349:
The progression from wing-assisted incline running to flight can be seen in the growth of birds, from when they are hatchlings to fully grown. They begin with wing-assisted incline running and slowly alter their wing strokes for flight as they grow and are able to make enough force. These transitional stages that lead to flight are both physical and behavioral. The transitions over a hatchling's life can be correlated with the evolution of flight on a macro scale. If protobirds are compared to hatchlings their physical traits such as wing size and behavior may have been similar. Flapping flight is limited by the size and muscle force of a wing. Even while using the correct model of arboreal or cursorial, protobirds' wings were not able to sustain flight, but they did most likely gain the behaviors needed for the arboreal or cursorial model like today's birds do when hatched. There are similar steps between the two. Wing-assisted incline running can also produce a useful lift in babies but is very small compared to that of juveniles and adult birds. This lift was found responsible for body acceleration when going up an incline and leads to flight as the bird grows.
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toward a substrate during WAIR indicated the onset of flight ability was constrained by neuromuscular control or power output rather than by external wing morphology itself and that partially developed wings not yet capable of flight could indeed provide useful lift during WAIR. Additionally, examination of the work and power requirements for extant bird pectoralis contractile behavior during WAIR at different angles of substrate incline demonstrated incremental increases in these requirements, both as WAIR angles increased and in the transition from WAIR to flapping flight. This provides a model for an evolutionary transition from terrestrial to aerial locomotion as transitional forms incrementally adapted to meet the work and power requirements to scale steeper and steeper inclines using WAIR and the incremental increases from WAIR to flight.
38:
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symmetrical motion would be costly in the cursorial model because it would be difficult while running on the ground, compared to the arboreal model where it is natural for an animal to move both arms together when falling. There is also a large fitness reduction between the two extremes of asymmetrical and symmetrical flapping motion so the theropods would have evolved to one of the extremes. However, new research on the mechanics of bipedal running has suggested that oscillations produced by the running motion could induce symmetrical flapping of the wings at the natural frequency of the oscillation.
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on the body. Many animals, even those which do not fly, demonstrate the ability to right themselves and face the ground ventrally, then exhibiting behaviors that act against aerodynamic forces to slow their rate of descent in a process known as parachuting. Arboreal animals that were forced by predators or simply fell from trees that exhibited these kinds of behaviors would have been in a better position to eventually evolve capabilities that were more akin to flight as we know them today.
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134:. The aerodynamic body of a bird can reduce drag, but when stopping or slowing down a bird will use its tail and feet to increase drag. Weight is the largest obstacle birds must overcome in order to fly. An animal can more easily attain flight by reducing its absolute weight. Birds evolved from other theropod dinosaurs that had already gone through a phase of size reduction during the
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arboreal parachuting, WAIR, and horizontal flap-leaping. Other research also shows that ancestral avialans were not necessarily exclusively arboreal or cursorial, but rather lived on a spectrum of habitats. The capability for powered flight evolved due to a multitude of selective advantages of incipient wings in navigating a more complex environment than previously thought.
255:
Ostrom to describe the use of wings as an insect-foraging mechanism which then evolved into a wing stroke. Research was conducted by comparing the amount of energy expended by each hunting method with the amount of food gathered. The potential hunting volume doubles by running and jumping. To gather the same volume of food,
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Birds use wing-assisted inclined running from the day they hatch to increase locomotion. This can also be said for birds or feathered theropods whose wing muscles cannot generate enough force to fly, and shows how this behavior could have evolved to help these theropods then eventually led to flight.
385:
The evolutionary path between arboreality and flight has been proposed through a number of hypotheses. Dudley and
Yanoviak proposed that animals that live in trees generally end up high enough that a fall, purposeful or otherwise, would generate enough speed for aerodynamic forces to have an effect
377:
exerts energy climbing the tree, it is able to achieve higher velocities and cover greater distances during the gliding phase, which conserves more energy in the long run than a cursorial bipedal runner. Conserving energy during the gliding phase makes this a more energy-efficient model. Therefore,
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versus the distance needed for minimum velocity to obtain liftoff speed is proportional, therefore, as mass increases, the energy required for takeoff increases. Other research has shown that the physics involved in cursorial flight would not make this a likely answer to the origin of avian flight.
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in 1879. This theory states that "flight evolved in running bipeds through a series of short jumps". As the length of the jumps extended, the wings were used not only for thrust but also for stability, and eventually eliminated the gliding intermediate. This theory was modified in the 1970s by John
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Some researchers have suggested that treating arboreal and cursorial hypotheses as mutually exclusive explanations of the origin of bird flight is incorrect. Researchers in support of synthesizing cite studies that show incipient wings have adaptive advantages for a variety of functions, including
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by which modern birds' wings produce swift, powerful upstrokes; since the downforce on which WAIR depends is generated by upstrokes, it seems that early birds were incapable of WAIR. However, a study that found lift generated from wings to be the primary factor for successfully accelerating a body
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bird. This characteristic allows for more strength and stability in the hindlimbs. Thrust produced by the wings coupled with propulsion in the legs generates the minimum velocity required to achieve flight. This wing motion is thought to have evolved from asymmetrical propulsion flapping motion.
