Lufengosaurus Huenei
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''Lufengosaurus'' (, meaning "Lufeng lizard") is a
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
massospondylid Massospondylidae is a family (biology), family of early massopod dinosaurs that existed in Asia, Africa, North America, South America and AntarcticaHellert, Spencer M. "A New Basal Sauropodomorph from The Early Jurassic Hanson Formation of Antarc ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
which lived during the Early
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately 143.1 Mya. ...
period in what is now southwestern
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 ...
."Lufengosaurus." In: Dodson, Peter & Britt, Brooks & Carpenter, Kenneth & Forster, Catherine A. & Gillette, David D. & Norell, Mark A. & Olshevsky, George & Parrish, J. Michael & Weishampel, David B. ''The Age of Dinosaurs''. Publications International, LTD. p. 38. .


History of Discovery

The first remains of ''Lufengosaurus'' were found during the late 1930s by
geologist A geologist is a scientist who studies the structure, composition, and History of Earth, history of Earth. Geologists incorporate techniques from physics, chemistry, biology, mathematics, and geography to perform research in the Field research, ...
Bien Meinian in the
Lower Lufeng Formation The Lufeng Formation (formerly Lower Lufeng Series) is a Lower Jurassic sedimentary rock formation found in Yunnan, China. It has two units: the lower Dull Purplish Beds/Shawan Member are of Hettangian age, and Dark Red Beds/Zhangjia'ao Member ar ...
at Shawan, near Lufeng in the
Yunnan Province Yunnan; is an inland Provinces of China, province in Southwestern China. The province spans approximately and has a population of 47.2 million (as of 2020). The capital of the province is Kunming. The province borders the Chinese provinces ...
of China. From 1938 onwards, Meinian was joined by paleontologist C. C. Young (Chinese: 杨钟健, Yáng Zhōngjiàn). Young would go on to name the animal's
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 ...
, ''Lufengosaurus huenei,'' in 1940, with the generic name referring to Lufeng where it was found and the specific name honoring Young's old tutor, the German paleontologist
Friedrich von Huene Baron Friedrich Richard von Hoyningen-Huene (22 March 1875 – 4 April 1969) was a German nobleman paleontologist who described a large number of dinosaurs, more than anyone else in 20th-century Europe. He studied a range of Permo-Carbonife ...
.Young, C.-C. 1940.
Preliminary notes on the Lufeng vertebrate fossils
. ''Bulletin of the Geological Society of China'' 20(3-4): 235-239
Young would go on to publish a detailed description of the animal the following year and would assign specimen
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 sugges ...
V15 as 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 ...
of the species, this specimen consisting of a nearly complete
skeleton A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fra ...
with a well-preserved
skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...
. A second species of ''Lufengosaurus'', ''L. magnus'', was also named by Young in 1940 in the same paper as ''Lufengosaurus huenei'' and would be fully described by him in 1947. As its specific name suggests, with the word magnus meaning "the large one" in
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
, this species was considered separate from ''L. huenei'' due to its significantly larger size, with it being up to a third larger than ''L. huenei'' in length. However, most authors have regarded it as a
junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...
of ''L. huenei'', with its specimens being said to simply represent larger individuals of the type species. Another "
prosauropod Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the Sauropoda, sauropods and their ancestral relatives. Sauropods generally grew to very large siz ...
" Young named in 1940, '' Gyposaurus sinensis'', was noted by him to be distinct from ''Lufengosaurus'' on behalf of the limbs being at least 50% smaller, though similarities in overall form were noted. This genus has subsequently been referred to ''Lufengosaurus'' on two separate occasions: once by
Peter Galton Peter Malcolm Galton (born 14 March 1942 in London) is a British vertebrate paleontologist who has to date written or co-written about 190 papers in scientific journals or chapters in paleontology textbooks, especially on ornithischian and prosau ...
in 1976 and once in a 2017 SVP presentation by Wang and colleagues. The referral is however doubted by some. ''Lufengosaurus'' would be referred to another genus yet again in the 1980s, with
Michael Cooper Michael Jerome Cooper (born April 15, 1956), nicknamed "Coop", is an American basketball coach and former player. He played for the Los Angeles Lakers during his entire career in the National Basketball Association (NBA), winning five NBA champ ...
suggesting that ''Lufengosaurus'' and ''
Yunnanosaurus ''Yunnanosaurus'' ( ) is an extinct genus of sauropodomorph dinosaur that lived approximately 199 to 183 million years ago in what is now the Yunnan Province, in China, for which it was named. ''Yunnanosaurus'' was a large sized, moderately-buil ...
'' were species of the
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
n genus ''
Massospondylus ''Massospondylus'' ( ) is a genus of sauropodomorph dinosaur from the Early Jurassic of southern Africa. It was described by Sir Richard Owen in 1854 from remains discovered in South Africa, and is thus one of the first dinosaurs to have been ...
'' 1981. However, a reanalysis in 2005 by
Paul Barrett Paul Franklyn "Legs" Barrett (14 December 1940 – 20 January 2019) was a UK agent and manager of 1950s style rock and roll artists, an author and previously a singer, songwriter and film actor. Barrett was the discoverer, mentor and first man ...
and colleagues, performed on the skull of ''Lufengosaurus huenei'' established it firmly as a valid genus separate from both ''Massospondylus'' and ''Yunnanosaurus'' based on craniodental characteristics. This same 2005 study would provide an extensive and detailed redescription of the cranial osteology of the holotype skull of the animal. The year 1985 saw
Zhao Xijin Zhao Xijin (赵喜进; born c. 1935 died July 21, 2012) was a Chinese paleontologist notable for having named numerous dinosaurs. He was a professor at Beijing's Institute of Vertebrate Paleontology and Paleoanthropology. Biography Zhao Xijin was ...
name a new species of the animal in a list of Jurassic reptiles from China. Zhao named it ''Lufengosaurus'' "changduensis", with the naming of the animal being prompted by the recovery of a specimen from
Tibet Tibet (; ''Böd''; ), or Greater Tibet, is a region in the western part of East Asia, covering much of the Tibetan Plateau and spanning about . It is the homeland of the Tibetan people. Also resident on the plateau are other ethnic groups s ...
. The species has however remained undescribed and thus a ''
nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...
,'' with neither the reported Tibetan specimen nor any other material being formally assigned to it after it was named. In 2015, preserved
collagen Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residue (biochemistry), residues. Proteins perform a vast array of functions within organisms, including Enzyme catalysis, catalysing metab ...
was found in a ''Lufengosaurus'' fossil by an international team led by Yao-Chang Lee of
Taiwan Taiwan, officially the Republic of China (ROC), is a country in East Asia. The main geography of Taiwan, island of Taiwan, also known as ''Formosa'', lies between the East China Sea, East and South China Seas in the northwestern Pacific Ocea ...
's National Synchrotron Radiation Research Center. The protein, described in ''Nature Communications'' (2017 January 31), was over 100 million years older than any previously recorded fossil protein, having sizeable implications for our understanding of protein preservation.


