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Fossils explained 53
Titans of the skies: azhdarchid pterosaurs
Pterosaurs, the flying reptiles of the Mesozoic, often play second fiddle in popularity to their
contemporaries, the dinosaurs. Such treatment conceals the remarkable diversity and biology of this
group: not only were pterosaurs the first vertebrates to achieve powered flight, but they also existed
for 160 million years—longer than any other flying vertebrates. Named after the Uzbek mythical dragon
‘azhdarkho’, the Azhdarchidae are among the most enigmatic of all pterosaurs. As with most pterosaurs,
azhdarchid remains are rare, and their fossil record is largely represented by isolated bones or
incomplete skeletons. Despite the collection of azhdarchid fossils over the last 100 years, recognition of
these pterosaurs as a distinct group was not achieved until relatively recently. It is now clear that the
azhdarchids were a highly successful group that probably first appeared in the Early Cretaceous,
gradually spreading across the globe until the latest Cretaceous when they, as one of the last remaining
groups of pterosaurs, became extinct. Although most notable for achieving wingspans comparable with
light aircraft, other aspects of azhdarchid morphology and ecology make them not just aberrant animals
but also unusual pterosaurs.

Azhdarchid origins
Some controversy surrounds the ancestry of
pterosaurs, but most agree that they should be
included within Archosauria, the reptilian group that
includes crocodiles, dinosaurs and birds. Exactly
where they fall in this category is still debated
because of the highly specialised nature of even the
most basal pterosaurs: their highly modified skeletons
leave few clues to their ancestry. The first pterosaur
fossils are found in the late Triassic, and these basal
groups are typically characterized by long tails and
toothed jaws. These groups dominated pterosaur
evolution until the late Jurassic but ultimately yielded
control of the skies to a diverse group of derived, tailless pterosaurs: the Pterodactyloidea. This group
diversified in the Early Cretaceous and includes the

Mark Witton
Palaeobiology Research
Group, School of Earth and
Environmental Sciences,
University of Portsmouth,
Portstmouth PO1 3QL, UK.

Azhdarchidae as the last and most derived family to
evolve (Fig. 1). Cretaceous forms such as Tapejara and
Tupuxuara are perhaps the closest related taxa to the
azhdarchids, sharing similarities in their toothless
jaws, relatively short wings and subequal limbs.

Fig. 1. Relationships of the Pterodactyloidea (based on Unwin
2003—see Suggested reading). ‘Basal pterosaurs’ is used here to
refer to all pterosaur taxa basal to Pterodactyloidea. Although
pterosaur systematics are controversial, the Azhdarchidae are
frequently suggested to be the most derived clade within
Pterodactyloidea. Basal pterosaurs are represented by the skull of
Rhamphorhynchus, Ornithocheiroidea by Coloborhynchus,
Ctenochasmatoidea by Gnathostoma, Dsungaripteridae by
Germanodactylus, Tapejaridae by Tapejara, and the Azhdarchidae by

