Tongtianlong limosus Lü, Chen, Brusatte, Zhu & Shen, 2016, gen. et sp. nov

Tongtianlong limosus gen. et sp. nov.

(Figs 2, 3 and 4)

Etymology. Tongtian, Chinese Pinyin, refers to Tongtianyan of Ganzhou, the first grotto south of the Yangtze River. Tongtian also means the road to heaven, a fitting epitaph for a deceased dinosaur preserved with outstretched arms. Long, Chinese Pinyin for dragon. Limosus, Latin for muddy, refers to the holotype specimen being found in an unusual posture in a mudstone (Fig.5).

Holotype. A nearly complete, three-dimensionally preserved skeleton with skull and lower jaws (DYM-2013-8). The specimen is accessioned at the Dongyang Museum, Dongyang City, Zhejiang Province.

Type locality and horizon. The building site of the No. 3 high school of Ganxian (GPS coordinates are provided on request from the first author); Nanxiong Formation (Maastrichtian, Upper Cretaceous)35.

Diagnosis. Oviraptorid dinosaur with the following unique combination of characters, with autapomorphies among all oviraptorosaurs indicated with an asterisk and autapomorphies among oviraptorids indicated with a double asterisk (these latter features are present in some caenagnathids): dome-like skull roof with highest point located above the posterodorsal corner of the orbit*; anterior margin of the premaxilla highly convex in lateral view*; distinct process at the middle of the anterior margin of the parietal on the skull roof*; plate-like lacrimal shaft that is anteroposteriorly long in lateral view, with a flat lateral surface*; foramen magnum smaller than the occipital condyle**; absence of symphyseal ventral process of the dentary**; absence of distinct lateral xiphoid process of the sternum posterior to the costal margin**.

Tongtianlong differs from other Ganzhou oviraptorids with preserved skull material (Banji, Huanansaurus) in that the anteroventral corner of the external naris is far above a horizontal line tangent with the posterodorsal corner of the antorbital fenestra, an unusual feature otherwise only seen in Nemegtomaia 36 – 38 and Rinchenia (Barsbold 39) (= Oviraptor mongoliensis 1, 40). Tongtianlong also differs from other Ganzhou oviraptorids in numerous ways that are encapsulated in the character scores in our phylogenetic analysis. Tongtianlong differs from Banji 26 in possessing a postorbital process of the jugal that is posterodorsally inclined relative to the ventral ramus (not perpendicular), lacking a downturned symphyseal portion of the dentary (see also Supplementary Information Fig. S1a), lacking a prominent process on the posteroventral surface of the dentary symphysis, and possessing a more anteroposteriorly elongate external mandibular fenestra.

Tongtianlong is distinguished from Ganzhousaurus 27 in lacking a downturned symphyseal portion of the dentary, possessing a more anteroposteriorly elongate external mandibular fenestra, having a dentary that contributes to the ventral border of the external mandibular fenestra (not excluded from the border by the anterior extension of the angular), lacking a depression on the lateral surface of the dentary immediately anterior to the external mandibular fenestra, and possessing a metatarsal III that is anteroposteriorly flattened with a concave posterior surface (not ovoid or subtriangular in cross section).

Tongtianlong is different from Jiangxisaurus 28 in lacking a downturned symphyseal portion of the dentary. Furthermore, the ratio of radius length to humerus length (78%) and the height-to-length ratio of the lower jaw (34%) are greater than those of Jiangxisaurus (70% and 20%, respectively).

Tongtianlong differs from Nankangia 9 in lacking a prominent process on the posteroventral surface of the dentary symphysis, lacking a deep fossa on the lateral surface of the dentary, having a dentary that contributes to the dorsal border of the external mandibular fenestra (not excluded from the border by the anterior extension of the surangular), and lacking a depression on the lateral surface of the dentary immediately anterior to the external mandibular fenestra.

