DINOSAURS THE ENCYCLOPEDIA by Donald P. Glut Foreword by KENNETH Canresten, PH.D.“Toall my friends at The Field Museum {Geology Department. Libsaty and Photography Department) «specially 10 Nina Cummings Frowe cover art Tife restoration of Chusmasaurie Bell by aati Boris Revie Glut, Donald F Dinessur: the eneylopedin : supplement five / by Danali F Gls forever by Kenatsh Carpenter. Pa. Incas ibiographicalrelerenees sd index Bpr.1s, 980-7863. (brats binding: $02 alkaline pare) & 1. Dinoiaun Encyclopedia. 1. Title QEse2. D652 2008 Sor 9103 a2 5.47668 Brvnh Library cataloguing dara are alll (©2008 Donald F. Glut, All igs seservee (Ne pare of ty ba vay be neproacd oy mented Ina firm fon any mens etre sr mech. bcludlag phosceping ‘or cording 0 any norman nigel retrieval 9a, Teithour perma in wg fm ale pb Masulaeired in the United States af Ameries McFarland &> Company, Jus. Publishers Bos 61, Jffovion, Novth Covalinn 28640 ewe mcfarlandpadseornhe author thanks the following vertebrate paleon- "LT ise se ical ne manuscript, for their invaluable cvricisms. suge gestions, and other comments. and for genevally helping me co improve the text (or primitive saurschisns and cherapods) Ralph Molnar, Museum of Northern Arizona, Flagstaff (aval sauropodomorphs) Peter M. Galton, College of Nazuro- pathic Medicine, University of Bridaepore. Bridgeport Connecticut: (sauropods) John S. Melntosh, Westeyan University, Middletown. Connecticut: (ths:cophorans) Kenneth Carpenter, Denver Museum of Nature andl Sei- cence, Denver, Colorado; (primitive onithischians,iguan ‘odontians, and hadroseurs) Michaeb K. Bree Surman, National Museum af Natural Histor Smithsonian Lnstx tation, Washington, D.C., also my ehie siersifie advisor, who reviewed the entire work and made myriad sugges~ tions for improving the overall manuscripts. (pachy~ cephalosauriansand more primitive “margivocephalians’) Rober: M Sullivan, Siete Museum of Pen ceratopsians) Peter Dodson. School of Vererinary Medi- cine, Univessty of Pennsylvania, Philadelph: Thanks co the following veriebeate palzontologists, who sent me reprints of articles, copies of journals, pho- tographs, sides, and drawings of specimens, pusanal com- municavions, advice, or othersise contrihuced directly in some way to the production of tis volume: Laweence G. Bames, Luis M. Chiappe. Doug Goodreau, John M, Har- ris, Gary Takeuchi, Howell W. Thomas, znd Xiaoming Wang, Vertebrate Paleontology Department and the Di- nosaur Instiuse, Natural History Museum of Los Ange~ les County, Los Angeles, California; Marthew T. Casrano, Department of Paleomioiogys National Muszum of Navaral Hiscory, Smithsonian Insiuion, Washington. D.C: Ger- ald Grellet'Tinner, Museum of Geology. South Dakota School of Mines and Technology: Rapid City, Souch Acknowledgments Dakota; David W, Krause, Depariment of Anacomical Sci- knees, Stony: Brook University, Stony Brook, New York: Gerhard Maies, formerly Royal Tyrrell Muscum of Palacontoloes; Calgary. Alberta, Canadas Poter J, Mako= vicky and William F, Simpson, Geology Department, The Field Museum; Ruben D, Martiner, Universidad Nacional ée la Patagonia, Argentina; Dareen Naish, University of Portsmouth: and Seow D. Sampson. Urth Museum of Natural History and Department of Geology and Geo- physics, Universiey of Utah, Salt Lake City, Utah Also thanks to the “paleovartists” Berislay Krzie. ‘Mark Hallet, and Gregory S. Paul, who contributed di- hrosaurillustravicns fand who retain all copyrights and aay ‘other rights to their respective ilkastrations, whether es pliciely stated or not) to this volume. Finally. hanks to Jerice Barrios and Nix photography depastment, and Sean O. Bober. exhibirs The Field Museum; Allen A, Debus and Diane E. Debus Hell Creck Creations: Iracy L. Ford (Dino Hunter”): Mike Fredericks, Pivhistorie Times: my mother. Julia Glut Don Lesser, DinoDon, Inc. Luci Betti Nash. Depart- ment of Anatomical Sciences. Stony Brook University, ‘Stony Brook, New York: George Olshevsky. Publications Requiring Research, Sam Diegos Adam Spektar, Spekror Manor: Pete Von Sholly, Fossil Records: and Beverly and Bill Wareen| apologize 10 anyone and any insticution | may have Inissed Cummings, Once 2auin, have done my best 10 exec all iluse ions sad to obtain permissions for their use in this book, if permissions are so requited. Copyrights