© Blackwell Publishing Ltd, Geology Today, Vol. 23, No. 1, January–February 2007


although controversial material from Tanzania may extend azhdarchid origins down into the Upper Jurassic of Africa. The earliest occurrences of azhdarchid fossils occur in Lower Cretaceous deposits of China and Brazil. No. Structures on the middle-neck vertebrae suggest a stiff neck with very limited articulation. azhdarchids walked on the distal end of their wing metacarpal (further supported by three small digits) and. Europe and North America (Fig. Unlike most other pterosaurs. January–February 2007 . torso and leg. Reported occurrences of azhdarchid fossils across the globe. Skeletal anatomy Fig. 34 Azhdarchid skeletons possess many features that clearly distinguish them from other pterosaurs (Fig. whilst the cranial condyle is reduced to a poorly developed. a highly diagnostic feature achieved through hyperelongation of the mid-series neck vertebrae. but remains in Australia and New Zealand indicate that azhdarchids were also present in southern regions by Late Cretaceous times. hemispherical structure allowing restricted movement of the skull. 3. may have held their © Blackwell Publishing Ltd. 3). When grounded. a large azhdarchid from Texas. robust metacarpal which in turn supports a relatively reduced wing finger. This finger supported the main flight membrane. with the radius and ulna leading to a long. 2). with a third set of membranes between the tail and hindlimbs. 1. 2. the torso is short (perhaps less than half the length of the skull) but strongly fused to anchor the flight muscles of the arms and legs. their location of origin is unclear. The skull is lightened by a large pneumatized foramen known as the nasoantorbital fenestra (within which the nostrils were placed) and reduced jaw muscles. the neck of azhdarchids articulates with the underside of the cranium and orientates the long axis of the skull. foramina lining the jaws of some taxa indicate that a bird-like horny beak was present. Geology Today. Restoration of the skeleton of Quetzalcoatlus. The wings comprise a short but powerfully built humerus. almost 90° to the shaft of the neck in some species. The neck in some taxa is almost twice as long as the head (making them the longest necks of any pterosaurs). Because the first well-preserved azhdarchid material occurs in geographically distant regions. Azhdarchids appear to have become more widespread in the Northern Hemisphere than the Southern. 23. with numerous finds from Upper Cretaceous rocks across Asia. Vol. In all pterosaurs the wing area is increased through additional membranes supported between the pteroid (a long bony shaft projecting from the wrist) and shoulder. Instead. As with all pterosaurs. a thin but responsive organ also attached to the forearm.FOSSILS Fig. Their jaws are straight and elongate (over 2 m long in some species) but possess no teeth. because of unusually long hindlimbs.

freakishly big individuals have almost no chance of entering the fossil record but are very likely to have existed in the same way that giant individuals are seen in modern animal populations. In the case of Arambourgiania. Estimates for Quetzalcoatlus with a 10– 11 m wingspan range from 70 to 250 kg. Equally. the largest estimate is figured here. despite its fragmentary nature. clearly enormous. although establishing pterosaur mass is controversial to the point where some workers claim it impossible.5 m off the ground—a height comparable with a mature Asian elephant. but no other pterosaur group obtained gigantic size as frequently as the azhdarchids. Azhdarchid wing configuration (demonstrated by Quetzalcoatlus—bottom left) compared to Pteranodon (bottom right). This contrasts with other large pterosaurs that had disproportionately long forelimbs. with one. but it is © Blackwell Publishing Ltd. 23. but an improved understanding of azhdarchid anatomy now favours the lower estimate. 4. An extensive pneumatic system and thin bone histology makes azhdarchid skeletons deceptively lightweight and. but their size increased steadily throughout the Cretaceous until they had the largest wingspans of any flying vertebrates (Fig. However. body almost parallel with the ground whilst standing and walking. it seems quite plausible that the azhdarchids were indeed the largest fliers of all time. 1. 4). 35 . forcing the body into a more erect posture whilst walking and perhaps limiting their terrestrial capability. suggesting a terrestrial prowess better than most pterosaurs. such was its size and peculiar features. Despite their enormous size. the probability of fossilization indicates that the azhdarchid individuals known to palaeontologists are very likely to be of average sizes: truly enormous. It was not until the 1970s and the discovery of more complete azhdarchid material in Upper Cretaceous deposits of Texas that the true size of the azhdarchids was appreciated. and very likely to the dissatisfaction of aeronautical engineers everywhere. The shorter. Geology Today. azhdarchids possess typical pterosaur features of hollow and extremely thin-walled bones. The discovery of Quetzalcoatlus prompted some aeronautical engineers to state that the genus represented the largest flying creature possible. Early forms held modest wingspans of 2. 5. new discoveries in Europe and a reappraisal of fossil material of the genus Arambourgiania hint at forms with wingspans of 12 m or more. Being so much larger than all other known flying animals. Their size was the cause of some confusion in the early twentieth century: an extremely elongate neck-vertebra discovered in Jordan in the 1920s (now known to belong to the azhdarchid Arambourgiania) was originally thought to be a pterosaur wing element. Initial estimates gave this giant a wingspan of somewhere between 11 and 21 m. Vol. January–February 2007 Fig. No. wingspans of 11–13 m are predicted. Such limb disproportions are not seen in azhdarchids however.5 m. Size ranges of azhdarchids. These sediments yielded three azhdarchid individuals belonging to the genus Quetzalcoatlus. The largest flying animals ever? Many pterosaurs grew to sizes far in excess of any living birds. When standing.FOSSILS Fig. the shoulder of Quetzalcoatlus would have stood some 2. most workers agree that even the largest azhdarchids would had weighed relatively little. broader wings of the azhdarchid are more akin to the statically soaring Andean condor (top left) than the dynamically soaring wandering albatross (top right). The fragmentary remains of many giant azhdarchids (such as Arambourgiania) mean that their sizes can only be estimated based on more complete material from other azhdarchid taxa.

