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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.<ref>Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.</ref>
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.<ref>Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.</ref>


It had been suggested azhdarchids were skimmers,<ref name="nesov1984">{{cite journal | last1 = Nesov | first1 = L. A. | year = 1984 | title = Upper Cretaceous pterosaurs and birds from Central Asia. | url = http://www.azhdarcho.com/Art/Paleoart/azhdarch3.htm| journal = Paleontologicheskii Zhurnal | volume = 1984 | issue = 1| pages = 47–57 }}</ref><ref name="kellner&langston1996">{{cite journal | last1 = Kellner | first1 = A. W. A. | last2 = Langston | first2 = W. | year = 1996 | title = Cranial remains of ''Quetzalcoatlus'' (Pterosauria, Azhdarchidae) from Late Cretaceous sediments of Big Bend National Park, Texas | url = | journal = Journal of Vertebrate Paleontology | volume = 16 | issue = 2| pages = 222–231 | doi = 10.1080/02724634.1996.10011310 }}</ref> 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 [[stork]]s and [[ground hornbill]]s.<ref name="chatterjee&templin2003">{{cite journal | last1 = Chatterjee | first1 = S. | last2 = Templin | first2 = R. J. | year = 2004 | title = Posture, locomotion, and paleoecology of pterosaurs | url = | journal = Geological Society of America Special Publication | volume = 376 | issue = | pages = 1–64 | doi=10.1130/0-8137-2376-0.1}}</ref><ref name="osietal2005">{{cite journal | last1 = Ősi | first1 = A. | last2 = Weishampel | first2 = D.B. | last3 = Jianu | first3 = C.M. | year = 2005 | title = First evidence of azhdarchid pterosaurs from the Late Cretaceous of Hungary | url = | journal = Acta Palaeontologica Polonica | volume = 50 | issue = 4| pages = 777–787 }}</ref><ref name="humphriesetal2007">{{cite journal|last1=Humphries |first1=S. |last2=Bonser |first2=R.H.C. |last3=Witton |first3=M.P. |last4=Martill |first4=D.M. |year=2007 |title=Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method |url=http://biology.plosjournals.org/archive/1545-7885/5/8/pdf/10.1371_journal.pbio.0050204-L.pdf |format=PDF |journal=PLoS Biology |volume=5 |issue=8 |page=e204 |doi=10.1371/journal.pbio.0050204 }}{{dead link|date=October 2016 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref name="Witton et al 2008">{{cite journal|last=Witton|first=Mark P.|author2=Naish, Darren|author3= McClain, Craig R.|title=A Reappraisal of Azhdarchid Pterosaur Functional Morphology and Paleoecology|journal=PLoS ONE|date=28 May 2008|volume=3|issue=5|pages=e2271|doi=10.1371/journal.pone.0002271|url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0002271|pmid=18509539|pmc=2386974}}</ref><ref>{{cite book|url=https://books.google.com/books?id=NVLkATBBtZ0C&pg=PA116&dq=foraged+on+the+ground+in+a+stork-+or+ground+hornbill-like+manner&hl=no&sa=X&ei=0w9TVPG1O8fsO82bgbgB&ved=0CB4Q6AEwAA#v=onepage&q=foraged%20on%20the%20ground%20in%20a%20stork-%20or%20ground%20hornbill-like%20manner&f=false|title=Pterosaurs|publisher=}}</ref> Bony protrusions in the jaw of ''[[Alanqa]]'' may possibly be an adaptation to crushing shellfish and other hard foodstuffs.<ref name="martillandibrahim2015">Martill, D.M., and Ibrahim, N. (2015). "An unusual modification of the jaws in cf. ''Alanqa'', a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco." ''Cretaceous Research'', '''53''': 59-67. {{doi|10.1016/j.cretres.2014.11.001}}</ref> ''[[Hatzegopteryx]]'' bears shorter and stronger jaws and neck than other azhdarchids; combined with its role as the largest carnivore in [[Hateg Island]], it might suggest a status as a macro-predator.<ref>Witton, Mark et al (2013). "Pterosaur overlords of Transylvania: short-necked giant azhdarchids in Late Cretaceous Romania". The Annual Symposium of Vertebrate Palaeontology and Comparative Anatomy</ref>
