Jeholopterus
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''Jeholopterus'' was a small anurognathid
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 ...
from the Middle to
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 ...
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 stages). It is known for its exceptionally preserved fossils, including those of ...
of
Inner Mongolia Inner Mongolia, officially the Inner Mongolia Autonomous Region, is an autonomous region of the People's Republic of China. Its border includes most of the length of China's border with the country of Mongolia. Inner Mongolia also accounts for a ...
,
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
, preserved with hair-like pycnofibres and skin remains.


Naming

The genus was named in 2002 by Wang Xiaolin, Zhou Zhonghe and Xu Xing. 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 specim ...
, the only known, is ''Jeholopterus ninchengensis''. The genus name is derived from its place of discovery, Jehol in China, and a Latinized Greek ''pteron'', "wing". The specific name refers to Nincheng County.


Description

The type species is based on
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 seve ...
IVPP The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name suggest ...
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. It is the world's List of countries and dependencies by population, most populous country, with a Population of China, population exceeding 1.4 billion, slig ...
.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. 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 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 (plural: patagia) is a membranous body part that assists an animal in obtaining lift when gliding or flight. The structure is found in extant and extinct groups of flying and gliding animals including bats, birds, some droma ...
.


Phylogeny

''Jeholopterus'' was by the authors assigned to the Anurognathidae. 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 (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
conducted by Xuefang Wei and colleagues recovered ''Jeholopterus'' within the subfamily Anurognathinae, 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'' and '' Vesperopterylus'' 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 species known of the group, might also have been a piscivore, a fish-eater.


See also

* List of pterosaur genera *
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

{{Taxonbar, from=Q2924821 Monofenestratans 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