Azhdarchidae

Last updated

Azhdarchids
Temporal range: Cretaceous, 108–66  Ma
O
S
D
C
P
T
J
K
Pg
N
Possible Early Cretaceous record [1]
Quetzalcoatlus 1.JPG
Reconstructed skeleton of Quetzalcoatlus northropi
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Suborder: Pterodactyloidea
Clade: Azhdarchiformes
Family: Azhdarchidae
Nesov, 1984
Type species
Azhdarcho lancicollis
Nesov, 1984
Genera
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 included some of the largest known flying animals of all time, 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. By the time of the end-Cretaceous mass extinction, most pterosaur families except for the Azhdarchidae disappear from the fossil record, but recent studies indicate a wealth of pterosaurian fauna, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms. [4] In several analyses, some taxa such as Navajodactylus , Bakonydraco and Montanazhdarcho were moved from Azhdarchidae to other clades. [5] [6] [7]

Contents

Description

Artist Mark Witton's reconstruction of Hatzegopteryx hunting the ornithopod Zalmoxes. Hatzegopteryx.png
Artist Mark Witton's reconstruction of Hatzegopteryx hunting the ornithopod Zalmoxes .
Hatzegopteryx (A-B) compared with Arambourgiania (C) and Quetzalcoatlus (D-E). This illustrates the difference between the "blunt-beaked" azhdarchids and the "slender-beaked" forms. Hatzegopteryx-Witton-and-Naish-2017.png
Hatzegopteryx (A-B) compared with Arambourgiania (C) and Quetzalcoatlus (D-E). This illustrates the difference between the "blunt-beaked" azhdarchids and the "slender-beaked" forms.

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]

Systematics

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 .[ citation needed ]

Taxonomy

Classification after Unwin 2006, except where noted. [21]

Reconstructed feeding posture of an azhdarchid with sagittally aligned limbs. Azhdarchfeedingwittonnaish2008.png
Reconstructed feeding posture of an azhdarchid with sagittally aligned limbs.

Phylogeny

The most complete cladogram of azhdarchids is presented by Andres (2021): [29]

Azhdarchidae
Azhdarchinae

Azhdarcho

Albadraco

Aerotitan

Mistralazhdarcho

Quetzalcoatlinae

Phosphatodraco

Aralazhdarcho

Eurazhdarcho

Zhejiangopterus

Wellnhopterus

Cryodrakon

Hatzegopteryx

Arambourgiania

Quetzalcoatlus

In the analysis Cretornis and Volgadraco were recovered as pteranodontians, Alanqa was recovered as a thalassodromine, and Montanazhdarcho was recovered just outside Azhdarchidae. [29]

An alternate phylogeny of Azhdarchidae was presented by Ortiz David et al. (2022) in their description of Thanatosdrakon: [30]

Azhdarchidae

Eurazhdarcho

Phosphatodraco

Aralazhdarcho

Zhejiangopterus

Azhdarcho

Quetzalcoatlinae

Cryodrakon

Thanatosdrakon

Quetzalcoatlus

Hatzegopteryx

Albadraco

Arambourgiania

Mistralazhdarcho

Aerotitan

In this analysis, Alanqa is interpreted as a non-azhdarchid azhdarchoid closely related to Keresdrakon . [30]

Related Research Articles

<i>Quetzalcoatlus</i> Genus of azhdarchid pterosaurs from the Late Cretaceous

Quetzalcoatlus is a genus of pterosaur known from the Late Cretaceous period of North America ; its members were among the largest known flying animals of all time. Quetzalcoatlus is a member of the Azhdarchidae, a family of advanced toothless pterosaurs with unusually long, stiffened necks. Its name comes from the Aztec feathered serpent god Quetzalcoatl. The type species is Q. northropi, named by Douglas Lawson in 1975; the genus also includes the smaller species Q. lawsoni, which was known for many years as an unnamed species before being named by Brian Andres and Wann Langston Jr. (posthumously) in 2021.

<i>Azhdarcho</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Azhdarcho is a genus of azhdarchid pterosaur from the late Cretaceous Period of the Bissekty Formation of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan. It is known from fragmentary remains including the distinctive, elongated neck vertebrae that characterizes members of the family Azhdarchidae, a family that includes many giant pterosaurs such as Quetzalcoatlus. The name Azhdarcho comes from the Persian word azhdar (اژدر), a dragon-like creature in Persian mythology. The type species is Azhdarcho lancicollis. The specific epithet lancicollis is derived from the Latin words lancea and collum ("neck").

<i>Arambourgiania</i> Genus of large azhdarchid pterosaur from the Late Cretaceous

Arambourgiania is an extinct genus of azhdarchid pterosaur from the Late Cretaceous period of Jordan, and possibly the United States. Arambourgiania was among the largest members of its family, the Azhdarchidae, and it is also one of the largest flying animals ever known. The incomplete left ulna of the "Sidi Chennane azhdarchid" from Morocco may have also belonged to Arambourgiania.

