Azhdarchidae: Difference between revisions

| fossil_range = [[Cretaceous]], {{fossilrange|10892|66|earliest=143}} <small>Possible [[Early Cretaceous]] record<ref name="Dykeetal"/></small>

'''Azhdarchidae''' (from the [[Persian language|Persian]] word ''{{lang|fa-Latn|[[Azhdahaazhdaha|azhdar]]''}}, ({{lang|fa|اژدر}}), a dragon-like creature in [[Persian mythology]]) is a [[Familyfamily (biology)|family]] of [[pterosaur]]s 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).<ref name="Dykeetal">{{cite journal | last1 = Dyke | first1 = G. | last2 = Benton | first2 = M. | last3 = Posmosanu | first3 = E. | last4 = Naish | first4 = D. | year = 2010 | title = Early Cretaceous (Berriasian) birds and pterosaurs from the Cornet bauxite mine, Romania | journal = Palaeontology | volume = 54| pages = 79–95| doi = 10.1111/j.1475-4983.2010.00997.x | s2cid = 15172374 | doi-access = free }}</ref> Azhdarchids includedare mainly known for including some of the largest known flying animals of all timediscovered, but smaller cat-size members have also been found.<ref>[https://web.archive.org/web/20160901013854/http://news.nationalgeographic.com/2016/08/cat-size-pterosaurs-dinosaurs-fossils-evolution-paleontology-science/ Cat-Size Flying Reptile Shakes Up Pterosaur Family Tree]</ref> Originally considered a sub-family of [[Pteranodon]]tidae, Nesov (1984)<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 |url-status = dead |archive-url=https://web.archive.org/web/20090105174323/http://www.azhdarcho.com/Art/Paleoart/azhdarch3.htm |archive-date=2009-01-05}}</ref> 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. ByPreviously theit timewas ofthought that by the end-Cretaceous massof extinctionthe Cretaceous, most pterosaur families except for the Azhdarchidae disappeardisappeared from the fossil record,<ref>{{Cite web |vauthors=Slack KE, Jones CM, Ando T, et al. |date=June 2006 |title=Early Penguin Fossils, Plus Mitochondrial Genomes, Calibrate Avian Evolution |journal=Molecular Biology and Evolution |url=https://academic.oup.com/mbe/article/23/6/1144/1055321/Early-Penguin-Fossils-Plus-Mitochondrial-Genomes |access-date=2023-07-25 |volume=23 |issue=6 |pages=1144–1155 |publication-place=Molecular Biology and Evolution. 23 (6): 1144–55 |doi=10.1093/molbev/msj124|pmid=16533822 }}</ref> but recent studies indicate a wealth of pterosaurian fauna, including [[Pteranodontidae|pteranodontids]], [[Nyctosauridae|nyctosaurids]], [[Tapejaridae|tapejarids]] and several indeterminate forms.<ref>{{cite journal|url=http://www.mnhn.fr/museum/front/medias/publication/48099_g2012n4a10.pdf |title=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 |doi=10.5252/g2012n4a10 |journal=Geodiversitas |volume=34 |author1=Agnolin, Federico L. |author2=Varricchio, David |name-list-style=amp|issue=4 |pages=883–894 |year=2012 |s2cid=56002643 |url-status = dead|archive-url=https://web.archive.org/web/20130115164102/http://www.mnhn.fr/museum/front/medias/publication/48099_g2012n4a10.pdf |archive-date=2013-01-15 }}</ref> In several analyses, some taxa such as ''[[Navajodactylus]]'', ''[[Bakonydraco]]'' and ''[[Montanazhdarcho]]'' were moved from Azhdarchidae to other clades.<ref name="ReferenceA">{{Cite journal|last=Carroll|first=Nathan|date=2015|title=Reassignment of ''Montanazhdarcho minor'' as a non-azhdarchid member of the Azhdarchoidea|url=http://vertpaleo.org/PDFS/SVP-2015-Program-and-Abstract-Book-9-22-2015.aspx|journal=[[Journal of Vertebrate Paleontology]]|volume=35|pages=104|via=|access-date=2021-01-21|archive-date=2019-12-24|archive-url=https://web.archive.org/web/20191224042705/http://vertpaleo.org/PDFS/SVP-2015-Program-and-Abstract-Book-9-22-2015.aspx|url-status=dead}}</ref><ref name=LoneStarPterosaurs>{{Cite journal | last1 = Andres | first1 = B. | last2 = Myers | first2 = T. S. | doi = 10.1017/S1755691013000303 | title = Lone Star Pterosaurs | journal = Earth