Cretornis Hlavaci
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''Cretornis'' is a
pterosaur Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
from the late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...
period (
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
stage) of what is now the Jizera Formation in the
Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Central Europe. The country is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the south ...
, dating to about 92 million years ago. It only contains a single species, ''Cretornis hlavaci''.


Discovery and naming

The fossils were discovered in
1880 Events January *January 27 – Thomas Edison is granted a patent for the incandescent light bulb. Edison filed for a US patent for an electric lamp using "a carbon filament or strip coiled and connected ... to platina contact wires." gr ...
by workers at a
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
quarry in
Zářecká Lhota Zářecká Lhota is a municipality and village in Ústí nad Orlicí District in the Pardubice Region of the Czech Republic. It has about 200 inhabitants. Zářecká Lhota lies approximately west of Ústí nad Orlicí, east of Pardubice, and e ...
near the town of
Choceň Choceň (; ) is a town in Ústí nad Orlicí District in the Pardubice Region of the Czech Republic. It has about 8,600 inhabitants. Administrative division Choceň consists of seven municipal parts (in brackets population according to the 2021 c ...
, who were getting gravel to repair a local road. A certain Mrs. Tomková, a croupier from Choceň, then alerted František Hlaváč, a Choceň pharmacist and fossil collector, to the find. He recognised exceptionality of that one, secured the rest of the fossils and then sent them to naturalist Professor Antonín Frič in Prague. In 1881, Antonín Frič identified it as a prehistoric toothed bird the size of a recent
swan Swans are birds of the genus ''Cygnus'' within the family Anatidae. The swans' closest relatives include the goose, geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form the tribe (biology) ...
and named it as the
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
''Cretornis Hlaváči''. The generic name is derived from
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''creta'', "chalk", in reference to the Cretaceous, and Greek ὄρνις, ''ornis'', "bird", as Frič originally thought that the fossil bones belonged to some kind of ancient toothed bird (similar to the genus ''
Ichthyornis ''Ichthyornis'' (meaning "fish bird", after its fish-like vertebrae) is an extinct genus of toothy seabird-like ornithuran from the late Cretaceous period of North America. Its fossil remains are known from the chalks of Alberta, Alabama, Kansa ...
''). The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
honors Hlaváč. Today, the holotype fossils of this pterosaur are stored in the National Museum in Prague. There is also unconfirmed information that similar bones (perhaps also pterosaur bones) were found in other places around the quarry before and during this discovery, but later they were apparently lost. Afterwards it was realized that the find represented some pterosaur. The name was incorrectly emended by
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was a British naturalist, geologist and writer of numerous books on natural history. He was known for his contributions to zoology, paleontology, and biogeography. He worked extensively in cata ...
into ''Ornithochirus hlavatschi'' Fritsch 1881, in 1888. As scientific species names are not allowed to contain diacritical signs, the specific name had to be transcribed. Lydekker chose to write it as if it, and Frič himself, were German, as
Bohemia Bohemia ( ; ; ) is the westernmost and largest historical region of the Czech Republic. In a narrow, geographic sense, it roughly encompasses the territories of present-day Czechia that fall within the Elbe River's drainage basin, but historic ...
at the time was under a strong German cultural influence. Frič, an ethnic Czech, in 1905 ultimately used the correct transcription ''Ornithocheirus hlavaci''. Since 1905, the
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
was typically considered a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium,'' it may be impossible to determine whether a ...
''. In 2010 however, Russian paleontologist Alexandr Averianov concluded that it should be possible to determine unique traits. In 2015 he and Czech paleontologist Boris Ekrt published a new description, concluding that ''Cretornis'' was a valid taxon. ''Cretornis'' is known from the
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
, presently preserved in the collection of the '' Národní muzeum'' (National Museum) in
Prague Prague ( ; ) is the capital and List of cities and towns in the Czech Republic, largest city of the Czech Republic and the historical capital of Bohemia. Prague, located on the Vltava River, has a population of about 1.4 million, while its P ...
as "Object 10". It was found in what Frič called the ''Mittlere Iserschichten'', today known as the Jizera Formation, dating from the Turonian. It is a partial skeleton lacking the skull. It contains a complete
humerus The humerus (; : humeri) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
(upper arm bone), an
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
, radius, wrist and two phalanges of the wing finger.Averianov, A.O. (2010).
The osteology of ''Azhdarcho lancicollis'' Nessov, 1984 (Pterosauria, Azhdarchidae) from the Late Cretaceous of Uzbekistan
" ''Proceedings of the Zoological Institute of the Russian Academy of Sciences'', 314(3): 246–317.


