Phosphatodraco

Phosphatodraco is a genus of azhdarchid pterosaur that lived

during the Late Cretaceous of what is now Morocco. In 2000, a Phosphatodraco
pterosaur specimen consisting of five cervical (neck) vertebrae Temporal range: Late Cretaceous,
was discovered in the Ouled Abdoun Phosphatic Basin. The
specimen was made the holotype of the new genus and species
Phosphatodraco mauritanicus in 2003; the genus name means
"dragon from the phosphates", and the specific name refers to
the region of Mauretania. Phosphatodraco was the first Late
Cretaceous pterosaur known from North Africa, and the second
pterosaur genus described from Morocco. It is one of the only
known azhdarchids preserving a relatively complete neck, and
was one of the last known pterosaurs. Additional cervical
vertebrae have since been assigned to the genus, and it has been
suggested that fossils of the pterosaur Tethydraco represent wing
elements of Phosphatodraco.

Due to the fragmentary nature of the holotype cervical vertebrae,

there has been controversy over their order. The describers
considered them as cervicals (abbreviated as C) C5–C9 in the
series, the first preserved vertebra (C5) being broken in two, but
others consider them C3–C8, C3 and C4 being two different Assigned C5 cervical (neck) vertebra
vertebrae. The interpretation followed has consequences for how in multiple views
Phosphatodraco is distinguished from other azhdarchids and
how large it is thought to have been; the describers considered it Scientific classification
to have had a wingspan of 5 m (16 ft); the alternate interpretation Kingdom: Animalia
would lead to a 4 m (13 ft) wingspan. The complete neck may
have been 865 mm (2 ft 10 in) long. Phosphatodraco is mainly Phylum: Chordata
distinguished by its C8 (or C7) vertebra being very elongated, Order: †Pterosauria
50% longer than the C5, and in having a prominent neural spine
that is almost as tall as the centrum (the main part of the Suborder: †Pterodactyloidea
vertebra), truncated in a square shape at the top, and located far Family: †Azhdarchidae
back. As an azhdarchid, it would have had a proportionally long
neck, small body, and long limbs, compared to other pterosaurs. Subfamily: †Quetzalcoatlinae
Genus: †Phosphatodraco
The closest relatives of Phosphatodraco appear to have been
Aralazhdarcho and Eurazhdarcho. Azhdarchids have Pereda-Suberbiola et
historically been considered skim-feeders that caught prey from al., 2003
water in coastal settings, but it has since been suggested that the
Type species
context in which their fossils are found and their morphology  –
such as their long, stiffened necks (informed by for example the †Phosphatodraco mauritanicus
neck of Phosphatodraco) – is more consistent with them having Pereda-Suberbiola et al., 2003
foraged terrestrially like storks or ground hornbills, but this is still
debated. Although pterosaurs were thought to have declined in
diversity towards the time of their extinction 66 million years ago, the diversity in taxa, including
Phosphatodraco, in the Ouled Abdoun Basin, which dates to the late Maastrichtian, right before the
Cretaceous-Paleogene extinction event, indicates their extinction happened abruptly.
Contents
Discovery
Interpretations of cervical vertebra order
Description
Cervical vertebrae
Classification
Paleobiology
Feeding and ecological niche
Locomotion
Paleoenvironment
See also
References
Bibliography

Discovery
Location of phosphate mines in Morocco (A), the Sidi Daoui
mine where Phosphatodraco was found (B), and stratigraphic
location of the discovery site, "couche III" (C)
place since 1997.[1][2]
During the late 1990s, remains of
pterosaurs began to be discovered in
different fossil localities of Morocco, all
dating to the Cretaceous period. In 2000,
pterosaur fossils were found by the Office
Chérifien des Phosphates (OCP, located in
Casablanca) during paleontological field
work in the eastern part of the Ouled
Abdoun Phosphatic Basin near the city of
Khouribga in central Morocco. They were
collected from "site 1" in the Sidi Daoui
mine in the northern part of Grand Daoui,
an area in which phosphate is quarried. The
fossils were found in the upper part of the
stratigraphic unit which miners called
"couche III". These excavations were part
of a collaboration between the OCP, the
Ministère de L'Energie et des Mines, Rabat,
and the French Centre National de la
Recherche Scientifique, which had taken

