Microraptor - Knowledge

767: 548: 5372: 1752: 1227: 1335: 683: 5361: 5367: 147: 5378: 129: 1315: 1096: 1464: 2014: 877: 5401: 2101: 1166: 1856:, although later discoveries of more primitive dromaeosaurids with short forelimbs unsuitable for gliding have cast doubt on this view. Work done on the question of flight ability in other paravians, however, showed that most of them probably would not have been able to achieve enough lift for powered flight, given their limited flight strokes and relatively smaller wings. These studies concluded that 1883: 989: 1567: 2357:, the holotype had a wing length of 42 centimetres (1.38 ft), resulting in a wingspan of more than 88 centimetres (2.89 ft). The larger specimen QV1002 measured 3 centimetres (1.2 in) longer than the holotype in precaudal length, and had wing length of 47.2 centimetres (1.55 ft), resulting in a wingspan of more than 99 centimetres (3.25 ft). 1545: 2312: 1151: 2092:), the authors argued that the current body of evidence, both from morphology and phylogeny, suggests that bird flight did shift at some point from shared limb dominance to front-limb dominance and that all modern birds may have evolved from four-winged ancestors, or at least ancestors with unusually long leg feathers relative to the modern configuration. 2047: 1976:. Brougham and Brusatte criticized the anatomy of the model used by Alexander and his team, noting that the hip anatomy was not consistent with other dromaeosaurs. In most dromaeosaurids, features of the hip bone prevent the legs from splaying horizontally; instead, they are locked in a vertical position below the body. Alexander's team used a specimen of 1200: 1788:

shoulder socket faced downward and slightly backward, making it impossible for the animals to raise their arms vertically, a prerequisite for the flapping flight stroke in birds. Studies of maniraptoran anatomy have suggested that the shoulder socket did not shift into the bird-like position of a high, upward orientation close to the

1256: 2152:, although the lack of other definitive body parts consumed may suggest the low-muscle mass foot may have been eaten during a late stage of carcass consumption, possibly through scavenging. The find is a rare example of a theropod definitively consuming a Mesozoic mammal, the only other being a specimen of the 785:), a co-ossified sternum, a tail consisting of 28 to 30 vertebrae and a third finger with a short phalanx III-3. Some of the feathers Czerkas described as primary were actually attached to the leg, rather than the arm. This, along with most of the other diagnostic characters, is also present in the genus 1048:

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism.

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when on the ground, due to the limited range of motion in the wrist and the extreme length of the wing feathers. A 2010 study by Corwin Sullivan and colleagues showed that, even with the wing folded as far as possible, the feathers would still have dragged along the ground if the arms were held in a

1963:

described the construction of a lightweight three-dimensional physical model used to perform glide tests. Using several hindleg configurations for the model, they found that the biplane model, while not unreasonable, was structurally deficient and needed a heavy-headed weight distribution for stable

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was an arboreal glider relies on it to have regularly climbed or even lived in trees, when study of its anatomy have shown that its limb proportions fall in line with modern ground birds rather than climbers, and its skeleton shows none of the expected adaptations in animals specialized for climbing

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would have hindered their ability to run on the ground. Some paleontologists have suggested that feathered dinosaurs used their wings to parachute from trees, possibly to attack or ambush prey on the ground, as a precursor to gliding or true flight. In their 2007 study, Chatterjee and Templin tested

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had the anatomical features expected of a flying animal, though it would have been a less advanced form of flight compared to birds. For example, some studies suggest the shoulder joint was too primitive to allow a full flapping flight stroke. In the ancestral anatomy of theropod dinosaurs, the

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measuring 77 centimetres (2.53 ft) in length, 88–94 centimetres (2.89–3.08 ft) in wingspan and weighing 0.5–1.4 kilograms (1.1–3.1 lb). There are larger specimens which would have measured at least 80 centimetres (2.6 ft) in length, more than 99 centimetres (3.25 ft) in

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hunted only in an arboreal environment, proposing that it was also an adept hunter of fish as well. They further argued that the specimen showed a probable adaptation to a fish-eating diet, pointing to the first three teeth of the mandible being inclined anterodorsally, a characteristic often

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anchored to the upper foot bones as well as the upper and lower leg. Though not apparent in most fossils under natural light, due to obstruction from decayed soft tissue, the feather bases extended close to or in contact with the bones, as in modern birds, providing strong anchor points.

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in an attempt to remove the name from the paleornithological record by assigning it to the part least likely to be a bird. However, Xu had discovered the remains of the specimen from which the tail had been taken and published a description of it later that year, giving it the name

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Larsson, Hans, Hone, David, Dececchi, T. Alexander, Sullivan, Corwin, Xu, Xing. "THE WINGED NON-AVIAN DINOSAUR MICRORAPTOR FED ON MAMMALS: IMPLICATIONS FOR THE JEHOL BIOTA ECOSYSTEM" "Program and Abstracts. 70th Anniversary Meeting Society of Vertebrate Paleontology October 2010"

2080:) stage. Chatterjee and Templin did not take a strong stance on this possibility, noting that both a conventional interpretation and a tetrapteryx stage are equally possible. However, based on the presence of unusually long leg feathers in various feathered dinosaurs, 919:

were published by Xu and colleagues, showing a distinctive "hindwing" completely separate from the forelimb wing. The first of these specimens was discovered in 2001, and between 2001 and 2003 four more specimens were bought from private collectors by Xu's museum, the

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in this book, Czerkas strongly denounces Norell's conclusions; "The misinterpretation of the primary wing feathers as being from the hind legs stems directly to seeing what one believes and wants to see". Czerkas also denounced Norell for failing to conclude that

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Both of these studies regarded each gut contents as instances of predation. However, Hone and colleagues (2022) questioned the reliability of these interpretations and wrote that both could just as equally be attributed to scavenging. Further, they argued against

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dinosaurs in its ecosystem, and the genus is represented by more fossils than any other dromaeosaurid, with possibly over 300 fossil specimens represented across various museum collections. One specimen in particular shows evidence of active primary feather

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has led many to question this assumption on the grounds that no modern birds that have iridescent plumage are known to be nocturnal, but this argument is itself questionable as there in fact are modern nocturnal birds with iridescent plumage, such as the

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is one of the few known bird precursors to sport long flight feathers on the legs as well as the wings. Their bodies had a thick covering of feathers, with a diamond-shaped fan on the end of the tail (possibly for added stability during flight). Xu

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are not unique, but are present to varying degrees across various specimens. In a review by Phil Senter and colleagues in 2004, the scientists suggested that all these features represented individual variation across various age groups of a single

1142:. Though the researchers state that the true function of the iridescence is yet unknown, it has been suggested that the tiny dromaeosaur was using its glossy coat as a form of communication or sexual display, much as in modern iridescent birds. 1987:, which is known from complete hip bones preserved in three dimensions, also shows hip sockets directed partially upward, possibly allowing the legs to splay more than in other dromaeosaurs. However, Hartman and colleagues suggested that 1734:

could have parachuted short distances, as between closely spaced tree branches. Wind tunnel experiments have demonstrated that sustaining a high-lift coefficient at the expense of high drag was likely the most efficient strategy for

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revealed preserved probable gut contents, and a full study on them was later published in 2022 by David Hone and colleagues. These consisted of the remains of a mammal, primarily a complete and articulated right foot (including all

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being a specialist in either or both arboreal or aquatic hunting, citing the broad range of vertebrate gut contents (i.e. fish, mammals, lizards, birds) as evidence for a generalist hunting strategy, and that neither required that

840:, BMNHC PH881, showed several features previously unknown in the animal, including the probably glossy-black iridescent plumage coloration. The new specimen also featured a bifurcated tailfan, similar in shape to previously known 1804:, is curved in such a way that the shoulder joint could only have been positioned high on the back, allowing for a nearly vertical upstroke of the wing. This possibly advanced shoulder anatomy, combined with the presence of a 844:

tailfans except sporting a pair of long, narrow feathers at the center of the fan. The new specimen also showed no sign of the nuchal crest, indicating that the crest inferred from the holotype specimen may be an artifact of

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which was crushed flat to make their model, which Brougham and Brusatte argued did not reflect its actual anatomy. Later in 2010, Alexander's team responded to these criticisms, noting that the related dromaeosaur

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were secondarily flightless (Czerkas, 2002). The current consensus is that there is not enough evidence to conclude whether dromaeosaurs descended from an ancestor with some aerodynamic abilities. The work of Xu

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style of gliding: launching itself from a perch, the animal would have swooped downward in a deep U-shaped curve and then lifted again to land on another tree. The feathers not directly employed in the biplane

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was controversial, because of the unusual circumstances of its first description. The first specimen to be described was part of a chimeric specimen—a patchwork of different feathered dinosaur species (

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were large enough to generate the lift necessary for powered launching into flight even without a fully vertical flight stroke. A 2016 study of incipient flight ability in paravians demonstrated that

1130:, these cells were shaped in a manner consistent with black, glossy coloration in modern birds. These rod-shaped, narrow melanosomes were arranged in stacked layers, much like those of a modern 2137:. It corresponds to an animal with an estimated snout to vent length of 80 mm (3.1 in) and a mass of 13–43 g (0.46–1.52 oz). The unguals of the foot are less curved than in 4522: 921: 589: 1040:. Bands of dark and light present on some specimens may indicate color patterns present in life, though at least some individuals almost certainly possessed an iridescent black coloration. 1964:

gliding, which they deemed unlikely. The study indicated that a laterally abducted hindwing structure represented the most biologically and aerodynamically consistent configuration for

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fused sternum, asymmetrical feathers, and features of the shoulder girdle indicated that it could fly under its own power, rather than merely gliding. Today, most scientists agree that

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Pei, Rui; Pittman, Michael; Goloboff, Pablo A.; Dececchi, T. Alexander; Habib, Michael B.; Kaye, Thomas G.; Larsson, Hans C.E.; Norell, Mark A.; Brusatte, Stephen L.; Xu, Xing (2020).