<span class="mw-page-title-main">Pteranodontidae</span> Family of pteranodontian pterosaurs

The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head. The spectacularly-crested Nyctosaurus is sometimes included in this family, though usually placed in its own family, the Nyctosauridae.

<span class="mw-page-title-main">Tapejaridae</span> Family of azhdarchoid pterosaurs from the Cretaceous period

Tapejaridae are a family of pterodactyloid pterosaurs from the Cretaceous period. Members are currently known from Brazil, England, Hungary, Morocco, Spain, the United States, and China. The most primitive genera were found in China, indicating that the family has an Asian origin.

<i>Hatzegopteryx</i> Genus of large azhdarchid pterosaur from the Late Cretaceous

Hatzegopteryx is a genus of azhdarchid pterosaur found in the late Maastrichtian deposits of the Densuş Ciula Formation, an outcropping in Transylvania, Romania. It is known only from the type species, Hatzegopteryx thambema, named by Buffetaut et al. in 2002 based on parts of the skull and humerus. Additional specimens, including a neck vertebra, were later placed in the genus, representing a range of sizes. The largest of these remains indicate it was among the biggest pterosaurs, with an estimated wingspan of 10 to 12 metres.

<i>Montanazhdarcho</i> Genus of azhdarchoid pterosaur from the Late Cretaceous

Montanazhdarcho is a genus of azhdarchoid pterosaur from the Late Cretaceous period of what is now the state of Montana, United States. Montanazhdarcho is known from only one species, M. minor.

<i>Bennettazhia</i> Genus of tapejaromorph pterosaur from the Early Cretaceous

Bennettazhia is a genus of tapejaromorph pterosaur from the Early Cretaceous period of what is now the Hudspeth Formation of the state of Oregon in the United States. Although originally identified as a species of the pteranodontoid pterosaur Pteranodon, Bennettazhia is now thought to have been a different animal. The type and only species is B. oregonensis.

Eoazhdarcho is a genus of azhdarchoid pterodactyloid pterosaur named in 2005 by Chinese paleontologists Lü Junchang and Ji Qiang. The type and only known species is Eoazhdarcho liaoxiensis. The fossil was found in the Aptian-age Lower Cretaceous Jiufotang Formation of Chaoyang, Liaoning, China.

<i>Phosphatodraco</i> Late Cretaceous genus of pterosaur

Phosphatodraco is a genus of azhdarchid pterosaur that lived during the Late Cretaceous of what is now Morocco. In 2000, a pterosaur specimen consisting of five cervical (neck) vertebrae was discovered in the Ouled Abdoun Phosphatic Basin. The specimen was made the holotype of the new genus and species Phosphatodraco mauritanicus in 2003; the genus name means "dragon from the phosphates", and the specific name refers to the region of Mauretania. Phosphatodraco was the first Late Cretaceous pterosaur known from North Africa, and the second pterosaur genus described from Morocco. It is one of the only known azhdarchids preserving a relatively complete neck, and was one of the last known pterosaurs. Additional cervical vertebrae have since been assigned to the genus, and it has been suggested that fossils of the pterosaur Tethydraco represent wing elements of Phosphatodraco.

<i>Aralazhdarcho</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Aralazhdarcho is a genus of azhdarchid pterosaur from the Santonian to the early Campanian stages of the Late Cretaceous period of Bostobe Svita in Kazakhstan. The type and only known species is Aralazhdarcho bostobensis.

<span class="mw-page-title-main">Azhdarchoidea</span> Superfamily of ornithocheiroid pterosaurs

Azhdarchoidea is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, Tendaguripterus, that lived in the Late Jurassic period. The largest azhdarchoids include members of the family Azhdarchidae, examples of these are Quetzalcoatlus, Hatzegopteryx, and Arambourgiania. The Azhdarchoidea has been recovered as either closely related to the Ctenochasmatoidea, as the sister taxon of the Pteranodontoidea within the Ornithocheiroidea, or within the Tapejaroidea, which in turn was also within the Ornithocheiroidea.

<span class="mw-page-title-main">Pterosaur size</span>

Pterosaurs included the largest flying animals ever to have lived. They are a clade of prehistoric archosaurian reptiles closely related to dinosaurs. Species among pterosaurs occupied several types of environments, which ranged from aquatic to forested. Below are the lists that comprise the smallest and the largest pterosaurs known as of 2022.

<i>Alanqa</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Alanqa is a genus of pterodactyloid pterosaur from the Late Cretaceous period of what is now the Kem Kem Beds of southeastern Morocco. The name Alanqa comes from the Arabic word العنقاءal-‘Anqā’, for a mythical bird of Arabian culture.

