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Zoological Journal of the Linnean Society SCELIDOSAURUS HARRISONII OWEN, 1861 FROM THE EARLY JURASSIC OF DORSET, ENGLAND: PART 1. CRANIAL ANATOMY Journal: Zoological Journal of the Linnean Society Manuscript ID ZOJ-10-2018-3496.R2 Manuscript Type:ForOriginal Review Article Only monograph < Taxonomy, alpha taxonomy < Taxonomy, Reptilia < Taxa, Dinosauria < Taxa, ontogeny < Growth, vertebrate palaeontology < Keywords: Palaeontology, Jurassic < Palaeontology, English South Coast < Geography, mechanical function < Biomechanics, cranial osteology < Anatomy The skull of Scelidosaurus harrisonii is described. Scelidosaurus is an early (late Sinemurian) armoured ornithischian dinosaur from Charmouth, Dorset. It is the first largely complete and articulated dinosaur ever discovered. Much of the exterior of the skull has a textured patina of exostotic bone that anchored keratinous scales. A small edentulous beak precedes a row of five premaxillary teeth, and there would have been a minimum of 22 maxillary teeth and 27 dentary teeth in the largest known individuals. Tooth wear is discontinuous along the dentition and jaw action appears to have been tightly constrained. Abstract: There is no physical evidence of a predentary but its existence is inferred from the structure of the dentary. A sclerotic ring was present. A prominent supraorbital brow ridge overhangs the orbital cavity. A crater- shaped osteoderm is attached to the lateral surface of the postorbital. Paired osteodermal horns project dorsally from the occiput, and long bladed styloid bones project obliquely from the posterior of the skull. The nasal chambers are roofed by epivomers that are unique to this taxon. A vertical, conical epipterygoid is attached to the pterygoid. A deep pit on the posterior surface of the quadrate suggests the existence of remnant cranial pneumatism. Page 1 of 215 Zoological Journal of the Linnean Society 1 2 3 4 SCELIDOSAURUS HARRISONII OWEN, 1861 FROM THE EARLY JURASSIC 5 6 OF DORSET, ENGLAND: PART 1. CRANIAL ANATOMY 7 8 9 10 11 12 DEDICATION 13 14 I wish to offer a personal note of thanks to Ron Croucher (former Head of 15 16 the Palaeontological Preparation-Conservation Laboratory at the Natural 17 History Museum, London) by dedicating this monograph to him. Ron 18 19 committed an enormous amount of time and effort, across several 20 21 decades, to the laboriousFor andReview painstaking Onlypreparation of the lectotype 22 skull and postcranial skeleton of Scelidosaurus (NHMUK R1111). His 23 24 having to watch as numerous researchers subsequently mishandled (and 25 26 damaged) some of these bones, hard-won with such skill and dedication, 27 must have severely tested his patience and goodwill. Ron is indeed a 28 29 wonderfully kind and incredibly patient man. While superintending the 30 31 museum laboratory he could always find time to give advice and 32 assistance to ignorant and cack-handed PhD students who, from time to 33 34 time, ‘messed about’ with fossil specimens in his laboratory … I know this 35 36 all too well, I was one of them. 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Zoological Journal of the Linnean Society Page 2 of 215 1 2 3 4 EXTENDED ABSTRACT 5 6 Scelidosaurus harrisonii Owen, 1861 is an early (late Sinemurian) 7 armoured ornithischian dinosaur whose remains have, to date, only been 8 recovered from a restricted location on the south coast of Dorset 9 (Charmouth), England. This dinosaur has been known since 1859, but 10 only on the basis of a partial description found in two articles published in 11 12 the early 1860s by Richard Owen. The original material, discovered in 13 1858, comprised the majority of the skull and its associated postcranial 14 skeleton, and represents the first ever more or less complete dinosaur 15 discovered. In addition to the original material, a number of further 16 discoveries have been made at Charmouth; these latter supplement the 17 information that can be gleaned from the original specimen. This article 18 describes the skull of Scelidosaurus. 