for pho= tographs (except for those privately eaken, sich ac the ones shor by the author) of specimens, whether stated or not, are assumed 10 he held hy the institution housing those specimens or by the publications in which those pictures inidally appeacedsince Dinosaurs: The Encyclopedia, Supplement 4 came out. But what a difference those few years have made in dinosaur pateoncology. The field has exploded interna~ tionally, with new discoveries announced monthly from a variety of couneries and continents, We are truly fiving in the Golden Age of dinosaur paleontology, as evidenced by the need for these supplements to keep abreast with all che now discoveries I 1 is sugprising chat only a couple yeats have passed For me, these supplements are not simply another addicion co my bookshelf, but an important tool regularly. Discov tase ies happen so fequendly that [ cannot keep up with chem, and [doubt many professional di- nosaur paleontologists can either, These supplements pro- vide us with an encapsulation of the specimens, their in~ terprotation, and most importantly, references to the original descriptions. The vase majority of chese original descriptions occur in sechnical literature that is noc easily accessed by the nonprofessional, because few non-univer~ sicy libraries catry chem. Thus, Don Glut’ encyclopedia and supplements provide an imporranc bridge between the scientific and amaccur communities, Foreword If you area dinosaur aficionado, rest assured hae this latest compendium has che latest on the greatest. This i formation not only includes the ‘atest new dinosaur, but also information on old favorites. As a science, dinosaur palzonrology tweaks its ideas as new informacion becomes available, You can trace some of rhese changing ideas through the supplements. So, how much longer will these supplements be needed? Thus fae, the Encylopedia and Supplemencs fexclude ing the present volume] total 3,678 pages For about 527, genera. Recent extimates by Steve Wang and Peter Dod- son suggest thac over 70 percent of che dinosaur genera that may have existed have yet to be discovered. That means we can see the need for these supplements inco the far distant Fuvure Kes ncTH Caneenter, PH.D. Curator of Lowey Vertebraxe Paleontology Denver Musewra of Nature and ScienceTable of Contents Foreword, by Kenneth Carpenter, Ph.D. Acknowledgments Proface 1 Introduction The Mesozoie Era New Discoveries, Ideas and Studies More on Ectothermy Versus Endothermy Dinosaurs and Birds Dinosaur Extinctions If Dinosaurian Systematics HI Dinosaurian Genera IV Nomen Nudum and Excluded Genera A Lise of Abbreviations Appendix: Main-Ensry Catalog of Genera Glossary Bibliography Index 152 155 168 173 245 695 699 703 721 733 781he present supplement is the fifth in this writer's I unofficial and open-ended “series” of reference books chat started almost a decade ago with Mc- Farland!’s publication of Dinosaurs: The Encyclopedia in 1997. (That book is referred co throughout this text as DcTE; the succeeding supplementary volumes are referred to herein as Si, 52, $3, and 4.) ‘As before, the intent ofthese books isto keep the sub- ject of dinosaur paleontology reasonably current (che term dinosaur” usually referring to “non-avian dinosaurs,” taxa traditionally regarded as dinosaurs; for information per- taining specifically to “avian dinosaurs.” i.,bitds, see the second appendix in 54). The present supplement offers, as did its predecessors, new information based upon the more recent research of vertebrate paleoncologists. It also corrects previously published errors or emends “old info: mation” no longer regarded as cortect. Like the earlicr supplements, this volume builds upon the foundation established in D-TE, the purpose of which was to make available, between «wo covers, a reasonably ridy compendium of dinosaur-relaced information based upon the original work of vertebrate paleontologists. Fol- lowing that fist book with « series of supplementary vol- umes is a way ¢o keep the subject of dinosaur paleontol- ogy relatively current ‘This book —like the original volume and its ist four supplements —is intended 0 serve as a handy reference tool useful co both the professional scientist and the seu- dent, while at the same time providing less technical in- formation that should be of interest to