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The first well-preserved coelophysoid theropod dinosaur from Asia

Authors:
  • Institute of Vertebrate Paleontology and Paleoanthropology,Chinese Academy of Sciences, Chinese Academy of Sciences
  • The Geological Museum of China

Abstract and Figures

Coelophysoid dinosaurs represent the earliest major radiation of neotheropods. These small-to-medium-sized agile bipeds lived throughout much of Pangaea during the Late Triassic-arly Jurassic. Previously reported coelophysoid material from Asia (excluding the Gondwanan territory of India) is limited to two specimens that comprise only limb fragments. This paper describes a new genus and species of coelophysoid, Panguraptor lufengensis, from the Lower Jurassic Lufeng Formation of Yunnan Province, China. The new taxon is represented by a well-preserved skeleton, including the skull and lower jaw, the presacral vertebral column and partial ribs, the right scapula, a partial forelimb, part of the pelvic girdle, and an almost complete hind limb. It is distinguished from other coelophysoid theropods by the unique combination of the following three character states: 1) diagonal (rostrodorsal-caudoventral) ridge on lateral surface of maxilla, within antorbital fossa, 2) elliptical, laterally facing fenestra caudodorsal to aforementioned diagonal ridge, and 3) hooked craniomedial corner of distal tarsal IV. Cladistic analysis recovers Panguraptor lufengensis deeply nested within Coelophysoidea as a member of Coelophysidae, and it is more closely related to Coelophysis than to "Syntarsus". Panguraptor represents the first well-preserved coelophysoid theropod dinosaur from Asia, and provides fresh evidence supporting the hypothesis that terrestrial tetrapods tended to be distributed pan-continentally during the Early Jurassic.
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Accepted by S. Brusatte: 9 Sept. 2014; published: 16 Oct. 2014
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http://dx.doi.org/10.11646/zootaxa.3873.3.3
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The first well-preserved coelophysoid theropod dinosaur from Asia
HAI-LU YOU
1,5
, YOICHI AZUMA
2
, TAO WANG
3
, YA-MING WANG
4
& ZHI-MING DONG
1
1
Key Laboratory of Vertebrate Evolution and Human Origins of Chinese Academy of Sciences, Institute of Vertebrate Paleontology
and Paleoanthropology, Chinese Academy of Sciences, 142 Xizhimenwai Dajie, Beijing, 100044, P. R. China.
Email: youhailu@ivpp.ac.cn
2
Fukui Prefectural University, 4-1-1 Kenjojima, Matsuoka, Eiheiji-cho, Yoshida-gun, Fukui 910-1195, Japan
3
Bureau of Land and Resources of Lufeng County, Yunnan Province, 651207, P. R. China
4
School of Earth Sciences and Resources, China University of Geosciences (Beijing), Beijing 100083, P. R. China
5
Corresponding author
Abstract
Coelophysoid dinosaurs represent the earliest major radiation of neotheropods. These small-to-medium-sized agile bipeds
lived throughout much of Pangaea during the Late Triassic–arly Jurassic. Previously reported coelophysoid material from
Asia (excluding the Gondwanan territory of India) is limited to two specimens that comprise only limb fragments. This
paper describes a new genus and species of coelophysoid, Panguraptor lufengensis, from the Lower Jurassic Lufeng For-
mation of Yunnan Province, China. The new taxon is represented by a well-preserved skeleton, including the skull and
lower jaw, the presacral vertebral column and partial ribs, the right scapula, a partial forelimb, part of the pelvic girdle,
and an almost complete hind limb. It is distinguished from other coelophysoid theropods by the unique combination of
the following three character states: 1) diagonal (rostrodorsal-caudoventral) ridge on lateral surface of maxilla, within an-
torbital fossa, 2) elliptical, laterally facing fenestra caudodorsal to aforementioned diagonal ridge, and 3) hooked cranio-
medial corner of distal tarsal IV. Cladistic analysis recovers Panguraptor lufengensis deeply nested within
Coelophysoidea as a member of Coelophysidae, and it is more closely related to Coelophysis than to “Syntarsus”. Pan-
guraptor represents the first well-preserved coelophysoid theropod dinosaur from Asia, and provides fresh evidence sup-
porting the hypothesis that terrestrial tetrapods tended to be distributed pan-continentally during the Early Jurassic.
Key words: Theropoda, Coelophysoidea, new genus and species, Early Jurassic, Lufeng Formation, Lufeng
Introduction
Coelophysoid dinosaurs are small-to-medium-sized agile bipedal meat-eaters that lived throughout much of
Pangaea during the Late Triassic and Early Jurassic (Tykoski 2005; Tykoski & Rowe 2004). They are among the
earliest well documented dinosaurs and represent the earliest major radiation of neotheropods (Brusatte et al. 2010;
Colbert 1989; Cope 1889; Sereno 1999). A recent study indicated that late Norian–Rhaetian theropod assemblages
were dominated by basal (early diverging) coelophysoids, whereas Early Jurassic ones were composed of
coelophysids (Coelophysis bauri + "Syntarsus" kayentakatae and all descendents of their most recent common
ancestor), dilophosaurids and basal averostrans (Ezcurra 2012). However, despite the well-documented discoveries
of derived coelophysoids in North America and Africa, the coelophysoid material that has previously been reported
from Asia is limited to two specimens comprising only limb fragments and perhaps belonging to one individual
(Irmis 2004). Here we describe a new genus and species of coelophysoid based on a well-preserved skeleton from
the same rock unit, the Lower Jurassic Lufeng Formation of Yunnan Province, China, that yielded both previously
reported specimens. Our cladistic analysis shows that the new taxon is a coelophysid coelophysoid, and is more
closely related to Coelophysis than to "Syntarsus". This new taxon represents the most basal theropod dinosaur
currently known in China, and provides fresh evidence supporting the hypothesis that terrestrial tetrapods tended to
be distributed pan-continentally during the Early Jurassic.
Institutional abbreviations: AMNH, American Museum of Natural History, New York, New York, USA;
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FMNH, Field Museum of Natural History, Chicago, Illinois, USA; LFGT, Bureau of Land and Resources of
Lufeng County, Lufeng, Yunnan, China; MNA, Museum of Northern Arizona, Flagstaff, Arizona, USA;
NMMNH, New Mexico Museum of Natural History and Science, Albuquerque, New Mexico, USA; QG, Natural
History Museum of Zimbabwe, Bulawayo, Zimbabwe.
Systematic palaeontology
Dinosauria Owen, 1842
Saurischia Seeley, 1887
Theropoda Marsh, 1881
Neotheropoda Bakker, 1986, sensu Sereno, 1998
Coelophysoidea Nopcsa, 1928, sensu Sereno, 1998
Coelophysidae Nopcsa, 1928, sensu Holtz, 1994
Panguraptor lufengensis gen. et sp. nov.
Holotype: Bureau of Land and Resources of Lufeng County LFGT-0103, an articulated partial skeleton that
includes the skull and lower jaw, the presacral vertebral column, part of the ribs, the right scapula and partial right
forelimb, part of the pelvic girdle and parts of both hind limbs, the right hind limb being almost complete.
Type locality and horizon. Lufeng County, Yunnan Province, China. Shawan Member of the Lufeng
Formation (Fang et al. 2000). This is equivalent to the Dull Purplish Beds of the Lower Lufeng Formation (Series)
as defined by Bien (1940). In the Lufeng area, the Lufeng Formation overlies the Lower Jurassic Yubacun
Formation and is disconformably overlain by the Middle Jurassic Chuanjie Formation (Cheng et al. 2004). The age
of the Lufeng Formation is probably early - middle Early Jurassic (Hettangian - Pliensbachian), based on
biostratigraphic correlations (Luo & Wu 1995). Magnetostratigraphic analysis has indicated late Sinemurian –
?Toarcian for the age of the Lufeng Formation (Huang et al. 2005).
Etymology. The genus name is from “Pangu” (Chinese), well known in Chinese mythology as the first living
being and the creator of all reality, and “raptor” (Latin), meaning “thief” or “robber”. The specific name refers to
“Lufeng County”, one of the world’s richest sources of Early Jurassic terrestrial vertebrate fossils.
Differential diagnosis (for genus and species by monotypy). A coelophysid theropod with the unique
combination of the following three character states (autapomorphies marked with *): 1*) diagonal (rostrodorsal-
caudoventral) ridge on lateral surface of maxilla, within antorbital fossa; 2) elliptical, laterally facing fenestra
caudodorsal to diagonal ridge mentioned in previous character state, also present in Zupaysaurus rougieri (Ezcurra
2006); 3*) distal tarsal IV with hooked craniomedial corner.
