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High diversity of the Ganzhou Oviraptorid Fauna increased by a new “cassowary-like” crested species


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A new oviraptorid dinosaur from the Late Cretaceous of Ganzhou, bringing oviraptrotid diversity of this region to seven taxa, is described. It is characterized by a distinct cassowary-like crest on the skull, no pleurocoels on the centra from the second through fourth cervical vertebrae, a neck twice as long as the dorsal vertebral column and slightly longer than the forelimb (including the manus). Phylogenetic analysis recovers the new oviraptorid taxon, Corythoraptor jacobsi, as closely related to Huanansaurus from Ganzhou. Osteochronology suggests that the type specimen of Corythoraptor had not reached stationary growth stage but died while decreasing growth rates. The histology implies that it would correspond to an immature individual approximately eight years old. We hypothesize, based on the inner structure compared to that in modern cassowaries, that the prominent casque of Corythoraptor was a multifunction-structure utilized in display, communication and probably expression of the fitness during mating seasons.
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SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
High diversity of the Ganzhou
Oviraptorid Fauna increased by
a new “cassowary-like” crested
Junchang Lü1,2, Guoqing Li3, Martin Kundrát4, Yuong-Nam Lee5, Zhenyuan Sun6, Yoshitsugu
Kobayashi7, Caizhi Shen1,2, Fangfang Teng8 & Hanfeng Liu3
A new oviraptorid dinosaur from the Late Cretaceous of Ganzhou, bringing oviraptrotid diversity of this
region to seven taxa, is described. It is characterized by a distinct cassowary-like crest on the skull, no
pleurocoels on the centra from the second through fourth cervical vertebrae, a neck twice as long as
the dorsal vertebral column and slightly longer than the forelimb (including the manus). Phylogenetic
analysis recovers the new oviraptorid taxon, Corythoraptor jacobsi, as closely related to Huanansaurus
from Ganzhou. Osteochronology suggests that the type specimen of Corythoraptor had not reached
stationary growth stage but died while decreasing growth rates. The histology implies that it would
correspond to an immature individual approximately eight years old. We hypothesize, based on the
inner structure compared to that in modern cassowaries, that the prominent casque of Corythoraptor
was a multifunction-structure utilized in display, communication and probably expression of the tness
during mating seasons.
Oviraptorosaurs, a well-defined group of coelurosaurian dinosaurs, are characterized by short, deep skulls
with toothless jaws (teeth are present in primitive forms such as Incisivosaurus and Caudipteryx), pneuma-
tized caudal vertebrae, anteriorly concave pubic shas, and posteriorly curved ischia13. In recent years, diverse
oviraptorid-like eggs (and clutches) as well as oviraptorid skeletons that have been unearthed from the Upper
Cretaceous deposits of Ganzhou, Jiangxi Province, southern China, have made the Ganzhou area one of the most
productive oviraptorosaurian regions of the world. At present, six oviraptorosaurian dinosaurs have been named
from Ganzhou, including Banji long4, Jiangxisaurus ganzhouensis5, Nankangia jiangxiensis6, Ganzhousaurus nan-
kangensis7, Huanansaurus ganzhouensis8, and Tongtianlong limosus9. All taxa are from the Upper Cretaceous
Nanxiong Formation, comprising a distinct “Ganzhou Dinosaurian Fauna8. Here we describe a new ovirap-
torid dinosaur Corythoraptor jacobsi gen. et sp. nov. from the Nanxiong Formation beds exposed near the
Ganzhou Railway Station, Ganzhou City, Jiangxi Province. Corythoraptor jacobsi gen. et sp. nov. bears a distinct
cassowary-like crest (helmet) and has a long neck, which exhibits convergent morphology to the modern ight-
less cassowary bird from Queensland in Australia. e discovery of Corythoraptor jacobsi provides unprecedented
evidence that oviraptorid dinosaurs were morphologically and taxonomically far more diverse in the Ganzhou
area than in any other known region of the world.
Systematic palaeontology.
Oviraptorosauria Barsbold, 1976.
1Institute of Geology, Chinese Academy of Geological Sciences, Beijing, 100037, China. 2Key Lab of Stratigraphy
and Paleontology, Ministry of Land and Resources of China, Beijing, 100037, China. 3Jiangxi College of Applied
Technology, Ganzhou, 341000, Jiangxi Province, China. 4Center for Interdisciplinary Biosciences, Faculty of Science,
University of Pavol Jozef Safarik, Kosice, 04154, Slovak Republic. 5School of Earth and Environmental Sciences,
Seoul National University, Seoul, 08826, South Korea. 6Jinzhou Paleontological Museum, Jinzhou, 121000, Liaoning
Province, China. 7Hokkaido University Museum, Hokkaido University, Sapporo, Hokkaido, 060-0810, Japan. 8Xinghai
Paleontological Museum of Dalian, Dalian, 116000, Liaoning Province, China. Correspondence and requests for
materials should be addressed to J.L. (email:
Received: 1 March 2017
Accepted: 6 June 2017
Published: xx xx xxxx
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
Oviraptoridae Barsbold, 1976.
Corythoraptor jacobsi gen. et sp. nov.
Etymology. e generic name Corythoraptor refers to a raptor bearing a “cassowary-like crest” on its head,
and the specic name is in honor of Professor Louis L. Jacobs, who has contributed to dinosaur research and
has given excellent mentoring to three authors (JLü, YL and YK) when they were Ph.D. students at Southern
Methodist University, Dallas, Texas, USA.
Holotype. Almost complete skeleton with the skull and lower jaw (JPM-2015-001) (Figs1 and 2) is housed at
the Jinzhou Paleontological Museum, Jinzhou, Liaoning Province, China.
Type locality and horizon. A site in the vicinity of the Ganzhou Railway Station (GPS coordinates are pro-
vided on request from the rst author), Ganzhou City; Campanian-Maastrichtian; Nanxiong Formation (Upper
Diagnosis. An oviraptorosaurian dinosaur with the following unique combination of characters: ratio of the
length of the tomial margin of the premaxilla to the premaxilla height (ventral to the external naris) is 1.0–1.4;
inclination of the anteroventral margin of the premaxilla relative to the horizontally positioned ventral mar-
gin of the jugal posterodorsal; antorbital fossa bordered anteriorly by the maxilla; narial opening much longer
Figure 1. e holotype of Corythoraptor jacobsi gen. et sp. nov. (JPM-2015-001). (a) Photograph. (b) Outline
drawings. (c) Close up of the skull and lower jaw, showing the pneumatic cassowary-like crest (Only skull and
lower jaw elements are labeled). (d) Skeletal reconstruction (missing parts are in grey). Abbreviations: aof,
antorbital fenestra; cav. caudal vertebrae; cr. cervical ribs; cv. cervical vertebrae; dv. dorsal vertebrae; fe, femur;
. bula; h, humerus; il, ilium; is, ischium; l, lacrimal; lj, lower jaw; ltf, lower temporal fenestra; m, maxilla; n,
nasal; nar, narial opening; o, orbit; oc, occipital condyle; p, parietal; pm, premaxilla; po, postorbital; ps, pes; psc,
pneumatic skull crest; pu. pubis; q, quadrate; ra, radius; sk, skull; sq, squamosal; stf, super temporal fenestra; ti,
tibia; ul, ulna. Scale bar = 8 cm in (c) and 100 cm in (d).
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
Figure 2. e cranial casque of Corythoraptor jacobsi and recent cassowaries. (ac) the crested skull of Corythoraptor
and head appearance restorations. (d) a close-up (see dotted rectangle in a and b) of eroded bony shell in the
posterolateral casque of Corythoraptor. (e) the crested skull of the recent cassowary (Casuarius uniappendiculatus;
Museum für Naturkunde in Berlin, Germany: MfN-ZMB 93274). (f) a keratinous helmet over the skull of the recent
cassowary (unnumbered specimen of Casuarius casuarius from the osteological collections of ZOO Protivín, Czech
Republic). (g,h) coronal cuts through the cassowary skull – (g) Casuarius casuarius MfN-ZMB 36820, (h) Casuarius
casuarius: MfN-ZMB 36885 (see dotted lines in f); note transition in strut-like trabecular arrangement. (i) close-up
to contact between keratinous and skeletal components of the casque in recent cassowary, unnumbered specimen of
Casuarius sp. from the osteological collections of Field Museum of Natural History, Chicago, USA. Abbreviations: cq,
casque; cr, cranium; exs, external surface; kesh, keratinous sheath; or, orbit; tr, trabeculae.
