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''Hamipterus'' is an extinct
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 ...
of pteranodontoid
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 ...
s from 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 ...
Shengjinkou Formation The Shengjinkou Formation is an Early Cretaceous-aged Konservat-Lagerstätte composed of "interbedded red green and yellow variegated mudstones and siltstones" that is part of the larger Tugulu Group of China. Dinosaur and pterosaur remains have ...
of northwestern China. It is known from a single species, 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 ...
, ''H. tianshanensis''.


Discovery and naming

In 2006 from the
Hami Hami ( zh, c=哈密) or Kumul () is a prefecture-level city in eastern Xinjiang, China. It is well known for sweet Hami melons. In early 2016, the former Hami county-level city merged with Hami Prefecture to form the Hami prefecture-level city ...
region in
Xinjiang Xinjiang,; , SASM/GNC romanization, SASM/GNC: Chinese postal romanization, previously romanized as Sinkiang, officially the Xinjiang Uygur Autonomous Region (XUAR), is an Autonomous regions of China, autonomous region of the China, People' ...
, the
Shengjinkou Formation The Shengjinkou Formation is an Early Cretaceous-aged Konservat-Lagerstätte composed of "interbedded red green and yellow variegated mudstones and siltstones" that is part of the larger Tugulu Group of China. Dinosaur and pterosaur remains have ...
, a Konservat-Lagerstätte was reported, in this case lake sediments allowing for an exceptional preservation of fossils. The same year, Qiu Zhanxiang and Wang Banyue started official excavations. Part of the finds consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. The site represented a nesting colony that storm floods had covered with mud. Dozens of individuals could be secured from a total that in 2014 was estimated to run into the many hundreds. * In 2014, 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 ...
''Hamipterus tianshanensis'' was named and described by Wang Xiaolin,
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 ...
, Jiang Shunxing, Wang Qiang, Ma Yingxia, Yahefujiang Paidoula, Cheng Xin, Taissa Rodrigues, Meng Xi, Zhang Jialiang, Li Ning, and Zhou Zhonghe. The generic name combines that of the Hami region with a Latinised Greek πτερόν, ''pteron'', "wing". 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 ...
refers to the provenance from the
Tian Shan The Tian Shan, also known as the Tengri Tagh or Tengir-Too, meaning the "Mountains of God/Heaven", is a large system of mountain ranges in Central Asia. The highest peak is Jengish Chokusu at high and located in Kyrgyzstan. Its lowest point is ...
, a mountain range. 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 ...
, IVPP V18931.1, has been found in a layer of the
Tugulu Group The Tugulu Group () is a geological Group in Xinjiang, China whose strata date back to the Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the ...
dating from the Lower Cretaceous. It consists of a skull, probably of a female. The
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...
is IVPP V18935.1, the skull of a male individual. The inventory number IVPP V18931 does not pertain to a single skeleton, but to a block containing various bones of different individuals. Eleven such blocks had in 2014 been secured, numbered IVPP V18931 to V18941. Together, they comprise the remains of at least 40 animals, both bones and soft tissue remnants such as the horn sheaths of skull crests. Exceptionally for pterosaur fossils, the bones have not been crushed, but were preserved three-dimensionally in good condition. Five uncrushed eggs were also found. The finds in 2014 represented the largest known concentration of pterosaur fossils, with the exception of the ''
Pterodaustro ''Pterodaustro'' (from Greek language, Greek , and Latin , ) is a genus of ctenochasmatid Pterodactyloidea, pterodactyloid pterosaur from South America. Its fossil remains dated back to the Early Cretaceous period, about 105 million years ago. ...
'' nesting colonies of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
.


Description

The wingspan of the individuals described in 2014 ranged from . The describing authors indicated some distinguishing traits, all of them
autapomorphies 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 ...
, unique derived characters. The dentary, the front bone of the lower jaw, has a hook-shaped process. The ascending branch of the jugal bone, running to the lacrimal bone, is thin, inclined to the front, and expanded at the top. The central supraoccipital of the top rear skull bears a well-developed crest. The humerus is perforated by a pneumatic foramen near the base of the deltopectoral crest. The outer lower carpal bone of the wrist has a spike-shaped process pointing to below. Also present is a unique combination of traits that in themselves are not unique. The front snout bone, the premaxilla, bears a crest with ridges and grooves that curve to the front. The groove on the dentary reaches the highest point of that bone. Both the snout tip and the tip of the lower jaws are slightly expanded. The deltopectoral crest is moderately twisted around the longitudinal axis of the humerus.


