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Unravelling the biology of t h e S outhern African Sauropodomorph dinosaurs , Plateosauravus a n d t h e ‘Maphutseng dinosaur’ . Em il Dariu s K rupandan ThesisUniversity presented for the of degree Cape of Doctor Town of Philosophy I n t h e Department of Biological Sciences University of Cape Town F e b r u a r y , 2 0 1 9 The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town i Declaration of Free Licence I hereby grant the University free license to reproduce the above thesis in whole or in part, for the purpose of research. I declare that the above thesis is my own unaided work, both in conception and execution, and that apart from the normal guidance of my supervisor s , I have received no assistance apart from that stated below. E x c e p t a s stated below, neither the substance or any part of the thesis has been submitted in the past, or is being, or is to be submitted for a degree at this University or any other University. I am now presenting the thesis for examination for the Degree o f P h D . _________________ Emil Darius Krupandan 11th d a y o f F e b r u a r y 2 0 1 9 at the University of Cape Town ii I confirm that I have been granted permission by the University of Cape Town’s Doctoral Degrees Board to include the f o l l o w i n g publication(s) in my PhD thesis, and where co - authorships are involved, my co - authors have agreed that I may include the publication(s): 1 . Krupandan, E., Chinsamy ‐ Turan, A. & Pol, D. 2018. The long bone histology of the sauropodomorph, Antetonitrus ingenipes. The Anatomical Record. 301(9):1506 - 1 5 1 8 . SIGNATURE: DATE: 0 3 / 07/ 2 0 1 9 STUDENT NAME: Emil Darius Krupandan STUDENT NUMBER: K R P E M I 0 0 1 iii A b s t r a c t T h e nomen dubium “ Euskelosaurus ” has functioned as a waste - basket taxon for decades , with large bodied sauropodomorph material from South Africa assigned to it without proper analysis. Following the collapse of Euskelosaurus , material p r e v i ously assigned to it was summarily referred to Plateosauravus cullingworthi . Based on this, material referred t o Euskelosaurus from Maphutseng, Lesotho (NMQR 1705) would technically also be referr a b l e t o Plateosauravus . In light of the tenuous nature of th e taxonomic affinity of these materials, a thorough analysis of material associated with both taxa follows. The validity of Plateosauravus was evaluated and the taxonomic affinity of material previously reffered to Euskelosaurus (NMQR 1705) tested. T h e Plateosauravus cullingworthi hypodigm was subject to a rigorous anatomical and phylogenetic a n a l y s es in order to test its taxonomic validity. Undescribed material from Maphutseng, Lesotho (NMQR 1705) was also analysed. It was hypo t h e s i s e d t h a t Plateosauravus was a valid non - Massopodan dinosaur and that NMQR 1705 represented a basal sauropod, with concomitant growth dynamics. Skeletal elements from Plateosauravus and NMQR 1705 were anatomically described and taxonomically identifie d based on comparisons with other sauropodomorph taxa. Anatomical data was scored in a character matrix and subject to phylogenetic analyses. The growth dynamics of NQMR 1705 was investigated through histological analysis of thin sections of long bones. T he material referred to Plateosauravus shows that some of the material belongs to a non - Massopodan basal sauropodomorph t h a t , but there also exists material that should iv be excluded from this taxon based on doubtful provenance and association. The Maphutseng material (NMQR 1705) can be referred to Antetonitrus ingenipes . New elements describe d , including a complete sacrum, reveal additional adaptations highlighting Antetonitrus ’ position as a transitional basal sauropod, exhibiting a mosaic o f basal sauropomorph and derived sauropod characterisitcs. The phylogenetic analysis corroborates the position of Antetonitrus as the sister taxon of Lessemsaurus sauropoides (Bonaparte, 1999), but the new information from the pes