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
than width; infratemporal fenestra dorsoventrally elongate, narrow anteroposteriorly; the supranarial process of
the premaxilla bears two processes: a short posterodorsally extending process, forming the anterodorsal margin
of the external nasal opening, and a long process, forming most of the anterodorsal process of the premaxilla;
distinct cassowary-like helmet on the skull; long axis of the external narial opening parallel to the dorsal mar-
gin of antorbital fenestra; straight anterodorsal margin of dentary in lateral view; a deep fossa, sometimes with
associated pneumatopore on lateral surface of dentary; no pleurocoels on the centra from the second through
fourth cervical vertebrae; the length of the neck twice as long as the dorsal vertebral column, and slightly longer
than the entire forelimb length (including the manus); less pronounced deltopectoral crest of humerus, forming
an arc rather than being quadrangular; ratio of the length of the manus to the length of the humerus plus the
radius between 0.50 and 0.65; the ungual of digit III less curved than other unguals; lesser trochanter (cranial
trochanter) completely fused with the greater trochanter and distal ends of shas of metatarsal II straight and
metatarsal IV laterally deected.
Corythoraptor jacobsi gen. et sp. nov. is assigned to oviraptorid dinosaurs based on the following characters:
proximal caudals with pneumatized centra; ischium with its posterior prole concave3; premaxilla pneumatized;
the subantorbital portion of the maxilla inset medially; the palate extending below the cheek margin; the external
naris overlapping most of the antorbital fossa rostrodorsally; the bones of the skull roof pneumatized; the pubic
sha concave cranially, the mandibular symphysis tightly sutured; the shortened preorbital region, and the tooth-
less jaws2.
Corythoraptor jacobsi diers from all other oviraptorosaurs containing skulls, such as Incisivosaurus gauthieri11,
Caudipteryx zoui12, Conchoraptor gracilis13, Wulatelong gobiensis14, Banji long4, Khaan mckennai15, Citipati osmol-
skae15, 16, Huanansaurus ganzhouensis8, Yulong mini17, Oviraptor philoceratops16, 18, Nemegtomaia barsboldi19, 20,
Rinchenia mongoliensis21 ( = Oviraptor mongoliensis13) and Heyuannia huangi22, in that Corythoraptor jacobsi
bears a cassowary-like, very thin highly pneumatic cranial crest, the highest point of the crest projecting above
the orbit, and an elongated narial opening. Although Banji long, Citipati osmolskae, Oviraptor philoceratops,
Nemegtomaia barsboldi and Rinchenia mongoliensis also bear crests, their morphology and positions at the skull
are quite dierent from that of Corythoraptor.
Corythoraptor jacobsi diers from Heyuannia huangi22 in that Heyuannia huangi has no skull crest, the pneu-
matic foramina on the neural arches and ribs of cervical vertebrae, no infradiapophyseal fossa in the anterior cau-
dal vertebrae, the strongly reduced metacarpal III, and the length ratio 1.25 of tibia to femur, whilst Corythoraptor
jacobsi does not have.
Corythoraptor jacobsi diers from Shixinggia oblita23 in that Shixinggia oblita has the preacetabular process
of the ilium distinctively shorter than the postacetabular process of the ilium, the distal end of the preacetabular
process higher than the dorsal margin of the acetabulum, the ischial peduncle almost the same depth with that of
the pubic peduncle, and a large opening in the medial surface near the proximal end of the femur.
Corythoraptor jacobsi diers from Jiangxisaurus ganzhouensis5, which has a weakly downturned mandibular
symphysis, pleurocoels in all cervical vertebrae except atlas, the radius-humerus length ratio of about 70%, and
slender metacarpal III.
Corythoraptor jacobsi diers from Ganzhousaurus nankangensis7, which has an acute angle formed by the
mandibular symphysis and the dorsal margin of dentary, and a relatively short and nearly straight rst pedal
Corythoraptor jacobsi mainly diers from Huanansaurus ganzhouensis8 in the skull morphology and the struc-
ture of the cervical vertebrae: Huanansaurus ganzhouensis has no skull crest, the anterior margin of the premax-
illa is nearly vertical to the ventral margin of the skull, the posterior margin of the lower temporal fenestra is
oblique, and the dorsal margin of dentary above the external mandibular fenestra is strongly concave ventrally.
Corythoraptor jacobsi diers from Tongtianlong limosus9 in the skull morphology. e anteroventral corner of
the external naris is far above a horizontal line tangent with the posterodorsal corner of the antorbital fenestra,
and the skull is dome-like in Tongtianlong. Whist the anteroventral corner of the external naris is below a horizon-
tal line tangent with the posterodorsal corner of the antorbital fenestra, and the skull bears a large cassowary-like
helmet in Corythoraptor.
Corythoraptor jacobsi also diers from Wulatelong gobiensis14 in that the anteroventral corner of the external
naris is far below a horizontal line tangent with the posterodorsal corner of the antorbital fenestra, whilst the
anteroventral corner of the external naris is slightly below a horizontal line tangent with the posterodorsal corner
of the antorbital fenestra in Corythoraptor jacobsi; the ischium/pubis ratio length is much smaller in Wulatelong
gobiensis (IS/PU = 0.45) than in Corythoraptor jacobsi (0.61) and the tibia/femur length ratio is 1.10 in Wulatelong
gobiensis, which is smaller than that in Corythoraptor jacobsi (1.19).
Corythoraptor jacobsi diers from Machairasaurus leptonychus24, which has elongate and blade-like manual
unguals I–III in lateral view, metacarpal I is proportionately short, about 41 per cent the length of metacarpal II,
which is shorter than that of Machairasaurus leptonychus (about 50% the length of metacarpal II; the combined
length of phalanges II-1 and II-2 is about 133% the length of metacarpal II in Corythoraptor jacobsi, which is
larger than that of Machairasaurus leptonychus24, where combined length of phalanges II-1 and II-2 does not
exceed 110% the length of metacarpal II.
Corythoraptor jacobsi diers from Rinchenia mongoliensis21 ( = Oviraptor mongoliensis13) in their skull mor-
phologies. the anterior margin of the premaxilla is strongly concave above the level at the postodorsal corner of
the antorbital fenestra and the highest point of the crest would project far above the orbit in Corythoraptor jacobsi,
whilst the anterior margin of the premaxilla is nearly straight and the highest point of the crest is above the orbit2.
ere is no pleurocoels on the centra from the second through fourth cervical vertebrae, the anterodorsal mar-
gin of ilium (above the acetabulum) is moderately convex and the pubic peduncle of the ilium is larger than the
the ischial peduncle in Corythoraptor jacobsi, pleurocoels are present on all the cervical centra, the anterodorsal
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
margin of ilium is strongly convex and both peduncles are equal in length in Rinchenia mongoliensis2(personal
Description. e skeleton including skull and lower jaws is almost complete except for the coracoid, and
middle-posterior caudal vertebrae (Fig.1, see also Supplementary Information TableS1). Most of the skull is pre-
served except for the quadratojugal, a portion of the skull roof and braincase. Some parts of the skull are overlain
by the right manus. e skull bears a high crest. Although the anterodorsal part of the crest is missing, its apparent
continuous outline implies that the highest point of the crest would project far above the orbit (Fig. 1b, 1c). e
internal structure of the crest is similar to the casque of Casuarius unappendiculatus. e dorsal part of crest is
very thin (about 3.5 mm, measured through its broken surfaces) and was lled with empty spaces. ese spaces
(pneumatic diverticula) are irregular in shape and dier in size, reminiscent of conditions seen in the modern
cassowary. e inner, bony core, casque of the cassowary25 is made by irregularly-arranged, slender bony struts
called trabeculae25. Corythoraptor jacobsi possessed an extensive cranial casque that was probably composed of
the skull roong bones: nasals, frontals and parietals (Fig.2a–c). Only a basal portion, likely a lower half, of the
bony core of the casque is preserved and exposed on the le side. e bony core is either obscured by sediment or
is exposed due to erosion of the external bony surface. e inner structure is best exposed on the postero-lateral
side of the casque. e inner structure consists of randomly branching, sparse, trabeculae of variable thickness
ranging from 0.3 (rod-like trabeculae) to 1.2 mm (lamellar trabeculae) (Fig.2d). ese delicate trabeculae outline
empty cavities, the largest exposed of which is 6.6 × 14.3 mm large, which implies that the inner core was light,
fragile (perhaps pliable), and hence, not suitable for percussive behavior including intraspecic combat. e larg-
est opening (called concavity by Naish and Perron25) is in the posterior part of the crest located above the external
narial opening (see Supporting Information Fig.S1). e crest extends posteriorly to reach the nuchal margin of
the skull. e orbit is circular and probably accommodated a relatively large eyeball. e lower temporal fenestra
is rectangular with its long axis vertical. e antorbital fenestra is triangular in lateral view, and larger than the
external narial opening. e narial opening is elongate and parallel to the long axis of the antorbital opening.