and Environmental Science Transactions of the Royal Society of Edinburgh | pages = 383–398 | year = 2013 | volume=103| issue = 3–4| s2cid = 84617119 }}</ref><ref>{{Cite book| title = Pterosaurs: Natural History, Evolution, Anatomy|author=Wilton, Mark P. |isbn=978-0691150611|year=2013|publisher=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 |url-status = dead|archive-url=https://web.archive.org/web/20090105174323/http://www.azhdarcho.com/Art/Paleoart/azhdarch3.htm |archive-date=2009-01-05}}</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 | journal = Journal of Vertebrate Paleontology | volume = 16 | issue = 2| pages = 222–231 | doi = 10.1080/02724634.1996.10011310| bibcode = 1996JVPal..16..222K }}</ref> but further research has cast doubt on this idea, demonstrating that azhdarchids lacked the necessary adaptations for a [[Rynchops|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 book | last1 = Chatterjee | first1 = S. | last2 = Templin | first2 = R. J. |title=Posture, locomotion, and paleoecology of pterosaurs | year = 2004 |series=GSA Special Papers | volume = 376 | pages = 1–64 | doi=10.1130/0-8137-2376-0.1| isbn = 9780813723761}}</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 | 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 |journal=PLOS Biology |volume=5 |issue=8 |page=e204 |doi=10.1371/journal.pbio.0050204 |pmid=17676976 |pmc=1925135 |doi-access=free }}</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|pmid=18509539|pmc=2386974|bibcode=2008PLoSO...3.2271W|doi-access=free}}</ref><ref>{{cite book|url=https://books.google.com/books?id=NVLkATBBtZ0C&q=foraged+on+the+ground+in+a+stork-+or+ground+hornbill-like+manner&pg=PA116|title=Pterosaurs|isbn=9789088900938|last1=Veldmeijer|first1=Andre J.|last2=Witton|first2=Mark|last3=Nieuwland|first3=Ilja|year=2012|publisher=Sidestone Press }}</ref> Most large azhdarchids probably fed on small prey, including hatchling and small dinosaurs; in an unusual modification of the azhdarchid bauplanbodyplan, 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 | pmid = 28133577 | journal = PeerJ | date = 2017 | page=e2908| pmc = 5248582 | doi-access = free }}</ref> In another departure from typical azhdarchid lifestyles, the jaw of ''[[Alanqa]]'' may possibly be an adaptation to crushing shellfish and other hard foodstuffs.<ref name="martillandibrahim2015">{{cite journal | last1 = Martill | first1 = D.M. | last2 = Ibrahim | first2 = N. | year = 2015 | title = An unusual modification of the jaws in cf. ''Alanqa'', a mid-Cretaceous azhdarchid pterosaur from the Kem Kem beds of Morocco | url = https://researchportal.port.ac.uk/portal/files/8204327/An_unusual_modification_of_the_jaws.pdf| journal = Cretaceous Research | volume = 53 | pages = 59–67 | doi = 10.1016/j.cretres.2014.11.001| bibcode = 2015CrRes..53...59M }}</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 | pmid=21085624 | pmc=2981443| bibcode = 2010PLoSO...513982W| doi-access = free }}</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 | journal = Royal Society Open Science | volume = 3 | issue = 8| page = 160333 | doi = 10.1098/rsos.160333| pmid = 27853614 | pmc = 5108964 | bibcode = 2016RSOS....360333M}}</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 = http://real.mtak.hu/21860/1/Prondvai_et_al.2014_reposit1_u_110445.946242.pdf| journal = Paleobiology | volume = 40 | issue = 2| pages = 288–321 | doi = 10.1666/13030| bibcode = 2014Pbio...40..288P | s2cid = 85673254 }}</ref> 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 {{convert|12-20|m|abbr=on}}, although similarities to the contemporary azhdarchid ''[[Hatzegopteryx]]'' have also been noted.<ref>{{Cite news | url=https://www.thelocal.de/20180323/worlds-largest-pterodachtyl-dracula-museum-altmuehltal | title=World's largest pterodactyl skeleton goes on show in Germany| newspaper=The Local Germany| date=2018-03-23}}</ref>