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We propose that birds evolved from predators that specialized in ambush from elevated sites, using their raptorial hindlimbs in a leaping attack. Drag-based, and later lift-based, mechanisms evolved under selection for improved control of body position and locomotion during the aerial part of the
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Prior to their existence on birds, feathers were present on the bodies of many dinosaur species. Through natural selection, feathers became more common among the animals as their wings developed over the course of tens of millions of years. The smooth surface of feathers on a bird's body helps to
165:
The mechanics of an avian's wings involve a complex interworking of forces, particularly at the shoulder where most of the wings' motions take place. These functions depend on a precise balance of forces from the muscles, ligaments, and articular cartilages as well as inertial, gravitational, and
169:
Birds have two main muscles in their wing that are responsible for flight: the pectoralis and the supracoracoideus. The pectoralis is the largest muscle in the wing and is the primary depressor and pronator of the wing. The supracoracoideus is the second largest and is the primary elevator and
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The decrease in efficiency when looking at the cursorial model is caused by the flapping stroke needed to achieve flight. This stroke motion needs both wings to move in a symmetrical motion, or together. This is opposed to an asymmetrical motion like that in humans' arms while running. The
138:, combined with rapid evolutionary changes. Flying birds during their evolution further reduced relative weight through several characteristics such as the loss of teeth, shrinkage of the gonads out of mating season, and fusion of bones. Teeth were replaced by a lightweight bill made of
324:, a version of the "cursorial model" of the evolution of avian flight, in which birds' wings originated from forelimb modifications that provided downforce, enabling the proto-birds to run up extremely steep slopes such as the trunks of trees, was prompted by observation of young
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Although the evidence in favor of this model is scientifically plausible, the evidence against it is substantial. For instance, a cursorial flight model would be energetically less favorable when compared to the alternative hypotheses. In order to achieve
158:, that is hollow or filled with air sacs, has often been seen as an adaptation reducing weight, but it was already present in non-flying dinosaurs, and birds on average do not have a lighter skeleton than mammals of the same size. The same is true for the
409:, which has the most complete foot of any known, showed that the hallux was not in fact reversed, limiting the creature's ability to perch on branches and implying a terrestrial or trunk-climbing lifestyle. Another skeletal feature that is similar in
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attack. Selection for enhanced lift-based control led to improved lift coefficients, incidentally turning a pounce into a swoop as lift production increased. Selection for greater swooping range would finally lead to the origin of true flight.
425:, perching characteristics were present, signifying an arboreal habitat. The ability for takeoff and flight was originally thought to require a supracoracoideus pulley system (SC). This system consists of a tendon joining the
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would have a very short and ineffective flight. In contrast to Ostrom's theory regarding flight as a hunting mechanism, physics again does not support this model. In order to effectively trap insects with the wings,
847:"Bird Flight: How Did It Begin? Did birds begin to fly "from the trees down" or "from the ground up"? Reexamination of Archaeopteryx adds plausibility to an "up from the ground" origin of avian flight"
126:) must be favorably combined. In order for birds to balance these forces, certain physical characteristics are required. Asymmetrical wing feathers, found on all flying birds with the exception of
437:. Based on experiments performed by M. Sy in 1936, it was proven that the SC pulley system was not required for flight from an elevated position but was necessary for cursorial takeoff.
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for fine motor coordination. These were gradual changes, though, and not strict conditions for flight: the first birds had teeth, at best a small keel and relatively unfused bones.
401:, or modified of the first digit of the foot, was long thought to have pointed posterior to the remaining digits, as in perching birds. Therefore, researchers once concluded that
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which lived over 150 million years ago, debates still persist regarding the evolution of flight. There are three leading hypotheses pertaining to avian flight: Pouncing
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functions as a result of the need to run quickly up very steep slopes such as tree trunks, for example to escape from predators. Note that in this scenario birds need
213:-like animal, with a skeleton more or less identical to terrestrial theropods, with few adaptations to flapping, but very advanced aerodynamic asymmetrical feathers.
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Lee, Michael S.Y.; Cau, Andrea; Naish, Darren; Dyke, Gareth J. (2014). "Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds".
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is in the air, drag would cause the velocity to instantaneously decrease; balance could not be maintained due to this immediate reduction in velocity. Hence,
241:-bearing feathers began with simple forms that produced a benefit by increasing drag. Later, more refined feather shapes could begin to also provide lift.
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would expend less energy by running and jumping than by running alone. Therefore, the cost/benefit ratio would be more favorable for this model. Due to
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would require a mechanism such as holes in the wings to reduce air resistance. Without this mechanism, the cost/benefit ratio would not be feasible.
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possessed skeletal features similar to those of modern birds. The first such feature to be noted was the supposed similarity between the foot of
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Odontornithes: A monograph on the extinct toothed birds of North
America. Report of the geological exploration of the fortieth parallel, Vol. 7
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the benefits gained by gliding outweigh the energy used in climbing the tree. A modern behavior model to compare against would be that of the
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Talori, Yaser Saffar; Zhao, Jing-Shan; Liu, Yun-Fei; Liu, Wen-Xiu; Li, Zhi-Heng; O'Connor, Jingmai
Kathleen (2 May 2019).
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Zhou, Z (2004). "The origin and early evolution of birds: discoveries, disputes, and
Perspectives from fossil evidence".
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Baier, D.B.; Gatesy, S.M.; Jenkins Jr, F.A. (2007). "A critical ligamentous mechanism in the evolution of avian flight".
382:. In addition to energy conservation, arboreality is generally associated positively with survival, at least in mammals.
17:
1301:
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Feduccia, A. 1999. The Origin and
Evolution of Birds. Yale University Press. London. p. 95, 97, 101, 103–104, 136.
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Heers, Ashley; Dial, Kenneth (2012). "From extant to extinct: locomotor ontogeny and the evolution of avian flight".
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possessed the same claw curvature of the foot to that of perching birds. However, the claw curvature of the hand in
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Elzanowoski, A. 2000. "The Flying
Dinosaurs." Ed. Paul, G. The Scientific American Book of Dinosaurs. p. 178.
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Chatterjee, S. 1997. The Rise of Birds. The Johns
Hopkins University Press. Baltimore. p. 150-151, 153, 158.
983:"Identification of avian flapping motion from non-volant winged dinosaurs based on modal effective mass analysis"
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reduce friction while in flight. The tail, also consisting of feathers, helps the bird to maneuver and glide.
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would have to run faster than modern birds by a factor of three, due to its weight. Furthermore, the mass of
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1236:"The broad range of contractile behaviour of the avian pectoralis: functional and evolutionary implications"
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Scientists believe they could be a step closer to solving the mystery of how the first birds took to the air
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used the hallux as a balancing mechanism on tree limbs. However, study of the
Thermopolis specimen of
130:, help in the production of thrust and lift. Anything that moves through the air produces drag due to
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Tobalske, B. W. & Dial, K. P. (2007). "Aerodynamics of wing-assisted incline running in birds".