Description and comparisons. LFGT-0103 is an articulated partial skeleton, exposed mainly in right lateral
view (Fig. 1). The specimen is small, with a preserved skull length (taken from the rostral end of the maxilla to the
ventral end of the quadrate because the premaxilla is missing) of 11.1 cm, presacral vertebral column length of ~58
cm (trunk/neck length ratio about 1.3), scapula length of 8.6 cm, femur length of 16.4 cm, and tibia length of 18.2
cm (Table 1). Early theropods normally had a long tail contributing more than half of the total body length,
implying that this dinosaur may have measured around two metres long at the time of death. Height at the hip was
probably about half a metre.
LFGT-0103 probably represents a sub-adult individual, as indicated by the relatively small body size of the
specimen, the large orbit, the lack of fusion between the scapula and coracoid, the lack of ossification of distal tips
of the ischia, and the separation of the astragalus and calcaneum. However, the neurocentral sutures are fused,
cervical ribs are fused to their respective centra, the ilium is fused to the ischium, and distal tarsal 3 is probably
fused to metatarsal III, indicating that the specimen may be close to an adult individual. Growth curves of femur
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length vs. age for C. rhodesiensis (Chinsamy 1994) and C. bauri (Rinehart et al. 2009) suggest, respectively, that
this dinosaur died at a sub-adult age of around four years or around two years.
TABLE 1 . Measurements of selected elements of Panguraptor lufengensis gen. et sp. nov. LFGT-0103. All right side, in
mm.
skull length (from preserved rostral end to ventral end of quadrate) 111
preorbital length (preserved) 60
length of internal antorbital fenestra 37
height of antorbital fossa along at caudal margin along lacrimal 20
orbit length 30
orbit height 35
maximum height of skull across orbit midpoint 48
length of infratemporal fenestra 23
height of infratemporal fenestra 31
width of skull above middle antorbital fossa 12
narrist width of skull above orbit 10
length of low jaw (preserved) 111
Cervicals centrum length total height
axis 18.5 22
C3 24 14.5
C4 28 16
C5 29 15
C6 31 23
C7 30.5 22
C8 28.5 -
C9 28 -
C10 23 -
Dorsals centrum length total height
D9 24 32
D10 26 36
D11 26 35
D12 25 34
D13 23 23+
sacral 1 length 22+
scapula length 86
maximum width of scapular blade 20
minimum width of scapular blade 11
humerus length 58+
metacarpal I length 10+
metacarpal II length 36
metacarpal II width at midportion 5
metacarpal III length 34
metacarpal III width at midportion 3.5
metacarpal IV length 22
metacarpal IV width at midportion 3
......continued on the next page
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FIGURE 1. Skeleton of Panguraptor lufengensis gen. et sp. nov. (LFGT-0103). a, photo; b, interpretive line drawing.
Abbreviations: as, astragalus; ax, axis; ca, calcaneum; cer 3–10, cervical 3-10; dor 7, dorsal 7; dt IV, distal tarsal IV; fdt III,
fused distal tarsal III; fi, fibula; hu, humerus; il, ilium; lfe, left femur; lj, lower jaw; is, ischia; ma, manus; mt IV, metatarsal IV;
mt V, metatarsal V; pd III, pedal digit III; ph IV-1, phalanx 1 of pedal digit IV; ra, radius; rfe, right femur; sac 1, sacral 1; sc,
scapula; sk, skull; ti, tibia; ul, ulna.
TABLE 1. (Continued)
Dorsals centrum length total height
manual phalanx I-1 length 17
manual phalanx I-2 (Ungual) length 21
manual phalanx II-1 length 18
manual phalanx II-2 length 22
manual phalanx II-3 length 12+
manual phalanx III-1 length 10
manual pahlanx III-2 length 8
manual phalanx III-3 length 9
manual phalanx IV-1 length 4+
Iliac blade length 63+
iliac blade height above acetabulum 44
femur length 164
tibia length 182
tibia length with astralas 188
fibula length 178
metatarsal IV length 98
metatarsal V length 41
pedal phalanx III-2 length 22
pedal phalanx III-3 length 23
pedal phalanx III-4 length 17
pedal phalanx IV-1 length 18
pedal phalanx IV-2 length 17
pedal phalanx IV-3 length 13.5
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FIGURE 2. Skull and lower jaw of Panguraptor lufengensis gen. et sp. nov. (LFGT-0103) in right lateral view. a, photo; b,
interpretive line drawing. Abbreviations: af, additional fenestra; ar, alveolar ridge; at, atlas; ax, axis; dr, diagonal ridge; emf,
external mandibular fenestra; f, frontal; gd, groove on dentary; iaf, internal antorbital fenestra; itf, infratemporal fenestra; j,
jugal; l, lacrimal; m, maxilla; n, nasal; or, orbit; pa, parietal; pf, promaxillary fenestra; po, postorbital; pr, prefrontal; q,
quadrate; qj, quadratojugal; rsur, ridge on surangular; sq, squamosal; stf, supratemporal fossa.
Skull and lower jaw. The skull is almost complete, and exposed mainly in right lateral view (Fig. 2).
Assuming that the missing premaxillary body contributed ~10% of the total skull length (based on skull
reconstruction of the “Syntarsus”c kayentakatae holotype of Tykoski 1998, figure 6), the intact skull of
Panguraptor would have measured 12.4 cm, about half as long as adult skulls of C. bauri (AMNH 7224: 20.7 cm)
(Rinehart et al. 2001), C. rhodesiensis (QG 193: 22.0 cm) (Raath 1977), and “S.” kayentakatae (MNA V2623:
23.0 cm) (Rowe 1989). The preorbital length without the premaxilla (6.0 cm) is slightly more than half the
preserved skull length (54%), and the maximum rostrocaudal length of the large triangular internal antorbital
fenestra is about two thirds of the preserved preorbital length. In contrast, the internal antorbital fenestra is
relatively small (about half as high as the orbit and half as long as the maxilla) in a similar-sized (skull length 12.3
cm) juvenile specimen of C. bauri NMMNH P-42200 (Rinehart et al. 2009). The orbit is subcircular, though with a
relatively straight rostral edge, and has a maximum length of 3.0 cm and maximum height of 3.5 cm. The
maximum height of the skull at the mid-orbital level is 4.8 cm, or 39% of the estimated skull length. The
postorbital portion of the lateral side of the skull is almost completely occupied by the rectangular infratemporal
fenestra, which is 2.3 cm long and 3.1 cm high.
Most of the lateral surface of the right maxilla is exposed. The rostral process of the maxilla and the
rostrodorsal edge of the maxillary body proper (the portion between the rostral process of the maxilla and the
rostral rim of the internal antorbital fenestra, but not including the ascending process) have been lost, and the
preserved rostrodorsal margin appears to represent the caudoventral rim of the external naris. The maxillary body
proper is rostrocaudally longer than dorsoventrally high as in C. rhodesiensis (QG 193), but the opposite is true in
“S.” kayentakatae (MNA V2623). The lateral surface of the maxillary body proper bears a diagonally aligned
(rostrodorsal-caudoventral) ridge between the rims of the internal and external antorbital fenestrae. Caudodorsal to
this ridge an elliptical fenestra opens laterally, while rostroventral to it are two small fenestrae separated by another
short vertical ridge: a rostrally positioned fenestra that is more or less round in lateral view, and a caudally
positioned one that is triangular. We interpret the rostral one as the promaxillary fenestra, which occupies a similar
position in “S.” kayentakatae. However, the part of the antorbital fossa that lies just caudal to the promaxillary
fenestra is smooth in “S.” kayentakatae, rather than interrupted by additional fenestrae. The longitudinal and very
prominent alveolar ridge runs along the entire ventral border of the external antorbital fenestra and extends onto the
rostral process of the jugal under the orbit. In “S.” kayentakatae the alveolar ridge extends further caudally,
continuing onto the caudal process of the jugal. The ascending process of the maxilla is broken away from the rest
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of the bone, and judging from the orientation of the nasal and the shape of the antorbital fenestra, it would have
been elevated about 30 degrees above the horizontal in the intact skull. The orientation of the ascending process is
similar in C. rhodesiensis, but steeper in “S.” kayentakatae.
FIGURE 3. Cervical vertebrae 2-6 of Panguraptor lufengensis gen. et sp. nov. (LFGT-0103) in right lateral view.
Abbreviations: a rib, axial rib; C4-C6, cervicals 4–6; cp, caudal pleurocoel; rib of cer 3, rib of cervical 3; rp, rostral pleurocoel;
sg, shallow groove on axis.
