Azhdarchidae
Azhdarchids | |
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Reconstructed skeleton of Quetzalcoatlus northropi
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | †Pterosauria |
Suborder: | †Pterodactyloidea |
Clade: | † Azhdarchiformes
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Family: | †Azhdarchidae Nesov, 1984 |
Type species | |
† Azhdarcho lancicollis Nesov, 1984
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Genera | |
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Synonyms | |
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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]
Description
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.[9]
It had been suggested azhdarchids were skimmers,[3][10] 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.[11][12][13][14][15] 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.[16] In another departure from typical azhdarchid lifestyles, the jaw of Alanqa may possibly be an adaptation to crushing shellfish and other hard foodstuffs.[17]
Azhdarchids are generally medium- to large-sized pterosaurs, with the largest achieving wingspans of 10–12 metres (33–39 ft),[18] but several small-sized species have recently been discovered.[19][20] 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.[21]
Systematics
Azhdarchids were originally classified as close relatives of
Taxonomy
Classification after Unwin 2006, except where noted.[22]
- Family Azhdarchidae
- Aerotitan
- Alanqa[23]
- Aralazhdarcho[24]
- Arambourgiania
- Azhdarcho
- Cretornis[25]
- Eurazhdarcho[26]
- Hatzegopteryx
- Mistralazhdarcho[27]
- Montanazhdarcho?[6]
- Palaeocursornis
- Phosphatodraco
- Quetzalcoatlus
- Volgadraco[28]
- Zhejiangopterus
- Possible valid azhdarchid genera
- Argentinadraco? (azhdarchoid, tentatively classified as an azhdarchid)[29]
- Bennettazhia
- Bogolubovia
- Navajodactylus?
Phylogeny
The most complete cladogram of azhdarchids is presented by Andres (2021):[30]
Azhdarchidae |
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In the analysis Cretornis and Volgadraco were recovered as pteranodontians, Alanqa was recovered as a thalassodromine, and Montanazhdarcho was recovered just outside Azhdarchidae.[30]
An alternate phylogeny of Azhdarchidae was presented by Ortiz David et al. (2022) in their description of Thanatosdrakon:[31]
Azhdarchidae | |
In this analysis, Alanqa is interpreted as a non-azhdarchid azhdarchoid closely related to Keresdrakon.[31]
References
- ^ S2CID 15172374.
- ^ Cat-Size Flying Reptile Shakes Up Pterosaur Family Tree
- ^ a b Nesov, L. A. (1984). "Upper Cretaceous pterosaurs and birds from Central Asia". Paleontologicheskii Zhurnal. 1984 (1): 47–57. Archived from the original on 2009-01-05.
- )
- S2CID 56002643. Archived from the original(PDF) on 2013-01-15.
- ^ a b Carroll, Nathan (2015). "Reassignment of Montanazhdarcho minor as a non-azhdarchid member of the Azhdarchoidea". Journal of Vertebrate Paleontology. 35: 104. Archived from the original on 2019-12-24. Retrieved 2021-01-21.
- S2CID 84617119.
- ISBN 978-0691150611.
- ^ Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.
- .
- ISBN 9780813723761.
- ^ Ősi, A.; Weishampel, D.B.; Jianu, C.M. (2005). "First evidence of azhdarchid pterosaurs from the Late Cretaceous of Hungary". Acta Palaeontologica Polonica. 50 (4): 777–787.
- PMID 17676976.
- PMID 18509539.
- ISBN 9789088900938.
- PMID 28133577.
- .
- PMID 21085624.
- PMID 27853614.
- S2CID 85673254.
- ^ "World's largest pterodactyl skeleton goes on show in Germany". The Local Germany. 2018-03-23.
- ISBN 0-13-146308-X.
- PMID 20520782.
- S2CID 128637719.
- ^ Averianov, A.O. (2010). "The osteology of Azhdarcho lancicollis Nessov, 1984 (Pterosauria, Azhdarchidae) from the Late Cretaceous of Uzbekistan" (PDF). Proceedings of the Zoological Institute of the Russian Academy of Sciences. 314 (3): 246–317.
- PMID 23382886.
- S2CID 91265861.
- S2CID 129558986.
- PMID 29166530.
- ^ S2CID 245078533.
- ^ S2CID 248140163. Retrieved 12 April 2022.
- 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. .
- Bennett, S. C. (2000). "Pterosaur flight: the role of actinofibrils in wing function". Historical Biology. 14 (4): 255–284. 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.