e premaxillae are completely fused, no sutural remnants are visible. e lower part of the premaxilla is con-
vex outwardly with an almost smooth surface sculptured with irregularly distributed pits. ese pits are probably
neurovascular foramina, and may indicate the presence of a keratinous sheath (such as rhamphotheca in modern
birds) over the premaxillary beak as in ornithomimid dinosaurs26, 27. e ventral margin of the premaxilla is sharp
and forms a crenulated tomial edge. e broken surface of the ventral margin of the premaxilla shows that the
bone consists of many chambers separated by thin bony struts, and is thus largely pneumatized and lightweight.
Anterior to the ventral corner of the external narial opening, the lateral surface of the premaxilla is moderately
concave. e supranarial (nasal) and subnarial (maxillary) processes of the premaxilla2 contact the nasal as in
Figure 3. Strict consensus of 4151738 most parsimonious trees obtained by TNT, based on analysis of
45 taxa and 257 characters, showing the phylogenetic position of Corythoraptor jacobsi gen. et sp. nov.
(Tree length = 623). Numbers adjacent to each node are Bremer support values. Oviraptors from southern
China are in red.
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
other derived oviraptorids. e subnarial process of the premaxilla is slightly convex with a smooth surface,
strap-like, and extends posterodorsally at about 40 degrees with the ventral margin of the skull. is process
forms the ventral margin of the external narial opening. Unlike other oviraptorids, the supranarial process of the
premaxilla bears two processes: a short posterodorsally extending process, forming the anterodorsal margin of
the external nasal opening, and a long process, forming most of the anterodorsal process of the premaxilla. ere
is a distinct ridge on the lateral surface near the anterior margin of the premaxilla. e external narial opening
is elongate and located above the antorbital fenestra. e long axis of the narial opening is parallel to the long
axis of the antorbital fenestra. In ventral view, the fused premaxillae are U-shaped, with three longitudinal ridges
on the ventral surface. e middle ridge is much thicker than the lateral and medial ones. Two deep grooves
are developed between the ridges. A large fossa is present on the ventral surface of the premaxillae near the
premaxillary-maxillary suture.
e short maxilla contacts the premaxilla anteriorly. e lateral margin of the maxilla is missing. e maxilla
bears a slender process closely appressed to the subnarial (maxillary) process of the premaxilla, which extends
posterodorsally. is process ends before reaching the posterodorsal corner of the antorbital fenestra. us, the
anterior corner of the antorbital fenestra is demarcated by the maxilla. ree small openings are present on the
lateral surface of the maxilla near the anteroventral corner of the antorbital fenestra. e one nearest the anterior
margin of the antorbital opening is a maxillary fenestra, and other two are pneumatopores as in other derived
oviraptorids such as Citipati and Conchoraptor2. e lateroventral surface of the maxilla is convex and smooth. It
extends ventrally toward the mid-line near the suture with the premaxilla, forming a distinct tooth-like process
as in other oviraptorids.
e anterior margin of the lacrimal is straight. e broken surface shows heavily pneumatized internal bone.
e ascending process of the jugal is curved posteriorly in lateral view. e ascending process of the jugal
occupies about two-thirds of the length of the postorbital bar.
e postorbital is T-shaped with anterior, posterior, and ventral processes. e posterior process extends pos-
teromedially and covers the anterior part of the squamosal. With the anterior process, the postorbital forms the
lateral margin of the supratemporal fenestra. e elongate supratemporal fenestra is much smaller than the orbit
and the infratemporal fenestra. e ventral process of the postorbital is relatively long, but it cannot be deter-
mined whether it reaches the posteroventral corner of the circular orbit due to its incomplete distal end. e pre-
served portion of the ventral process occupies at least two-thirds the length of the postorbital bar. In lateral view,
the posterior margin of the ventral process is straight and forms the anterior margin of the infratemporal fenestra.
e preserved portion of the nasals exhibits highly pneumatized bone structure. e suture between the pari-
etal and frontal is not clear, but it seems the bones project dorsally and formed a distinct crest together with the
e occipital condyle is preserved and extends posteroventrally. A large opening on the lateral surface on the
occipital condyle is probably for cranial nerve XII.
e mandible is toothless. e lateral surface of the dentary is slightly concave and covered with foramina.
e symphyseal suture is straight. e mandibular symphysis is U-shaped in dorsal view. e rostral end of the
mandibular symphysis is slightly downturned in lateral view. e external mandibular fenestra is large, longer
anteroposteriorly than dorsoventrally. e cranial margin of the fenestra is deeply incised and divides the dentary
into two long, shallow dorsal and ventral caudal processes. e angular and surangular bones are separated by a
gap and the posterior rim of the external mandibular fenestra. e dentary is W-shaped in ventral view because
of a distinct process at the ventral end of the mandibular symphysis with large concavities on each side. is con-
cavity may have accommodated the splenial.
e cervical series is complete, and almost naturally articulated. e circular curl of the neck is similar to
that of Heyuannia22, 28, which doesn’t seem like the typical death pose of theropods, although what caused this is
unclear. Including the atlas, there are twelve cervical vertebrae (Fig.1d). e sixth and eleventh cervical vertebrae
are the longest among the series. ere is no pleurocoel on the second through fourth cervical vertebrae, but one
is present in the h through twelh cervical vertebrae. e pleurocoel is nearly circular on the h cervical
vertebra (about 4.8 mm in diameter). It is oval in the sixth cervical vertebra (about 5 mm long). All pleurocoels
are located in the middle of the centra. e pleurocoels have sharp lower margins but their upper boundaries
are less distinct. e anterior articular surfaces of centra are strongly concave and the posterior articular surface
becomes moderately convex and oblique anteroventrally to posterodorsally. e anterior articular surfaces are
almost square and wider than the posterior articular surfaces. e widest part is situated between the parapo-
physes. e cervical ribs are fused with the vertebrae and bear distinct anterior processes. e weathered surface
of neural arch shows many small chambers separated by thin bony struts within the neural arch, which indicate
that the neural arches are densely pneumatized. e exposed neural spine is low and triangular in lateral view. A
centrodiapophyseal lamina is well-developed on the fourth to seventh cervical vertebrae. A distinct concavity lies
above the centrodiapophyseal lamina and below the postzygodiapophyseal lamina at their junction.
Only the rst six dorsal vertebrae are exposed, thus the total number of the dorsal vertebrae is not determined.
e dorsal vertebrae are shorter in length than the cervicals. e rst hypapophysis is a distinctive plate-like pro-
jection, which extends anteroventrally. e hypapophyseal body is anteriorly placed, but its posterior extension
reaches almost to the posterior edge of the centrum. Its distal end is rounded. Part of the second dorsal vertebra
is covered by a dorsal rib. e hypapophysis of the third dorsal vertebra is also distinct, similar to that of the rst
dorsal vertebra. e second and third dorsal vertebrae bear larger pleurocoels than those of cervical vertebrae.
e anterior surfaces of the dorsal vertebrae are slightly concave and their posterior articular surfaces are nearly
at.Only the posterior two sacral vertebrae are observable and they have a small pleurocoel. e rib of the last
sacral vertebra is stout and contacts the postacetabular process of the ilium medially. Both the ventral and lateral
surfaces of the sacral centra are smooth and round.
