''Panguraptor'' ("

Pangu Pangu or Pan Gu (also sometimes spelled Peng Gu and P’an-ku) ( zh, t=盤古, ) is a primordial being and creation figure in Chinese mythology and in Taoism. According to legend, Pangu separated heaven and earth, and his body later became ge ...

Chinese godplunderer") 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 ...

coelophysid Coelophysoidea is an extinct clade of theropod dinosaurs common during the Late Triassic and Early Jurassic periods. They were widespread geographically, probably living on all continents. Coelophysoids were all slender, carnivore, carnivorous fo ...

theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...

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 ...

known from fossils discovered in

Lower Jurassic The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic–Jurassic extinction event, 201.3 Ma&nb ...

rocks of southern

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 ...

. The

type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * ...

and only known species is ''Panguraptor lufengensis''. The generic name refers to the deity Pangu but also to the

supercontinent In geology, a supercontinent is the assembly of most or all of Earth's continent, continental blocks or cratons to form a single large landmass. However, some geologists use a different definition, "a grouping of formerly dispersed continents", ...

Pangaea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous period approximately 335 mi ...

for which in a geological context the same characters are used: 盘古. ''Raptor'' means "seizer", "robber" 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 ...

. 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 ...

is a reference to the Lufeng Formation. It was described in

2014 The year 2014 was marked by the surge of the Western African Ebola epidemic, West African Ebola epidemic, which began in 2013, becoming the List of Ebola outbreaks, most widespread outbreak of the Ebola, Ebola virus in human history, resul ...

by You Hai-Lu and colleagues.

History and naming

The specimen that would be named ''Panguraptor'' was discovered by a survey team of the Bureau of Land and Resources of

Lufeng County Lufeng (; Yi script, Chuxiong Yi script: , International Phonetic Alphabet, IPA：) is a county-level city located in Yunnan province, China, administered as a part of Chuxiong Yi Autonomous Prefecture. Geography Lufeng is located in the east of Ch ...

, China, on the hillside behind Xiaolishu Village, Lufeng Dinosaur Mountain Town. The team found the partially articulated skeleton exposed on the slope on . Due to weathering, there was not time to slowly clean the skeleton, so it was glued and

plaster Plaster is a building material used for the protective or decorative coating of walls and ceilings and for moulding and casting decorative elements. In English, "plaster" usually means a material used for the interiors of buildings, while "re ...

ed on location over the course of two days before being shipped to Lufeng World Dinosaur Valley. After four years it was inventoried and discovered to be a complete

skeleton, at which point it was cleaned and prepared over the course of a year. In 2014 the fossil, catalogued as Bureau of Land and Resources of Lufeng County LFGT-0103, was described by Chinese paleontologist Hailu You and colleagues as the

type specimen In biology, a type is a particular wikt:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally associated. In other words, a type is an example that serves to ancho ...

of the new theropod ''Panguraptor lufengensis''. The name is derived from the creator of all reality in Chinese mythology

, the

word ''raptor'' for "thief" or "robber", and the Lufeng County. Sediments that ''Panguraptor'' were found in correspond to the Shawan Member of the

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 a ...

. These beds have also been known as the Dull Purplish Beds of the Lower Lufeng Formation, and have been dated using

magnetostratigraphy Magnetostratigraphy is a geophysical correlation technique used to date sedimentary and volcanic sequences. The method works by collecting oriented samples at measured intervals throughout the section. The samples are analyzed to determine their ' ...

to the late

Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...

to possibly

Toarcian The Toarcian is, in the International Commission on Stratigraphy, ICS' geologic timescale, an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic. It spans the time between 184.2 Megaannum, Ma (million ...

. Only the single specimen of ''Panguraptor'' has been found, which includes a partially complete articulated skeleton with the

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 ...

, neck, and back, most of the right

forelimb A forelimb or front limb is one of the paired articulated appendages ( limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used inst ...

, and both

hindlimb A hindlimb or back limb is one of the paired articulated appendages ( limbs) attached on the caudal ( posterior) end of a terrestrial tetrapod vertebrate's torso.http://www.merriam-webster.com/medical/hind%20limb, Merriam Webster Dictionary-H ...

s. It can be seen from the right side, and after being on display at Lufeng World Dinosaur Valley, it was exhibited at the

Hong Kong Science Museum The Hong Kong Science Museum is a public science museum in Tsim Sha Tsui East, Kowloon, Hong Kong, located next to the Hong Kong Museum of History. History The Hong Kong Science Museum was first conceived by the Urban Council in 1976. The co ...

from October 2013 to April 2014, before being returned to Lufeng World Dinosaur Valley for display in the exhibition hall.

Description

The holotype of ''Panguraptor,'' LFGT-0103, is the partial articulated skeleton of a subadult individual, including the skull, lower jaws, presacral vertebrae, first sacral vertebra, parts of the

pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...

and

pelvic girdle The hip bone (os coxae, innominate bone, pelvic bone or coxal bone) is a large flat bone, constricted in the center and expanded above and below. In some vertebrates (including humans before puberty) it is composed of three parts: the Ilium (bone) ...

