Open Paleo

Archaeopteryx

Meyer, 1861 AHR-kee-OP-ter-iks /ˌɑːrkiːˈɒptərɪks/

From Greek 'archaios' meaning 'ancient' and 'pteryx' meaning 'feather' or 'wing', meaning 'ancient wing'.

149.2–143.1 million years ago Tithonian, Late Jurassic
Late Jurassic stages

Archaeopteryx , sometimes referred to by its German name, "Urvogel" (lit. 'Primeval Bird') is a genus of bird-like dinosaurs. The genus name derives from the Ancient Greek ἀρχαῖος (archaîos), meaning 'ancient', and πτέρυξ (ptérux), meaning 'feather, wing'. Between the late 19th century and the early 21st century, Archaeopteryx was generally accepted by paleontologists and popular reference books as the oldest known bird (member of the group Avialae). Older potential avialans have since been identified, including Anchiornis, Xiaotingia, Aurornis, and Baminornis.

Under dispute

Archaeopteryx is traditionally the basalmost avialan (Turner and colleagues, 2012), or an avialan just crownward of the anchiornithines (Pei and colleagues, 2020), but a number of analyses instead recover it among the basal deinonychosaurs, close to the anchiornithids and troodontids (Xu and colleagues, 2011; Godefroit and colleagues, 2013; Hartman and colleagues, 2019). It is placed at Avialae following the prevailing recent view.

Diagnosis

  1. Jugal with a longitudinal groove on the medial side of the suborbital process.
  2. Infratemporal fenestra with a sharply angled incision at its anteroventral margin, separating the quadratojugal and postorbital processes of the jugal.
  3. Quadratojugal process of the jugal very slender, less than half the minimum height of the suborbital process.
  4. Depressed rim around the posterior margin of the trigeminal foramen in the prootic.
  5. Mandibular fenestra absent.
  6. Maxilla with only eight to nine tooth positions.
  7. 10 to 12 dentary teeth.
  8. Premaxillary and anteriormost dentary teeth with a straight basal part and a bulbous, strongly recurved apical part.
  9. Nine cervical and 14 dorsal vertebrae.
  10. Fewer than 25 caudal vertebrae.
  11. Pubic obturator foramen opens ventrally.
  12. Pubic apron divided along parts of the shaft.
  13. Ischium with a well-developed intermediate process posteriorly at about mid-shaft.
  14. Metatarsal III with a smooth distal end.

Species

Archaeopteryx lithographica Meyer, 1861

Type species

Named for the lithographic limestone of the Solnhofen deposits in Germany from which the specimens were recovered.

Neotype
BMNH 37001 , Natural History Museum
Material
Feather from the Solnhofen lithographic limestone; near-complete skeleton of a feathered animal from the lithographic limestone
Horizon
Altmühltal Formation, Tithonian, Late Jurassic , 149.2–143.1 million years ago
Locality
Ottmann quarry, Langenaltheimer Haardt, Bavaria, Germany

Synonyms

  • Jurapteryx junior subjective

    Senter (2007) demonstrated that the proposed diagnostic features could be explained by ontogenetic variation, treating it as a junior synonym of Archaeopteryx lithographica.

  • Wellnhoferia grandis junior subjective

    Named by Elżanowski (2001) on the Solnhofen specimen, which Mayr and colleagues (2007) concluded had not been convincingly shown to differ from the London specimen of A. lithographica. Foth and Rauhut (2026) place it within Archaeopteryx but leave species-level taxonomy unresolved.

Synonyms of the genus

Name Type Reason
Archaeornis junior subjective Originally described as a separate genus for the Berlin specimen; subsequent review treated it as another species of Archaeopteryx (A. siemensii).
Griphornis junior subjective Formally rejected by the ICZN in favor of Archaeopteryx.
Griphosaurus junior subjective Formally rejected by the ICZN in favor of Archaeopteryx.
Wellnhoferia junior subjective Erected by Elżanowski (2001) for the Solnhofen specimen on the basis of a shortened tail and a four-phalanx fourth pedal digit. Mayr and colleagues (2007) found the specimen not convincingly distinct from the London specimen, and the specimen-based phylogeny of Foth and Rauhut (2026) recovers it within Archaeopteryx.