the more nontech= ically oriented dinosaur enthusiast. As the present author has often stated, these volumes do not constituce a substi- ture for the original paleontological literature, nor are they used for formal taxonomic purposes. Again, the writer en- courages those readers lacking a more technical back- ¢ground in the subject matter to use the glossary as an aid in understanding the more difcule passages in the texts also, urge the reader to make use ofthe bibliography and seek out the cited original publications Most of the information appearing in this volume is based on the original research of vertebrate paleontolo- gists that has been published —as peer-reviewed technical articles printed in scientific journals — and was available to the present writer in time for inclusion, Some of this in- Formation was first published as “abstracts” —ie., shore, concise, and often preliminary summaries of works-in Preface progress — thar were also presented as talks at a scientific conference or symposium, the Final detailed results (some~ times differing, to varying degrees, in content from that appearing in the abstract) of which could subsequently appear in the Form of a fully realized article. Caution, ho ever, should be used when evaluating the information pre~ sented in these short publications. Note also that abstracts are nor peer reviewed: their content, therefore, should not necessarily be afforded the same weight as a reviewed ar- ticle published in an accrediced journal Paleontology isa science that depends on the collec sion, preparation, and then study and analysis of fossil materials housed in institutions certified by the American [Association of Museums. What we know about dinosaurs {as well as other extinet organisms) depends not only on the number of fossils presenting these animals, but also ‘on the completeness and quality of preservation of those specimens. With modern prospecting and collecting meth- ‘ods, new discoveries of dinosaur specimens continue to be made throughout the world, such finds often leading not only to the naming of new dinosaurian genera and species, but also to the reinterpretation of known taxa. As did its predecessors, this supplement presents many new di- nosautian generic and specific names, and also additional informarion concerning taxa that fist appeared in eatlicr volumes. [As stared in the past books, I ask that the reader be tolerant in dealing with 2 multi-volume book series with so many words preserved between theit covers. Spelling and other errors are bound ta infiltrate the text despite the ‘myriad human eyes and computer spell-checks combing the manuscript and proof pages before the final maverials ate shipped off to the princer. Again [ask the reader to di- rect any and all cortections to me in care of MeFarland, or via email (dinosaue@frontlinefilms.com), so thac such er- rors may be corrected in a future volume. I thankfully welcome in advance all articles, pho- tographs of specimens, arowork, and related items senc co me for possible use in preparing the next supplement. Continuing the Formac of che earlier supplements {and to keep these volumes at 2 manageable size), I have avoided much redundane information (eg., background data, definitions of taxa above the level of genus level, il- lustrations, ev.) chat can be found in che earlier volumes. ‘As already staced, the content of the text is based on in formation that has published in the paleontologicalPreface lierature before this volume went to press. Therefore, in~ from fossil remains complete enough co produce a rea- formation and illustrations acquired by the author beyond sonably accusate depiction the publisher's cut-off dace (April 15, 2007) will have to aswait the next supplement for inclusion DoNaw . GuuT As before; life restorations of dinosaues featured inthe Burbauk, California USA “genera” section ofthis book are included only if—in the May 15, 2007 estimation ofthe present writer — they portray taxa knownI: Introductionince the publication of the fourth supplement inthe specimens have continued to be found, col- catalogued, and prepared, while new di- nosatirian genera and species ae erected and described iy vertebrate paleontologists. Indeed, some of these more recent discoveries, discussed inthis volume, have n¢ series of reference books, dinosaur