No. As such. slopes and cliffs may have assisted azhdarchids in becoming airborne. This low aspect ratio has implications for other basic flight calculations. estimates of pterosaur wing loading are lower than flying vertebrates of equivalent dimensions. making for a less stable but more manoeuvrable configuration. flapping flight. A figure between 70 and 85 kg conforms more to our understanding of pterosaur anatomy and flight dynamics. but more data regarding pterosaur soft tissues is needed before these figures can be verified. In the air As seems to be the case with many fossil groups. manipulating air currents to travel with minimal expenditure of energy on flapping. 6). an idea of wing shape or aspect ratio (AR = wingspan2/wing area) can be ascertained through relative limb lengths (Fig. with flight modelling of large pterosaurs during the 1970s casting doubts over their flight capabilities. Such locomotion was probably possible: pterosaur trackways indicate that pterosaurs may have been competent terrestrial animals. Pterosaur aerodynamics were particularly criticised. but several features of azhdarchid skeletons indicate that their method of flight may have been different.FOSSILS Fig. 36 probable that the latter figure is far too high. suggesting that azhdarchids may have flown relatively slowly. Generally. Although smaller forms may have been capable of powered. 5). Shoulder and wing structures in pterosaurs (based on Frey et al. This possibility has led to thoughts that pterosaurs were active. The stable ‘top-decker’ configuration with the centre of gravity (indicated by crosses) below the wings (top) seen in ornithocheiroid pterosaurs. narrow wings (high aspect ratio). resulting in relatively short and broad wings (low aspect ratio). Wing loading (WL = mass/ wing area) is indicative of flying style: high wing loading is typical of faster flight whereas low wing loading suggests a gentler pace. the energy requirements for flapping flight in larger azhdarchids probably excluded the largest forms from this activity. the unique ‘middle-decker’ system of azhdarchids may have been a compromise © Blackwell Publishing Ltd. it seems unlikely that they would be mandatory for take-off. However. powerfully flying creatures able to become airborne regardless of weather or topographical conditions. 6. Because of their broad wings azhdarchids have lower wing loading values than other large pterosaurs. with either a headwind or a long drop from a cliff edge required to become airborne. 1. bats and flying insects possess methods of temperature regulation). palaeontologists did not portray an attractive picture of pterosaurs for much of the twentieth century. 23. Because the pterosaur wing incorporates both sets of limbs in its construction. Many large pterosaurs are thought to have been dynamic soarers akin to modern gulls and albatross. Most large pterosaurs had disproportionately long forelimbs compared to their hindlimbs and consequently have long. ‘Top-decker’ pterosaurs carried their centre of gravity beneath the wings for stable flight (akin to a modern cargo plane) whereas ‘bottom-deckers’ acted more like fighter planes with the wings below the centre of gravity. Geology Today. azhdarchids have relatively long hind limbs and truncated distal components of the wing finger. the characteristic azhdarchid ‘middle-decker’ configuration with the centre of gravity between the wings. necks and skulls. It was even suggested that if conditions were too adverse the pterosaurs would have to remain grounded as their clumsy flight capability would not have withstood the flight stresses of blustery conditions. Because of their size many azhdarchids probably required a brief run-up to assist with take-off. It was thought that take-off was particularly difficult. 2003—see Suggested reading). Vol. although there are no modern flying animals large enough to compare the biggest azhdarchids against. January–February 2007 . but instead forms a ‘middle decker’ construction with the glenoid positioned halfway up the shoulder girdle (Fig. Although headwinds. perhaps suggesting that pterosaurs controlled their core body temperatures in a manner similar to all modern actively flying animals (birds. with both quadrupedal galloping and bipedal running possibly used during take-off. and the unstable ‘bottom-decker’ configuration seen in tapejarid pterosaurs with the centre of gravity above the wings. as seen in larger species of modern birds. This attitude has now changed to one more understanding of this highly successful and diverse group: evidence from numerous pterosaur fossils indicates that many species had a hair-like integument covering their bodies. It is neither in the ‘topdecker’ configuration seen in ornithocheiroid pterosaurs or the ‘bottom-decker’ condition known from tapejarid pterosaurs. The position of the shoulder joint (glenoid) is also unusual in azhdarchids.