A large portion of the right nasal is visible, exposed in dorsal view. A large and subcircular hole penetrates the
rostral half of the bone, probably not a true feature of the animal in life. The dorsal surface of the nasal is smooth,
and lacks any evidence of a parasagittal vertical crest. The transverse width of the nasal above the antorbital
fenestra exceeds the width of the frontal above the orbit. The transverse width of the prefrontal exceeds its
rostrocaudal length. A depression is visible rostrodorsal to the orbit, at the junction between the nasal, prefrontal
and frontal.
The robust and almost vertical ventral ramus of the lacrimal separates the caudal border of the antorbital fossa
from the rostral border of the orbit. The caudodorsal corner of the lateral surface of the lacrimal bears a
rostrodorsal-caudoventrally running groove, the caudal edge of which leads to a small foramen, which should be
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the exit of the naso-lacrimal duct. The lateral lamina seems to be restricted to the caudal margin of the ventral half
of the ventral ramus of the lacrimal, and lacks an expansion at the dorsal end. The rostral process seems to
contribute at least one third to the dorsal border of the antorbital fenestra because its rostral end appears to be
broken off and incomplete.
The jugal borders the orbit ventrally, and seems to contact the maxilla ventral to the lacrimal as in C.
rhodesiensis (QG 165) (Bristowe & Raath 2004). The caudal edge of the postorbital process of the jugal forms an
angle of approximately 70
0
with the dorsal edge of the caudal process, rather than a right angle as in “S.”
kayentakatae. The ventral branch of the short and forked caudal process is slightly longer than the dorsal branch,
and the slot for the quadratojugal formed by the two branches extends rostrally to a point located ventral to the base
of the postorbital process.
The quadratojugal forms the ventral half of the caudal border of the infratemporal fenestra, and contacts the
ventral process of the squamosal. This squamosal-quadratojugal contact excludes the quadrate from the border of
the infratemporal fenestra, as in “S.” kayentakatae (Tykoski 1998) but in contrast to the condition in C.
rhodesiensis (Raath 1977). The frontal-parietal contact region is damaged. The dorsal surface of the frontals bulges
along the midline at the level of the caudodorsal corners of the orbits. The rostral portion of the well-developed
supratemporal fossa is bounded ventrolaterally by a sharp rimmed edge on the rostral process of the postorbital.
As preserved, the lower jaw is oriented almost perpendicular to the long axis of the skull, and the mandibular
glenoid fossa is preserved close to the quadrate head and considerably displaced from the distal condyles of the
quadrate. Judging from the position of the external mandibular fenestra, which is normally located underneath the
orbit, the rostral end of the lower jaw comparable to the premaxilla is missing; in actuality the preserved lengths of
the skull and the lower jaw are same. The caudal end of the lower jaw is overlapped by the atlas and axis. It is
difficult to discern the sutures among the bones of the lower jaw. A longitudinal groove is visible on the lateral
surface of the middle portion of the dentary, and a longitudinal ridge is present on the lateral surface of the
surangular.
Six maxillary teeth are visible, four in the rostral one-third of the maxilla and two at the caudal end of the bone
under the caudal part of the orbit. Ten dentary teeth are preserved. All maxillary and dentary teeth are slightly
recurved, and bear few or no serrations.
Vertebral column and ribs. Ten cervicals, 13 dorsals and the first sacral vertebra are preserved (Fig. 1). The
atlas is disarticulated in such a way that its individual components are hard to identify (Fig. 2). The axis is the
shortest of all the cervicals, except for the atlas. The exposed right lateral side of the axial centrum does not bear
any clearly visible pneumatic excavations (pleurocoels), although a shallow longitudinal groove is present (Fig. 3).
Although the neural spine is not complete, it is clearly tall and craniocaudally long.
The centrum of the third cervical is about 130% longer than that of the axis. Centrum lengths continue
increasing steadily until cervical 7, then decrease gradually so that the last cervical is about equal in centrum length
to the third one. All postaxial cervicals are elongate, with mid-central constrictions and stout prezygapophyses. The
lateral surfaces of the centra of cervicals 4-5 bear deep, elongate pneumatic foramina (“pleurocoels) (Fig. 3). In
cervical 4, the rostral part of the pleurocoel is larger and deeper than the caudal part, but in cervical 5 the opposite
is the case. Only weak pleurocoels are present on cervicals 6-7, and the condition in the rest of the cervicals is
unknown due to poor preservation.
The shafts of the axial ribs are preserved ventral to cervical 3, and one of them extends far enough to lie partly
ventral to cervical 4 (Fig. 3). Although it is hard to trace individual ribs in the remainder of the cervical series, the
existence of bundles of elongated ribs is evident. The third cervical rib has a small capitulum and tuberculum. In
contrast, the remaining cervical ribs have well-developed capitula and tubercula, and are fused to their
corresponding vertebrae.
The centra of the well-preserved caudal dorsals are less than twice as long as they are high, whereas the
corresponding centra are about twice as long as high in C. bauri and C. rhodesiensis. The neural arches of the
cranialmost seven dorsal vertebrae have sheet-like transverse processes that are strongly backswept and triangular
when viewed from above. In contrast, the transverse processes of the eighth and ninth dorsals are rostrocaudally
narrow and taper laterally (Fig. 1). The parapophysis is short, and situated cranioventral to the transverse process.
The neural spines are longer than high and so close together that they almost form a thin, continuous wall along the
dorsal midline of the axial column. Only the first sacral is exposed, while the others are covered by the right ilium.
This vertebra is slightly shorter than the caudal dorsals, and has a round, flat cranial articular surface.
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Appendicular skeleton. An almost complete right scapula is exposed in lateral view, and is broken only near
the articulation with the coracoid (Fig. 1). The scapula is long. Its blade is expanded distally, and constricted
proximally. The caudal edge of the blade is straight, while the cranial edge is concave. The coracoid is not
preserved, indicating that it is not fused with the scapula.
In describing the forelimb, we orient it so that it is held vertically with the radius positioned cranial to the ulna
and the flexor surface of the manus facing medially. The distal portion of the right humerus, including what seems
to be the part of the deltopectoral crest that lies distal to the apex, is preserved (Fig. 4). The distal end of the
humerus curves slightly cranially. In distal view, the medial condyle is slightly mediolaterally wider than the lateral
one.
The right radius and ulna are preserved, and are exposed mainly in caudal view (Fig. 4). The well-preserved
proximal end of the radius is well preserved, and can be seen in proximal view. The proximal surface is
subrectangular in shape, being craniocaudally elongated, and protrudes slightly caudally, unlike the strongly
caudally protruded condition in the Lufeng specimen FMNH CUP 2089 (Irmis 2004). The distal end of the radius
is not exposed. The ulna is broken into four pieces, the distalmost of which is detached from the others (Fig. 1).
FIGURE 4. Forelimb of Panguraptor lufengensis gen. et sp. nov. LFGT-0103. Abbreviations: dhu, distal end of humerus; dpc,
deltopectoral crest; mtc I, metacarpal I; mtc IV, metacarpal IV; ph I-1, phalanx 1 of digit I; ph III-3, phalanx 3 of digit III; ph IV-
1, phalanx 1 of digit IV; pra, proximal end of radius.
The right manus is represented by four digits, exposed mainly in medial and caudal views (Fig. 4). Metacarpals
I and II are more than twice as wide as metacarpals III and IV. Metacarpal I is less than half the length of
metacarpal II, which is about the same length as metacarpal III. The slender metacarpal IV is preserved between
metacarpals II and III, and measures about two thirds the length of metacarpal III. Phalanx 1 of digit 1 is not
completely exposed, but seems to be as long as phalanx 2 of digit 2, one of the longest phalanges in the manus. The
claw of digit 1 is almost the same length as phalanx 2 of digit 2, and is transversely compressed, strongly recurved,
and equipped with a large flexor tubercle. The claw of digit 2 is not completely exposed, but seems less well
developed than that of digit 1. Three slender phalanges of digit 3 are visible, but there is no trace of a claw. Only
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one small phalanx appears to be present in digit 4, although an additional small terminal nub may also exist as in
“S.” kayentakatae (Tykoski 1998). Therefore, the manual phalangeal formula is 2-3-3-1/2-X.
Part of the right ilium is preserved (Fig. 1). Most of the pre- and postacetabular processes are missing, but the
portion surrounding the acetabulum is well preserved. The pubic peduncle is much stouter than the ischial
peduncle, and bears an approximately triangular articular surface for the pubis in distal view. The acetabulum is
bordered by a prominent supracetabular crest craniodorsally. The bone surface that forms the caudal and
caudodorsal parts of the wall of the acetabulum is relatively narrow transversely and faces slightly laterally. The
ischial peduncle is fused to the ischium, although a suture can be vaguely discerned.
