FOSSILS

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 yearslonger 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.
mark.witton@port.ac.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

2003see 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
Quetzalcoatlus.

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Fig. 2. Reported occurrences of

azhdarchid fossils across the
globe.

The earliest occurrences of azhdarchid fossils

occur in Lower Cretaceous deposits of China and
Brazil, although controversial material from Tanzania
may extend azhdarchid origins down into the Upper
Jurassic of Africa. Because the first well-preserved
azhdarchid material occurs in geographically distant
regions, their location of origin is unclear.
Azhdarchids appear to have become more widespread
in the Northern Hemisphere than the Southern, with
numerous finds from Upper Cretaceous rocks across
Asia, Europe and North America (Fig. 2), but remains
in Australia and New Zealand indicate that
azhdarchids were also present in southern regions by
Late Cretaceous times.

Skeletal anatomy
Fig. 3. Restoration of the
skeleton of Quetzalcoatlus, a
large azhdarchid from Texas.

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Azhdarchid skeletons possess many features that

clearly distinguish them from other pterosaurs
(Fig. 3). Their jaws are straight and elongate (over

2 m long in some species) but possess no teeth.

Instead, foramina lining the jaws of some taxa
indicate that a bird-like horny beak was present. 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 in some taxa is almost twice as long as the
head (making them the longest necks of any
pterosaurs); a highly diagnostic feature achieved
through hyperelongation of the mid-series neck
vertebrae. Unlike most other pterosaurs, the neck of
azhdarchids articulates with the underside of the
cranium and orientates the long axis of the skull,
almost 90 to the shaft of the neck in some species.
Structures on the middle-neck vertebrae suggest a
stiff neck with very limited articulation, whilst the
cranial condyle is reduced to a poorly developed,
hemispherical
structure
allowing
restricted
movement of the skull.
As with all pterosaurs, 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. The wings comprise a short but powerfully built
humerus, with the radius and ulna leading to a long,
robust metacarpal which in turn supports a relatively
reduced wing finger. This finger supported the main
flight membrane; a thin but responsive organ also
attached to the forearm, torso and leg. 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, with a third set of membranes between the
tail and hindlimbs. When grounded, azhdarchids
walked on the distal end of their wing metacarpal
(further supported by three small digits) and, because
of unusually long hindlimbs, may have held their

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Fig. 4. Size ranges of

azhdarchids. 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. In the case of
Arambourgiania, wingspans of
1113 m are predicted; the
largest estimate is figured here.

body almost parallel with the ground whilst standing

and walking. This contrasts with other large
pterosaurs that had disproportionately long forelimbs,
forcing the body into a more erect posture whilst
walking and perhaps limiting their terrestrial
capability. Such limb disproportions are not seen in
azhdarchids however, suggesting a terrestrial
prowess better than most pterosaurs.

The largest flying animals ever?

Many pterosaurs grew to sizes far in excess of any
living birds, but no other pterosaur group obtained
gigantic size as frequently as the azhdarchids. Early
forms held modest wingspans of 2.5 m, but their size
increased steadily throughout the Cretaceous until
they had the largest wingspans of any flying
vertebrates (Fig. 4). 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, such was its size and
peculiar features. 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. These sediments
yielded three azhdarchid individuals belonging to the
genus Quetzalcoatlus, with one, despite its
fragmentary nature, clearly enormous. Initial
estimates gave this giant a wingspan of somewhere
between 11 and 21 m, but an improved
understanding of azhdarchid anatomy now favours
the lower estimate. When standing, the shoulder of
Quetzalcoatlus would have stood some 2.5 m off the
grounda height comparable with a mature Asian
elephant. The discovery of Quetzalcoatlus prompted
some aeronautical engineers to state that the genus

represented the largest flying creature possible.