Description

''Lufengosaurus'' is often described as a rather small early
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
, about long. However, when the ''L. magnus'' specimens are included, its size is more considerable:
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology. He is best known for his work and research on theropoda, theropod dinosaurs and his detailed illustrations, both l ...
estimated a length of and a weight of in 2010, while Benson ''et al''. (2014) estimated a mass of . For an early sauropodomorph, its neck is rather long and the forelimbs are relatively short. From these it was inferred that the species was bipedal, even before it became common to assume this for all basal sauropodomorphs. Yang published a full osteology of ''Lufengosaurus'' in 1941, but was severely hampered in his diagnosis by the war conditions, preventing a full access to literature and making an adequate comparison with related forms impossible. Of the skull a modern description exists. The skull of the holotype is long.


Skull

''Lufengosaurus'' snout was deep and broad, and it had distinctive bony bumps just behind its large
nostril A nostril (or naris , : nares ) is either of the two orifices of the nose. They enable the entry and exit of air and other gasses through the nasal cavities. In birds and mammals, they contain branched bones or cartilages called turbinates ...
s and on its
cheek The cheeks () constitute the area of the face below the eyes and between the nose and the left or right ear. ''Buccal'' means relating to the cheek. In humans, the region is innervated by the buccal nerve. The area between the inside of th ...
s. A bony ridge on the side of its
upper jaw In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxillar ...
might have helped anchor
soft tissue Soft tissue connective tissue, connects and surrounds or supports internal organs and bones, and includes muscle, tendons, ligaments, Adipose tissue, fat, fibrous tissue, Lymphatic vessel, lymph and blood vessels, fasciae, and synovial membranes. ...
. If so, then ''Lufengosaurus'' must have had larger cheeks than most other
sauropodomorph Sauropodomorpha ( ; from Greek, meaning "lizard-footed forms") is an extinct clade of long-necked, herbivorous, saurischian dinosaurs that includes the sauropods and their ancestral relatives. Sauropods generally grew to very large sizes, had lo ...
s. Its closely spaced, serrated teeth suited a diet of leaves. About thirty major specimens have been discovered, including those of juveniles. In 1958 a specimen of ''Lufengosaurus'' was the first complete dinosaur skeleton mounted in China; a commemorative postage stamp of 8 yuan was issued on 15 April 1958 to celebrate the event, the first time ever a dinosaur was depicted on a stamp. The skeleton is now on display in the
Paleozoological Museum of China The Paleozoological Museum of China (PMC; ) is a museum in Beijing, China. The same building also houses the Institute of Vertebrate Paleontology and Paleoanthropology of the Chinese Academy of Sciences. The museum contains exhibition halls with s ...
in
Beijing Beijing, Chinese postal romanization, previously romanized as Peking, is the capital city of China. With more than 22 million residents, it is the world's List of national capitals by population, most populous national capital city as well as ...
.


