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Arambourgiania

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Not to be confused with Arambourgia.

Arambourgiania
Temporal range: Maastrichtian
Holotype fossil cast at Museum Histoire Naturelle, Paris
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Order: †Pterosauria
Suborder: †Pterodactyloidea
Family: †Azhdarchidae
Subfamily: †Quetzalcoatlinae
Genus: †Arambourgiania
Nessov vide Nessov & Yarkov, 1989
Species:
†A. philadelphiae
Binomial name
†Arambourgiania philadelphiae
(Arambourg, 1959)
Synonyms
Genus synonymy
Species synonymy
  • Titanopteryx philadelphiae
    Arambourg, 1959

Arambourgiania (meaning "Camille Arambourg's", with -iania being a suffix indicating possession) is a genus of pterosaur, an extinct group of flying reptiles, that inhabited Jordan during the Maastrichtian age of the Cretaceous period, around 72 to 66 million years ago. Additional fossil remains from the United States and Morocco have also been found, but their assignment to Arambourgiania is only tentative. The holotype (name-bearing) specimen was discovered in 1943 by a railway worker near Russeifa, Jordan. After examination of the specimen by paleontologist Camille Arambourg, a new genus and species was named in 1959, Titanopteryx philadelphiae. The generic name means "titan wing", as the fossil was initially misidentified as a wing metacarpal (it would be later identified as a cervical (neck) vertebra), while the specific name refers to the ancient name of Amman (the capital of Jordan), Philadelphia. The genus "Titanopteryx" would later be problematic, as it had already been taken by a fly. Because of this, paleontologist Lev Nessov in 1989 named a new genus, Arambourgiania, in honor of Arambourg. Since 1943, additional isolated remains including vertebrae, wing bones, and hindlimb bones have been referred to the genus.

Due to the fragmentary nature of Arambourgiania fossils, little direct information is known about its anatomy. Its cervical vertebrae are extremely elongated, with the holotype vertebra measuring 77–78 cm (2 ft 6.31 in – 2 ft 6.71 in) in length. Based on the complete neck of its relative Quetzalcoatlus, Arambourgiania has a total neck length of 3 m (9 ft 10 in), a length longer than that of the contemporary giraffe. Its vertebrae are also more lightly built and weakly muscled than its robust, short-necked relative Hatzegopteryx. Arambourgiania is one of the largest flying animals ever discovered. Initial wingspan estimates ranged from 11 to 13 m (36 to 43 ft), which would have made it the largest known pterosaur. However, due to Arambourgiania only presenting fragmentary specimens, recent research has suggested more moderate wingspan estimates. Some of the latest studies put the wingspan anywhere between 8 to 10 m (26 to 33 ft), which would still be among the largest known flying creatures. Arambourgiania has often been compared to other gigantic pterosaurs such as Quetzalcoatlus and Hatzegopteryx in terms of size.

Arambourgiania is a member of the family Azhdarchidae, which includes some of the largest known pterosaurs ever. Azhdarchids are thought to have had a terrestrial stalker lifestyle, similar to that of extant storks, however some studies have pointed out that azhdarchids are frequently found in marine deposits. This includes Arambourgiania, whose fossils are exclusively known from oceanic or coastal localities, indicating that azhdarchids may have consumed aquatic prey as well. The bones of Arambourgiania are extremely hollow and lightly built with adaptations for soaring. One of the closest relatives of Arambourgiania is Quetzalcoatlus, as multiple studies have found both pterosaurs to be grouped together within Azhdarchidae. In Jordan, Arambourgiania lived in the Ruseifa Formation which was a deep marine deposit during the Maastrichtian. At this time, Jordan was submerged under the Tethys Sea and had a diverse composition of marine life, including mosasaurids, elasmosaurids, bony fish, and selachians. Arambourgiania is one of two pterosaurs known from Jordan, the other being Inabtanin.