stiff necks and strongly downturned skulls of azhdarchids bear little resemblance to their analogues in modern dip-feeders and would surely impair their ability to feed in this manner. they probably did so over land. This strongly suggests a more land-based ecology for azhdarchids than other pterosaurs: from palaeoenvironmental analysis of azhdarchid bearing sediments. ranging from near-shore lagoons to shallow seaways. The neck also articulates with the skull at an angle slightly below horizontal. 7). smaller azhdarchids may have been capable of flapping flight as well as soaring. Most authors currently cite aerial fishing as the most likely method of azhdarchid feeding. However. nor do they possess the horizontally orientated skulls or short. Some controversy has surrounded the ecological significance of the apparent azhdarchid preference for terrestrial environments. it seems that azhdarchids like Quetzalcoatlus inhabited arid environments with few large bodies of water. whilst others such as Zhejiangopterus occupied wooded settings crossed with small rivers and streams. These birds manipulate thermal updrafts to gain high altitude. Vol. The unusual abundance of azhdarchid remains in continental deposits may support this notion. strong neck of skimming animals. a useful © Blackwell Publishing Ltd. plucking fish and other pelagic organisms from bodies of water by either dip feeding or skimming. Skim-feeding like that seen in the modern bird Rhyncops is also unlikely. they do conform to some types of extant birds. Quetzalcoatlus has been suggested to be a scavenger of large dinosaur carcasses. implying a different environmental preference for azhdarchids. No. Feeding habits The majority of pterosaur fossils are found in marine or coastal sediments. trans-continental migrations. azhdarchids are often associated with material derived from continental settings such as plants or dinosaur fossils. most are found in lucustrine deposits and. but many aspects of their skeleton imply that wading was certainly a potential feeding option. it seems unlikely that large azhdarchids were capable of long-distance migration over bodies of water as suggested by some authors—if azhdarchids did perform long.FOSSILS between stability and manoeuvrability. Azhdarchids may have not been dynamic soarers like gulls. suggesting that these lifestyles are not viable for azhdarchids. 7. Although many of these characters make azhdarchids unique amongst pterosaurs. but their wing loading and aspect ratios indicate that they may have been static soarers akin to modern vultures and storks. when found in marine deposits. Because of their lower masses and lower energy requirements to achieve flapping flight. flexible necks capable of curving ventrally to minimize drag and resistance when submerging their jaws into the water during feeding. Geology Today. holding the skull close to the ground. then slowly glide to the next thermal column. With a sub-horizontally held neck and ventrally-orientated occipital condyle the head is naturally directed downwards. Although some azhdarchids are found in such settings. modern avian scavengers and probers have flexible necks able to probe deep into carcasses or investigate narrow burrows. A Quetzalcoatlus seizes an unwary crocodile in a Cretaceous swamp. large azhdarchids probed for invertebrate infauna. the mandibles of azhdarchids do not show the extreme degree of lateral compression seen in modern skimmers. Other workers have argued that. although laterally compressed. The relatively long hindlimbs position the torso sub-horizontally. January–February 2007 Fig. particularly concerning their feeding habits. because of the richness of trace fossils and burrows in the Quetzalcoatlus horizon. Such a lifestyle explains many otherwise problematic anatomical features of azhdarchid skeletons. Modern birds that partake in dip feeding have short. The increased manoeuvrability offered by the ‘middle-decker’ shoulder configuration may have enabled large azhdarchids to control their position within thermals more efficiently. As such updrafts are not generated readily on open water. rigid neck of azhdarchids. 37 . A model for azhdarchid feeding that complies more with their anatomy is that azhdarchids acted in a similar manner to some modern storks. Because of its unusual habitat and in situ association with numerous sauropod dinosaur remains. 23. the structure of the neck also makes these interpretations problematic. extending the neck forward along a horizontal plane. obtaining most of their food when foraging terrestrially or wading (Fig. facing the long-axis of the skull essentially forwards. most large pterosaur remains are preserved in open marine or coastal sediments. Both these suggestions ignore the long. 1. Whether azhdarchids fed in this manner is controversial. The long.