The pterosaur material, catalogued as specimen OCP DEK/GE 111, consists of five disarticulated but
closely associated cervical (neck) vertebrae and an indeterminate bone, most likely belonging to a single
individual. The vertebrae are crushed and damaged, and the surface of the bone is missing in some areas,
with some infilling of phosphate sediments, and the fossils have therefore not been removed from the
matrix. The block containing the bones is 98 cm (39 in) long and 34 cm (13 in) wide. During mechanical
preparation of the specimen fossil remains of other animals were also found in association, including of
several types of fish and mosasaurs.[1]
The specimen was made the holotype of the new genus and species Phosphatodraco mauritanicus by
paleontologist Xabier Pereda-Suberbiola and colleagues in 2003. The genus name derives from the words
phosphate and the Latin draco, meaning "dragon from the phosphates", and the specific name refers to the
region of Mauretania where the fossils were found.[1] The describers gave the etymology of Mauretania as
Latin for North Africa; other sources specify it as an area stretching from Algeria to Morocco.[1][3][4]
Phosphatodraco was the first Late Cretaceous pterosaur known from North Africa (and thus the first
known member of the family Azhdarchidae of this age from the region), and only the second pterosaur
genus described from Morocco (the first being Siroccopteryx). At the time it was described, it was one of
the only known azhdarchids preserving a relatively complete neck (the others being Zhejiangopterus and
Quetzalcoatlus[5]), and was one of the last known pterosaurs.[1] Complete neck vertebral series are rare for
azhdarchids, but such vertebrae are some of the most commonly found and best known remains of the
group.[6]

Humerus (left) and ulnae (right) of Tethydraco, which may represent wing elements of Phosphatodraco

In 2018 paleontologist Nicholas R. Longrich and colleagues reported pterosaur fossils collected from
"couche III" in cooperation with the Moroccan fossil industry the previous three years; until that point, only
the single specimen of Phosphatodraco was known from the assemblage. At the time, the collection was
the largest and most diverse collection of pterosaurs from the Maastrichtian age of the Late Cretaceous, and
included two cervical vertebrae they assigned to Phosphatodraco, based on similarity with the holotype in
size and proportions. One of the vertebrae, specimen FSAC-OB 12, was identified as a C5 (though stated
in the description to be similar to C6 of the holotype); the other, FSAC-OB 13, was identified only as a
cervical vertebra. The cervical ribs (ribs of the neck vertebrae) of FSAC-OB 12 do not appear to have yet
fused to the centrum (the main part of the vertebra), so the animal may not have been completely mature.
These specimens are housed at Faculté des Sciences Aïn Chock in Casablanca.[2]

In 2020 paleontologists Claudio Labita and David M. Martill described an articulated (where the bones are
connected as in life) pterosaur wing from "couche III" (specimen FSAC CP 251, bought from fossil
dealers), which they assigned to Tethydraco, a genus also described by Longrich and colleagues in 2018
based on a humerus (upper arm bone). Tethydraco was originally considered a pteranodontid, but Labita
and Martill found it to be an azhdarchid, and to possibly represent the wing elements of Phosphatodraco.
They noted that more associated and articulated pterosaur fossils were being collected from these deposits
due to improving methods used by fossil diggers, and that azhdarchid fossils were becoming abundant.
They also cautioned that the provenance of some of the Moroccan fossils was difficult to establish, due to
the commercial nature of their collection.[7]