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
e anterior ve caudal vertebrae are preserved, and the rst three are complete. e rst caudal vertebra
abuts the posterior sacral vertebra. Both the anterior and posterior articular surfaces of caudal vertebrae are
nearly at. e anterior articular surface is circular in cranial view. e pleurocoel is small and elongate on the
rst and second caudal vertebrae. e pleurocoels of the third through h caudal vertebrae are somewhat larger,
but do not match the size of those of the dorsal vertebrae. e transverse process of the rst caudal vertebra is
long and projects posterolaterally. ere is no infraprezygapophyseal fossa on the rst caudal vertebra. ere are
three fossae on the other caudal vertebra as in Nankangia6: the largest fossa (called infraprezygapophyseal fossa)
is located on the anterior surface near the junction of the prezygapophysis and the transverse process; the second
fossa (called the infradiapophyseal fossa) is exposed ventral to the base of the transverse process; and the third
fossa is the pleurocoel. e neural arches of the caudal vertebrae are similar to those of Nankangia6. Only a small
proximal portion of haemal arch, which starts between the second and third caudal vertebrae, is preserved.
Only a thin strap-like distal end of the scapula is exposed. e very thin sternum is present, but not well
The left humerus is almost complete except for its broken proximal portion. The humerus is approxi-
mately 27% of the entire forelimb length including the manus and 48% of the forearm length without the hand
(Supplementary Information Table S1), and is weakly twisted as in other oviraptorids. e deltopectoral crest is
short and located at the proximal portion of the humerus (ratio of deltopectral crest to total length of humerus,
measured along the humeral sha, = 31%). e distal humerus is expanded and bears well-developed rounded
condyles. Both lateral condyles and the medial epicondyle are broken, but the former seems larger than the latter.
e lateral epicondyle is well developed. In anterior view, a distinct concave surface is visible between the lateral
condyle and epicondyle.
e ulna is slightly shorter than the humerus. It is short, approximately 26% of the length of the forelimb
including the manus, bowed and convex caudally with a poorly developed olecranon. e proximal end is more
expanded than its distal. e medial surface of the distal portion is nearly at.
e radius is rod-like and slightly shorter than the ulna. It is moderately curved cranially, thus leaving a space
between the ulna and radius as in Heyuannia22, 28. e radial sha exhibits a uniform width along its length, but is
much narrower than the ulna. e radius has an expanded distal end.
e proximal ends of metacarpals are closely appressed. e rst metacarpal is the shortest. It is slightly
stouter and about 41% the length of the second metacarpal. e ventral surface of the rst metacarpal is slightly
concave. Metacarpal II is moderately robust, with a sha diameter about 13% of total length. e second meta-
carpal has a circular sha. e distal articular surface of the second metacarpal has a well-developed trochlea.
e medial condyle is more developed and extends more medioventrally than the lateral condyle. In ventral view,
the shas of the second and third metacarpals are convex medially, thus leaving a gap between the distal ends of
the second and third metacarpals. e third metacarpal is narrower than the second; the former is about 68% of
the width of the latter. Its proximal portion is expanded dorsoventrally rather than mediolaterally. e middle
sha of metacarpal III is slender, the diameter being about 9% of the bone length. Metacarpal II and III are equal
in length.
e manual phalanges are long and robust. Manual phalanx I-1 is stout and the longest among the phalanges,
being about 72% of the length of the metacarpal II, and strongly constructed, with diameter about 15% of its
length. A large collateral ligament pit occurs laterally and medially.
e distal portion of le manual ungual I is missing. is laterally compressed ungual is the largest of the
manual unguals. e weakly curvedclaw has a simple, single groove on its lateral and medial surfaces, with the
medial groove being deeper. e exor tubercle is developed as a large, mound-like rugosity. e proximal artic-
ular surface extends dorsally and forms a well-developed tongue-like projection. Manual phalanges (MP) II-1 and
II-2 are long and moderately robust, each being about 67% and 66% of length of the metacarpal II, respectively.
e collateral ligament pits are strongly developed. e proximal articular surface of MP II-2 is concave with a
moderate-sized articular heel and tongue. e second manual ungual is smaller, proportionately more elongate,
and more weakly curved than the rst manual ungual. Its proximodorsal lip is similar to that of the rst manual
ungual, but the exor tubercle is smaller. Manual phalanges III-1 and III-2 are shorter than MP III-3. MP III-2
is the shortest among the phalanges. e third manual ungual resembles the ungual of digit II in shape, but is
slightly smaller than the unguals of digit I and digit II. e ungual of digit III is not as strongly curved as other
unguals. Its proximodorsal lip is developed, as in other unguals.
Both ilia are preserved, but the postacetabular process of the right ilium is incomplete. e postacetabular and
the preacetabular processes of the le ilium are missing their distal ends, but the impression of the preacetabular
process and the outline of the postacetabular process provide a precise shape and size of the ilium. e dorsal
margins of both iliac blades are close to each other medially. e distal ends of the postacetabular processes are
more greatly separated than the preacetabular processes. e preacetabular and postacetabular processes are
similar in length. e lateral surface of the postacetabular process is convex, and the middle portion of the ilium
above the acetabulum is slightly concave. A distinct brevis fossa is present on the ventral surface of the distal end
of the postacetabular process. e hook-like distal end of the preacetabular process extends below the level of
the dorsal margin of the acetabulum. e pubic peduncle is deeper than the ischial peduncle. A distinct process
occurs on the posterior margin near the base of the pubic peduncle. Anterior to the pubic peduncle, a shallow
elongate cuppedicus fossa is situated at the ventral surface of the preacetabular process. e brevis fossa, located
on the ventral surface of the distal end of the ischial peduncle, is shallow and short. A weak antitrochanter occurs
on the ischial peduncle. e pubic peduncle is deeper dorsoventrally and narrower anteroposteriorly than that of
the ischial peduncle.
Both pubes are well preserved and naturally articulated with ilium and ischium. As in most other derived
oviraptorids, the pubis is concave cranially, but diers from Nomingia, where the pubic sha is almost staight3.
e sha is mediolaterally compressed distally and becomes more rounded posteriorly near its middle portion
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
but becomes mediolaterally compressed again toward to the proximal end. A pubic apron extends medially from
the proximal end along the middle margin of the sha, although some parts are broken. It occupies the 53% the
whole length of the pubis. Distally, pubic aprons are completely fused to form a symphysis (pubic boot). e pubic
boot has a well-developed cranial process and a caudal process. e cranial process (7 cm long) is longer than
the caudal process (5 cm long), which is dierent from that of Nomingia, which bears an equally long processes3.
Both ischia are preserved, but are partially overlain by the right pes. As in other oviraptorids, the ischium has
a medially positioned triangular obturator process. e obturator process is located approximately midway along
the sha. Its lateral surface is concave with the sha mediolaterally compressed and concave caudally. e distal
ends of both ischia are not fused.
e femur is longer than the ilium as in oviraptorids and forms about 30% of the total hindlimb length. It
is straight in both lateral and posterior views. e femoral neck is directed dorsomedially at an angle of about
115° to the sha. A distinct short ridge-like structure occurs on the posterior surface of the femoral head. e
greater trochanter is massive and extends craniocaudally. It is separated from the femoral head by a slightly
constricted femoral neck. ere is no sign of lesser trochanter (cranial trochanter), which may be fused into the
greater trochanter. ere is no distinct fourth trochanter, but a muscular scar lies along the sha in the region
where a fourth trochanter would be expected to reside. is coarse surface (41 mm long and 15 mm wide) is
also present in Citipati osmolskae and Khaan mckennai29. e dorsal surface of the greater trochanter is 113 mm
from the middle of the coarse area. e distal end is of the femur broad with a nearly at cranial surface. On the
posterior surface, the distal femoral condyles are well separated, with the lateral condyle projecting well below the
medialone. A deep popliteal fossa is present between the two distal condyles. Caudally, a shallow fossa separates a
well-developed tibiobular crest from the large caudal surface of the medial condyle on the posterolateral surface
of the distal end of the femur. An extensive tibiobular crest projects posteriorly beyond the level of the medial
e tibia is 19% longer than the femur. In medial view, the dorsal margin of the proximal end is convex.
e pronounced cnemial crest is inected anteriorly and slightly medially. It constitutes about half of the entire
craniocaudal length of the proximal surface of the tibia. ere is no distinct boss on the distal end of the crest,
similar to the condition in Khaan mckennai29. e cnemial crest of the tibia is 52.9 mm long and it extends to
approximately the proximal 14% of the tibia, and ends at the proximal margin of the bular crest. e proximal
portion of the bular crest is ridge-like, extending posterolaterally, ending 130 mm from the dorsal margin of the
proximal end of the tibia. Lateral to the distal end of the bular crest, there is a small foramenwith a short dorsal
groove associated it. is foramen is likely a nutrient foramen, which is similar to that of Khaan mckennai (IGM
100/973)29. e distal tibiae are not well preserved but appear more attened anteroposteriorly. Distal ends of the
tibiae are not well-preserved.
e distal portion of the bula is missing. e proximal head of the bula expands anteroposteriorly. e
bular head is weakly concave medially as in most oviraptorosaurs2.
