Azhdarchidae

Azhdarchids Temporal range: Cretaceous, 92–66 Ma PreꞒ Ꞓ O S D C P T J K Pg N Possible Early Cretaceous record[1]
Reconstructed skeleton of Quetzalcoatlus northropi
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Order:	†Pterosauria
Suborder:	†Pterodactyloidea
Clade:	†Azhdarchiformes
Family:	†Azhdarchidae Nesov, 1984
Type species
†Azhdarcho lancicollis Nesov, 1984
Genera
†Alanqa? †Argentinadraco? †Montanazhdarcho? †Navajodactylus? †Palaeocursornis †Tethydraco? †Volgadraco? †Azhdarchinae †Aerotitan? †Albadraco? †Azhdarcho †Mistralazhdarcho? †Quetzalcoatlinae †Aralazhdarcho? †Arambourgiania †Cryodrakon †Eurazhdarcho? †Hatzegopteryx †Nipponopterus †Phosphatodraco? †Quetzalcoatlus †Thanatosdrakon †Wellnhopterus †Zhejiangopterus?
Synonyms
"Titanopterygiidae" Padian, 1984 (preoccupied)

Azhdarchidae (from the Persian word azhdar, اژدر, a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cretaceous as well (late Berriasian age, about 140 million years ago).^[1] Azhdarchids are mainly known for including some of the largest flying animals discovered, but smaller cat-size members have also been found.^[2] Originally considered a sub-family of Pteranodontidae, Nesov (1984)^[3] named the Azhdarchinae to include the pterosaurs Azhdarcho, Quetzalcoatlus, and Titanopteryx (now known as Arambourgiania). They were among the last known surviving members of the pterosaurs, and were a rather successful group with a worldwide distribution. Previously it was thought that by the end of the Cretaceous, most pterosaur families except for the Azhdarchidae disappeared from the fossil record,^[4] but recent studies indicate a wealth of pterosaurian fauna, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms.^[5] In several analyses, some taxa such as Navajodactylus, Bakonydraco and Montanazhdarcho were moved from Azhdarchidae to other clades.^[6][7][8]

Azhdarchids are characterized by their long legs and extremely long necks, made up of elongated neck vertebrae which are round in cross section. Most species of azhdarchids are still known mainly from their distinctive neck bones and not much else. The few azhdarchids that are known from reasonably good skeletons include Zhejiangopterus and Quetzalcoatlus. Azhdarchids are also distinguished by their relatively large heads and long, spear-like jaws. There are two major types of azhdarchid morphologies: the "blunt-beaked" forms with shorter and deeper bills and the "slender-beaked" forms with longer and thinner jaws.^[8]

It had been suggested azhdarchids were skimmers,^[3][9] but further research has cast doubt on this idea, demonstrating that azhdarchids lacked the necessary adaptations for a skim-feeding lifestyle, and that they may have led a more terrestrial existence similar to modern storks and ground hornbills.^{[10][11][12][13][14]} Most large azhdarchids probably fed on small prey, including hatchling and small dinosaurs; in an unusual modification of the azhdarchid bodyplan, the robust Hatzegopteryx may have tackled larger prey as the apex predator in its ecosystem.^[15] In another departure from typical azhdarchid lifestyles, the jaw of Alanqa may possibly be an adaptation to crushing shellfish and other hard foodstuffs.^[16]

Azhdarchids are generally medium- to large-sized pterosaurs, with the largest achieving wingspans of 10–12 metres (33–39 ft),^[17] but several small-sized species have recently been discovered.^[18][19] Another azhdarchid that is currently unnamed, recently discovered in Transylvania, may be the largest representative of the family thus far discovered. This unnamed specimen (nicknamed "Dracula" by paleontologists), currently on display in the Altmühltal Dinosaur Museum in Bavaria is estimated to have a wingspan of 12–20 m (39–66 ft), although similarities to the contemporary azhdarchid Hatzegopteryx have also been noted.^[20]

Azhdarchids were originally classified as close relatives of Pteranodon due to their long, toothless beaks. Others have suggested they were more closely related to the toothy ctenochasmatids (which include filter-feeders like Ctenochasma and Pterodaustro). Currently it is widely agreed that azhdarchids were closely related to pterosaurs such as Tupuxuara and Tapejara.^[21][22][23]

Classification after Averianov 2014, except where noted.^[21]

• Family Azhdarchidae
  • Aerotitan
  • Albadraco^[22]
  • Aralazhdarcho
  • Arambourgiania
  • Azhdarcho
  • Cryodrakon^[22]
  • Eurazhdarcho
  • Hatzegopteryx
  • Mistralazhdarcho^[24]
  • Nipponopterus^[25]
  • Palaeocursornis^[5]
  • Phosphatodraco
  • Quetzalcoatlus
  • Thanatosdrakon^[26]
  • Wellnhopterus^[22]
  • Zhejiangopterus

The following genera may belong in Azhdarchidae, but may alternatively represent non-azhdarchid pterosaurs:

• Alanqa^[21]
• Argentinadraco^[27]
• Bakonydraco^[21]
• Montanazhdarcho^[6]
• Navajodactylus^[28]
• Tethydraco^[29]
• Volgadraco^[21]

One of the most complete cladograms of azhdarchids is presented by Andres (2021):^[22]

In the analysis Cretornis and Volgadraco were recovered as pteranodontians, Alanqa was recovered as a thalassodromine, and Montanazhdarcho was recovered just outside Azhdarchidae.^[22]

The other most complete azhdarchid cladogram so far is that of Zhou et al. (2024):,^[25] based on Pêgas (2024):^[23]

• Astibia, H.; Buffetaut, E.; Buscalioni, A.D.; Cappetta, H.; Corral, C.; Estes, R.; Garcia-Garmilla, F.; Jaeger, Mazin; Jimenez-Fuentes, J.J.; Loeuff, J. Le; Mazin, J.M.; Orue-Etxebarria, X.; Pereda-Suberbiola, J.; Powell, J.E.; Rage, J.C.; Rodriguez-Lazaro, J.; Sanz, J.L.; Tong, H.; et al. (1991). "The fossil vertebrates from Lafio (Basque Country, Spain); new evidence on the composition and affinities of the Late Cretaceous continental fauna of Europe". Terra Nova. 2 (5): 460–466. doi:10.1111/j.1365-3121.1990.tb00103.x.
• Bennett, S. C. (2000). "Pterosaur flight: the role of actinofibrils in wing function". Historical Biology. 14 (4): 255–284. Bibcode:2000HBio...14..255B. doi:10.1080/10292380009380572. S2CID 85185457.
• Nesov, L.A. (1990). "Flying reptiles of the Jurassic and Cretaceous of the USSR and the significance of their remains for the reconstruction of palaeogeographic conditions". Bulletin of Leningrad University, Series 7, Geology and Geography (in Russian). 4 (28): 3–10.
• Nesov, L.A. (1991). "Giant flying reptiles of the family Azhdarchidae: 11. Environment, sedirnentological conditions and preservation of remains". Bulletin of Leningrad University, Series 7, Geology and Geography (in Russian). 3 (21): 16–24.