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Dinosauria R. Owen 1842 [M Dinosauria Dinosauria Dinosauria R. Owen 1842 [M. C. Langer, F. E. Novas, J. S. Bittencourt, M. D. Ezcurra, and J. A. Gauthier], converted clade name Registration Number: 194 al. (2008; supplementary Fig. 2) was selected as the primary reference phylogeny. Note that the Definition: The smallest clade contain- definition uses clade addresses (e.g., Theropoda: ing Iguanodon bernissartensis Boulenger in Megalosauroidea) from Lloyd et al. (2008) to Beneden 1881 (Ornithischia/Euornithopoda) facilitate finding specifiers (e.g., Megalosaurus Megalosaurus bucklandii Mantell 1827 bucklandii) on this densely branched supertree. (Theropoda/Megalosauroidea) and Cetiosaurus Only species, not more inclusive taxa, can be oxoniensis Phillips 1871 (Sauropodomorpha). specifiers in this kind of definition. We accord- This is a minimum-clade definition. Abbreviated ingly do not regard these additional taxon definition: min∇ (Iguanodon bernissartensis names as parts of the formal definition, nor do Boulenger in Beneden 1881 & Megalosaurus we mean to endorse their use for these clades. bucklandii Mantell 1827 & Cetiosaurus oxoni- ensis Phillips 1871). Composition: According to most phylogenetic hypotheses, Dinosauria is composed of two pri- Etymology: Derived from the ancient Greek: mary subclades—Ornithischia and Saurischia δεινός = fearful, terrible; σαύρος = lizard-like (this volume)—with the latter composed of (Liddell and Scott, 1882); seemingly in reference Sauropodomorpha (this volume) and Theropoda to the large size of the fossils originally assigned (including Aves, this volume) (e.g., Gauthier et to this group. Indeed, Owen (1842) translated al., 1989; Sereno, 1999; Ezcurra, 2006; Langer δειυός as “fearfully great,” conceivably employ- and Benton, 2006; Irmis et al., 2007; Lloyd et ing the term as “the quality of objects which, al., 2008; Apaldetti et al., 2011; Martínez et al., from their vastness, magnitude, etc., inspire 2011; Sereno et al., 2013; Niedźwiedzki et al., fear, awe, reverence, power, etc.” (Pickering, 2014; Pretto et al., 2015; Cabreira et al., 2016). 1873; in Creisler, 1996). There is some contention over relations of a few very early species—e.g., some have proposed Reference Phylogeny: There are a plethora of that herrerasaurs (e.g., Gauthier et al., 1989; phylogenies depicting relationships within and Novas, 1992; Fraser et al., 2002) and silesaurids among various parts of the host of extinct clades (e.g., Langer and Ferigolo, 2013) are, respec- that diverged from along the avian stem dur- tively, outside and within Dinosauria as defined ing the Mesozoic. More comprehensive stud- here. The composition and relations of those ies (e.g., Sereno, 1999) are rare. No one has yet primary dinosaurian subclades have otherwise attempted to simultaneously analyze a broad been remarkably stable, and alternative topolo- range of species from across the avian total gies proposed among basalmost branches (e.g., clade, which includes more than 11,000 species Cooper, 1985; Baron et al., 2017; but see Langer ranging from mid-Triassic to Recent (see Aves, et al., 2017) do not alter the circumscription of this volume). Because it contains the largest Dinosauria as that taxon name is defined here. sample of non-avian dinosaurs, the phylogeny Starting with the compilation of Weishampel derived from the supertree analysis of Lloyd et et al. (2004), Benton (2008) identified 726 Dinosauria valid dinosaur species of Mesozoic age. This pelvic and hind-limb modifications thought count includes early winged dinosaurs such to reflect acquisition of an upright striding as Archaeopteryx lithographica, with early bipedal stance and gait (e.g., Gauthier et al., avialans comprising about 10% of Mesozoic 2011) figured prominently in the diagnosis of dinosaur species diversity. The International Dinosauria from Owen’s time to the dawn of Ornithological Union recognizes 10,672 extant the Hennigian (phylogenetic) era (e.g., Bakker species of Aves (Gill and Donsker, 2017); hence, and Galton, 1974; Gauthier, 1986). This col- there are accordingly at least 11,398 species lection of apomorphies remains diagnostic of currently assigned to Dinosauria as that taxon Dinosauria relative to the last ancestor it shared name is defined here. with Alligator mississippiensis (Archosauria, this Starrfelt and Liow (2016) used a Poisson volume). Nevertheless, these distinctly bird-like sampling model to more accurately estimate “dinosaurian” synapomorphies likely originated species richness among Mesozoic dinosaurs, before the divergence amongst Ornithischia, more than doubling the estimated diversity to Theropoda, and Sauropodomorpha (= Dinosauria around 2,000 species. Considering the episodic as defined here). A growing understanding of nature