Notable specimens

  • Thermopolis specimen

    WDC CSG 100 Wyoming Dinosaur Center exceptional preservation

    Has the best-preserved head and feet. Showed Archaeopteryx lacked a reversed toe, limiting perching, and confirmed a hyperextensible second toe, interpreted as evidence of theropod ancestry. Regarded as the most complete and best-preserved Archaeopteryx remains.

    Mayr et al. (2005)

  • Berlin specimen

    HMN 1880/81 Museum für Naturkunde, Berlin Found 1874 by Jakob Niemeyer most complete

    The most complete specimen and the first with a complete head. Its body plumage, including leg feathering, is the best researched of any Archaeopteryx. Basis for the name siemensii and a key reference for the genus.

    Dames (1884)

  • Chicago specimen

    FMNH PA 830 Field Museum of Natural History exceptional preservation

    The fourteenth specimen, described in 2025 by O'Connor and colleagues, preserving the first known tertials and other novel features. A second study the same year found oral papillae under UV light, a basihyal suggesting a mobile tongue, and jaw-tip foramina suggesting a bill-tip organ.

    O'Connor et al. (2025a)O'Connor et al. (2025b)

  • Solnhofen specimen

    BMMS 500 Bürgermeister-Müller-Museum largest individual

    The largest known specimen, missing only portions of the neck, tail, backbone, and head. Originally misidentified as Compsognathus, it became the holotype of a separate genus and species, Wellnhoferia grandis (Elżanowski, 2001), now treated as a junior synonym of Archaeopteryx.

    Wellnhofer (1988)Elżanowski (2001)Mayr et al. (2007)Foth & Rauhut (2026)

  • Munich specimen

    BSP 1999 I 50 Bayerische Staatssammlung für Paläontologie und Geologie Found 1992 exceptional preservation

    A nearly complete specimen missing only the front of the face. Basis for the proposed species A. bavarica; a possible cartilaginous sternum and its juvenile status have figured in debates over Archaeopteryx flight ability.

    Wellnhofer (1993)

References

  1. Meyer, H. V. (1861). [Die Feder von Solenhofen] [The feather of Solnhofen]. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde 1861: 678-679. meyer1861a
  2. Dames, W. (1884). Ueber Archaeopteryx. Palaeontologische Abhandlungen 2(3): 119-196. G. Reimer. dames1884a
  3. Wellnhofer, P. (1993). Das siebte Exemplar von Archaeopteryx aus den Solnhofener Schichten [The Seventh Specimen of Archaeopteryx from the Solnhofen Layers]. Archaeopteryx 11: 1-48. wellnhofer1993a
  4. Elżanowski, A. (2001). A new genus and species for the largest specimen of Archaeopteryx. Acta Palaeontologica Polonica 46(4): 519-532. elżanowski2001a
  5. Mayr, G.; Pohl, B.; Peters, D. S. (2005). A well-preserved Archaeopteryx specimen with theropod features. Science 310(5753): 1483-1486. doi:10.1126/science.1120331 mayr2005a
  6. O'Connor, J.; Clark, A.; Kuo, P. C.; Kiat, Y.; Fabbri, M.; Shinya, A.; Van Beek, C.; Lü, J.; Wang, M.; Hu, H. (2025). Chicago Archaeopteryx informs on the early evolution of the avian bauplan. Nature 641(8065): 1201-1207. doi:10.1038/s41586-025-08912-4 o'connor2025a
  7. O'Connor, J. K.; Clark, A. D.; Kuo, P.-C.; Wang, M.; Shinya, A.; Van Beek, C.; Chang, H. (2025). Avian features of Archaeopteryx feeding apparatus reflect elevated demands of flight. The Innovation 7(2): 101086. doi:10.1016/j.xinn.2025.101086

    Reports choanal papillae, an ossified basihyal and rostral neurovascular foramina in the Chicago specimen, the first evidence of these features in Archaeopteryx.

    o'connor2025b
  8. Jones, B. (2026). The Princeton Encyclopedia of Dinosaurs: Theropods. Princeton University Press.

