Azhdarchidae (from the
Persian
Persian may refer to:
* People and things from Iran, historically called ''Persia'' in the English language
** Persians, the majority ethnic group in Iran, not to be conflated with the Iranic peoples
** Persian language, an Iranian language of the ...
word , , a dragon-like creature in
Persian mythology
Persian mythology or Iranian mythology (Persian:اساطیرشناسی ایرانی) is the body of the myths originally told by ancient Persians and other Iranian peoples, and a genre of Ancient Persian folklore. These stories concern the origi ...
) is a
family
Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
pterosaur
Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 to ...
s known primarily from the
Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
Period, though an isolated vertebra apparently from an azhdarchid is known from the
Early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
as well (late
Berriasian
In the geological timescale, the Berriasian is an age/stage of the Early/Lower Cretaceous. It is the oldest subdivision in the entire Cretaceous. It has been taken to span the time between 145.0 ± 4.0 Ma and 139.8 ± 3.0 Ma (million years ago) ...
age, about 140 million years ago).
Azhdarchids included some of the largest known flying animals of all time, but smaller cat-size members have also been found. Originally considered a sub-family of
Pteranodon
''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
tidae, Nesov (1984)
named the Azhdarchinae to include the pterosaurs ''
Azhdarcho
''Azhdarcho'' is a genus of azhdarchid pterosaur from the late Cretaceous Period of the Bissekty Formation (middle Turonian stage, about 92 million years ago) of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan. It is known from ...
'', ''
Quetzalcoatlus
''Quetzalcoatlus'' is a genus of pterosaur known from the Late Cretaceous period of North America (Maastrichtian stage); its members were among the largest known flying animals of all time. ''Quetzalcoatlus'' is a member of the Azhdarchidae, ...
'', and ''Titanopteryx'' (now known as ''
Arambourgiania
''Arambourgiania'' is an extinct genus of azhdarchid pterosaur from the Late Cretaceous period (Maastrichtian stage) of Jordan, and possibly the United States.Harrell, T. Lynn Jr.; Gibson, Michael A.; Langston, Wann Jr. (2016). "A cervical ver ...
''). 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
Nyctosauridae (meaning "night lizards" or "bat lizards") is a Family (biology), family of specialized soaring pterosaurs of the late Cretaceous Period of North America, Africa, and possibly Europe. It was named in 1889 by Henry Alleyne Nicholson ...
,
tapejarids
Tapejaridae (from a Tupi word meaning "the old being") 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 primit ...
and several indeterminate forms. In several analyses, some taxa such as ''
Navajodactylus
''Navajodactylus'' (meaning "Navajo finger") is an extinct genus of pterodactyloid pterosaur from Late Cretaceous (late Campanian stage) deposits of the San Juan Basin in New Mexico, United States.
Discovery
The holotype specimen of ''Navajoda ...
'', ''
Bakonydraco'' and ''
Montanazhdarcho
''Montanazhdarcho'' is a genus of azhdarchoid pterosaur from the Late Cretaceous period (Campanian stage) of what is now the state of Montana, United States. ''Montanazhdarcho'' is known from only one species, ''M. minor''.
Discovery
The holo ...
'' were moved from Azhdarchidae to other clades.
Description
![Hatzegopteryx-Witton-and-Naish-2017](https://upload.wikimedia.org/wikipedia/commons/5/5a/Hatzegopteryx-Witton-and-Naish-2017.png)
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
''Quetzalcoatlus'' is a genus of pterosaur known from the Late Cretaceous period of North America (Maastrichtian stage); its members were among the largest known flying animals of all time. ''Quetzalcoatlus'' is a member of the Azhdarchidae, ...
''. 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.
It had been suggested azhdarchids were skimmers,
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
Storks are large, long-legged, long-necked wading birds with long, stout bills. They belong to the family called Ciconiidae, and make up the order Ciconiiformes . Ciconiiformes previously included a number of other families, such as herons an ...
s and
ground hornbill
The ground hornbills (Bucorvidae) are a family of the order Bucerotiformes, with a single genus ''Bucorvus'' and two extant species. The family is endemic to sub-Saharan Africa: the Abyssinian ground hornbill occurs in a belt from Senegal east ...
s.
Most large azhdarchids probably fed on small prey, including hatchling and small dinosaurs; in an unusual modification of the azhdarchid bodyplan, the robust ''
Hatzegopteryx
''Hatzegopteryx'' ("Hațeg basin wing") 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 ...
'' may have tackled larger prey as the
apex predator
An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own.
Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...
in its ecosystem.