of deposition on land, the dearth of species diversity and anatomical disparity of dinosaur-bearing beds of Mesozoic age around Triassic dinosaurs and their close kin reveals the world, and the uneven geographic distribu- that many of these traditional “dinosaurian” tion of palaeontologists prospecting for them, apomorphies did indeed evolve earlier on the this is still likely to be an underestimate. Extant avian stem (e.g., Novas, 1996; Sereno, 1999; dinosaurs (Aves) may be far more diverse in Langer and Benton, 2006; Ezcurra, 2006; Irmis terms of species, doubtless reflecting their rel- et al., 2007; Nesbitt, 2011; Cabreira et al., 2016; atively small body size, but large-bodied stem Nesbitt et al., 2017). avians (traditional “dinosaurs”) are far more Several authors have reviewed the synapo- disparate morphologically. Among the aston- morphies of Dinosauria (e.g., Brusatte et al., ishing variety of dinosaurs alive at the end of 2010; Langer et al., 2010; Nesbitt et al., 2010; the Cretaceous, only the very earliest branches Nesbitt, 2011; Nesbitt et al., 2012; Cabreira of crown Aves survived the Chicxulub impact et al., 2016; Baron et al., 2017). They indicate at 66 Ma (Longrich et al., 2011; see Aves this that the precise diagnosis of the clade turns on volume). whether it contains silesaurids (Dzik, 2003; Langer and Ferigolo, 2013; Niedźwiedzki et Diagnostic Apomorphies: With respect to al., 2014; Cabreira et al., 2016) or not. In addi- other “reptiles”, Owen (1842) characterized his tion to some uncertainty from character con- three species of Dinosauria—now recognized flict, incomplete preservation in Triassic taxa as two ornithischians and a theropod sauris- (e.g., Nyasasaurus parringtoni; Nesbitt et al., chian—by their large size and unusual combina- 2012) remains a serious problem in that impor- tion of osteological traits, including a distinctive tant characters of the skull and manus are very sacral construction and an upright limb posture poorly known among non-dinosaur dinosauro- resembling those of “bulky terrestrial mam- morphs (Langer et al., 2013). mals.” Owen’s conception was developed in the A list of traditional dinosaur synapomor- pre-Darwinian era and was burdened further phies based on reviews noted above is presented by there being so few extinct dinosaurs known below. It is mainly composed of uncontrover- in the mid-nineteenth century. Nevertheless, sial dinosaur apomorphies, but it also includes 1210 Dinosauria some apomorphies with uncertain distributions with ilium and pubis separated by broad con- among a few non-dinosaur dinosauromorphs cave surface** (Irmis et al., 2007); (17) ischia from the Triassic (indicated by *). Others are with extensive contact between their shafts‡ not present in all early dinosaurs, but might (Nesbitt, 2011); (18) femoral head distinctly be synapomorphic for the clade depending off-set at a sharp angle from the shaft‡ (Bakker on assumptions about the frequency of inde- and Galton, 1974); (19) femur with dorsolat- pendent origin vs. secondary loss in evolution eral trochanter‡ (Gauthier, 1986); (20) fourth (indicated by **). Finally, some are also known trochanter with distal margin forming steeper in Nyasaurus parringtoni (indicated by †) and angle relative to femoral shaft** (Langer and in some silesaurids (indicated by ‡), taxa that Benton, 2006); (21) tibia cnemial crest arcs cra- are either sister to, or just inside of, Dinosauria. niolaterally‡ (Benton and Clark, 1988); (22) They nevertheless suffice to distinguish dino- caudal surface of tibia with proximodistally saurs from more distant outgroups near the base oriented ridge** (Nesbitt, 2011); (23) flange on of the avian stem such as Lagosuchus talampay- distal portion of tibia overlapping caudally the ensis (= Marasuchus lilloensis) and Lagerpeton ascending process of the astragalus‡ (Novas, chanarensis (authorship credited to first explicit 1992); (24) cranial edge of proximal portion identification as an apomorphy): (1) postfron- of fibula tapers to point and arches medially‡ tal absent* (Gauthier, 1986); (2) jugal with (Nesbitt, 2011); (25) broad astragalar ascending bifurcated quadratojugal process* (Sereno and process‡ (Gauthier, 1986); (26) astragalus with Novas, 1992); (3) supratemporal fossa extends fibular facet occupying less than 1/3 of trans- onto frontal rostral to supratemporal fenestra verse width‡ (Langer and Benton, 2006); (27) (Gauthier, 1986); (4) post-temporal opening concave articular surface of the fibula in the much smaller than foramen magnum (Sereno calcaneum‡ (Novas, 1996); (28) calcaneal tuber and Novas, 1993); (5) exoccipitals fail to meet absent** (Gauthier, 1986); (29) distal tarsal IV on the floor of endocranial cavity* (Nesbitt, with flat proximal surface (Nesbitt, 2011); (30)
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