Only a part of the astragalus, which is articulated with the tibia, is preserved. e calcaneum is damaged, thus
detailed description of the astragalus and the calcaneum is not available.
e pedes are exposed ventrally. e foot accounts for approximately 29% of the length of the hindlimb. e
proximal portion of the le foot is missing. Two separate distal tarsals were found; the lateral distal tarsal IV is
larger than the medial distal tarsal III. e lateral tarsal covers the proximal articular surfaces of the metatarsal
II and III. e medial metatarsal covers the proximal articular surface of the metatarsal IV. Each tarsal fuses with
the corresponding metatarsal, which is similar to those of some caenagnathids (Elmisaurus, Leptorhynchos)30, 31.
Metatarsals I through V are preserved. e stout metatarsal III is longest. It is exposed along its entire length and
exhibits almost constant width. Metatarsal IV is slightly shorter than metatarsal III but longer than metatarsal II.
Metatarsal V is a thin splint of bone; its distal portion is missing. Deep ligament foveae are present on medial and
lateral surfaces of the distal ends of the metatarsals II-IV. e foveae are much larger and deeper on metatarsal III
than others. In ventral view, 56.6 mm away from the distal ends of metatarsals II-IV, a coarse area occurs on each
metatarsal. e proximal ends of metatarsals III and V expand mediolaterally and are wider than that the prox-
imal end of the metatarsal II, which expands anteroposteriorly. e distal ends of the metatarsals bear distinct
trochleae, with medial margins stronger than lateral. e distal ends of metatarsals II-IV are twisted medially; the
distal end of the metatarsal II is not prominent.
Corythoraptor jacobsi has the typical theropod phalangeal formula of 2-3-4-5. e le pes is preserved in a
natural pose, exposing its ventral surface. Digit 3 is longest. Digit 4 is longer than the digit 2. Digit 1 is shortest,
being 2/3 the length of the rst phalanx of the digit 2. All phalanges possess a deep ligament fovea on the lateral
and medial surfaces. ese pits are circular in shape. e phalangeal joints are symmetrical and ginglymoid.
e unguals are moderately curved and bear a shallow groove running along the medial and lateral surfaces; the
groove becomes shallower more distally. A long tongue-like dorsal lip occurs near the proximal end.
Phylogenetic analysis of Corythoraptor jacobsi. e strict consensus tree (Fig.3) shows all oviraptorid
dinosaurs from southern China are mainly distributed in three main clades of Oviraptoridae. Corythoraptor
jacobsi and Huanansaurus ganzhouensis form one clade, and they share the following seven synapomorphies:
1) posterodorsally inclined postorbital process of the jugal (character 37, state 0); 2) dentary, anterodorsal tip of
beak: projecting anterodorsally, tip of beak projecting at an angle of 45° or less relative to the ventral margin of the
symphysis (character 191, state 1); 3) pneumatized dentaries (character 199, state 1); 4) posteroventral branch of
dentary twisted so that lateral surface of branch faces somewhat ventrally (character 226, state 1); 5) development
of symphyseal shelf of mandible: length of symphysis (as measured on midline) greater than 20% but less than
25% length of mandible (character 229, state 1); 6) prominent proximodorsal extensor ‘lip’ on manual unguals
(‘set o’ from remainder of dorsal surface by distinct change in slope immediately distal to ‘lip’) (character 234,
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
state 1); and 7) external mandibular fenestra: anteriorly constricted by posteroventral ramus of dentary (character
254, state 1).
e strict consensus tree also recovers that oviraptorid dinosaurs from Ganzhou area are nested in three
subclades of Oviraptoridae, and better resolved than the tree obtained by Lü et al.9. It is worth mentioning
that Nangkangia, Yulong and Nomingia form a clde. However, Nankangia and Yulong were regarded as being
plant-eating6, 17, thus, these oviraptorid dinosaurs nested in dierent clades may imply dierent ecological niches.
Bone microstructures of Corythoraptor jacobsi. ree samples from rib, right bula and le radius of
the holotype of Corythoraptor jacobsi for histological study were acquired (see also Supplementary Information).
The preserved fibular histology exhibits intensive bone remodeling that removed early growth marks. The
remaining bular growth marks would suggest that the holotype of Corythoraptor jacobsi was an individual with
an age of more than 6 years.
Radius (Fig. 4, Supplementary Information Fig.S4) – Bone microstructure of the radius midsha has pro-
vided valuable information about the growth stage and helped to estimate the histological age of the type speci-
men of Corythoraptor jacobsi. e cortex is composed of both primary and secondary bone tissue; the latter being
dominant throughout the section. e cortex is interrupted by ve (A to E revealed in reected light; Fig.4a) or
six growth lines (A to F revealed in polarized light; Supplementary Information Fig.S4a), although more may
have been lost to extensive remodeling and erosion. e primary bone is formed by bro-lamellar tissue (indi-
cating rapid osteogenesis) with osteons typically arranged in a laminar pattern. Osteonal canals are oriented
longitudinally. is tissue remained mostly external to the growth line D (Fig.4b).
e growth lines are composed of the dark component referred herein to the line of arrested growth (LAG)
and two adjacent brighter avascular layers considered to represent the annulus. Only growth line A is slightly
modied, having a band of several distinct lamellae deposited aer LAG (Fig.4c). e three-component growth
lines indicate that during seasonal retardation of growth, rapid osteogenesis slowed down (to form pre-annulus),
and aer temporary interruption of bone deposition (expressed by LAG) osteogenesis resumed at slower rates
rst (to form post-annulus). Annular depositions are variably thick. Pre-annulus of the growth line D and E is
47 µm and 27 µm wide, respectively, whereas corresponding post-annuli are 23 µm and 28 µm wide (Fig.4b).
However, pre-annulus of growth line A is only 13 µm wide while post-annulus (lamellar bone) is much wider
(49 µm, Fig.4c).
In contrast to the bula, the spacing between concentric growth lines in the radius diminishes moderately
towards the periphery according to zonal measurements collected in reected light: AB-zone = 357 µm; BC-zone:
242 µm, CD-zone: 187 µm, DE-zone 180 µm, and E-periosteal surface zone (lacking any growth mark near exter-
nal periphery) 213 µm (Fig.4a). e fastest recorded radial growth rate of 0.96 µm is calculated for a 371-day-long
Cretaceous year32. However, daily bone deposition rate might be higher earlier in life and its estimation also
depends on how long the rapid growth season lasted in each year. Zonal measurements collected from another
portion of the cortex in polarized light (Supplementary Information Fig.S4a) are slightly dierent but correspond
to gradual slowing of somatic growth: AB-zone: 145 µm, BC-zone: 284 µm, CD-zone: 136 µm, DE-zone: 125 µm,
EF- 117 µm. ese measurements imply that the minimum age estimated based on preserved histology might be
around seven years.
Most peripheral secondary osteons occur within the CD-zone (Fig.4c,d; Supplementary Information
Fig.S4b). Internal to growth line B, the Haversian bone tissue is dense due to repeated remodeling that makes
it impossible to discern tissue discontinuities referable to early growth marks. Secondary osteons are variable
in size, ranging from 100 × 109 µm to 131 × 137 µm. e earliest growth marks likely disappeared by medullary
cavity expansion that considerably eroded the innermost cortical surface.
e cortex encloses a large medullary cavity devoid of cancellous bone (Fig.4e). Several deposition cycles of
endosteal bone line the entire margin of the medullary cavity (Fig.4e, Supplementary Information Fig.S4c). is
internal circumferential layer is composed of avascular parallel-bered bone and is variable in thickness (193 to
292 µm). It is up to almost seven times thicker than the bular endosteal bone.
e animal perished at the beginning of a new season as is exemplied by resumed bone deposition nearest to
the periosteal surface. We suggest that the holotype of Corythoraptor jacobsi had not reached maximum body size
and was still growing at the time of death. e ontogenetic stage of the holotype individual probably corresponds
to a young adult that was approaching a stationary stage of development33. It is reasonable to assume that the
prominent casque served as a sexual signaling ornament, what would imply that these oviraptorids were repro-
ductive before they nished growing. Finally, the osteochronology exhibits that Corythoraptor jacobsi required
more than 8 years to reach somatic maturity.