History of research

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Arambourgiania was named after paleontologist Camille Arambourg, who first studied the specimen

In 1943, a railway worker during repairs on the Amman-Damascus railroad, near the city of Russeifa in Jordan, an incomplete fossilized cervical (neck) vertebra measuring 60.96 cm (2 ft). This vertebra was acquired in 1943 by the director of a nearby phosphate mine, Amin Kawar, who brought it to the attention of British archaeologist Fielding after the Second World War. This generated some publicity — the vertebra was even shown to Abdullah I of Jordan — but more importantly, it made the scientific community aware of the find.[1] These phosphates belong to the Ruseifa Formation, which dates to the Maastrichtian age (72-66 ma) of the Late Cretaceous period.[2][3]

In 1953, the vertebra was sent to the National Museum of Natural History in Paris, France, where it was examined by French paleontologist Camille Arambourg. In 1954, he misinterpreted the vertebra as the wing metacarpal of a giant pterosaur, though he did not give the specimen a genus or species name. Afterwards, in 1959, he named the new genus and species: Titanopteryx philadelphiae. The generic name comes from the ancient Greek roots titan, in reference to the giant Greek Titans, and pteryx meaning "wing", while the specific name refers to the ancient name of Amman that was used by the Greeks: Philadelphia. The holotype and only known specimen was chosen to be this wing metacarpal, although several more elements were known but undescribed and unidentified at the time.[3][4] Arambourg let a plaster cast be made and then sent the fossil back to the phosphate mine; this last aspect was later forgotten and the bone was assumed lost.[1]

This fossil indicated the largest pterosaur known to science, but remained largely ignored, with the pteranodontid Pteranodon still being recognized as the largest pterosaur known to science, until the description of Quetzalcoatlus.[5] While studying and describing the closely related pterosaur Quetzalcoatlus from Texas in 1975, American paleontologist Douglas A. Lawson concluded that the bone was not a metacarpal but a cervical vertebra, the correct identification.[6] In the 1980s, Russian paleontologist Lev Nesov was informed by an entomologist that the name Titanopteryx had already been given by Günther Enderlein to a fly from the Simuliidae family in 1935. Therefore, in 1989, he created the new genus name Arambourgiania, honoring Arambourg.[7] However, the name "Titanopteryx" was informally kept in use in the West, partially because the new name Arambourgiania was assumed by many to be a nomen dubium (dubious name).[2]

In early 1995, paleontologists David Martill and Eberhard Frey traveled to Jordan in an attempt to clarify matters. In a cupboard of the office of the Jordan Phosphate Mines Company they discovered some other pterosaur bones: a smaller vertebra and the proximal and distal extremities of a wing phalanx — but not the original material of Arambourgiania.[4] However, after their departure to Europe, engineer of the mine Rashdie Sadaqah investigated further and in 1996 he established that the original fossil had been bought from the company in 1969 by geologist Hani N. Khoury, who had then donated it in 1973 to the University of Jordan. It was still present in the collection of this institute and now it could be restudied by Martill and Frey.[3][8][2] Later, Frey and Martill rejected the suggestion that Arambourgiania was a nomen dubium or an identical pterosaur to Quetzalcoatlus, affirming its validity to replace the preoccupied name "Titanopteryx".[3][2]

Additional specimens

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Afro-Arabia

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Map of sites preserving Arambourgiania.
Navesink Formation
Coon Creek Formation
Jordan Phosphates
Khouribga Phosphates (type locality)
Merchantville Formation
Mishash Formation
class=notpageimage|
Fossil localities of Arambourgiania Legend: A. philadelphiae A. sp. Formerly referred specimens

In 2018, a set of topotype specimens from Arambourgiania that had been located in the Bavarian State Collection for Palaeontology and Geology in Munich, Germany were described after having been in the museum's collections since 1966. These fossils were unearthed or purchased by fossil collector J. Otto Haas in 1955 and were catalogued in his personal collection. The azhdarchid fossils in the collection include six fragmentary specimens, only one of which, a cervical vertebra fragment, is confidently referable to A. philadelphiae. The other five fragments, which were referred to cf. Arambourgiania philadelphiae, include another cervical vertebra fragment, a potential neural arch, a left femur (thighbone) fragment, a possible radius (forearm bone), and an incomplete metacarpal IV, as well as other indeterminate fragments. It is possible due to their same origin, lack of overlap, and size these topotype specimens come from the same individual as the holotype. However, due to the lack of detailed date on their discovery, it is impossible to verify this. Additionally, the holotype was unearthed in 1943 whereas the topotype material was unearthed or purchased 12 years later.[4]