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1813:"On the origins of birds: the sequence of character acquisition in the evolution of avian flight"
806:"On the origins of birds: the sequence of character acquisition in the evolution of avian flight"
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bones, allowing rotation of the humerus during the upstroke. However, this system is lacking in
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Tobalske, B. W.; Dial, K. P. (2007). "Aerodynamics of wing-assisted incline running in birds".
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1357:"A fundamental avian wing-stroke provides a new perspective on the evolution of flight"
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Gill, F. 2007. Ornithology. W.H. Freeman and
Company. New York. p. 25, 29, 40–41.
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1175:"Scapular orientation in theropods and basal birds, and the origin of flapping flight"
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supinator. In addition, there are distal wing muscles that assist the bird in flight.
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934:"Forelimb Posture In Dinosaurs And The Evolution Of The Avian Flapping Flight-Stroke"
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885:"Forelimb posture in dinosaurs and the evolution of the avian flapping flight-stroke"
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was similar to that in basal birds. Based upon the comparisons of modern birds to
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to give their feet increased grip. It has been argued that early birds, including
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Sy, Maxheinz (1936). "Funktionell-anatomische Untersuchungen am VogelflĂĽgel".
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It predicts the observed sequence of character acquisition in avian evolution.
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Videler, J. 2005. Avian Flight. Oxford University Press. Oxford. P. 2, 91–98.
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would then use its wings as a balancing mechanism. According to this model,
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and other philosophers of the time attempted to explain the aerodynamics of
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Proceedings of the Royal Society of London. Series B: Biological Sciences
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MAYR, GERALD; POHL, BURKHARD; HARTMAN, SCOTT; PETERS, D. STEFAN (2007).
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developed a gliding method to conserve energy. Even though an arboreal
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565:"This feathery dinosaur probably flew, but not like any bird you know"
478:, a four-winged stage proposed by William Beebe; hindlimb feathers on
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162:, a bone which enhances skeletal bracing for the stresses of flight.
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was a reptilian bird that soared from tree to tree. After the leap,
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644:"A critical ligamentous mechanism in the evolution of avian flight"
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long and erect leg, supporters of this model say the species was a
146:. Other advanced physical characteristics evolved for flight are a
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Baier, David B.; Gatesy, Stephen M.; Jenkins, Farish A. (2006).
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Bennet, Michael B.; Glen, Christopher L. (November 6, 2007).
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The authors believed that this theory had four main virtues:
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1302:"Wing-assisted incline running and the evolution of flight"
1042:"Wing-Assisted Incline Running and the Evolution of Flight"
506:"Wing bone geometry reveals active flight in Archaeopteryx"
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A cursorial, or "running" model was originally proposed by
48:
1619:"Foraging modes of Mesozoic birds and non-avian theropods"
1198:
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was first proposed by Garner, Taylor, and Thomas in 1999:
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Lewin, R (1983). "How did vertebrates take to the air?".
1612:
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1483:"Animal Aloft: The Origins of Aerial Behavior and Flight"
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have been interpreted as evidence of four-winged gliding.
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Researchers in support of this model have suggested that
1234:
Jackson, B. E., Tobalske, B. W. and Dial, K. P. (2011).
1454:. Washington: Government Printing Office. p. 189.
810:
Proceedings of the Royal Society B: Biological Sciences
804:
Garner, J. P.; Taylor, G. K.; Thomas, A. L. R. (1999).
94:, but in a manner substantially different from that of
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Chatterjee, S.; Templin, R. J. (2002). "The flight of
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for the attachment of flight muscles and an enlarged
30:"Theory of flight" redirects here. For the film, see
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Voeten, Dennis F.A.E.; et al. (13 March 2018).
1668:"The origin of avian flight: finding common ground"
1123:"Mechanics of wing-assisted incline running (WAIR)"
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216:It explains that primitive pouncers (perhaps like
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63:. Even after the discovery of the ancestral bird
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2042:Origin of the propatagium in non-avian dinosaurs
1666:Segre, Paolo S.; Banet, Amanda I. (2018-09-18).
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413:and modern birds is the curvature of the claws.
328:chicks, and proposes that wings developed their
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222:) could coexist with more advanced fliers (like
1471:
1355:Dial, K. P.; Jackson, B. E.; Segre, P. (2008).
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272:was able to achieve flight from the ground up.
234:) since they did not compete for flying niches.
357:This model was originally proposed in 1880 by
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1285:: CS1 maint: multiple names: authors list (
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237:It explains that the evolution of elongated
27:Evolution of birds from non-flying ancestors
1811:Garner, J.; Taylor, G.; Thomas, A. (1999).
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1672:Biological Journal of the Linnean Society
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84:In March 2018, scientists reported that
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397:and that of modern perching birds. The
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1890:: early bird catches a can of worms".
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3071:Dialogues Concerning Natural Religion
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1965:
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24:
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289:Once flight speed is obtained and
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2011:
1093:"Secrets of bird flight revealed"
743:Trends in Ecology & Evolution
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951:10.1111/j.1558-5646.2009.00613.x
902:10.1111/j.1558-5646.2009.00613.x
845:Ostrom, John H. (Jan–Feb 1979).
268:Thus, through these mechanisms,
110:to occur, four physical forces (
3133:Extended evolutionary synthesis
2322:Gene-centered view of evolution
1933:The Quarterly Review of Biology
1572:
1406:
1348:
1293:
925:
876:
777:"Bird Anatomy & Bird Parts"
706:Journal of Experimental Biology
166:aerodynamic loads on the wing.
3261:Hologenome theory of evolution
3128:History of molecular evolution
2354:Evolutionarily stable strategy
2243:Last universal common ancestor
769:
734:
693:
584:
563:Guarino, Ben (13 March 2018).
13:
1:
3055:Renaissance and Enlightenment
1701:
1182:Acta Palaeontologica Polonica
702:"Biomechanics of bird flight"
316:Wing-assisted incline running
310:Wing-assisted incline running
177:
3266:Missing heritability problem
2893:Gamete differentiation/sexes
1912:10.1126/science.259.5096.764
1008:10.1371/journal.pcbi.1006846
883:Nudds, R.; Dyke, G. (2009).
