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Paraves ( Tšekki )

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Paraves je klad, do kterého patří všichni ptáci (Avialae) i jejich nejbližší příbuzní ze skupiny teropodních dinosaurů (skupina Deinonychosauria). Mezi zástupci bychom tedy mohli jmenovat jak vrabce domácího, rod Archaeopteryx, tak i rody Velociraptor nebo Troodon. Název Paraves poprvé zavedl americký paleontolog Paul Sereno v roce 1997.

Ancestrální forma paravianů měla zřejmě podobu malého, několik stovek gramů vážícího opeřeného teropoda. Mohl mít také čtyřkřídlou stavbu těla (jak ukazují rody Anchiornis, Microraptor nebo Pedopenna). Byl zřejmě schopen aktivního nebo alespoň klouzavého letu z větví stromů. Z této hypotetické formy se pak vyvinuli všichni odvozenější paraviani, včetně pravých ptáků.

Zástupci

Zdroje

  • Sereno, P. C., 1997, "The origin and evolution of dinosaurs", Annual Review of Earth & Planetary Sciences 25:435- 489. (21)
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Paraves: Brief Summary ( Tšekki )

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Paraves je klad, do kterého patří všichni ptáci (Avialae) i jejich nejbližší příbuzní ze skupiny teropodních dinosaurů (skupina Deinonychosauria). Mezi zástupci bychom tedy mohli jmenovat jak vrabce domácího, rod Archaeopteryx, tak i rody Velociraptor nebo Troodon. Název Paraves poprvé zavedl americký paleontolog Paul Sereno v roce 1997.

Ancestrální forma paravianů měla zřejmě podobu malého, několik stovek gramů vážícího opeřeného teropoda. Mohl mít také čtyřkřídlou stavbu těla (jak ukazují rody Anchiornis, Microraptor nebo Pedopenna). Byl zřejmě schopen aktivního nebo alespoň klouzavého letu z větví stromů. Z této hypotetické formy se pak vyvinuli všichni odvozenější paraviani, včetně pravých ptáků.

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Paraves ( englanti )

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Paraves are a widespread group of theropod dinosaurs that originated in the Late Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and scansoriopterygids, the group also contains the avialans, among which are the over ten thousand species of living birds.[1] Primitive members of Paraves are well known for the possession of an enlarged claw on the second digit of the foot, which was held off the ground when walking in some species.[2]

Description

Like other theropods, all paravians are bipedal, walking on their two hind legs.[3]

The teeth of primitive paravians were curved and serrated, but not blade-like except in some specialized species such as Dromaeosaurus albertensis. The serrations on the front edge of dromaeosaurid and troodontid teeth were very small and fine, while the back edge had serrations which were very large and hooked.[4]

Most of the earliest paravian groups were carnivorous, though some smaller species (especially among the troodontids and early avialans) are known to have been omnivores, and it has been suggested that an omnivorous diet was the ancestral state for this group, with strict carnivory evolving in some specialized lineages.[5][4] Fossils also suggest that legs and feet covered with feathers was an ancestral condition, possibly having originated in the Coelurosauria, even if this trait was later lost in more advanced birds.[6]

Wings

Paravians generally have long, winged forelimbs, though these have become smaller in many flightless species and some extinct lineages that evolved before flight. The wings usually bore three large, flexible, clawed fingers in early forms.[4] The fingers became fused and stiffened and the claws highly reduced or lost in some advanced lineages. An increasingly asymmetric wrist joint, a trend that can be traced back to primitive coelurosaurs, allowed the forelimbs to elongate and an elaboration of their plumage, traits that made the evolution of flapping flight possible.[7]

Many early members of Paraves had both well-developed wings and long feathers on the hind legs, which in some cases, formed a second set of airfoils. These species, most famously represented by Microraptor gui, have often been referred to as "four winged dinosaurs".[8][9][10] Though it has been suggested that these hind wings would have prevented some paravians from getting around on the ground, and that they must have lived in trees, there is very little evidence that any of the earliest paravians were capable of climbing. This apparent paradox was addressed by later studies which showed that early paravians like Microraptor were capable of flapping flight and powered launching from the ground into the air without relying on climbing. Microraptor in particular also seems to represent a case of flight evolving independently of the bird lineage within Paraves.[11]

Feet and sickle claw

Most theropods walked with three toes contacting the ground, but fossilized footprint tracks confirm that many basal paravians, including dromaeosaurids, troodontids, and some early avialans, held the second toe off the ground in a hyperextended position, with only the third and fourth toes bearing the weight of the animal. This is called functional didactyly.[2] The enlarged second toe bore an unusually large, curved sickle-shaped claw (held off the ground or 'retracted' when walking). This claw was especially large and flattened from side to side in the large-bodied predatory eudromaeosaurs.[12] In these early species, the first toe (hallux) was usually small and angled inward toward the center of the body, but only became fully reversed in more specialized members of the bird lineage.[4] One species, Balaur bondoc, possessed a first toe which was highly modified in parallel with the second. Both the first and second toes on each foot of B. bondoc were held retracted and bore enlarged, sickle-shaped claws.[13]

 src=
Deinonychus "sickle claw"

One of the best-known features of paravians is the presence of an enlarged and strongly curved "sickle claw" on a hyper-extendible second toe, modified to hold the sickle claw clear of the ground when walking, most notably developed in the dromaeosaurids and troodontids. While this characteristic claw and its associated modifications to the anatomy of the foot (such as a shortened metatarsus in eudromaeosaurs) had been known since the mid-20th century, their possible functions were the subject mainly of speculation, and few actual studies were published. Initial speculation regarded the claws as slashing implements used to disembowel large prey. In this scenario, the shortened upper foot would serve as an anchor point for powerful tendons to improve kicking ability. However, subsequent studies of the actual claw shape showed that the underside of the claw was only weakly keeled and would not have been an effective cutting instrument. Instead, it appeared to be more of a hooking implement. Manning et al. suggested in 2006 that the claws were similar to crampons and were used for climbing, and in the case of larger species or individuals, climbing up the flanks of very large prey.[14]