Interpretations of cervical vertebra order

Pereda-Suberbiola and colleagues originally interpreted the five preserved cervical vertebrae of
Phosphatodraco as cervicals C5–C9. The frontmost preserved vertebra they interpreted as C5 consisted of
two fragments; they found it unlikely that these belonged to two different vertebrae, since they lay in
continuity with no sediment in between, and overlapped each other in some areas. They considered the
sideways expansion at the front of this vertebra to be due to crushing, and pointed out that such
 preservation where fragile, yet well-
preserved bones are associated with
damaged material of the same individual is
known from other vertebrate fossils in the
same level. They identified the frontmost
vertebra as a C5 because this is usually the
longest cervical vertebra in pterosaurs, their
Illustration of the holotype cervical vertebra series as length decreasing hindward.[1]
preserved, with interpretation of its order by Pereda-Suberbiola
and colleagues, 2003 (above, C5–C9), and Kellner, 2010 In 2007 paleontologist Alexander W. A.
(below, C3–C8) Kellner and colleagues noted that
Phosphatodraco was one of the most
interesting azhdarchids found in Africa, but
used cautious language about the original interpretation of the vertebrae.[8] Kellner suggested in 2010 that
interpretation of the holotype specimen had been affected by taphonomy (changes during decay and
fossilisation); he instead proposed that the element originally described as a fifth cervical vertebra was
actually the third and fourth vertebrae which had been compressed together, giving the impression that they
were a single vertebra broken in the middle. This would shift the order of the vertebrae to C3–C8, and
though this did not change the validity of the taxon (its distinctness being based on its stratigraphy,
geography, and morphology), Kellner noted that the diagnosis (the suite of features that distinguish a taxon)
and size estimate had to be reevaluated.[9] The frontmost cervical vertebrae which are missing from
Phosphatodraco in either scheme are the atlas (C1, which connects with the back of the skull) and the axis
(C2).[9][10]

Subsequent articles from 2011 and 2015 with Kellner among the co-authorship have concurred with
Kellner's interpretation.[11][12] Paleontologist Alexander Averianov disagreed with Kellner's
reinterpretation of the cervical vertebrae in 2014, and considered the original description accurate.[13] A
2015 article by paleontologist Mátyás Vremir and colleagues called the issue "controversial" and
considered the specimen too crushed for proper comparison,[14] and Martill and Markus Moser concurred
with this in 2018.[15] Paleontologists Darren Naish and Mark P. Witton (the co-authors of Vremir's article)
followed Kellner's interpretation in 2017.[1][16] Paleontologist Rodrigo V. Pêgas and colleagues also
followed Kellner's order in 2021.[17] Though the palaeontologist Alexandru A. Solomon and colleagues
noted the suggested change in interpretation of the holotype order in 2019, they stated that even if the
reinterpretation was correct, the specimen was too damaged for comparison with the single known cervical
vertebra of their new genus Albadraco.[18]

Description
In their 2003 description, Pereda-Suberbiola and colleagues
estimated Phosphatodraco to have had a wingspan close to 5  m
(16 ft), based on comparison with other azhdarchids with preserved
cervical vertebrae, and referred to it as a "large azhdarchid
pterosaur". This is larger than azhdarchids such as Zhejiangopterus
and Montanazhdarcho, and comparable to the smaller species of
Quetzalcoatlus, Q. lawsoni; the larger Q. northropi is thought to
have reached 10–11 metres (33–36  ft), thereby being the largest
known flying animal.[1][19] Witton grouped Phosphatodraco with
Estimated size (upper middle in
"midsized" azhdarchids based on this size estimate in 2013.[20] In
turquoise) compared to contemporary
2010, Kellner suggested this size estimate too large, based on his
pterosaurs, birds, and a human
reinterpretation of the neck vertebra order.[9] Naish and Witton, who followed Kellner's interpretation, listed
a neck-length of 865 mm (2 ft 10 in) for Phosphatodraco in 2017, and a wingspan of 4 m (13 ft).[16]