980:(2007) suggested that the ancestral dromaeosaur could not glide or fly, but that there was good evidence that it was small-bodied (around 65 cm long and 600–700 g in mass). 1860:

probably evolved flight and its associated features (fused sternum, alula, etc.) independently of the ancestors of birds. In 2024, Kiat and O'Connor analyzed that Mesozoic birds and

1053:, such as a combination of unserrated and partially serrated teeth with constricted 'waists', and unusually long upper arm bones, are shared with both primitive avians and primitive 4527: 1909:), and that the latter posture would have been anatomically impossible. Using this biplane model, Chatterjee was able to calculate possible methods of gliding and determined that 2176:

bird in its abdomen, specifically a partial wing and feet. Their position implies the bird was swallowed whole and head-first, which the authors interpreted as implying that the

4483: 912:, the explicit definition of the term "bird" that Norell employs, would definitely exclude BPM 1 3-13. However, he does not consider the specimen to belong to Avialae either. 636:. However, there is some doubt whether Olson in fact succeeded in meeting all the formal requirements for establishing a new taxon. Namely, Olson designated the specimen as a 2716:

Czerkas, Sylvia J. ed. (2002) "Feathered Dinosaurs and the Origin of Flight" The Dinosaur Museum Journal Volume 1. Blanding, Utah, USA. The Dinosaur Museum, August 1, 2002

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is the most abundant non-avialan dinosaur fossil type found in this formation. In 2010, it was reported that there were over 300 undescribed specimens attributable to

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was significantly different ecologically compared to extant corvids and icterids with dark, iridescent plumage, which are social omnivores of more open habitats.

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Czerkas (2002) mistakenly described the fossil as having no long feathers on its legs, but only on its hands and arms, as he illustrated on the cover of his book

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Federico L. Agnolín & Fernando E. Novas (2013). "Avian ancestors. A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria,

1852:, were secondarily flightless. The work of Xu and colleagues also suggested that the ancestors of dromaeosaurids were probably small, arboreal, and capable of 2799:

Feduccia, A.; Lingham-Soliar, T; Hinchliffe, JR (2005). "Do feathered dinosaurs exist? Testing the hypothesis on neontological and paleontological evidence".

662:) was subsequently used more frequently than the original name; as such, this and the chimeric nature of the specimen would render the name "Archaeoraptor" a 1968:. A further analysis by Brougham and Brusatte, however, concluded that Alexander's model reconstruction was not consistent with all of the available data on 1730:

was too narrow to successfully parachute to the ground without injury from any significant height. However, the authors did leave open the possibility that

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had both primary (anchored to the hand) and secondary (anchored to the arm) flight feathers. This standard wing pattern was mirrored on the hindlegs, with

2608: 2145:, indicating that the mammal could climb but less effectively than in the two latter genera and so was likely not arboreal but potentially scansorial. 4554: 3298:"A new tiny dromaeosaurid dinosaur from the Lower Cretaceous Jehol Group of western Liaoning and niche differentiation among the Jehol dromaeosaurids" 763:

LPM 0159, a smaller skeleton. Both individuals are preserved as articulated compression fossils; they are reasonably complete but partially damaged.

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neutral position, or extended forward as in a predatory strike. Only by keeping the wings elevated, or the upper arm extended fully backward, could

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Li, Q.; Gao, K.-Q.; Meng, Q.; Clarke, J.A.; Shawkey, M.D.; D'Alba, L.; Pei, R.; Ellision, M.; Norell, M.A.; Vinther, J. (2012). "Reconstruction of

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linking the wrist to the shoulder (which fills the space in front of the flexed wing and may support the wing against drag in modern birds) and an

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The cladogram below follows a 2012 analysis by paleontologists Phil Senter, James I. Kirkland, Donald D. DeBlieux, Scott Madsen and Natalie Toth.

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or its close relatives among the collections of several Chinese museums, though many had been altered or composited by private fossil collectors.

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on arms and hands 10–20 cm long (3.9–7.9 in) with legs and feet 11–15 cm long (4.3–5.9 in). The long feathers on the legs of

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raised the question of whether the evolution of flight in modern birds went through a four-winged stage, or whether four-winged gliders like

706:, and referred two additional specimens (the first to show well-preserved feathers) to this species. The generic name was derived from Greek 2086:, and some modern birds such as raptors, as well as the discovery of further dinosaurs with long primary feathers on their feet (such as 1956:

did have the requirements to sustain level powered flight, so it is theoretically possible that the animal flew, as opposed to gliding.

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Czerkas's interpretation of the hindleg feathers noted by Norell proved to be incorrect the following year when additional specimens of

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have been described in detail, from which most feather impressions are known. Unusual even among early birds and feathered dinosaurs,

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represented a small individual and included faint feather remnants, but was otherwise not well preserved and lacked a skull. In 2002

4046:"Reply to Brougham and Brusatte: Overall anatomy confirms posture and flight model offers insight into the evolution of bird flight" 2968: 2405:

Senter, P.; Barsold, R.; Britt, B.B.; Burnham, D.A. (2004). "Systematics and evolution of Dromaeosauridae (Dinosauria, Theropoda)".

1940:, etc. The orientation of the hindwings would also have helped the animal control its gliding flight. Chatterjee also used computer 771: 2692:

Norell, Mark, Ji, Qiang, Gao, Keqin, Yuan, Chongxi, Zhao, Yibin, Wang, Lixia. (2002). "'Modern' feathers on a non-avian dinosaur".

953:, the animal usually referred to as the earliest known bird. He cited the fused sternum and asymmetrical feathers, and argued that 3040:

Norell, Mark; Ji, Qiang; Gao, Keqin; Yuan, Chongxi; Zhao, Yibin; Wang, Lixia (2002). "'Modern' feathers on a non-avian dinosaur".

1685:, one on each of its forelimbs and hindlimbs, somewhat resembling one possible arrangement of the quartet of flight surfaces on a 2577: 5371: 3618:

Dyke, Gareth; de Kat, Roeland; Palmer, Colin; van der Kindere, Jacques; Naish, Darren; Ganapathisubramani, Bharathram (2013).

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in 2003 to describe the first specimen to preserve this feature as a "four-winged dinosaur" and to speculate that it may have

3669:"Assessing Arboreal Adaptations of Bird Antecedents: Testing the Ecological Setting of the Origin of the Avian Flight Stroke" 936:, most of the differences appeared to correspond with size, and likely age differences. Two further specimens, classified as 5360: 2994:

Xu, X.; Norell, M.A. (2006). "Non-Avian dinosaur fossils from the Lower Cretaceous Jehol Group of western Liaoning, China".

1959:

Some paleontologists have doubted the biplane hypothesis, and have proposed other configurations. A 2010 study by Alexander

3423:"New Dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the Evolution of the Dromaeosaurid Tail" 2125:) as well as the shafts of additional long bones and potentially other fragments. The foot skeleton is similar to those of 829:

has been accepted as a distinct species with the specimen reported in 2013 being distinguishable from the type specimen of

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Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (2019-07-10).

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Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (2019-07-10).

2353:(2016) which states that the wingspan of the paravians in the study would have been 2.1 times the wing length. In case of 1832: 5377: 462:), though further study has suggested that all of them represent variation in a single species, which is properly called 5550: 1123: 698:

et al. described another specimen, BPM 1 3-13, which they did not name or refer to an existing species. Later that year

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Zhou, Zhonghe; Zhang, Xiaomei; Wang, Yan; Wang, Xiaoli; Dong, Liping; Zheng, Xiaoting; O’Connor, Jingmai (2019-07-11).

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due to revisions in the ICZN rules that took place on December 31, 1999. In addition, Xu's name for the type specimen (

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was capable of true, powered flight, as opposed to or in addition to passive gliding. The resulting data showed that

146: 4496: 888:(2002) described BPM 1 3-13 as the first dinosaur known to have flight feathers on its legs as well as on its arms. 3297: 2025:

would have been awkward during normal ground movement or running. The front wing feathers would also have hindered

1945: 961:. Czerkas cited the fact that this possibly volant animal is also very clearly a dromaeosaurid to suggest that the 576:

and an as-of-yet undescribed third species) assembled from multiple specimens in China and smuggled to the USA for

924:. Xu also considered these specimens, most of which had hindwings and proportional differences from the original 5545: 5535: 4651: 4540: 625: 581: 502: 266: 2494:"Rarity of molt evidence in early pennaraptoran dinosaurs suggests annual molt evolved later among Neornithes" 2296:

as well as various night-feeding waterfowl; furthermore, as a forest-dwelling, presumably solitary carnivore,

1751: 4105:"A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight" 3800:"A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight" 1743:

did not require a sophisticated, 'modern' wing morphology to be an effective glider. However, the idea that

976:(2003) suggested that basal dromaeosaurs were probably small, arboreal, and could glide. The work of Turner 781:(2002) diagnosed the genus on the basis of having primary feathers (which in the authors' opinion made it a 718: 601: 2787: 5560: 5421: 3734:

Senter, P (2006). "Scapular orientation in theropods and basal birds, and the origin of flapping flight".