A larger study of sickle-claw function, published in 2011 by Fowler and colleagues, concluded that the earlier study by Manning and colleagues was correct and that the "sickle claws" would have been ineffective as cutting weapons. They compared the claw and overall foot anatomy of various primitive species with modern birds to shed light on their actual function. Fowler and colleagues showed that many modern predatory birds also have enlarged claws on the second toes. In modern raptors, these claws are used to help grip and hold prey of sizes smaller than or equal to the predator, while the birds use their body weight to pin their prey to the ground and eat it alive.[4] Fowler and colleagues suggested that this behavior is entirely consistent with the anatomy of advanced dromaeosaurids like Deinonychus, which had slightly opposing first toes and strong tendons in the toes and foot. This makes it likely that advanced dromaeosaurids also used their claws to puncture and grip their prey to aid in pinning it to the ground, while using shallow wing beats and tail movements to stabilize themselves.[4] Other lines of evidence for this behavior include teeth which had large, hooked serrations only on the back edge (useful in pulling flesh upward rather than slicing it) and large claws on the wings (for greater maneuvering of prey while mantling it with the wings).[4]

In more primitive dromaeosaurids and in troodontids, the feet were not as specialized and the claws were not as large or as hooked. Additionally, the toe joints allowed more range of motion than the simple up-down movements of advanced dromaeosaurids. This makes it likely that these species specialized in smaller prey that could be pinned using only the inner toes, not requiring the feet to be as strong or sturdy.[4]

Dichotomy in body sizes

The most extreme examples of miniaturization and progenesis are found in Paraves.[15] The ancestors of Paraves first started to shrink in size in the early Jurassic 200 million years ago, and fossil evidence shows that this theropod line evolved new adaptations four times faster than other groups of dinosaurs,[16] and was shrinking 160 times faster than other dinosaur lineages were growing.[17] Turner et al. (2007) suggested that extreme miniaturization was ancestral for the clade, whose common ancestor has been estimated to have been around 65 centimeters long and 600-700 grams in mass. In Eumaniraptora, both Dromaeosauridae and Troodontidae went later through four independent events of gigantism, three times in dromaeosaurids and once in troodontids, while the body mass continued to decrease in many forms within Avialae.[18] Fossils show that all the earliest members of Paraves found to date started out as small, while Troodontidae and Dromaeosauridae gradually increased in size during the Cretaceous period.[18]

Evolution

Relationships

Pennaraptora

Oviraptorosauria

  Paraves    

Scansoriopterygidae

     

Xiaotingia

   

Yixianosaurus

             

Pedopenna

   

Aurornis

       

Serikornis

   

Eosinopteryx

         

Anchiornis

Eumaniraptora    

Troodontidae

   

Dromaeosauridae

     

Avialae

            Cladogram following the results of a phylogenetic study by Lefèvreet al., 2017.[19]

Paraves is a branch-based clade defined to include all dinosaurs which are more closely related to birds than to oviraptorosaurs. The ancestral paravian is the earliest common ancestor of birds, dromaeosaurids, and troodontids which was not also ancestral to oviraptorosaurs. Paravians comprises three major sub-groups: Avialae, including Archaeopteryx and modern birds, as well as the dromaeosaurids and troodontids, which may or may not form a natural group.

The name 'Paraves' (Greek par(a):near + Latin aves, plural of avis:bird) was coined by Paul Sereno in 1997.[20] The clade was defined by Sereno in 1998 as a branch-based clade containing all Maniraptora closer to Neornithes (which includes all the birds living in the world today) than to Oviraptor.[21]

In 1997, a node-based clade called Eumaniraptora ("true maniraptorans") was named by Padian, Hutchinson and Holtz. They defined their clade to include only avialans and deinonychosaurs. Paraves and Eumaniraptora are generally considered to be synonyms, though a few phylogenetic studies suggest that the two groups have a similar but not identical content; Agnolín and Novas (2011) recovered scansoriopterygids and alvarezsaurids as paravians that were not eumaniraptorans,[22] while Turner, Makovicky and Norell (2012) recovered Epidexipteryx as the only known non-eumaniraptoran paravian.[1] A nearly identical definition, "the theropod group that includes all taxa closer to Passer than to Dromaeosaurus", was used by Agnolín and Novas (2013) for their clade Averaptora.[23]

Since the 1960s, the dromaeosaurids and troodontids have often been classified together in a group or clade named the Deinonychosauria, initially based primarily on the presence of a retractable second toe with sickle-claw (now also known to be present in some avialans). The name Deinonychosauria was coined by Ned Colbert and Dale Russell in 1969, and defined as a clade (all theropods closer to dromaeosaurids than to birds) by Jacques Gauthier in 1986. However, several more recent studies have cast doubt on the hypothesis that dromaeosaurids and troodontids were more closely related to each other than either was to birds, instead finding that troodontids were more closely related to birds than to dromaeosaurids.[24][25] Because Deinonychosauria was originally defined as all animals closer to dromaeosaurids than to birds without specific reference to troodontids, this would render Deinonychosauria a synonym of Dromaeosauridae.[25]

In 2015 Chatterjee created Tetrapterygidae in the second edition of his book The Rise of Birds: 225 Million Years of Evolution, where he included Microraptor, Xiaotingia, Aurornis, and Anchiornis; together they were proposed to be the sister group of the Avialae.[26] Paraves, Eumaniraptora and Averaptora are generally considered to be synonyms. Deinonychosauria is a synonym of Dromaeosauridae if troodontids are closer to Avialae as now seems likely. Tetrapterygidae is a polyphyletic grouping of four-winged basal paravian genera.