However, and very likely to the dissatisfaction of
aeronautical engineers everywhere, new discoveries
in Europe and a reappraisal of fossil material of the
genus Arambourgiania hint at forms with wingspans
of 12 m or more. Equally, the probability of
fossilization indicates that the azhdarchid individuals
known to palaeontologists are very likely to be of
average sizes: truly enormous, 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. Being so much larger than all
other known flying animals, it seems quite plausible
that the azhdarchids were indeed the largest fliers of
all time.
Despite their enormous size, azhdarchids possess
typical pterosaur features of hollow and extremely
thin-walled bones. An extensive pneumatic system
and thin bone histology makes azhdarchid skeletons
deceptively lightweight and, although establishing
pterosaur mass is controversial to the point where
some workers claim it impossible, most workers agree
that even the largest azhdarchids would had weighed
relatively little. Estimates for Quetzalcoatlus with a 10
11 m wingspan range from 70 to 250 kg, but it is

Blackwell Publishing Ltd, Geology Today, Vol. 23, No. 1, JanuaryFebruary 2007

Fig. 5. Azhdarchid wing

configuration (demonstrated by
Quetzalcoatlusbottom left)
compared to Pteranodon
(bottom right). The shorter,
broader wings of the azhdarchid
are more akin to the statically
soaring Andean condor (top left)
than the dynamically soaring
wandering albatross (top right).

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Fig. 6. Shoulder and wing

structures in pterosaurs (based
on Frey et al. 2003see
Suggested reading). The stable
top-decker configuration with
the centre of gravity (indicated
by crosses) below the wings
(top) seen in ornithocheiroid
pterosaurs; the characteristic
azhdarchid middle-decker
configuration with the centre of
gravity between the wings; and
the unstable bottom-decker
configuration seen in tapejarid
pterosaurs with the centre of
gravity above the wings.

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probable that the latter figure is far too high. 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,
palaeontologists did not portray an attractive picture
of pterosaurs for much of the twentieth century.
Pterosaur aerodynamics were particularly criticised,
with flight modelling of large pterosaurs during the
1970s casting doubts over their flight capabilities. It
was thought that take-off was particularly difficult,
with either a headwind or a long drop from a cliff
edge required to become airborne. 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. 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, necks and skulls, perhaps suggesting that
pterosaurs controlled their core body temperatures in
a manner similar to all modern actively flying
animals (birds, bats and flying insects possess
methods of temperature regulation). This possibility
has led to thoughts that pterosaurs were active,
powerfully flying creatures able to become airborne
regardless of weather or topographical conditions.
Because of their size many azhdarchids probably
required a brief run-up to assist with take-off, as seen
in larger species of modern birds. Such locomotion
was probably possible: pterosaur trackways indicate
that pterosaurs may have been competent terrestrial
animals, with both quadrupedal galloping and bipedal
running possibly used during take-off. Although
headwinds, slopes and cliffs may have assisted
azhdarchids in becoming airborne, it seems unlikely
that they would be mandatory for take-off.
Many large pterosaurs are thought to have been
dynamic soarers akin to modern gulls and albatross,

manipulating air currents to travel with minimal

expenditure of energy on flapping, but several
features of azhdarchid skeletons indicate that their
method of flight may have been different. Because the
pterosaur wing incorporates both sets of limbs in its
construction, an idea of wing shape or aspect ratio
(AR = wingspan2/wing area) can be ascertained
through relative limb lengths (Fig. 5). Most large
pterosaurs had disproportionately long forelimbs
compared to their hindlimbs and consequently have
long, narrow wings (high aspect ratio). However,
azhdarchids have relatively long hind limbs and
truncated distal components of the wing finger,
resulting in relatively short and broad wings (low
aspect ratio).
This low aspect ratio has implications for other
basic flight calculations. 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. Generally, estimates
of pterosaur wing loading are lower than flying
vertebrates of equivalent dimensions, although there
are no modern flying animals large enough to
compare the biggest azhdarchids against. Because of
their broad wings azhdarchids have lower wing
loading values than other large pterosaurs,
suggesting that azhdarchids may have flown
relatively slowly. Although smaller forms may have
been capable of powered, flapping flight, the energy
requirements for flapping flight in larger azhdarchids
probably excluded the largest forms from this activity.
The position of the shoulder joint (glenoid) is also
unusual in azhdarchids. It is neither in the topdecker configuration seen in ornithocheiroid
pterosaurs or the bottom-decker condition known
from tapejarid pterosaurs, but instead forms a middle
decker construction with the glenoid positioned
halfway up the shoulder girdle (Fig. 6). 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,
making for a less stable but more manoeuvrable
configuration. As such, the unique middle-decker
system of azhdarchids may have been a compromise

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between stability and manoeuvrability.