** ''[[Aralazhdarcho]]''<ref name="averianov2007">{{cite journal | last1 = Averianov | first1 = A.O. | year = 2007 | title = New records of azhdarchids (Pterosauria, Azhdarchidae) from the late Cretaceous of Russia, Kazakhstan, and Central Asia | journal = Paleontological Journal | volume = 41 | issue = 2| pages = 189–197 | doi = 10.1134/S0031030107020098| bibcode = 2007PalJ...41..189A | s2cid = 128637719 | url = http://doc.rero.ch/record/16205/files/PAL_E3413.pdf }}</ref>

**''[[Volgadraco]]''<ref name=AAetal08>{{cite journal|doi=10.1134/S0031030108060099|last=Averianov|first=A.O.|author2=Arkhangelsky, M.S.|author3= Pervushov, E.M.|year=2008|title=A New Late Cretaceous Azhdarchid (Pterosauria, Azhdarchidae) from the Volga Region|journal=Paleontological Journal|volume=42|issue=6|pages=634–642|bibcode=2008PalJ...42..634A |s2cid=129558986}}</ref>

The most complete cladogram of azhdarchids is presented by Andres (2021):<ref name=":0">{{Cite journal|last=Andres|first=Brian|date=2021-12-07|title=Phylogenetic systematics of ''Quetzalcoatlus'' Lawson 1975 (Pterodactyloidea: Azhdarchoidea)|url=https://www.tandfonline.com/doi/pdf/10.1080/02724634.2020.1801703|journal=Journal of Vertebrate Paleontology|volume=41|issue=sup1|pages=203–217|doi=10.1080/02724634.2020.1801703|s2cid=245078533|issn=0272-4634|doi-access=free|bibcode=2021JVPal..41S.203A }}</ref>

|2=''[[Quetzalcoatlus]]'' [[File:Quetzalcoatlus07.jpg|50 px]] }} }} }} }} }} }} }} }}

An alternate phylogeny of Azhdarchidae was presented by Ortiz David ''et al.'' (2022) in their description of ''Thanatosdrakon'':<ref name=":1">{{cite journal |author1last1=Ortiz David |firstfirst1=Leonardo D. |last2=González Riga |first2=Bernardo J. |last3=Kellner |first3=Alexander W. A. |date=12 April 2022 |title=''Thanatosdrakon amaru'', gen. ET SP. NOV., a giant azhdarchid pterosaur from the upper Cretaceous of Argentina |url=https://www.sciencedirect.com/science/article/abs/pii/S0195667122000921 |journal=Cretaceous Research |volume=135 |page=105228 |doi=10.1016/j.cretres.2022.105228 |bibcode=2022CrRes.13705228O |s2cid=248140163 |access-date=12 April 2022}}</ref>

|2=''[[Quetzalcoatlus]]'' [[File:Quetzalcoatlus07.jpg|50 px]] }} }}

* {{cite journal | last1 = Bennett | first1 = S. C. | year = 2000 | title = Pterosaur flight: the role of actinofibrils in wing function | journal = Historical Biology | volume = 14 | issue = 4| pages = 255–284 | doi = 10.1080/10292380009380572| bibcode = 2000HBio...14..255B | s2cid = 85185457 }}

[[Category:Prehistoric reptilePterosaur families]]