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''Aurorazhdarcho'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of ctenochasmatoid
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 to ...
known from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
period (early
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
stage) of what is now
Bavaria Bavaria ( ; ), officially the Free State of Bavaria (german: Freistaat Bayern, link=no ), is a state in the south-east of Germany. With an area of , Bavaria is the largest German state by land area, comprising roughly a fifth of the total lan ...
, southern
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
.


History


First discovery

A specimen, originally classified as ''
Pterodactylus ''Pterodactylus'' (from Greek () meaning 'winged finger') is an extinct genus of pterosaurs. It is thought to contain only a single species, ''Pterodactylus antiquus'', which was the first pterosaur to be named and identified as a flying rept ...
micronyx'' (now ''Aurorazhdarcho micronyx''), was one of the earliest, if not the earliest, documented pterosaur fossil ever found. 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 several ...
specimen of ''P. micronyx'', also known as the "Pester Exemplar", was originally part of the private fossil collection held by Archduchess Maria Anna of Austria. Evidence suggest that the Pester Exemplar was unearthed at some point between 1757, when Maria Anna was recovering from serious cases of pneumonia and tuberculosis, after which she began collecting fossils, and 1779, when the specimen was first studied by scientists. This overlaps with the possible time of discovery of the holotype specimen of ''Pterodactylus antiquus'', often considered the first pterosaur found, which was unearthed sometime between 1767 and 1784.Ősi, A., Prondvai, E., & Géczy, B. (2010). The history of Late Jurassic pterosaurs housed in Hungarian collections and the revision of the holotype of Pterodactylus micronyx Meyer 1856 (a ‘Pester Exemplar’). Geological Society, London, Special Publications, 343(1), 277–286. The Pester Exemplar consists of a jumbled and partially dis-articulated juvenile pterosaur skeleton lacking a skull. Because of this, and the unusual (and at the time totally unknown) anatomy of pterosaurs, the specimen was originally misidentified as a decapod
crustacean Crustaceans (Crustacea, ) form a large, diverse arthropod taxon which includes such animals as decapods, seed shrimp, branchiopods, fish lice, krill, remipedes, isopods, barnacles, copepods, amphipods and mantis shrimp. The crustacean group ...
when it was first studied by
Ignaz von Born Ignaz Edler von Born, also known as Ignatius von Born ( hu, Born Ignác, ro, Ignațiu von Born, cs, Ignác Born) (26 December 1742 in Alba Iulia, Grand Principality of Transylvania, Habsburg monarchy – 24 July 1791 in Vienna), was a mineral ...
, a prominent Enlightenment naturalist. In 1871, the specimen was sold, along with the rest of Maria Anna's collection, to the Royal Hungarian University of Buda in
Hungary Hungary ( hu, Magyarország ) is a landlocked country in Central Europe. Spanning of the Carpathian Basin, it is bordered by Slovakia to the north, Ukraine to the northeast, Romania to the east and southeast, Serbia to the south, Croatia a ...
, from which it was later transferred to the collection of Pest University when the university relocated. (In 1950, the name of the university changed again to
Eötvös Loránd University Eötvös Loránd University ( hu, Eötvös Loránd Tudományegyetem, ELTE) is a Hungarian public research university based in Budapest. Founded in 1635, ELTE is one of the largest and most prestigious public higher education institutions in Hung ...
). In 1856,
Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born at Frankfurt am Ma ...
illustrated the Pester Exemplar, and designated it the type specimen of his new species ''Pterodactylus micronyx'' after comparison with the newly recognized and better-preserved pterosaur specimens, all of which were lumped together at that time into the single genus ''Pterodactylus''. However, by 1960,
Peter Wellnhofer Peter Wellnhofer (born Munich, 1936) is a German paleontologist at the Bayerische Staatssammlung fur Paläontologie in Munich. He is best known for his work on the various fossil specimens of ''Archaeopteryx'' or "Urvogel", the first known bird. W ...
was not able to locate the specimen in the
Hungarian National Museum The Hungarian National Museum ( hu, Magyar Nemzeti Múzeum) was founded in 1802 and is the national museum for the history, art, and archaeology of Hungary, including areas not within Hungary's modern borders, such as Transylvania; it is not to ...
or in the collections of Eötvös Loránd University. Wellnhofer therefore considered the specimen lost, and designated a
neotype In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
. The Pester Exemplar was re-discovered in 1982, during a re-organization of the collections in Eötvös Loránd University, and so the status of type specimen for ''P. micronyx'' reverted to it.