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Schmitz L, Motani R (2011). "Nocturnality in dinosaurs inferred from scleral ring and orbit morphology".

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nationalgeographic.com 2012-03-08 Ed Yong, A shiny dinosaur four-winged Microraptor gets color and gloss

4164:"The asymmetry of the carpal joint and the evolution of wing folding in maniraptoran theropod dinosaurs" 3572:

Xu, X., Zhou, Z., Wang, X., Kuang, X., Zhang, F. and Du, X. (2003). "Four-winged dinosaurs from China."

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Due to the extent of the hindwings onto most of the animal's foot, many scientists have suggested that

1226: 3372:"The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents" 1334: 1118:

based on the new specimen BMNHC PH881, which also showed several other features previously unknown in

759:; LPM 0201, its counterslab (slab and counterslab together represent the earlier BPM 1 3-13); and the 5468: 4705: 4431: 4378: 4316: 4265: 4057: 3998: 3929: 3870: 3859:"Potential for Powered Flight Neared by Most Close Avialan Relatives, but Few Crossed Its Thresholds" 3680: 3631: 3620:"Aerodynamic performance of the feathered dinosaur Microraptor and the evolution of feathered flight" 3524: 3434: 3250: 3187: 3129: 3049: 3003: 2922: 2742: 2446: 2265: 825:. Many other researchers, including Alan Feduccia and Tom Holtz, have since supported its synonymy. 501:

that formed aerodynamic surfaces on the arms and tail but also on the legs. This led paleontologist

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passed indigestible fur, feathers, and bits of bone in its droppings instead of producing pellets.

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wingspan and weighed 1.25–1.88 kilograms (2.8–4.1 lb). Aside from their extremely small size,

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was an opportunistic feeder, hunting the most common prey in both arboreal and aquatic habitats.

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In 2013 researchers announced that they had found fish scales in the abdominal cavity of another

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have avoided damaging the wing feathers. Therefore, it may have been anatomically impossible for

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are birds, accusing him of succumbing to "...the blinding influences of preconceived ideas." The

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Senter, P.; Kirkland, J. I.; Deblieux, D. D.; Madsen, S.; Toth, N. (2012). Dodson, Peter (ed.).

2431:"Model tests of gliding with different hindwing configurations in the four-winged dromaeosaurid 1761:

under two different UV light filters, revealing the extent of preserved feathers and soft tissue

528:, which is one of the few known fossil evidence of such behavior among pennaraptoran dinosaurs. 472:, initially described as another four-winged dinosaur, is usually considered to be a synonym of 5512: 5494: 5319: 5210: 5146: 5042: 4835: 4447: 4404: 4396: 4344: 4193: 4144: 4126: 4085: 4026: 3963: 3945: 3898: 3839: 3821: 3770: 3716: 3698: 3649: 3598: 3552: 3462: 3403: 3278: 3205: 3145: 3065: 2938: 2876: 2816: 2758: 2642: 2602: 2525: 2474: 1933: 1890: 1813: 642: 604:, published a description of the Microraptor's tail in an obscure journal, giving it the name 577: 5499: 5459: 674:(as it's been used in more published works than "Archaeoraptor" and was properly described). 5243: 5189: 5049: 5027: 4528:

nationalgeographic.com 2008-10-08 Ed Yong, Microraptor–the dinosaur that flew like a biplane

4439: 4386: 4334: 4324: 4273: 4183: 4175: 4134: 4116: 4075: 4065: 4016: 4006: 3953: 3937: 3888: 3878: 3829: 3811: 3762: 3706: 3688: 3639: 3542: 3532: 3489: 3452: 3442: 3393: 3383: 3268: 3258: 3195: 3137: 3057: 3011: 2930: 2866: 2808: 2750: 2515: 2505: 2464: 2454: 2317: 1561: 1310: 1037: 41: 3339:"High flyer or high fashion? A comparison of flight potential among small-bodied paravians" 2726:

Xu, Xing; Zhou, Zhinghe; Wang, Xiaolin; KUang, Xuewen; Zhang, Fucheng; Du, Xiangke (2003).

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Turner, Alan H.; Pol, Diego; Clarke, Julia A.; Erickson, Gregory M.; Norell, Mark (2007).

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Dececchi, T. Alexander; Larsson, Hans C. E.; Pittman, Michael; Habib, Michael B. (2020).

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Xu, X., Zhou, Z., and Wang, X. (2000). "The smallest known non-avian theropod dinosaur."

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and argued that the study was insufficient for determining a likely flight pattern for

1898: 1853: 1793: 1718: 1706: 1701:, with asymmetrical vanes on the arm, leg, and tail feathers. As in modern bird wings, 617: 506: 405: 397: 3114: 2785:

Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,

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Dececchi, T. Alexander; Larsson, Hans C. E. (2011-08-09). Farke, Andrew Allen (ed.).

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had argued that the evolution of bird flight may have gone through a four-winged (or

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techniques, the researchers compared their arrangements to those of modern birds. In

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died shortly after eating the lizard and before significant digestion had occurred.

2100: 1721:, and probably lived mainly in trees, because the hindwings anchored to the feet of 1165: 1083:

are very primitive members of two closely related groups, and both are close to the

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dinosaurs discovered with the impressions of feathers and wings. Seven specimens of

5341: 5252: 5173: 5166: 5131: 5110: 5095: 5034: 4901: 4894: 4887: 4877: 4824: 4809: 3484:(Dinosauria: Dromaeosauridae) from the Jiufotang Formation in northeastern China". 3231: 3157: 3077: 2770: 2213: 1522: 1379: 1222: 1054: 726: 278: 4277: 3493: 2349:

The wingspan estimate is based on the equation presented in the study by Dececchi

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appeared. However, other scientists have argued that the shoulder girdle in some

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or fly, the forewings and hindwings must have been on different levels (as on a

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Proceedings of the National Academy of Sciences of the United States of America

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specimens that had been described by 2002, which have 24 to 26 tail vertebrae.

5289: 5266: 5218: 4969: 4938: 4816: 4794: 4602: 4593: 4391: 4367:"Microraptor with Ingested Lizard Suggests Non-specialized Digestive Function" 4366: 3883: 3858: 3766: 2307: 2256: 2148:

It is ambiguous whether the mammal had been predated upon or scavenged by the

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might actually be a basal bird group, and that later, larger, species such as

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In 2010 researchers announced that further preparation of the type fossil of

2069:

were an evolutionary side-branch that left no descendants. As early as 1915,

2038:

to have used its clawed forelimbs in capturing prey or manipulating objects.

1122:. By analyzing the fossilized melanosomes (pigment cells) in the fossil with 5327: 5259: 4954: 4946: 4852: 4663: 4611: 4443: 4329: 4070: 4011: 3941: 3263: 3200: 3175: 2934: 2670: 2622: 2459: 2234: 2122: 2088: 1941: 1906: 947:

Czerkas also believed that the animal may have been able to fly better than

846: 637: 217: 158: 104: 48: 17: 4451: 4408: 4348: 4248:

Hone, D. W.; Dececchi, T. A.; Sullivan, C.; Xu, X.; Larsson, H. C. (2022).

4197: 4179: 4148: 4089: 4044:

Alexander, D.E.; Gong, E.; Martin, L.D.; Burnham, D.A.; Falk, A.R. (2010).

4030: 3967: 3902: 3843: 3720: 3653: 3556: 3466: 3407: 3282: 3209: 3149: 3069: 2942: 2880: 2820: 2762: 2680:." Message to the Dinosaur Mailing List, 1 Jan 2001. accessed 30 Sept 2014. 2529: 2478: 2429:

Alexander, D.E.; Gong, E.; Martin, L.D.; Burnham, D.A.; Falk, A.R. (2010).

1864:

had remex morphologies that are consistent with modern volant birds, while

1544: 944:

had not yet been named), have also been described by Hwang and colleagues.

932:. However, Senter also questioned this classification, noting that as with 751:

and now belong to the collection of the Paleontology Museum of Beipiao, in

732:, who had long proposed that dromaeosaurids evolved from flying ancestors. 2632:." Message to the Dinosaur Mailing List, 4 Jan 2001. accessed 23 Sep 2009. 2180:

had caught and consumed the bird in the trees, rather than scavenging it.