In 1992, Lewy et al described a pair of natural endocranial endocasts (an internal cast) which had been unearthed from Upper Campanian or Early Maastrichtian-aged phosphates belonging to the Mishash Formation located near Mitzpe Ramon, Israel. Lewy referred these endocasts to Titanopteryx (=Arambourgiania) sp. on the basis of their similarity to the endocasts of pterosaurs.[9] However, these endocasts have no overlap with any described fossils of Arambourgiania and, according to a 2014 study, most likely belong to birds instead on the basis of their length of around 3 centimetres (1.2 in) and bird-like structure.[10]

Ulna bone from the Ouled Abdoun Basin of Morocco that may belong to Arambourgiania

In 2018, paleontologists Nicholas Longrich, David Martill, and Brian Andres described an incomplete left ulna of the "Sidi Chennane azhdarchid" from the Ouled Abdoun Basin of Morocco which they tentatively identified as ?Arambourgiania. This ulna came from the Sidi Chennane mine located in the basin, a group of phosphatic sandstones dating to the Maastrichtian, which contains a host of other pterosaur genera from several different families.[11]

In 2024, the describers of the genus Inabtanin discovered a partial right humerus of a large pterosaur in the Ruseifa Phosphate Mines, near the Jordanian capital of Amman, which was where the holotype of Arambourgiania was recovered. They concluded that the specimen belonged to A. philadelphiae and that it is comparable in size and shape to the humerus of the type species of Quetzalcoatlus, Q. northropi.[12]

North America

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On 15 August 1982, Ralph Johnson unearthed an azhdarchid cervical vertebra (YPM VPPU 023497) from rock layers of the Maastrichtian-aged Navesink Formation located in Monmouth County, New Jersey in the United States.[13] In 1983, this vertebra was referred to Titanopteryx (=Arambourgiania) sp. on the basis of its elongation and lateral compression, though it is extremely poorly preserved.[14] The assignment of this specimen came into question in the 2025 description of Infernodrakon, where the authors considered the specimen as aff. Arambourgiania sp. due to its cross section and the lack of further study of the fossil.[15] Between 1971-73, two American fossil collectors collected a set of three pterosaur specimens, YPM-PU 21820, YPM-PU 22359 and YPM-PU 21821, consisting of a cervical vertebra, a humerus, and a paired femur and tibia respectively. All of these fossils had been found in the Early Campanian-aged Merchantville Formation of New Castle County, Delaware in the United States.[16] Initially described as indeterminate pterosaur or later Pteranodon, in 1993 they were tentatively attributed to as cf. ?"Titanopteryx".[17] In 2014, these specimens were later labeled as Azhdarchidae indet.,[10] but a 2021 study argued that the cervical vertebra (YPM-PU 21820) belongs to a pteranodontid or even Pteranodon itself based on its pneumaticity.[18]

In 1999, a fragmentary azhdarchid cervical vertebra (MPPM 2000.23.1) was discovered by Memphis local Wendy Melton-Beeson and collected by paleontologist T. Lynn Harrell Jr. that same year. The cervical vertebra had been taken from strata in Selmer, Tennessee belonging to the lower Coon Creek Formation. These strata date to the Late Campanian age of the Late Cretaceous,[19] making them older than the Maastrichtian-aged remains found in Jordan.[2] In 2016, it was referred to A. philadelphiae on the basis of its size, morphology, and differences between it and the cervical vertebrae of Quetzalcoatlus, which may potentially extend the geographic range of Arambourgiania to North America.[19] However, the assignment of this specimen to A. philadelphiae has come under scrutiny; in 2021, it was referred to as cf. Arambourgiania, indicating a species of uncertain affinities[20] whereas in 2022, American paleontologist Gregory S. Paul stated that it may not even belong to Arambourgiania itself.[21] Finally in 2025, the describers of Infernodrakon considered this specimen as Arambourgiania sp. due to the morphological differences and temporal separation from the holotype. This same study also noted morphological similarities between MPPM 2000.23.1 and YPM VPPU 023497 from the Navesink Formation.[15]

Description

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Speculative life reconstruction

