Jeholopterus
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''Jeholopterus'' was a small anurognathid
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 ...
known from the
Middle Middle or The Middle may refer to: * Centre (geometry), the point equally distant from the outer limits. Places * Middle (sheading), a subdivision of the Isle of Man * Middle Bay (disambiguation) * Middle Brook (disambiguation) * Middle Creek ...
to
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
Daohugou Beds of the
Tiaojishan Formation The Tiaojishan Formation is a geological formation in Hebei and Liaoning, People's Republic of China, dating to the middle-late Jurassic period (Bathonian-Oxfordian (stage), Oxfordian stages). It is known for its Lagerstätte, exceptionally preser ...
of
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an Autonomous regions of China, autonomous region of China. Its border includes two-thirds of the length of China's China–Mongolia border, border with the country of Mongolia. ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
, and possibly the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
Sinuiju Formation of
North Korea North Korea, officially the Democratic People's Republic of Korea (DPRK), is a country in East Asia. It constitutes the northern half of the Korea, Korean Peninsula and borders China and Russia to the north at the Yalu River, Yalu (Amnok) an ...
.


Naming

The genus was named in 2002 by Wang Xiaolin,
Zhou Zhonghe Zhonghe Zhou (; born 19 January 1965 in Jiangdu, Jiangsu province) is a Chinese palaeontologist. He described the ancient bird '' Confuciusornis''.. Zhonghe graduated from Nanjing University and earned a Ph.D. in Biology in 1999 from the Univer ...
and Xu Xing. 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 ...
, the only known, is ''Jeholopterus ningchengensis''. The genus name is derived from its place of discovery, Jehol in China, and a Latinized Greek ''pteron'', "wing". The specific name refers to
Ningcheng County Ningcheng County ( Mongolian: ; zh, s=宁城县) is a county of southeastern Inner Mongolia, China, bordering Liaoning province to the east. It is under the administration of Chifeng City. The daohugouthallus extinct genus of lichen was found ...
.


Description

The type species is based on
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 ...
IVPP V12705, a nearly complete specimen from the Daohugou beds of Ningcheng County in the Neimongol (Inner Mongolia) Autonomous Region of
China China, officially the People's Republic of China (PRC), is a country in East Asia. With population of China, a population exceeding 1.4 billion, it is the list of countries by population (United Nations), second-most populous country after ...
.Wang, X., Zhou, Z., Zhang, F., and Xu, X. (2002). "A nearly completely articulated rhamphorhynchoid pterosaur with exceptionally well-preserved wing membranes and 'hairs' from Inner Mongolia, northeast China." ''Chinese Science Bulletin'' 47(3), 226 – 232. The specimen is crushed into a slab and counterslab pair, so that parts of the specimen are preserved on one side of a split stone and some on the other. This includes exquisite preservation of carbonized skin fibers and
pycnofibres Pterosaurs are an extinct clade of flying reptiles in the Order (biology), 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). Pterosau ...
. The fibers are preserved around the body of the specimen in a "halo." Wing tissue is preserved, though its extent is debatable, including the exact points of attachment to the legs (or if it is attached to the legs at all). In 2009
Alexander Kellner Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
published a study reporting the presence of three layers of fibers in the wing, allowing the animal to precisely adapt the wing profile.Alexander W. A. Kellner, Xiaolin Wang, Helmut Tischlinger, Diogenes de Almeida Campos, David W. E. Hone, and Xi Meng. (2009). "The soft tissue of ''Jeholopterus'' (Pterosauria, Anurognathidae, Batrachognathinae) and the structure of the pterosaur wing membrane", ''Proc. R. Soc. B'' As an anurognathid, ''Jeholopterus'' shows the skull form typical for this group, being wider than it was long, at , with a very broad mouth. Most teeth are small and peg-like, but some are longer and recurved. The neck was short with seven or eight cervical vertebrae. Twelve or thirteen dorsal vertebrae are present and three sacrals. There are five pairs of belly ribs. The tail vertebrae have not been preserved. The describers argue that ''Jeholopterus'' had a short tail, a feature seen in other anurognathids but unusual for "
rhamphorhynchoid The Rhamphorhynchoidea forms one of the two suborders of pterosaurs and represents an evolutionary grade of primitive members of flying reptiles. This suborder is paraphyletic unlike the Pterodactyloidea, which arose from within the Rhamphorhync ...
" (i.e. basal) pterosaurs that typically have a long tail. Wang ''et al.'' cited the presence of a fringe of pycnofibres in the region of the tail to infer the presence of a short tail. However, a subsequent study by Dalla Vecchia argued that gleaning any information about the tail is impossible, given that the tail is "totally absent" in the fossil.Dalla Vecchia, F.M. (2002). "Observations on the non-pterodactyloid pterosaur ''Jeholopterus ningchengensis'' from the Early Cretaceous of Northeastern China." ''Natura Nascosta'', 24: 8 - 27. The wing bones are robust. The
metacarpals In human anatomy, the metacarpal bones or metacarpus, also known as the "palm bones", are the appendicular skeleton, appendicular bones that form the intermediate part of the hand between the phalanges (fingers) and the carpal bones (wrist, wris ...
are very short. A short pteroid, supporting a propatagium, is pointing towards the body. The hand claws are long and curved. The wings of ''Jeholopterus'' show evidence that they attached to the ankle, according to Wang ''et al.''. They are relatively elongated with a wingspan of ninety centimetres. The legs are short but robust. The toes bear well-developed curved claws, but these are not as long as the hand claws. The fifth toe is elongated, according to the authors supporting a membrane between the legs, the
uropatagium The patagium (: patagia) is a membranous body part that assists an animal in obtaining lift when gliding or flying. The structure is found in extant and extinct groups of flying and gliding animals including bats, theropod dinosaurs (including b ...
. Another articulated specimen was discovered no later than 2009 in the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
Sinuiju Formation of
North Korea North Korea, officially the Democratic People's Republic of Korea (DPRK), is a country in East Asia. It constitutes the northern half of the Korea, Korean Peninsula and borders China and Russia to the north at the Yalu River, Yalu (Amnok) an ...
and it represents the first
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 ...
record of ''Jeholopterus''; the specimen was assigned to ''Jeholopterus
cf. The abbreviation cf. (short for either Latin or , both meaning 'compare') is generally used in writing to refer the reader to other material to make a comparison with the topic being discussed. However some sources offer differing or even contr ...
ningchengensis'' by So ''et al.'' (2024).