Although many of these characters make
azhdarchids unique amongst pterosaurs, they do
conform to some types of extant birds. Azhdarchids
may have not been dynamic soarers like gulls, but
their wing loading and aspect ratios indicate that
they may have been static soarers akin to modern
vultures and storks. These birds manipulate thermal
updrafts to gain high altitude, then slowly glide to the
next thermal column. 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, it
seems unlikely that large azhdarchids were capable of
long-distance migration over bodies of water as
suggested by some authorsif azhdarchids did
perform long, trans-continental migrations, they
probably did so over land. The unusual abundance of
azhdarchid remains in continental deposits may
support this notion; most large pterosaur remains are
preserved in open marine or coastal sediments,
implying a different environmental preference for
azhdarchids. Because of their lower masses and lower
energy requirements to achieve flapping flight,
smaller azhdarchids may have been capable of
flapping flight as well as soaring.

Feeding habits
The majority of pterosaur fossils are found in marine
or coastal sediments, ranging from near-shore
lagoons to shallow seaways. Although some
azhdarchids are found in such settings, most are
found in lucustrine deposits and, when found in
marine deposits, azhdarchids are often associated
with material derived from continental settings such
as plants or dinosaur fossils. This strongly suggests a
more land-based ecology for azhdarchids than other
pterosaurs: from palaeoenvironmental analysis of
azhdarchid bearing sediments, 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.
Some controversy has surrounded the ecological
significance of the apparent azhdarchid preference for
terrestrial environments, particularly concerning
their feeding habits. Because of its unusual habitat
and in situ association with numerous sauropod
dinosaur remains, Quetzalcoatlus has been suggested
to be a scavenger of large dinosaur carcasses. Other
workers have argued that, because of the richness of
trace fossils and burrows in the Quetzalcoatlus
horizon, large azhdarchids probed for invertebrate
infauna. Both these suggestions ignore the long, rigid

neck of azhdarchids; modern avian scavengers and

probers have flexible necks able to probe deep into
carcasses or investigate narrow burrows, suggesting
that these lifestyles are not viable for azhdarchids.
Most authors currently cite aerial fishing as the most
likely method of azhdarchid feeding, plucking fish and
other pelagic organisms from bodies of water by
either dip feeding or skimming. However, the
structure of the neck also makes these interpretations
problematic. Modern birds that partake in dip feeding
have short, flexible necks capable of curving ventrally
to minimize drag and resistance when submerging
their jaws into the water during feeding. The neck
also articulates with the skull at an angle slightly
below horizontal, facing the long-axis of the skull
essentially forwards. The long, 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. Skim-feeding like that seen in the modern
bird Rhyncops is also unlikely; although laterally
compressed, the mandibles of azhdarchids do not
show the extreme degree of lateral compression seen
in modern skimmers; nor do they possess the
horizontally orientated skulls or short, strong neck of
skimming animals.
A model for azhdarchid feeding that complies more
with their anatomy is that azhdarchids acted in a
similar manner to some modern storks, obtaining
most of their food when foraging terrestrially or
wading (Fig. 7). Such a lifestyle explains many
otherwise problematic anatomical features of
azhdarchid skeletons. The relatively long hindlimbs
position the torso sub-horizontally, extending the
neck forward along a horizontal plane, holding the
skull close to the ground. With a sub-horizontally
held neck and ventrally-orientated occipital condyle
the head is naturally directed downwards, a useful

Blackwell Publishing Ltd, Geology Today, Vol. 23, No. 1, JanuaryFebruary 2007

Fig. 7. A Quetzalcoatlus seizes

an unwary crocodile in a
Cretaceous swamp. Whether
azhdarchids fed in this manner is
controversial, but many aspects
of their skeleton imply that
wading was certainly a potential
feeding option.