-M. 1981. (eds). Langston. Ösi. These environments are less likely to be preserved in the geological record and do not provide many settings conducive to fossilization. D. as with storks. E. The length of the neck may have allowed azhdarchids to extend their feeding range into areas where they could otherwise not reach—such as the deeper waters of rivers or lakes. J. v. 2003. v. 23. Evolution and Palaeobiology of Pterosaurs. the latter idea seems most satisfactory. pp. Geology Today. 1998. First evidence of azhdarchid pterosaurs from the Late Cretaceous of Hungary.244. a group which appear to have gradually replaced pterosaurs in a variety of roles throughout the Cretaceous. Evolution and Palaeobiology of Pterosaurs. pp. v. Pi Press. Geological Society Special Publication. (eds).139–190. they clearly demonstrate that azhdarchids must have been fantastic sights to behold. The extinction of the azhdarchids and pterosaurs may be attributed to competition from birds. Geological Society Special Publication. pp.217. Frey. In: Buffetaut.16. Pterosaurs. and the status of Arambourgiania and Quetzalcoatlus. © Blackwell Publishing Ltd. Weishampel. & Martill. E.J.1–64. Abhandlungen.M.M. & Langston. on a theoretical level at least.376. Buchy. A dying breed? The azhdarchids are among some of the last pterosaurs known.W. Jr. Vol.. Azhdarchidae) from Late Cretaceous sediments of Big Bend National Park.222–231. pp. Scientific American. S.92–102. The disappearance of azhdarchids from the fossil record at the end of the Mesozoic is often attributed to their size: larger animals are more prone to extinction through slower rates of reproduction and higher energy requirements. v. C. Jr.A. & Frey. possibly hiding true Late Cretaceous pterosaur diversity. It is not inconceivable to imagine azhdarchids standing in shallow water or along riverbanks with open jaws suspended in the water column. Geological Society of America Special Publication. W. snagging fish and other forms as they swam between the jaws or perhaps even collecting prey whilst walking across plains and woodlands. Journal of Vertebrate Paleontology. J. by the latest Cretaceous.FOSSILS adaptation for large animals feeding at ground level. 2005. The Pterosaurs from Deep Time.267–274. New York. Kellner. A. No. Discovery of the holotype of the giant pterosaur Titanopteryx philadelphiae Arambourg 1959. As the continental environments that the azhdarchids 38 had apparently become well adapted-to changed at the termination of the Mesozoic it appears that birds—and not azhdarchids—possessed the necessary diversity and numbers to progress into the Tertiary. & Jianu. E. pp. v.-C. but their remains are rare). Locomotion and Palaeoecology of the Pterosaurs. v. Acta Palaeontologica Polonica. 1. Unwin. D. although we now only have fossils as evidence of their presence.50. In: Buffetaut. It has been suggested that the apparent decline of pterosaurs may merely be a taphonomic artefact. Posture. M. 1996.M.207. & Mazin. 2005. The elongate skull positions the jaw tips close to feeding level. It seems that pterosaur diversity declined towards the end of the Mesozoic and. 2003. a vast range of small vertebrates may have been eaten. perhaps instigated by a shift of pterosaur habit from marine environments towards continental settings. whilst the peculiar stiffness of the neck may have served to minimize the energy requirements to hold the head in feeding position. M. R. 2003. & Templin. Martill. On the phylogeny and evolutionary history of pterosaurs. and Mazin. pp. D. pp. D. Neues Jahrbuch für Geologie und Paläontologie. E. January–February 2007 . only the azhdarchids remained in any abundance (a few other pterosaur lineages may have been present at this time. No flying animals before or since the azhdarchids have managed to capture their size or grandeur.. Cranial remains of Quetzalcoatlus (Pterosauria. More work is needed before any of these hypotheses can be verified but. The straight nature of azhdarchid beak morphology suggests that they did not specialize in one food item and. Middleand bottom-decker Cretaceous pterosaurs: unique designs in active flying vertebrates. v. Suggestions for further reading Chatterjee.217. Unwin.M.777–787.57–76. A. W. D.M.B.

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