19 20 The external surface of individual skull bones in ontogenetically relatively 21 mature individualsFor displays Review exostoses, a patina Only of fibrous or granular- 22 textured bone that anchored an external shielding of keratinous scales. 23 There is a small, edentulous rostral beak, behind which is found a row of 24 five heterodont premaxillary teeth. There is a minimum of 22 maxillary 25 teeth and 27 dentary teeth in jaws of the largest well-preserved 26 individuals known to date. Both dentitions (upper and lower) are bowed 27 28 medially and are sinuous longitudinally. Maxillary and dentary crowns are 29 tilted lingually on their roots, trapezoidal in outline and have crenellate 30 (coarsely denticulate) margins. Adjacent crowns of teeth have mesio- 31 distally (anteroposteriorly) expanded bases that overlap slightly and are 32 consequently arranged en echelon. The dentitions are flanked by deep 33 cheek pouches. Tooth abrasion is usually discontinuous along the 34 dentition. In one individual nearly all teeth seem to be fully emerged and 35 there is little evidence of abrasion. There is no physical evidence of a 36 37 predentary, but the presence of this (typically ornithischian) element may 38 be inferred from the structure of the symphyseal region of the dentary. 39 The external narial and antorbital fenestrae are comparatively small, 40 whereas the orbit and temporal fenestrae are large and open. A sclerotic 41 ring was undoubtedly present and supported the eyeball, but it is too 42 poorly preserved to allow it to be reconstructed with accuracy. A 43 prominent supraorbital brow ridge overhangs the orbit. There are three 44 osteoderms: palpebral, middle supraorbital and posterior supraorbital, 45 sutured to the dorsal margin of the orbit. The occiput provides an area for 46 47 attachment of a pair of curved, keratin-sheathed, osteodermal horns. 48 Epistyloid bones project from the ventrolateral region of the braincase; 49 their distal ends flank the anterolateral region of the neck. Rugose facets 50 on either side of the basioccipital are suggested to have provided 51 attachment sites for the epistyloid bones. Internally, the skull has a 52 53 deeply vaulted snout and the nasal chambers are roofed by what are here 54 named epivomer bones that appear to have been sutured to the 55 dorsolateral edges of the vomers. Unusually, among dinosaurs generally, 56 an epipterygoid is preserved attached to the dorsolateral surface of the 57 pterygoid; there is no obvious point of articulation for the epipterygoid 58 against the lateral wall of the braincase. There is also a narrow, slot-like 59 pocket on the medial wall of the quadrate wing of the pterygoid. A deep 60 2 Page 3 of 215 Zoological Journal of the Linnean Society 1 2 3 pit on the posterior surface of the quadrate of an immature specimen is 4 suggestive of the existence of a remnant of cranial pneumatism; this pit 5 becomes occluded in larger, more mature specimens. 6 7 8 9 NOTE CONCERNING THE LECTOTYPE 10 11 In 1955 Bill (William Elgin) Swinton (1900-1994) approved a programme 12 13 of work for Arthur Rixon, focused upon the acetic acid-mediated 14 15 preparation of a split marlstone (=argillaceous limestone) nodule 16 17 (‘dogger’) that had been collected by J.F. Jackson from the foreshore 18 beneath Black Ven, Charmouth and acquired by the Natural History 19 20 Museum (London). The dogger contained an incomplete articulated 21 For Review Only 22 postcranial skeleton of a small individual of Scelidosaurus. Rixon (1949) 23 had experimented with the use of dilute mineral acids to prepare fossil 24 25 vertebrates and with Harry Toombs (Toombs & Rixon 1950) had 26 27 developed the ‘Transfer Method’ for preparing such fossils in the round 28 (Toombs & Rixon 1959). The scelidosaur project resulted in the successful 29 30 extraction of this small dinosaur skeleton (Rixon 1968) and the 31 32 development of further plans to prepare the lectotype skeleton; these 33 plans were however abandoned after the departure of Swinton. 