Putative functions of the cassowary-like crest of Corythoraptor. Recent cassowaries, the ight-
less birds from New Guinea, Australia and the Aru archipelago34, also evolved cranial casques of diverse shapes
(Fig.2e, 2f). However, the bony core of the cassowary casque incorporate much denser web of microtrabeculae
that is able to deform when subjected to pressure35, 36. Many of the trabecuale are rod-like with diameter about
0.1 mm (Fig.2g), whereas trabeculae situated close to peripheries of the casque are lamellar or plate-like measur-
ing across up to 3 mm (Fig.2h). e empty spaces are considerably smaller in cassowaries and bony shell of the
casque is sheated by keratin (Fig.2i). Rugose external surface suggests that cranial casque was likely covered by
outer keratinous sheath in Corythoraptor as well. e shell-like outer layer of the casque is approximately 2-3 mm
thick in the cassowary25 and 2 mm thick in Corythoraptor.
ese peculiar morphological similarities make the cassowaries the closest living analogue that provides some
clues for assuming putative functions of the casque in the oviraptorid Corythoraptor. Based on comparisons with the
cassowary model, we put forward three hypotheses to explain plausible functions of the Corythoraptor casque: 1)
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
termoregulatory hypothesis – larger inner cavities overlying could eectively dissipate heat produced inside of
the braincase; the function demonstrated for the casque of the cassowary37; 2) acoustic hypothesis – Kundrát and
Janáček38 suggested the supraencephalic pneumatic pathway connecting contralateral middle ear cavities indi-
cate enhancements of acoustic perception in the lower-frequency registers in the oviraptorid Conchoraptor. e
casque microstructure and positional proximity to tympanic recesses enabled the Corythoraptor´s helmet could
function as a resonator amplifying or/and pointing low frequency signals over a much greater range. Increased
perception could be then utilized for predator avoidance or prey capture. and 3) sociosexual hypothesis partly
overlap with the acousting hypothesis as more ecient low-frequency communication linked to more strongly
Figure 4. Radius microstructure of the holotype of Corythoraptor jacobsi gen. et sp. nov. (JPM-2015-001)
viewed with reected light microscopy. (a) Transversal section of the midsha showing ve growth lines (red
arrows A to E) present in the middle-to-outer cortex. Note that growth line spacing decreases periosteally. (b)
Close-up of the outer cortex with three growth lines (C to E). Note avascular layers (annuli) deposited prior
to and aer darker undulating line (line of arrested development, LAG). Primary bone is mostly preserved
external to the growth line D. Secondary osteons partly obliterate the growth line C. (c) Close-up of the middle
cortex with two growth lines (A and B). Growth line A consists of an ill-dened LAG followed by laminar bone
(annulus). Growth line B comprises a darker line (LAG) and lighter avascular layer (annulus). Note several
generations of overlapping secondary osteons are present. (d) e last growth cycles delimited by growth lines
C through E. (e) Interior margin of the cortical bone with a thick deposition of endosteal laminar bone. e
blue arrow points to the resorptive margin. Abbreviations: an, annulus; elb, endosteal laminar bone; LAG, line
of arrested growth; lb, laminar bone; mc, medullary cavity; nvc, neurovascular canal; osla, osteonal laminae; po,
primary canal; rl, resorption line; so, secondary osteon.
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
resonating casque evolved under sexual selection. Recent cassowaries produce low-frequency casque-directed
vocalizations towards a partner during mating season25, 39. Low-frequency sounds, however, are made inside the
throat apparatus in the cassowaries25 and there is no morphological evidence for such behavior in oviraptorids.
Moreover, this behaviour would require a casque structure to evolve in both sexes of the same species at least.
us there is at the moment a little support for the Corythoraptor´s casque was used solely for acoustic-sexual
signalling. Larger and probably more ornamented casque rather provided members of one or both sexes with
display structure of hierarchic intraspecic status. However, we have to stress here the fact, the male and female
cassowaries exhibit similarly ornamented with large casque25.
e cassowary-like crest in the skull is similar to the casque of cassowaries (Fig.5), which serves a sociosexual
role and functions in visual and acoustic display25. It is therefore reasonable to assume that the cassowary-like crest
of Corythoraptor jacobsi was probably utilized in a similar way. e sharp claw and long neck of Corythoraptor
jacobsi may also indicate that its living behavior is perhaps similar to the modern ightless cassowary. It seems
more reasonable to assume that the cassowary-like crest of Corythoraptor was more likely the multifunctional
structure that conspicuously expressed tness, and probably sex, of each individual of Corythoraptor.
e histological study revealed that the type specimen of Corythoraptor jacobsi was probably at least eight
years old but still not a fully grown individual. Finally, the discovery of Corythoraptor jacobsi, the seventh ovirap-
tosaurian taxon from the region, provides evidence of unprecedented morphological and taxonomic diversity of
this clade in the Ganzhou area, China.
Corythoraptor jacobsi gen. et sp. nov. represents the first oviraptorid dinosaur with a highly developed
cassowary-like skull crest from China. Phylogenetic analysis indicates that Corythoraptor jacobsi belongs to the
clade of Oviraptoridae, and shows a close relationship with Huanansaurus.
Phylogenetic analysis. We conducted a phylogenetic analysis to investigate the taxonomic anities of
Corythoraptor within Oviraptorosauria, using the modied data matrix9, 40 (see also Supplementary Information)
of Lamanna et al.41. With the addition of Corythoraptor into the modied data matrix, 45 taxa (Herrerasaurus,
Velociraptor and Archaeopteryx as outgroups; 42 taxa as ingroups) and 257 osteological characters was analyzed
using TNT (Tree Analysis Using New Technology) version 1.1 (Willi Hennig Society Edition)42. A traditional
search (tree bisection-reconnection swapping algorithm, 1,000 random seeds, 1,000 replicates, 10 trees saved per
replication) yielded 4151738 most parsimonious trees with 623 steps.
Analysis of the bone mircostructures. ree samples for histological study were acquired as small frag-
ments extracted from rib, right bula and le radius of the holotype of Corythoraptor jacobsi. e samples were
taken from near midsha of the bones. in sections of the samples revealed that original histostructure was
petrographically altered and is barely visible in transmitted light (Supplementary Information Fig.S2A). Some
histological information, however, became accessible when we investigated polished bone samples using reected
light microscopy (e.g., Supplementary Information Fig.S2B–D), as well as we examined the thin sections with
circular polarized light (e.g., compare Supplementary Information Fig.S3A,B).
e petrographic thin sections were prepared according to the following methodology: 1) bone samples
were embedded in bicomponent epoxy resin (Lamit 109; Kittfort); 2) the embedded samples were ground on a
Montasupal grinder (Germany) using SiC (grain size: 400–600 nm); 3) warm re-impregnation of the ground sur-
face with EpoFix (Struers); 4) xation of the samples to slides using epoxy resin (type 109); 5) xed samples were
sectioned using a diamond knife (diameter 150 mm, Struers); 6) xed samples were thinned on the Montasupal
Figure 5. e living scene of Corythoraptor jacobsi gen. et sp. nov. (Drawn by Zhao Chuang).
SCIentIFIC RepORTS | 7: 6393 | DOI:10.1038/s41598-017-05016-6
grinder using the abrasives of 240, 400 and 600 grits combined with ultrasound cleaning to reach a thickness of
0.2 mm; 7) nal manual abrasion using 1000 grit SiC to reach a thickness of 30 microns; and nally 8) the sections
were cover-slipped using a synthetic resin or polished on the Planopol TS (Struers). Preparation of polished bone
samples followed the above steps: 1 through 3 and 5 through 7.
e cover-slipped sections and polished samples were examined using crossed polarized light and reected
light microscopy. Photography was carried out using digital camera Olympus XC50 (operating soware: Olympus
Stream Start; Olympus So Imaging Solutions GmbH) mounted on the Olympus BX51 microscope. e images
were processed using Adobe Photoshop and CorelDRAW X5 soware. Histological measurements were taken
from digitized cross sections using ImageJ.
