstroke (when the bird often draws its wing in close to its body), the primaries are separated and rotated, reducing air resistance while still helping to provide some thrust. The flexibility of

11374-454: The remainder of the body—although some feathers did not fossilize and others were obliterated during preparation, leaving bare patches on specimens—and the lower neck. There is no indication of feathering on the upper neck and head. While these conceivably may have been nude, this may still be an artefact of preservation. It appears that most Archaeopteryx specimens became embedded in anoxic sediment after drifting some time on their backs in

11495-448: The remaining "plumage patterns of Archaeopteryx remain unknown". Today, fossils of the genus Archaeopteryx are usually assigned to one or two species, A. lithographica and A. siemensii , but their taxonomic history is complicated. Ten names have been published for the handful of specimens. As interpreted today, the name A. lithographica only referred to the single feather described by Meyer . In 1954 Gavin de Beer concluded that

11616-585: The remiges (and alulae) of nestling hoatzins is much delayed compared to the development of these feathers in other young birds, presumably because young hoatzins are equipped with claws on their first two digits . They use these small rounded hooks to grasp branches when clambering about in trees, and feathering on these digits would presumably interfere with that functionality. Most youngsters shed their claws sometime between their 70th and 100th day of life, but some retain them— though callused -over and unusable— into adulthood. Rectrices (singular rectrix) from

11737-410: The remiges on the wingtips of large soaring birds also allows for the spreading of those feathers, which helps to reduce the creation of wingtip vortices , thereby reducing drag . The barbules on these feathers, friction barbules, are specialized with large lobular barbicels that help grip and prevent slippage of overlying feathers and are present in most of the flying birds. Species vary somewhat in

11858-403: The rest of the species of Archaeopteryx . In the Berlin specimen, there are "trousers" of well-developed feathers on the legs; some of these feathers seem to have a basic contour feather structure, but are somewhat decomposed (they lack barbicels as in ratites ). In part they are firm and thus capable of supporting flight. A patch of pennaceous feathers is found running along its back, which

11979-524: The same small, stiff, slightly curved feathers as are found on the rest of their bodies. The ground-dwelling kākāpō , which is the world's only flightless parrot, has remiges which are shorter, rounder and more symmetrically vaned than those of parrots capable of flight; these flight feathers also contain fewer interlocking barbules near their tips. Once they have finished growing, feathers are essentially dead structures. Over time, they become worn and abraded, and need to be replaced. This replacement process

12100-404: The same species. It has been noted that the feather, the first specimen of Archaeopteryx described, does not correspond well with the flight-related feathers of Archaeopteryx . It certainly is a flight feather of a contemporary species, but its size and proportions indicate that it may belong to another, smaller species of feathered theropod , of which only this feather is known so far. As

12221-415: The same species. Because all juvenile feathers are grown at once—a tremendous energy burden to the developing bird—they are softer and of poorer quality than the equivalent feathers of adults, which are moulted over a longer period of time (as long as several years in some cases). As a result, they wear more quickly. As feathers grow at variable rates, these variations lead to visible dark and light bands in

12342-430: The sea—the head, neck and the tail are generally bent downward, which suggests that the specimens had just started to rot when they were embedded, with tendons and muscle relaxing so that the characteristic shape ( death pose ) of the fossil specimens was achieved. This would mean that the skin already was softened and loose, which is bolstered by the fact that in some specimens the flight feathers were starting to detach at

12463-492: The second pair of rectrices (both R2 feathers) are the first to drop. (In some species in the genera Celeus and Dendropicos , the third pair is the first dropped.) The pattern of feather drop and replacement proceeds as described for passerines (above) until all other rectrices have been replaced; only then are the central tail rectrices moulted. This provides some protection to the growing feathers, since they're always covered by at least one existing feather, and also ensures that

12584-408: The situation in modern birds and also had firm vanes. The thumb did not yet bear a separately movable tuft of stiff feathers . The body plumage of Archaeopteryx is less well-documented and has only been properly researched in the well-preserved Berlin specimen . Thus, as more than one species seems to be involved, the research into the Berlin specimen's feathers does not necessarily hold true for

