Arambourgiania Philadelphiae
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''Arambourgiania'' is an extinct genus of
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 ...
pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...
from 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 interval ...
stage) of
Jordan Jordan ( ar, الأردن; tr. ' ), officially the Hashemite Kingdom of Jordan,; tr. ' is a country in Western Asia. It is situated at the crossroads of Asia, Africa, and Europe, within the Levant region, on the East Bank of the Jordan Rive ...
, and possibly the
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
.Harrell, T. Lynn Jr.; Gibson, Michael A.; Langston, Wann Jr. (2016). "A cervical vertebra of ''Arambourgiania philadelphiae'' (Pterosauria, Azhdarchidae) from the Late Campanian micaceous facies of the Coon Creek Formation in McNairy County, Tennessee, USA" ''Bull. Alabama Mus. Nat. Hist.'' 33:94–103 ''Arambourgiania'' was among the largest members of its family, the Azhdarchidae, and it is also one of the largest flying animals ever known. The incomplete left
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
of the "Sidi Chennane azhdarchid" from
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 t ...
may have also belonged to ''Arambourgiania''.


History of discovery

In the early 1940s, a railway worker during repairs on the Amman- Damascus railroad near
Russeifa Russeifa, also spelled ''Russiefa'', ( ar, الرصيفة) is a city in Zarqa Governorate in Jordan. It had a population of 472,604 inhabitants in 2015, making it the fourth-largest city in Jordan, after Amman, Irbid, and Zarqa. Geography The ...
found a two foot long fossil bone. In 1943 this was acquired by the director of a nearby
phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...
mine, Amin Kawar, who brought it to the attention of a British archeologist, Fielding, after the war. This generated some publicity — the bone was even shown to
Abdullah I of Jordan AbdullahI bin Al-Hussein ( ar, عبد الله الأول بن الحسين, translit=Abd Allāh al-Awwal bin al-Husayn, 2 February 1882 – 20 July 1951) was the ruler of Jordan from 11 April 1921 until his assassination in 1951. He was the Emi ...
— but more importantly, it made the scientific community aware of the find. In 1953 the fossil was sent to
Paris Paris () is the Capital city, capital and List of communes in France with over 20,000 inhabitants, most populous city of France, with an estimated population of 2,165,423 residents in 2019 in an area of more than 105 km² (41 sq mi), ma ...
, where it was examined by Camille Arambourg of the
Muséum national d'histoire naturelle The French National Museum of Natural History, known in French as the ' (abbreviation MNHN), is the national natural history museum of France and a ' of higher education part of Sorbonne Universities. The main museum, with four galleries, is loc ...
. In 1954, he concluded the bone was the wing
metacarpal In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ar ...
of a giant pterosaur. In 1959, he named a new genus and species: ''Titanopteryx philadelphiae''. The genus name meant "titan wing" in Greek; the specific name refers to the name of Amman in Antiquity: Philadelphia. 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. In 1975
Douglas A. Lawson Douglas A. Lawson (born 1947) is a geologist, paleontologist, and computer scientist. In 1971 Lawson discovered wing bone fossils from a giant pterosaur embedded in a sandstone outcropping at Big Bend National Park, Texas. At the time the fossil ...
, studying the related '' Quetzalcoatlus'', concluded the bone was not a metacarpal but a cervical vertebra. In the eighties, Russian
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Lev Nesov Lev may refer to: Common uses * Bulgarian lev, the currency of Bulgaria *an abbreviation for Leviticus, the third book of the Hebrew Bible and the Torah People and fictional characters * Lev (given name) *Lev (surname) Places *Lev, Azerbaijan, ...
was informed by an entomologist that the name ''
Titanopteryx ''Titanopteryx'' is a genus of fly in the family Simulidae. Simuliidae Nematocera genera {{Chironomidae-stub ...
'' had already been given by Günther Enderlein to a fly from the Simulidae family in 1935. Therefore, in 1989 he renamed the genus into ''Arambourgiania'', honoring Arambourg. However, the name "Titanopteryx" was informally kept in use in the West, partially because the new name was assumed by many to be a ''nomen dubium''. 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 find. However, after their departure to Europe engineer Rashdie Sadaqah of the mine investigated further and in 1996 established it had been bought from the company in 1969 by geologist Hani N. Khoury who had donated it in 1973 to the University of Jordan; it was still present in the collection of this institute and now could be restudied by Martill and Frey. Frey and Martill rejected the suggestion that ''Arambourgiania'' was a ''nomen dubium'' or identical to ''Quetzalcoatlus'' and affirmed its validity in relation to "Titanopteryx". Nesov in 1984 had placed the species within Azhdarchinae, back then part of the Pteranodontidae; the same year Kevin Padian placed it within Titanopterygidae. Both concepts have fallen into disuse now that such forms are commonly assigned to the Azhdarchidae. In 2016, an Azhdarchid cervical vertebra was described from the Coon Creek Formation of McNairy County, Tennessee and referred to ''Arambourgiania philadelphiae''. This find extends ''Arambourgianias geographic range to North America. In 2018, topotype specimens were located in Bavarian State Collection for Palaeontology and Geology in Munich, Germany that were placed there in 1966 from Jordan and probably represent additional elements of the holotype individual. These include the "fragments of two cervical vertebrae, a neural arch, a left femur, a ?radius, and a metacarpal IV" and other indeterminate fragments.