34 35 Alan Charig (1927-1997) was appointed curator in succession to Bill 36 37 Swinton in 1961. He eventually revisited Swinton’s original project, in the 38 39 light of the success of the technique applied to the small scelidosaur 40 skeleton, and so the extraction of the entire skeleton of the lectotype of 41 42 Scelidosaurus harrisonii (NHMUK R1111) began, under the supervision of 43 44 Ron Croucher. The latter skeleton was preserved in a more-or-less 45 contiguous series of substantial marlstone blocks. As the work continued 46 47 Croucher noticed the fragility of some of the acid-extracted bones and 48 49 modified the original preparation protocol by adding a phosphate buffer to 50 the acid solution. This latter modification reduced the extent of acid- 51 52 induced erosion of the fabric of the fossil bones (and their consequent 53 54 extreme fragility) that has seriously jeopardised the earlier extracted 55 bones – in particular the skull. Charig announced his intention to describe 56 57 Scelidosaurus but he had made little progress by the time of his death. 58 59 60 3 Zoological Journal of the Linnean Society Page 4 of 215 1 2 3 REPOSITORY ABBREVIATIONS: BRSMG - Bristol City Museum UK, CAMSM 4 5 – Sedgwick Museum, University of Cambridge UK; DORCM - Dorset 6 7 County Museum, Dorchester, Dorset UK; FMNH – Field Museum of Natural 8 History, Chicago USA; IVPP – Institute of Vertebrate Palaeontology and 9 10 Palaeoanthropology, Beijing, China; LYMPH – The Lyme Regis (Philpot) 11 12 Museum, Dorset UK; NHMUK – Natural History Museum, London British 13 Museum; MNA – Museum of Northern Arizona, Flagstaff USA; SGDS – St 14 15 George Dinosaur Discovery Site, Utah USA; SGWG – Sektion Geologische 16 17 Wissenshaften der Ernst-Moritz-Arndt-Universität Griefswald, Germany; 18 TMM – Texas Memorial Museum (Vertebrate Paleontology Laboratory of 19 20 the University of Texas at Austin), USA; UCMP – University of California 21 For Review Only 22 Museum of Paleontology, Berkeley, USA; ZDM – Zigong Dinosaur 23 Museum, Zigong, China.
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  • A Stable Isotopic Investigation of Resource Partitioning Among Neosauropod Dinosaurs of the Upper Jurassic Morrison Formation

    A Stable Isotopic Investigation of Resource Partitioning Among Neosauropod Dinosaurs of the Upper Jurassic Morrison Formation

    A stable isotopic investigation of resource partitioning among neosauropod dinosaurs of the Upper Jurassic Morrison Formation Benjamin T. Breeden, III SID: 110305422 [email protected] GEOL394H University of Maryland, College Park, Department of Geology 29 April 2011 Advisors: Dr. Thomas R. Holtz1, Jr., Dr. Alan Jay Kaufman1, and Dr. Matthew T. Carrano2 1: University of Maryland, College Park, Department of Geology 2: National Museum of Natural History, Department of Paleobiology ABSTRACT For more than a century, morphological studies have been used to attempt to understand the partitioning of resources in the Morrison Fauna, particularly between members of the two major clades of neosauropod (long-necked, megaherbivorous) dinosaurs: Diplodocidae and Macronaria. While it is generally accepted that most macronarians fed 3-5m above the ground, the feeding habits of diplodocids are somewhat more enigmatic; it is not clear whether diplodocids fed higher or lower than macronarians. While many studies exploring sauropod resource portioning have focused on differences in the morphologies of the two groups, few have utilized geochemical evidence. Stable isotope geochemistry has become an increasingly common and reliable means of investigating paleoecological questions, and due to the resistance of tooth enamel to diagenetic alteration, fossil teeth can provide invaluable paleoecological and behavioral data that would be otherwise unobtainable. Studies in the Ituri Rainforest in the Democratic Republic of the Congo, have shown that stable isotope ratios measured in the teeth of herbivores reflect the heights at which these animals fed in the forest due to isotopic variation in plants with height caused by differences in humidity at the forest floor and the top of the forest exposed to the atmosphere.