Zhenyuanopterus
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''Zhenyuanopterus'' is a
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
boreopterid Boreopteridae (meaning "northern wings") is a group of pterodactyloid pterosaurs from the Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. Classification In 2006, Lü Junchang and colleagues named the clade Boreopteridae for the ...
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 ...
which is known from
Lower Cretaceous Lower may refer to: *Lower (surname) *Lower Township, New Jersey *Lower Receiver (firearms) *Lower Wick Lower Wick is a small hamlet located in the county of Gloucestershire, England. It is situated about five miles south west of Dursley, eight ...
(early
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...
stage)
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its ex ...
of
Liaoning Liaoning () is a coastal province in Northeast China that is the smallest, southernmost, and most populous province in the region. With its capital at Shenyang, it is located on the northern shore of the Yellow Sea, and is the northernmost ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. It contains one species, ''Zhenyuanopterus longirostris'', which was first described and named by
Lü Junchang Lü Junchang (; 1965 – 9 October 2018) was a Chinese palaeontologist and professor at the Institute of Geology, Chinese Academy of Geological Sciences. An expert on Mesozoic reptiles, he described and named dozens of dinosaur and pterosaur taxa ...
.


Discovery and naming

''Zhenyuanopterus longirostris'' was named by paleontologist Lü Junchang in 2005. The genus is named after Sun Zhenyuan, who gave Lü Junchang the fossils along with Zi Fan. The specific name ("long-snouted") refers to the long snout of the pterosaur. One specimen is known; a complete skeleton, it was found in the Huangbanjigou locality of the
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its ex ...
, and it is catalogued as GLGMV 0001 in the Guilin Longshan Geological Museum in
Guangxi Guangxi (; ; Chinese postal romanization, alternately romanized as Kwanghsi; ; za, Gvangjsih, italics=yes), officially the Guangxi Zhuang Autonomous Region (GZAR), is an Autonomous regions of China, autonomous region of the People's Republic ...
, China. A second, larger partial specimen, consisting of the rear half of the body, was found at the same locality and catalogued as XHPM1088 at the Dalian Xinghai Paleontological Museum. In 2014, Teng Fangfang and colleagues described it.


Description

The skull of the type specimen of ''Zhenyuanopterus'' measures long. It possesses a rectangular crest extending along the top of the snout, and another small crest at the back of the head. The eye socket is in the shape of an inverted triangle. Typical of the
Boreopteridae Boreopteridae (meaning "northern wings") is a group of pterodactyloid pterosaurs from the Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. Classification In 2006, Lü Junchang and colleagues named the clade Boreopteridae for the ...
, ''Zhenyuanopterus'' had a long snout filled with needle-like teeth, which are longest at the front of the mouth; a distinguishing characteristic is that the longest teeth are more than 10 times the length of the shortest. The teeth are smooth and triangular, with a slight curve. There are 86 teeth in the upper and another 86 teeth in the lower jaw. Additional distinguishing characteristics include the third phalanx of the wing digit, the humerus, and femur having the same length; and the feet being very small. The second specimen indicated that the humerus and femur grew at the same rate. The forelimb was more robust than the hindlimb. It was thought that the
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
was longer than the
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
or shoulder blade, unlike most advanced pterosaurs, but the type specimen had a poorly-preserved coracoid and the second specimen showed this to not be the case. ''Zhenyuanopterus'' had relatively short
cervical vertebrae In tetrapods, cervical vertebrae (singular: vertebra) are the vertebrae of the neck, immediately below the skull. Truncal vertebrae (divided into thoracic and lumbar vertebrae in mammals) lie caudal (toward the tail) of cervical vertebrae. In ...
compared to ''
Feilongus ''Feilongus'' is an extinct genus of ctenochasmatid pterodactyloid pterosaur from the Barremian–Aptian-age Lower Cretaceous Yixian Formation of Beipiao, Liaoning, China. Discovery and naming The genus was named and described in 2005 by Wang X ...
''. Several of the
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
are fused into a
notarium Notarium or os dorsale is a bone consisting of the fused vertebra of the shoulder in birds and some pterosaurs. The structure helps brace the chest against the forces generated by the wings. In birds, the vertebrae are only in contact with adjac ...
, an indicator of maturity. ''Zhenyuanopterus'' was estimated to have a wingspan of , making it one of the larger boreopterids.