<span class="mw-page-title-main">Thalassodrominae</span> Subfamily of azhdarchoid pterosaurs from the Cretaceous period

Thalassodrominae or Thalassodromidae is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional members come from Brazil, however, other possible members also come from other places, including the United States, Morocco, and Argentina. Thalassodrominae is considered either to be a subfamily within the pterosaur family Tapejaridae, or as a distinct family, Thalassodromidae, within the clade Neoazhdarchia, closely related to dsungaripterids or azhdarchids.

<i>Aerotitan</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Aerotitan is a genus of large azhdarchid pterosaur known from the Late Cretaceous period of what is now the Allen Formation of the Neuquén Basin in northern Patagonia, Argentina.

<i>Eurazhdarcho</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Eurazhdarcho is a genus of azhdarchid pterosaur from the Late Cretaceous period of what is now the Transylvanian Basin of Romania. Its fossil remains dated back 69 million years ago.

<i>Argentinadraco</i> Genus of azhdarchoid pterosaur from the Late Cretaceous

Argentinadraco is an extinct genus of azhdarchoid pterosaur from the Late Cretaceous Portezuelo Formation of Argentina. It contains a single species, A. barrealensis, named in 2017 by Alexander Kellner and Jorge Calvo. Argentinadraco is unusual for bearing a bottom jaw with a concave bottom edge, as well as a pair of ridges and depressions on the top surface. These features distinguish it from all other azhdarchoid groups, complicating its assignment, but it may belong to the Chaoyangopteridae. The ridges on the lower jaw may have been used to feed on small invertebrates in loose sediment within the system of lakes and rivers that it resided in.

<i>Wellnhopterus</i> Genus of azhdarchid pterosaur from the Late Cretaceous

Wellnhopterus is an azhdarchid pterosaur recovered from the Late Cretaceous Javelina Formation in Texas that was previously identified as a thalassodromine. It consists of a set of upper and lower jaws, as well as some cervical vertebrae and a fragmentary long bone. In July 2021, the jaws were given the genus name "Javelinadactylus", with the type and only species as "J. sagebieli"; however, this article has now been retracted. In a paper published in December 2021, the complete holotype was independently named Wellnhopterus, with the only species being W. brevirostris. As of 2022, this is the formal name of this pterosaur.