    Source of the hip height estimate, and the appearance features.

    jones2026c
  9. Turner, A. H.; Makovicky, P. J.; Norell, M. A. (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

    Paravian phylogeny recovering Archaeopteryx as the basalmost avialan.

    turner2012a
  10. Pei, R.; Pittman, M.; Goloboff, P. A.; Dececchi, T. A.; Habib, M. B.; Kaye, T. G.; Larsson, H. C. E.; Norell, M. A.; Brusatte, S. L.; Xu, X. (2020). Potential for Powered Flight Neared by Most Close Avialan Relatives, but Few Crossed Its Thresholds. Current Biology 30(20): 4033-4046.e8. Elsevier BV. doi:10.1016/j.cub.2020.06.105

    Recovered most close avialan relatives, including Archaeopteryx, as near-flight-capable basal avialans.

    pei2020a
  11. Godefroit, P.; Demuynck, H.; Dyke, G.; Hu, D.; Escuillié, F.; Claeys, P. (2013). Reduced plumage and flight ability of a new Jurassic paravian theropod from China. Nature Communications 4(1). Springer Science and Business Media LLC. doi:10.1038/ncomms2389

    New paravian description recovering Archaeopteryx among basal avialan-grade paravians.

    godefroit2013a
  12. Hartman, S.; Mortimer, M.; Wahl, W. R.; Lomax, D. R.; Lippincott, J.; Lovelace, D. M. (2019). A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ 7: e7247. PeerJ. doi:10.7717/peerj.7247

    Recovered Archaeopteryx as a basal avialan in a broad paravian phylogeny.

    hartman2019a
  13. Xu, X.; You, H.; Du, K.; Han, F. (2011). An Archaeopteryx-like theropod from China and the origin of Avialae. Nature 475(7357): 465-470. Springer Science and Business Media LLC. doi:10.1038/nature10288

    Phylogenetic analysis shifted Archaeopteryx into Deinonychosauria near anchiornithids and troodontids.

    xu2011d
  14. Senter, P. (2007). A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology 5(4): 429-463. doi:10.1017/S1477201907002143 senter2007a
  15. Wellnhofer, P. (1988). A new specimen of Archaeopteryx. Science 240(4860): 1790-1792. American Association for the Advancement of Science. doi:10.1126/science.240.4860.1790

    Original description of the Solnhofen specimen, then the largest of six known skeletons.

    wellnhofer1988a
  16. Mayr, G.; Pohl, B.; Hartman, S.; Peters, D. S. (2007). The tenth skeletal specimen of Archaeopteryx. Zoological Journal of the Linnean Society 149(1): 97-116. Oxford University Press (OUP). doi:10.1111/j.1096-3642.2006.00245.x

    Found the Solnhofen specimen, holotype of Wellnhoferia grandis, not convincingly distinct from the London specimen.

    mayr2007a
  17. Foth, C.; Rauhut, O. W. M. (2026). Considerations on the taxonomy of Late Jurassic Avialae (Theropoda) from the Solnhofen Archipelago. Fossil Record 29(2): 731-758. Pensoft Publishers. doi:10.3897/fr.29.200245

    Source of the revised generic diagnosis; its specimen-based phylogeny recovers the Solnhofen specimen within Archaeopteryx.

    foth2026a
  18. O'Connor, J.; Marugán-Lobón, J. (2026). Evaluating variation in Solnhofen avialans. Biology Letters 22(3): 20250601. doi:10.1098/rsbl.2025.0601

    Morphometrics of all 15 Solnhofen avialans suggest a single taxon along a growth curve, referring the Haarlem specimen and Alcmonavis to Archaeopteryx.

    o'connor2026a