Phylogeny

''Hamipterus'' was within the
Pterodactyloidea Pterodactyloidea ( ; derived from the Greek words ''πτερόν'' (''pterón'', for usual ''ptéryx'') "wing", and ''δάκτυλος'' (''dáktylos'') "finger") is one of the two traditional suborders of pterosaurs ("wing lizards"), and contai ...
, placed into the group
Pteranodontoidea Pteranodontoidea (or pteranodontoids, from Greek meaning "toothless wings") is an extinct clade of ornithocheiroid pterosaurs from the Early to Late Cretaceous (early Valanginian to late Maastrichtian stages) of Asia, Africa, Europe, North Amer ...
. An exact
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
analysis could not resolve the relationship with ''
Istiodactylus ''Istiodactylus'' is a genus of pterosaur that lived during the Early Cretaceous period, about 120 million years ago. The first fossil was discovered on the English Isle of Wight in 1887, and in 1901 became the holotype specimen of a new species ...
'', ''
Ludodactylus ''Ludodactylus'' (meaning "play finger") is a genus of anhanguerid pterodactyloid pterosaur from the Early Cretaceous period (Aptian stage) of what is now the Crato Formation of the Araripe Basin in Ceará, Brazil. The type species, type and only ...
'' and the
Anhangueridae Anhangueridae (alternatively called Ornithocheiridae, meaning "bird hands") is a group of pterosaurs within the suborder Pterodactyloidea. These pterosaurs were among the last to possess teeth. Members that belong to this group lived from the ...
. The cladogram below is a topology recovered by Pêgas et al. (2019). In the analyses, they recovered this genus within the family Hamipteridae.Rodrigo V. Pêgas, Borja Holgado & Maria Eduarda C. Leal (2019) On ''Targaryendraco wiedenrothi'' gen. nov. (Pterodactyloidea, Pteranodontoidea, Lanceodontia) and recognition of a new cosmopolitan lineage of Cretaceous toothed pterodactyloids, Historical Biology,


Paleobiology

The large number of individuals found allowed the establishment of a growth series, showing how individuals developed through their
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the ovum, egg to adult. The term can also be used to refer to t ...
. Larger animals feature a number of changes. Their snout tips become relatively wider. The snout crest becomes more robust and expands its base towards the front, beginning at the level of the fifth tooth instead of the sixth. The pattern of grooves and ridges on the crest grows more prominent. The snout tip also starts to straighten in side view, no longer curving upwards. The groove in the dentary deepens and lengthens, as well. No change, however, takes place in the number of teeth, the degree of fusion in the symphysis of the lower jaws, or the shape of the postcranial skeleton, as far as can be ascertained, given the fact that the elements behind the skull were not found articulated. It was assumed that a clear
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
was discovered, with the largest specimens sporting the largest crests being the males, while smaller individuals were females with smaller crests. This was seen as a refutation of the hypothesis that with pterosaurs, only the males possessed crests.
Tomography Tomography is imaging by sections or sectioning that uses any kind of penetrating wave. The method is used in radiology, archaeology, biology, atmospheric science, geophysics, oceanography, plasma physics, materials science, cosmochemistry, ast ...
scans of fossilized ''Hamipterus'' eggs suggests that young ''Hamipterus'' had well-developed thigh bones for walking, but weak chests for flight. With the close proximity of the nests and adults, as well as how underdeveloped the hatchlings were for flying, it has also been suggested that ''Hamipterus'' practiced some form of
parental care Parental care is a behavioural and evolutionary strategy adopted by some animals, involving a parental investment being made to the evolutionary fitness of offspring. Patterns of parental care are widespread and highly diverse across the animal k ...
. However, this study has since been criticized by some paleontologists. A 2021 further disagrees by demonstrating that the young of ''Hamipterus'' don't differ significantly from the other flight capable flaplings.


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, China Pteranodontoidea Early Cretaceous pterosaurs of Asia Cretaceous China Fossils of China Fossil taxa described in 2014 Taxa named by Alexander Kellner Pterosaur genera