References

  1. 1 2 Dyke, G.; Benton, M.; Posmosanu, E.; Naish, D. (2010). "Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania". Palaeontology. 54: 79–95. doi:10.1111/j.1475-4983.2010.00997.x. S2CID   15172374.
  2. Cat-Size Flying Reptile Shakes Up Pterosaur Family Tree
  3. 1 2 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.
  4. Agnolin, Federico L. & Varricchio, David (2012). "Systematic reinterpretation of Piksi barbarulna Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of Western USA (Montana) as a pterosaur rather than a bird" (PDF). Geodiversitas. 34 (4): 883–894. doi:10.5252/g2012n4a10. S2CID   56002643. Archived from the original (PDF) on 2013-01-15.
  5. 1 2 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.
  6. Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103 (3–4): 383–398. doi:10.1017/S1755691013000303. S2CID   84617119.
  7. Wilton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. ISBN   978-0691150611.
  8. Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.
  9. Kellner, A. W. A.; Langston, W. (1996). "Cranial remains of Quetzalcoatlus (Pterosauria, Azhdarchidae) from Late Cretaceous sediments of Big Bend National Park, Texas". Journal of Vertebrate Paleontology. 16 (2): 222–231. doi:10.1080/02724634.1996.10011310.
  10. Chatterjee, S.; Templin, R. J. (2004). Posture, locomotion, and paleoecology of pterosaurs. GSA Special Papers. Vol. 376. pp. 1–64. doi:10.1130/0-8137-2376-0.1. ISBN   9780813723761.
  11. 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.
  12. Humphries, S.; Bonser, R.H.C.; Witton, M.P.; Martill, D.M. (2007). "Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method". PLOS Biology. 5 (8): e204. doi:10.1371/journal.pbio.0050204. PMC   1925135 . PMID   17676976.
  13. Witton, Mark P.; Naish, Darren; McClain, Craig R. (28 May 2008). "A Reappraisal of Azhdarchid Pterosaur Functional Morphology and Paleoecology". PLOS ONE. 3 (5): e2271. Bibcode:2008PLoSO...3.2271W. doi: 10.1371/journal.pone.0002271 . PMC   2386974 . PMID   18509539.
  14. Veldmeijer, Andre J.; Witton, Mark; Nieuwland, Ilja (2012). Pterosaurs. ISBN   9789088900938.
  15. Naish, D.; Witton, M.P. (2017). "Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators". PeerJ. 5: e2908. doi:10.7717/peerj.2908. PMC   5248582 . PMID   28133577.
  16. Martill, D.M.; Ibrahim, N. (2015). "An unusual modification of the jaws in cf. Alanqa, a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco" (PDF). Cretaceous Research. 53: 59–67. doi:10.1016/j.cretres.2014.11.001.
  17. Witton, M.P.; Habib, M.B. (2010). "On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness". PLOS ONE. 5 (11): e13982. Bibcode:2010PLoSO...513982W. doi: 10.1371/journal.pone.0013982 . PMC   2981443 . PMID   21085624.
  18. Martin-Silverstone, Elizabeth; Witton, Mark P.; Arbour, Victoria M.; Currie, Philip J. (2016). "A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants". Royal Society Open Science. 3 (8): 160333. Bibcode:2016RSOS....360333M. doi:10.1098/rsos.160333. PMC   5108964 . PMID   27853614.
  19. Prondvai, E.; Bodor, E. R.; Ösi, A. (2014). "Does morphology reflect osteohistology-based ontogeny? A case study of Late Cretaceous pterosaur jaw symphyses from Hungary reveals hidden taxonomic diversity" (PDF). Paleobiology. 40 (2): 288–321. doi:10.1666/13030. S2CID   85673254.
  20. "World's largest pterodactyl skeleton goes on show in Germany". The Local Germany. 2018-03-23.
  21. Unwin, David M. (2006). The Pterosaurs: From Deep Time. New York: Pi Press. p. 273. ISBN   0-13-146308-X.
  22. Ibrahim, N.; Unwin, D.M.; Martill, D.M.; Baidder, L.; Zouhri, S. (2010). Farke, Andrew Allen (ed.). "A New Pterosaur (Pterodactyloidea: Azhdarchidae) from the Upper Cretaceous of Morocco". PLOS ONE. 5 (5): e10875. Bibcode:2010PLoSO...510875I. doi: 10.1371/journal.pone.0010875 . PMC   2877115 . PMID   20520782.
  23. Averianov, A.O. (2007). "New records of azhdarchids (Pterosauria, Azhdarchidae) from the late Cretaceous of Russia, Kazakhstan, and Central Asia". Paleontological Journal. 41 (2): 189–197. doi:10.1134/S0031030107020098. S2CID   128637719.
  24. 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.
  25. Vremir, M. T. S.; Kellner, A. W. A.; Naish, D.; Dyke, G. J. (2013). Viriot, Laurent (ed.). "A New Azhdarchid Pterosaur from the Late Cretaceous of the Transylvanian Basin, Romania: Implications for Azhdarchid Diversity and Distribution". PLOS ONE. 8 (1): e54268. Bibcode:2013PLoSO...854268V. doi: 10.1371/journal.pone.0054268 . PMC   3559652 . PMID   23382886.
  26. Romain Vullo; Géraldine Garcia; Pascal Godefroit; Aude Cincotta; Xavier Valentin (2018). "Mistralazhdarcho maggii, gen. et sp. nov., a new azhdarchid pterosaur from the Upper Cretaceous of southeastern France". Journal of Vertebrate Paleontology. 38 (4): (1)-(16). doi:10.1080/02724634.2018.1502670. S2CID   91265861.
  27. Averianov, A.O.; Arkhangelsky, M.S.; Pervushov, E.M. (2008). "A New Late Cretaceous Azhdarchid (Pterosauria, Azhdarchidae) from the Volga Region". Paleontological Journal. 42 (6): 634–642. doi:10.1134/S0031030108060099. S2CID   129558986.
  28. Kellner, A.W.A.; Calvo, J.O. (2017). "New azhdarchoid pterosaur (Pterosauria, Pterodactyloidea) with an unusual lower jaw from the Portezuelo Formation (Upper Cretaceous), Neuquén Group, Patagonia, Argentina". Anais da Academia Brasileira de Ciências. 89 (3 suppl): 2003–2012. doi: 10.1590/0001-3765201720170478 . PMID   29166530.
  29. 1 2 Andres, Brian (2021-12-07). "Phylogenetic systematics of Quetzalcoatlus Lawson 1975 (Pterodactyloidea: Azhdarchoidea)". Journal of Vertebrate Paleontology. 41 (sup1): 203–217. doi:10.1080/02724634.2020.1801703. ISSN   0272-4634. S2CID   245078533.
  30. 1 2 Ortiz David, Leonardo D.; González Riga, Bernardo J.; Kellner, Alexander W. A. (12 April 2022). "Thanatosdrakon amaru, gen. ET SP. NOV., a giant azhdarchid pterosaur from the upper Cretaceous of Argentina". Cretaceous Research. 135: 105228. doi:10.1016/j.cretres.2022.105228. S2CID   248140163 . Retrieved 12 April 2022.