Description

Based on comparison with its relatives, the wingspan of ''C. hlavaci'' has by Averianov been estimated at . The humerus is in length. In 2015, Averianov indicated a single
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a Synapomorphy, derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or Outgroup (cladistics), outgroup taxa, not ...
, unique derived trait: the distal part of the humerus has a diamond-shaped cross-section.


Classification

''Cretornis'' was classified as a species in the genus ''
Ornithocheirus ''Ornithocheirus'' (from Ancient Greek "ὄρνις", meaning bird, and "χεῖρ", meaning hand) is a pterosaur genus known from fragmentary fossil remains uncovered from sediments in the United Kingdom and possibly Morocco. Several species ha ...
'' in the nineteenth century. In 1997 Coralia-Maria Jianu suggested it belonged to the
Pteranodontidae The Pteranodontidae are a Family (biology), family of large pterosaurs from the Late Cretaceous of North America and possibly other continents including Europe and Africa. The family was named in 1876 by Othniel Charles Marsh. Pteranodontids had a ...
. In 2010, Averianov thought is it was a member of the
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 ...
. More detailed comparisons of the wing bones led Averianov to conclude in 2015 that they belonged to a non-azhdarchid
azhdarchoid Azhdarchoidea (International Phonetic Alphabet, , meaning "azhdarchid-like forms") is a group of pterosaurs within the suborder Pterodactyloidea. Pterosaurs belonging to this group lived throughout the Early Cretaceous, Early and Late Cretaceous ...
, probably a member of the group
Neoazhdarchia Azhdarchoidea ( , meaning " azhdarchid-like forms") is a group of pterosaurs within the suborder Pterodactyloidea. Pterosaurs belonging to this group lived throughout the Early and Late Cretaceous periods, with one tentative member, '' Tendagu ...
with which it has many features in common. It is probably more advanced than the
Thalassodromidae Thalassodromidae (meaning "sea runners") is a group of azhdarchoid pterosaurs from the Early Cretaceous (Albian) of Brazil. All known definitive members come from the Romualdo Formation of Brazil, which include the type genus '' Thalassodromeus' ...
, and shares with ''
Montanazhdarcho ''Montanazhdarcho'' is a genus of azhdarchoid pterosaur from the Late Cretaceous period (Campanian stage) of what is now the state of Montana, United States. ''Montanazhdarcho'' is known from only one species, ''M. minor''. Discovery The holot ...
'' the trait that the distal part of the ulna has a joint surface that is placed more proximal than the tubercle. In 2018, paleontologist Nicholas Longrich and colleagues had recovered ''Cretornis'' within the family
Nyctosauridae Nyctosauridae (meaning "night lizards" or "bat lizards") is a family of specialized soaring pterosaurs of the late Cretaceous Period of North America, Africa, and possibly other continents including South America. It was named in 1889 by Henry ...
, in a more derived position than ''
Alamodactylus ''Alamodactylus'' is an extinct genus of pteranodontian pterosaur known from the Late Cretaceous of Texas, southern United States. It contains a single species, ''Alamodactylus byrdi''. Discovery ''Alamodactylus'' is known solely from its holoty ...
'' and ''
Volgadraco ''Volgadraco'' ("Volga River dragon") is a genus of pteranodontian pterosaur from the Upper Cretaceous of European Russia. Discovery and naming ''Volgadraco'' is known from lower beak (holotype SGU, no. 46/104a) and postcranial fragments from the ...
'', this contradicts its position within the Azhdarchoidea. The cladogram of their analysis is shown below: However, subsequent phylogenetic analyses placed ''Cretornis'' as a pteranodontian outside
Nyctosauridae Nyctosauridae (meaning "night lizards" or "bat lizards") is a family of specialized soaring pterosaurs of the late Cretaceous Period of North America, Africa, and possibly other continents including South America. It was named in 1889 by Henry ...
.


See also

*
List of pterosaur genera This list of pterosaurs is a comprehensive listing of all Genus, genera that have ever been included in the order Pterosauria, excluding purely vernacular terms. The list includes all commonly accepted genera, but also genera that are now considere ...
*
Timeline of pterosaur research This timeline of pterosaur research is a chronologically ordered list of important fossil discoveries, controversies of interpretation, and Biological taxonomy, taxonomic revisions of pterosaurs, the famed flying reptiles of the Mesozoic Era (ge ...


References

{{Portal bar, Paleontology, Czech Republic Late Cretaceous pterosaurs of Europe Pteranodontia Fossil taxa described in 1881