indicates that both taxa a r e m o r e derived than Blikanasaurus cromptoni (Galton & Van Heerden, 1 9 9 8 ) . Growth patterns in the youngest individuals of NMQR 1705 exhibit uninterrupted fibrolamellar bone without any growth m a r k s . S u b - adult individuals, also exhibit highly vascularised fibrolamellar bone throughout the cortex, as in more derived sauropods and t he basal sauropodiforme M u s s a u r u s patagonicus , but growth lines occur intermittently throughout the cortex as in Lessemsaurus . Antetonitrus (NMQR 1705) does not exhibit the growth dynamics previously considered characteristic of Sauropoda. Growth marks a re decoupled from bone size, indicating a level of developmental plasticity in this taxon. Modulations in the pattern of vascular canal arrangements throughout the fibrolamellar bone in the cortex may be related to resource availability. Localised bands of radial fibrolamellar bone, followed by resumption of normal growth in two samples indicate disease infliction, and subsequent recovery. v “ If it looks like a duck, and quacks like a duck, we have at least to consider the possibility that we have a small aquatic bird of the family anatidae on our hands.” - Douglas Adams vi Acknowledgements Firstly, I’d like to express my thanks to my two supervisors, A. C h i n s a m y - Turan and D. Pol, without whom this work would not h a v e b e e n possible. Both of them have provided me with countless opportunities and funding to learn new techniques, visit collections across South Africa and Argentina and generally become a fully - fledged palaeontologist. I am grateful to J . B o t h a - Brink and E. Butl er (National Museum, Bloemfontein, South Africa) for allowing for the histological analysis of NMQR 1705; as well as t o B. Rubidge and B. Zipfel (Evolutionary Studies Institute, Johannesburg, South Africa) for access to the holotype of Antetonitrus ingenip es a n d o t h e r comparative material on countless occasions. I’d also like to extend thanks to S. Kaal , Z Skosana and R . S m i t h at Iziko Natural History Museum for access to the collections throughout my work. Further acknowledgments go t o C. Apaldett i and A. Otero for advice, discussion and assistance in South Africa and Argentina throughout my s t u d i e s . L a s t l y , I’d like to thank my family and friends for supporting me throughout this endeavour. This research was supported by a South African National R esearch Foundation (NRF) Scarce Skills Bursary and Palaeontological and Scientific Trust (PAST) grant to E. K r u p a n d a n . v i i T A B L E O F C ONTENTS List of Tables v i i i List of Figures ix C h a p t e r s P a g e 1. Introduction 1 2. S t u d y 1 : The taxonomic validity of P lateosauravus 1 1 cullingworthi (Haughton, 1924) 3. S t u d y 2 : The taxonomic affinity of the “Maphutseng Dinosaur” (NMQR 1705) 53 4. S t u d y 3 : The long bone histology of Antetonitrus 101 i n g e n i p e s . 5. Concluding remarks and future work 1 30 R e f e r e n c e s 138 A p p e n d i c e s A p p endix 1: Character List 219 Appendix 2: D e t a i l s o f Phylogenetic Analysis 254 Appendix 3: Chapter 2 a n d 3 Phylogenetic Analys i s 261 OTU S c o r e s v i i i LIST OF TABLES P a g e T a b l e 1 : Phylogenetic nomenclature for clades 3 mentioned throughout the text T a b l e 2 : Sources of comparative taxa 1 6 T a b l e 3 : Elements present in Antetonitrus h o l o t y p e 61 ( BPI/1/4952a), previously referred material (BPI/1/4952a, b, c; BPI/1/5091; NMQR 1545) and Maphutseng material (NMQR 1705). T a b l e 4 : List of specimens studied, including s u m m a r y 1 22 of histological f e a t u r e s . ix L I S T O F FIGURES P a g e C h a p t e r 2 : F i g u r e 1 . S AM PK 3356, Dorsal centrum. 2 2 F i g u r e 2 . SAM PK 3348. V e n t r a l portion of left 2 3 s c a p u l a F i g u r e 3 . SAM PK 3342. & SAM PK 3350. Left humeri. 2 5 F i g u r e 4 . SAM PK 3347. left radius 2 6 F i g u r e 5 . SAM PK 3341. Partial proximal right 2 7 i s c h i u m . F i g u r e 6 . S A M P K 3 3 4 1 .