The distal portions of the ischia are preserved (Fig. 5). They are straight and mutually appressed, with
moderately enlarged ends that are about two times broader craniocaudally than the shaft.
FIGURE 5. Distal portions of ischia and right hind limb of Panguraptor lufengensis gen. et sp. nov. LFGT-0103.
Abbreviations: as, astragalus; ca, calcaneum; dt IV, distal tarsal IV; fdt III, fused distal tarsus III; fi, fibula; lis, left ischium; mt
IV, metatarsal IV; mt V, metatarsal V; ris, right ischium; ti, tibia.
The right femur is broken into two portions. The proximal portion is exposed in cranial and lateral views, and
the distal portion in lateral view (Fig. 1). The femoral head is enlarged, well offset from the shaft, and directed
craniomedially, with its end turned slightly downward. In proximal view, no clear constriction is apparent, and the
proximal surface tapers caudolaterally. A longitudinal bulge is present near the proximal end of the caudolateral
surface of the shaft. The anterior trochanter begins at the level of the ventral edge of this bulge, centered
transversely on the craniolateral surface of the femur, as a prominent boss that extends ventrally. A tubercle lying
caudal to the ventral end of the anterior trochanter may be homologous to the trochanteric shelf. The area proximal
to the anterior trochanter is shallowly depressed.
The right tibia and fibula are preserved, and exposed mainly in proximal and lateral views (Fig. 1). In proximal
view, they combine to form a flat and roughly square articular surface for the femur. The cnemial crest forms the
craniolateral corner of this surface, while the craniomedial corner consists of two small tubercles of the tibia that
protrude slightly above the surface. The shafts of both the tibia and fibula are straight, and the fibula is closely
appressed to the lateral surface of the tibia. The distal ends of both bones are enlarged.
The astragalus and calcaneum are clearly not fused to each other, or to the tibia and the fibula (Fig. 5). The
astragalus is visible in caudodistal view, and the calcaneum is slightly displaced from its original position. The
articular surfaces between the astragalus and calcaneum are flat. In lateral view, the calcaneum acquires a
semilunate shape.
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One distal tarsal is visible, and is very likely to be distal tarsal IV based on its position and shape (Fig. 5). Its
proximal surface is slightly depressed, and it has an irregular outline with an expanded cranial portion. The
craniomedial corner of the bone is hooked, rather than a simple right angle as in “S.” kayentakatae (Tykoski 1998).
Right metatarsals III, IV and V are partially exposed (Fig. 5). The proximal end of metatarsal IV is positioned
slightly distal and dorsal to that of metatarsal III, indicating that the latter incorporates a fused distal tarsal III. The
proximal end of metatarsal III extends ventrally well beyond the metatarsal shaft, and bears a clear ventrolateral
process. The proximal surface of metatarsal IV has a triangular shape matching that of distal tarsal IV, with an
expanded dorsal portion and a tapering ventral end. The fifth metatarsal is splint-like, with a proximal end at the
same level as that of metatarsal III and a distal end that tapers abruptly to a point. The four phalanges of pedal digit
III are preserved, as are the proximal three phalanges of pedal digit IV (Fig. 1).
Cladistic analysis
A cladistic analysis was performed in order to assess the phylogenetic position of Panguraptor. The data matrix
was based on that of Ezcurra & Brusatte (2011), which consists of 339 characters and 43 dinosaur and other
archosauriform taxa. In our analysis, Panguraptor was added, and two characters (222 and 316) were recoded from
state ‘0’ to state ‘1’ for “S.” kayentakatae (see Appendix for data matrix). Erythrosuchus was used as an outgroup.
A traditional search in TNT (Goloboff et al. 2008) with 100 Wagner tree replications and tree bisection and
reconnection (TBR) branch swapping (saving 100 trees per replicate) recovered 8 MPTs (most parsimonious trees)
of 1011 steps each. The CI (consistency index) was 0.400, and the RI (retention index) was 0.691. The strict
consensus of these eight trees is shown in Fig. 6, with absolute bootstrap frequencies and all decay index values
equal to or greater than one.
FIGURE 6. Phylogenetic placement of Panguraptor lufengensis gen. et sp. nov., based on strict consensus tree obtained in this
study. Absolute bootstrap frequencies and decay index values are indicated.
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The strict consensus recovered Panguraptor as a member of Coelophysidae, and placed Panguraptor more
closely related to the grouping of C. bauri + (C. rhodesiensis + Camposaurus arizonensis) than to “S.”
kayentakatae. The strict consensus also recovered Liliensternus liliensterni and Zupaysaurus rougieri as members
of Coelophysoidea.
The monophyly of Coelophysoidea is supported by four unambiguous synapomorphies: 17(1>2): more than 18
maxillary teeth; 24(0>1): sharp longitudinal ridge present on lateral surface of the maxilla; 194(0>1): caudal
margin of postacetabular process of ilium notched or indented; and 320(3>2): sublacrimal part of jugal squared off
rostrally with small dorsally directed prong, slightly overlapping lacrimal. Of these character states, only 24(1) is
clearly present in Panguraptor.
Coelophysidae is supported by eight unambiguous synapomorphies: 47(0>1): width of ventral process of
squamosal less than one quarter of its length; 129(0>1): cervical vertebrae with rimmed depression on caudal part
of centrum; 139(0>1): sacral centra co-ossified at ventral edges; 230(1>2): dorsolateral margin of proximal portion
of femur bearing rounded ridge; 234(0>1): trochanteric shelf present proximal to fourth trochanter on femur;
240(0>1): infrapopliteal ridge present between medial distal condyle and crista tibiofibularis on femur; 265(0>1):
distinct deep fossa on medial side of fibula; and 323(1>0): small contact between squamosal and quadratojugal. Of
these character states, 47(1), 129(1), 230(2), 234(1), and 323(0) are present in Panguraptor.
The Panguraptor + Coelophysis clade is supported by four unambiguous synapomorphies: 316(0>1): angle
between ascending and horizontal processes of maxilla less than 35; 318(0>1): lateral surface of maxillary
antorbital fossa at base of ascending process bears at least one deep, large, blind pocket that is subcircular or oval in
shape; 319(3>2): lateral lamina of lacrimal with no interruption of the lacrimal antorbital fossa and restricted to the
caudal margin of the ventral ramus, and 321(0>1): angle between ascending process and longitudinal axis of jugal
less than 75, with ascending process strongly caudodorsally oriented. All four of these features are present in
Panguraptor.
The Coelophysis bauri + (Coelophysis rhodesiensis + Camposaurus arizonensis) clade is supported by five
unambiguous synapomorphies: 27(1>0): absence of promaxillary foramen; 322(1>0): tapering ventral process of
squamosal; 323(0>2): no squamosal-quadratojugal contact; 324(0>1): no postorbital participation in supratemporal
fossa; and 329(1>2): significantly elongated caudal dorsal vertebrae, with centrum length at least 1.33 times height
of cranial articular surface. All these features are absent in Panguraptor.
Discussion
There are three other Early Jurassic coelophysoid specimens not included in the above cladistic analysis, and we
compare them with Panguraptor lufengensis.
Irmis (2004) referred FMNH CUP 2089 (distal humerus, proximal radius and ulna, fragments of metacarpals
and phalanges) and FMNH CUP 2090 (right distal tarsals II and III, and proximal metatarsals II and III) from the
Early Jurassic Lufeng Formation in Lufeng Basin to cf. Megapnosaurus (=Syntarsus). Comparable portions of
FMNH CUP 2089 and the holotype of Panguraptor lufengensis are similar in morphology, and both specimens
share a cranially curved distal end of the humerus and uneven distal humeral condyles. Distal tarsal III of the
Panguraptor lufengensis holotype possesses a ventrolateral process that is not present in FMNH CUP 2090, but
this feature may have been lost in the latter specimen due to poor preservation. However, pending further
discoveries here we still refer FMNH CUP 2089 and 2090 to cf. Megapnosaurus.
Carrano et al. (2005) restudied the holotype and only known specimen of Segisaurus halli Camp 1936, a small
coelophysoid from the Lower Jurassic of North America. Segisaurus is distinctive in having a very long and
slender scapula, measuring 93 mm long and 20 mm wide at the distal end. The equivalent measurements are 86 mm
and 20 mm in Panguraptor lufengensis. Histological study indicates that the holotype individual of Segisaurus
halli was a subadult at the time of death (Carrano et al. 2005).
