Cretornis Hlavaci
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''Cretornis'' is a
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 ...
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 ...
from the late
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
period (
Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...
stage) of what is now the Jizera Formation in the
Czech Republic The Czech Republic, or simply Czechia, is a landlocked country in Central Europe. Historically known as Bohemia, it is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the southeast. The ...
, dating to about 92 million years ago. It only contains a single species, ''Cretornis hlavaci''.


Discovery and naming

The fossils were discovered in 1880 by workers at a quarry in Zářecká Lhota near the town of Choceň, 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 Antonín Jan Frič (in German: Anton Johann Fritsch, 30 June 1832 – 15 November 1913) was a Czech paleontologist, biologist and geologist, living during the Austria-Hungary era. Professor at the Charles University and later became director of th ...
in Prague. In 1881,
Antonín Frič Antonín Jan Frič (in German: Anton Johann Fritsch, 30 June 1832 – 15 November 1913) was a Czech paleontologist, biologist and geologist, living during the Austria-Hungary era. Professor at the Charles University and later became director of th ...
identified it as a pterosaur and named it as 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 ...
''Cretornis Hlaváči''. 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 ...
''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 honors Hlaváč. Today, the holotype 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 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 an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
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 ( ; cs, Čechy ; ; hsb, Čěska; szl, Czechy) is the westernmost and largest historical region of the Czech Republic. Bohemia can also refer to a wider area consisting of the historical Lands of the Bohemian Crown ruled by the Bohem ...
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''; plural 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 nam ...
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 s ...
''. In 2010 however, Alexandr Averianov concluded that it should be possible to determine unique traits. In 2015 he published a new description, concluding that ''Cretornis'' was a valid taxon. ''Cretornis'' is known from 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 ...
, presently preserved in the collection of the ''
Národní muzeum The National Museum (NM) ( Czech: ''Národní muzeum'') is a Czech museum institution intended to systematically establish, prepare, and publicly exhibit natural scientific and historical collections. It was founded in 1818 by Kašpar Maria Šte ...
'' (National Museum) in
Prague Prague ( ; cs, Praha ; german: Prag, ; la, Praga) is the capital and largest city in the Czech Republic, and the historical capital of Bohemia. On the Vltava river, Prague is home to about 1.3 million people. The city has a temperate ...
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 (; ) 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 and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
(upper arm bone), an
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 ...
, 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 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 taxa, not even those most closely related to t ...
, 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 UK and possibly Morocco. Several species have ...
'' in the nineteenth century. In 1997 Coralia-Maria Jianu suggested it belonged to the
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 ( ...
.Jianu C-M., Weishampel D.B., Stiuca E., 1997, "Old and new pterosaur material from the Hateg basin (Late Cretaceous) of western Romania, and comments about pterosaur diversity in the Late Cretaceous of Europe", ''Abstracts of the Second European Workshop on Vertebrate Palaeontology'' 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 Cret ...
. More detailed comparisons of the wing bones led Averianov to conclude in 2015 that they belonged to a non-azhdarchid
azhdarchoid 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 and Late Cretaceous periods, with ...
, probably a member of the group
Neoazhdarchia 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 and Late Cretaceous periods, with o ...
with which it has many features in common. It is probably more advanced than the
Thalassodromidae Thalassodrominae or Thalassodromidae (meaning "sea runners", due to previous misconceptions of skimming behavior; they are now thought to be terrestrial predators) is a group of azhdarchoid pterosaurs from the Cretaceous period. Its traditional ...
, 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 holo ...
'' 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 Europe. It was named in 1889 by Henry Alleyne Nicholson and Richard Lydek ...
, in a more derived position than '' Alamodactylus'' and ''
Volgadraco ''Volgadraco'' ("Volga River dragon") is a genus of pterodactyloid pterosaur from the Upper Cretaceous of European Russia. ''Volgadraco'' was originally classified as an azhdarchid. However, recent studies have concluded that it may belong to e ...
'', this contradicts its position within the Azhdarchoidea. The cladogram of their analysis is shown below:


See also

*
List of pterosaur genera This list of pterosaurs is a comprehensive listing of all 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 considered inval ...
*
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, Cretaceous, Czech Republic Late Cretaceous pterosaurs of Europe Ornithocheiroids Fossil taxa described in 1881