Tongtianlong differs from Huanansaurus 29 in skull morphology and forelimb proportions. There is a crest in Huanansaurus but not in Tongtianlong. The dorsal margin of the lower jaw, from the anterior tip to the coronoid eminence, is smoothly convex in Tongtianlong, whilst it is wave-like in Huanansaurus. The antorbital fenestra is sub-oval in Tongtianlong, but triangular in Huanansaurus. The ratio of radius length to humerus length in Tongtianlong (0.78) is much smaller than that of Huanansaurus (0.97). The anteroventral corner of the external naris is slightly below the horizontal line projected through the posterodorsal corner of the antorbital fenestra in Huanansaurus, whilst it is far above this line in Tongtianlong (Fig.6).

The specimen is very well preserved in three dimensions, with the bones in natural articulation (Figs2 and 3 a; see also Supplementary Information Table S1). The limbs are splayed out sideways relative to the trunk, and the neck is curved upwards, such that the head is elevated relative to the remainder of the body. Because the specimen was collected by a farmer and construction workers, and it was not mapped in situ while being excavated, it is difficult to interpret what biological and/or taphonomic processes caused this strange posture. Judging by the fine state of preservation, the specimen probably was originally complete or nearly complete. However, some portions of the skeleton are missing, such as the distal regions of the arms, the right pelvic girdle and hind leg, and parts of the tail. This is because the specimen was collected by workers at an active construction site. The specimen was exposed after workmen blasted away some of the surrounding rocks with TNT; a drill hole where TNT was placed can be seen near the pelvic girdle.

The skull is almost completely preserved. It is missing only small portions of the anterior end of the premaxilla and nasals, and a small part of the right lower jaw. The most salient feature of the skull is that the cranial roof is dome-like, with its highest point above the posterodorsal corner of the orbit. Many other oviraptorosaurs possess cranial ornaments, which in some cases are elaborate and highly pneumatic (Fig.4). However, in other taxa these crests are usually thinner (such as in Nemegtomaia 36 – 38, it is 6 mm) than the dome-like condition in Tongtianlong. Furthermore, in other taxa these crests are peaked further anteriorly relative to Tongtianlong, either at the anterior end of the snout above the external naris and antorbital fenestra (as in Banji, Citipati, Oviraptor, and Nemegtomaia), or at approximately the midpoint of the cranium above the orbit (as in Rinchenia, Huanansaurus, and caenagnathids like Anzu 41) (Fig.6). Therefore, the posteriorly-peaked dome-like crest of Tongtianlong is autapomorphic among oviraptorosaurs, and a novel type of cranial ornamentation in this highly variable clade. The fine three-dimensional preservation of the specimen ensures that the shape of the dome-like crest is not an artefact of crushing or deformation.

There are five main openings in the cranium, as is standard for oviraptorosaurs and other dinosaurs. Anteriorly a large, oval-shaped external naris is positioned above a slightly smaller, triangular antorbital fenestra (Fig.4). The anteroventral corner of the naris is located far above the level of the posterodorsal corner of the antorbital fenestra, which is also the case in Nemegtomaia and Rinchenia, but differs from the condition in most other oviraptorosaurs, in which the naris extends further ventrally so that it reaches past the posterodorsal corner of the antorbital fenestra (Fig.6). The orbit is large and nearly circular, as is typical for oviraptorosaurs. The lateral temporal fenestra is the largest opening in the skull. It is rectangular, with a long axis that extends slightly anteroventrally, which differs from the more circular or square fenestrae of many other oviraptorosaurs. The supratemporal fenestra is positioned above the lateral temporal fenestra and is partially visible in lateral view. It is much smaller than the orbit and lateral temporal fenestra.