It had been suggested azhdarchids were skimmers,<ref name="nesov1984">{{cite journal | last1 = Nesov | first1 = L. A. | year = 1984 | title = Upper Cretaceous pterosaurs and birds from Central Asia. | url = http://www.azhdarcho.com/Art/Paleoart/azhdarch3.htm| journal = Paleontologicheskii Zhurnal | volume = 1984 | issue = 1| pages = 47–57 }}</ref><ref name="kellner&langston1996">{{cite journal | last1 = Kellner | first1 = A. W. A. | last2 = Langston | first2 = W. | year = 1996 | title = Cranial remains of ''Quetzalcoatlus'' (Pterosauria, Azhdarchidae) from Late Cretaceous sediments of Big Bend National Park, Texas | url = | journal = Journal of Vertebrate Paleontology | volume = 16 | issue = 2| pages = 222–231 | doi = 10.1080/02724634.1996.10011310 }}</ref> 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 [[stork]]s and [[ground hornbill]]s.<ref name="chatterjee&templin2003">{{cite journal | last1 = Chatterjee | first1 = S. | last2 = Templin | first2 = R. J. | year = 2004 | title = Posture, locomotion, and paleoecology of pterosaurs | url = | journal = Geological Society of America Special Publication | volume = 376 | issue = | pages = 1–64 | doi=10.1130/0-8137-2376-0.1}}</ref><ref name="osietal2005">{{cite journal | last1 = Ősi | first1 = A. | last2 = Weishampel | first2 = D.B. | last3 = Jianu | first3 = C.M. | year = 2005 | title = First evidence of azhdarchid pterosaurs from the Late Cretaceous of Hungary | url = | journal = Acta Palaeontologica Polonica | volume = 50 | issue = 4| pages = 777–787 }}</ref><ref name="humphriesetal2007">{{cite journal|last1=Humphries |first1=S. |last2=Bonser |first2=R.H.C. |last3=Witton |first3=M.P. |last4=Martill |first4=D.M. |year=2007 |title=Did pterosaurs feed by skimming? Physical modelling and anatomical evaluation of an unusual feeding method |url=http://biology.plosjournals.org/archive/1545-7885/5/8/pdf/10.1371_journal.pbio.0050204-L.pdf |format=PDF |journal=PLoS Biology |volume=5 |issue=8 |page=e204 |doi=10.1371/journal.pbio.0050204 }}{{dead link|date=October 2016 |bot=InternetArchiveBot |fix-attempted=yes }}</ref><ref name="Witton et al 2008">{{cite journal|last=Witton|first=Mark P.|author2=Naish, Darren|author3= McClain, Craig R.|title=A Reappraisal of Azhdarchid Pterosaur Functional Morphology and Paleoecology|journal=PLoS ONE|date=28 May 2008|volume=3|issue=5|pages=e2271|doi=10.1371/journal.pone.0002271|url=http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0002271|pmid=18509539|pmc=2386974}}</ref><ref>{{cite book|url=https://books.google.com/books?id=NVLkATBBtZ0C&pg=PA116&dq=foraged+on+the+ground+in+a+stork-+or+ground+hornbill-like+manner&hl=no&sa=X&ei=0w9TVPG1O8fsO82bgbgB&ved=0CB4Q6AEwAA#v=onepage&q=foraged%20on%20the%20ground%20in%20a%20stork-%20or%20ground%20hornbill-like%20manner&f=false|title=Pterosaurs|publisher=}}</ref> Most large azhdarchids probably fed on small prey, including hatchling and small dinosaurs; in an unusual modification of the azhdarchid bauplan, the unusually robust ''[[Hatzegopteryx]]'' may have tackled larger prey as the [[apex predator]] in its ecosystem.<ref name="witton2017">{{cite journal | first2 = M.P. | last2 = Witton | first1 = D. | last1 = Naish | title = Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators | volume = 5 | doi = 10.7717/peerj.2908 | journal = PeerJ | date = 2017 | url = https://peerj.com/articles/2908/}}</ref> In another departure from typical azhdarchid lifestyles, in the jaw of ''[[Alanqa]]'' may possibly be an adaptation to crushing shellfish and other hard foodstuffs.<ref name="martillandibrahim2015">Martill, D.M., and Ibrahim, N. (2015). "An unusual modification of the jaws in cf. ''Alanqa'', a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco." ''Cretaceous Research'', '''53''': 59-67. {{doi|10.1016/j.cretres.2014.11.001}}</ref>