Origin and early evolution

Paravians diverged from other maniraptorans no later than 160 Mya; some 65 Mya later, the ancestors of modern birds diverged from other paravians at approximately 95 Mya.[27]

Other than the crown group of modern birds, which are direct descendants in the stem lineage of Paraves, there are no extant survivors or genetic material, so their entire phylogeny is inferred only from the fossil record.[28] The prototypical fossil is Archaeopteryx, of which 11 whole or partial examples have been found.[29]

See also

References

  1. ^ a b Alan Hamilton Turner, Peter J. Makovicky and Mark Norell (2012). "A review of dromaeosaurid systematics and paravian phylogeny". Bulletin of the American Museum of Natural History. 371: 1–206. doi:10.1206/748.1. hdl:2246/6352.
  2. ^ a b Li, Rihui; Lockley, M.G.; Makovicky, P.J.; Matsukawa, M.; Norell, M.A.; Harris, J.D.; Liu, M. (2007). "Behavioral and faunal implications of Early Cretaceous deinonychosaur trackways from China". Naturwissenschaften. 95 (3): 185–91. Bibcode:2008NW.....95..185L. doi:10.1007/s00114-007-0310-7. PMID 17952398.
  3. ^ Mayr, G. (Oct 2016). Avian Evolution: The Fossil Record of Birds and its Paleobiological Significance (1 ed.). p. Ch. 2. ISBN 978-1119020769.
  4. ^ a b c d e f g h Fowler, D.W.; Freedman, E.A.; Scannella, J.B.; Kambic, R.E. (2011). "The Predatory Ecology of Deinonychus and the Origin of Flapping in Birds". PLOS ONE. 6 (12): e28964. Bibcode:2011PLoSO...628964F. doi:10.1371/journal.pone.0028964. PMC 3237572. PMID 22194962.
  5. ^ Zanno, L.E.; Makovicky, P.J. (2011). "Herbivorous ecomorphology and specialization patterns in theropod dinosaur evolution". Proc Natl Acad Sci USA. 108 (1): 232–237. Bibcode:2011PNAS..108..232Z. doi:10.1073/pnas.1011924108. PMC 3017133. PMID 21173263.
  6. ^ 125-Million-Year-Old Biplanes: New Evidence Suggests the Earliest Bird Species Had Feathers on their Hind Limbs
  7. ^ The asymmetry of the carpal joint and the evolution of wing folding in maniraptoran theropod dinosaurs
  8. ^ Hu, Dongyu; Lianhi, Hou; Zhang, Lijun; Xu, Xing (2009). "A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus". Nature. 461 (7264): 640–643. Bibcode:2009Natur.461..640H. doi:10.1038/nature08322. PMID 19794491.
  9. ^ Xing, X.; Zhou, Z.; Wang, X.; Kuang, X.; Zhang, F.; Du, X. (2003). "Four-winged dinosaurs from China". Nature. 421 (6921): 335–340. Bibcode:2003Natur.421..335X. doi:10.1038/nature01342. PMID 12540892.
  10. ^ Xu, X.; Zhang, F. (2005). "A new maniraptoran dinosaur from China with long feathers on the metatarsus". Naturwissenschaften. 92 (4): 173–177. Bibcode:2005NW.....92..173X. doi:10.1007/s00114-004-0604-y. PMID 15685441.
  11. ^ Dececchi, T.A.; Larsson, H.C.E.; Habib, M.B. (2016). "The wings before the bird: an evaluation of flapping-based locomotory hypotheses in bird antecedents". PeerJ. 4: e2159. doi:10.7717/peerj.2159. PMC 4941780. PMID 27441115.
  12. ^ Longrich, N.R.; Currie, P.J. (2009). "A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America". PNAS. 106 (13): 5002–7. Bibcode:2009PNAS..106.5002L. doi:10.1073/pnas.0811664106. PMC 2664043. PMID 19289829.
  13. ^ Z., Csiki; Vremir, M.; Brusatte, S. L.; Norell, M. A. (2010). "An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania". Proceedings of the National Academy of Sciences of the United States of America. 107 (35): 15357–61. Bibcode:2010PNAS..10715357C. doi:10.1073/pnas.1006970107. PMC 2932599. PMID 20805514. Supporting Information
  14. ^ Manning, P.L.; Payne, D.; Pennicott, J.; Barrett, P.M.; Ennos, R.A. (2006). "Dinosaur killer claws or climbing crampons?". Biology Letters. 2 (1): 110–112. doi:10.1098/rsbl.2005.0395. PMC 1617199. PMID 17148340.
  15. ^ Bhullar, B.A.; et al. (2012). "Birds have paedomorphic dinosaur skulls". Nature. 487 (7406): 223–226. Bibcode:2012Natur.487..223B. doi:10.1038/nature11146. PMID 22722850.
  16. ^ Dinosaurs 'shrank' regularly to become birds
  17. ^ How Dinosaurs Shrank and Became Birds
  18. ^ a b Turner, Alan H.; Pol, Diego; Clarke, Julia A.; Erickson, Gregory M.; Norell, M. (2007). "A basal dromaeosaurid and size evolution preceding avian flight". Science. 317 (5843): 1378–1381. Bibcode:2007Sci...317.1378T. doi:10.1126/science.1144066. PMID 17823350.
  19. ^ Ulysse Lefèvre, Andrea Cau, Aude Cincotta, Dongyu Hu, Anusuya Chinsamy, François Escuillié & Pascal Godefroit (2017). A new Jurassic theropod from China documents a transitional step in the macrostructure of feathers. The Science of Nature, 104: 74 (advance online publication). doi:10.1007/s00114-017-1496-y
  20. ^ Sereno, P. C., 1997, "The origin and evolution of dinosaurs", Annual Review of Earth & Planetary Sciences 25:435- 489. (21)
  21. ^ Sereno, P. C., 1998, "A rationale for phylogenetic definitions, with application to the higher level taxonomy of Dinosauria", Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 210:41-83. (23)
  22. ^ Agnolín, Federico L.; Novas, Fernando E. (2011). "Unenlagiid theropods: are they members of the Dromaeosauridae (Theropoda, Maniraptora)?". Anais da Academia Brasileira de Ciências. 83 (1): 117–162. doi:10.1590/S0001-37652011000100008. PMID 21437379.
  23. ^ Federico L. Agnolín and Fernando E. Novas (2013). Avian ancestors. A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. SpringerBriefs in Earth System Sciences. pp. 1–96. doi:10.1007/978-94-007-5637-3. ISBN 978-94-007-5636-6.
  24. ^ Godefroit, Pascal; Cau, Andrea; Hu, Dong-Yu; Escuillié, François; Wu, Wenhao; Dyke, Gareth (2013). "A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds". Nature. 498 (7454): 359–362. Bibcode:2013Natur.498..359G. doi:10.1038/nature12168. PMID 23719374.
  25. ^ a b Mortimer, M. (2012): The Theropod Database: Phylogeny of Theropoda. Retrieved 2013-AUG-15.
  26. ^ Chatterjee, S. (2015). The rise of birds: 225 million years of evolution. Johns Hopkins University Press, 45-48.
  27. ^ Claramunt, S.; Cracraft, J. (Dec 2015). "A new time tree reveals Earth history's imprint on the evolution of modern birds". Sci Adv. 1 (11): e1501005. Bibcode:2015SciA....1E1005C. doi:10.1126/sciadv.1501005. PMC 4730849. PMID 26824065.
  28. ^ Toby, M; et al. (Feb 2005). "The quality of the fossil record of Mesozoic birds". Proc Biol Sci. 272 (1560): 289–294. doi:10.1098/rspb.2004.2923. PMC 1634967. PMID 15705554.
  29. ^ Foth, C.; et al. (Jul 2014). "New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers". Nature. 511 (7507): 79–82. Bibcode:2014Natur.511...79F. doi:10.1038/nature13467. PMID 24990749.