490:
440:
47:, one of the earliest known
7:
1931:and the origin of flight".
1871:10.1126/science.221.4605.38
449:
10:
3782:
3581:Flying and gliding animals
3417:Fin and flipper locomotion
2898:Life cycles/nuclear phases
2450:Trivers–Willard hypothesis
987:PLOS Computational Biology
755:10.1016/j.tree.2011.12.003
532:10.1038/s41467-018-03296-8
466:Flying and gliding animals
313:
29:
3692:
3631:
3571:
3474:
3463:
3392:
3316:
3216:
3141:
3045:
2972:
2928:
2783:
2687:
2504:
2463:
2396:Parent–offspring conflict
2332:
2201:Earliest known life forms
2122:
2089:
2032:How Birds Got their wings
1988:10.1007/s00114-004-0570-4
1782:10.1007/s00114-002-0385-0
1684:10.1093/biolinnean/bly116
1636:10.1016/j.cub.2007.09.026
3249:Cultural group selection
3113:The eclipse of Darwinism
3085:On the Origin of Species
3060:Transmutation of species
1581:Journal fĂĽr Ornithologie
1239:(Automatic PDF download)
1178:(Automatic PDF download)
252:Samuel Wendell Williston
3254:Dual inheritance theory
3093:History of paleontology
1460:10.5962/bhl.title.61298
1326:10.1126/science.1078237
1066:10.1126/science.1078237
700:Tobalske, Bret (2007).
613:10.1126/science.1252243
187:A theory of a pouncing
3756:Evolution by phenotype
3720:Terrestrial locomotion
3664:Evolution of cetaceans
3659:Origin of avian flight
3644:Evolution of tetrapods
2942:Punctuated equilibrium
2263:Non-adaptive radiation
2211:Evolutionary arms race
1829:10.1098/rspb.1999.0772
822:10.1098/rspb.1999.0772
198:
183:Pouncing Proavis model
102:Flight characteristics
90:was likely capable of
52:
3735:Undulatory locomotion
3684:Homologous structures
3234:Evolutionary medicine
3108:Mendelian inheritance
2816:Biological complexity
2804:Programmed cell death
2496:Phenotypic plasticity
2216:Evolutionary pressure
2206:Evidence of evolution
2104:Timeline of evolution
1448:Marsh, O. C. (1880).
511:Nature Communications
193:
40:
3679:Analogous structures
3674:Convergent evolution
3208:Teleology in biology
3103:Blending inheritance
2481:Genetic assimilation
2344:Artificial selection
2083:Evolutionary biology
2048:. February 23, 2023.
2038:. February 24, 2023.
1300:Dial, K. P. (2003).
1249:(Pt 14): 2354–2361.
1207:(Pt 10): 1742–1751.
1136:(Pt 24): 4553–4564.
361:. The theory states
32:The Theory of Flight
3730:Rotating locomotion
3669:Comparative anatomy
3271:Molecular evolution
3229:Ecological genetics
3098:Transitional fossil
2888:Sexual reproduction
2728:endomembrane system
2657:pollinator-mediated
2613:dolphins and whales
2391:Parental investment
1980:2004NW.....91..455Z
1968:Naturwissenschaften
1927:Ostrom, J (1974). "
1904:1993Sci...259..764M
1886:Morell, V (1993). "
1863:1983Sci...221...38L
1823:(1425): 1259–1266.
1774:2003NW.....90...27C
1762:Naturwissenschaften
1730:10.1038/nature05435
1722:2007Natur.445..307B
1384:10.1038/nature06517
1376:2008Natur.451..985D
1318:2003Sci...299..402D
1173:Senter, P. (2006).
1091:(24 January 2008).
1058:2003Sci...299..402D
1040:Dial, K.P. (2003).
999:2019PLSCB..15E6846T
816:(1425): 1259–1266.
671:10.1038/nature05435
663:2007Natur.445..307B
605:2014Sci...345..562L
570:The Washington Post
524:2018NatCo...9..923V
18:Evolution of flight
3649:Evolution of birds
3402:Aquatic locomotion
3244:Cultural evolution
2359:Fisher's principle
2288:Handicap principle
2278:Parallel evolution
2142:Adaptive radiation
2046:Zoological Letters
1593:10.1007/BF01906709
1503:10.1093/icb/icr002
1427:10.1242/jeb.001701
1256:10.1242/jeb.052829
1213:10.1242/jeb.001701
851:American Scientist
719:10.1242/jeb.000273
342:shoulder mechanism
53:
3743:
3742:
3700:Animal locomotion
3639:Evolution of fish
3519:facultative biped
3344:
3343:
2960:Uniformitarianism
2913:Sex-determination
2418:Sexual dimorphism
2413:Natural selection
2317:Unit of selection
2283:Signalling theory
2027:Arboreal argument
2023:(journal article)
1898:(5096): 764–765.
1716:(7125): 307–310.
1629:(21): R911–R912.
1143:10.1242/jeb.00673
1052:(5605): 402–404.
712:(18): 3135–3146.
657:(7125): 307–310.
599:(6196): 562–566.