1150: 5438: 5311: 5232: 5203: 4977: 4787: 4587: 4493: 3095:

Hwang, S.H., Norell, M.A., Ji, Q., and Gao, K. (2002). "New Specimens of

1842:

could fly and yet is also very clearly a dromaeosaurid suggests that the

1805: 1789: 1758: 1099: 760: 756: 752: 572: 525: 428: 420: 191: 178: 94: 89: 74: 69: 59: 3893: 3232:"Biplane wing planform and flight performance of the feathered dinosaur 3141: 2754: 2046: 1199: 5486: 5334: 4620: 4121: 3816: 3644: 3619: 3388: 2812: 2252: 2233:

apparently swallowed its prey head first, a behavior typical of modern

1992: 1915: 1902: 1838:

Stephen Czerkas, Gregory S. Paul, and others have argued that the fact

1797: 1773:

may have been a powered flier, and indeed possibly a better flyer than

1653:

In a 2024 paper which reported the smallest known juvenile specimen of

1013: 909: 585: 520: 489:

provide important evidence about the evolutionary relationship between

306: 109: 84: 4211:

Beebe, C. W. A. (1915). "Tetrapteryx stage in the ancestry of birds".

2871: 2850: 2378: 509:

using all four limbs for lift. Subsequent studies have suggested that

4909: 4801: 4581: 2216: 2127: 1765:

Describing specimens originally referenced as a distinctive species (

1726:

this hypothesis as well, and found that the combined wing surface of

1250: 793:. However, BPM 1 3-13 has a longer tail, proportionately, than other 755:, China. They are referred to by the inventory numbers LPM 0200, the 748: 443: 424: 168: 5415: 4249: 3061: 3015: 1835:, as well as wing-assisted leaping and even ground-based launching. 3987:

specimen is not ideal for understanding the origin of avian flight"

3918:"Functional constraints on the number and shape of flight feathers" 3113:

Xing, X.; Zhou, Z.; Wang, X.; Kuang, X.; Zhang, F.; Du, X. (2003).

4532: 3509:"The extent of the preserved feathers on the four-winged dinosaur 2099: 2045: 2012: 1925: 1881: 1809: 1750: 1255: 1164: 1149: 1094: 987: 875: 765: 711: 681: 546: 432: 413: 136:

Fossil specimen, with white arrows pointing at preserved feathers

3176:"A basal dromaeosaurid and size evolution preceding avian flight" 5473: 2237: 1996: 1929: 1921: 1698: 1682: 782: 490: 447: 5419: 4574: 4536: 1075:

note that this is not particularly surprising, given that both

2244:

bones lacked marked pitting and scarring, indicating that the

2272:

specimens that preserve stomach contents. This suggests that

2187:

specimen. The authors contradicted the prior suggestion that

3486:

Historical Biology: An International Journal of Paleobiology

632:

and the latter, if valid, would have had priority under the

3480:

Wang, R.; Pei, R. (2024). "The smallest known specimen of

2225:) was described from a specimen found in the stomach of a 1886:

Wind tunnel experiments with different wing configurations

922:

Institute of Vertebrate Paleontology and Paleoanthropology

590:

Institute of Vertebrate Paleontology and Paleoanthropology

3507:

Hone, D.W.E.; Tischlinger, H.; Xu, X.; Zhang, F. (2010).

2647:"So why hasn't Tyrannosaurus been renamed Manospondylus?" 2168:, Jingmai O'Connor and coauthors described a specimen of 856:

have been uncovered, all from the Shangheshou Bed of the

616:

Since the two names designate the same individual as the

2383:

Student Showcase Journal, University of Alaska Anchorage

2287:

hunted at night. The discovery of iridescent plumage in

957:

has modern bird features that make it more derived than

666:(as it was improperly described) and the junior synonym 4305:

provides unique evidence of dinosaurs preying on birds"

3099:(Theropoda: Dromaeosauridae) from Northeastern China." 4228: 4226: 4162:

Sullivan, C.; Hone, D.W.E.; Xu, X.; Zhang, F. (2010).

3916:

Kiat, Yosef; O’Connor, Jingmai K. (20 February 2024).

4518:

Microraptor's glossy black feather coat reconstructed

4299:

Jingmai O'Connor; Zhonghe Zhou & Xing Xu (2011).

3370:

Dececchi, T.A.; Larsson, H.C.E.; Habib, M.B. (2016).

3365: 3363: 3361: 3359: 1932:, could have been used to control drag and alter the 4050:

Proceedings of the National Academy of Sciences, USA

3991:

Proceedings of the National Academy of Sciences, USA

2969:"Feathers Worth a 2nd Look Found on a Tiny Dinosaur" 2439:

Proceedings of the National Academy of Sciences, USA

332: 315: 285: 5428: 5242: 5156: 5121: 5059: 4987: 4876: 4834: 4771: 4716: 4689: 4662: 4635: 2424: 2422: 2420: 2209:being a specialist for hunting in either habitats. 1792:column until relatively advanced avialans like the 804:) have shown that the features used to distinguish 3346:Bulletin of the American Museum of Natural History 1823:Other studies have demonstrated that the wings of 3225: 3223: 3221: 3219: 3091: 3089: 3087: 2400: 2398: 2396: 1169:Specimen in the Shandong Tianyu Museum of Nature 1061:is particularly similar to the basal troodontid 3922:Proceedings of the National Academy of Sciences 3243:Proceedings of the National Academy of Sciences 3169: 3167: 3035: 3033: 2962: 2960: 2688: 2686: 2407:Bulletin of the Gunma Museum of Natural History 2192:associated with piscivory. They concluded that 2166:Proceedings of the National Academy of Sciences 1106:with coloration based on fossilized melanosomes 852:Numerous further specimens likely belonging to 3981:Brusatte, Stephen L.; Brougham, Jason (2010). 2904: 2902: 2900: 2898: 2712: 2710: 2708: 2706: 2704: 2702: 2542:Olson, S.L. (2000). "Countdown to Piltdown at 1087:split between dromaeosaurids and troodontids. 928:specimen, to be a new species, which he named 4548: 3588: 3586: 3568: 3566: 1846:, including later and larger species such as 634:International Code of Zoological Nomenclature 8: 2017:Restoration of two individuals by the ground 1697:were true flight feathers as seen in modern 992:Wingspan and body size compared with a human 893:Feathered Dinosaurs and the Origin of Flight 3597:. London: Dorling Kindersley. p. 332. 5416: 5065: 4779: 4768: 4695: 4668: 4641: 4632: 4571: 4555: 4541: 4533: 2913:and the Evolution of Iridescent Plumage". 800:Subsequent studies (and more specimens of 686:Skeletal restorations of various specimens 127: 31: 4494:Bird wings evolved from biplane dinosaurs 4390: 4338: 4328: 4243: 4241: 4239: 4187: 4138: 4120: 4079: 4069: 4020: 4010: 3957: 3892: 3882: 3833: 3815: 3710: 3692: 3643: 3546: 3536: 3456: 3446: 3397: 3387: 3272: 3262: 3199: 2870: 2844: 2842: 2840: 2838: 2519: 2509: 2468: 2458: 1769:), paleontologist Stephen Czerkas argued 789:, which was first described earlier than 2005:based on phylogenetic analyses in 2019. 5566:Multispecific non-avian dinosaur genera 3759:SpringerBriefs in Earth System Sciences 2369: 2342: 513:was capable of powered flight as well. 4488:/Cryptovolans pauli by Boban Filipovic 3593:Benson, R.B.J.; Brussatte, S. (2012). 3332: 3330: 3328: 3326: 3230:Chatterjee, S.; Templin, R.J. (2007). 2607:: CS1 maint: archived copy as title ( 2600: 1991:is not a dromaeosaur, but actually an 4360: 4358: 2851:"Piscivory in the feathered dinosaur 2061:The unique wing arrangement found in 1893:suggested in 2005 that, in order for 1820:was capable of true, powered flight. 1739:when gliding between low elevations. 1114:determined the plumage coloration of 1049:Several anatomical features found in 1029:(2003) compared the longer plumes on 7: 2967:Wilford, John Noble (8 March 2012). 2379:"A new phylogeny of Dromaeosauridae" 580:. After the forgery was revealed by 450:ago. Three species have been named ( 1812:, much like a "thumb-like" form of 1514: 1450: 1426: 1419: 1395: 1371: 1303: 1281: 1271: 1242: 1215: 1186: 1176: 1067:; in their 2002 description of two 908:definition of Aves, as a subset of 427:specimens have been recovered from 5541:Early Cretaceous dinosaurs of Asia 4258:Journal of Vertebrate Paleontology 4168:Proceedings of the Royal Society B 3115:"Four-winged dinosaurs from China" 2728:"Four-winged dinosaurs from China" 2492:Kiat, Y.; O'Connor, J. K. (2023). 2331:Timeline of dromaeosaurid research 2283:of the eye, it has been suggested 1665:within a new clade Serraraptoria. 1044:Distinguishing anatomical features 1001:smallest-known non-avian dinosaurs 598:National Museum of Natural History 25: 2623:Archaeoraptor still a nomen nudum 2164:In the December 6, 2011 issue of 772:Beijing Museum of Natural History 5400: 5399: 5376: 5370: 5365: 5359: 3296:Xu, Xing; Qin, Zi-Chuan (2017). 2696:, 416: pp. 36. 7 March 2002.> 2310: 2268:, despite the existence of four 2172:containing bones of an arboreal 1948:capacity to test whether or not 1565: 1543: 1462: 1333: 1313: 1254: 1225: 1198: 519:was among the most abundant non- 446:stage), 125 to 120 million 145: 2849:Lida Xing; et al. (2013). 1713:It was originally thought that 1689:aircraft of today. It had long 690:The first specimen referred to 650:, in which the former became a 555:" fossil; the tail belongs to 1: 5556:Fossil taxa described in 2000 4278:10.1080/02724634.2022.2144337 3580:(6921): 335-340, 23 Jan 2003. 3494:10.1080/08912963.2024.2385604 1924:structure, like those on the 1874:were secondarily flightless. 1833:wing-assisted incline running 860:in Liaoning, China. In fact, 3694:10.1371/journal.pone.0022292 3538:10.1371/journal.pone.0009223 3448:10.1371/journal.pone.0036790 2783:Holtz, Thomas R. Jr. (2011) 1905:) and not overlaid (as on a 1124:scanning electron microscope 485:, well-preserved fossils of 2671:Archaeoraptor v Microraptor 630:Archaeoraptor liaoningensis 606:Archaeoraptor liaoningensis 435:. They date from the early 5582: 3757:and Scansoriopterygidae". 2511:10.1038/s42003-023-05048-x 2264:has never been found with 1110:In March 2012, Quanguo Li 702:et al. named the specimen 596:, curator of birds in the 540: 423:. Numerous well-preserved 27:Extinct genus of dinosaurs 5389: 5357: 5068: 4782: 4767: 4748: 4698: 4671: 4644: 4631: 4570: 4392:10.1016/j.cub.2019.06.020 3884:10.1016/j.cub.2020.06.105 3767:10.1007/978-94-007-5637-3 3101:American Museum Novitates 2279:Based on the size of the 1558: 1536: 1519: 1512: 1477: 1455: 1448: 1431: 1424: 1417: 1400: 1393: 1376: 1369: 1328: 1308: 1301: 1286: 1279: 1269: 1247: 1240: 1220: 1213: 1191: 1184: 1012:were among the first non- 813:species, making the name 412:: "one who seizes") is a 359: 352: 312: 304: 284: 277: 142:Scientific classification 140: 135: 126: 34: 4507:The Four-Winged Dinosaur 4301:"Additional specimen of 3513:under ultraviolet light" 2212:In 2019, a new genus of 1657:, Wang and Pei included 1158:(1) compared with other 747:, dating from the early 743:were collected from the 626:junior objective synonym 4478:Relationship with birds 4444:10.1126/science.1200043 4330:10.1073/pnas.1117727108 4071:10.1073/pnas.1004977107 4012:10.1073/pnas.1004977107 3942:10.1073/pnas.2306639121 3264:10.1073/pnas.0609975104 3201:10.1126/science.1144066 2935:10.1126/science.1213780 2546:: the rise and fall of 2460:10.1073/pnas.0911852107 2057:" with four wings, 1915 1913:most likely employed a 1036:s head to those of the 1003:, with the holotype of 895:. In his discussion of 602:Smithsonian Institution 493:and earlier dinosaurs. 4509:- PBS website for the 4264:(e2144337). e2144337. 4180:10.1098/rspb.2009.2281 3305:Vertebrata PalAsiatica 2669:Williams, T. (2002). " 2621:Creisler, B. (2002). " 2498:Communications Biology 2105: 2058: 2018: 1887: 1762: 1170: 1162: 1107: 993: 881: 774: 687: 562:The initial naming of 559: 416:of small, four-winged 5508:Paleobiology Database 4252:Microraptor zhaoianus 3624:Nature Communications 3097:Microraptor zhaoianus 2801:Journal of Morphology 2788:Winter 2010 Appendix. 