Phylogeny

''Jeholopterus'' was by the authors assigned to the
Anurognathidae Anurognathidae is a family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Eight genera are definitively known: '' Anurognathus'', from the Late Jurassic of Germ ...
. An analysis by Lü Junchang in 2006 resolved its position as being the sister taxon of ''Batrachognathus''. Kellner and colleagues in 2009 proposed Batrachognathinae for the clade comprising these two genera and ''Dendrorhynchoides''. In 2021, a
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
conducted by Xuefang Wei and colleagues recovered ''Jeholopterus'' within the subfamily
Anurognathinae Anurognathidae is a family (biology), family of small, short-tailed pterosaurs that lived in Europe, Asia, and possibly North America during the Jurassic and Cretaceous periods. Eight genera are definitively known: ''Anurognathus'', from the Late ...
, which is unlike the former study by Kellner and colleagues where it was recovered as closely related to ''Batrachognathus''. ''Jeholopterus'' was found to have been the sister taxon to an unnamed clade formed by both ''
Anurognathus ''Anurognathus'' (from the Greek ''ανоυρα γναθος'' "frog jaw") is an extinct genus of small pterosaur from the Late Jurassic Altmühltal Formation of Germany. ''Anurognathus'' was first named and described by Ludwig Döderlein in 19 ...
'' and ''
Vesperopterylus ''Vesperopterylus'' (meaning "dusk wing") is a genus of anurognathidae, anurognathid pterosaur from the Early Cretaceous Jiufotang Formation of China, the geologically youngest member of its group. Notably, ''Vesperopterylus'' appears to have a r ...
'' in Wei and colleagues' study. Below is a cladogram representing their phylogenetic analysis:


Lifestyle

Anurognathids are normally considered insectivores. Wang ''et al'' hypothesized that ''Jeholopterus'', being the largest genus known of the group, might also have been a
piscivore A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
, a fish-eater.


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

{{Taxonbar, from=Q2924821 Anurognathidae Middle Jurassic pterosaurs of Asia Late Jurassic pterosaurs of Asia Taxa named by Zhou Zhonghe Fossil taxa described in 2002 Taxa named by Xu Xing Paleontology in Inner Mongolia Fossils of North Korea