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    Dinosaur Species List E to M E F G • Echinodon becklesii • Fabrosaurus australis • Gallimimus bullatus • Edmarka rex • Frenguellisaurus • Garudimimus brevipes • Edmontonia longiceps ischigualastensis • Gasosaurus constructus • Edmontonia rugosidens • Fulengia youngi • Gasparinisaura • Edmontosaurus annectens • Fulgurotherium australe cincosaltensis • Edmontosaurus regalis • Genusaurus sisteronis • Edmontosaurus • Genyodectes serus saskatchewanensis • Geranosaurus atavus • Einiosaurus procurvicornis • Gigantosaurus africanus • Elaphrosaurus bambergi • Giganotosaurus carolinii • Elaphrosaurus gautieri • Gigantosaurus dixeyi • Elaphrosaurus iguidiensis • Gigantosaurus megalonyx • Elmisaurus elegans • Gigantosaurus robustus • Elmisaurus rarus • Gigantoscelus • Elopteryx nopcsai molengraaffi • Elosaurus parvus • Gilmoreosaurus • Emausaurus ernsti mongoliensis • Embasaurus minax • Giraffotitan altithorax • Enigmosaurus • Gongbusaurus shiyii mongoliensis • Gongbusaurus • Eoceratops canadensis wucaiwanensis • Eoraptor lunensis • Gorgosaurus lancensis • Epachthosaurus sciuttoi • Gorgosaurus lancinator • Epanterias amplexus • Gorgosaurus libratus • Erectopus sauvagei • "Gorgosaurus" novojilovi • Erectopus superbus • Gorgosaurus sternbergi • Erlikosaurus andrewsi • Goyocephale lattimorei • Eucamerotus foxi • Gravitholus albertae • Eucercosaurus • Gresslyosaurus ingens tanyspondylus • Gresslyosaurus robustus • Eucnemesaurus fortis • Gresslyosaurus torgeri • Euhelopus zdanskyi • Gryponyx africanus • Euoplocephalus tutus • Gryponyx taylori • Euronychodon
  • High European Sauropod Dinosaur Diversity During Jurassic–Cretaceous Transition in Riodeva (Teruel, Spain)

    High European Sauropod Dinosaur Diversity During Jurassic–Cretaceous Transition in Riodeva (Teruel, Spain)

    CORE Metadata, citation and similar papers at core.ac.uk Provided by RERO DOC Digital Library [Palaeontology, Vol. 52, Part 5, 2009, pp. 1009–1027] HIGH EUROPEAN SAUROPOD DINOSAUR DIVERSITY DURING JURASSIC–CRETACEOUS TRANSITION IN RIODEVA (TERUEL, SPAIN) by RAFAEL ROYO-TORRES*, ALBERTO COBOS*, LUIS LUQUE*, AINARA ABERASTURI*, , EDUARDO ESPI´LEZ*, IGNACIO FIERRO*, ANA GONZA´ LEZ*, LUIS MAMPEL* and LUIS ALCALA´ * *Fundacio´n Conjunto Paleontolo´gico de Teruel-Dino´polis. Avda. Sagunto s ⁄ n. E-44002 Teruel, Spain; e-mail: [email protected] Escuela Taller de Restauracio´n Paleontolo´gica II del Gobierno de Arago´n. Avda. Sagunto s ⁄ n. E-44002 Teruel, Spain Typescript received 13 December 2007; accepted in revised form 3 November 2008 Abstract: Up to now, more than 40 dinosaur sites have (CPT-1074) referring to the Diplodocidae clade. New been found in the latest Jurassic – earliest Cretaceous remains from the RD-28, RD-41 and RD-43 sites, of the sedimentary outcrops (Villar del Arzobispo Formation) of same age, among which there are caudal vertebrae, are Riodeva (Iberian Range, Spain). Those already excavated, assigned to Macronaria. New sauropod footprints from the as well as other findings, provide a large and diverse Villar del Arzobispo Formation complete the extraordinary number of sauropod remains, suggesting a great diversity sauropod record coming to light in the area. The inclusion for this group in the Iberian Peninsula during this time. of other sauropods from different contemporaneous expo- Vertebrae and ischial remains from Riodevan site RD-13 sures in Teruel within the Turiasauria clade adds new evi- are assigned to Turiasaurus riodevensis (a species described dence of a great diversity of sauropods in Iberia during in RD-10, Barrihonda site), which is part of the the Jurassic–Cretaceous transition.