Discovery of adult specimen

In 1999, amateur paleontologist Peter Katschmekat uncovered a fossil of a pterosaur in the Blumenberg Quarry, northwest of
Eichstätt Eichstätt () is a town in the federal state of Bavaria, Germany, and capital of the district of Eichstätt. It is located on the Altmühl river and has a population of around 13,000. Eichstätt is also the seat of the Roman Catholic Diocese ...
, in layers of the Solnhofen limestone. The fossil was prepared by Gerd Stübener and acquired by the Swiss ''
Naturhistorisches Museum Basel Natural History Museum Basel (german: Naturhistorisches Museum Basel) is a natural history museum in Basel, Switzerland that houses wide-ranging collections focused on the fields of zoology, entomology, mineralogy, anthropology, osteology and pal ...
''. Based on the specimen, the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
''Aurorazhdarcho primordius'' was named and described by
Eberhard Frey Eberhard is an old Germanic name meaning the strength or courage of a wild boar. People First name *Eberhard of Friuli (815–866), Duke and key figure in the Carolingian Empire *Eberhard of Béthune (died 1212), Flemish grammarian *Eberhard I, Du ...
, Christian A. Meyer and Helmut Tischlinger in 2011. The generic name is derived from
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 of the ...
''aurora'', "dawn" and Kazakh ''Azhdarcho'', the name of a mythical dragon. The specific name means "primordial" or "the very first" 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 several ...
of ''Aurorazhdarcho'', NMB Sh 110, was found in the Upper Eichstätt Formation dating to the early
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
stage of the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
, 150
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...
. It consists of a nearly complete
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
, three-dimensionally preserved on a single slab with the counterplate having been removed, lacking the skull and neck. However, the head and neck are present as impressions, movement of these elements on the bottom of the lagoon the carcass descended on leaving a natural mould covered by organic remains in the form of
calcium phosphate The term calcium phosphate refers to a family of materials and minerals containing calcium ions (Ca2+) together with inorganic phosphate anions. Some so-called calcium phosphates contain oxide and hydroxide as well. Calcium phosphates are white ...
flakes made visible by the UV-photography of Tischlinger. That the head and neck are now absent is explained by the animal being killed by a partially failed attack by a predator, almost ripping off these parts, after which the body quickly sank. Subsequent bloating would then have caused the head to drift away. The individual was probably subadult and, as indicated by the fusion of the ischiopubes in the pelvis, a male as it lacks a wide birth canal. In 2013, Bennett compared the holotype specimen of ''Aurorazhdarcho primordius'' to the specimens of the species ''"
Pterodactylus ''Pterodactylus'' (from Greek () meaning 'winged finger') is an extinct genus of pterosaurs. It is thought to contain only a single species, ''Pterodactylus antiquus'', which was the first pterosaur to be named and identified as a flying rept ...
" micronyx'' (named by
Christian Erich Hermann von Meyer Christian Erich Hermann von Meyer (3 September 1801 – 2 April 1869), known as Hermann von Meyer, was a German palaeontologist. He was awarded the 1858 Wollaston medal by the Geological Society of London. Life He was born at Frankfurt am Ma ...
in 1856). He found that some of the supposed distinguishing features of ''A. primordius'', such as length of the metacarpal IV equal to the length of the radius and ulna, or first wing finger phalanx being the longest element in the wing, are in fact also present in the skeletons of the specimens of ''"P." micronyx''; in addition, limb proportions in the holotype specimen of ''A. primordius'' matched those of ''"P." micronyx''. Bennett concluded that ''Aurorazhdarcho primordius'' and ''"Pterodactylus" micronyx'' are in fact conspecific; he retained ''Aurorazhdarcho'' as a genus distinct from ''Pterodactylus'', but he considered the specific epithet ''primordius'' to be a junior synonym of the epithet ''micronyx''. As noted by Bennett, the taxonomy of ''Aurorazhdarcho micronyx'' is further complicated by the author's earlier suggestion that ''"Pterodactylus" micronyx'' specimens are in fact juveniles of ''
Gnathosaurus ''Gnathosaurus'' (meaning "jawed lizard") is a genus of ctenochasmatid pterosaur containing two species: ''G. subulatus'', named in 1833 from the Solnhofen Limestone of Germany, and ''G. macrurus'', known from the Purbeck Limestone of the UK. ...
subulatus'', which if confirmed would make the former a junior synonym of the latter. ''G. subulatus'' is only known from the holotype incomplete mandible and the isolated skull; this makes comparison difficult, because skulls are only known in small, juvenile specimens of ''A. micronyx'', while the large specimens, including NMB Sh 110, lack it. Thus, Bennett stated that until a large specimen with an associated skull and skeleton is discovered, which would make it possible to establish or reject the synonymy of ''Gnathosaurus'' and ''Aurorazhdarcho'', the genus ''Aurorazhdarcho'' should be used to contain the species ''A. micronyx''. Additional likely synonyms of ''P. micronyx'' are ''Pterodactylus nettecephaloides'' and ''P. redenbacheri''. While both of these names are older than ''A. micronyx'', Olshevsky in 1991 considered them '' nomina oblita'' ("forgotten names"), and therefore not senior synonyms.


