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A New Clade of Asian Late Cretaceous Long-Snouted Tyrannosaurids

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A New Clade of Asian Late Cretaceous Long-Snouted Tyrannosaurids ARTICLE Received 30 Dec 2013 | Accepted 1 Apr 2014 | Published 7 May 2014 DOI: 10.1038/ncomms4788 A new clade of Asian Late Cretaceous long-snouted tyrannosaurids Junchang Lu¨1,2, Laiping Yi3, Stephen L. Brusatte4, Ling Yang5, Hua Li3 & Liu Chen3 The iconic tyrannosaurids were top predators in Asia and North America during the latest Cretaceous, and most species had deep skulls that allowed them to generate extreme bite forces. Two unusual specimens of Alioramus from Mongolia seem to indicate a divergent long-snouted body plan among some derived tyrannosaurids, but the rarity and juvenile nature of these fossils leaves many questions unanswered. Here, we describe a remarkable new species of long-snouted tyrannosaurid from the Maastrichtian of southeastern China, Qianzhousaurus sinensis. Phylogenetic analysis places Qianzhousaurus with both species of Alioramus in a novel longirostrine clade, which was geographically widespread across latest Cretaceous Asia and formed an important component of terrestrial ecosystems during this time. The new specimen is approximately twice the size as both Alioramus individuals, showing that the long-snouted morphology was not a transient juvenile condition of deep-snouted species, but a characteristic of a major tyrannosaurid subgroup. 1 Institute of Geology, Chinese Academy of Geological Sciences, Beijing 100037, China. 2 Key Lab of Stratigraphy and Paleontology, Ministry of Land and Resources of China, Beijing 100037, China. 3 Ganzhou Museum, Ganzhou 341000, Jiangxi Province, China. 4 School of GeoSciences, Grant Institute, University of Edinburgh, The King’s Buildings, West Mains Road, Edinburgh EH9 3JW, UK. 5 Geological Museum of Jiangxi Province, Nanchang 330002, Jiangxi Province, China. Correspondence and requests for materials should be addressed to J.L. (email: [email protected]). NATURE COMMUNICATIONS | 5:3788 | DOI: 10.1038/ncomms4788 | www.nature.com/naturecommunications 1 & 2014 Macmillan Publishers Limited. All rights reserved. ARTICLE NATURE COMMUNICATIONS | DOI: 10.1038/ncomms4788 yrannosauroids are among the most iconic animals in the Definition. The most inclusive clade containing A. fossil record. This group of large-bodied carnivorous remotus9, but not Tyrannosaurus rex18, Albertosaurus Tdinosaurs, which includes Tyrannosaurus rex and its sarcophagus18 or Proceratosaurus bradleyi19. Olshevsky20 closest relatives, were the apex predators in Asia and North used the name Alioramini to refer to a monotypic tribe-level America during the final 20 million years of the Cretaceous, and Linnean taxon for A. remotus, but we first use this name to were at the top of the food chain in these regions when a bolide refer to a novel clade of long-snouted tyrannosauroids. impact (and perhaps other environmental changes) suddenly Diagnosis. Alioramini differs from all other tyrannosaur- ended the age of non-avian dinosaurs 66 million years ago1–3. oids in possessing an extremely long and low skull, in which The vast majority of derived tyrannosauroids (tyrannosaurids) the snout comprises 2/3 or more of the total skull length; an are characterized by deep skulls, thick teeth and large jaw anteroposteriorly elongate maxillary fenestra (ratio of muscles, features which allowed these dinosaurs to employ bite maximum anteroposterior length to maximum dorsoventral forces strong enough to crunch through the bones of even large- depth greater than 1.9); a series of pronounced, discrete bodied prey species4–7. This unusual body plan is thought to have rugosities on the nasal; 18 or more dentary teeth. Full been the key to tyrannosaurid success8. synapomorphy is listed in Supplementary Note 3. A few enigmatic specimens from the Late Cretaceous Included species. A. altai10, A. remotus9, Qianzhousaurus (Maastrichtian) of Asia, however, suggest that a very different sinensis gen. et sp. nov. type of tyrannosaurid may have coexisted with these muscular, deep-skulled species during the final few million years of dinosaur Qianzhousaurus sinensis gen. et sp. nov. evolutionary history. The holotypes of Alioramus remotus9 and Alioramus altai10,11 from Mongolia are both gracile animals with Etymology. From Qianzhou, an ancient name of Ganzhou, extremely long skulls relative to body size, with elongated snouts where the fossils were discovered. Sin (Greek) China, refers and thin teeth. It has been suggested recently that these fossils to the specimen found from China. may represent a peculiar body plan among a handful of derived Holotype. A nearly complete skull and most of the left lower tyrannosaurids, indicative of different feeding styles and a lower jaw (GM F10004-1); nine cervical vertebrae (GM F10004-2); position