There were two main types of azhdarchid skulls; very long, low skulls that were up to ten times longer than
wide, and some that were much shorter than that, closer to those of other pterosaurs. Some had crests and
some did not. Azhdarchids had necks that were proportionally longer than those of other pterosaurs, and
their vertebral column and much of the rest of the skeleton was pneumatized (filled with air-sacs that
lightened it). The body skeleton of azhdarchids was small but robust, and their upper arm bones were
solidly built. Their wing-metacarpals (the hand bones that connect with the fingers) were relatively the
largest among pterosaurs, and the longest bones in the wings; their wing-fingers (which supported the
wing-membranes) were relatively short. The pelvises were relatively robust, and the hindlimbs long.
Combined, the long wing metacarpals and legs made azhdarchids relatively taller when standing than other
pterosaurs, though their feet were narrow and short.[20] As a pterosaur, Phosphatodraco was covered with
hair-like pycnofibers.[21]

Cervical vertebrae

The suite of features that distinguish a

taxon from other related taxa is called a
diagnosis, and in the case of
Phosphatodraco, these features are all
found in the cervical vertebrae. Since
Pereda-Suberbiola and colleagues
considered the preserved vertebrae to be
C5–C9 of the series in their description,
that is the diagnosis and description
Azhdarchid neck-lengths compared to their estimated
followed here.[1] Note that if Kellner's
wingspans. Phosphatodraco is fourth from the top, following
suggestion that the series actually represents
Kellner's interpretation. (The last preserved vertebra is not
vertebrae C3–C8 is correct, the diagnostic
shown)
features listed by Pereda-Suberbiola and
colleagues may possibly be inaccurate, and
the following description would refer to different vertebrae in the series.[9] Pereda-Suberbiola and
colleagues found Phosphatodraco to differ from other azhdarchids in that the hindmost vertebra, C8 in their
order, is very elongated, 50% longer than C5, and has a prominent neural spine (the spine that projects
upwards from the vertebra) that is almost as tall as the centrum, truncated in a square shape at the top, and
located far back. Phosphatodraco is also distinct in that the ratio between the maximum length of the
vertebrae and the front width between prezygapophyses (processes at the sides of the centrum which
connected with the postzygapophyses of the previous vertebrae) of the middle cervical vertebrae is about
4.3 in C5 and 4.1 in C6.[1]

The five preserved cervical vertebrae have hollow centra and their cortical bone (the outer, thick layer) is
about 1  mm (0.039  in) thick. The vertebrae vary in length, the longest being the frontmost of those
preserved, a C5 broken in two according to Pereda-Suberbiola and colleagues (C3–C4 according to
Kellner[9]), which they estimated to have been about 300 mm (0.98 ft) long when complete. The first of
these fragments is 110 mm (4.3 in), the second is about 190 mm (7.5 in). When viewed from the side, the
hind end of this vertebra shows a developed left postzygapophysis, and the convex articular condyle (the
condyle that connected with the following vertebra) and left postexapophysial process (which connected
with the preexapophys at the front of the preceding vertebra) is in front of it, all lying in the same plane due
to crushing.[1]
The C6 (Kellner's C5[9]) is the best-preserved of the vertebrae, and
is shorter than the preceding C5, about 225 mm (8.9 in) long. It is
distorted so that the front part is visible in lower side view, and the
hind part is visible in left side view. Its centrum is procoelous
(concave at the front surface), its prezygapophyses are horn-like,
and nearly concave and parallel when seen from below. The right
prezygapophys has a small tubercle (a rounded projection) at the
midline, and there is no indication of additional processes at the
front end of the vertebra or pneumatic foramina (holes) on the side
surface of the centrum. The front cotyle (the concave front end of
the centrum) is distorted, but it appears to be twice as wide as high,
ovoid (or egg-shaped), and with a slightly concave upper margin.
The lower margin has a prominent hypapophysis (a downwards
projection), and this keel's height diminishes towards the centrum's
mid-length. There is a longitudinal oval sulcus (a groove) on the
right side of the lower surface, near the base of the C3–C5 vertebrae of Quetzalcoatlus
prezygapophysis. The lower surface of the centrum is nearly flat, (A–C) compared with those of
and the postexapophys is well-developed at the lower side of the Phosphatodraco (D)
condyle, like in the preceding vertebrae.[1]