Tykoski (2005) argued for the existence of a distinct ‘Shake-N-Bake’ coelophysoid species from the Kayenta
Formation, which also yielded “S.” kayentakatae, and showed that the ‘Shake-N-Bake’ species was a small
coelophysoid. The proximal tarsals, the astragalus and calcaneum, are clearly well fused to the tibia, and to each
other in the small ‘Shake-N-Bake’ species, but such fusion is not apparent in the holotype of Panguraptor
lufengensis despite the larger size of this specimen.
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The Lufeng Saurischian Fauna (Young 1951), or more appropriate the Early Jurassic Lufeng Dinosaur Fauna,
represents one of the richest dinosaur faunas in the world, and provides critical information regarding dinosaur
evolution and biogeography during the Early Jurassic. Eight species of sauropodomorphs (Lufengosaurus huenei
Young, 1941, L. magnus Young, 1947, “Gryposaurus” sinensis Young, 1941, Yunnanosaurus huangi Young, 1942,
Y. ro bus t u s Young, 1951, Jinshanosaurus xinwaensis Zhang and Yang, 1995, Chuxiongosaurus lufengensis L et al.,
2010, Xixiposaurus suni Sekiya, 2010) and two species of ornithischians (Tatisaurus oehleri Simmons, 1965 and
Bienosaurus lufengensis Dong, 2001) have been reported from the Lower Jurassic Lufeng Formation in the Lufeng
area. In contrast, only one theropod, Sinosaurus sinensis Young, 1948 has been previously confirmed to exist in
these strata (Xing 2012). The discovery of Panguraptor lufengensis adds a second theropod to the known fauna of
the Lufeng Formation.
Acknowledgements
We are grateful to the crew of the Lufeng World Dinosaur Valley for discovering, excavating, and preparing the
specimen, to Ms. Jin-Ling Huang for producing the line drawings, and to Dr. Corwin Sullivan for detailed review
of the first draft. Thanks also go to reviewers (Dr. Ronald Tykoski and Mr. Martin Ezcurra) and editor Dr. Stephen
Brusatte. Funding was provided by the Hundred Talents Project of the Chinese Academy of Sciences, the National
Natural Science Foundation of China, the National Basic Research Program of China (973 Program), and the
Bureau of Land and Resources of Lufeng County.
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APPENDIX. Phylogenetic data matrix.
Erythrosuchus_africanus
00010001200002000100000000000000000010000000000000001010000000001000000000000000000000001000000000000
100000000000110000000?00000000010?000000000000000??1000?001?000101?000?00??01??0???000?0000001000000000
000??000000000010001000000000001??00000000000000000000010000010000000?00?00?0000000000?000000000001000
?0???????000010110101001000001000
Euparkeria_capensis
000100000000?000100000000100000000000000000100000000[01]0002000000010100000000000001000000000000000000
011000000000001000000000000000010000010000000000000001000?00000010000000000??010?000?00000000000000000
000000??0000000000100010001000000000000000000000000000000000?00000000000?00?0000000000010?000000000000
0000101000001002100000110??000001000
Revueltosaurus_callenderi
000000002000030110101001020101000000000010010100011012002000000011211000000?20????0?001?001?0000000011
0000110000?1110000000001000020100011000000000000?01001010000000110110?0???000?0?0?????00000000000000000
00??000000000000011000100000000000000000000010000001000010000?000100?00?0001101101021010?0000?000000??
10012112?2022001??01???00000100?
Aetosaurus_ferratus
10100000111000010110011002010?00?10000000011010?011012002011000??12?1?00000?????1???001000110??21??1??00
01000010?111??????????????????????0?000000???0001?01010000000110110?00??00000???00000?[01]000?00000000000?
??00?0000000?0011000100000000000000?00000?10?000010??0????0??????????????????1?1?21?????????0??0???2100121
12210221??0110???000001000
Arizonasaurus_babbitti
??????1????00?00??00011000000100???????000?100000000?000200000001001100000000011?010001?001?000001101100
000000000111000010?0000000010?0000?11000100???0010?1010?????????????????????????????00100010020000?0011100
21111100011?10000100000001001000100?0????????????????????????????????????????????????????????????0???????200?
20??00?10110????????
Effigia_okeeffeae
011000104??0?20???01?111000000000000000011?000120010???000?1000001211010?????111?0?0001?001000?010010001
02????????11???000?01?0?1101??000111111011?0110?00?1001??0?10?1?000100??00?000??????10110000021010?0011110
2122100001??30100??0000011??101110000001?0100010010100010000100?0??00011011?1?11?10000001010100?10??????
00102200??10000?000001000
Batrachotomus_kuperferzellensis
2?0?000?10010100011001100000111101001001100110011000?00020000001112110011000?01111110011101?00000110110
000000000011100000000000000211000100010001001000010?101000000011110??????????????????0010001000000010011
00010010100011010000100000001001010000000010000001010010000??????0??1????110?101021?100000??000?10?2111
?10?012021001000101100000100?
Postosuchus_kirkpatricki
0001000010000101100000020101111101001001101110011100111031000001112110?110012011??1?0011111100020110110
0000000000111??00???????000?1?00000000000000??0??1011011001000111?00100??00???0??0000001?0010000000??????
??1??1110001?010000100000000001010?00000010000001011011000?????00?01?0001102101111?10000001010110?0111?1
00?120031010110011000001000
Dromicosuchus_grallator
2000000?200011111100000001001?1000010001111110?1?10011102110????????????100?????????001?011?000?021011000
000000001?1????????0??0000?1???000?0000000?00201011111001110011?00100?10???????????00110010000100?00????01
??????????010000100000100001010?00000010000001001011000?????00??1????11021011?1?11?00???????????1110100012
0220??01?1?210????????
Eudimorphodon_ranzii
1110???010100?0?2000011000000??0??0?000???0?????0?00000?0??1????????????????????????10??0?100?000000?00?001
?0000?101????1???0000000?1?0?0?00?????0?01000????110000110?0?001??????00000110?110?0100???0?00?00000??00??
0?0000001????????0?00??2??????0?0??????0?????????????1?0???0?1??1????1?01?????11?001?1??0110000??????1200031?
211000210000???0?
Dimorphodon_macronyx
2100??1?2010000?11?0011000000?????00000???010?0?00000??00??0????????????????????????101?20?00??00??0?00??01
?0000?1?1????1?1??????00?1??000?????????010??????110000110?????1?0???00000?110?1?000100?000100?00000??00???
?0000?0110000?00000000??001020000000?00?000?0??1?00?11000?0?1??101??1??1?????110001?001??1?000???????2000
?0?21???011000011000
Lagerpeton_chanarensis
??????????????????????????????????????????????????????????????????????????????????????????????????????????????????
?????????????????????0??000000000?????????????????????????????????????????0000000000000000000??00000000000011
11001100010?0100010?0010100?0000000010100011100110?1001011?1?01???2011100000000001200????????????????????
210?0000001
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Dromomeron_romeri
??????????????????????????????????????????????????????????????????????????????????????????????????????????????????
???????????????????????????????????????????????????????????????????????????????????????????????????????????111100
1100010102?001110111100000000000101?00??????10?1001011?1?01???2?1???????????????0???????????????????????0000
010?
Dromomeron_gregorii
??????????????????????????????????????????????????????????????????????????????????????????????????????????????????
???????????????????????????????????????????????????????????????????????????????????????????????????????????111100
1?01010100000111011110000000000?????00??????1????????1??????????????????????????????????????????????????000???
??
Marasuchus_lilloensis
????????????????????????????????????????????????????????????????01210000?001100????0????????????????????????????
???101000?0000000000?00000000000000000??1?????1000?101???0??????????????????0000000000100000010??11000000
01011100001001001100000?0?00001000000000101010000110010101001001?010000100111000000000012?0???????00???
???????010000011000
Eucoelophysis_baldwini
??????????????????????????????????????????????????????????????????????????????????????????????????????????????????
???????????????????????????????????????????????????????????????????????????????????10????????????????????0??12001
??1110001???0??1100010?100??????????????????????????????????????????????0???????????????????????????????????????
Sacisaurus_agudoensis
??????????????010?00000000000?????????????????????????????????????????????????????????1?????0?101????????101001
110?1????????????????????????????????????1?????1?????????????????????????????002???010?0000100?0??1????????????
100111011100010000???100010010110001?????????????????????????????????????????????????????????200??????????010
000?????