Description

''Aurorazhdarcho'' was a relatively small pterosaur. The combined elements of a single wing of the type specimen have the extended length of . The impressions indicate the head was elongated with the eye socket in a high position. The snout might have carried a high, rounded, crest. The impression of the neck is relatively short and shows signs of a throat pouch. The shoulder girdle was by the describers reconstructed with a low shoulder joint. The legs were relatively long with the
shinbone The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
over a third longer than the
thighbone The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
. In the wing the
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 ...
s were very elongated, the fourth metacarpal being longer than the lower arm. The first to fourth phalanges of the wing finger have the respective length of , , , and . As mentioned before, the skull is only known in small, juvenile specimens. The skull was elongate, with its superior margin markedly concave upward, and slender, elongated rostrum. Its jaws had up to 18 closely spaced, long, gently curving teeth per jaw side. The anterior teeth were angled anteriorly, while the posterior teeth were shorter and more upright. The upper and lower teeth interlocked; according to Bennett (2013) they formed a basket for sieving food items from water, though the lack of a pumping mechanism suggests a function more akin to
spoonbill Spoonbills are a genus, ''Platalea'', of large, long-legged wading birds. The spoonbills have a global distribution, being found on every continent except Antarctica. The genus name ''Platalea'' derives from Ancient Greek and means "broad", refe ...
jaws, wading with the jaws open and closing them when catching individual prey, much as in other ctenochasmatoids.Wilton, Mark P. (2013). Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press. .


Phylogeny

''Aurorazhdarcho'' was by the describers assigned to the
Azhdarchoidea Azhdarchoidea (or azhdarchoids) is a group of pterosaurs within the suborder Pterodactyloidea, more specifically within the group Ornithocheiroidea. Pterosaurs belonging to this group lived throughout the Early Cretaceous, Early and Late Cretaceo ...
and placed in a new family, the Protazhdarchidae, of which it is the only named member. It would then be the oldest known azhdarchoid with the dubious exception of the "
Doratorhynchus ''Doratorhynchus'' is a generic replacement name for ''Pterodactylus validus'', in 1875 suggested by Harry Govier Seeley. Today it is considered a '' nomen vanum''. In 1870 Richard Owen named ''Pterodactylus validus'' based on holotype NHMUK 406 ...
" material. The describers rejected the use of a
cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...
analysis to establish the
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
. Using the comparative method they found that ''Aurorazhdarcho'' was closest in proportions to the
ctenochasmatid Ctenochasmatidae is a group of pterosaurs within the suborder Pterodactyloidea. They are characterized by their distinctive teeth, which are thought to have been used for filter-feeding. Ctenochasmatids lived from the Late Jurassic to the Early ...
''
Ctenochasma ''Ctenochasma'' (meaning "comb jaw") is a genus of Late Jurassic ctenochasmatid pterosaur belonging to the suborder Pterodactyloidea. Three species are currently recognized: ''C. roemeri'' (named after Friedrich Adolph Roemer), ''C. taqueti'', a ...
elegans'', a species also known from the Solnhofen, but they concluded that ''Aurorazhdarcho'' belonged to the Azhdarchoidea because it shared certain evolutionary key adaptations with that group, the most decisive of which they considered to be the low shoulder joint. The position of the shoulder joint with ''Ctenochasma'' could not be established but the describers assumed it was higher because of the closer relation of that taxon with ''
Pterodactylus ''Pterodactylus'' (from Greek () meaning 'winged finger') is an extinct genus of pterosaurs. It is thought to contain only a single species, ''Pterodactylus antiquus'', which was the first pterosaur to be named and identified as a flying rept ...
''. Bennett in 2013, based on the analysis of all known specimens of ''Aurorazhdarcho micronyx'' and taking its short neck and pedal morphology into account, considered this species to be a ctenochasmatid. 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 d ...
following Vidovic and Martill, 2017, highlighting the positions of several possible aurorazhdarchians, including ''Aurorazhdarcho'':


See also

*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic era. Although pterosaurs w ...


References

{{Portal bar, Paleontology, Germany Late Jurassic pterosaurs of Europe Ctenochasmatoids Fossil taxa described in 2011 Tithonian life