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''Lingwulong'' is a genus of
dicraeosaurid Dicraeosauridae is a family of diplodocoid sauropods who are the sister group to Diplodocidae. Dicraeosaurids are a part of the Flagellicaudata, along with Diplodocidae. Dicraeosauridae includes genera such as ''Amargasaurus'', ''Suuwassea'', ' ...
sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
from the
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
of what is now
Lingwu Lingwu (, Xiao'erjing: لِئٍ‌وُ شِ) is a county-level city of Ningxia Hui Autonomous Region, Southwest China, it is under the administration of the prefecture-level city of Yinchuan. It is the most important industrial city of Ningxia. Li ...
,
Yinchuan Yinchuan (, ; ) is the capital of the Ningxia Hui Autonomous Region, China, and was the capital of the Tangut-led Western Xia dynasty. It has an area of and a total population of 2,859,074 according to the 2020 Chinese census, and its built- ...
,
Ningxia Ningxia (,; , ; alternately romanized as Ninghsia), officially the Ningxia Hui Autonomous Region (NHAR), is an autonomous region in the northwest of the People's Republic of China. Formerly a province, Ningxia was incorporated into Gansu in ...
, China. The type and only species is ''L. shenqi'', known from several partial skeletons. It is the earliest-aged
neosauropod Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentina, Argentine paleontologist José Bonaparte and currently described as ''Saltasaurus loricatus'', ''Diplodocus longus'', and all animals directly descended from their most recen ...
ever discovered, as well as the only definite
diplodocoid Diplodocoidea is a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like ''Supersaurus'', ''Diplodocus'', ''Apatosaurus'', and ''Amphicoelias''. Most had very long necks and long, ...
from east Asia.


Discovery

In 2004,
sheep herder A shepherd or sheepherder is a person who tends, herds, feeds, or guards flocks of sheep. ''Shepherd'' derives from Old English ''sceaphierde (''sceap'' 'sheep' + ''hierde'' 'herder'). ''Shepherding is one of the world's oldest occupations, i ...
Ma Yun found dinosaur bones, notifying local administrators Yang Huozhu and Liu Hongan. In the spring of 2005 these showed the fossils to dinosaur expert Xu Xing who sent out a team to investigate the find. From 2005 onwards several quarries ware excavated at Ciyaopu, near
Lingwu Lingwu (, Xiao'erjing: لِئٍ‌وُ شِ) is a county-level city of Ningxia Hui Autonomous Region, Southwest China, it is under the administration of the prefecture-level city of Yinchuan. It is the most important industrial city of Ningxia. Li ...
, in
Ningxia Ningxia (,; , ; alternately romanized as Ninghsia), officially the Ningxia Hui Autonomous Region (NHAR), is an autonomous region in the northwest of the People's Republic of China. Formerly a province, Ningxia was incorporated into Gansu in ...
. The remains were discovered of about seven to ten sauropod skeletons. The excavations and the subsequent preparation were carried out by Wang Haijun, Xiang Lishi, He Sicai, Cao Renfang, Tang Zhilu and Tao Yu. In 2018, Xu Xing, Paul Upchurch, Philip D. Mannion, Paul M. Barrett, Omar R. Regalado-Fernandez, Mo Jinyou, Ma Jinfu and Liu Hongan named and described 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 ...
''Lingwulong shenqi''. The generic name combines a reference to Lingwu with the Mandarin ''long'', "dragon". The specific name ''shenqi'' (神奇) means "amazing" or "magical" in Mandarin, reflecting the unexpected appearance of a member of the
Dicraeosauridae Dicraeosauridae is a family of diplodocoid sauropods who are the sister group to Diplodocidae. Dicraeosaurids are a part of the Flagellicaudata, along with Diplodocidae. Dicraeosauridae includes genera such as '' Amargasaurus'', '' Suuwassea' ...
in East Asia, a group never before identified in the region. 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 ...
, LM V001a, consists of the rear of a skull associated with a series of dentary teeth found in their original position. It is part of the collection of the
Lingwu Museum Lingwu (, Xiao'erjing: لِئٍ‌وُ شِ) is a county-level city of Ningxia Hui Autonomous Region, Southwest China, it is under the administration of the prefecture-level city of Yinchuan. It is the most important industrial city of Ningxia. Li ...
. 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 nor a syntype). Of ...
, LGP V001b, is a partial skeleton lacking the skull. It contains the rear vertebrae of the back, the sacrum, the pelvis, the first tail vertebra and elements of the right hindlimb. Paratype and holotype possibly represent a single individual. The paratype is displayed in the
Lingwu Geopark Lingwu (, Xiao'erjing: لِئٍ‌وُ شِ) is a county-level city of Ningxia Hui Autonomous Region, Southwest China, it is under the administration of the prefecture-level city of Yinchuan. It is the most important industrial city of Ningxia. Li ...
. Several specimens have been referred to the species. IVPP V23704 is a series of twenty-nine dentary teeth. LGP V002 is a partial skeleton including vertebrae of the back and tail, the shoulder girdle, and elements of the forelimbs and pelvis. LGP V003 is a partial skeleton containing a series of vertebrae including dorsals, sacrals and the first two caudals, and both ilia. LGP V004 consists of a front neck vertebra, a front back vertebra and a right shinbone of a small individual. LGP V005 is a partial skeleton containing the sacrum, the pelvis and a series of twenty-five front and middle tail vertebrae. LGP V006 contains neck vertebrae, the shoulder girdle and forelimb elements. Additionally numerous disarticulated bones from the quarries were referred.