Silesaurus_opolensis
0?0?000?20?0000100000000000001?000?0???00?0100100000?10020001??00021010000011001?001001?001001121001010
00100001010111100100000000001000000000000110000??110101100001010?000?00??0???????????002000010100001001
0??110110000111012011101110[0
1]01000010110001001011000101110001?1???0101001010?0?01???2011110?00010101210????????00?20????000110000111
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Eocursor_parvus
???????????????????????????????????????????????0????????????????01210100??????????????????1??100?001??10?0?1011
?11?1????????0?0001?0??00?10?001?1?0?????11????11?0?101??????????????????????00210001020000?11????01??10?002
1?110002101200011110010?000020000?2010101?10??1??????1???????????????????11???000???????0?????????????????1?
???0100???0?
Pisanosaurus_mertii
???????????????11??01??0????????????????????????????????????????????????????????????0?????????000001?01200?1111
111?1?????0?0???0000???00??????????????????????????????????????????????????????????0?????????0?????????????2????
???????????????001??0000?0000010001011100?1?????01010?111??0?01???21111???0?01??????0????????0??????????12??
0000010?
Heterodontosaurus_tucki
000100000011101100001000000001000000000011010002000003001000101?0121?110000111???0??001101111?0[02]0001
00120[02]11100111111000000001000000?000211?0011100001?111?0?1110011010001011000100001120000?021000002100
011110??01??100002011?0??????20?010110010?0000200001101??0110???111000?1?10?1?1??1?01?????111100?00101?120
0??????20001100101000100?00???00
Lesothosaurus_diagnosticus
00010?0?3000001110001000000001000000000011010000000003000010101001210100000??1??10??0011011111000001011
2000101101111????000001000000??0001??0?111?0??1??110???1100?1000?01010???000?00??000000210?0102100011110?
?01??1000021111000210120001011?0100000020000020101?111?00?????001?100111??0?01???21111100000101???00?????
??0001010000000??010????00
Scutellosaurus_lawleri
??0??0?????????1?0?01??000?????????????00??10??2???????00??????0???????0?????1??????0?1?????110[02]0????1????01
011011?10???00?00??0?000??00?00??0???00?00?11??0?000001101000???????0???????????0?21000102?100111?????1??0?
????1111000?1?12000?0110010000002000002010?01?10001?????010100111????01???21110?????0??????01???000?????10
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Saturnalia_tupiniquim
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01?1???0???0??100001??0000000000001001??1000?111000111??010?????????????????00200001011000100110011001210
0211010002102100111110011?0000200001111010111001111110010101111??0?01???2011110000011101?00??????????????
?????020000001000
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Plateosaurus_engelhardti
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0110011001210021101000210210001111001100000200000111010111000111110010101111??0?01???201101000001110110
0???????2001010?000001000000[01]000
Efraasia_minor
0?001?10100101012?0000?000000100??11000001?1001????00??1?000????01210100000011??1???011?01??000000?1?000?
001001001?11??010?001100?01000000000000001001?110?0?1110011110001010?01011?111?000?0020000002100010010?
?1100121002110100021021000111100110000020000011101011100011111001010?111??0?01???2011010000011101100????
??120010?0?00000000000???00
Herrerasaurus_ischigualastensis
20010?00100101001000000000000?0000100000?10100000000030010001?1?012111000000?1??1???001101100??00000?10
0?000000001111000?01001100101??0020000000001011?11?????111011010000011201100110130101002000000210001001
110110112000201010102102100110110011?0000200100101010111001111110010101111??0?01???2011110000011101100??
?????0001301?0101001000001[01]00
Staurikosaurus_pricei
????????????????????????????????????????????????????????????????????????????????????????????00000?00??00?0000000
01?1???0????01?00?????000000000?00101???????????????????????????????????????002000000210001001110110111?00?0
11?000210110011111001??00002001001000101110001?????????????????????????????????????????0????????????????0??00
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Chindesaurus_bryansmalli
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00?0?
Eoraptor_lunensis
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?000?00?0101???00??001?0??0?0000200000?000?0?1?110?0?11110110100010111??1001110??1010020000112?00110011??
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Tawa_hallae
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10?101?1???2??12000220110001210001110000200110100010111000111110010101111??0?01???2111110000010101200???
???1000?3011?0010110????1?00
Coelophysis_bauri
1110111110111?0020000101100001?010111000010100100001130010?0101001211110000111???00?0011011000?00000?10
0?0000000011111001111111110010000001000111010111111?0?11110110100010112111001111301011121110212111110011
1011?11111021?12000210210011210011110000210111111010111011111010010100111??1?01???21111110100101012?0????
0??1212210210001201?0000011
Coelophysis_rhodesiensis
1110111111111000200001111000010010111000010100100001?3001000111?012111100001111?1???0011011000020000??00
000000000111110?1?1111111?010000001100111010011111010111101101000101121110011113010211211102121111100111
010011111121012000210210011210011110000210111111010111011111010010100111??1?01???2111111010010101200????
???111201021?00120101100111
Panguraptor_lufengensis
??????????????00?0000??1101?0?0???1110???10?0?100001130010?0????????????????????????001?01?0???????0?1001?00
000?0??11?0?1?11?1111?0?00??00??00???????1??11??????????0??????11???????111?0?0????????????111?????????????1??
???1??0?2?021?01??????????0??21011?11?0101110??1?10100???0??11?????1???211111??1?0?0?0120????????1212011000?
??10?0???????
'Syntarsus'_kayentakatae
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?1?111?21?120002?0210011210011110000210111111010110?111?101001010?111??1??1????1111110?00101???0???????102
0320100011?????0000011
Camposaurus_arizonensis
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Liliensternus_liliensterni
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FIRST COEL OPHYSOID DINOSAUR FROM ASIA
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??21?020002101200012100011?00002101111110101110001?????010100111??1??1???21111???000101???0?????????2??201
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Zupaysaurus_rougieri
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Dilophosaurus_wetherelli
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0000000011110000111111200010?0100000011101??1?111?1011110010100010112??10011112010111211002121011100?110
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Cryolophosaurus_ellioti
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Ceratosaurus_nasicornis
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Piatnitzkysaurus_floresi
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Allosaurus_fragilis
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01000000001111011101011120001011100000011111111110101011110110100010112111101111?1??10121100202101110011
0111021111121?02000220020001010001110000210111211010111010111010021100111??0?01???2111011000110101200???
???1020330110001001010000000
Velociraptor_mongoliensis
01001011100100101110000010100100111111100101001000000310000011100121?1001111111?????0011011000000000?10
0100000000111??11???????????10?01?01001111110?11101?1111100110100000012111000111?1??101211002021011110111
011?20010020102010220?200010100011100002101?12110?01?0010?11???02110?111??1?01????111011000010101200????
??10103110100?00??000000000
... Most dinosaur body fossils discovered in Yunnan are referable to sauropodomorphs (Young, 1941;Young, 1942;Young, 1947;Simmons, 1965;Yang, 1982;Chao, 1985;Bai, Yang & Wang, 1990;Dong, 1992;Zhang & Yang, 1995;Fang et al., 2000;Fang et al., 2004;Lü et al., 2006;Lü et al., 2007a;Lü et al., 2008;Upchurch et al., 2007;Sekiya, 2010;Xing et al., 2015;Wang et al., 2017;Zhang et al., 2018). At the same time the ornithischian and theropod records are relatively patchy (Young, 1948;Simmons, 1965;Xu, Zhao & Clark, 2001;Irmis, 2004;You et al., 2014;Wang et al., 2017). Yet, due to different preservation conditions, footprints and skeletons are often not preserved together or in the same proportions (Thulborn, 1990). ...
... Thus, the gap left by body fossils is often filled by trace fossils, which may provide an indicator of the diversity and distribution of a particular taxon in a given area. This is precisely the case for theropods in Yunnan Province: hitherto, merely seven theropod body fossils have been discovered, including relatively complete specimens such as Panguraptor lufengensis (You et al., 2014) and Sinosaurus triassicus (Young, 1948). Meanwhile, multiple theropod ichnotaxa were described, including: Changpeipus (Young, 1960), Eubrontes (Hitchcock, 1845), Grallator (Hitchcock, 1858) and Kayentapus (Welles, 1971). ...
... A total of six theropod species were unearthed from lower Jurassic deposits in Yunnan, namely: Eshanosaurus deguchiianus (Xu, Zhao & Clark, 2001), Lukousaurus yini (Young, 1940), which might be a crurotarsan (see Irmis, 2004), Megapnosaurus sp. (Irmis, 2004), Panguraptor lufengensis (You et al., 2014), Shuangbaisaurus anlongbaoensis (Wang et al., 2017) and Sinosaurus triassicus (formerly Dilophosaurus sinensis) (Young, 1948;Hu, 1993). In the Fengjiahe Formation where the Xiyang track site lies, three theropod genera are known: Sinosaurus, Eshanosaurus, and Shuangbaisaurus (Hu, 1993;Xu, Zhao & Clark, 2001;Wang et al., 2017). ...
