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Of the Jurassic Period

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Of the Jurassic Period VOLUMINA JURASSICA, 2016, XIV: 159–164 DOI: A review of the basal tyrannosauroids (Saurischia: Theropoda) of the Jurassic Period Changyu YUN Key words: tyrannosauroid, Saurischia, theropod, Jurassic Abstract. The well supported clade Tyrannosauroidea represents one of the most basal coelurosaurian theropods. Given that current fossil records of earliest coelurosaur theropods are extremely scarce, basal-most tyrannosauroid materials are key to understanding the origin and diversification of coelurosaurs. Here, I present a brief overview of currently known basal tyrannosauroids of Jurassic age, discussing their systematics and distribution. The currently oldest known Jurassic tyrannosauroids are from Europe continent, possibly suggesting the European origin of the superfamily. INTRODUCTION the oldest tyrannosauroid material plays an important role in understanding coelurosaurian origin and diversification. The Coelurosauria is the only dinosaurian clade which This paper aims to present a review of the currently survives today as birds, and it is one of the most diverse known Jurassic tyrannosauroids, and a general description theropod clades comprising many herbivorous and carnivo- of each taxon or subclade. My hope is that this paper will be rous taxa. Therefore, the diversification of the Coelurosauria a helpful guide to understanding tyrannosauroid or coeluro- is one of the most interesting subject of the evolutionary his- saurian origin and/or distribution. tory of the dinosaurs. Unfortunately, most of the oldest known coelurosaurian fossils are largely fragmentary so it is POSSIBLE CASES currently very dificult to understand the clade’s diversifica- tion. One putative record from the Early Jurassic of China might be the oldest known therizinosaurian (Zhao, Xu, Iliosuchus incognitus (Huene, 1932) 1998; Barrett, 2009) but its exact age and taxonomic status Age: Middle Bathonian, Middle Jurassic are controversial (Kirkland, Wolfe, 2001; Kirkland et al., 2005). Occurrence: Stonesfield Slate, England The Tyrannosauroidea is considered as a basal-most clade of Coelurosauria by a majority of recent phylogenetic Comments: Only 3 small ilia are the currently known analyses (e.g. Loewen et al., 2013; Brusatte et al., 2014) but material of this taxon. Though this material is very small in some analyses found them to be more derived than the size (being 9 to 10 cm long), a vertical iliac ridge is clearly Compsognathidae (e.g. Rauhut et al., 2010; Novas et al., present similar to other tyrannosauroids. So this taxon was 2013). What is clear is that the tyrannosauroids represent traditionally allied with the tyrannosauroids (Galton, 1976), a clade close to the origin of the Coelurosauria. Therefore though the fact that some other theropods belonging to the 1 Vertebrate Paleontological Institute of Incheon, Incheon 21974, Republic of Korea, Biological Sciences, Inha University, Incheon 22212, Republic of Korea; e-mail: [email protected]. 160 C. YUN Tetanurae such as Megalosaurus or Piatnitzkysaurus have tion. However, the ilium of Tanycolagreus is currently un- similar feature makes this referral uncertain (Benson, 2009). known so the direct comparison of the taxa is currently im- Recent studies have considered Iliosuchus to be an indeter- possible (the distinctive bones of Stokesosaurus being the minate avetheropod or a juvenile Megalosaurus (Benson, ilia). If this taxon indeed forms the clade Coeluridae and 2009; Carrano et al., 2012) but such fragmentary material is belongs to the tyrannosauroid, it is probably close to the not enough for accurate classification. base (Senter, 2007). The taxon had a large, enlongated skull If Iliosuchus is indeed a tyrannosauroid, it would be the and long legs, and gracile general morphology. earliest genus in the clade or maybe even in the whole Coe- lurosauria itself (Rauhut, 2003). CERTAIN CASES Coeluridae (Marsh, 1881) Definition: The most inclusive clade containing Coelu­ Proceratosauridae (Averianov et al., 2010) rus fragilis but not Proceratosaurus bradleyi, Tyrannosau­ rus rex, Allosaurus fragilis, Compsognathus longipes, Orni­ Definition: A node based taxon including Proceratosau­ thomimus edmontonicus and Deinonychus antirrhopus rus bradleyi and Kileskus aristotocus, their most recent (Hendrickx et al., 2015) common ancestor and all its decendants (Averianov et al., Comments: Whether or not this clade is monophyletic is 2010). controversial, and some taxon or whole members of the Comments: Rauhut et al. (2010) first suggested the defi- clade might not belong to the Tyrannosauroidea at all (Choi- nition for the Proceratosauridae as “all theropods that are niere et al., 2014). Many of the recent analyses found that more closely