The premaxilla is toothless like in all derived oviraptorosaurs. The left and right premaxillae appear to be unfused to each other, based on open sutures in the region of the broken dorsal surfaces of both bones, but the premaxilla and maxilla are fused together without a clear sutural trace. The anterior margin of the premaxilla is highly convex, which is an autapomorphy of Tongtianlong. Most other oviraptorosaurs have a straight anterior premaxilla (e.g., Citipati 42, Khaan 42), and this is also the case in the Ganzhou oviraptorid Huanasaurus 29. Yulong and the Ganzhou oviraptorid Banji have a slightly rounded anterior margin of the premaxilla in lateral view, but it is not nearly as convex as in Tongtianlong. The premaxilla is divided into two branches, both of which extend posterodorsally. The upper one forms the anterodorsal margin of the external naris, whereas the much wider lower one forms most of the anterodorsal margin of the antorbital fenestra, thus separating the naris from the antorbital fenestra and completely excluding the maxilla from the narial border. The posterior end of this branch overlaps the lateral surface of the lacrimal. The divergence of the two branches defines the shape of the external naris. Anteroventral to the naris, a deeply concave fossa extends on the lateral surface of the premaxilla, as in Huanasaurus 29, Yulong 10, and Nemegtomaia 37, 38, but unlike the slightly concave surface in Citipati 6. The maxilla is very small and exposed only as a tiny sliver of bone in lateral view. It forms the ventral margin of the antorbital opening and lacks teeth, but has a small triangular ‘tooth-like’ process on its ventral surface.

The lacrimal is divided into three branches: a short anterior process that is covered by the premaxilla, a bulbous posterior process that extends dorsally to define the anterodorsal corner of the orbit, and a large ventral shaft. The shape of the shaft is unique: whereas in other oviraptorosaurs the lacrimal shaft is gracile (thin anteroposteriorly) and has at least a partially convex lateral surface (Fig.6), in Tongtianlong it is robust (thick anteroposteriorly) with a flat lateral surface (Fig.4). In effect, the lacrimal shaft of Tongtianlong is plate-like, which is considered an autapomorphy of the taxon. In the region where the three processes meet, the lateral surface of the lacrimal is penetrated by a large opening (called the nasopharyngeal canal by Balanoff and Norell 43) that leads into an internal recess, which is further subdivided internally. Anteroventral to this pneumatic opening is an ovoid fossa on the lateral surface of the lacrimal, which is probably also pneumatic in origin, and which may also invade the bone internally, although poor preservation makes this difficult to confirm. Complex pneumaticity in this region is common in oviraptorosaurs 6, 43). However, the pneumatic openings in Tongtianlong are much larger and more elaborate than the corresponding pneumaticity in the two Ganzhou oviraptorids with well-preserved cranial material, Huanansaurus 29 and Banji 26.

The postorbital is triradiate, with short anterior and posterior processes and a very long ventral process that projects anteroventrally, terminating at the floor of the orbit. The slender and elongate jugal is divided into three branches. The rod-like anterior process contacts the lacrimal and maxilla. The short ascending process extends posterodorsally to make up approximately half of the postorbital bar separating the orbit and lateral temporal fenestra. The posterior process contacts the quadratojugal underneath the lateral temporal fenestra. Here, the jugal overlaps the quadratojugal laterally, and the two bones are sutured but not fused. The quadratojugal is tightly appressed to the lateral surface of the quadrate, and it does not appear that the two bones could move relative to each other. There is, however, a small fenestra between the small dorsal process of the quadratojugal and the lateral margin of the quadrate. The dorsal part of quadrate is bent backwards, and there is an opening on the anterior surface of the quadrate which indicates that the bone is pneumatized.

On the skull roof, the dorsal surface of the posterior portion of the nasal is smoothly convex. The left and right nasals are fused, without any sign of a suture between them. The lateral surface of the nasal is strongly concave, and although the surface is not well preserved, visible regions of original bone texture indicate extreme pneumaticity in this region, as is standard for derived oviraptorosaurs. The nasal-frontal suture is V-shaped in dorsal view. The frontals are short anteroposteriorly, and the left and right elements are not fused on the midline. The two parietals are fused to each other, but not to the frontals and there is no parietal crest (see also Supplementary Information, Fig. S1b). The frontal-parietal suture is mostly straight, but there is a distinct process extending forwards from the middle of the anterior margin of the parietal (Fig. 3 d, pp). This process is wedged between the frontals. It is considered an autapomorphy of Tongtianlong, as it is absent in other oviraptorosaurs.