Description

The holotype, VF 1, consists of a very elongated cervical vertebra, probably the fifth. Today the middle section is missing; the original find was about long, but had been sawed into three parts. Most of the fossil consists of an internal infilling or mold; the thin bone walls are missing on most of the surface. The find had not presented the whole vertebra; a piece was absent from its posterior end as well. Frey and Martill estimated the total length to have been , using for comparison the relative position of the smallest shaft diameter of the fifth cervical vertebra of '' Quetzalcoatlus''. From this again the total neck length was extrapolated at about . From the relatively slender vertebra the length dimension was then selected to be compared to that of ''Quetzalcoatlus'', estimated at long, resulting in a ratio of 1.18. Applying that ratio to the overall size, Frey and Martill in the late 1990s concluded that the wingspan of ''Arambourgiania'' had been , compared with the wingspan of ''Quetzalcoatlus'', and that ''Arambourgiania'' was thus the largest pterosaur then known. However, the estimate proposed by Frey and Martill was taken into question and the later estimates of wingspan have been more moderate, for the remains are too fragmentary to estimate a gigantic size. The researchers who described ''Phosphatodraco'' stated that the wingspan of ''Arambourgiania'' was more likely at , but this estimate was not given a rationale. In 2010, Mark Witton and Michael Habib argued that a wingspan is an underestimate, while a wingspan is an overestimate. In 2022, Gregory S. Paul proposed that ''Arambourgiania'' had a wingspan of , smaller than that of ''Quetzalcoatlus'' or ''Hatzegopteryx''.


Classification

Below is a cladogram showing the phylogenetic placement of ''Arambourgiania'' within the clade Neoazhdarchia. The cladogram is based on a topology recovered by Brian Andres and Timothy Myers in 2013.


See also

* List of pterosaur genera * Timeline of pterosaur research * Pterosaur size


Notes


References

* Arambourg, C. (1959). "''Titanopteryx philadelphiae'' nov. gen., nov. sp. Ptérosaurien géant." ''Notes Mém. Moyen-Orient'', 7: 229–234. * Frey, E. & Martill, D.M. (1996). "A reappraisal of ''Arambourgiania'' (Pterosauria, Pterodactyloidea): One of the world's largest flying animals." ''N.Jb.Geol.Paläont.Abh.'', 199(2): 221–247. * Martill, D.M., E. Frey, R.M. Sadaqah & H.N. Khoury (1998). "Discovery of the holotype of the giant pterosaur ''Titanopteryx philadelphiae'' Arambourg 1959, and the status of ''Arambourgiania'' and ''Quetzalcoatlus''." ''Neues Jahrbuch fur Geologie und Paläontologie'', Abh. 207(1): 57–76. * Nessov, L.A., and Yarkov, A.A. (1989). "New Cretaceous-Paleogene birds of the USSR and some remarks on the origin and evolution of the class Aves". ''Trudy Zoologicheskogo Instituta AN SSSR'', 197: 78–97. [In Russian] * Steel, L., D.M. Martill., J. Kirk, A. Anders, R.F. Loveridge, E. Frey, and J.G. Martin (1997). "''Arambourgiania philadelphiae'': giant wings in small halls." ''The Geological Curator'', 6(8): 305–313. {{Portal bar, Paleontology, Cretaceous, Jordan, United States Azhdarchids Fossil taxa described in 1989 Late Cretaceous pterosaurs of Asia Late Cretaceous pterosaurs of North America