in the food chain relative to their coexisting deep-skulled three anterior dorsal vertebrae (GM F10004-3); 18 middle cousins10,11. There is concern, however, because only two posterior caudal vertebrae (GM F10004-4); a complete specimens of these long-snouted tyrannosaurids have been right scapulocoracoid (GM F10004-5); part of the left recovered, and both are juveniles, leading some workers to scapulocoracoid (GM F10004-6); both partial ilia, a left worry that they may represent transient longirostrine growth femur and left tibia, a left partial fibula, a left astragalus stages during the ontogeny of deeper-skulled species12. The and calcaneum and left metatarsals (GM F10004-7, 8). The extreme rarity of these animals, and their restriction to a narrow specimens, which were found associated and represent a time interval in Mongolia, makes it difficult to test these single individual, are housed at the Ganzhou Museum, hypotheses. Ganzhou City, Jiangxi Province, China. Measurements are We report the discovery of an aberrant new tyrannosaurid provided in Supplementary Tables 1-5. from the Late Cretaceous of southeastern China that is known Locality and horizon. Nanxiong Formation (Upper Cretac- from a remarkably complete and well-preserved skull, along with eous, Maastrichtian) of Longling Town, Nankang, Ganzhou much of the vertebral column and appendicular skeleton. Other City, Jiangxi Province, China21. dinosaurs from the same horizon include the somphospondylan Diagnosis. Long-snouted tyrannosaurid tyrannosauroid sauropod Gannansaurus sinesis13 and several oviraptorosaurian with the following autapomorphies: large, discrete, win- theropods (Banji long14; Ganzhousaurus nankangensis15; dow-like pneumatic fenestra on the ascending ramus of the Jiangxisaurus ganzhouensis16; Nankangia jiangxiensis17). The maxilla; extremely reduced premaxilla (maximum antero- new tyrannosaurid was the top predator in this terrestrial posterior length of the main body of the bone is B2.2% of ecosystem. This tyrannosaurid possesses the characteristic long the total basal skull length, whereas other tyrannosaurids and low snout of both species of Alioramus, along with other have measures of B4.3–4.6%); absence of a pronounced features uniquely shared by these longirostrine taxa relative to vertical ridge on the lateral surface of the ilium (possibly deep-skulled tyrannosaurids. As a result, the three species are shared with Raptorex kriegsteini8). grouped together in their own distinct long-snouted clade by a phylogenetic analysis. The new species is larger (twice the body mass) and more skeletally mature than either specimen of Comparative description. The skull of Qianzhousaurus is well Alioramus, which demonstrates that a long-snouted morphology preserved and nearly complete, missing only a small portion of was present in larger individuals and not likely to be a transient the orbital region on the right side (Fig. 1). It is exceptionally long juvenile feature. The discovery of this species in southern China and low compared with other large-bodied tyrannosauroids like also shows that the long-snouted carnivores were geographically Tyrannosaurus and Albertosaurus12,22,23, but very similar in widespread across Asia during the latest Cretaceous, and formed proportions to the longirostrine skulls of both species of an important (but hitherto overlooked) component of terrestrial Alioramus9–11.AsinAlioramus, the preorbital (snout) region of ecosystems during this time. the skull is elongated such that it is B70% of the length of the entire skull (Fig. 1), whereas this is B60% in other large tyrannosaurids. Also as in Alioramus, the individual bones of the Results snout such as the maxilla and nasal (but not premaxilla) are Systematic palaeontology. elongated, whereas those of the posterior skull are of similar Dinosauria Owen 1842 proportions to other large tyrannosaurids. The skull of Theropoda Marsh 1881 Qianzhousaurus is 90 cm long anteroposteriorly when measured Tetanurae Gauthier 1986 between the anterior edge of the premaxilla and the Coelurosauria von Huene 1914 posteroventral corner of the quadrate in lateral view, compared Tyrannosauroidea Osborn, 1905 with B63 cm in the holotype of A. altai and a similar Alioramini Olskevsky, 1995, clade nov. reconstructed measurement for the type of A. remotus10. 2 NATURE COMMUNICATIONS | 5:3788 | DOI: 10.1038/ncomms4788 | www.nature.com/naturecommunications & 2014 Macmillan Publishers Limited. All rights reserved. NATURE COMMUNICATIONS | DOI: 10.1038/ncomms4788 ARTICLE lh lpr ph pcr nrug arp n sq en or pm la po antor.f mf ltf pmf m j jcp qj q Figure 1 | Holotype skull of Qianzhousaurus sinensis gen. et sp. nov. (GM F10004-1). (a) Photograph in left lateral view, (b) line drawing in left lateral view, (c) photograph in dorsal view, (d) photograph in ventral view. antor.f, antorbital
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