2377:Miller, Z.M. (2004). 2103: 2049: 2016: 1885: 1800:theropods, including 1754: 1481:Microraptor zhaoianus 1168: 1153: 1098: 991: 879: 823:Microraptor zhaoianus 769: 685: 622:Microraptor zhaoianus 611:Microraptor zhaoianus 550: 290:Microraptor zhaoianus 4377:(14): 2423–2429.e2. 4250:"Generalist diet of 3869:(20): 4033–4046.e8. 3736:Acta Palaeontol. Pol 2575:(December): 705-708. 1816:, may indicate that 1779:. He noted that the 678:Additional specimens 5551:Feathered dinosaurs 5123:Saurornitholestinae 4436:2011Sci...332..705S 4383:2019CBio...29E2423O 4321:2011PNAS..10819662O 4315:(49): 19662–19665. 4270:2022JVPal..42E4337H 4174:(1690): 2027–2033. 4062:2010PNAS..107E.155B 4003:2010PNAS..107E.155B 3934:2024PNAS..12106639K 3875:2020CBio...30E4033P 3685:2011PLoSO...622292D 3636:2013NatCo...4.2489D 3529:2010PLoSO...5.9223H 3439:2012PLoSO...736790S 3255:2007PNAS..104.1576C 3192:2007Sci...317.1378T 3186:(5843): 1378–1381. 3142:10.1038/nature01342 3134:2003Natur.421..335X 3054:2002Natur.416...36N 3008:2006GeolJ..41..419X 2927:2012Sci...335.1215L 2921:(6073): 1215–1219. 2755:10.1038/nature01342 2747:2003Natur.421..335X 2544:National Geographic 2451:2010PNAS..107.2972A 2326:Dinosaur coloration 1691:pennaceous feathers 1289:Saurornitholestinae 858:Jiufotang Formation 821:junior synonyms of 745:Jiufotang Formation 648:Manospondylus gigas 499:pennaceous feathers 440:Jiufotang Formation 4499:2007-01-27 at the 4122:10.7717/peerj.7247 3817:10.7717/peerj.7247 3645:10.1038/ncomms3489 3389:10.7717/peerj.2159 2996:Geological Journal 2973:The New York Times 2813:10.1002/jmor.10382 2676:2016-03-03 at the 2628:2008-07-25 at the 2251:Unlike its fellow 2121:, and most of the 2106: 2104:Close-up of a cast 2059: 2019: 1888: 1767:Cryptovolans pauli 1763: 1171: 1163: 1138:in the plumage of 1108: 994: 882: 836:A new specimen of 815:Cryptovolans pauli 775: 704:Cryptovolans pauli 688: 624:would have been a 560: 537:Naming controversy 5523: 5522: 5495:Open Tree of Life 5422:Taxon identifiers 5413: 5412: 5385: 5384: 5355: 5354: 5351: 5350: 5320:Saurornitholestes 5211:Saurornitholestes 5147:Saurornitholestes 4836:Halszkaraptorinae 4763: 4762: 4759: 4758: 4744: 4743: 4740: 4739: 4736: 4735: 4476:Dromaeosauridae, 4254:included mammals" 3776:978-94-007-5636-6 3604:978-0-7566-9910-9 3128:(6921): 335–340. 2872:10.1111/evo.12119 2741:(6921): 335–340. 2558:(2) (April): 1–3. 2240:and lizards. The 2053:'s hypothetical " 1891:Sankar Chatterjee 1814:leading edge slot 1663:eudromaeosaurians 1650: 1649: 1641: 1640: 1632: 1631: 1623: 1622: 1614: 1613: 1605: 1604: 1596: 1595: 1587: 1586: 1578: 1577: 1501: 1500: 1492: 1491: 1355: 1354: 1346: 1345: 1085:deinonychosaurian 1071:specimens, Hwang 654:and the latter a 643:Tyrannosaurus rex 389: 388: 383: 374:"Tetrapterornis" 371: 346: 329: 273: 16:(Redirected from 5573: 5516: 5515: 5503: 5502: 5490: 5489: 5477: 5476: 5464: 5463: 5462: 5449: 5448: 5447: 5417: 5403: 5402: 5380: 5374: 5369: 5363: 5244:Velociraptorinae 5190:Dromaeosauroides 5066: 5050:Zhongjianosaurus 5028:Sinornithosaurus 4780: 4769: 4696: 4669: 4642: 4633: 4626: 4625: 4572: 4557: 4550: 4543: 4534: 4464: 4463: 4419: 4413: 4412: 4394: 4362: 4353: 4352: 4342: 4332: 4296: 4290: 4289: 4245: 4234: 4230: 4221: 4220: 4208: 4202: 4201: 4191: 4159: 4153: 4152: 4142: 4124: 4100: 4094: 4093: 4083: 4073: 4041: 4035: 4034: 4024: 4014: 3978: 3972: 3971: 3961: 3913: 3907: 3906: 3896: 3886: 3854: 3848: 3847: 3837: 3819: 3795: 3789: 3788: 3750: 3744: 3743: 3731: 3725: 3724: 3714: 3696: 3664: 3658: 3657: 3647: 3615: 3609: 3608: 3595:Prehistoric Life 3590: 3581: 3570: 3561: 3560: 3550: 3540: 3504: 3498: 3497: 3477: 3471: 3470: 3460: 3450: 3418: 3412: 3411: 3401: 3391: 3367: 3354: 3353: 3343: 3334: 3321: 3320: 3302: 3293: 3287: 3286: 3276: 3266: 3249:(5): 1576–1580. 3240: 3227: 3214: 3213: 3203: 3171: 3162: 3161: 3119: 3110: 3104: 3093: 3082: 3081: 3037: 3028: 3027: 3002:(3–4): 419–437. 2991: 2985: 2984: 2982: 2980: 2964: 2955: 2954: 2906: 2893: 2892: 2874: 2865:(8): 2441–2445. 2846: 2833: 2832: 2796: 2790: 2781: 2775: 2774: 2732: 2723: 2717: 2714: 2697: 2690: 2681: 2667: 2661: 2660: 2658: 2657: 2651:The Dinosaur FAQ 2639: 2633: 2619: 2613: 2612: 2606: 2598: 2596: 2595: 2589: 2583:. Archived from 2582: 2565: 2559: 2540: 2534: 2533: 2523: 2513: 2489: 2483: 2482: 2472: 2462: 2445:(7): 2972–2976. 2426: 2415: 2414: 2402: 2391: 2390: 2374: 2358: 2347: 2320: 2318:Dinosaurs portal 2315: 2314: 2313: 1995:close to modern 1878:Hindwing posture 1674:Wings and flight 1569: 1562:Sinornithosaurus 1547: 1515: 1466: 1451: 1427: 1420: 1396: 1372: 1337: 1317: 1311:Velociraptorinae 1304: 1282: 1272: 1258: 1243: 1229: 1216: 1202: 1187: 1177: 1134:, and indicated 1038:Philippine eagle 1035: 872:Study and debate 770:Specimen at the 717:, "flying". The 710:, "hidden", and 375: 365: 340: 334: 323: 317: 287: 265: 258: 245: 229: 150: 149: 131: 121: 51: 42:Early Cretaceous 40:Temporal range: 32: 21: 5581: 5580: 5576: 5575: 5574: 5572: 5571: 5570: 5546:Microraptorians 5536:Jiufotang fauna 5526: 5525: 5524: 5519: 5511: 5506: 5498: 5493: 5485: 5480: 5472: 5467: 5458: 5457: 5452: 5443: 5442: 5437: 5424: 5414: 5409: 5381: 5347: 5304:Luanchuanraptor 5238: 5158:Dromaeosaurinae 5152: 5117: 5061:Eudromaeosauria 5055: 4983: 4872: 4830: 4776: 4774:Dromaeosauridae 4755: 4752:Dromaeosauridae 4732: 4718:Dromaeosauridae 4712: 4685: 4658: 4652:Avemetatarsalia 4637:Avemetatarsalia 4627: 4577: 4576: 4566: 4564:Dromaeosauridae 4561: 4503:COSMOS magazine 4501:Wayback Machine 4472: 4467: 4430:(6030): 705–8. 4421: 4420: 4416: 4371:Current Biology 4364: 4363: 4356: 4298: 4297: 4293: 4247: 4246: 4237: 4231: 4224: 4210: 4209: 4205: 4161: 4160: 4156: 4102: 4101: 4097: 4043: 4042: 4038: 3980: 3979: 3975: 3915: 3914: 3910: 3863:Current Biology 3856: 3855: 3851: 3797: 3796: 3792: 3777: 3752: 3751: 3747: 3733: 3732: 3728: 3666: 3665: 3661: 3617: 3616: 3612: 3605: 3592: 3591: 3584: 3571: 3564: 3511:Microraptor gui 3506: 3505: 3501: 3479: 3478: 3474: 3420: 3419: 3415: 3369: 3368: 3357: 3341: 3336: 3335: 3324: 3300: 3295: 3294: 3290: 3238: 3234:Microraptor gui 3229: 3228: 3217: 3173: 3172: 3165: 3117: 3112: 3111: 3107: 3094: 3085: 3062:10.1038/416036a 3039: 3038: 3031: 3016:10.1002/gj.1044 2993: 2992: 2988: 2978: 2976: 2966: 2965: 2958: 2908: 2907: 2896: 2848: 2847: 2836: 2798: 2797: 2793: 2782: 2778: 2730: 2725: 2724: 2720: 2715: 2700: 2691: 2684: 2678:Wayback Machine 2668: 2664: 2655: 2653: 2641: 2640: 2636: 2630:Wayback Machine 2620: 2616: 2599: 2593: 2591: 2587: 2580: 2578:"Archived copy" 2576: 2566: 2562: 2541: 2537: 2491: 2490: 2486: 2433:Microraptor gui 2428: 2427: 2418: 2404: 2403: 2394: 2376: 2375: 2371: 2367: 2362: 2361: 2348: 2344: 2339: 2316: 2311: 2309: 2306: 2266:gastric pellets 2174:enantiornithean 2170:Microraptor gui 2158:Sinosauropteryx 2098: 2044: 2011: 2009:Ground movement 1880: 1844:Dromaeosauridae 1831:was capable of 1707:flight feathers 1676: 1671: 1659:microraptorians 1651: 1642: 1633: 1624: 1615: 1606: 1597: 1588: 1579: 1502: 1493: 1459:Microraptor gui 1435:Microraptor sp. 1356: 1347: 