The following C7 vertebra (Kellner's C6[9]) is visible in bottom view, is missing the hind part, the
preserved part being 190 mm (7.5 in) long, and the complete length perhaps being the same or shorter as
the preceding C6. Its prezygapophyses are similar to those of the preceding vertebra, and the cotyle is oval
and compressed from top to bottom. There is a sulcus below the left prezygapophys, apparently without a
ridge extending hindward. The centrum bulges slightly hindward, becoming narrower at its mid-length.
The postzygapophyses are well-developed, and diverge widely from the longitudinal midline of the
centrum. A small protuberance between the postzygapophyses perhaps indicates where the upper margin of
the neural canal (through which the spinal cord passed) was located, though its features cannot be
accurately determined.[1]

The next to last vertebra is C8 (Kellner's

C7[9][16]) which is visible in side view, and
despite missing some of the front, the
centrum is very elongated, measuring
150 mm (5.9 in). The most notable feature
of this vertebra is its tall neural spine, which
is placed at its hindmost part. The neural
spine is 40 mm (1.6 in) high measured from
the upper surface of the postzygapophys to
the top, almost the same height as the
centrum, which is 45  mm (1.8  in). The
front and hind margins of the neural spine
are vertically parallel to each other, and its
Characteristics of azhdarchid cervical vertebrae; J is the C7 of
top is truncated in a square shape, and
Phosphatodraco following Kellner's order (it is C8 in Pereda-
perpendicular to the side edges. The left
Suberbiola and colleagues' order)
postzygapophys is located at the base of the
hind end of the neural arch (which forms
the arch of bone through which the spinal
cord passed). It is similar to the same vertebra of Quetzalcoatlus in that the neural spine is square on top,
but differs in being placed so far back. The left postexaphophyseal process is well-developed at the back
and below, but does not extend past the condyle as it does in the preceding vertebrae.[1]
The last vertebra is the C9 according to Pereda-Suberbiola and colleagues (Kellner's C8,[9] which is visible
in hind view, and its preserved part is 75 mm (3.0 in) high. Its neural arch has a large neural spine and well-
developed transverse processes (which projected from the sides of the centrum and acted as attachment
points for muscles and ligaments). The neural spine terminates in a blunt process above, and the hind side
of the neural spine has an oval depression, which has thick vertical edges at its sides. The transverse
processes are long and slender, and project to the sides and slightly downwards. The neural canal is small
and nearly circular, is about 22 mm (0.87 in) in diameter, and there are no pneumatic foramina near it. Its
condyle is broad, around five times wider than high, and is crescent-shaped in cross-section. The left
postexapophysis is placed at the side of the condyle and almost vertical. Though none of the vertebrae
preserve cervical ribs, the development of the transverse processes of the last vertebra indicates that it
probably had ribs.[1] The indeterminate bone fragment associated with the two last vertebrae has a similar
texture to them, is flat and crescent-shaped, and is about 9 mm (0.35 in) wide and 44 mm (1.7 in) long.[1]

Pereda-Suberbiola and colleagues found that the frontmost

preserved cervical vertebrae of Phosphatodraco (their C5–C7)
were similar in form to those of the mid-series cervical vertebrae of
other long-necked pterodactyloid pterosaurs (the group consisting
of short-tailed pterosaurs). The last hindmost preserved vertebrae
(their C8–C9) have some features in common with the rest of the
vertebrae, including broad, ovoid cotyles and condyles, as well as
postexapophyses, but they differ in their neural canal being
demarcated from the centrum and in having a prominent neural
spine. Pereda-Suberbiola and colleagues suggested the C8–C9
could be cervicalized dorsal vertebrae, back vertebrae which have
been incorporated into the neck. The total number of cervical Life restoration showing
vertebrae in pterosaurs varies between seven and nine, and the first Phosphatodraco in terrestrial pose
dorsal vertebra is considered to be the first one that connects with
the sternum (breast bone). Early pterosaurs like Rhamphorhynchus
had eight cervical vertebrae with cervical ribs on at least C3–C8; later pterodactyloids had seven vertebrae
and no ribs. In later pterdactyloid groups, nine cervical vertebrae are present, two of them being
cervicalized dorsals, and adults have a notarium (a structure consisting of fused vertebrae in the shoulder-
region, also seen in birds).[1]