Data archiving. Specimen measurements, the phylogenetic character scores for Corythoraptor, and the phy-
logenetic topology with synapomorphies are available as Supplementary Information.
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2. Osmólsa, H., Currie, P.J. & Brasbold, . In e Dinosauria. 2nd edn (eds Weishampel, D., Dodson, P., & Osmólsa, H.) 165–183
(University of California Press) (2004).
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We thank Professors Louis L. Jacobs, Dale Winkler of Southern Methodist University (Dallas, TX USA) for
providing careful comments on an earlier version. e Willi Hennig Society provided the version of TNT used
for phylogenetic analysis. Zhang Y. Q. prepared the specimens. Mr. Sun Deyu and Ha Tao gave a great help when
the rst author was staying in Jinzhou for describing the specimen. is research is supported by National Natural
Science Foundation of China (grant no.: 41672019; 41688103), the Fundamental Research Funds for the Chinese
Academy of Geological Sciences (grant No.: JB1504) and the China Geological Survey (grant no.: DD 20160126)
to J.C. Lü, and National Research Foundation of Korea (grant no.: 2016R1A2B2015012) to Y.-N. Lee.
Author Contributions
J.L. designed the project. J.L., G.L., Z.S., C.S., F.T., and H.L. organized the curation and preparation of the
specimen and oversaw all research at the Jinzhou Paleontolgical Museum. J.L. performed the anatomical
descriptive research. J.L., Y.L., Y.K., performed the phylogenetic analyses. M.K. prepared gure 4. J.L. and M.K.
wrote the manuscript. All authors reviewed the manuscript.
Additional Information
Supplementary information accompanies this paper at doi:10.1038/s41598-017-05016-6
Competing Interests: e authors declare that they have no competing interests.
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Supplementary resource (1)

... For example, several North American caenagnathid species are based solely on lower jaw remains whereas others are based on postcranial holotypes (Longrich et al., 2013), complicating the testing of taxonomic assignments and reconstruction of quantitative phylogenetic hypotheses (but see Funston and Currie, 2016, for an alternative view). Incisivosaurus has been recovered as the earliest-diverging oviraptorosaurian by a number of analyses that excluded oviraptorosaurian taxa known from poorly preserved specimens, e.g., Protarchaeopteryx, Ningyuansaurus, and Luoyanggia (Longrich et al., 2013;Brusatte et al., 2014;Lamanna et al., 2014;Lü et al., 2016Lü et al., , 2017Pei et al., 2020) (figs. 1, 2A, C). ...
... 2B, D). Oviraptorosauria has two main lineages, the Caenagnathidae and the Oviraptoridae, that are consistently recovered as monophyletic sister taxa (Longrich et al., 2013;Lamanna et al., 2014;Lü et al., 2015;Funston and Currie, 2016;Lü et al., 2016;Lü et al., 2017;Yu et al., 2018; fig. 2A-D). ...
... 1). Within Caenagnathidae, Microvenator and the giant Gigantoraptor are consistently recovered as early-diverging members, whereas the interrelationships between later-diverging members remain unresolved largely due to the issue of nonoverlapping elements between specimens of different species (Lamanna et al., 2014;Lü et al., 2015;Lü et al., 2016;Lü et al., 2017;fig. 2C). ...
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New and important pennaraptoran specimens continue to be discovered on a regular basis. Yet, with these discoveries the number of viable phylogenetic hypotheses has increased, including ones that challenge the traditional grouping of dromaeosaurids and troodontids within a monophy-letic Deinonychosauria. This chapter will cover recent efforts to address prevailing phylogenetic uncertainties and controversies, both between and within key clades, including deinonychosaurian monophyly, the phylogenetic position of anchiornithines and scansoriopterygids, and the interrelationships of enantiornithines. While recent discoveries mainly from Asia have created much of the latest uncertainty and controversy, new material, particularly from Asia, promises to rather fittingly address these issues. Further curatorship of long-standing phylogenetic datasets and more prevalent use of extended analytical protocols will be essential to meeting this challenge, especially for groups whose boundaries have been blurred. As it becomes increasingly difficult to study all fossil materials, owing to their growing numbers and ever disparate locations, broader use of digital fossils and online character databases for character coding is acutely needed to ensure that errors arising from remote, rather than firsthand, scoring are reduced as far as possible, particularly at this time of rapid data accumulation.
... This allows us to visualize the evolutionary trends of different characters in Pennaraptora, especially across those well-sampled lineages such as Oviraptoridae and Caenagnathidae. As there is no single phylogeny that includes all the pennaraptorans involved in the study, we have produced a hypothetical phylogenetic tree by integrating the trees of different pennaraptoran clades (Lü et al., 2017;Pei et al., in press). ...
... 2B, D). However, some late-diverging oviraptorosaurians have a taller skull because of the presence of a tall crest (Lamanna et al., 2014;Funston et al., 2017;Lü et al., 2017). Oviraptorids, such as Rinchenia, have a skull length and height that are nearly identical (Tsuihiji et al., 2016: fig. ...
... Six functional characters were measured and subjected to ancestral-state reconstruction analysis using squared-change parsimony and a tree topology ( fig. 4; table 2) based on Lü et al. (2017) and Pei et al. (in press). The reconstructed nodal value of mechanical advantage (MA) of the jawclosing system (average of AMA and PMA) of scansoriopterygids (~0.179) is lower than those of oviraptorosaurians. ...
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Oviraptorosauria and Scansoriopterygidae are theropod clades that include members suggested to have partially or fully herbivorous diets. Obligate herbivory and carnivory are two ends of the spectrum of dietary habits along which it is unclear how diet within these two clades might have varied. Clarifying their diet is important as it helps understanding of dietary evolution close to the dinosaur-bird transition. Here, diets are investigated by conventional comparative anatomy, as well as measuring mandibular characteristics that are plausibly indicative of the animal's feeding habit, with reference to modern herbivores that may also have nonherbivorous ancestry. In general, the skulls of scansoriopterygids appear less adapted to herbivory compared with those of oviraptorids because they have a lower dorsoventral height, a smaller lateral temporal fenestra, and a smaller jaw-closing mechanical advantage and they lack a tall coronoid process prominence. The results show that oviraptorid mandibles are more adapted to herbivory than those of caenagnathids, early-diverging oviraptorosaurians and scansoriopterygids. It is notable that some caenagnathids possess features like an extremely small articular offset, and low average mandibular height may imply a more carnivorous diet than the higher ones of other oviraptorosaurians. Our study provides a new perspective to evaluate different hypotheses on the diets of scansoriopterygids and oviraptorosauri-ans, and demonstrates the high dietary complexity among early-diverging pennaraptorans.
... Modern, ornamented analogues-living animals with comparable structures, lifestyles, and biology-have been proposed in order to aid inferences about extinct, ornamented taxa, particularly those among non-avian dinosaurs. These include artiodactyl mammals, squamate lizards, neognathous birds, and palaeognathous birds (Dodson 1975;Farlow and Dodson 1975;Bubenik and Bubenik 1990;Hieronymus et al. 2009;Snively and Theodor 2011;Lü et al. 2017;Eastick et al. 2019;Angst et al. 2020). However, bony cranial ornament function is little explored in extant tetrapods outside of mammals, including for birds, despite the fact that birds are living dinosaurs. ...
... Because they are flightless, large-bodied, and generally resemble non-avian theropods, modern Casuarius is one of the most commonly referenced avian analogues for the ornaments of extinct non-avian dinosaurs (e.g. Dodson 1975;Hone et al. 2012;Farke et al. 2013;Lü et al. 2017). Nonetheless, volant, smaller bodied birds with osseous cranial ornaments may also represent valuable comparative systems (Angst et al. 2020). ...
