Reconstruction of the Early Cretaceous Jeholornis prima by Matt Martyniuk.

Belongs within: Maniraptora.
Contains: Ornithothoraces.

Though the specific definition has varied from author to author, the clade Avialae has always effectively referred to the total group of all 'birds', i. e. the union of living birds with other flying fossil dinosaurs. Basal members of the clade such as Archaeopteryx and Jeholornis retained a long skeletal tail. Eleven specimens of Archaeopterygidae are known from the Late Jurassic of Europe; some authors have assigned all these specimens to a single species Archaeopteryx lithographica whereas others have recognised multiple species.

More derived avialans in the clade Pygostylia had the tail reduced as in modern birds. This clade includes the Confuciusornithidae from the lower Cretaceous Yixian Formation of China, which are characterised by toothless jaws with a hooked rostrum (Padian 2004).

Synapomorphies (from Holtz 1998): Vertebral foramen to cranial articular facet ratio of dorsals 0.4 or greater; less than 25 caudal vertebrae present; glenoid oriented laterally; proximal ulnar shaft arched; preacetabular portion of ilium significantly longer than postacetabular portion; ischial proximodorsal process just distal to iliac process; ischiadic terminal processes separate; proximal region of fibular medial face flat; metatarsal IV subequal in length to metatarsal II; metatarsal I completely reverted.

<==Avialae [Archaeornithes, Palaeoaves, Sauriurae]
    |--Aurornis Godefroit, Cau et al. 2013 GC13
    |    `--*A. xui Godefroit, Cau et al. 2013 GC13
    `--+--Anchiornis huxleyi GC13
       `--+--Archaeopterygidae [Archaeopterygiformes, Archaeopteryginae] P04
          |    |--Wellnhoferia Elżanowski 2001 P04
          |    |    `--W. grandis Elżanowski 2001 P04
          |    `--Archaeopteryx von Meyer 1861 GC13, P04 (see below for synonymy)
          |         |--A. bavarica Wellnhofer 1993 P04
          |         `--A. lithographica von Meyer 1861 (see below for synonymy) P04
          `--+--Xiaotingia zhengi GC13
             `--+--+--Rahonavis Forster, Sampson et al. 1998 GC13, P04 (see below for synonymy)
                |  |    `--R. ostromi (Forster, Sampson et al. 1998) [=Rahona ostromi Forster, Sampson et al. 1998] P04
                |  `--Shenzhouraptor Zhou & Zhang 2002 GC13, D07
                |       `--S. sinensis D07
                `--+--Balaur GC13
                   |--Jeholornis Zhou & Zhang 2002 ZZ02
                   |    `--*J. prima Zhou & Zhang 2002 ZZ02
                   `--+--Sapeornis Zhou & Zhang 2002 GC13, P04
                      |    `--S. chaoyangensis Zhou & Zhang 2002 P04
                      `--Pygostylia P04
                           |--Ornithothoraces GC13
                           `--Confuciusornithidae [Confuciusornithiformes] P04
                                |--Changchengornis Ji, Chiappe & Ji 1999 P04
                                |    `--C. hengdaoziensis Hi, Chiappe & Ji 1999 P04
                                `--Confuciusornis Hou, Zhou et al. 1995 GC13, P04
                                     |--C. dui Hou, Martin et al. 1999 P04
                                     `--C. sanctus Hou, Zhou et al. 1995 (see below for synonymy) P04

Avialae incertae sedis:
  Songlingornis linghensis ZBH03
  Jibeinia Hou 2000 P04
    `--J. luahera Hou 2000 P04
  Wileyia valdensis Harrison & Walker 1973 (n. d.) P04
  Ilerdopteryx viai Lacasa-Ruiz 1991 (n. d.) P04
  ‘Ornithomimus’ minutus Marsh 1892 P04
  Platannavis nana Nessov 1992 (n. d.) P04
  Protoavis texensis Chatterjee 1991 (n. d.) P04
  Cuspirostrisornis Hou 1997 D07
    `--C. houi D07
  Yandangornis Cai & Zhao 1999 D07
    `--Y. longicaudatus D07

Archaeopteryx von Meyer 1861 GC13, P04 [incl. Archaeornis Petronievics in Petronievics & Woodward 1917 P04, Griphosaurus Wagner 1861 P04, Jurapteryx Howgate 1985 P04]

Archaeopteryx lithographica von Meyer 1861 [incl. Pterodactylus crassipes von Meyer 1857, Archaeopteryx crassipes, Rhamphorhynchus crassipes, Scaphognathus crassipes, Griphornis longicaudatus Woodward 1862, Griphosaurus longicaudatus, Archaeopteryx macrura Owen 1862, Archaeopteryx oweni Petronievics 1921, Griphosaurus problematicus Woodward 1862, Archaeopteryx recurva Howgate 1984, Jurapteryx recurva, Archaeopteryx siemensii Dames 1897, Archaeornis siemensi] P04

Confuciusornis sanctus Hou, Zhou et al. 1995 [incl. C. chuonzhous Hou 1997, C. suniae Hou 1997] P04

Rahonavis Forster, Sampson et al. 1998 GC13, P04 [incl. Rahona Forster, Sampson et al. 1998 P04; Rahonavidae, Rahonaviformes]

*Type species of generic name indicated


[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[GC13] Godefroit, P., A. Cau, Hu D.-Y., F. Escuillié, Wu W. & G. Dyke. 2013. A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds. Nature 498: 359–362.

Holtz, T. R., Jr. 1998. A new phylogeny of the carnivorous dinosaurs. Gaia 15: 5-61.

[P04] Padian, K. 2004. Basal Avialae. In: Weishampel, D. B., P. Dodson & H. Osmólska (eds) The Dinosauria 2nd ed. pp. 210–231. University of California Press: Berkeley.

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807–814.

[ZZ02] Zhou, Z., & F. Zhang. 2002. A long-tailed, seed-eating bird from the Early Cretaceous of China. Nature 418: 405–409.

Last updated: 6 August 2017.

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