1331:Dromaeosaurinae 1276:Eudromaeosauria 1181:Dromaeosauridae 1148: 1093: 1046: 1033: 986: 963:Dromaeosauridae 930:Microraptor gui 880:Fossil specimen 874: 819:Microraptor gui 730:Gregory S. Paul 700:Stephen Czerkas 680: 672:nomen protectum 652:nomen protectum 594:Storrs L. Olson 545: 539: 534: 300: 293: 264: 256: 243: 231:Dromaeosauridae 227: 144: 122: 120: 119: 118: 117: 112: 107: 102: 97: 92: 87: 82: 77: 72: 67: 62: 57: 46: 45: 38: 28: 23: 22: 15: 12: 11: 5: 5579: 5577: 5569: 5568: 5563: 5558: 5553: 5548: 5543: 5538: 5528: 5527: 5521: 5520: 5518: 5517: 5504: 5491: 5478: 5465: 5450: 5434: 5432: 5426: 5425: 5420: 5411: 5410: 5408: 5407: 5397: 5390: 5387: 5386: 5383: 5382: 5358: 5356: 5353: 5352: 5349: 5348: 5346: 5345: 5338: 5331: 5324: 5316: 5308: 5300: 5293: 5286: 5279: 5271: 5263: 5256: 5248: 5246: 5240: 5239: 5237: 5236: 5229: 5222: 5215: 5207: 5200: 5193: 5186: 5178: 5170: 5162: 5160: 5154: 5153: 5151: 5150: 5143: 5135: 5127: 5125: 5119: 5118: 5116: 5115: 5107: 5100: 5092: 5085: 5078: 5069: 5063: 5057: 5056: 5054: 5053: 5046: 5039: 5031: 5024: 5017: 5009: 5002: 4994: 4992: 4985: 4984: 4982: 4981: 4974: 4966: 4962:Unquillosaurus 4958: 4951: 4943: 4935: 4928: 4920: 4913: 4906: 4898: 4891: 4883: 4881: 4874: 4873: 4871: 4870: 4863: 4856: 4849: 4841: 4839: 4832: 4831: 4829: 4828: 4821: 4813: 4806: 4798: 4791: 4783: 4777: 4772: 4765: 4764: 4761: 4760: 4757: 4756: 4749: 4746: 4745: 4742: 4741: 4738: 4737: 4734: 4733: 4731: 4730: 4729: 4728: 4722: 4720: 4714: 4713: 4711: 4710: 4709: 4708: 4699: 4693: 4687: 4686: 4684: 4683: 4682: 4681: 4672: 4666: 4660: 4659: 4657: 4656: 4655: 4654: 4645: 4639: 4629: 4628: 4624: 4623: 4614: 4605: 4596: 4590: 4584: 4575: 4568: 4567: 4562: 4560: 4559: 4552: 4545: 4537: 4531: 4530: 4525: 4520: 4514: 4504: 4492:Jacqui Hayes: 4490: 4481: 4471: 4470:External links 4468: 4466: 4465: 4414: 4354: 4291: 4235: 4222: 4203: 4154: 4095: 4036: 3973: 3908: 3849: 3790: 3775: 3745: 3726: 3659: 3610: 3603: 3582: 3562: 3499: 3472: 3413: 3355: 3322: 3311:(2): 129–144. 3288: 3215: 3163: 3105: 3083: 3048:(6876): 36–7. 3029: 2986: 2956: 2894: 2834: 2807:(2): 125–166. 2791: 2776: 2718: 2698: 2682: 2662: 2645:(2002-08-27). 2634: 2614: 2560: 2535: 2484: 2416: 2392: 2368: 2366: 2363: 2360: 2359: 2341: 2340: 2338: 2335: 2334: 2333: 2328: 2322: 2321: 2305: 2302: 2097: 2094: 2043: 2040: 2010: 2007: 1879: 1876: 1866:anchiornithids 1794:enantiornithes 1675: 1672: 1670: 1667: 1648: 1647: 1644: 1643: 1639: 1638: 1635: 1634: 1630: 1629: 1626: 1625: 1621: 1620: 1617: 1616: 1612: 1611: 1608: 1607: 1603: 1602: 1599: 1598: 1594: 1593: 1590: 1589: 1585: 1584: 1581: 1580: 1576: 1575: 1572: 1571: 1557: 1554: 1553: 1550: 1549: 1535: 1532: 1531: 1528: 1527: 1518: 1513: 1511: 1508: 1507: 1504: 1503: 1499: 1498: 1495: 1494: 1490: 1489: 1486: 1485: 1476: 1473: 1472: 1469: 1468: 1454: 1449: 1447: 1444: 1443: 1440: 1439: 1430: 1425: 1423: 1418: 1416: 1413: 1412: 1409: 1408: 1399: 1394: 1392: 1389: 1388: 1385: 1384: 1375: 1370: 1368: 1362: 1361: 1358: 1357: 1353: 1352: 1349: 1348: 1344: 1343: 1340: 1339: 1327: 1324: 1323: 1320: 1319: 1307: 1302: 1300: 1297: 1296: 1293: 1292: 1285: 1280: 1278: 1270: 1268: 1265: 1264: 1261: 1260: 1246: 1241: 1239: 1236: 1235: 1232: 1231: 1219: 1214: 1212: 1209: 1208: 1205: 1204: 1190: 1185: 1183: 1175: 1147: 1146:Classification 1144: 1092: 1089: 1045: 1042: 999:was among the 985: 982: 873: 870: 737:type specimens 727:paleontologist 679: 676: 541:Main article: 538: 535: 533: 530: 387: 386: 385: 384: 376:Miller, 2004 ( 372: 357: 356: 350: 349: 348: 347: 330: 310: 309: 302: 301: 294: 282: 281: 275: 274: 254: 250: 249: 241: 234: 233: 225: 221: 220: 215: 208: 207: 202: 195: 194: 189: 182: 181: 176: 172: 171: 166: 162: 161: 156: 152: 151: 138: 137: 133: 132: 124: 123: 115: 114: 113: 108: 103: 98: 93: 88: 83: 78: 73: 68: 63: 58: 53: 52: 39: 26: 24: 14: 13: 10: 9: 6: 4: 3: 2: 5578: 5567: 5564: 5562: 5561:Animal flight 5559: 5557: 5554: 5552: 5549: 5547: 5544: 5542: 5539: 5537: 5534: 5533: 5531: 5514: 5509: 5505: 5501: 5496: 5492: 5488: 5483: 5479: 5475: 5470: 5466: 5461: 5455: 5451: 5446: 5440: 5436: 5435: 5433: 5431: 5427: 5423: 5418: 5406: 5398: 5396: 5392: 5391: 5388: 5379: 5375: 5373: 5368: 5362: 5344: 5343: 5339: 5337: 5336: 5332: 5330: 5329: 5325: 5322: 5321: 5317: 5314: 5313: 5309: 5306: 5305: 5301: 5299: 5298: 5294: 5292: 5291: 5287: 5285: 5284: 5283:Kansaignathus 5280: 5277: 5276: 5272: 5269: 5268: 5264: 5262: 5261: 5257: 5255: 5254: 5250: 5249: 5247: 5245: 5241: 5235: 5234: 5230: 5228: 5227: 5223: 5221: 5220: 5216: 5213: 5212: 5208: 5206: 5205: 5201: 5199: 5198: 5197:Dromaeosaurus 5194: 5192: 5191: 5187: 5184: 5183: 5179: 5176: 5175: 5171: 5169: 5168: 5164: 5163: 5161: 5159: 5155: 5149: 5148: 5144: 5141: 5140: 5136: 5134: 5133: 5129: 5128: 5126: 5124: 5120: 5113: 5112: 5108: 5106: 5105: 5101: 5098: 5097: 5093: 5091: 5090: 5089:Dineobellator 5086: 5084: 5083: 5079: 5076: 5075: 5071: 5070: 5067: 5064: 5062: 5058: 5052: 5051: 5047: 5045: 5044: 5040: 5037: 5036: 5032: 5030: 5029: 5025: 5023: 5022: 5018: 5015: 5014: 5013:Hesperonychus 5010: 5008: 5007: 5006:Graciliraptor 5003: 5001: 5000: 4999:Changyuraptor 4996: 4995: 4993: 4990: 4989:Microraptoria 4986: 4980: 4979: 4975: 4972: 4971: 4967: 4964: 4963: 4959: 4957: 4956: 4952: 4949: 4948: 4944: 4941: 4940: 4936: 4934: 4933: 4929: 4926: 4925: 4924:Ornithodesmus 4921: 4919: 4918: 4917:Neuquenraptor 4914: 4912: 4911: 4907: 4904: 4903: 4899: 4897: 4896: 4892: 4890: 4889: 4885: 4884: 4882: 4879: 4875: 4869: 4868: 4864: 4862: 4861: 4857: 4855: 4854: 4850: 4848: 4847: 4846:Halszkaraptor 4843: 4842: 4840: 4837: 4833: 4827: 4826: 4822: 4819: 4818: 4814: 4812: 4811: 4807: 4804: 4803: 4799: 4797: 4796: 4792: 4790: 4789: 4785: 4784: 4781: 4778: 4775: 4770: 4766: 4754: 4753: 4747: 4726: 4725: 4724: 4723: 4721: 4719: 4715: 4707: 4703: 4702: 4701: 4700: 4697: 4694: 4692: 4688: 4680: 4676: 4675: 4674: 4673: 4670: 4667: 4665: 4661: 4653: 4649: 4648: 4647: 4646: 4643: 4640: 4638: 4634: 4630: 4622: 4618: 4615: 4613: 4609: 4606: 4604: 4600: 4597: 4595: 4591: 4589: 4585: 4583: 4579: 4578: 4573: 4569: 4565: 4558: 4553: 4551: 4546: 4544: 4539: 4538: 4535: 4529: 4526: 4524: 4521: 4519: 4516:NewScientist 4515: 4512: 4508: 4505: 4502: 4498: 4495: 4491: 4489: 4487: 4482: 4480: 4479: 4474: 4473: 4469: 4461: 4457: 4453: 4449: 4445: 4441: 4437: 4433: 4429: 4425: 4418: 4415: 4410: 4406: 4402: 4398: 4393: 4388: 4384: 4380: 4376: 4372: 4368: 4361: 4359: 4355: 4350: 4346: 4341: 4336: 4331: 4326: 4322: 4318: 4314: 4310: 4306: 4304: 4295: 4292: 4287: 4283: 4279: 4275: 4271: 4267: 4263: 4259: 4255: 4253: 4244: 4242: 4240: 4236: 4229: 4227: 4223: 4218: 4214: 4207: 4204: 4199: 4195: 4190: 4185: 4181: 4177: 4173: 4169: 4165: 4158: 4155: 4150: 4146: 4141: 4136: 4132: 4128: 4123: 4118: 4114: 4110: 4106: 4099: 4096: 4091: 4087: 4082: 4077: 4072: 4067: 4063: 4059: 4055: 4051: 4047: 4040: 4037: 4032: 4028: 4023: 4018: 4013: 4008: 4004: 4000: 3996: 3992: 3988: 3986: 3977: 3974: 3969: 3965: 3960: 3955: 3951: 3947: 3943: 3939: 3935: 3931: 3927: 3923: 3919: 3912: 3909: 3904: 3900: 3895: 3890: 3885: 3880: 3876: 3872: 3868: 3864: 3860: 3853: 3850: 3845: 3841: 3836: 3831: 3827: 3823: 3818: 3813: 3809: 3805: 3801: 3794: 3791: 3786: 3782: 3778: 3772: 3768: 3764: 3760: 3756: 3749: 3746: 3741: 3737: 3730: 3727: 3722: 3718: 3713: 3708: 3704: 3700: 3695: 3690: 3686: 3682: 3679:(8): e22292. 