Azhdarchids are generally large sized pterosaurs, but several small-sized species have recently been discovered.<ref>{{cite journal | last1 = Martin-Silverstone | first1 = Elizabeth | last2 = Witton | first2 = Mark P. | last3 = Arbour | first3 = Victoria M. | last4 = Currie | first4 = Philip J. | year = 2016 | title = A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants | url = | journal = Royal Society Open Science | volume = 3 | issue = 8| page = 160333 | doi = 10.1098/rsos.160333 }}</ref><ref>{{cite journal | last1 = Prondvai | first1 = E. | last2 = Bodor | first2 = E. R. | last3 = Ösi | first3 = A. | year = 2014 | title = Does morphology reflect osteohistology-based ontogeny? A case study of Late Cretaceous pterosaur jaw symphyses from Hungary reveals hidden taxonomic diversity | url = | journal = Paleobiology | volume = 40 | issue = 2| pages = 288–321 | doi = 10.1666/13030 }}</ref>
Azhdarchids are generally medium-to-large sized pterosaurs, with the largest achieving wingspans of {{convert|10|-|12|m}},<ref name=witton2010b>{{cite journal | last1 = Witton | first1 = M.P. | last2 = Habib | first2 = M.B. | date = 2010 | title = On the Size and Flight Diversity of Giant Pterosaurs, the Use of Birds as Pterosaur Analogues and Comments on Pterosaur Flightlessness | journal = PLoS ONE | volume = 5 | issue = 11 | page = e13982 | doi = 10.1371/journal.pone.0013982 | url = http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0013982}}</ref> but several small-sized species have recently been discovered.<ref>{{cite journal | last1 = Martin-Silverstone | first1 = Elizabeth | last2 = Witton | first2 = Mark P. | last3 = Arbour | first3 = Victoria M. | last4 = Currie | first4 = Philip J. | year = 2016 | title = A small azhdarchoid pterosaur from the latest Cretaceous, the age of flying giants | url = | journal = Royal Society Open Science | volume = 3 | issue = 8| page = 160333 | doi = 10.1098/rsos.160333 }}</ref><ref>{{cite journal | last1 = Prondvai | first1 = E. | last2 = Bodor | first2 = E. R. | last3 = Ösi | first3 = A. | year = 2014 | title = Does morphology reflect osteohistology-based ontogeny? A case study of Late Cretaceous pterosaur jaw symphyses from Hungary reveals hidden taxonomic diversity | url = | journal = Paleobiology | volume = 40 | issue = 2| pages = 288–321 | doi = 10.1666/13030 }}</ref>


==Systematics==
==Systematics==

Revision as of 18:00, 19 January 2017

Azhdarchids
Temporal range: Late Cretaceous, 108–66 Ma Possibly earlier
Reconstructed skeleton of Quetzalcoatlus northropi
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: †Pterosauria
Suborder: †Pterodactyloidea
Clade: †Azhdarchiformes
Family: †Azhdarchidae
Nesov, 1984
Type species
†Azhdarcho lancicollis
Nesov, 1984
Genera

See text

Synonyms

"Titanopterygiidae"
Padian, 1984 (preoccupied)