Birds, along with their dinosaurian precursors, possess a variety of bony cranial expansions. A deep understanding of the phenotypic complexity of these structures would be useful for addressing the development, evolution, and function of hard-tissue cranial ornamentation. Yet, the evolutionary significance and function of these structures have gone largely unaddressed because no unifying conceptual framework for interpreting bony cranial expansions currently exists. To provide such a framework, we examine osseous ornament variation in modern birds, using µ-CT imaging to examine the cranial casque components, structural composition, and developmental changes of two neognathous (Numida meleagris, Macrocephalon maleo) and one palaeognathous species (Casuarius casuarius) and survey the avian osteology literature of the 11 orders containing members with osseous cranial ornamentation. Our anatomical analyses suggest two broad configuration categories: (i) geminal, in which ornaments consist of paired elements only (i.e. within Neognathae) and (ii) disunited, in which ornaments consist of unpaired, midline elements along with paired bones (i.e. within Palaeognathae). Ornament bones contribute to casque elevation (proximal ornament support), elaboration (distal ornament shape), or both. Our results hold utility for unravelling the selection processes, particularly in difficult-to-decipher display roles, that shaped modern avian casques, as well as for the use of extant avians as comparative analogues of non-avian dinosaurs with ornamental head structures.
... The Upper Cretaceous Nanxiong Formation of Jiangxi Province, southern China has yielded a diverse array of vertebrates in recent years, including theropods (Xu and Han 2010;Wang et al. 2013;Wei et al. 2013;Lü et al. 2013aLü et al. , 2014Lü et al. , 2015Lü et al. , 2016Lü et al. , 2017Mo and Xu 2015), ornithopods (Xing et al. 2021(Xing et al. , 2022, crocodiles (Li et al. 2019), turtles (Tong and Mo 2010), lizards (Mo et al. , 2012, and mammals (Jin et al. 2022), as well as a vast number of dinosaur eggs (Sato et al. 2005;Cheng et al. 2008;Ji 2009;Shao et al. 2014;Zhao et al. 2015;Wang et al. 2016;Jin et al. 2019;Bi et al. 2021;Fang et al. 2022). Only one sauropod taxon, Gannansaurus sinensis, has been recorded in this area (Lü et al. 2013b). ...
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This paper represents the first time that I propose to correlate all fossil quarries of Dinosaur Provincial Park by identifying marker beds from a combination of 3D models, digital elevation models and orthomosaics of the badlands landscape of this famous fossil locality, with the raw images all obtained from drone flights that I lead with my crew during every field season in the Park.
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An unabated surge of new and important discoveries continues to transform knowledge of pen-naraptoran biology and evolution amassed over the last 150+ years. This chapter summarizes progress made thus far in sampling the pennaraptoran fossil record of the Mesozoic and Paleocene and proposes priority areas of attention moving forward. Oviraptorosaurians are bizarre, nonparavian pennaraptorans first discovered in North America and Mongolia within Late Cretaceous rocks in the early 20th century. We now know that oviraptorosaurians also occupied the Early Cretaceous and their unquestionable fossil record is currently limited to Laurasia. Early Cretaceous material from China preserves feathers and other soft tissues and ingested remains including gastroliths and other stomach contents, while brooding specimens and age-structured, single-species accumulations from China and Mongolia provide spectacular behavioral insights. Less specialized early oviraptorosaurians like Incisivosaurus and Microvenator remain rare, and ancestral forms expected in the Late Jurassic are yet to be discovered, although some authors have suggested Epidexipteryx and possibly other scansoriopterygids may represent early-diverging oviraptorosaurians. Long-armed scansoriopterygids from the Middle-Late Jurassic of Laurasia are either early-diverging oviraptorosaurians or paravians, and some have considered them to be early-diverging avialans. Known from five (or possibly six) feathered specimens from China, only two mature individuals exist, representing these taxa. These taxa, Yi and Ambopteryx, preserve stylopod-supported wing membranes that are the only known alternative to the feathered, muscular wings that had been exclusively associated with dinosaurian flight. Thus, scansoriopterygid specimens-particularly those preserving soft tissue-remain a key priority for future specimen collection. Dromaeosaurids and troodontids were first discovered in North America and Mongolia in Late Cretaceous rocks. More recent discoveries show that these animals originated in the Late Jurassic, were strikingly feathered, lived across diverse climes and environments, and at least in the case of dromaeosaurids, attained a global distribution and the potential for aerial locomotion at small size.
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Oviraptorosaurs are a bizarre group of bird-like theropod dinosaurs, the derived forms of which have shortened, toothless skulls, and which diverged from close relatives by developing peculiar feeding adaptations. Although once among the most mysterious of dinosaurs, oviraptorosaurs are becoming better understood with the discovery of many new fossils in Asia and North America. The Ganzhou area of southern China is emerging as a hotspot of oviraptorosaur discoveries, as over the past half decade five new monotypic genera have been found in the latest Cretaceous (Maastrichtian) deposits of this region. We here report a sixth diagnostic oviraptorosaur from Ganzhou, Tongtianlong limosus gen. et sp. nov., represented by a remarkably well-preserved specimen in an unusual splayed-limb and raised-head posture. Tongtianlong is a derived oviraptorid oviraptorosaur, differentiated from other species by its unique dome-like skull roof, highly convex premaxilla, and other features of the skull. The large number of oviraptorosaurs from Ganzhou, which often differ in cranial morphologies related to feeding, document an evolutionary radiation of these dinosaurs during the very latest Cretaceous of Asia, which helped establish one of the last diverse dinosaur faunas before the end-Cretaceous extinction.
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Our understanding of caenagnathids has benefited from recent discoveries, including nearly complete skeletons from the Hell Creek Formation of Montana. However, their phylogenetic relationships remain unclear. A new specimen from the Horseshoe Canyon Formation of Alberta has implications for the phylogeny and paleobiology of these creatures. The partial skeleton is articulated and includes a mandible, a full cervical and dorsal series of vertebrae, a right pectoral girdle and arm, a sternum, gastralia, a partial ilium, and a partial hind limb. The mandible is edentulous and the articular ridge is intermediate in form between Caenagnathus collinsi and Chirostenotes pergracilis. The neck is long and composed of at least 11 well-pneumatized cervical vertebrae with fused cervical ribs. The dorsal ribs have finger-like uncinate processes dissimilar in shape to those of other oviraptorosaurs. The pectoral girdle is large and typically maniraptoran, except that the glenoid of the scapulocoracoid faces laterally instead of posteroventrally. The arm is well muscled and can be interpreted to have been a pennibrachium, as indicated by ulnar papillae on the ulna. The manus is characterized by a short first metacarpal but an elongate phalanx I-1 and oviraptorid-like phalangeal proportions in the second digit. These and other features indicate that the specimen represents a new taxon, Apatoraptor pennatus, gen. et sp. nov. Phylogenetic analysis resolves the complicated relationships of Caenagnathidae and allows the evolution of display features to be traced throughout Oviraptorosauria. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at Citation for this article: Funston, G. F., and P. J. Currie. 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1160910.
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A description of Nomingia gobiensis gen. et sp. n., the first known dinosaur with a pygostyle, the structure known so far only in birds, is presented. The specimen comes from the Late Cretaceous strata at Bugin Tsav, Trans-Altai Gobi, Mongolia. N. gobiensis is assigned within the Oviraptorosauria based on the following characters: pneumatized caudal vertebrae, posteriorly concave ischium, and deep cervicodorsal hypapophyses. This specimen has been previously partially described without being formally named (Barsbold et al. 2000).
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The Ganzhou area of Jiangxi Province, southern China is becoming one of the most productive oviraptorosaurian localities in the world. A new oviraptorid dinosaur was unearthed from the uppermost Upper Cretaceous Nanxiong Formation of Ganzhou area. It is characterized by an anterodorsally sloping occiput and quadrate (a feature shared with Citipati), a circular supratemporal fenestra that is much smaller than the lower temporal fenestra, and a dentary in which the dorsal margin above the external mandibular fenestra is strongly concave ventrally. The position of the anteroventral corner of the external naris in relation to the posterodorsal corner of the antorbital fenestra provides new insight into the craniofacial evolution of oviraptorosaurid dinosaurs. A phylogenetic analysis recovers the new taxon as closely related to the Mongolian Citipati. Six oviraptorid dinosaurs from the Nanxiong Formation (Ganzhou and Nanxiong) are distributed within three clades of the family. Each of the three clades from the Nanxiong Formation has close relatives in Inner Mongolia and Mongolia, and in both places each clade may have had a specific diet or occupied a different ecological niche. Oviraptorid dinosaurs were geographically widespread across Asia in the latest Cretaceous and were an important component of terrestrial ecosystems during this time.