Classification
In their 2003 description Pereda-Suberbiola and colleagues considered Phosphatodraco a member of
Azhdarchidae based on features such as its mid-series cervical vertebrae being elongated, with low vestigial
(almost evolutionarily lost) or absent neural spines, the presence of prezygapophyseal tubercles, a pair of
lower sulci near the prezygapophyses, and the lack of oval pneumatic foramina on the lower surfaces of the
centra. These features are especially similar to those of Quetzalcoatlus and Azhdarcho. Other features
distinguishing the group could not be identified in Phosphatodraco due to the preservation of its fossils.[1]

Longrich and colleagues performed a phylogenetic analysis that included Moroccan pterosaurs in 2018,
and found Phosphatodraco to be an azhdarchid, and the sister taxon of Aralazhdarcho from Kazakhstan.[2]
A 2021 analysis by Rodrigo and colleagues also found these two genera to be sister taxa, joined in a clade
by Eurazhdarcho from Romania. This clade was supported by one clear synapomorphy (a distinct,
ancestrally shared feature), that the side margin of the mid-cervical vertebrae is straight when viewed from
above and below, with almost parallel sides. These researchers noted that previous studies had defined
Azhdarchidae as a node based clade with Azhdarcho and Quetzalcoatlus as internal specifiers, but
cautioned that in their new phylogeny, Phosphatodraco, Zhejiangopterus, and Eurazhdarcho would fall
outside the group. They found this undesirable, as those genera had otherwise consistently been considered
azhdarchids, and that for stability's sake, Phosphatodraco should be added as a third internal specifier for
the group, since this would result in all these taxa being included.[17]

In 2021 paleontologist Brian Andres and colleagues also found Phosphatodraco and Aralazhdarcho to be
sister taxa, supported by the reduction of pneumatic foramina on the side of the neural canal. This clade
was recovered as part of the azhdarchid subclade Quetzalcoatlinae. The cladogram below shows the
placement of Phosphatodraco within Azhdarchiformes according to Andres and colleagues, 2021:[22]

Reconstructed skeleton of
Quetzalcoatlus in a flying pose

Azhdarchiformes
   
  Radiodactylus langstoni
 
   
  Montanazhdarcho minor
Azhdarchidae
   
  Azhdarcho lancicollis

 
  Albadraco tharmisensis
Azhdarchinae
 
   
    Aerotitan sudamericanus
 
 
 
  Mistralazhdarcho maggii
Quetzalcoatlinae
   
  Aralazhdarcho bostobensis
 
 
 
  Phosphatodraco mauritanicus
 
    Eurazhdarcho langendorfensis
 
 
    Zhejiangopterus linhaiensis
 
 
    Cryodrakon boreas
 

 
  Wellnhopterus brevirostris
   
     Hatzegopteryx thambema

 
  Arambourgiania philadelphiae

   
    Quetzalcoatlus lawsoni
 
 
 
  Quetzalcoatlus northropi

Paleobiology

Feeding and ecological niche

Skeletal diagrams showing Zhejiangopterus (left, as preserved without skull, right, as reconstructed), one of
the two other azhdarchids known preserved with a relatively complete neck (the other is Quetzalcoatlus)