In another departure from typical azhdarchid lifestyles, the jaw of ''
Alanqa
''Alanqa'' is a genus of pterodactyloid pterosaur from the Late Cretaceous period ( Cenomanian stage) of what is now the Kem Kem Beds of southeastern Morocco. The name ''Alanqa'' comes from the Arabic word العنقاء ''al- ‘Anqā’'', for ...
'' may possibly be an adaptation to crushing shellfish and other hard foodstuffs.
Azhdarchids are generally medium- to large-sized pterosaurs, with the largest achieving wingspans of ,
but several small-sized species have recently been discovered. Another azhdarchid that is currently unnamed, recently discovered in
Transylvania
Transylvania ( ro, Ardeal or ; hu, Erdély; german: Siebenbürgen) is a historical and cultural region in Central Europe, encompassing central Romania. To the east and south its natural border is the Carpathian Mountains, and to the west the Ap ...
, may be the largest representative of the family thus far discovered. This unnamed specimen (nicknamed "
Dracula
''Dracula'' is a novel by Bram Stoker, published in 1897. As an epistolary novel, the narrative is related through letters, diary entries, and newspaper articles. It has no single protagonist, but opens with solicitor Jonathan Harker taking ...
" by paleontologists), currently on display in the Altmühltal Dinosaur Museum in
Bavaria
Bavaria ( ; ), officially the Free State of Bavaria (german: Freistaat Bayern, link=no ), is a state in the south-east of Germany. With an area of , Bavaria is the largest German state by land area, comprising roughly a fifth of the total lan ...
is estimated to have a wingspan of , although similarities to the contemporary azhdarchid ''
Hatzegopteryx
''Hatzegopteryx'' ("Hațeg basin wing") 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 ...
'' have also been noted.
Systematics
Azhdarchids were originally classified as close relatives of ''
Pteranodon
''Pteranodon'' (); from Ancient Greek (''pteron'', "wing") and (''anodon'', "toothless") is a genus of pterosaur that included some of the largest known flying reptiles, with ''P. longiceps'' having a wingspan of . They lived during the late Cr ...
'' due to their long, toothless beaks. Others have suggested they were more closely related to the toothy
ctenochasmatids (which include filter-feeders like ''
Ctenochasma
''Ctenochasma'' (meaning "comb jaw") is a genus of Late Jurassic ctenochasmatid pterosaur belonging to the suborder Pterodactyloidea. Three species are currently recognized: ''C. roemeri'' (named after Friedrich Adolph Roemer), ''C. taqueti'', a ...
'' and ''
Pterodaustro
''Pterodaustro'' is a genus of ctenochasmatid pterodactyloid pterosaur from South America. Its fossil remains dated back to the Early Cretaceous period, about 105 million years ago. The most distinctive characteristic that separates ''Pterodaus ...
''). Currently it is widely agreed that azhdarchids were closely related to pterosaurs such as ''
Tupuxuara
''Tupuxuara'' is a genus of large, crested, and toothless Pterodactyloidea, pterodactyloid pterosaur from the Early Cretaceous period (Albian stage) of what is now the Romualdo Formation of the Santana Group, Brazil, about 125 to 112 million ye ...
'' and ''
Tapejara''.
Taxonomy
Classification after Unwin 2006, except where noted.
![Azhdarchfeedingwittonnaish2008](https://upload.wikimedia.org/wikipedia/commons/b/bc/Azhdarchfeedingwittonnaish2008.png)
* Family Azhdarchidae
**''
Aerotitan
''Aerotitan'' is a genus of large azhdarchid pterosaur known from the Late Cretaceous period (Maastrichtian stage) of what is now the Allen Formation of the Neuquén Basin in northern Patagonia, Argentina.
Discovery and naming
The type speci ...
''
** ''
Alanqa
''Alanqa'' is a genus of pterodactyloid pterosaur from the Late Cretaceous period ( Cenomanian stage) of what is now the Kem Kem Beds of southeastern Morocco. The name ''Alanqa'' comes from the Arabic word العنقاء ''al- ‘Anqā’'', for ...
''
** ''
Aralazhdarcho
''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''.
Etymology
''A ...
''
** ''
Arambourgiania
''Arambourgiania'' is an extinct genus of azhdarchid pterosaur from the Late Cretaceous period (Maastrichtian stage) of Jordan, and possibly the United States.Harrell, T. Lynn Jr.; Gibson, Michael A.; Langston, Wann Jr. (2016). "A cervical ver ...
''
**''
Azhdarcho
''Azhdarcho'' is a genus of azhdarchid pterosaur from the late Cretaceous Period of the Bissekty Formation (middle Turonian stage, about 92 million years ago) of Uzbekistan, as well as the Zhirkindek Formation of Kazakhstan. It is known from ...