Description

''Lingwulong''s remains belong to 7 to 10 individuals at different ontogenetic stages, and even include skull bones. Overall, nearly the whole skeletal anatomy is known. Autapomorphies (unique traits) that distinguish ''Lingwulong'' from other diplodocoids include highly elaborated ornamentation along the upper margin of the orbital area, occipital condyle with transversely wide articular surface and anterior dorsal vertebrae with slightly twisted metapophyses presenting a sub-circular pseudofacet on their tip. Some traits, such as morphology of cervical vertebrae metapophyses look intermediate between the derived dicraeosaurids condition and the plesiomorphic condition spread among
flagellicaudata Flagellicaudata is a clade of Dinosauria. It belongs to Sauropoda and includes two families, the Dicraeosauridae and the Diplodocidae. Phylogeny The clade Flagellicaudata was erected by Harris and Dodson (2004) for the diplodocoid clade formed ...
ns. Unlike most other diplodocoids, which have square-shaped snouts in dorsal view, ''Lingwulong'' had a U-shaped snout.


Classification

Simplified cladogram of Flagellicaudata after Xu and colleagues (2018) is shown below:


Significance


Geologic age

''Lingwulong'' has been considered the earliest known
neosauropod Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentina, Argentine paleontologist José Bonaparte and currently described as ''Saltasaurus loricatus'', ''Diplodocus longus'', and all animals directly descended from their most recen ...
and the oldest known diplodocoid. Prior to its discovery, the oldest known possible diplodocoids included "''Cetiosaurus''" ''glymptonensis'' from the late Bathonian and '' Cetiosauriscus stewarti'' from the
Callovian In the geologic timescale, the Callovian is an age and stage in the Middle Jurassic, lasting between 166.1 ± 4.0 Ma (million years ago) and 163.5 ± 4.0 Ma. It is the last stage of the Middle Jurassic, following the Bathonian and preceding the ...
, both from the United Kingdom. At the time of ''Lingwulongs naming, it was regarded as having been found in the Yanan Formation. At the time, the age of the Yanan Formation was poorly understood, but was considered highly unlikely to be older than the late
Toarcian The Toarcian is, in the ICS' geologic timescale, an age and stage in the Early or Lower Jurassic. It spans the time between 182.7 Ma (million years ago) and 174.1 Ma. It follows the Pliensbachian and is followed by the Aalenian. The Toarcian ...
or younger than the
Bajocian In the geologic timescale, the Bajocian is an age and stage in the Middle Jurassic. It lasted from approximately 170.3 Ma to around 168.3 Ma (million years ago). The Bajocian Age succeeds the Aalenian Age and precedes the Bathonian Age. Stratig ...
, yielding a midpoint age estimate of 174.15 Ma for ''Lingwulong''. Subsequent research on the Yanan Formation dated it more precisely as
Aalenian The Aalenian () is a subdivision of the Middle Jurassic Epoch/Series of the geologic timescale that extends from about 174.1 Ma to about 170.3 Ma (million years ago). It was preceded by the Toarcian and succeeded by the Bajocian. Stratigraphic ...
in age, largely between 174.0 and 171.7 Ma. However, the Lingwu dinosaur fauna has been argued to have actually come from the Zhiluo Formation, which would make ''Lingwulong'' substantially younger,
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Strat ...
to early Oxfordian in age.


Paleobiogeography

Diplodocoids were long thought to have been absent from East Asia due to the presence of the
Turgai Sea The Turgai Sea, also known as the Turgay Sea, Turgai Strait, Obik Sea, Ural Sea or West Siberian Sea, was a large shallow body of salt water (an epicontinental or epeiric sea) during the Mesozoic through Cenozoic Eras. It extended north of the pre ...
, and the East Asian Isolation Hypothesis was used to explain the presence of
mamenchisaurid Mamenchisauridae is a family of sauropod dinosaurs belonging to Eusauropoda known from the Jurassic and Early Cretaceous of Asia and Africa. Some members of the group reached gigantic sizes, amongst the largest of all sauropods. Classification T ...
s in East Asia but not elsewhere in the world. The discovery of ''Lingwulong'' casts doubt on these paleobiogeographical assumptions by showing that
diplodocoid Diplodocoidea is a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like ''Supersaurus'', ''Diplodocus'', ''Apatosaurus'', and ''Amphicoelias''. Most had very long necks and long, ...
s were present in East Asia during the Jurassic.


See also

*
2018 in paleontology Flora Plants Fungi Cnidarians Research * New three dimensionally phosphatized microfossils of coronate scyphozoan '' Qinscyphus necopinus'', including a new type of fossil embryo, are described from the Cambrian (Fortunian) Kuanchuanpu F ...


References

{{Portal bar, Dinosaurs, Cretaceous, China Middle Jurassic dinosaurs of Asia Dicraeosaurids Fossil taxa described in 2018 Taxa named by Xu Xing