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Armored Dinosaurs of Mongolia T.A The Armored Dinosaurs of Mongolia T.A. Tumanova 1987 The Joint Soviet-Mongolian Paleontological Expedition Transaction vol. 32, 76p. Translated by Ruth Griffith, edited by Kenneth Carpenter and T. A. Tumanova i Editor's Comments There are two major philosophies regarding translations, one which literally translates the words (transcription) and the other the implied meaning. The problem with transcription is that words, sentence structure, punctuation, grammar, etc. (i.e., everything that makes each language unique), is often unintelligible or confusing. This is especially true from Russian to English. Anyone who has taken a foreign language knows that a word-by-word transcription to English seldom makes sense. To avoid this problem and to make this translation of " The Armored Dinosaurs of Mongolia " readable, I have edited the translation by my collaborator Ruth Griffith as I would a manuscript written by a colleague for whom American English is not a first language. I do so on the basis that American English rules of grammar are not identical to those of Russian. In a few places, words have been added for clarification; these are inserted in brackets, [ ]. In addition, most Russian names for localities and formations have been standardized to Jerzykiewicz and Russell (1991) because these names are more readily understood among English-speakers. As parts of the translation were completed, they were checked by Tatyana Tumanova for accuracy. The resulting translation tries to be true to the original. Kenneth Carpenter Denver, 1999 Reference: Jerzykiewicz, T., and Russell, D.A., 1991, Late Mesozoic stratigraphy and vertebrates of the Gobi Basin. Cretaceous Research 12 :345-377. [Originally, this translation was sold and the proceeds used to pay for Dr. Tumanova's membership in the Society of Vertebrate Paleontology. Now, funds are available elsewhere, so this important monograph is being released more broadly - Kenneth Carpenter, 2008] ii CONTENTS Introduction ................................................... ............................. 1 Material ................................................... ................................ 3 Chapter 1 A brief history of ankylosaur studies ................................................... 5 Chapter 2 Systematic description ................................................... .................... 8 Suborder Ankylosauria .............................. .......................................... 8 Family Nodosauridae Marsh, 1890 ................... ....................................... 9 Family Ankylosauridae Brown, 1908 . ....................................... 9 Subfamily Ankylosaurinae Brown, 1908 . .................................. 11 Subfamily Shamosaurinae Tumanova, 1983 . .................................. 19 Chapter 3 Morphology ................................................... ............................ 22 Skull.............................................. ........................................ 22 Shape and external features ....................... ........................................ 22 Cranial armor ...................................... ..................................... 23 Structure of the Skull roof ........................ ......................................... 27 Morphology of the Nasal Cavity ..................... ....................................... 29 Palatal area ....................................... ...................................... 31 Ossifications of the quadrate complex . ......................................... 31 Braincase .......................................... .................................... 33 Ethmosphenoidal section ............................ .................................. 33 Otico-occipital section ............................ .................................... 33 Maxilla ............................................ .................................... 38 Mandible ........................................... ................................... 41 Teeth.............................................. .................................... 41 Hyoid apparatus ................................... ..................................... 42 Muscles, glands, nerves and vessels of the head ................................................. 43 Musculature of' the head ........................... ....................................... 43 Lateral nasal gland and Jacobson's organ . .......................................... 44 Circulatory system ................................. ...................................... 45 Cranial nerves ..................................... ..................................... 46 Chapter 4 Dermal armor ................................................... .......................... 48 Chapter 5 Phylogeny of ankylosaurs ................................................... ................ 51 Chapter 6 Some morphological features of ankylosaurs and their systematic significance ...................... 55 Chapter 7 Ecology ................................................... ................................ 59 Chapter 8 Geological and geographical distribution of ankylosaurs ........................................ 63 Localites of the Mongolian ankylosaurs . ............................................ 67 Lower Cretaceous sites ............................. ...................................... 67 Upper Cretaceous sites ............................. ...................................... 68 Summary ................................................... .............................. 72 References ................................................... ............................. 72 Plate Captions ................................................... .......................... 81 iii INTRODUCTION The Ankylosaurs are a separate group of dinosaurs known from the Cretaceous Period and widespread throughout practically the entire World. Ankylosaur morphology reveals their extreme specialization, as well as primitiveness compared to the rest of the dinosaurs. Until recently, the discoveries of these animals consisted, as a rule, of scutes and spines from armor or of fragments of the postcranial skeleton; characteristically, cranial material was poorly preserved. Because of this, knowledge of the group remained rather limited and descriptions were basically of a superficial character. Only in recent years have a monograph and some separate papers appeared in which the morphology of Mongolian (Marya ńska, 1971, 1977) and North American (Coombs, 1972a,b,c, 1979) armored dinosaurs were treated in more detail. A major obstacle to the study of ankylosaur cranial morphology was found to be the massive overlapping of the cranial surface by osteodermal plates which are firmly fused with the underlying bones. In this connection, was a lucky find by the Polish Mongolian Paleontological Expedition to Bayn Dzak (MPR) of a young specimen of Pinacosaurus grangeri (Marya ńska, 1971, 1977); on the skull sutures were still visible. A detailed study of ankylosaur morphology and comparisons with other dinosaurs and reptiles enable us to solve more definitely the problems of their systematic position, relationships, and origins of both North American and Asian specimens, and allows us to raise questions concerning dinosaur evolution as a whole. The aim of the present work is an attempt to make a detailed study of ankylosaur morphology, especially of the skull, so it provide us with the key to clarifying the origin and phylogeny of this independent group of animals. Because of the poor state of previous ankylosaur morphology studies, there were still problems concerning their origin and connection with related groups, and even about relationships within the group. In the present work, only the least studied specimens of ankylosaurs from Mongolia are considered in detail, including Talarurus known almost solely by the postcranial skeleton, and the new genus Amtosaurus , the same age as Talarurus . A revised diagnosis of the genus Maleevus is also presented. A new Early Cretaceous genus of ankylosaurid, Shamosaurus , is also established. The family Ankylosauridae is divided into two subfamilies, the Shamosaurinae and the Ankylosaurinae. Shamosaurinae, the more primitive, is represented at present by only one genus, Shamosaurus , from the Lower Cretaceous; the Ankylosaurinae includes all the rest of the known ankylosaurids. Sauroplites , from the Lowest Cretaceous of China, possibly belongs to the Shamosaurinae, but we can not resolve the problem definitively because its remains are incomplete (Bohlin, 1953). As criteria for this division [into two subfamilies], we have the following: difference in form of the preorbital section of the skull, orientation of the orbits, relative distance of the orbits from the occiput, and also some structures of the skull of Shamosaurus that are intermediate between 1 Ankylosauridae and Nodosauridae. These include the construction of the premaxillary section of the muzzle, the large curves anterior to the pterygoid flanges, contact of the pterygoids with the basisphenoid, the structure of the quadrates and paroccipital process, and the circular shape of the occipital condyle. All of these intermediate traits are obviously due to the very great age of Shamosaurus and its proximity to the ancestry of both families. The study of the discoveries, including some preliminary material we received, supports the claims of W. Coombs for the extremely isolated position of the ankylosaurs within the ornithischian
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