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Paraves: Brief Summary ( englanti )

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Paraves are a widespread group of theropod dinosaurs that originated in the Late Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and scansoriopterygids, the group also contains the avialans, among which are the over ten thousand species of living birds. Primitive members of Paraves are well known for the possession of an enlarged claw on the second digit of the foot, which was held off the ground when walking in some species.

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Paraves ( kastilia )

tarjonnut wikipedia ES

Paraves es un clado de dinosaurios terópodos manirraptores, que se originó en el Jurásico medio (hace aproximadamente 167 millones de años, en el Bathoniense) y continúa hasta el presente en la forma de las aves, teniendo distribución mundial.

Historia

Paraves fue creado y definido por Paul Sereno, en 1997 y 1998 respectivamente. Es un clado troncal que incluye a todos los terópodos cercanos a las aves, a estas pero excluye a los ovirraptorosaurianos.

Sistemática

Paraves es el clado más inclusivo que contiene al Passer domesticus (Linneo, 1758) pero no al Oviraptor philoceratops (Osborn, 1924). Corresponde a todos los manirraptores más emparentados con las aves que con el ovirráptor.

Cladograma según un estudio de Zhang y colaboradores de 2008:[1]

Paraves Avialae

Epidendrosaurus (=Scansoriopteryx)


Aves

Archaeopteryx




Jeholornis


Avebrevicauda

Sapeornis


Pygostylia

Confuciusornis


Ornithothoraces

Enantiornithes


Ornithurae

Yanornis




Hesperornis



Neornithes










Deinonychosauria Troodontidae

Sinovenator




Mei



Troodon




Dromaeosauridae

Unenlagia




Microraptoria

Bambiraptor



Microraptor



Eudromaeosauria

Velociraptor




Deinonychus




Dromaeosaurus









Otro cladograma alternativo según Turner, Makovicky & Norell (2012) excluyendo a Pyroraptor y Limenavis:[2]

Paraves

Epidexipteryx


Eumaniraptora Deinonychosauria

Troodontidae



Dromaeosauridae



Avialae

Archaeopteryx




Sapeornis




Jeholornis




Jixiangornis


Pygostylia

Confuciusornis




Enantiornithes


Euornithes

Patagopteryx




Hongshanornis




Yixianornis




Songlingornis




Yanornis















Cladograma según un estudio de Pascal Godefroit y colegas en 2013:[3]

Paraves

Scansoriopterygidae




Eosinopteryx


Eumaniraptora

Dromaeosauridae




Troodontidae




Avialae







Referencias

  1. Zhang, F.; Zhou, Z.; Xu, X.; Wang, X.; Sullivan, C. (2008). «A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers». Nature 455 (7216): 1105-1108. Bibcode:2008Natur.455.1105Z. PMID 18948955. doi:10.1038/nature07447.
  2. Alan Hamilton Turner, Peter J. Makovicky and Mark Norell (2012). «A review of dromaeosaurid systematics and paravian phylogeny». Bulletin of the American Museum of Natural History 371: 1-206.
  3. Pascal Godefroit, Andrea Cau, Hu Dong-Yu, François Escuillié, Wu Wenhao and Gareth Dyke (2013). «A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds». Nature. in press. doi:10.1038/nature12168.
  • Sereno, P. C., 1997, "The origin and evolution of dinosaurs", Annual Review of Earth & Planetary Sciences 25:435- 489. (21)
  • Sereno, P. C., 1998, "A rationale for phylogenetic definitions, with application to the higher level taxonomy of Dinosauria", Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 210:41-83. (23)
  • Xing, X., Zhou, Z., Wang, X., Kuang, X., Zhang, F., and Du, X. (2003). "Four-winged dinosaurs from China." Nature, 421: 335–340.
  • Turner, Alan H.; Pol, Diego; Clarke, Julia A.; Erickson, Gregory M.; and Norell, Mark.. (2007). " A basal dromaeosaurid and size evolution preceding avian flight". Science, 317: 1378-1381. doi:10.1126/science.1144066

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Paraves: Brief Summary ( kastilia )

tarjonnut wikipedia ES

Paraves es un clado de dinosaurios terópodos manirraptores, que se originó en el Jurásico medio (hace aproximadamente 167 millones de años, en el Bathoniense) y continúa hasta el presente en la forma de las aves, teniendo distribución mundial.