Arambourgiania was among the largest azhdarchids, though was rivalled in size by Quetzalcoatlus and Hatzegopteryx[11][21][22] (and possibly Cryodrakon).[21] Azhdarchids were split into two primary categories: short-necked taxa with short, robust beaks (i.e. Hatzegopteryx[23] and Wellnhopterus), and long-necked taxa with longer, slenderer beaks (i.e. Zhejiangopterus). Of these, Arambourgiania is of uncertain affiliation.[24] Based on the limb morphology related azhdarchids such as Zhejiangopterus and Quetzalcoatlus and pterosaurs at large, in addition to azhdarchid tracks from South Korea, Arambourgiania was likely quadrupedal.[25][26][22] As a pterosaur, Arambourgiania would have been covered in hair-like pycnofibres, and had extensive wing-membranes, which would have been distended by a long wing-finger.[24] There have been various models of the morphology of pterodactyloid wings,[22] though based on multiple well-preserved pterosaur specimens, it is likely that azhdarchids had broad wings, with a brachiopatagium extending down to the ankle. The aspect ratio of azhdarchid wings is 8.1, similar to that of storks and birds of prey that engage in static soaring (relying on air currents to gain altitude and remain aloft).[25]

Anatomy

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Neck vertebra of Arambourgiania (F) alongside those of other azdarchids

Arambourgiania is known from fragmentary remains, with no confirmed associated specimens known. As of 2025, it is only confidently known from incomplete cervical (neck) vertebrae, a portion of a dorsal (back) vertebra or posterior (towards the body) cervical vertebra, parts of the wings, and a fragment of the femur (thighbone). The holotype of Arambourgiania, UJA VF1, consists of a very elongated cervical vertebra, probably the fifth one in the cervical series. Today the middle section is missing; the original material combined with its plaster was about 62 cm (2 ft 0.41 in) long, but had been sawed into three parts. However, later, more accurate estimates based on the related Quetzalcoatlus put its estimated length at 77–78 cm (2 ft 6.31 in – 2 ft 6.71 in). Most of the fossil now consists of an internal infilling or mold; the thin bone walls are missing on most of the surface. The holotype does not present the whole vertebra; a piece is absent from its posterior end as well.[2]

The posterior (back) portion of the vertebra is 44 mm (1.73 in) horizontally at its smallest point and 55 mm (2.17 in) vertically at its smallest point with a near circle-shaped cross section. As for the anterior (front) portion, the carina sagitalis ventralis is blunt and merges into the spinous process, making it have a low and slim profile. Arambourgiania preserves a circular, internal tube inside of the cervical vertebrae, a feature observed in other azhdarchids and the Dsungaripteridae.[3] The pneumatophores of the holotype cervical vertebra are significantly larger than its neural canal, a trait corresponding with maturity indicating adulthood, though to an extent unseen in other azhdarchids. Arambourgiania lacks a dorsally (top) positioned pneumatic foramen above its neural canal, a trait typically present in relatives like Cryodrakon, distinguishing it from other azhdarchids.[27] Fragments of other cervical vertebrae have been unearthed from Jordan as well as in the United States, though none as complete as the holotype. A fragment of a neural arch from either an anterior dorsal vertebra or a cervical 7, but it is too incomplete and worn to surmise much information.[4]

Of the topotype specimens, several come from the limbs. The proximal portion of a left metacarpal IV is known, with a preserved length of 128 mm (5.04 in). The internal structure has a D-shaped cross section and a thin covering of cortical bone. The distal (part away from body) portion of an abraded left femur was also unearthed, with a total preserved length of 142 mm (5.59 in). Its shaft has an ovular to compressed ovular cross section from shaft to distal end. The bone is gently curved, as is typical of femora. A probable distal radius fragment is 108 mm (4.25 in), though is extremely worn. Its posterior end is more expanded at its proximal end than the anterior end, a trait typical of pterosaur wings.[4] An incomplete ulna unearthed in Morocco was referred to Arambourgiania and has a preserved length of 362 mm (1 ft 2.25 in) leading to an estimated complete length of 600–700 mm (1 ft 11.62 in – 2 ft 3.56 in). As for its anatomy, it is laterally compressed with a broad tubercule on its distal end and low flange, similar to the condition seen in Azhdarcho.[11] The incomplete right humerus shaft referred to Arambourgiania measures 185 mm (7.28 in) in preserved length and 80 mm (3.15 in) in minimum dorsoventral (top down) width, making it smaller than that of Quetzalcoatlus. It bears both an ovular and D-shaped cross section at different points of the humerus' width as well as thin, 2 mm (0.08 in) cortical bone. This indicates a humerus composed of 90% air by volume, making it extremely well adapted for flight.[12]