S been unusual, surprising, unique, and even spectac~ lar These discoveries include what may be the largest Jurassic theropod specimen yet collected (see Tor- zesourus entey); “pigantic” compsognathid theropod with long filamentous integument, and with the re- rains of a dromacosaurid preserved in the stomach auca (see Sinoclliopterys entry); the fist examples of ed apparent Rexible vascular tissue and dullary bone in a dinosaur (see Tyrannosaurus prese entry); hundreds of specimens representing the first verifably basalmose therizinosauroid theropod (see Falcarias entty); the first undisputed evidence of a deinonychosaurian theropod in she Southern Hemi sphere (see Nequenraptor entry) the frst South Amer- ican deomacosaurid represented by reasonably com- plete skelecal remains (sce Buitrerapeor entry); the first evidence of a dinosaur (ie, che dromaeosaurid Bam Siraptor) possessing opposable fingers (see below, sec tion on theropods); a modest-sized Mongolian sauro- pod possessing an extremely long neck (see Erkete centey)s a “dwarf” macronarian sauropod from Ger- many celebrated as the smallest adule neosaurpod yet discovered (see Europascurus entry): “tiranosaurid” eggs containing preserved membrana testacea (see boclow); the first embryonic “prosauropod” skeletons, indeed the oldest embryonic dinosaur skeletons yet discovered (see Massospondylus entry); pethaps the most massive animal known from Europe and one of the largest of all known sauropods (see Turiasaurus entry); the frst cartilage Found preserved in a sauro- podomorph (see Cetiosauriseus entry); an atypically very short-necked sauropod from South America (sce Introduction Museums and other insticu tions continue co add dino- sur skeletons to their x bit halls, such a5 thie mounted cat specimen of the Tyrannosaurus rex skele- ton known as“ cently installed versity of Manchester,Introduction Brachyrrachhelopan entry): the fst report ofan tooth”-like suuevure ina ritanosaurian sauropod (3 Ineroduction") the fist trace and body fossil evi- dence indicating dinosaurian burrowing and denning behavior (see Oreytedramens entey): an iguanodontisn possessing che largest seth ofall known plant-cating dinosaurs (see Lanchowanrus entry: the frst North American spiked yet fat-headed pachycephalossurid Gee Di found in Alaska (see Alavkacephale entry); the basal most ceratopsian yet found, dating back to the Late Jurasic period (see Vinlong entry); and the smalles (subadule) Tricerazops (se entey) skull yet discovered, surprisingly possessing fill and horas. Among the more dramatic changes, and one with salient phylogenetic implications, isthe “sinking” by Robert M. Sullivan (20066) of the clade Pachy- cephalosautia and the various clades within Pachy- assessment ofthis grup ‘rex enity); the frst pachycephalosaurid Selena casts of the ther pod dinosaur Monalopho ead wl We ep Ste Tugfangosaurns tigi offered 0 ccephalosauridae. Sullivan ‘of bone-headed dinosauts’ also, ifcorreet, would un- ermine the validity of the “Marginocephalia,” a clade huced by Paul C. Sereno in 1986—to date, aur Dynasty” exhibition, widely accepeed, is validity only rarely challenged — to embrace all “pachvcephalosaurian” and ceratopsian dlimosaurs. Nevertheless, another recent study by Xu, Forster, Clark and Mo (2006) seems to reinforce che validity of the Marginocepbalia (see “System chapter) Dinosaurs continue to captivate the publics in. terest as well as that of the scientist Ie is not surpris: ing, there ponents of their world (e.., nondinosaurian animals, € organisms, plants, geography; and so in the premiere attractions in navutal his- pptian mummies di- rnasaur exhibitions — usually presenting to the public authentic fossil specimens, mounted skeletons (both original ma ing paincin mated fu! that dinosaurs, as well as other com- forth), re and science museums (with arguably occupying second pl ice status). Traveli and casts) life restorations (inchad= models, and somerimes full-scale ani and other attraction: tour museums. Such exhibitions usually provide both, education and entertainment to a dinosaur-conscious publicTwo such recent exhibitions, “Chinssaurs: Di- saosaut Dynasty” and “DINOWorld,” were organized and are offered by Don Lessem, one of the founders of the now defunct Dinosaur Society (sce S/). Lessem pur this exhibition