16:(Redirected from
3773:
3710:Robot locomotion
3484:Limb development
3469:
3442:Lobe-finned fish
3371:
3364:
3357:
3348:
3347:
3334:
3324:
3323:
3123:Modern synthesis
2883:Multicellularity
2878:Mosaic evolution
2763:auditory ossicle
2445:Social selection
2428:Flowering plants
2423:Sexual selection
2076:
2069:
2062:
2053:
2052:
2007:
1956:
1923:
1882:
1842:
1840:
1801:
1749:
1695:
1694:
1692:
1690:
1663:
1657:
1656:
1638:
1614:
1605:
1604:
1576:
1570:
1569:
1567:
1549:
1536:
1530:
1529:
1527:
1526:
1520:
1514:. Archived from
1505:
1487:
1478:
1469:
1463:
1445:
1439:
1438:
1410:
1404:
1403:
1361:
1352:
1346:
1345:
1297:
1291:
1290:
1284:
1276:
1258:
1240:
1231:
1225:
1224:
1196:
1190:
1189:
1179:
1170:
1164:
1163:
1145:
1127:
1118:
1112:
1111:
1109:
1107:
1097:
1089:Morelle, Rebecca
1085:
1037:
1031:
1030:
1020:
1010:
978:
972:
971:
953:
929:
923:
922:
904:
880:
874:
873:
871:
869:
842:
836:
835:
833:
801:
792:
791:
789:
787:
773:
767:
766:
738:
732:
731:
721:
697:
691:
690:
648:
639:
633:
632:
588:
582:
581:
579:
577:
560:
554:
553:
543:
501:
359:Othniel C. Marsh
55:Around 350 BCE,
21:
3781:
3780:
3776:
3775:
3774:
3772:
3771:
3770:
3746:
3745:
3744:
3739:
3688:
3654:Origin of birds
3627:
3567:
3489:Limb morphology
3470:
3461:
3447:Ray-finned fish
3412:Fish locomotion
3388:
3375:
3345:
3340:
3312:
3239:Group selection
3212:
3137:
3041:
2968:
2930:Tempo and modes
2924:
2779:
2683:
2500:
2459:
2335:
2328:
2305:Species complex
2118:
2109:History of life
2085:
2080:
2014:
1974:(10): 455–471.
1857:(4605): 38–39.
1704:
1699:
1698:
1688:
1686:
1664:
1660:
1623:Current Biology
1615:
1608:
1577:
1573:
1547:
1537:
1533:
1524:
1522:
1518:
1485:
1479:
1472:
1446:
1442:
1421:(10): 1742–51.
1411:
1407:
1370:(7181): 985–9.
1359:
1353:
1349:
1312:(5605): 402–4.
1298:
1294:
1278:
1277:
1238:
1232:
1228:
1197:
1193:
1177:
1171:
1167:
1125:
1119:
1115:
1105:
1103:
1095:
1038:
1034:
993:(5): e1006846.
979:
975:
944:(4): 994–1002.
930:
926:
895:(4): 994–1002.
881:
877:
867:
865:
843:
839:
802:
795:
785:
783:
775:
774:
770:
739:
735:
698:
694:
646:
640:
636:
589:
585:
575:
573:
561:
557:
502:
498:
493:
456:Origin of birds
452:
443:
380:flying squirrel
355:
322:WAIR hypothesis
318:
312:
261:Archaeopteryx's
248:
246:Cursorial model
207:It predicts an
185:
180:
136:Middle Jurassic
104:
35:
28:
23:
22:
15:
12:
11:
5:
3779:
3769:
3768:
3763:
3758:
3741:
3740:
3738:
3737:
3732:
3727:
3722:
3717:
3712:
3707:
3702:
3696:
3694:
3690:
3689:
3687:
3686:
3681:
3676:
3671:
3666:
3661:
3656:
3651:
3646:
3641:
3635:
3633:
3629:
3628:
3626:
3625:
3620:
3618:Pterosaur wing
3615:
3610:
3609:
3608:
3603:
3598:
3588:
3583:
3577:
3575:
3569:
3568:
3566:
3565:
3560:
3555:
3554:
3553:
3543:
3538:
3533:
3532:
3531:
3526:
3521:
3516:
3511:
3506:
3501:
3496:
3486:
3480:
3478:
3472:
3471:
3464:
3462:
3460:
3459:
3454:
3449:
3444:
3439:
3434:
3429:
3424:
3419:
3414:
3409:
3407:Cephalopod fin
3404:
3398:
3396:
3390:
3389:
3374:
3373:
3366:
3359:
3351:
3342:
3341:
3339:
3338:
3328:
3317:
3314:
3313:
3311:
3310:
3305:
3300:
3295:
3290:
3289:
3288:
3278:
3273:
3268:
3263:
3258:
3257:
3256:
3251:
3246:
3236:
3231:
3226:
3220:
3218:
3214:
3213:
3211:
3210:
3205:
3204:
3203:
3198:
3193:
3192:
3191:
3181:
3176:
3171:
3166:
3161:
3151:
3145:
3143:
3139:
3138:
3136:
3135:
3130:
3125:
3120:
3115:
3110:
3105:
3100:
3095:
3090:
3089:
3088:
3079:Charles Darwin
3076:
3075:
3074:
3062:
3057:
3051:
3049:
3043:
3042:
3040:
3039:
3034:
3029:
3024:
3019:
3017:Non-ecological
3014:
3009:
3004:
2999:
2994:
2989:
2984:
2978:
2976:
2970:
2969:
2967:
2966:
2957:
2948:
2934:
2932:
2926:
2925:
2923:
2922:
2917:
2916:
2915:
2910:
2905:
2900:
2895:
2885:
2880:
2875:
2870:
2865:
2860:
2855:
2850:
2845:
2840:
2835:
2834:
2833:
2823:
2818:
2813:
2808:
2807:
2806:
2801:
2790:
2788:
2781:
2780:
2778:
2777:
2776:
2775:
2770:
2768:nervous system
2765:
2760:
2755:
2747:
2746:
2745:
2740:
2735:
2730:
2725:
2720:
2710:
2705:
2700:
2694:
2692:
2685:
2684:
2682:
2681:
2676:
2671:
2666:
2661:
2660:
2659:
2649:
2648:
2647:
2642:
2641:
2640:
2635:
2625:
2620:
2615:
2610:
2605:
2604:
2603:
2598:
2588:
2578:
2573:
2572:
2571:
2561:
2556:
2551:
2546:
2545:
2544:
2534:
2529:
2528:
2527:
2517:
2511:
2509:
2502:
2501:
2499:
2498:
2493:
2488:
2483:
2478:
2473:
2467:
2465:
2461:
2460:
2458:
2457:
2452:
2447:
2442:
2441:
2440:
2435:
2430:
2420:
2415:
2410:
2405:
2400:
2399:
2398:
2393:
2383:
2378:
2373:
2372:
2371:
2361:
2356:
2351:
2346:
2340:
2338:
2330:
2329:
2327:
2326:
2325:
2324:
2314:
2309:
2308:
2307:
2302:
2292:
2291:
2290:
2280:
2275:
2270:
2268:Origin of life
2265:
2260:
2255:
2253:Microevolution
2250:
2248:Macroevolution
2245:
2240:
2235:
2234:
2233:
2223:
2218:
2213:
2208:
2203:
2198:
2193:
2188:
2186:Common descent
2183:
2182:
2181:
2171:
2166:
2164:Baldwin effect
2161:
2160:
2159:
2154:
2144:
2139:
2134:
2128:
2126:
2120:
2119:
2117:
2116:
2111:
2106:
2101:
2096:
2090:
2087:
2086:
2079:
2078:
2071:
2064:
2056:
2050:
2049:
2039:
2029:
2024:
2018:Flight of the
2013:
2012:External links
2010:
2009:
2008:
1963:
1960:
1957:
1945:10.1086/407902
1924:
1883:
1846:
1843:
1808:
1805:
1802:
1753:
1750:
1703:
1700:
1697:
1696:
1678:(2): 452–454.