3678: 3674: 3670: 3663: 3660: 3655: 3651: 3646: 3641: 3637: 3633: 3629: 3625: 3621: 3614: 3611: 3606: 3600: 3596: 3589: 3587: 3583: 3579: 3575: 3569: 3567: 3563: 3558: 3554: 3549: 3544: 3539: 3534: 3530: 3526: 3522: 3518: 3514: 3512: 3503: 3500: 3495: 3491: 3487: 3483: 3476: 3473: 3468: 3464: 3459: 3454: 3449: 3444: 3440: 3436: 3433:(5): e36790. 3432: 3428: 3424: 3417: 3414: 3409: 3405: 3400: 3395: 3390: 3385: 3381: 3377: 3373: 3366: 3364: 3362: 3360: 3356: 3351: 3347: 3340: 3333: 3331: 3329: 3327: 3323: 3318: 3314: 3310: 3306: 3299: 3292: 3289: 3284: 3280: 3275: 3270: 3265: 3260: 3256: 3252: 3248: 3244: 3237: 3235: 3226: 3224: 3222: 3220: 3216: 3211: 3207: 3202: 3197: 3193: 3189: 3185: 3181: 3177: 3170: 3168: 3164: 3159: 3155: 3151: 3147: 3143: 3139: 3135: 3131: 3127: 3123: 3116: 3109: 3106: 3103:, 3381: 44pp. 3102: 3098: 3092: 3090: 3088: 3084: 3079: 3075: 3071: 3067: 3063: 3059: 3055: 3051: 3047: 3043: 3036: 3034: 3030: 3025: 3021: 3017: 3013: 3009: 3005: 3001: 2997: 2990: 2987: 2974: 2970: 2963: 2961: 2957: 2952: 2948: 2944: 2940: 2936: 2932: 2928: 2924: 2920: 2916: 2912: 2905: 2903: 2901: 2899: 2895: 2890: 2886: 2882: 2878: 2873: 2868: 2864: 2860: 2856: 2854: 2845: 2843: 2841: 2839: 2835: 2830: 2826: 2822: 2818: 2814: 2810: 2806: 2802: 2795: 2792: 2789: 2786: 2780: 2777: 2772: 2768: 2764: 2760: 2756: 2752: 2748: 2744: 2740: 2736: 2729: 2722: 2719: 2713: 2711: 2709: 2707: 2705: 2703: 2699: 2695: 2689: 2687: 2683: 2679: 2675: 2672: 2666: 2663: 2652: 2648: 2644: 2638: 2635: 2631: 2627: 2624: 2618: 2615: 2610: 2604: 2590:on 2008-12-17 2586: 2579: 2574: 2570: 2564: 2561: 2557: 2553: 2549: 2548:Archaeoraptor 2545: 2539: 2536: 2531: 2527: 2522: 2517: 2512: 2507: 2503: 2499: 2495: 2488: 2485: 2480: 2476: 2471: 2466: 2461: 2456: 2452: 2448: 2444: 2440: 2436: 2434: 2425: 2423: 2421: 2417: 2412: 2408: 2401: 2399: 2397: 2393: 2388: 2384: 2380: 2373: 2370: 2364: 2356: 2352: 2346: 2343: 2336: 2332: 2329: 2327: 2324: 2323: 2319: 2308: 2303: 2301: 2299: 2295: 2290: 2286: 2282: 2277: 2275: 2271: 2267: 2263: 2259: 2258: 2254: 2249: 2247: 2243: 2239: 2236: 2232: 2228: 2224: 2223: 2218: 2215: 2214:scleroglossan 2210: 2208: 2203: 2197: 2195: 2190: 2186: 2181: 2179: 2175: 2171: 2167: 2162: 2160: 2159: 2155: 2154:compsognathid 2151: 2146: 2144: 2140: 2136: 2135: 2130: 2129: 2124: 2120: 2116: 2111: 2102: 2095: 2093: 2091: 2090: 2085: 2084: 2083:Archaeopteryx 2079: 2075: 2074:William Beebe 2072: 2068: 2064: 2056: 2052: 2051:William Beebe 2048: 2041: 2039: 2037: 2033: 2028: 2024: 2015: 2008: 2006: 2004: 2003: 2002:Balaur bondoc 1998: 1994: 1990: 1989:Hesperonychus 1986: 1985: 1984:Hesperonychus 1979: 1975: 1971: 1967: 1962: 1957: 1955: 1951: 1947: 1946:animal flight 1943: 1939: 1935: 1931: 1927: 1923: 1918: 1917: 1912: 1908: 1904: 1900: 1896: 1892: 1884: 1877: 1875: 1873: 1872: 1867: 1863: 1859: 1855: 1851: 1850: 1845: 1841: 1836: 1834: 1830: 1826: 1821: 1819: 1815: 1811: 1807: 1803: 1799: 1795: 1791: 1786: 1782: 1781:Microraptor's 1778: 1777: 1776:Archaeopteryx 1772: 1768: 1760: 1757: 1753: 1749: 1746: 1742: 1738: 1733: 1729: 1724: 1720: 1716: 1711: 1708: 1704: 1700: 1696: 1692: 1688: 1684: 1680: 1673: 1668: 1666: 1664: 1660: 1656: 1646: 1645: 1637: 1636: 1628: 1627: 1619: 1618: 1610: 1609: 1601: 1600: 1592: 1591: 1583: 1582: 1574: 1573: 1570: 1568: 1564: 1563: 1556: 1555: 1552: 1551: 1548: 1546: 1542: 1541: 1540:Graciliraptor 1534: 1533: 1530: 1529: 1526: 1525: 1524: 1517: 1516: 1510: 1509: 1506: 1505: 1497: 1496: 1488: 1487: 1484: 1483: 1482: 1475: 1474: 1471: 1470: 1467: 1465: 1461: 1460: 1453: 1452: 1446: 1445: 1442: 1441: 1438: 1437: 1436: 1429: 1428: 1422: 1421: 1415: 1414: 1411: 1410: 1407: 1406: 1405: 1404:Hesperonychus 1398: 1397: 1391: 1390: 1387: 1386: 1383: 1382: 1381: 1374: 1373: 1367: 1366:Microraptoria 1364: 1363: 1360: 1359: 1351: 1350: 1342: 1341: 1338: 1336: 1332: 1326: 1325: 1322: 1321: 1318: 1316: 1312: 1306: 1305: 1299: 1298: 1295: 1294: 1291: 1290: 1284: 1283: 1277: 1274: 1273: 1267: 1266: 1263: 1262: 1259: 1257: 1253: 1252: 1245: 1244: 1238: 1237: 1234: 1233: 1230: 1228: 1224: 1218: 1217: 1211: 1210: 1207: 1206: 1203: 1201: 1197: 1196: 1189: 1188: 1182: 1179: 1178: 1174: 1167: 1161: 1157: 1152: 1145: 1143: 1141: 1137: 1133: 1129: 1125: 1121: 1117: 1113: 1105: 1101: 1097: 1090: 1088: 1086: 1082: 1078: 1074: 1070: 1066: 1065: 1060: 1056: 1052: 1043: 1041: 1039: 1032: 1028: 1023: 1019: 1015: 1011: 1006: 1002: 998: 990: 983: 981: 979: 975: 970: 969: 964: 960: 959:Archaeopteryx 956: 952: 951: 950:Archaeopteryx 945: 943: 939: 935: 931: 927: 923: 918: 913: 911: 907: 903: 898: 894: 889: 887: 878: 871: 869: 867: 863: 859: 855: 850: 848: 843: 839: 834: 832: 828: 824: 820: 816: 812: 807: 803: 798: 796: 792: 788: 784: 780: 773: 768: 764: 762: 758: 754: 750: 746: 742: 738: 733: 731: 728: 724: 720: 719:specific name 716: 713: 709: 705: 701: 697: 693: 684: 677: 675: 673: 669: 665: 661: 657: 656:nomen oblitum 653: 649: 645: 644: 639: 635: 631: 627: 623: 619: 614: 612: 607: 603: 599: 595: 591: 587: 583: 579: 575: 574: 569: 565: 558: 554: 553:Archaeoraptor 549: 544: 543:Archaeoraptor 536: 531: 529: 527: 522: 518: 514: 512: 508: 504: 500: 496: 492: 488: 484: 483: 482:Archaeopteryx 477: 475: 471: 470: 465: 461: 457: 453: 449: 445: 441: 438: 434: 430: 426: 422: 419: 418:dromaeosaurid 415: 411: 407: 403: 399: 395: 394: 381: 380: 373: 369: 364: 361: 360: 358: 355: 351: 344: 339: 338: 331: 327: 322: 321: 314: 313: 311: 308: 303: 298: 292: 291: 283: 280: 276: 271: 268: 263: 262: 255: 252: 251: 248: 247:Microraptoria 242: 239: 236: 235: 232: 226: 223: 222: 219: 216: 213: 210: 209: 206: 203: 200: 197: 196: 193: 190: 187: 184: 183: 180: 177: 174: 173: 170: 167: 164: 163: 160: 157: 154: 153: 148: 143: 139: 134: 130: 125: 111: 106: 101: 96: 91: 86: 81: 76: 71: 66: 61: 56: 50: 43: 37: 33: 30: 19: 5429: 5364: 5342:Velociraptor 5340: 5333: 5326: 5318: 5310: 5302: 5295: 5288: 5281: 5273: 5265: 5258: 5253:Acheroraptor 5251: 5231: 5224: 5217: 5209: 5202: 5195: 5188: 5180: 5174:Dakotaraptor 5172: 5167:Achillobator 5165: 5145: 5137: 5132:Atrociraptor 5130: 5111:Zhenyuanlong 5109: 5102: 5096:Tianyuraptor 5094: 5087: 5080: 5072: 5048: 5041: 5035:Tianyuraptor 5033: 5026: 5020: 5019: 5011: 5004: 4997: 4976: 4968: 4960: 4953: 4945: 4937: 4930: 4922: 4915: 4908: 4902:Dakotaraptor 4900: 4895:Buitreraptor 4893: 4888:Austroraptor 4886: 4878:Unenlagiinae 4865: 4858: 4851: 4844: 4825:Zhenyuanlong 4823: 4815: 4810:Tianyuraptor 4808: 4800: 4793: 4786: 4750: 4616: 4607: 4598: 4510: 4485: 4477: 4427: 4423: 4417: 4374: 4370: 4312: 4308: 4302: 4294: 4261: 4257: 4251: 4216: 4212: 4206: 4171: 4167: 4157: 4112: 4108: 4098: 4056:(40): E155. 4053: 4049: 4039: 3997:(40): E155. 3994: 3990: 3984: 3976: 3925: 3921: 3911: 3894:11336/143103 3866: 3862: 3852: 3807: 3803: 3793: 3758: 3754: 3748: 3739: 3735: 3729: 3676: 3672: 3662: 3627: 3623: 3613: 3594: 3577: 3573: 3523:(2): e9223. 3520: 3516: 3510: 3502: 3485: 3481: 3475: 3430: 3426: 3416: 3379: 3375: 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