, a left

femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...

and most of the right limb. This specimen is likely a sub-adult due to its small size (approximately 2 meters long in life), large orbit, and unfused

scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...

s and astragalocalcaneum. However, it may have been close to adulthood due to having other bones which have fused. The rather short skull is almost complete, although the

premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...

and rostral edge of the

maxilla 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 maxil ...

are missing and the nasals are partially obscured. The

orbit In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an artificial satellite around an ...

is quite large but the

antorbital fenestra An antorbital fenestra (plural: fenestrae) is an opening in the skull that is in front of the eye sockets. This skull character is largely associated with Archosauriformes, archosauriforms, first appearing during the Triassic Period. Among Extant ...

is quite small. The exposed portions of the nasal are wide and smooth and do not show any sign of a sagittal crest present in some other basal theropods. The

jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...

is positioned similarly to that of '' C. rhodesiensis'', although the

quadratojugal The quadratojugal is a skull bone present in many vertebrates, including some living reptiles and amphibians. Anatomy and function In animals with a quadratojugal bone, it is typically found connected to the jugal (cheek) bone from the front and ...

is positioned more akin to that of '' Coelophysis ("Megapnosaurus") kayentakatae.'' The rostral end of the lower jaw is missing. Teeth are preserved in the

dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...

and maxilla, and are slightly recurved yet unserrated. The centra of the cervical vertebrae gradually increase in length from the third to seventh cervical, then decrease once more to the tenth (last) cervical. The dorsal vertebrae are more compressed than those of ''

Coelophysis bauri ''Coelophysis'' ( traditionally; or , as heard more commonly in recent decades) is a genus of coelophysid theropod dinosaur that lived approximately 215 to 201.4 million years ago during the Late Triassic period from the middle Norian to ...

'' and ''C. rhodesiensis'', and their neural spines are longer than they are high and so close to each other that they form a continuous wall along the dorsal vertebral column. The

scapula The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...

is long, with a scapular blade with a straight caudal edge and concave cranial edge. The hand has four digits, with metacarpals I and II being the widest and metatarsals II and III being the longest. The first digit also has a flattened and recurved claw, the largest found in the holotype. The right ilium, though incomplete, has a stout pubic peduncle a prominent supracetabular crest. Distal portions of both

ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from the city of Naples. It is the largest of the Phlegrean Islands. Although inhabited since the Bronze Age, as a Ancient G ...

are preserved, and are straight with broad ends. The femur has a large and offset head and a longitudinal bulge on the caudolateral surface of the shaft. The

tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...

and

fibula The fibula (: fibulae or fibulas) or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. ...

are straight while the

astragalus Astragalus may refer to: * ''Astragalus'' (plant), a large genus of herbs and small shrubs *Astragalus (bone) The talus (; Latin for ankle or ankle bone; : tali), talus bone, astragalus (), or ankle bone is one of the group of foot bones known ...

and

calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel; : calcanei or calcanea) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is t ...

are unfused. Only one tarsal is exposed (likely tarsal IV) along with right metatarsals III, IV, and V and a few pedal digits. Metatarsal III is very long while IV and V taper distally. Panguraptor can be distinguished from other coelophysids by the following traits: * A diagonal ridge on the lateral surface of the maxilla, within the antorbital fossa. * An elliptical fenestra (also known in '' Zupaysaurus'') caudodorsal to the above-mentioned ridge. * a distal tarsal IV with a hooked craniomedial corner.

Classification

You ''et al.'' performed 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 ...

and found ''Panguraptor'' to be a

coelophysoid, in a clade with ''

, Coelophysis rhodesiensis,'' and ''

Camposaurus ''Camposaurus'' ( ) is a coelophysid dinosaur genus from the Norian stage of the Late Triassic period of North America. The pertinent fossil remains date back to the early to middle Norian stage, and is widely regarded as the oldest known neothe ...

'' but not ''" Megapnosaurus"'' ''kayentakatae''. Panguraptor was placed in this clade due to having a very acute angle between the horizontal and ascending processes of the maxilla, a blind pocket within the antorbital fossa, a short lateral lamina of the lacrimal, and the ascending process of the jugal making an angle less than 75 degrees with its longitudinal axis. Per this analysis, ''Panguraptor'' would be the first coelophysoid known from Asia. It is also the second definite theropod genus known from the Lufeng Formation, after ''

Sinosaurus ''Sinosaurus'' (meaning "Chinese lizard") is an extinct genus of basal theropod dinosaur which lived during the Early Jurassic (Hettangian-Sinemurian). Fossils of the animal have been found in the Lufeng Formation, in the Yunnan Province of Chi ...

''. Other small coelophysoid specimens such as FMNH CUP 2089 (some forelimb bones) and FMNH CUP 2090 (some hindlimb bones) recovered from the Lufeng formation may belong to this species, although they have been provisionally referred to '' "Megapnosaurus" kayetakatae'' until further analysis. The phylogenetic analysis of Martínez and Apaldetti in 2017 added ''Panguraptor'', with it being recovered as a member of Coelophysidae. The recovery of ''Panguraptor'' and ''Coelophysis rhodesiensis'' within Coelophysidae expands the content of the group in

to Asia and Africa, particularly in the Early Jurassic. Though the fossil record is scarce, the distribution of coelophysids

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

to the Americas in the Late Triassic before becoming globally distributed in the Early Jurassic may support the change from

fauna Fauna (: faunae or faunas) is all of the animal life present in a particular region or time. The corresponding terms for plants and fungi are ''flora'' and '' funga'', respectively. Flora, fauna, funga and other forms of life are collectively ...

l regionalism to faunal

homogeneity Homogeneity and heterogeneity are concepts relating to the Uniformity (chemistry), uniformity of a Chemical substance, substance, process or image. A homogeneous feature is uniform in composition or character (i.e., color, shape, size, weight, ...

at the beginning of the Jurassic.

References

{{Taxonbar, from=Q18325158 Coelophysoidea Dinosaur genera Early Jurassic dinosaurs Lufeng Formation Taxa named by Hailu You Taxa named by Dong Zhiming Fossil taxa described in 2014 Dinosaurs of China