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Paraves ( Ranska )

tarjonnut wikipedia FR

Paraves est un groupe de dinosaures théropodes qui contient les oiseaux et leurs plus proches parents. Il a été défini par Paul Sereno comme étant le groupe de tous les maniraptoriens plus proches des Neornithes que d'Oviraptor[1].

Caractéristiques

 src=
Photo d'un Paraves actuel de la classe des oiseaux : le Touraco de Pauline.
 src=
Squelette d'un Paraves droméosauridé du genre Bambiraptor.

Selon une étude de 2012, une configuration à quatre ailes pourrait représenter l'état primitif des Paraves qui pouvaient utiliser cette caractéristique pour le vol plané, les oiseaux ayant perdu par la suite cet empennage postérieur[2].

Systématique

Liste des genres basaux et familles, d'après Godefroit et al. (2013) :

Phylogénie

Place au sein des Théropodes

Phylogénie simplifiée des groupes de théropodes, d'après Hendrickx et al., 2015[3] :

Theropoda

Coelophysoidea




Dilophosauridae



AverostraCeratosauria
TetanuraeMegalosauroidea
AvetheropodaAllosauroidea
Coelurosauria

Tyrannosauroidea




Compsognathidae



ManiraptoriformesOrnithomimosauria
Maniraptora

Alvarezsauroidea



Therizinosauria
PennaraptoraOviraptorosauria
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Phylogénie interne des Paraves

 src=
Squelette d'un Paraves droméosauridé
de l'espèce Microraptor gui avec les empreintes de ses plumes.
La barre horizontale bicolore mesure 5 centimètres.
 src=
Deinonychus : Griffe en faucille ou « griffe tueuse » redressée,
courante sur le deuxième orteil chez les Paraves basaux ainsi que chez les droméosauridés, les troodontidés et même certains Avialae basaux.
Ici chez le genre Deinonychus.

La phylogénie des Paraves n'est pas encre stabilisée. Un analyse phylogénétique complète, réalisée en 2017 par Ulysse Lefèvre et ses collègues[4], montrent des différences sensibles, surtout pour les Paraves basaux, avec la principale étude précédente réalisée en 2013 par P. Godefroit et ses collègues[5] (voir plus bas). Ce cladogramme de 2017, n’intègre pas la création de la famille des Anchiornithidae intervenue juste après[6] :

Paraves

Scansoriopterygidae




Xiaotingia



Yixianosaurus








Pedopenna



Aurornis





Serikornis



Eosinopteryx






Anchiornis


Eumaniraptora

Troodontidae



Dromaeosauridae




AvialaeOiseaux






Phylogénie des Paraves, d'après Pascal Godefroit et ses collègues en 2013[5]:

Paraves

Epidendrosaurus



Epidexipteryx





Eosinopteryx



Eumaniraptora

Dromaeosauridae


Averaptora

Troodontidae



AvialaeOiseaux







Voir aussi

Références taxinomiques

(en) Référence Paleobiology Database : Paraves Sereno, 1997

Annexes

Notes et références

Références

  1. (en) P. C. Sereno. 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences 25:435-489
  2. (en) X. Xing, Z. Zhou, X. Wang, X. Kuang, F. Zhang, X. Du, « Four-winged dinosaurs from China », Nature, vol. 421, no 6921,‎ 2003, p. 335–340 (DOI )
  3. (en) C. Hendrickx, S.A. Hartman et O. Mateus, « An Overview of Non- Avian Theropod Discoveries and Classification », PalArch’s Journal of Vertebrate Palaeontology, vol. 12, no 1,‎ 2015, p. 1–73 (lire en ligne).
  4. (en) Ulysse Lefèvre, Andrea Cau, Aude Cincotta, Dongyu Hu, Anusuya Chinsamy, François Escuillié et Pascal Godefroit, « A new Jurassic theropod from China documents a transitional step in the macrostructure of feathers », The Science of Nature, vol. 104, no 74,‎ 2017 (DOI , lire en ligne)
  5. a et b (en) Pascal Godefroit, Andrea Cau, Dong-Yu Hu, François Escuillié, Wenhao Wu et Gareth Dyke, « A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds », Nature, vol. 498, no 7454,‎ 2013, p. 359–362 (PMID , DOI , Bibcode )
  6. (en) C. Foth et O.W.M. Rauhut, « Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs », BMC Evolutionary Biology, vol. 17,‎ 2017, p. 236 (DOI , lire en ligne)
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Paraves: Brief Summary ( Ranska )

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Paraves est un groupe de dinosaures théropodes qui contient les oiseaux et leurs plus proches parents. Il a été défini par Paul Sereno comme étant le groupe de tous les maniraptoriens plus proches des Neornithes que d'Oviraptor.

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Paraves ( galicia )

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Paraves é un clado de dinosauros terópodos manirraptores, que se orixinaron no Xurásico medio (hai aproximadamente 167 millóns de anos, no Batoniano) e continua até o presente na forma das aves, tendo distribución mundial.

Historia

Paraves foi creado e definido por Paul Sereno, en 1997 e 1998 respectivamente. É un clado troncal que inclúe a todos os terópodos próximos ás aves, a este pero exclúe aos ovirraptorosaurianos.