In 2008 Witton and Naish pointed out that although azhdarchids have historically been considered to have
been scavengers, probers of sediment, swimmers, waders, aerial predators, or stork-like generalists, most
researchers until that point had considered them to have been skim-feeders living in coastal settings, which
fed by trawling their lower jaws through water while flying and catching prey from the surface (like
skimmers and some terns). In general, pterosaurs have historically been considered marine piscivores (fish-
eaters), and despite their unusual anatomy, azhdarchids have been assumed to have occupied the same
ecological niche. Witton and Naish noted that evidence for this mode of feeding lacked support from
azhdarchid anatomy and functional morphology; they lacked cranial features such as sideways compressed
lower jaws and the shock-absorbing adaptations required, and their jaws instead appear to have been almost
triangular in cross-section, unlike those of skim-feeders and probers.[6]

Witton and Naish instead stated that azhdarchids probably inhabited inland environments, based on the
taphonomic contexts their fossils have been found in (more than half the fossils surveyed were from for
example fluvial or alluvial deposits, and most of the marine occurrences also had fossils of terrestrial
lifeforms), and their morphology made them ill-suited for lifestyles other than wading and foraging
terrestrially, though their feet were relatively small, slender, and had pads, not suited for wading either.
These researchers instead argued that azhdarchids were similar to storks or ground hornbills, generalists
they termed "terrestrial stalkers" that foraged in different kinds of environments for small animals and
carrion, supported by their apparent proficiency on the ground and relatively inflexible necks, for example
the well-preserved neck of Phosphatodraco providing information about their morphology. Witton and
Naish suggested that their more generalist lifestyle could explain the group's resilience compared to other
pterosaur lineages, which were not thought to have survived until the late Maastrichtian like the
azhdarchids did (pterosaurs went extinct along with the non-bird dinosaurs during the Cretaceous-
Paleogene extinction event 66 million years ago).[6][2]
Witton elaborated in a 2013 book that the proportions of azhdarchids would have been consistent with them
striding through vegetated areas with their long limbs, and their downturned skull and jaws reaching the
ground. Their long, stiffened necks would be an advantage as it would help lowering and raising the head
and give it a vantage point when searching for prey, and enable them to grab small animals and fruit.[23] In
their 2021 study that reinterpreted Tethydraco as an azhdarchid, and possibly the same as Phosphatodraco,
Labita and Martill noted that azhdarchids might have been less terrestrial than suggested by Witton and
Naish, since the Moroccan fossils were from marine strata, as was Arambourgiania from the phosphates of
Jordan. They noted that no azhdarchids had been found in truly terrestrial strata, and proposed they could
instead have been associated with aquatic environments, such as rivers, lakes, marine and off-shore
settings.[7]

Pterosaurs are generally thought to have gone gradually extinct by decreasing in diversity towards the end
of the Cretaceous, but Longrich and colleagues suggested this impression could be a result of the poor
fossil records for pterosaurs (the Signor-Lipps Effect). Since they found multiple lineages (Pteranodontidae,
Nyctosauridae, and Azhdarchidae, the latter including Phosphatodraco and others) to have co-existed
during the late Maastrichtian of Morocco, this is the most diverse pterosaur assemblage known from the
Late Cretaceous. Pterosaurs during this time thereby had increased niche-partitioning compared to earlier
faunas from the Santonian and Campanian ages, and they were able to outcompete birds in large size based
niches, and birds therefore remained small, not exceeding 2  m (6.6  ft) wingspans during the Late
Cretaceous (most pterosaurs during this time had larger wingspans, and thereby avoided the small-size
niche). To these researchers, this indicated that the extinction of pterosaurs was abrupt instead of gradual,
caused by the catastrophic Chicxulub impact. Their extinction freed up more niches that were then filled by
birds, which led to their evolutionary radiation in the Early Cenozoic.[2]

Locomotion

Possible azhdarchid trackway Haenamichnus from Korea (left) and feeding posture inferred from the tracks