Size

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Size of Arambourgiania (center left) compared to a human, Nyctosaurus (beside the human), and Quetzalcoatlus northropi (right). Note the very similar size of Arambourgiania and Q. northropi

Frey and Martill estimated the total length of the holotype to have been 77–78 cm (2 ft 6.31 in – 2 ft 6.71 in), using for comparison the relative position of the smallest diameter of the shaft of the fifth cervical vertebra of Quetzalcoatlus. The total neck length was extrapolated at about 3 m (9 ft 10 in) using the same method. From the relatively slender vertebra, the length dimension was then selected to be compared to that of Quetzalcoatlus as well, estimated at 66 cm (2 ft 2 in) long, which results in a ratio of 1.18. This neck length is among the largest known for a non-sauropod tetrapod, exceeding those of animals like the extant giraffe, the extinct mammals Paraceratherium, the extinct reptile Tanystropheus, and dinosaurs like Gigantoraptor and Therizinosaurus.[5][28] Applying that 1.18 ratio to the overall size, Frey and Martill in the late 1990s concluded that the wingspan of Arambourgiania was 11 to 13 m (36 to 43 ft), larger than the estimated wingspan of Quetzalcoatlus, which measured 10 to 11 m (33 to 36 ft). This would have made Arambourgiania the largest pterosaur ever known.[3][2] In 1997, paleontologist Lorna Steel and colleagues reconstructed a life-sized skeleton of Arambourgiania based on better-known related pterosaurs. They set its wingspan at 11.5 m (38 ft), within the range of Frey and Martill's estimate.[8]

Subsequently, the estimates proposed by Frey and Martill in the late 1990s were taken into question, with later estimates of the wingspan of Arambourgiania being more moderate. This was due to the remains being too fragmentary to estimate a gigantic size. In 2003, the researchers who described the related pterosaur Phosphatodraco stated that the wingspan of Arambourgiania was more likely at 7 m (23 ft), though this measurement was not given a rationale.[29] In 2010, paleontologists Mark Witton and Michael Habib argued that a 7 m (23 ft) wingspan is an underestimate for Arambourgiania, while a 11 to 13 m (36 to 43 ft) wingspan would be too much.[30]

In his 2022 pterosaur book, Paul proposed that Arambourgiania had a wingspan of 8 to 9 m (26 to 30 ft), making it smaller than that of Quetzalcoatlus northropi, which he kept at 10 to 11 m (33 to 36 ft). Arambourgiania would have also had a smaller wingspan than that of the related Hatzegopteryx from Romania, which Paul situated at 10 to 12 m (33 to 39 ft). Just like both Arambourgiania and Quetzalcoatlus, Hatzegopteryx is also among the largest known flying animals to ever exist.[31] In a 2024 study, the wingspan of Arambourgiania was estimated to be around 10 m (33 ft) based on a large humerus comparable in size to that of Q. northropi. This new estimate for the wingspan of Arambourgiania is slightly larger than Paul's 2022 estimate, but does not surpass the wingspan of Quetzalcoatlus.[12] In 2018, "Sidi Chennane Azhdarchid" ulna that was referred to Arambourgiania was estimated to be smaller than other Arambourgiania individuals, with a wingspan of around 9 m (30 ft).[11]

Classification

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Skeletal reconstructions of Arambourgiania (C) and the related Hatzegopteryx (A), and Quetzalcoatlus lawsoni (D), with known parts in gray

Arambourgiania was initially assigned to a newly named subfamily called Azhdarchinae by Nesov in 1984, though it was still known as "Titanopteryx" at the time of the assignment. Azhdarchinae also included the pterosaurs Azhdarcho and Quetzalcoatlus. Nesov assigned the subfamily as part of the family Pteranodontidae, based on its members featuring toothless beaks just like the pteranodontids.[32] Unaware of the creation of Azhdarchinae, American paleontologist Kevin Padian created the family Titanopterygiidae, which included both "Titanopteryx" and Quetzalcoatlus, which was created on the basis of cervical form and proportions, and it was differentiated from Pteranodontidae, which also received a definition by Padian the same year.[33][34] Two years later, in 1986, Padian would become aware of the existence of Azhdarchinae and would make Titanopterygiidae a junior synonym of it, as he believed that the diagnoses of the cervical vertebrae for both groups were identical. He removed Azhdarchinae from Pteranodontidae based on his previous diagnoses, and he would further elevate it to family level, creating Azhdarchidae as it is known today.[35][34] Since the 1980s, many genera have been named or reassigned to the genus, with fossils of azhdarchids known from every continent besides Antarctica.[10][15]