Sistemática

Paraves é o clado máis inclusivo que contén ao Passer domesticus (Linneo, 1758) pero non ao Oviraptor philoceratops (Osborn, 1924). Corresponde a todos os manirraptores máis emparentados coas aves que co ovirráptor.

Notas

  • Sereno, P. C., 1997, "The origin and evolution of dinosaurs", Annual Review of Earth & Planetary Sciences 25:435- 489. (21)
  • Sereno, P. C., 1998, "A rationale for phylogenetic definitions, with application to the higher level taxonomy of Dinosauria", Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 210:41-83. (23)
  • Xing, X., Zhou, Z., Wang, X., Kuang, X., Zhang, F., and Du, X. (2003). "Four-winged dinosaurs from China." Nature, 421: 335–340.
  • Turner, Alan H.; Pol, Diego; Clarke, Julia A.; Erickson, Gregory M.; and Norell, Mark.. (2007). " A basal dromaeosaurid and size evolution preceding avian flight". Science, 317: 1378-1381. doi:10.1126/science.1144066

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Paraves: Brief Summary ( galicia )

tarjonnut wikipedia gl Galician

Paraves é un clado de dinosauros terópodos manirraptores, que se orixinaron no Xurásico medio (hai aproximadamente 167 millóns de anos, no Batoniano) e continua até o presente na forma das aves, tendo distribución mundial.

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Paraves ( Italia )

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I Paraves sono un diffuso gruppo di dinosauri teropodi originatisi durante il Giurassico superiore. Oltre agli estinti dromaeosauridi, troodontidi e scansoriopterygidi, il gruppo contiene anche gli avialae, clade che contiene anche le novemila specie di uccelli viventi.[1] I membri più primitivi di Paraves sono ben noti per avere un grande artiglio a falce sul secondo dito di ciascun piede, tenuto sollevato durante la locomozione in alcune specie.[2]

Classificazione

I Paraves comprendono due grandi sottogruppi: Avialae, che comprendono Jeholornis e gli uccelli veri e propri, e i deinonicosauri, che comprendono i dromeosauridi e i troodontidi. Queste ultime due famiglie potrebbero non rappresentare un gruppo monofiletico.

Il nome Paraves fu coniato da Paul Sereno nel 1997, e venne definito dallo stesso Sereno nel 1998 come un clade contenente tutti i Maniraptora più vicini ai neorniti (ovvero tutti gli uccelli moderni e i loro stretti parenti) che a Oviraptor. Sempre nel 1997, un clade definito Eumaniraptora venne descritto da Padian, Hutchinson e Holtz. Questo clade venne istituito per comprendere uccelli e deinonicosauri. Il termine Eumaniraptora è solitamente considerato un sinonimo di Paraves, anche se alcuni studi filogenetici suggeriscono che i due gruppi potrebbero essere stati simili ma non identici. Uno studio (Agnolín e Novas, 2011) ha proposto che gli scansoriotterigidi e gli alvarezsauridi erano paraviani ma non eumaniraptori, mentre un altro studio (Turner, Makovicky e Norell, 2012) ha ipotizzato che Epidexipteryx fosse l'unico paraviano non Eumaniraptora noto.

Il paraviano ancestrale, un animale ipotetico, è il primo antenato comune di uccelli, dromeosauridi e troodontidi, che non fosse anche ancestrale agli oviraptorosauri. Poco può essere detto con certezza di questo animale. Uno studio (Turner et al., 2007) ha suggerito che questo animale non poteva volare o planare, e che fosse probabilmente piccolo (forse 65 cm di lunghezza e 600 - 700 grammi di peso). Altri studi, tuttavia, hanno proposto esempi di paraviani antichi e basali dotati di quattro ali (Xu et al., 2003; Xu et al., 2005; Hu et al., 2009).

Di seguito è raffigurato il cladogramma che segue l'analisi filogenetica degli studi di Cau et al. (2015):[3]

Oviraptorosauria

Paraves

Scansoriopterygidae

     

Eosinopteryx

Eumaniraptora

Jinfengopteryx

   

Aurornis

   

Dromaeosauridae

   

Troodontidae

Avialae

Anchiornis

     

Archaeopteryx

   

Uccelli moderni e altri avialae

           

Note

  1. ^ Alan Hamilton Turner, Peter J. Makovicky and Mark Norell, A review of dromaeosaurid systematics and paravian phylogeny, in Bulletin of the American Museum of Natural History, vol. 371, 2012, pp. 1–206, DOI:10.1206/748.1.
  2. ^ Rihui Li, Lockley, M.G., Makovicky, P.J., Matsukawa, M., Norell, M.A., Harris, J.D. e Liu, M., Behavioral and faunal implications of Early Cretaceous deinonychosaur trackways from China , in Naturwissenschaften, vol. 95, n. 3, 2007, pp. 185–91, Bibcode:2008NW.....95..185L, DOI:10.1007/s00114-007-0310-7, PMID 17952398.
  3. ^ Andrea Cau, Tom Brougham e Darren Naish, The phylogenetic affinities of the bizarre Late Cretaceous Romanian theropod Balaur bondoc(Dinosauria, Maniraptora): Dromaeosaurid or flightless bird?, in PeerJ, vol. 3, 2015, pp. e1032, DOI:10.7717/peerj.1032, PMC 4476167, PMID 26157616.

