Lingwulong shenqi, the “Amazing Dragon”, and the dispersal of Sauropods.

Skeletal reconstruction and exemplar skeletal remains of Lingwulong shenqi. Scale bars = 100 cm for a and 5 cm for b–o. From Xu et al., 2018

Sauropods were the largest terrestrial vertebrates. Their morphology is easy recognizable: a long, slender neck and a tail at the end of a large body supported by four columnar limbs. Sauropods dominated many Jurassic and Cretaceous terrestrial faunas. Although they were globally distributed, the absence of Diplodocoidea from East Asia has been interpreted as a biogeographic pattern caused by the Mesozoic fragmentation of Pangea. However, a newly discovered dinosaur from the Middle Jurassic of northern China suggests that Sauropods dispersed and diversified earlier than previously thought.

Lingwulong shenqi — literally the “amazing dragon from Lingwu” — is the first well-preserved confirmed diplodocoid from East Asia (23 synapomorphies support the placement of Lingwulong within Diplodocoidea with 10 of these being unequivocal). The holotype, (LM) V001a, is a partial skull comprising the braincase, skull roof, and occiput, and an associated set of dentary teeth. The paratype, (LGP) V001b, comprises a semi-articulated partial skeleton including a series of posterior dorsal vertebrae, complete sacrum, the first caudal vertebra, partial pelvis, and incomplete right hind limb.

An artist’s interpretation of what Lingwulong shenqi (Image: Zhang Zongda)

The Lingwulong specimens were found in the Yanan Formation at Ciyaopu, in northwest China. This formation has been divided in four or five members. Although, no radiometric constraints have been obtained for the Yanan Formation, its age has been estimated on the basis of biostratigraphy. The presence of a conchostracans assemblage (including Palaeoleptoestheria, Triglypta, and Euestheria) is indicative of a Middle Jurassic age.

The East Asian Isolation Hypothesis (EAIH) has become a well-established explanation of profound differences between Jurassic (and sometimes Early Cretaceous) Asian terrestrial faunas, that resulted in the evolution of endemic groups such as mamenchisaurid sauropods, and the early diverging lineage of tetanurans, oviraptorosaurs, therizinosaurs. In this model, the isolation ended in the Early Cretaceous when marine regressions allowed the invasion of groups from elsewhere in Pangaea, and the dispersal of Asian endemics (e.g., oviraptorosaurs, marginocephalians) into Europe and North America. However, it was claimed that diplodocoids never took part in these dispersals because the end-Jurassic extinction that greatly reduced their diversity and geographic range in the Early Cretaceous. The discovery of Lingwulong undermines the EAIH, forcing a significant revision of hypotheses concerning the origins and early radiation of Neosauropoda.



Xing Xu, Paul Upchurch, Philip D. Mannion, Paul M. Barrett, Omar R. Regalado-Fernandez, Jinyou Mo, Jinfu Ma and Hongan Liu. 2018. A New Middle Jurassic Diplodocoid Suggests An Earlier Dispersal and Diversification of Sauropod Dinosaurs. Nature Communications.9, 2700.  DOI:  10.1038/s41467-018-05128-1 





Introducing Akainacephalus johnsoni

Skeletal reconstructions of Akainacephalus johnsoni. From Wiersma and Irmis, 2018

The Ankylosauria is a group of herbivorous, quadrupedal, armoured dinosaurs subdivided in two major clades, the Ankylosauridae and the Nodosauridae. The group is predominantly recorded from the Late Cretaceous (Turonian—late Maastrichtian) of Asia and the last Cretaceous (early Campanian—late Maastrichtian) of western North America (Laramidia). Ankylosauridae were present primarily in Asia and North America, and the most derived members of this clade are characterized by shortened skulls, pyramidal squamosal horns, and tail clubs.

Akainacephalus johnsoni, a new genus and species of an ankylosaurid dinosaur from the upper Campanian Kaiparowits Formation of southern Utah, represents the most complete ankylosaurid specimen from southern Laramidia to date, and reveals new details about the diversity and evolution of this clade. The genus name is derived from the Greek akaina, meaning “thorn” or “spine,” referring to the thorn-like cranial caputegulae of the holotype; and “cephalus,” the Greek meaning for head. The specific epithet honors Randy Johnson, volunteer preparator at the Natural History Museum of Utah.

Skull of Akainacephalus johnsoni. From Wiersma and Irmis, 2018

The holotype (UMNH VP 20202) is a partial skeleton comprising a complete skull, both mandibles, predentary, four dorsal, four dorsosacral, three sacral, one caudosacral, and eight caudal vertebrae, dorsal ribs, a complete tail club, both scapulae, left coracoid, right humerus, right ulna, partial left ilium, left femur, left tibia, left fibula, phalanx, two partial cervical osteoderm half rings, and 17 dorsal and lateral osteoderms of various sizes and morphologies.