1658:
1606:
1587:(2): 199–296.
1571:
1531:
1496:(6): 926–936.
1470:
1440:
1405:
1347:
1292:
1226:
1191:
1165:
1113:
1086:Summarized in
1032:
973:
924:
875:
837:
793:
768:
749:(5): 296–305.
733:
692:
634:
583:
555:
495:
494:
492:
489:
488:
487:
473:
468:
463:
458:
451:
448:
442:
439:
354:
353:Arboreal model
351:
314:Main article:
311:
308:
247:
244:
243:
242:
235:
225:Confuciusornis
214:
205:
184:
181:
179:
176:
156:Pneumatic bone
103:
100:
26:
9:
6:
4:
3:
2:
3778:
3767:
3764:
3762:
3759:
3757:
3754:
3753:
3751:
3736:
3733:
3731:
3728:
3726:
3723:
3721:
3718:
3716:
3713:
3711:
3708:
3706:
3703:
3701:
3698:
3697:
3695:
3691:
3685:
3682:
3680:
3677:
3675:
3672:
3670:
3667:
3665:
3662:
3660:
3657:
3655:
3652:
3650:
3647:
3645:
3642:
3640:
3637:
3636:
3634:
3630:
3624:
3621:
3619:
3616:
3614:
3611:
3607:
3604:
3602:
3599:
3597:
3594:
3593:
3592:
3589:
3587:
3584:
3582:
3579:
3578:
3576:
3574:
3570:
3564:
3561:
3559:
3556:
3552:
3549:
3548:
3547:
3544:
3542:
3539:
3537:
3534:
3530:
3527:
3525:
3522:
3520:
3517:
3515:
3512:
3510:
3507:
3505:
3502:
3500:
3497:
3495:
3492:
3491:
3490:
3487:
3485:
3482:
3481:
3479:
3477:
3473:
3468:
3458:
3455:
3453:
3452:Pectoral fins
3450:
3448:
3445:
3443:
3440:
3438:
3435:
3433:
3430:
3428:
3425:
3423:
3420:
3418:
3415:
3413:
3410:
3408:
3405:
3403:
3400:
3399:
3397:
3395:
3391:
3387:
3383:
3379:
3372:
3367:
3365:
3360:
3358:
3353:
3352:
3349:
3337:
3333:
3329:
3327:
3319:
3318:
3315:
3309:
3306:
3304:
3301:
3299:
3296:
3294:
3291:
3287:
3284:
3283:
3282:
3281:Phylogenetics
3279:
3277:
3274:
3272:
3269:
3267:
3264:
3262:
3259:
3255:
3252:
3250:
3247:
3245:
3242:
3241:
3240:
3237:
3235:
3232:
3230:
3227:
3225:
3222:
3221:
3219:
3215:
3209:
3206:
3202:
3199:
3197:
3194:
3190:
3187:
3186:
3185:
3184:Structuralism
3182:
3180:
3177:
3175:
3172:
3170:
3167:
3165:
3162:
3160:
3159:Catastrophism
3157:
3156:
3155:
3152:
3150:
3147:
3146:
3144:
3140:
3134:
3131:
3129:
3126:
3124:
3121:
3119:
3118:Neo-Darwinism
3116:
3114:
3111:
3109:
3106:
3104:
3101:
3099:
3096:
3094:
3091:
3087:
3086:
3082:
3081:
3080:
3077:
3073:
3072:
3068:
3067:
3066:
3063:
3061:
3058:
3056:
3053:
3052:
3050:
3048:
3044:
3038:
3035:
3033:
3032:Reinforcement
3030:
3028:
3025:
3023:
3020:
3018:
3015:
3013:
3010:
3008:
3005:
3003:
3000:
2998:
2995:
2993:
2990:
2988:
2985:
2983:
2980:
2979:
2977:
2975:
2971:
2965:
2964:Catastrophism
2961:
2958:
2956:
2955:Macromutation
2952:
2951:Micromutation
2949:
2947:
2943:
2939:
2936:
2935:
2933:
2931:
2927:
2921:
2918:
2914:
2911:
2909:
2906:
2904:
2901:
2899:
2896:
2894:
2891:
2890:
2889:
2886:
2884:
2881:
2879:
2876:
2874:
2871:
2869:
2866:
2864:
2861:
2859:
2858:Immune system
2856:
2854:
2851:
2849:
2846:
2844:
2841:
2839:
2836:
2832:
2829:
2828:
2827:
2824:
2822:
2819:
2817:
2814:
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2721:
2719:
2718:symbiogenesis
2716:
2715:
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2416:
2414:
2411:
2409:
2406:
2404:
2401:
2397:
2394:
2392:
2389:
2388:
2387:
2386:Kin selection
2384:
2382:
2381:Genetic drift
2379:
2377:
2374:
2370:
2367:
2366:
2365:
2362:
2360:
2357:
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2209:
2207:
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2202:
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2177:
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2170:
2167:
2165:
2162:
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2143:
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2129:
2127:
2125:
2121:
2115:
2112:
2110:
2107:
2105:
2102:
2100:
2097:
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2092:
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2088:
2084:
2077:
2072:
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2065:
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2058:
2057:
2054:
2047:
2043:
2040:
2037:
2033:
2030:
2028:
2025:
2022:
2021:
2020:Archaeopteryx
2016:
2015:
2005:
2001:
1997:
1993:
1989:
1985:
1981:
1977:
1973:
1969:
1964:
1961:
1958:
1954:
1950:
1946:
1942:
1938:
1934:
1930:
1929:Archaeopteryx
1925:
1921:
1917:
1913:
1909:
1905:
1901:
1897:
1893:
1889:
1888:Archaeopteryx
1884:
1880:
1876:
1872:
1868:
1864:
1860:
1856:
1852:
1847:
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1779:
1775:
1771:
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1763:
1759:
1758:Archaeopteryx
1754:
1751:
1747:
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1735:
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1727:
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1613:
1611:
1602:
1598:
1594:
1590:
1586:
1582:
1575:
1566:
1561:
1558:(1): 97–116.