Bibliografia

  • Sereno, P. C., 1997, "The origin and evolution of dinosaurs", Annual Review of Earth & Planetary Sciences 25:435- 489. (21)
  • Sereno, P. C., 1998, "A rationale for phylogenetic definitions, with application to the higher level taxonomy of Dinosauria", Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 210:41-83.
  • Xing, X.; Zhou, Z.; Wang, X.; Kuang, X.; Zhang, F.; Du, X. (2003). "Four-winged dinosaurs from China". Nature 421 (6921): 335–340.
  • Xu, X.; Zhang, F. (2005). "A new maniraptoran dinosaur from China with long feathers on the metatarsus". Naturwissenschaften 92 (4): 173–177.
  • Turner, Alan H.; Pol, Diego; Clarke, Julia A.; Erickson, Gregory M.; Norell, M. (2007). "A basal dromaeosaurid and size evolution preceding avian flight". Science 317 (5843): 1378–1381.
  • Hu, Dongyu; Lianhi, Hou; Zhang, Lijun; Xu, Xing (2009). "A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus.". Nature 461: 640–643.
  • Agnolín, Federico L.; Novas, Fernando E. (2011). "Unenlagiid theropods: are they members of the Dromaeosauridae (Theropoda, Maniraptora)?". Anais da Academia Brasileira de Ciências 83 (1): 117–162.
  • Alan Hamilton Turner, Peter J. Makovicky and Mark Norell (2012). "A review of dromaeosaurid systematics and paravian phylogeny". Bulletin of the American Museum of Natural History 371: 1–206.
  • Agnolín, F. L.; Novas, F. E. (2013). "Systematic Palaeontology". Avian Ancestors. SpringerBriefs in Earth System Sciences. p. 9. doi:10.1007/978-94-007-5637-3_3. ISBN 978-94-007-5636-6. edit
  • Pascal Godefroit, Andrea Cau, Hu Dong-Yu, François Escuillié, Wu Wenhao and Gareth Dyke (2013). "A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds". Nature. in press.

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Paraves: Brief Summary ( Italia )

tarjonnut wikipedia IT

I Paraves sono un diffuso gruppo di dinosauri teropodi originatisi durante il Giurassico superiore. Oltre agli estinti dromaeosauridi, troodontidi e scansoriopterygidi, il gruppo contiene anche gli avialae, clade che contiene anche le novemila specie di uccelli viventi. I membri più primitivi di Paraves sono ben noti per avere un grande artiglio a falce sul secondo dito di ciascun piede, tenuto sollevato durante la locomozione in alcune specie.

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Paraves ( puola )

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Paraves ( portugali )

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Subgrupos Ver texto

Paraves é um clado baseado em ramo definido para incluir todos os dinossauros que são mais estreitamente relacionados às aves do que aos Oviraptorosauria. Os paravianos compreendem dois grandes sub-grupos: Avialae, incluindo Jeholornis e as aves modernas, e o Deinonychosauria, que inclui os Dromaeosauridae e Troodontidae, que podem ou não formar um grupo natural.[1]

Referências

  1. Li, Rihui; Lockley, M.G.; Makovicky, P.J.; Matsukawa, M.; Norell, M.A.; Harris, J.D.; Liu, M. (2007). «Behavioral and faunal implications of Early Cretaceous deinonychosaur trackways from China». Naturwissenschaften. 95 (3): 185–91. PMID 17952398. doi:10.1007/s00114-007-0310-7
Ícone de esboço Este artigo sobre animais é um esboço. Você pode ajudar a Wikipédia expandindo-o.
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Paraves: Brief Summary ( portugali )

tarjonnut wikipedia PT

Paraves é um clado baseado em ramo definido para incluir todos os dinossauros que são mais estreitamente relacionados às aves do que aos Oviraptorosauria. Os paravianos compreendem dois grandes sub-grupos: Avialae, incluindo Jeholornis e as aves modernas, e o Deinonychosauria, que inclui os Dromaeosauridae e Troodontidae, que podem ou não formar um grupo natural.

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近鳥類 ( kiina )

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近鳥類学名Paraves)或直接稱為近鳥型,是個演化支,包含恐龍之中親緣關係接近於鳥類而離偷蛋龍下目較遠的所有物種,例如鳥翼類始祖鳥熱河鳥和現代的鳥類)以及恐爪龍下目馳龍科傷齒龍科

分類

近鳥類是由保羅·塞里諾(Paul Sereno)在1997年所提出[1]。保羅·塞里諾在1998年將近鳥類定義為一個演化支,包含手盜龍類中,親緣關係較接近今鳥亞綱,而離偷蛋龍較遠的所有物種[2]

同樣在1997年,凱文·帕迪恩(Kevin Padian)等人建立真手盜龍類演化支,範圍包含鳥類與恐爪龍下目。由於這兩個分類單元的範圍幾乎一致,較晚命名的真手盜龍類,常被認為是近鳥類的異名。少數種系發生學研究則認為這兩個分類單元範圍相近,但不完全一致。2011年的一份研究,提出擅攀鳥龍科阿瓦拉慈龍科屬於近鳥類,但不屬於真手盜龍類[3]。同年的另一份研究,則提出耀龍是唯一屬於近鳥類、但不屬於真手盜龍類的分類單元[4]

就理論上,近鳥類的祖先應是馳龍科、傷齒龍科、鳥翼類的最近共同祖先,而離偷蛋龍類較遠的物種。但目前仍不能建構出其祖先的面貌與特徵。在2003年與2005年、2009年,徐星等人提出近鳥類的祖先應該具有氣動性,能夠滑翔,可能是四翼恐龍,例如鳥翼類的足羽龍、馳龍科的小盜龍、傷齒龍科的近鳥龍[5][6][7]。在2007年,艾倫·透納等人則認為近鳥類的祖先不能滑翔或飛行,而體重應該非常小,身長大約65公分,體重介於600到700公克[8]。在2009年,徐星指出早期、原始近鳥類是可採四翼飛行的小型恐龍,包含:鳥翼類足羽龍馳龍科小盜龍、以及傷齒龍科近鳥龍[7]

演化關係

以下演化樹是根據徐星等人在2008年的研究,各演化支定義是採保羅·賽里諾(Paul Sereno)的2005年版本[9][10]