Article
Full-text available
Yunnan Province is famous for its diversified Lufeng vertebrate faunas containing many saurischian dinosaur remains. In addition to the body fossil record, dinosaur ichnofossils have also been discovered in Yunnan, and the number of published track sites is on the rise. We report a theropod assemblage from the Lower Jurassic Fengjiahe Formation in Xiyang, central Yunnan. It is the third report and description of dinosaur footprints from the Fengjiahe Formation, and this new track site is the largest in number of footprints for theropods in Yunnan. Over one hundred footprints are preserved on different layers of a claystone-dominated succession close to the Lower-Middle Jurassic boundary. The track area is referred to as a lacustrine shallow-water paleoenvironment. Tracks vary in size, morphology, and preservation. All are tridactyl and digitigrade, and some are identified as undertracks. The best preserved footprints were divided into three morphotypes: morphotype A (>8 cm–27 cm–
... By contrast, Eo. murphi and other early neotheropods (e.g. D. wetherilli [64]; G. quayi [61]; Panguraptor lufengensis [67]) have proportionally taller middleposterior dorsal neural spines (ratio > 0.5). The anterior margin of the neural spine is located at the level of the parapophysis (figure 4a,b). ...
... Pendraig milnerae was incorporated into the data matrix of Novas et al. [92], which represents the most recent iteration of the matrix originally published by Nesbitt et al. [89], which has also been modified in various other studies [58,67,84,[93][94][95][96][97][98][99][100]. The Nexus and TNT files of the matrix, as well as a list of modifications made to the scoring of various characters for certain taxa and the script to calculate the consistency and retention indices (CI and RI, respectively), are included as electronic supplementary material. ...
... Open Sci. 8: 210915 [58,67,100]. Lepidus praecisio was previously considered as a coelophysid [58,95,99,100,110], but was found outside Coelophysoidea by Marsh et al. [96], in which it was found as the sister taxon to all other non-coelophysoid neotheropods, and by Marsh & Rowe [64], in which it was found as the sister taxon to L. liliensterni in a clade that is part of a polytomy at the base of Neotheropoda. ...
Article
Full-text available
We describe a new small-bodied coelophysoid theropod dinosaur, Pendraig milnerae gen. et sp. nov, from the Late Triassic fissure fill deposits of Pant-y-ffynnon in southern Wales. The species is represented by the holotype, consisting of an articulated pelvic girdle, sacrum and posterior dorsal vertebrae, and an associated left femur, and by two referred specimens, comprising an isolated dorsal vertebra and a partial left ischium. Our phylogenetic analysis recovers P. milnerae as a non-coelophysid coelophysoid theropod, representing the first-named unambiguous theropod from the Triassic of the UK. Recently, it has been suggested that Pant-y-ffynnon and other nearby Late Triassic to Early Jurassic fissure fill faunas might have been subjected to insular dwarfism. To test this hypothesis for P. milnerae , we performed an ancestral state reconstruction analysis of body size in early neotheropods. Although our results indicate that a reduced body size is autapomorphic for P. milnerae , some other coelophysoid taxa show a similar size reduction, and there is, therefore, ambiguous evidence to indicate that this species was subjected to dwarfism. Our analyses further indicate that, in contrast with averostran-line neotheropods, which increased in body size during the Triassic, coelophysoids underwent a small body size decrease early in their evolution.
... However, in the phylogenetic analyses of Maidment (2010), Raven and Maidment (2017) and Norman (2021), Scutellosaurus, Emausaurus and Scelidosaurus are regarded as successive basal thyreophorans (see also Thompson et al., 2012), so Family Scelidosauridae would then be a paraphyletic group. The first genus of Neornithischia, Stormbergia, appeared during the Hettangian-Sinemurian in South Gondwana (South Africa and Lesotho; Butler, 2005 (Yates, 2006;You et al., 2014;Wang et al., 2017b). The dilophosaurids occupied the top of the trophic chain, with Dracovenator and Dilophosaurus (5-6.5 m and 270-390 kg; Therrien & Henderson, 2007;Reolid et al., 2021b) being among the largest theropods in the Early Jurassic. ...
Article
Full-text available
The early Toarcian Jenkyns Event (~183 Ma) was characterized by a perturbation of the global carbon cycle, global warming, which at continental areas led to intensifi ed chemical weathering, enhanced soils erosion, and intensifi ed wildfi res. Warming and acid rain aff ected diversity and composition of land plant assemblages, caused a loss of forests and thereby impacted on trophic webs. The Jenkyns Event, triggered by volcanic activity of the Karoo-Ferrar Large Igneous Province, changed terrestrial ecosystems, and also aff ected the dinosaurs. Fossil macroplant assemblages and palynological data reveal reductions in the diversity and richness of plant communities. A substantial loss of land plant biomass and a shift to forests dominated by Cheiropelidiaceae conifers occurred as a consequence of seasonally dry and warm conditions. Major changes occurred to hervivore dinosaurs, with extinction of diverse basal families of Sauropodomorpha ('prosauropods') as well as some basal sauropods. Ornithischian dinosaurs show patchy records; some heterodontosaurids disappeared and the scelidosaurids (Thyreophora) went extinct during the Jenkyns Event. The dominant carnivorous dinosaurs, the Coelophysoidea (Theropoda), died out during the Jenkyns Event. We interpret the Jenkyns Event as a terrestrial crisis for ecosystems, marked especially by fl oral changes and the extinction of some dinosaur clades, both hervivores and carnivores. Resumen: El Evento Jenkyns del Toarciense inferior (~183 Ma) se caracterizó en ambientes continentales por una perturbación del ciclo del carbono, un calentamiento global, un aumento de la meteorización, la pérdida de suelos y la proliferación de incendios. El calentamiento y la lluvia ácida afectaron a la diversidad y composición de las asociaciones vegetales, causó la pérdida de masas forestales y tuvo un fuerte impacto en las redes trófi cas. El Evento Jenkyns, cuyo detonante fue la intensa actividad volcánica de la Provincia Ígnea de Karoo-Ferrar, cambió los ecosistemas continentales, afectando entre otros a los dinosaurios. Los datos palinológicos y de las asociaciones fósiles de macroplantas muestran una reducción de la diversidad y la riqueza de las comunidades vegetales, especialmente una pérdida de biomasa y la dominancia de coníferas cheirolepidiáceas en los bosques, en un contexto de condiciones cálidas estacionalmente áridas. Pueden observarse cambios importantes entre los dinosaurios herbívoros con la extinción de varias familias basales de sauropodomorfos ("prosaurópodos") y algunos saurópodos basales. Los dinosaurios ornitisquios, pese a su registro más incompleto, muestran la desaparición de algunas especies de heterodontosáuridos y la extinción de la familia Scelidosauridae (Thyreophora) en relación con el Evento Jenkyns. Los dinosaurios carnívoros de la superfamilia Coelophysoidea (Theropoda) también se extinguieron durante el Evento Jenkyns. Por lo tanto, se interpreta que el Evento Jenkyns contituyó una crisis biótica en los ecosistemas continentales de gran importancia, marcada especialmente por cambios en la fl ora y la extinción de algunos grupos de dinosaurios tanto herbívoros como carnívoros.
... As for sauropodomorphs, theropods extended during the Early Jurassic. Both families, Coelophysidae and Dilophosauridae colonized Asia (Panguraptor, Shuangbaisaurus) and Africa (Megapnosaurus, Dracovenator) during the Hettangian and Sinemurian (Yates, 2006;You et al., 2014;Wang et al., 2017b) (Figs. 5-7). ...
Article
Full-text available
The Early Jurassic Jenkyns Event (~183 Ma) was characterized in terrestrial environments by global warming, perturbation of the carbon cycle, enhanced weathering and wildfires. Heating and acid rain on land caused a loss of forests and affected diversity and composition of land plant assemblages and the rest of the trophic web. We suggest that the Jenkyns Event, triggered by the activity of the Karoo-Ferrar Large Igneous Province, was pivotal in remodelling terrestrial ecosystems, including plants and dinosaurs. Macroplant assemblages and palynological data show reductions in diversity and richness of conifers, cycadophytes, ginkgophytes, bennetitaleans, and ferns, and continuation of seasonally dry and warm conditions. Major changes occurred to sauropodomorph dinosaurs, with extinction of diverse basal families formerly called ‘prosauropods’ as well as some basal sauropods, and diversification of the derived Eusauropoda in the Toarcian in South America, Africa, and Asia, and wider diversification of new families, including Mamenchisauridae, Cetiosauridae and Neosauropoda (Dicraeosauridae and Macronaria) in the Middle Jurassic, showing massive increase in size and diversification of feeding modes. Ornithischian dinosaurs show patchy records; some heterodontosaurids and scelidosaurids disappeared, and major new clades (Stegosauridae, Ankylosauridae, Nodosauridae) emerged soon after the Jenkyns Event, in the Bajocian and Bathonian worldwide. Among theropod dinosaurs, Coelophysidae and Dilophosauridae died out during the Jenkyns Event and a diversification of theropods (Megalosauroidea, Allosauroidea, Tyrannosauroidea) occurred after this event with substantial increases in size. We suggest then that the Jenkyns Event terrestrial crisis was marked especially by floral changes and origins of major new sauropodomorph and theropod clades, characterized by increasing body size. Comparison with the end Triassic Mass Extinction helps to understand the incidence of climatic changes driven by activity of large igneous provinces on land ecosystems and their great impacts on early dinosaur evolution.