The placement of Arambourgiania within the family Azhdarchidae has been consistent in various studies, which is in a derived (advanced) position in the subfamily Quetzalcoatlinae. However, its specific location within the group has been somewhat disputed. One of the closest relatives of Arambourgiania is Quetzalcoatlus, as both pterosaurs have consistently been found together in multiple phylogenetic analyses, either as sister taxa or close to each other.[36][34][37] However, there have also been several studies opposing this placement and have instead favored a closer relationship between Arambourgiania and the azhdarchids Mistralazhdarcho and Aerotitan.[38][39][40]

Below are two cladograms showing different studies regarding the position of Arambourgiania within Azhdarchidae. The first one is based on the phylogenetic analysis by American paleontologist Brian Andres in 2021, which places Arambourgiania within Quetzalcoatlinae as the sister taxon to both species of Quetzalcoatlus, Q. northropi and Q. lawsoni.[34] The second cladogram is based on the 2023 study by paleontologist Rodrigo Pêgas and colleagues, in which they placed Arambourgiania in a trichotomy with Mistralazhdarcho and Aerotitan within Quetzalcoatlinae, contrasting its placement as the sister taxon of Quetzalcoatlus.[40]

In some studies, the azhdarchids Mistralazhdarcho (above) and Aerotitan (below) have been recovered as the closest relatives of Arambourgiania

Paleobiology

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Feeding and ecological niche

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Life reconstruction of two Arambourgiania fighting over a small theropod

In 2008, Mark Witton and Darren 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.[25]

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. 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).[25][11]

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.[25] The prevalence of Arambourgiania fossils in oceanic deposits suggests that it died at sea or was washed out.[3][41] This includes the oceanic deposits at the Coon Creek Formation,[19] the Ruseifa Formation,[1][2] and the Khouribga Phosphates.[11] However, in a 2021 study , Labita and Martill noted that azhdarchids might have been less terrestrial than suggested by Witton and Naish, since the Moroccan azhdarchid fossils were from marine strata, as was Arambourgiania. They noted that no azhdarchids had been found in truly terrestrial strata,[10] and proposed they could instead have been associated with aquatic environments, such as rivers, lakes, marine and off-shore settings.[42]

Locomotion

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Length of the holotype cervical vertebra of Arambourgiania (first above) compared to other azhdarchid cervical vertebrae

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]

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

Azhdarchids are the only group of pterosaurs to which trackways have been confidently assigned, such as Haenamichnus from Korea, which matches this group in shape, age, and size.[43] 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.[24] Terrestrial locomotion in azhdarchids like Quetzalcoatlus likely involved a pacing gait, wherein the limbs on one side of the body would move at roughly the same time, followed by those of the opposite side. For example, the forelimb on one side of the body would lift off the ground and move forward first, to avoid colliding with the hind foot, and the hind limb would follow suit. The forefoot would be planted in the ground just before the hind foot. Once the stride completed, the same process would repeat on the opposite side of the body.[24][22]

A 2024 study by paleontologist Kierstin Rosenbach and colleagues included the description of the humerus of Arambourgiania. They compared it to that of soaring birds and suggested that Arambourgiania itself was also a soarer. The humeral shaft displays several adaptations to soaring flight, such as its arrangement of diaphyseal ridges which are similar to those in vultures. The humerus also lacks struts, a trait associated with torsional loadings from soaring. Soaring is defined as sustained powered flight that needs launch and maintenance flapping, whereas gliding is sustained by gravity. This is in contrast to the contemporary pterosaur Inabtanin, which had a style of flight closer to those of continuously flapping birds. The ability for large azhdarchids like Arambourgiania to soar corresponds with geologic data, with Arambourgiania living in a marine environment suggesting that it and other giant azhdarchids were volant and could implement thermal soaring in their flight.[12]

Neck biomechanics

[edit]
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)