related to Proceratosaurus than to Tyranno­ this clade belongs to the basal tyrannosauroids (e.g. Senter, saurus, Allosaurus, Compsognathus, Coelurus, Ornithomi­ 2007; Brusatte et al., 2014). There are currently two genera mus, or Deinonychus”. Averianov et al. (2010) pointed out in this clade, which are Coelurus and Tanycolagreus. This that this definition suggested by Rauhut et al. (2010) does clade comprises small carnivorous coelurosaurs which are not meet ICZN’s requirements, so their “Proceratosauridae” likely have hunted small prey. is a nomen nudum. There are currently 5 valid taxa belong to this clade, based on the parsimony analysis of Brusatte, Carr Coelurus fragilis (Marsh, 1879) (2016) and 3 of them are from the Jurassic period. This clade comprises small to large basal tyranno- Age: Middle, Late Kimmeridgian, Late Jurassic sauroids (Brusatte, Carr, 2016) characterized by crests on Occurrence: Morrison Formation, Wyoming, USA their head. Since the crests of these dinosaurs are thin, they might have served as display organs. Comments: The exact position of this taxon’s theropoda phylogenetical tree is uncertain. Some consider Coelurus as Guanlong wucaii (Xu et al., 2006) more derived than the tyrannosauroids (Turner et al., 2007), or even than the basal maniraptoran (Zanno, 2010). If this Age: Oxfordian, Late Jurassic taxon indeed forms a clade Coeluridae and belongs to the Occurrence: Shishugou Formation, Xinjiang, China tyrannosauroid, it is probably close to the base (Senter, 2007). This taxon had a relatively long neck and trunk, and Comments: This is a taxon famous for its highly distinc- a long slender hindlimb. This suggests it was a small, gracile tive crest on its skull (Fig. 1A). Carr (2006) suggested that theropod with great speed ability. this taxon is actually a Carnosauria sister to Monolophosau­ rus or even possibly synonymous with it. This was support- Tanycolagreus topwilsoni (Carpenter et al., 2005) ed by Paul (2010) as he described this taxon as Monolopho­ saurus wucaii. However, no other analyses have found them Age: Middle, Late Kimmeridgian, Late Jurassic as closely related (e.g. Loewen et al., 2013; Brusatte, Carr, Occurrence: Morrison Formation, Wyoming, USA 2016). As this taxon had long forelimbs with a large manus and a small skull with small teeth, it is likely that its fore- Comments: This taxon could be synonymous with the limbs played an important role in hunting unlike the more tyrannosauroid Stokesosaurus clevelandi, since they are derived tyrannosauroids, as their forelimbs are very short- similar sized coelurosaurs from the same geological forma- ened. A review of the basal tyrannosauroids (Saurischia: Theropoda) of the Jurassic Period 161 Fig. 1. Skull reconstructions of the two Jurassic proceratosaurids A. Guanlong. B. Proceratosaurus. Reconstructions by Tracy Ford Proceratosaurus bradleyi Aviatyrannis jurassica (Rauhut, 2003) (Woodward, 1910; Huene, 1926) Age: Early Kimmeridgian, Late Jurassic Age: Middle Bathonian, Middle Jurassic Occurrence: Alcobaca Formation, Portugal Occurrence: Great Oolite, England Comments: This is one of the oldest tyrannosauroids Comments: Once considered as a taxon closely related ever found. The holotype ilium was once originally referred to Ceratosaurus (Huene, 1926), this is now considered as to Stokesosaurus, though later study described it as a new the currently oldest definite tyrannosauroid and coelurosaur genus (Rauhut, 2003). However, it is probable that this ge- of all time (Rauhut et al., 2010). The currently only known nus is in fact a Portuguese species of Stokesosaurus. Some skull has a small “horn” on the end of its snout, but this is material such as an ilium (now lost) and an isolated tooth probably the trace of a broken delicate crest similar that pre- from the North American Morrison Formation were once re- sent in Guanlong (Rauhut et al., 2010; Fig. 1B). It had ferred to Stokesosaurus, but might actually belong to Avia­ a small, subrectangular head and it probably had a similar tyrannis (Rauhut, 2003). These findings, and the cases of lifestyle to Guanlong. Portuguese Allosaurus or Torvosaurus (Mateus et al., 2006), support the Kimmeridgian land connections between North Kileskus aristotocus (Averianov et al., 2010) America and Europe. Age: Bathonian, Middle Jurassic Juratyrant langhami Occurrence: Itat Formation, Russia (Benson, 2008; Brusatte, Benson, 2013) Comments: This is only known from very fragmentary Age: Early Tithonian, Late Jurassic cranial material, but this is very similar to Proceratosaurus. Occurrence: Kimmeridge Clay, England This is one of the most basal coelurosaur and tyrannosauroid genera, and one of the oldest records of these clades. This Comments: Once thought to be a species of Stokeso­ taxon probably hunted small vertebrates such as fishes,
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