... There are two known theropods based on skeletal remains from the Lufeng area and its vicinity: the neotheropod Sinosaurus (Young, 1948, Hu, 1993Xing, 2012) and the coelophysid Panguraptor (You et al., 2014). Shuangbaisaurus described by Wang et al. (2017), was thought to fall within the range of specimens assigned to Sinosaurus triassicus (Currie et al., 2019). ...
Article
Full-text available
An increasing number of theropod-dominated tracksites have been reported from the Jurassic and Cretaceous of China. These include a significant number from the Lower Jurassic of the Lufeng Basin, famous for its Lufengosaurus fauna and known for a typical Lower Jurassic globally-distributed tetrapod footprint biochron. Here we report another localized theropod track occurrence regular of various scattered tracksites from the Lufeng Formation. The tracks are medium-sized tridactyl tracks from the basal member of the Zhangjia'ao Member, Lufeng Formation which shows an unusually wide divarication between the traces of digits III and IV, which suggest several possible interpretations.
... Young 1941aYoung , 1941bYe 1975;Zhao 1985;Dong 1992;Zhang and Yang 1995;Fang et al. 2000Fang et al. , 2004Fu and Zhang 2004;Lü et al. 2006Lü et al. , 2007Lü et al. , 2010Upchurch et al. 2007;Sekiya and Dong 2010;Sekiya 2010Sekiya , 2011Wang et al. 2017Wang et al. , 2019aZhang et al. 2018aZhang et al. , 2020Ren et al. 2020;Peyre De Fabrègues et al. 2020. Especially, the Early Jurassic sedimentary unit (LFF) in Lufeng Basin, which is widely recognised for the 'Lufeng Saurischian Fauna' (Young 1940(Young , 1941a(Young , 1942(Young , 1944(Young , 1947a(Young , 1947b(Young , 1948(Young , 1951(Young , 1974(Young , 1978(Young , 1982Chow and Hou 1959;Chow 1962;Rigney 1963;Simmons 1965;Ye 1973;Cui 1976Cui , 1981Carroll and Galton 1977;Zhang and Cui 1983;Sun et al. 1985;Wu 1986;Luo and Sun 1993;Wu 1994, 1995;Zhang and Yang 1995;Irmis 2004;Reisz et al. 2013;You et al. 2014;Wang et al. 2017Wang et al. , 2019aPeyre De Fabrègues et al. 2021). These units represent the major fossiliferous Early Jurassic 'Red Beds' exposed in China (Dong et al. 1983;Fang et al. 2008;Lü et al. 2010;Sekiya 2010;Wang et al. 2017Wang et al. , 2019aZhang et al. 2018aZhang et al. , 2020Xing et al. 2018;Peyre De Fabrègues et al. 2021). ...
Article
The age of Yunnanosaurus youngi, a non-sauropod basal sauropodomorph, was reported as the Middle Jurassic. However, no basal sauropodomorphs have been reported worldwide in the Middle Jurassic previously, and only one clade of sauropodomorphs, Sauropoda, existed in the Middle Jurassic and became extinct by the end of the Cretaceous. Here, we further investigated the locality and its horizon of Yunnanosaurus youngi in Jiangyi, Yuanmou County, Yunnan Province, China. We found the species horizon is actually from the Lower Jurassic Fengjiahe Formation, and its age is probably Pliensbachian. Furthermore, the three known Yunnanosaurus-bearing horizons in Chuxiong, Yunnan can be correlated lithostratigraphically. Abbreviations FJHF: Fengjiahe Formation; LFF: Lufeng Formation; P: profile; TJB: Triassic-Jurassic boundary; YNOQ: quarry of Yunnanosaurus youngi; ZHF: Zhanghe Formation.
... The specimen was molded with silicone rubber and cast with urethane casting resin; the actual specimen is currently on exhibit at the SGDS. Comparisons to some specimens below were made from the literature, specifically Lucianovenator bonoi (Martínez and Apaldetti, 2017), Panguraptor lufengensis (You et al., 2014), Notatesseraeraptor frickensis (Zahner and Brinkmann, 2019), Piatnitzkysaurus floresi (Bonaparte, 1986), Allosaurus fragilis (Madsen, 1976), Majungasaurus crenatissimus (O'Connor, 2007), Scelidosaurus harrisonii (Norman, 2020), and Heterodontosaurus tucki (Santa Luca, 1980). Comparisons to Eoraptor lunensis and Herrerasaurus ischigualastensis, were made from high-resolution photographs and casts housed at the TMM. ...
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... Aus der Toqui-Formation sind einige bisher noch undiagnostische Reste von Theropoden und Sauropoden (salgado et al. 2008Sauropoden (salgado et al. , 2015 Lufeng-Formation in China (z. B. luo et al. 2001a;lü et al. 2010;you et al. 2014), aber unterjurassische terrestrische Wirbeltiere sind auch aus zahlreichen anderen Lokalitäten bekannt (siehe z. B. Rauhut & lópez-aRBaRello 2008). ...
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Triassic beds from Argentina and Brazil provide the most relevant fossil record of early dinosauriforms in terms of numerical abundance and taxonomic diversity. This record currently represents the best source to understand the origin and early evolutionary radiation of dinosaurs. In the present paper we offer an updated review focused on the available evidence of Carnian dinosaurs from this continent, but we also discuss the record of Triassic dinosaur precursors and the evolution of Triassic dinosaurs in other continents. It is clear that, aside the agreed taxonomic composition of some particular dinosaurian subclades (e.g., Herrerasauridae, Neotheropoda), there is no consensus about early dinosaur phylogeny, and our paper is not the exception. Recent years witnessed the discovery of several new early dinosaurian taxa, as well as reviews of the taxonomic allocation of several renowned forms such as Lagerpeton, Lewisuchus, Pisanosaurus, and Eoraptor. New analyses demonstrate that evidence supporting the taxonomic referrals of pre-Norian dinosaurs to Theropoda, Sauropodomorpha and Ornithischia are tenuous, at best. Here we present new anatomical observations and comparisons for each of these South American early dinosauriforms with the aim to test previous phylogenetic interpretations. Evidence from South America allows reviewing the phylogenetic relationships of taxa from other continents, including Tawa, Chindesaurus, and Daemonosaurus, which are here suggested to nest within Herrerasauria. Evidence at hand indicates that herrerasaurs were a successful clade of archaic predatory saurischians that inhabited both South and North America, and probably also India and Europe.
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Not enough room Modern carnivore communities include species that span a range of body sizes. For example, on the African savannah, there are small species (mongooses), medium species (wild dogs), and large species (lions). This variation reflects available prey sources that best suit each group. Carnivorous dinosaur communities, however, were missing species that fall into the middle, or mesocarnivore, group as adults. Schroeder et al. looked across communities, space, and time and found that this absence appears to have been driven by the distinctive biology of dinosaurs, in which giant adults start out as tiny hatchlings. Growing juvenile dinosaurs thus filled the other niches and limited trophic species diversity. Science , this issue p. 941
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Several authors have drawn attention to the close similarities between the neotheropod dinosaurs Coelophyis and Syntarsus. Reconstruction and analysis of a skull from a juvenile specimen of Syntarsus (collected from the Forest Sandstone Formation of Zimbabwe) show that cranial characters previously used to distinguish these taxa and justify their generic separation (namely the presence of a 'nasal fenestra' in Syntarsus and the length of its antorbital fenestra), were based on erroneous reconstructions of disassociated cranial elements. On the basis of this reinterpretation we conclude that Syntarsus is a junior synonym of Coelophysis. Variations are noted in three cranial characters - the length of the maxillary tooth row, the width of the base of the lachrymal and the shape of the antorbital maxillary fossa - that taken together with the chronological and geographical separation of the two taxa justify separation at species level.