12705-400: 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

12826-505: 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

12947-426: 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, eleven more fossils of Archaeopteryx have surfaced. Despite variation among these fossils, most experts regard all the remains that have been discovered as belonging to

13068-587: The tail are called rectrices ( / ˈ r ɛ k t r ɪ s iː z / or / r ɛ k ˈ t r aɪ s iː z / ), singular rectrix ( / ˈ r ɛ k t r ɪ k s / ). The primary function of the flight feathers is to aid in the generation of both thrust and lift , thereby enabling flight . The flight feathers of some birds perform additional functions, generally associated with territorial displays, courtship rituals or feeding methods. In some species, these feathers have developed into long showy plumes used in visual courtship displays, while in others they create

13189-407: The team was able to detect the structure of melanosomes in the isolated feather specimen described in 1861. The resultant measurements were then compared to those of 87   modern bird species, and the original colour was calculated with a 95% likelihood to be black. The feather was determined to be black throughout, with heavier pigmentation in the distal tip. The feather studied was most probably

13310-510: The then-newly discovered Thermopolis specimen concluded that two distinct species of Archaeopteryx could be supported: A. lithographica (consisting of at least the London and Solnhofen specimens), and A. siemensii (consisting of at least the Berlin, Munich, and Thermopolis specimens). The two species are distinguished primarily by large flexor tubercles on the foot claws in A. lithographica (the claws of A. siemensii specimens being relatively simple and straight). A. lithographica also had

13431-466: The wing or tail and proceeds in a sequential manner in one or both directions from there. For example, most passerines have a focus between the innermost primary (P1, using the numbering scheme explained above) and outermost secondary (S1), and a focus point in the middle of the center pair of rectrices. As passerine moult begins, the two feathers closest to the focus are the first to drop. When replacement feathers reach roughly half of their eventual length,

13552-418: The wing. There are typically 11 primaries attached to the manus (six attached to the metacarpus and five to the phalanges), but the outermost primary, called the remicle, is often rudimentary or absent; certain birds, notably the flamingos, grebes, and storks, have seven primaries attached to the metacarpus and 12 in all. Secondary feathers are attached to the ulna. The fifth secondary remex (numbered inwards from

13673-469: The years, twelve body fossil specimens of Archaeopteryx have been found. All of the fossils come from the limestone deposits, quarried for centuries, near Solnhofen , Germany. These quarries excavate sediments from the Solnhofen Limestone formation and related units. The initial specimen was the first dinosaur to be discovered with feathers. The initial discovery, a single feather,

13794-487: 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. The Munich Specimen (BSP 1999 I 50, formerly known as the Solenhofer-Aktien-Verein Specimen )

13915-516: 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

14036-579: 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". Long in

14157-665: 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). 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. Another fragmentary fossil

14278-476: Was quite similar to the contour feathers of the body plumage of modern birds in being symmetrical and firm, although not as stiff as the flight-related feathers. Apart from that, the feather traces in the Berlin specimen are limited to a sort of "proto- down " not dissimilar to that found in the dinosaur Sinosauropteryx : decomposed and fluffy, and possibly even appearing more like fur than feathers in life (although not in their microscopic structure). These occur on

14399-461: 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 0.5 metres (1 ft 8 in) in body length and 0.7 metres (2 ft 4 in) in wingspan , with an estimated mass of 0.5 to 1 kilogram (1.1 to 2.2 lb). Archaeopteryx feathers, although less documented than its other features, were very similar in structure to modern-day bird feathers. Despite

14520-514: Was the favoured translation of Archaeopteryx among German scholars in the late nineteenth century. In English, 'ancient pinion' offers a rough approximation to this. Since then, twelve specimens have been recovered: The Berlin Specimen (HMN 1880/81) was discovered in 1874 or 1875 on the Blumenberg near Eichstätt , Germany, by farmer Jakob Niemeyer. He sold this precious fossil for

14641-411: Was unearthed in 1860 or 1861 and described in 1861 by Hermann von Meyer . It is currently located at the Natural History Museum of Berlin . 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

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