cogether through his company DinoDon, Inc., in cooperation with several Chinese ‘museums, The exhibition was designed by the Royal British Columbia Museum in Victoria, British Co- lumbia, where ie debused in spring, 2008, nasaurs: Dinosaur Dynasty” (also known as “Dragon Bones"; see Lessem 2005 for additional in- formation), is hailed as "the largest collection of au- thentic Chinese fossils ever eoured.” The exhibition is hosted hy the Children’s Muscum of Richmond, Vie- ginia, the Conneceicut Science Center, in West He ford, and che Pink Palace Mascura in Memphis, Ten anessee. Ceauinuing in 2003, it has couted such other institutions as rhe Arizona Science Center in Phoenix, the Pink Palace Mascum, and The Field Museum in Chicago, ending its run in 2007. Available compa- nents of the exhibition include cast skeletons of such Asian dinosaurs as the checopods Monolophosaurun Vangchuanosanrus, Velociraptor, and the putative spe cies ‘Dilophosaurus sinensis, the suvtopods Belluscurus, Introduction Mamenchisaurus, and Norotaura, stegosaur Taojaan- {goraurus, oenithopad Bactrosaurus, and badrosaut Biimtaosaeras, also the pterosaur Dsungaripterar, orig inal fossils including specimens of the theropod Ar chacornithomimus, "prosauropod” Lufengoraurts, the ceratopsians Puitcaconurus and Protoceratops, dincsaur epgs: a theropad nest with eggs, dinosaur teeth, di- nostur footprints, and nondinosaurian (eg. cute, salamander, fish, marine reptiles) skelecal material; life-sized robotic recreations of Velociraptor, the hadrosaur Seurolophus, chetopod Ovinaptor, and #o- oceratops: a life-sized (70 feet long) model of the sauropod Mamrnehisauras; murals, depicting the th Mesozoic periods (see section on “The Mesozoic Era, below), painted by Chinese American artists Ya Shans plus various video presentations, interaczive exhibits, and other dinosaur-relaced attractions. Much of che proceeds garnered fiom “Chinasauss” will go to help Finance paleontological work in China. “DINOWoxld,” billed as “THE largest traveling dinosaur exhibit inthe world,” includes numerous ex- hibits and actractions, some of them similar to those featured in “Chinasaurs.” Among the components offered in this exhibicion are skeletal casts of the Sheu (esto KMV HO se0 SP of ‘Dilophosave tion" Chinas: DingsaueIntroduction Velociraptor one of he robotic dinosaur figures salable in the exhibition *Chisntauras: Dinosaur Dynasty” from DinoDon,Introduction Robotic Protceratepr made by Crestve Presentations and subsequently reir Bished by Creature Craft for theropods Tjranmosaurns and Giganotosaurus, the ropods Argentinosauras and Amargaieurus, and the pachycephalosaur Pachyeephalosaurus, and a robotic “Tjrannorauras igure. The “DINOWorld” show played fiom September to December, 2004, then moved on to tour Europe Another exhibition, the “Gigantic Dinoseur Expo 2006” (or simply “Dino Expo’), held from July to September of that yar a the Messe Interoational Convention Complex, in Chiba City, Japan (see [Anonymous] 2006; Chan 2006). Presumably the largest exhibition of any kind ever held at the com- plex, the cheme of the Expo was “Life and Environ- ment — Wonder of Evolution.” Displays feacated at the exposition included more than 270 fosilspeci- mens (@. the theropods Lourimhanasaurns and Er- Wianosaurus, the sauropods Qzogoseurus, Mamen= ohitaurus, Camarasauras, Diplodecus, Kurmingoraurus, and the as yet undescribed “Moabosauras” [see “Nomen Nudum and Excluded Genera"), and the stegosaur Dacentrarus), both original material and cast, and ako numerous fll-scale models of dinosaurs and other Mesozoic taxa. Included in the exhibition ‘vasa life-sized diorama featuring fauna and flora fiom the Jehel Group of China. The main attraction atthe show was a reconstructed skeletal cast of the enor- ‘mous sauropod Supersaurs measuring 33 meters (ap- proximaely 110 feet in length “Dinosautios Argentinos: Giants of Patagonia" isa uaveling exhibit featuring an impressive collection of skull and skeletal cst specimens representing South American dinosaurs and other Mesozoic reptiles (see , Calvo and Frachtenberg 2006), The vwas assembled with the cooperation of veree- brate paleontologist Jorge O. Calvo. Most ofthe pieces in this display ate fiom Patagonia, principally from Neuguen Province ‘The exhibit, which opened in North America in