1557:
1553:
1546:
1544:
1543:Archaeopteryx
1535:
1521:on 2014-04-25
1517:
1513:
1509:
1504:
1499:
1495:
1491:
1484:
1477:
1475:
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1461:
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1248:
1244:
1237:
1230:
1222:
1218:
1214:
1210:
1206:
1202:
1195:
1188:(2): 305–313.
1187:
1183:
1176:
1169:
1161:
1157:
1153:
1149:
1144:
1139:
1135:
1131:
1124:
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1094:
1090:
1083:
1079:
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1028:
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961:
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952:
947:
943:
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916:
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908:
903:
898:
894:
890:
886:
879:
864:
860:
856:
852:
848:
841:
832:
827:
823:
819:
815:
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807:
800:
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778:
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764:
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756:
752:
748:
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737:
729:
725:
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715:
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703:
696:
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664:
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474:
472:
471:Insect flight
469:
467:
464:
462:
459:
457:
454:
453:
447:
438:
436:
435:Archaeopteryx
432:
428:
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423:Archaeopteryx
420:
419:Archaeopteryx
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415:Archaeopteryx
412:
411:Archaeopteryx
408:
407:Archaeopteryx
404:
403:Archaeopteryx
400:
396:
395:Archaeopteryx
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391:Archaeopteryx
387:
383:
381:
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375:Archaeopteryx
372:
371:Archaeopteryx
368:
367:Archaeopteryx
364:
363:Archaeopteryx
360:
350:
346:
343:
340:, lacked the
339:
338:Archaeopteryx
335:
331:
327:
323:
317:
307:
303:
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300:Archaeopteryx
296:
295:Archaeopteryx
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291:Archaeopteryx
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286:Archaeopteryx
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270:Archaeopteryx
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257:Archaeopteryx
253:
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210:Archaeopteryx
206:
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149:
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117:
113:
109:
99:
97:
93:
89:
88:
87:Archaeopteryx
82:
80:
76:
72:
68:
67:
66:Archaeopteryx
62:
58:
50:
46:
45:
44:Archaeopteryx
39:
33:
19:
3658:
3293:Polymorphism
3276:Astrobiology
3224:Biogeography
3179:Saltationism
3169:Orthogenesis
3154:Alternatives
3083:
3069:
3002:Cospeciation
2997:Cladogenesis
2946:Saltationism
2903:Mating types
2826:Color vision
2811:Avian flight
2810:
2733:mitochondria
2471:Canalisation
2349:Biodiversity
2094:Introduction
2045:
2035:
2019:
1971:
1967:
1936:
1932:
1928:
1895:
1891:
1887:
1854:
1850:
1820:
1816:
1768:(1): 27–32.
1765:
1761:
1757:
1713:
1709:
1687:. Retrieved
1675:
1671:
1661:
1626:
1622:
1584:
1580:
1574:
1555:
1551:
1542:
1534:
1523:. Retrieved
1516:the original
1493:
1489:
1450:
1443:
1418:
1414:
1408:
1367:
1363:
1350:
1309:
1305:
1295:
1281:cite journal
1246:
1242:
1229:
1204:
1200:
1194:
1185:
1181:
1168:
1133:
1129:
1116:
1104:. Retrieved
1099:
1049:
1045:
1035:
990:
986:
976:
941:
937:
927:
892:
888:
878:
866:. Retrieved
857:(1): 46–56.
854:
850:
840:
813:
809:
784:. Retrieved
780:
771:
746:
742:
736:
709:
705:
695:
654:
650:
637:
596:
592:
586:
574:. Retrieved
568:
558:
518:(923): 923.
515:
509:
499:
444:
434:
422:
418:
414:
410:
406:
402:
394:
390:
388:
384:
374:
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356:
347:
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333:
319:
304:
299:
294:
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285:
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274:
269:
260:
256:
249:
229:
223:
217:
208:
199:
194:
186:
172:
168:
164:
128:hummingbirds
105:
96:modern birds
85:
83:
64:
61:avian flight
54:
42:
3766:Bird flight
3761:Ornithology
3613:Insect wing
3563:Webbed foot
3504:unguligrade
3499:plantigrade
3494:digitigrade
3303:Systematics
3174:Mutationism
2992:Catagenesis
2920:Snake venom
2853:Eusociality
2831:in primates
2821:Cooperation
2749:In animals
2569:butterflies
2542:Cephalopods
2532:Brachiopods
2464:Development
2438:Mate choice
2191:Convergence
2174:Coevolution
2132:Abiogenesis
1689:14 November
868:14 November
480:Microraptor
476:Tetrapteryx
461:Bird flight
330:aerodynamic
265:terrestrial
219:Microraptor
77:model, and
41:The Berlin
3750:Categories
3541:Cephalopod
3457:Pelvic fin
3427:Dorsal fin
3422:Caudal fin
3164:Lamarckism
3142:Philosophy
3065:David Hume
3027:Peripatric
3022:Parapatric
3007:Ecological
2987:Anagenesis
2982:Allopatric
2974:Speciation
2938:Gradualism
2863:Metabolism
2723:chromosome
2713:Eukaryotes
2491:Modularity
2408:Population
2334:Population
2295:Speciation
2273:Panspermia
2226:Extinction
2221:Exaptation
2196:Divergence
2169:Cladistics
2157:Reciprocal
2137:Adaptation
1702:References
1525:2014-04-24
1466:Other link
1106:25 January
1102:. BBC News
484:Anchiornis
178:Hypotheses
152:cerebellum
3632:Evolution
3591:Bird wing
3536:Arthropod
3529:quadruped
3298:Protocell
3149:Darwinism
3037:Sympatric
2786:processes
2674:Tetrapods
2623:Kangaroos
2549:Dinosaurs
2486:Inversion
2455:Variation
2376:Gene flow
2369:Inclusive
2179:Mutualism
2124:Evolution
1939:: 27–47.