Portions of the braincase are visible in lateral and posterior view. The supraoccipital is triangular, which a concave posterior surface. The exoccipital-opisthotic forms the dorsal margin of the foramen magnum, thus separating the supraoccipital from the foramen margin. The exoccipital tapers as it extends lateroventrally. The occipital condyle is larger than the foramen magnum, a condition that is seen in some caenagnathids (e.g., Anzu 41), but differs from the proportionally smaller occipital condyles of other oviraptorids. The occipital condyle is located posterior to the articular condyles of the quadrate.

The mandible is nearly complete. In lateral view, the ventral margin of the lower jaw is straight. The anterior end of the dentary is not as strongly downturned as in other derived oviraptorids. There is no depressed fossa on the lateral surface of the dentary immediately anterior to the external mandibular fenestra, and there are no articular grooves for the dentary on the ventrolateral edge of the angular and the dorsal surface of the surangular. The dentary contributes widely to the dorsal and ventral margins of the external mandibular fenestra, which is more anteroposteriorly elongated than the circular fenestrae of many other oviraptorids. The posterior part of the surangular is strongly concave laterally, and is pierced by a small opening.

Postcranially, the neck is comprised of 11 cervical vertebrae. The first nine of these are preserved in natural articulation, with their dorsal surfaces exposed. In dorsal view the anterior-middle cervicals are roughly square shaped, as defined by lines drawn between the posterior margins of the postzygapophyses, the anterior margins of the prezygapophyses, and the lateral edge of the vertebra (Fig.3 e). They become more rectangular in shape, longer than wide, more posteriorly in the neck. The neural spines are very small, as they are reduced to tiny peg-like projections at the center of the neural arches. The epipophyses are well developed in the second, third and fourth cervical vertebrae, but they become smaller in the middle cervicals and then disappear posterior to the sixth vertebra. There is a pneumatic opening (=pleurocoel) visible on the slightly exposed lateral centrum surface of the second cervical, but the lateral surfaces of the remaining cervicals are covered by matrix. The isolated posterior cervical vertebra, which is not in close articulation with the rest of the neck and therefore more widely exposed than the others, has a concave anterior articular surface of the centrum and a slightly convex posterior articular surface.

The dorsal vertebrae were heavily damaged during collection, so few details of their morphology can be observed. The neural spines of the posterior dorsals are tall and slightly expanded anteroposteriorly. Some dorsal ribs are present on both sides of the specimen, none of which exhibit any pneumatic openings on their proximal ends. The sacrum is not well preserved, but the anterior neural spines are clearly unfused to each other and were closely appressed to the medial surface of the ilium in dorsal view. There appears to be a pneumatic foramen (=pleurocoel) on the final sacral vertebra, and the lateral ends of the fused transverse processes and sacral ribs are strongly expanded anteroposteriorly, with rounded dorsal surfaces. Part of the distal tail is missing, but there are at least 19 caudal vertebrae. The caudals are rectangular in dorsal view, with elongate transverse processes that extend laterally and slightly posteriorly. One laterally exposed anterior caudal has a small opening that appears to be pneumatic in nature. The haemal arches are very long.

Portions of the shoulder girdles and proximal forearms are present on both sides of the specimen. The scapula is slender and curved medially. Its proximal end is expanded but not fused to the coracoid, the two bones forming an angle of approximately 130 degrees when in articulation. The coracoid is quadrangular in shape and has a large distally tapering posteroventral process, which extends slightly past the glenoid and is rounded at its end. The lateral surface of the coracoid is convex, the distinct biceps tubercle is located anterior to the glenoid, and the small and elongated coracoid foramen is positioned between the dorsal margin of the bone and the biceps tubercle. The medial surface of the coracoid is deeply concave and the coracoid foramen is expressed as a much larger, more circular opening than on the lateral surface. The thin sternum is a single element consisting of fused left and right components. It lacks a lateral xiphoid process and there is no groove for the coracoids along its anterior margin. The furcula is a broadly U-shaped, with a short ventral process on the midline and flattened distal ends (Fig.3 b). The humerus has a long deltopectoral crest, which extends distally for nearly half the length of the shaft (Fig.3 c). The shaft is slightly twisted as in Heyuannia 22 and Nankangia 9. Part of the radius is preserved on the left side, but the ulnae and more distal forelimb elements are missing.