The most striking feature of Akainacephalus johnsoni is the skull ornamentation comprising several symmetrical rows of small pyramidal and conical caputegulae along the dorsolateral surface of the skull. The postorbital horns are dorsoventrally tall, backswept, and project laterally in dorsal view. The quadratojugal horns display an  asymmetrical triangular morphology with a vertically positioned apex. Only a partial squamosal horn is preserved, but is largely broken.

Life reconstruction of Akainacephalus johnsoni (Image credit: Andrey Atuchin and the Denver Museum of Nature & Science)

The unique anatomical features of Akainacephalus johnsoni indicate a close taxonomic relationship with Nodocephalosaurus kirtlandensis, that clearly distinguish them from other Late Cretaceous Laramidian (although both taxa are temporally separated by nearly three million years). Because both taxa a more closely related to Asian ankylosaurids, the geographic distribution of Late Cretaceous ankylosaurids throughout the Western Interior could be the result of several geologically brief intervals of lowered sea level that allowed Asian ankylosaurid dinosaurs to immigrate to North America several times during the Late Cretaceous. The dispersal of ankylosaurids into Laramidia is coeval with the dispersal of other dinosaur clades, like tyrannosaurids and ceratopsians. The climate gradients and the fluctuations in sea level, may have helped reinforced Campanian provincialism.



Wiersma JP, Irmis RB. (2018) A new southern Laramidian ankylosaurid, Akainacephalus johnsoni gen. et sp. nov., from the upper Campanian Kaiparowits Formation of southern Utah, USA. PeerJ 6:e5016

Arbour, V. M.; Currie, P. J. (2015). “Systematics, phylogeny and palaeobiogeography of the ankylosaurid dinosaurs”. Journal of Systematic Palaeontology: 1–60. doi: 10.1080/14772019.2015.1059985

Ingentia prima, the first giant

Skeletal anatomy of Ingentia prima (From Apaldetti et al., 2018)

During the Late Triassic period numerous extinctions, diversifications and faunal radiations changed the ecosystem dynamics throughout the world. Followed the extinction of rhynchosaurs in most, or all, parts of the world, there was a burst of dinosaurian diversity in the late Carnian, represented by the upper Ischigualasto Formation and coeval units, with mostly carnivorous small- to medium-sized dinosaurs. Then, the long span of the early Norian, from 228.5–218 Ma, during which dicynodonts and sauropodomorph dinosaurs were the major herbivores.

Sauropods evolved from small, gracile, bipedal forms, and it was long thought that acquisition of giant body size in this clade occurred during the Jurassic and was linked to several skeletal modifications. Ingentia prima — literally the “first giant” in Latin — from the Late Triassic of Argentina shed new lights on the origin of gigantism in this group. The holotype, PVSJ 1086, composed of six articulated posterior cervical vertebrae, glenoid region of right scapula and right forelimb lacking all phalanges, has been recovered from the southern outcrops of the Quebrada del Barro Formation, northwestern Argentina. Discovered in 2015 by Diego Abelín and a team led by Cecilia Apaldetti of CONICET-Universidad Nacional de San Juan, Argentina, this new fossil weighed up to 11 tons and measured up to 32 feet (10 meters) long.

Bones of Ingentia prima (Image credit: Cecilia Apaldetti, CONICET-Universidad Nacional de San Juan, Argentina)

Ingentia was unearthed with three new specimens of Lessemsaurus sauropoides. The four dinosaurs belongs to the clade Lessemsauridae, that differs from all other Sauropodomorpha dinosaurs in possessing robust scapulae with dorsal and ventral ends equally expanded; slit-shaped neural canal of posterior dorsal vertebrae; anterior dorsal neural spines transversely expanded towards the dorsal end; a minimum transverse shaft width of the first metacarpal greater than twice the minimum transverse shaft of the second metacarpal; and bone growth characterized by the presence of thick zones of highly vascularized fibrolamellar bone, within a cyclical growth pattern.

The age of the oldest lessemsaurid (mid-Norian) indicates the appearance of an early trend towards large body size at least 15 Myr earlier than previously thought. For a long time, gigantism in eusauropods has been proposed as the result of a complex interplay of anatomical, physiological and reproductive intrinsic traits. For example, the upright position of the limbs has been highlighted as a major feature of the sauropodomorph bauplan considered an adaptation to gigantism. However Lessemsaurids lacked the purported adaptations related to a fully erect forelimb and the marked modifications of the hindlimb lever arms in eusauropods, showing that these features were not strictly necessary for the acquisition of gigantic body size. Another feature interpreted as a key acquisition was the elongated neck. However, lessemsaurids also lacked an elongated neck as they had proportionately short cervical vertebrae, indicating that the neck elongation was not a prerequisite for achieving body sizes comparable to those of basal eusauropods or gravisaurians.



Cecilia Apaldetti, Ricardo N. Martínez, Ignacio A. Cerda, Diego Pol and Oscar Alcober (2018). An early trend towards gigantism in Triassic sauropodomorph dinosaurs. Nature Ecology & Evolution.