Classification

Young originally assigned ''Lufengosaurus'' to the Plateosauridae of the suborder Prosauropoda, noting that it stood close to ''Plateosaurus fraasianus''. It is still sometimes considered a member of Plateosauridae, though some cladistic analyses have found it to be a member of Massospondylidae instead. This matches with our current knowledge of ''Lufengosaurus''' anatomy, which has proven to be more similar to taxa like ''Coloradisaurus'' and ''Massospondylus'' than was previously thought, with ''Lufengosaurus'' formerly being considered to have more anatomical similarities with ''Plateosaurus''. ''Lufengosaurus'' has historically had a maximum of three named species, only one of these being almost unilaterally considered valid nowadays: ''L. huenei''. The other two species, ''L. magnus'' and ''L.'' "changduensis", are respectively considered a junior synonym and a ''nomen nudum''. Specimens assigned to ''L. magnus'' in the past are now referred to the type species instead. Barret, Upchurch and Wang recovered ''Lufengosaurus'' as being the sister taxon to ''Gyposaurus sinensis'' in their 2005 cladistic analysis. A differing result was recovered from a cladistic analysis done by Fernando E. Novas and colleagues in 2011, which recovered the animal as the sister taxon to ''Glacialisaurus hammeri''. This result was replicated by Oliver W. M. Rauhut and colleagues' 2020 analysis. A simplified version of the resulting cladogram, not showing the part including the Sauropodiform taxa, is shown below.


Palaeobiology

Like all early sauropodomorphs, ''Lufengosaurus'' had much longer hindlimbs than forelimbs and was probably bipedal. It was herbivorous, although it had sharp claws (with an especially large thumb claw) and teeth. These features have been used to support claims, the most recent by Cooper in 1981, that ''Lufengosaurus'' may have been at least partially omnivorous, but the sharp teeth witnessed in ''Lufengosaurus'' and other early sauropodomorphs are similar to those seen in
iguana ''Iguana'' (, ) is a genus of herbivorous lizards that are native to tropical areas of Mexico, Central America, South America, and the Caribbean. The genus was first described by Austrian naturalist Josephus Nicolaus Laurenti, J.N. Laurenti in ...
ian
lizard Lizard is the common name used for all Squamata, squamate reptiles other than snakes (and to a lesser extent amphisbaenians), encompassing over 7,000 species, ranging across all continents except Antarctica, as well as most Island#Oceanic isla ...
s – which are herbivorous. Alternatively, the claws may have been used for defense or raking foliage from trees. Embryos of this genus also represent the earliest evidence of vertebrate soft tissue preservation. ''Lufengosaurus'' likely engaged in parental feeding as a fully
altricial Precocial species in birds and mammals are those in which the young are relatively mature and mobile from the moment of birth or hatching. They are normally nidifugous, meaning that they leave the nest shortly after birth or hatching. Altricial ...
animal, given that the bone development in its femora is closer to that of altricial pigeons than precocious chickens.


Palaeoecology

While originally considered to be a Triassic locality, nowadays the Lufeng Formation is dated to the Lower Jurassic instead (Hettangian-Sinemurian), implying that ''Lufengosaurus'' existed in a more recent timeframe than previously thought.


References


Sources

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External links


Lufengosaurus on stamp


{{Taxonbar, from=Q131561 *
Paleontologiara Discover Nearly Complete Dinosaur Skeleton
Massospondylidae Dinosaur genera Sinemurian dinosaurs Lufeng Formation Fossil taxa described in 1941 Taxa named by Yang Zhongjian Dinosaurs of China