The lithe, thin-walled vertebrae of Arambourgiania indicate the neck was much weaker than that of Hatzegopteryx. This can be quantified using relative failure force, which is the bone failure force of a vertebra divided by the body weight of the pterosaur that it belongs to, estimated at 180 to 250 kg (400 to 550 lb) for Arambourgiania and Hatzegopteryx. While Arambourgiania's neck vertebrae fail at about half of its body weight, the posterior neck vertebrae of Hatzegopteryx can withstand anywhere between five and ten body weights, depending on the loading of the bone. Even the hypothetically longer anterior neck vertebrae of Hatzegopteryx would be able to withstand four to seven body weights.[23] Although the centrum of Arambourgiania is much more lighltly built than that of Hatzegopteryx, their ratios of bone radius to bone thickness (R/t)[26] are roughly the same (9.45 for Hatzegopteryx and 9.9 for Arambourgiania). This may represent a compromise between increasing bending strength and buckling strength. Higher R/t ratios lead to improved bending strength, but weaker buckling strength. To compensate for this, Hatzegopteryx shows a number of other adaptations to improve buckling strength, namely the distinctive internal structures of the bones and the large articular joints of the vertebrae, the latter of which helps to distribute stress.[23] In order to support the robust head, the neck of Hatzegopteryx was likely strongly muscled, in contrast to that of Arambourgiania with its fewer muscle attachments.[23]

Paleoecology

[edit]
Map of Europe and the Middle East during the Maastrichtian, showing the extent of the Tethys Sea.

Fossils of Arambourgiania are confidently known only from Ruseifa, Jordan, though specimens potentially belonging to the genus have been described from Morocco and the Southern United States as well. In Jordan and Morocco, Arambourgiania fossils come from sites that during the Maastricthian were covered by the Tethys Sea.[44] This was a large sea extending across Europe, the Middle East, and North Africa during the Late Cretaceous and Paleogene. Other Maastrichtian-aged azhdarchid pterosaurs are known from the Tethys Sea as well, including Inabtanin from Jordan, Phosphatodraco, a Quetzalocatlus-like azhdarchid, and possibly Arambourgiania itself from Morocco,[29][11] Hatzegopteryx and several other azhdarchids from Romania,[45][46][47] and indeterminate azhdarchids from Spain[48] and France.[49][10]

Jordan

[edit]

In Jordan, Arambourgiania fossils have been unearthed from a series of phosphate mines in Ruseifa dating to the Maastrichtian age of the Late Cretaceous period (72-66 Ma). During this time period, Jordan was underwater and located at the southern margin of the Mediterranean Tethys Sea.[50] These phosphate mines have been labeled as the Ruseifa Formation,[50] Amman Formation,[51] or the Al-Hasa Phosphorite Formation, with little consensus on a formal name.[52] However, the Ruseifa Formation is the most recently adopted and commonly used title. The Ruseifa Formation, the underlying Wadi Sir Formation, and the overlying Muwaqqar Formation make up the Balqa Group. The Ruseifa Formation is composed of four units, the first of which is the origin of many Arambourgiania fossils.[3] The phosphates where Arambourgiania is known from have been interpreted as a deep marine environment whereas the higher layers belong to shoreline environments.[12] However, other researchers have stated that these phosphates are derived from a shallow marine deposit.[53][54]

Fossils here are preserved through the infilling of bone by limestone matrix, resulting in well preserved, 3D fossils being available for study.[12] Contemporary fauna to Arambourgiania from the phosphates includes a diversity of marine reptiles including the mosasaurids Globidens, Platecarpus, Prognathodon, and Halisaurus, many of which are known from teeth, indeterminate elasmosaurids, indeterminate chelonoids, and an isolated crocodyliform tooth.[52] Fish are also prevalent in this formation, with bony fishes known from the enchodontid Enchodus, aulopiform Stratodus, pycnodontid Stephanodus, anguillioform Pseudoegertonia, and indeterminate pycnodontids. As for the cartilaginous fishes, the sharks Cretolamna, Scapanorhynchus, Squalicorax, Plicatoscyllium, and the ray Rhombodus have been found in the Ruseifa phosphates.[55] This faunal composition is very similar to that of the Maastrichtian of Syria and the rest of the Mediterranean Tethys Sea.[41][52]

See also

[edit]

References

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