近鳥類 恐爪龍下目

傷齒龍科

   

馳龍科

    鳥翼類

擅攀鳥龍科

     

始祖鳥科

   

今鳥亞綱

       

以下演化樹來自於周忠和的2008年手盜龍類研究[9]

近鳥類 鳥翼類

樹息龍 (=擅攀鳥龍)

鳥綱

始祖鳥

     

熱河鳥

短尾鳥類

會鳥

尾綜骨鳥目

孔子鳥

鳥胸骨類

反鳥亞綱

扇尾亞綱

燕鳥

     

黃昏鳥

   

今鳥亞綱

                  恐爪龍下目 傷齒龍科

中國獵龍

     

寐龍

   

傷齒龍

      馳龍科    

半鳥

      小盜龍類

斑比盜龍

   

小盜龍

    真馳龍類

伶盜龍

     

恐爪龍

     

馳龍

               

以下演化樹來自於2012年的近鳥類、馳龍科種系發生學重新研究[4]

近鳥類

耀龍

真手盜龍類 恐爪龍下目

傷齒龍科

   

馳龍科

       

始祖鳥

     

會鳥

     

熱河鳥

     

吉祥鳥

尾綜骨鳥目

孔子鳥

     

反鳥亞綱

Euornithes

巴塔哥尼鳥

     

紅山鳥

     

義縣鳥

     

松嶺鳥

     

燕鳥

     

神翼鳥

今鳥亞綱

Iaceornis

   

魚鳥

     

黃昏鳥

   

浸水鳥

     

鳥綱

                               

在2011年命名的曉廷龍,是種非常類似始祖鳥的小型獸腳類恐龍。曉廷龍的發現,衝擊目前的主流親緣分支分類法研究,始祖鳥與某些鳥翼類可能屬於恐爪龍下目。需要發現更多的相關化石,才能確定始祖鳥是否屬於恐爪龍下目[11]。在2012年的郊狼龍研究,提出不同版本的分類,將始祖鳥歸類於鳥類,將近鳥龍歸類於傷齒龍科,而將曉庭龍歸類於馳龍科的最基礎型物種,而不是近鳥龍的近親[12][4]

相關條目

參考資料

  1. ^ Sereno, P. C., 1997, "The origin and evolution of dinosaurs", Annual Review of Earth & Planetary Sciences 25:435- 489. (21)
  2. ^ Sereno, P. C., 1998, "A rationale for phylogenetic definitions, with application to the higher level taxonomy of Dinosauria", Neues Jahrbuch für Geologie und Paläontologie Abhandlungen 210:41-83. (23)
  3. ^ Agnolín, Federico L.; Novas, Fernando E. Unenlagiid theropods: are they members of the Dromaeosauridae (Theropoda, Maniraptora)?. Anais da Academia Brasileira de Ciências. 2011, 83 (1): 117–162. doi:10.1590/S0001-37652011000100008.
  4. ^ 4.0 4.1 4.2 Alan Hamilton Turner, Peter J. Makovicky and Mark Norell. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 2012, 371: 1–206.
  5. ^ Xing, X.; Zhou, Z.; Wang, X.; Kuang, X.; Zhang, F.; Du, X. Four-winged dinosaurs from China. Nature. 2003, 421 (6921): 335–340. PMID 12540892. doi:10.1038/nature01342.
  6. ^ Xu, X.; Zhang, F. A new maniraptoran dinosaur from China with long feathers on the metatarsus. Naturwissenschaften. 2005, 92 (4): 173–177. Bibcode:2005NW.....92..173X. PMID 15685441. doi:10.1007/s00114-004-0604-y.
  7. ^ 7.0 7.1 Hu, Dongyu, Lianhi, Hou, Zhang, Lijun, Xu, Xing. (2009) A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus. Nature, Vol 461 1 October 2009, pp. 640-643. doi:10.1038/nature08322.
  8. ^ Turner, Alan H.; Pol, Diego; Clarke, Julia A.; Erickson, Gregory M.; Norell, M. A basal dromaeosaurid and size evolution preceding avian flight. Science. 2007, 317 (5843): 1378–1381. Bibcode:2007Sci...317.1378T. PMID 17823350. doi:10.1126/science.1144066.
  9. ^ 9.0 9.1 Zhang, F.; Zhou, Z.; Xu, X.; Wang, X.; Sullivan, C. A bizarre Jurassic maniraptoran from China with elongate ribbon-like feathers. Nature. 2008, 455 (7216): 1105–1108. Bibcode:2008Natur.455.1105Z. PMID 18948955. doi:10.1038/nature07447.
  10. ^ Sereno, P. C., McAllister, S., and Brusatte, S. L. (2005). "TaxonSearch: a relational database for suprageneric taxa and phylogenetic definitions." PhyloInformatics, 8: 1-21.[1]
  11. ^ Xu, Xing; You, Hailu; Du, Kai; Han, Fenglu. An Archaeopteryx-like theropod from China and the origin of Avialae. Nature. 2011, 475 (7357): 465–470. PMID 21796204. doi:10.1038/nature10288.
  12. ^ {Senter, P.; Kirkland, J. I.; Deblieux, D. D.; Madsen, S.; Toth, N. (2012). Dodson, Peter. ed. "New Dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the Evolution of the Dromaeosaurid Tail". PLoS ONE 7 (5): e36790. {doi|10.1371/journal.pone.0036790}}
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近鳥類: Brief Summary ( kiina )

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近鳥類(学名:Paraves)或直接稱為近鳥型,是個演化支,包含恐龍之中親緣關係接近於鳥類而離偷蛋龍下目較遠的所有物種,例如鳥翼類始祖鳥熱河鳥和現代的鳥類)以及恐爪龍下目馳龍科傷齒龍科

lisenssi
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维基百科作者和编辑
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wikipedia 中文维基百科