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adaptation for large animals feeding at ground level.

The elongate skull positions the jaw tips close to
feeding level, whilst the peculiar stiffness of the neck
may have served to minimize the energy
requirements to hold the head in feeding position.
The length of the neck may have allowed azhdarchids
to extend their feeding range into areas where they
could otherwise not reachsuch as the deeper
waters of rivers or lakes. It is not inconceivable to
imagine azhdarchids standing in shallow water or
along riverbanks with open jaws suspended in the
water column, snagging fish and other forms as they
swam between the jaws or perhaps even collecting
prey whilst walking across plains and woodlands. The
straight nature of azhdarchid beak morphology
suggests that they did not specialize in one food item
and, as with storks, a vast range of small vertebrates
may have been eaten. More work is needed before
any of these hypotheses can be verified but, on a
theoretical level at least, the latter idea seems most
satisfactory.

A dying breed?
The azhdarchids are among some of the last
pterosaurs known. It seems that pterosaur diversity
declined towards the end of the Mesozoic and, by the
latest Cretaceous, only the azhdarchids remained in
any abundance (a few other pterosaur lineages may
have been present at this time, but their remains are
rare). It has been suggested that the apparent decline
of pterosaurs may merely be a taphonomic artefact,
perhaps instigated by a shift of pterosaur habit from
marine environments towards continental settings.
These environments are less likely to be preserved in
the geological record and do not provide many
settings conducive to fossilization, possibly hiding true
Late
Cretaceous
pterosaur
diversity.
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.
The extinction of the azhdarchids and pterosaurs
may be attributed to competition from birds, a group
which appear to have gradually replaced pterosaurs
in a variety of roles throughout the Cretaceous. As
the continental environments that the azhdarchids

38

had apparently become well adapted-to changed at

the termination of the Mesozoic it appears that
birdsand not azhdarchidspossessed the necessary
diversity and numbers to progress into the Tertiary.
No flying animals before or since the azhdarchids
have managed to capture their size or grandeur;
although we now only have fossils as evidence of
their presence, they clearly demonstrate that
azhdarchids must have been fantastic sights to
behold.

Suggestions for further reading

Chatterjee, S. & Templin, R.J. 2003. Posture,
Locomotion and Palaeoecology of the Pterosaurs.
Geological Society of America Special Publication,
v.376, pp.164.
Kellner, A.W.A. & Langston, W. Jr. 1996. Cranial
remains
of
Quetzalcoatlus
(Pterosauria,
Azhdarchidae) from Late Cretaceous sediments of
Big Bend National Park. Journal of Vertebrate
Paleontology, v.16, pp.222231.
Langston, W. Jr. 1981. Pterosaurs. Scientific American,
v.244, pp.92102.
Frey, E., Buchy, M.-C. & Martill, D.M. 2003. Middleand bottom-decker Cretaceous pterosaurs: unique
designs in active flying vertebrates. In: Buffetaut,
E. & Mazin, J.-M. (eds), Evolution and Palaeobiology
of Pterosaurs.
Geological
Society
Special
Publication, v.217, pp.267274.
Martill, D.M. & Frey, E. 1998. Discovery of the
holotype of the giant pterosaur Titanopteryx
philadelphiae Arambourg 1959, and the status of
Arambourgiania and Quetzalcoatlus. Neues Jahrbuch
fr Geologie und Palontologie, Abhandlungen, v.207,
pp.5776.
si, A., Weishampel, D.B. & Jianu, C.M. 2005. First
evidence of azhdarchid pterosaurs from the Late
Cretaceous of Hungary. Acta Palaeontologica
Polonica, v.50, pp.777787.
Unwin, D.M. 2003. On the phylogeny and
evolutionary history of pterosaurs. In: Buffetaut, E.
and Mazin, J. M. (eds), Evolution and Palaeobiology of
Pterosaurs, Geological Society Special Publication,
v.217, pp.139190.
Unwin, D.M. 2005. The Pterosaurs from Deep Time. Pi
Press, New York.

Blackwell Publishing Ltd, Geology Today, Vol. 23, No. 1, JanuaryFebruary 2007