Very few details of the pelvic girdle are apparent, due to damage that occurred during collecting. Parts of the ilium and pubis are present but little can be said of their morphology, although the preserved portions indicate that the pelvis is mesopubic and the distal ends of the left and right pubes are not fused together. The ischia are better preserved on the left side. The posterior margin of the shaft is deeply concave, the distal margin of the obturator process is straight, and the lateral surface of the bone is concave. The tibia is longer than the femur. It has a straight shaft, a well-developed cnemial crest, and an expanded distal end with a concave posterior surface. The astragalus is tightly appressed to the distal tibia. In posterior view, the ventral margin of the astragalus is concave dorsally, and in anterior view the ascending process is taller than wide. Two flattened distal tarsals are fused to each other and the proximal metatarsals. Distal tarsal III, which covers the proximal ends of metatarsals II and III, is larger than distal tarsal IV, which covers the proximal end of metatarsal IV. The left pes is partially preserved (See also Supplementary Information Fig. S2). Metatarsal III is longer than metatarsal II, which is longer than metatarsal IV. Metatarsal III remains visible along the length of the metatarsaus, with only a slight constriction near its proximal end. Metatarsal V is short and rod-like with a pointed distal end. It is approximately 35% of the length of metatarsal V. The single visible pedal ungual is slightly curved.

Phylogenetic analysis. Tongtianlong can clearly be assigned to Oviraptoridae based on numerous characters that are diagnostic of the clade (or proximal nodes within Oviraptorosauria), including: a pneumatic premaxilla; a medially inset subantorbital portion of the maxilla; fused nasals; a laterally projecting medial part of the lacrimal shaft that forms a flattened transverse bar in front of the eye; pneumatic skull roof bones; left and right iliac blades closely approaching or contacting each other on the midline 1; and proximal caudals with pneumatized centra 44.

We added Tongtianlong to a modified version of the phylogenetic dataset of Lü et al. 29, which itself was an updated version of the dataset of Lamanna et al. 41. We changed some characters that were previously multistate characters combining absence/presence and morphological differences into two separate characters, and also ordered multistate characters that describe a progressive sequence of size or morphological change. The data matrix now includes 43 taxa scored for 237 characters (see Methods and Supplementary Information).

The strict consensus of the 33,104 most parsimonious trees recovers Tongtianlong as deeply nested within Oviraptoridae (synapomorphies for Oviraptoridae and other major clades largely follow previous analyses of this dataset, and won’t be repeated here) (Fig.7). Tongtianlong is the sister taxon to a sister-taxon pair of the Ganzhou oviraptorid Banji 26 and Wulatelong from the Campanian of Inner Mongolia 18. The subclade comprised of these three taxa is united by three synapomorphies: the lack of a sagittal crest along the interparietal contact (character 30), a jugal process of the postorbital that extends far ventrally (character 36), and the presence of a surangular foramen (character 94). Tongtianlong is not recovered as a particularly close relative of any of the four other Ganzhou oviraptorids. Of these, Nankangia is placed within a polytomy as one of the most basal oviraptorids, Huanansaurus is recovered as an ‘intermediate’ grade oviraptorid that is outside of the clade consisting of Tongtianlong and more derived oviraptorids, and Jiangxisaurus and Ganzhousaurus are positioned as very highly nested oviraptorids, as successive outgroups to the specialized subclade centered on Ingenia.

The phylogenetic separation between Tongtianlong and other Ganzhou oviraptorids provides further evidence for their generic separation. It is not outside of the realm of possibility, however, that future work on oviraptorosaur ontogeny may show that Tongtianlong is synonymous with another Ganzhou taxon. If this is the case, we suggest that Banji would be the most likely con-specific, as it is the most closely related to Tongtianlong and is based on a much smaller holotype that conceivably could belong to a juvenile 26. With that said, we consider the phylogenetic separation of Tongtianlong and Banji, the possession of numerous autapomorphies in Tongtianlong that are not seen in Banji, and the many character differences between the holotypes of Tongtianlong and Banji to be strong evidence that the two are distinct taxa, based on our current understanding of oviraptorosaur ontogeny and morphology.