Synonymy with ''Boreopterus''

''Boreopterus'' and ''Zhenyuanopterus'' are both part of the Boreopteridae, and occur in the same depositional unit. Lü distinguished the two based on the greater number of teeth, the shorter
mandibular symphysis In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral halves ...
, and equal-length third wing finger phalanx, humerus, and femur in ''Zhenyuanopterus''. However, it has been suggested that this animal is actually the adult of ''Boreopterus'', which is known to be a juvenile; all known specimens of the crestless ''Boreopterus'' are smaller than the large, crested ''Zhenyuanopterus'', and most of the differences between them can be explained as age-related. ''Boreopterus'' may have been too young to develop a crest when it died.


Classification

Lü classified ''Zhenyuanopterus'' in the Boreopteridae based on the long and low skull, long and sharp teeth, long front teeth, and the femur and tibia being of equal length. In 2012, he further subdivided Boreopteridae by allying ''Zhenyuanopterus'' and ''Boreopterus'' in the Boreopterinae, based on the tooth row extending further back on the jaw, the shorter cervical vertebrae, and the weaker feet of these two genera. This concept was later followed by Brian Andres and colleagues, while also placing ''
Guidraco ''Guidraco'' (Chin. ''gui'' (鬼) "malicious ghost" + Lat. ''draco'' "dragon") is an extinct genus of toothed pterodactyloid pterosaur known from the Early Cretaceous of Liaoning Province, northeast China. According to many recent studies, ''Gu ...
'' as a basal member of the family. However, Jiang Shun-Xing later restricted Boreopteridae to ''Zhenyuanopterus'' and ''Boreopterus'' after finding that Lü's definition was not
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
. In 2018, a later study by Nicholas Longrich and colleagues found the Boreopteridae (consisting only of ''Boreopterus'' and ''Zhenyuanopterus'') as the sister taxon of the family
Lonchodectidae Lonchodectidae or LonchodraconidaePêgas, R.V., Holgado, B., Leal, M.E.C., 2019. "''Targaryendraco wiedenrothi'' gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pter ...
, while also placed as members of the more inclusive group
Ornithocheiromorpha Ornithocheiromorpha (from Ancient Greek, meaning "bird hand form") is a group of pterosaurs within the suborder Pterodactyloidea. Fossil remains of this group date back from the Early to Late Cretaceous periods (Valanginian to Turonian stages ...
. Topology 1: Andres ''et al.'' (2014). Topology 2: Longrich ''et al.'' (2018).


Paleoecology

In the Yixian Formation, ''Zhenyuanopterus'' lived with other pterosaurs such as ''Boreopterus'', ''Feilongus'', ''
Haopterus ''Haopterus'' is a genus of pterodactyloid pterosaur from the Barremian-Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. Its fossil remains dated back 124.6 million years ago. Discovery and naming It was in 2001 named by Wang Xi ...
'', ''
Yixianopterus ''Yixianopterus'' is a pterodactyloid pterosaur genus from the Barremian-Aptian-age Lower Cretaceous Yixian Formation of Liaoning, China. It is known from a single specimen, holotype JZMP-V-12, housed at the Benxi Geological Museum. This specimen ...
'', and ''
Gegepterus ''Gegepterus'' is a genus of ctenochasmatid pterosaur from the Early Cretaceous period of what is now the Yixian Formation of Liaoning, China. Only one species is known, ''G. changi''. History and etymology The genus was named in 2007 by Wang X ...
''. Teng and colleagues suggested in 2014 that ''Zhenyuanopterus'' spent less time on land and more time in the air or on cliffs, owing to the robust forelimbs and weak hindlimbs.


Diet

The Yixian Formation in the early Cretaceous was a lakeside forest ecosystem with the presence of volcanoes. These lakes and waterways would have provided many aquatic prey, like fish and amphibians, that ''Zhenyuanopterus'' would have likely hunted. It has been suggested that ''Zhenyuanopterus'' foraged while swimming, trapping prey within its needle-like teeth, a method similar to that of modern '' Platanista'' river dolphins, which display a similar dentition.


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, Cretaceous, China Early Cretaceous pterosaurs of Asia Pteranodontoids Yixian fauna Fossil taxa described in 2010 Taxa named by Lü Junchang