Witton summarized ideas about azhdarchid flight abilities in 2013, and noted they had generally been
considered adapted for soaring, although some have found it possible their musculature allowed flapping
flight like in swans and geese. Their short and potentially broad wings may have been suited for flying in
terrestrial environments, as this is similar to some large, terrestrially soaring birds. Albatross-like soaring has
also been suggested, but Witton thought this unlikely due to the supposed terrestrial bias of their fossils and
adaptations for foraging on the ground. Studies of azhdarchid flight abilities indicate they would have been
able to fly for long and probably fast (especially if they had an adequate amount of fat and muscle as
nourishment), so that geographical barriers would not present obstacles.[24]

Azhdarchids are also the only group of pterosaurs to which trackways have been assigned, such as
Haenamichnus from Korea, which matches this group in shape, age, and size. One long trackway of this
kind shows that azhdarchids walked with their limbs held directly underneath their body, and along with
the morphology of their feet indicates they were more proficient on the ground than other pterosaurs.
According to Witton, their proportions indicate they were not good swimmers on the other hand, and
though they could probably launch from water, they were not as good at this as some other pterosaur
groups.[24]

Paleoenvironment
Phosphatodraco is known from the "couche III" phosphatic unit of
the Ouled Abdoun Basin in Morocco, which was deposited during
the late Maastrichtian age of the Late Cretaceous period, which
ended 66 million years ago. The phosphatic series is condensed and
the Maastrichtian part is only 3–5 cm (1.2–2.0 in) thick. From the
bottom to the top, "couche III" consists of thin phosphatic levels
The contemporary Chenanisaurus
moving along the coastline
and marls, a grey limestone bed containing fish fossils, yellow, soft
phosphates at the lower level, a thick, yellow marly level,
separating the lower and upper "couche III", and gray, soft
phosphates with brown stripes overlay in a thick marl level at upper "couche III". Pterosaur fossils are
found in the lower part of the upper phosphatic unit.[1][2] The kind of fossils that are usually used in
biostratigraphy are rare, which complicates attempts at dating these beds, but "couche III" has been
correlated with the late Maastrichtian on the basis of shark teeth, which has also been confirmed by carbon
and oxygen istope stratigraphy.[25] The phosphates were deposited in an embayment of the eastern Atlantic
Ocean that flooded North Africa during the Late Cretaceous and Early Paleogene.[26] The area would have
been part of the Tethys Sea at the time.[2]

The phosphatic matrix of the original Phosphatodraco specimen is gray and mottled with orange, and
contained fossils including of the fish Serratolamna, Rhombodus, and Enchodus, and the mosasaur
Prognathodon, as well as small nodules. The specimen was found close to skeletal remains of an
indeterminate mosasaur, and remains of sharks, rays, fish such as Stratodus, mosasaurs such as
Platecarpus, Mosasaurus, and Halisaurus, indeterminate elasmosaurid plesiosaurs, and indeterminate
bothremydid turtles have also been found at the site. This assemblage of animals suggests the sediments
were deposited in a marine environment.[1] Other contemporary pterosaurs from the Ouled Abdoun Basin
include the pteranodontid Tethydraco (if it is not the same animal as Phosphatodraco), the nyctosaurids
Alcione, Simurghia, and Barbaridactylus, a small azhdarchid similar to Quetzalcoatlus, and a very large
azhdarchid which may be Arambourgiania. Dinosaurs are rare there, but the abelisaurid Chenanisaurus,
the hadrosaur Ajnabia, and sauropod fossils are also known. Longrich and colleagues suggested in 2018
that, although the fauna was overwhelmingly marine, the presence of terrestrial dinosaurs and azhdarchids
indicates the coast was nearby.[2][7][25][27]

See also
List of pterosaur genera
Timeline of pterosaur research
Pterosaur size

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Bibliography
Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs. Crescent Books.
ISBN 978-0-517-03701-0.
Witton, M. P. (2013). Pterosaurs: Natural History, Evolution, Anatomy (1st ed.). Princeton
University Press. ISBN 978-0-691-15061-1.

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