''
**''
Cretornis''
**''
Eurazhdarcho
''Eurazhdarcho'' is a genus of azhdarchid pterosaur from the Late Cretaceous period (Maastrichtian stage) of what is now the Transylvanian Basin of Romania. Its fossil remains dated back 69 million years ago.
Discovery and naming
In 2009, M ...
''
**''
Hatzegopteryx
''Hatzegopteryx'' ("Hațeg basin wing") 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 ...
''
**''
Mistralazhdarcho
''Mistralazhdarcho'' is a genus of azhdarchid pterosaur from the Late Cretaceous period (Campanian stage) of France. The type and only species is ''Mistralazhdarcho maggii''.
Discovery
In 1992, Xavier Valentin at Velaux–La Bastide Neuve, in ...
''
**''
Montanazhdarcho
''Montanazhdarcho'' is a genus of azhdarchoid pterosaur from the Late Cretaceous period (Campanian stage) of what is now the state of Montana, United States. ''Montanazhdarcho'' is known from only one species, ''M. minor''.
Discovery
The holo ...
''?
**''
Palaeocursornis
''Palaeocursornis'' is a monotypic genus of pterosaurs. The only known species, ''P. corneti'', was described in 1984 based on a single bone ( MTCO-P 1637) interpreted as the distal part of a left femur, found in Early Cretaceous (Berriasian roc ...
''
**''
Phosphatodraco
''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. ...
''
**''
Quetzalcoatlus
''Quetzalcoatlus'' is a genus of pterosaur known from the Late Cretaceous period of North America (Maastrichtian stage); its members were among the largest known flying animals of all time. ''Quetzalcoatlus'' is a member of the Azhdarchidae, ...
''
**''
Volgadraco
''Volgadraco'' ("Volga River dragon") is a genus of pterodactyloid pterosaur from the Upper Cretaceous of European Russia. ''Volgadraco'' was originally classified as an azhdarchid. However, recent studies have concluded that it may belong to e ...
''
**''
Zhejiangopterus''
** Possible valid azhdarchid genera
*** ''
Argentinadraco
''Argentinadraco'' (meaning "Argentina dragon") 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 Jo ...
''? (azhdarchoid, tentatively classified as an azhdarchid)
***''
Bennettazhia''
***''
Bogolubovia
''Bogolubovia'' is a genus of pterosaur from the Upper Cretaceous (early Campanian) Rybushka Formation of Petrovsk, Saratov Oblast, Russia. It is named for Nikolai Nikolaevich Bogolubov, the paleontologist who discovered the remains in 1914. It w ...
''
***''
Navajodactylus
''Navajodactylus'' (meaning "Navajo finger") is an extinct genus of pterodactyloid pterosaur from Late Cretaceous (late Campanian stage) deposits of the San Juan Basin in New Mexico, United States.
Discovery
The holotype specimen of ''Navajoda ...
''?
Phylogeny
The most complete cladogram of azhdarchids is presented by Andres (2021):
In the analysis ''
Cretornis'' and ''
Volgadraco
''Volgadraco'' ("Volga River dragon") is a genus of pterodactyloid pterosaur from the Upper Cretaceous of European Russia. ''Volgadraco'' was originally classified as an azhdarchid. However, recent studies have concluded that it may belong to e ...
'' were recovered as
pteranodontians, ''
Alanqa
''Alanqa'' is a genus of pterodactyloid pterosaur from the Late Cretaceous period ( Cenomanian stage) of what is now the Kem Kem Beds of southeastern Morocco. The name ''Alanqa'' comes from the Arabic word العنقاء ''al- ‘Anqā’'', for ...
'' was recovered as a
thalassodromine, and ''
Montanazhdarcho
''Montanazhdarcho'' is a genus of azhdarchoid pterosaur from the Late Cretaceous period (Campanian stage) of what is now the state of Montana, United States. ''Montanazhdarcho'' is known from only one species, ''M. minor''.
Discovery
The holo ...
'' was recovered just outside Azhdarchidae.
An alternate phylogeny of Azhdarchidae was presented by Ortiz David ''et al.'' (2022) in their description of ''Thanatosdrakon'':
In this analysis, ''Alanqa'' is interpreted as a non-azhdarchid azhdarchoid closely related to ''
Keresdrakon
''Keresdrakon'' is an extinct genus of tapejaromorph pterosaur from the Goio-Erê Formation of Brazil, which dates back to the Late Cretaceous period (Turonian-Campanian stages), 94 to 71 million years ago. ''Keresdrakon'' contains a single spe ...
''.
References
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*
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*
{{Taxonbar, from=Q134363
Berriasian first appearances
Maastrichtian extinctions
Prehistoric reptile families