Azhdarchidae (from 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 members no larger than a cat have also been found.[2] Originally considered a sub-family of Pteranodontidae, Nesov (1984) 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 in pterosaurian faunas, including pteranodontids, nyctosaurids, tapejarids and several indeterminate forms.[3] Some taxa like Navajodactylus, Bakonydraco and Montanazhdarcho were moved from Azhdarchidae to other clades.[4][5][6]

Description

Artist's reconstruction of Hatzegopteryx hunting the ornithopod Zalmoxes.
Hatzegopteryx (A-B) compared with Arambourgiana (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.[7]

It had been suggested azhdarchids were skimmers,[8][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 bauplan, the unusually robust Hatzegopteryx may have tackled larger prey as the apex predator in its ecosystem.[15] In another departure from typical azhdarchid lifestyles, in 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]

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.

Taxonomy

Classification after Unwin 2006, except where noted.[20]

Reconstructed feeding posture of an azhdarchid with sagittally aligned limbs.

References

  1. ^ Dyke, G., Benton, M., Posmosanu, E. and Naish, D. (2010). "Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania." Palaeontology, published online before print 15 September 2010. doi:10.1111/j.1475-4983.2010.00997.x
  2. ^ Cat-Size Flying Reptile Shakes Up Pterosaur Family Tree
  3. ^ 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. Archived from the original (PDF) on 2013-01-15. {{cite journal}}: Unknown parameter |deadurl= ignored (|url-status= suggested) (help); Unknown parameter |lastauthoramp= ignored (|name-list-style= suggested) (help)
  4. ^ a b Carroll, N. REASSIGNMENT OF MONTANAZHDARCHO MINOR AS A NON-AZHDARCHID MEMBER OF THE AZHDARCHOIDEA, SVP 2015
  5. ^ Andres, B.; Myers, T. S. (2013). "Lone Star Pterosaurs". Earth and Environmental Science Transactions of the Royal Society of Edinburgh. 103: 1. doi:10.1017/S1755691013000303.
  6. ^ Wilton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. ISBN 0691150613.
  7. ^ Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.
  8. ^ Nesov, L. A. (1984). "Upper Cretaceous pterosaurs and birds from Central Asia". Paleontologicheskii Zhurnal. 1984 (1): 47–57.
  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". Geological Society of America Special Publication. 376: 1–64. doi:10.1130/0-8137-2376-0.1.
  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" (PDF). PLoS Biology. 5 (8): e204. doi:10.1371/journal.pbio.0050204.{{cite journal}}: CS1 maint: unflagged free DOI (link)[permanent dead link]
  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. doi:10.1371/journal.pone.0002271. PMC 2386974. PMID 18509539.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  14. ^ Pterosaurs.
  15. ^ Naish, D.; Witton, M.P. (2017). "Neck biomechanics indicate that giant Transylvanian azhdarchid pterosaurs were short-necked arch predators". PeerJ. 5. doi:10.7717/peerj.2908.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  16. ^ Martill, D.M., and Ibrahim, N. (2015). "An unusual modification of the jaws in cf. Alanqa, a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco." 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. doi:10.1371/journal.pone.0013982.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  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. doi:10.1098/rsos.160333.
  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". Paleobiology. 40 (2): 288–321. doi:10.1666/13030.
  20. ^ Unwin, David M. (2006). The Pterosaurs: From Deep Time. New York: Pi Press. p. 273. ISBN 0-13-146308-X.
  21. ^ 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. doi:10.1371/journal.pone.0010875. PMC 2877115. PMID 20520782.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  22. ^ 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.
  23. ^ a b 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.
  24. ^ 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: e54268. doi:10.1371/journal.pone.0054268. PMC 3559652. PMID 23382886.{{cite journal}}: CS1 maint: unflagged free DOI (link)
  25. ^ 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.
  • 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. doi:10.1080/10292380009380572.
  • 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 Universitv Series 7, Geology and Geography (in Russian). 3 (21): 16–24.
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