938:Evolution
889:Evolution
781:All-Birds
491:Footnotes
441:Synthesis
334:downforce
231:Sapeornis
75:Cursorial
57:Aristotle
3623:Wingspan
3606:feathers
3601:skeleton
3586:Bat wing
3546:Tetrapod
3432:Fish fin
3326:Category
3201:Vitalism
3196:Theistic
3189:Spandrel
2873:Morality
2868:Monogamy
2743:plastids
2708:Flagella
2664:Reptiles
2645:sea cows
2628:primates
2537:Molluscs
2515:Bacteria
2403:Mutation
2336:genetics
2312:Taxonomy
2258:Mismatch
2238:Homology
2152:Cheating
2147:Altruism
2036:Phys.org
1996:15365634
1953:85396846
1920:17809336
1879:17738003
1798:25382695
1790:12545240
1738:17173029
1645:17983564
1601:36259402
1512:21558180
1435:17488937
1400:15166485
1392:18216784
1342:40712093
1334:12532020
1265:21697427
1221:17488937
1152:14610039
1082:40712093
1074:12532020
1027:31048911
968:29012467
960:19154383
919:29012467
911:19154383
863:27849060
763:22304966
728:17766290
679:17173029
629:37866029
621:25082702
576:13 March
550:29535376
450:See also
431:coracoid
132:friction
79:Arboreal
3693:Related
3551:dactyly
3437:Flipper
3217:Related
3047:History
2908:Meiosis
2843:Empathy
2838:Emotion
2738:nucleus
2679:Viruses
2669:Spiders
2581:Mammals
2564:Insects
2364:Fitness
2300:Species
2099:Outline
2004:3329625
1976:Bibcode
1900:Bibcode
1892:Science
1859:Bibcode
1851:Science
1838:1690052
1770:Bibcode
1746:4379208
1718:Bibcode
1372:Bibcode
1314:Bibcode
1306:Science
1273:7496862
1160:6323207
1054:Bibcode
1046:Science
1018:6497222
995:Bibcode
831:1690052
786:9 April
687:4379208
659:Bibcode
601:Bibcode
593:Science
541:5849612
520:Bibcode
427:humerus
278:liftoff
189:proavis
160:furcula
144:gizzard
140:keratin
81:model.
73:model,
71:Proavis
3715:Samara
3524:triped
3509:uniped
3336:Portal
3012:Hybrid
2848:Ethics
2690:organs
2652:Plants
2638:lemurs
2633:humans
2618:horses
2608:hyenas
2596:wolves
2591:canids
2525:origin
2002:
1994:
1951:
1918:
1877:
1835:
1796:
1788:
1744:
1736:
1710:Nature
1653:535424
1651:
1643:
1599:
1510:
1433:
1398:
1390:
1364:Nature
1340:
1332:
1271:
1263:
1219:
1158:
1150:
1080:
1072:
1025:
1015:
966:
958:
917:
909:
861:
828:
761:
726:
685:
677:
651:Nature
627:
619:
548:
538:
399:hallux
326:chukar
239:rachis
124:weight
112:thrust
108:flight
92:flight
3573:Wings
3558:Digit
3514:biped
3476:Limbs
3386:wings
3382:limbs
2799:Death
2794:Aging
2773:brain
2559:Fungi
2520:Birds
2433:Fungi
2231:Event
2114:Index
2000:S2CID
1949:S2CID
1794:S2CID
1742:S2CID
1649:S2CID
1597:S2CID
1548:(PDF)
1519:(PDF)
1486:(PDF)
1396:S2CID
1360:(PDF)
1338:S2CID
1269:S2CID
1156:S2CID
1126:(PDF)
1096:(Web)
1078:S2CID
964:S2CID
915:S2CID
859:JSTOR
683:S2CID
647:(PDF)
625:S2CID
49:birds
3705:Gait
3596:keel
3394:Fins
3384:and
3378:Fins
3286:Tree
2758:hair
2698:Cell
2601:dogs
2586:cats
2576:Life
2554:Fish
2507:taxa
1992:PMID
1916:PMID
1875:PMID
1786:PMID
1734:PMID
1691:2020
1641:PMID
1508:PMID
1431:PMID
1388:PMID
1330:PMID
1287:link
1261:PMID
1217:PMID
1148:PMID
1108:2008
1070:PMID
1023:PMID
956:PMID
907:PMID
870:2020
788:2016
759:PMID
724:PMID
675:PMID
617:PMID
578:2018
546:PMID
482:and
429:and
320:The
148:keel
122:and
120:lift
116:drag
114:and
106:For
2784:Of
2753:eye
2703:DNA
2688:Of
2505:Of
1984:doi
1941:doi
1908:doi
1896:259
1867:doi
1855:221
1833:PMC
1825:doi
1821:266
1778:doi
1760:".
1726:doi
1714:445
1680:doi
1676:125
1631:doi
1589:doi
1560:doi
1556:149
1498:doi
1456:doi
1423:doi
1419:210
1380:doi
1368:451
1322:doi
1310:299
1251:doi
1247:214
1209:doi
1205:210
1138:doi
1134:206
1062:doi
1050:299
1013:PMC
1003:doi
946:doi
897:doi
826:PMC
818:doi
814:266
751:doi
714:doi
710:210
667:doi
655:445
609:doi
597:345
536:PMC
528:doi
228:or
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1990:.
1982:.
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