''Tethydraco'' is a genus of
pterodactyloid
Pterodactyloidea (derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger" meaning "winged finger", "wing-finger" or "finger-wing") is one of the two traditional ...
pterosaur
Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the Order (biology), order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cre ...
that lived during 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'', ...
period (
Maastrichtian
The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interv ...
stage) of what is now the area of present
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria ...
, about 66 million years ago. ''Tethydraco'' was originally assigned to the family
Pteranodontidae
The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head ...
. Subsequently described material suggested that it was an
azhdarchid
Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cre ...
, and possibly synonymous with ''
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 ...
''. The
type and only species is ''T. regalis''.
Discovery and naming

Since 2015, a group of paleontologists has been acquiring pterosaur fossils from commercial Moroccan fossil traders, who obtain these from workers in the
phosphate mines on the
Khouribga
Khouribga ( ber, ⵅⵯⵔⵉⴱⴳⴰ, xʷribga; ar, خريبڭة, ḵurībga, ) is the capital of Khouribga Province in the Béni Mellal-Khénifra region, Morocco. With a population of 196,196 (2014 census), Khouribga owes its growth to the ph ...
plateau, which is located within the
Ouled Abdoun Basin
The Oulad Abdoun Basin (also known as the Ouled Abdoun Basin or Khouribga Basin) is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and a ...
. The purpose of this project is to determine pterosaur diversity in the latest Cretaceous. From this stage, no
Konservat-Lagerstätten are known, sites combining a large variety of species with exceptional preservation. It is in such sites that the vast majority of pterosaur fossils and taxa have been discovered. The latest Cretaceous had only produced some partial skeletons of
Azhdarchidae
Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cre ...
. Researchers usually have concluded from this fact that other pterosaur groups had already gone extinct. However, an alternative explanation could be that the poor
fossil record
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
caused a distorted image of the true situation through undersampling. To test this hypothesis, an effort was made to collect all pterosaurs bones brought to light by the massive and systematic commercial exploitation of the Khourigba phosphate layers. It transpired that indeed some finds could not be determined as azhdarchids and likely represented other groups. Four of the findings were described as new species in 2018, including Tethydraco.
In 2018, Nicholas R. Longrich, David M. Martill and Brian Andres described and named the type species ''Tethydraco regalis''. The generic name combines a reference to the
Tethys, the ocean in the Late Cretaceous separating Africa from Europe and Asia, with a
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
''draco'', "dragon". The
specific name means "royal" in Latin.
The
holotype
A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of seve ...
, FSAC-OB 1, was found in the middle ''Couche III'', itself the lowest phosphate layer complex at
Sidi Daoui
''Sidi'' or ''Sayidi'', also Sayyidi and Sayeedi, ( ar, سيدي, Sayyīdī, Sīdī (dialectal) " milord") is an Arabic masculine title of respect. ''Sidi'' is used often to mean "saint" or "my master" in Maghrebi Arabic and Egyptian Arabic. Wi ...
, dating from the late
Maastrichtian
The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interv ...
. It consists of a left humerus. The bone is relatively crushed. Other specimens have been referred to the species. FSAC-OB 199 is an ulna. FSAC-OB 200 is another ulna. FSAC-OB 201 is a thighbone. FSAC-OB 202 consists of a thighbone with shinbone. The describing authors admitted that a connection between the holotype and the referred specimens is hard to prove, in view of the lack of overlapping material. However, the wide ulnae fit the exceptional distal width of the humerus. The thighbones were, more tentatively, referred because they seemed to be
pteranodontid
The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the h ...
.
Description

''Tethydraco'' had a wingspan of and a body mass of .
The describing authors indicated some traits in which ''Tethydraco'' could be distinguished from known pteranodontids. In the humerus, the deltopectoral crest is placed rather proximally, closer to the torso of the animal, its closest border being positioned just proximal to the beginning of the opposite crest, the ''crista ulnaris''. Distally, away from the torso, the humerus has a broad triangular expansion. The bone ridge running to the outer joint condyle has a distinct process pointing to above, when the wing is in a stretched position. The ridge leading to the inner condyle is enlarged and extends towards the torso over a long distance. The ulna is relatively short and wide while its proximal end, towards the humerus, is massively expanded.
Phylogeny

''Tethydraco'' was placed in the family
Pteranodontidae
The Pteranodontidae are a family of large pterosaurs of the Cretaceous Period of North America and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a distinctive, elongated crest jutting from the rear of the head ...
; it would have been the youngest known member of that family, since its fossil remains dated back 66 million years ago. Its existence was seen as proof that pterosaur diversity in the Maastrichtian was higher than previously assumed. Apparent pterosaur decline would have been an illusion caused by the
Signor–Lipps effect
The Signor–Lipps effect is a paleontological principle proposed in 1982 by Philip W. Signor and Jere H. Lipps which states that, since the fossil record of organisms is never complete, neither the first nor the last organism in a given taxon w ...
, groups seeming to disappear earlier than a
mass extinction
An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. I ...
because their youngest fossils by chance have been found at somewhat older layers than the extinction event.
Below is a
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
showing the results of a
phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analysis first presented by Andres and colleagues in 2014, and updated with additional data by Longrich and colleagues in 2018. In this analysis, they found ''Tethydraco'' to be the sister taxon of the two species 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 C ...
'' (''P. longiceps'' and ''
P. sternbergi''); all three of them forming the family Pteranodontidae, which is found as the sister taxon of the family
Nyctosauridae.
In 2020, a wing from the same deposit was assigned to the genus; however, the morphology suggested that ''Tethydraco'' was an azhdarchid, rather than a pteranodontid as originally proposed, and possibly represented the wing-elements of ''
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 ...
''.
Paleoecology
''Tethydraco'' was discovered in the
Ouled Abdoun Basin
The Oulad Abdoun Basin (also known as the Ouled Abdoun Basin or Khouribga Basin) is a phosphate sedimentary basin located in Morocco, near the city of Khouribga. It is the largest in Morocco, comprising 44% of Morocco's phosphate reserves, and a ...
in
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria ...
. This basin is divided into layers called "Couches", and ''Tethydraco'' was discovered in Couche III. It coexisted with the pterosaurs ''
Alcione'', ''
Barbaridactylus'', ''
Simurghia'' and ''Phosphatodraco'' and the
abelisaurid
Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are fou ...
dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
''
Chenanisaurus
''Chenanisaurus'' is a genus of predatory abelisaurid dinosaur, with a single known species ''C. barbaricus''. It comes from the upper Maastrichtian phosphates of the Ouled Abdoun Basin in Morocco, North Africa. The animal is known from a holo ...
''.
References
{{Portal bar, Paleontology, Cretaceous, Morocco
Fossil taxa described in 2018
Pterosaurs of Africa
Ornithocheiroids