MEMOIRS OF THE PEABODY MUSEUM OF NATURAL HISTORY VOLUME III, PART A REVISION of th< GERATOPSIA OR HORNED DINOSAURS BY RICHARD SWANN LULL STERLING PROFESSOR OF PALEONTOLOGY AND DIRECTOR OF PEABODY MUSEUM, YALE UNIVERSITY LVX ET cHntt a'ipm VERITAS NEW HAVEN, CONN !933 h < 0* CO o O LLl Q Z < I- co UJ z I I- o < cr UJ o o Io oc a u O z g i< en O Ico UJ u = s E "So a noise, was was commanded: and as I prophesied, there came together, bone to his and behold a shaking, and the bones I prophesied as I bone "And when beheld, and the skin covered them them, the sinews and the flesh came up upcm above: but there was no breath in them." lo, Ezekiel XXXVII, 7, MEMOIRS OF THE PEABODY MUSEUM OF NATURAL HISTORY YALE UNIVERSITY Volume I America A Monograph on the Extinct Toothed Birds of North By Othniel Charles Marsh Pp i-ix, 1-201, pis 1-34, text 1-40 Odontornithes: figs Museum Volume II Part Brachiospongidae By III Part A Memoir on a Group of Silurian Sponges : To Pp 1-28, Pp i-xvi, 1-171, Price $1 To 1929 1-62 figs from the Yale University Press, obtained Museum By George Gaylord Simp- 1-32, text pis 1-6, text figs 1-4 pis be obtained from the Peabody American Mesozoic Mammalia son from the Peabody obtained be Charles Emerson Beecher 1889 Volume To 1880 Price $3 New be Haven, Conn Price $5 A Pp i-x, Part Remarkable Ground 1-20, pis from the Yale University Part A Part (bound The IV Part Pp be i-xii, obtained in cloth), Lull be obtained Haven, Conn Price $1 By Horned Dinosaurs 1-175, pis I-XVII, text figs from the Peabody Museum $4 (bound in paper) By Malcolm Rutherford Thorpe In preparation Brachiopod Genera of the Suborders Orthoides and Pentam- eroidea i-xii, New To Merycoidodontidae, an Extinct Group of Ruminant Mammals Volume To 1933 Price $5 Press, 1929 1-3 Revision of the Ceratopsia or Richard Swann Lull 1-42 By Richard Swann Sloth 1-9, text figs By Charles Schuchert and G Arthur Cooper 1-270, pis A and 1-29, text obtained from the Peabody $6 (bound in paper) Museum figs 1-36 1932 To Pp be Price $7 (bound in cloth), PUBLISHED ON THE OTHN1EL CHARLES MARSH PUBLICATION FUND PEABODY MUSEUM OF YALE UNIVERSITY THE TUTTLE, MOREHOUSE & TAYLOR NEW HAVEN, CONNECTICUT COMPANY ' CONTENTS Page Introduction ' History of discovery Resume of discoveries prior to 1907 Recent discoveries Geologic distribution Geographic distribution Alberta Saskatchewan 13 Montana 13 North Dakota 14 South Dakota 14 Wyoming ^ Colorado 16 New '° Mexico Texas ' Mexico I** 1" South America 1° Mongolia British Columbia Alphabetical 1° (footprints) genera and species list of Ecology, habits and manner of life 21 Feeding habits of 20 ^U Environment Methods 19 22 combat Progressive evolutionary changes and phylogeny 24 General morphology 10 Monoclones (Centrosaurus) flexus Axial skeleton 30 Skull Rostral Premaxillary 32 32 Nasal Lacrvmal Maxillary t Jugal i ' 33 33 CONTENTS viii General morphology —Cont Page Skull— Cont Squamosal 33 Parietal 34 Crest 34 Palatal aspect 35 Lower jaw 37 Predentary 37 Hyoid 37 39 Vertebral column 39 Coalesced cervicals Cervical 40 IV Cervicals V Cervical VII 40 and VI 41 41 Cervical VIII Cervical 42 IX 42 Dorsal vertebrae Dorsal VI 43 Dorsal IX XI 44 Dorsal 46 Sacrum 46 Caudals 49 50 Appendicular skeleton Coracoid 50 Scapula 51 Sternum 51 Humerus 53 Ulna 55 Radius 55 Manus 56 Ilium : 57 Pubis 57 Ischium 58 Femur 59 Tibia 60 Fibula 61 Pes 62 Mounted specimens 64 Integument Chasmosaurus , 66 66 Skull 66 Vertebral column 68 Integument 69 Brain and special sense organs 71 CONTENTS IX Page Systematic description of genera and species ' Family Protoceratopsidae 74 Genus Protoceratops ' Protoceratops andrewsi 7R Genus Leptoceratops Leptoceratops ' '° gracilis Q Family Ceratopsidae Genus Monoclones (Judith River phase) 8? oi Monoclonius crassus °-> Monoclonius recurvicornis 0-2 Monoclonius sphenocerus 8^ i Monoclonius dawsoni Sub-genus Centrosaurus (Belly River phase) Monoclonius (Centrosaurus) apertus Monoclonius (Centrosaurus) flexus Monoclonius (Centrosaurus) nasicornus Monoclonius (Centrosaurus) cutleri 87 Genus Styracosaurus 87 • Styracosaurus albertensis 90 Styracosaurus ovatus Genus Eoceratops Eoceratops canadensis 92 Genus Chasmosaurus Chasmosaurus belli 94" Chasmosaurus brevirostns Chasmosaurus 94 kaiseni 96 Genus Ceratops ^ Ceratops montanus 96 97 Genus Brachyceratops Brachyceratops montanensis 102 A Genus Anchiceratops Anchiceratops ornatus • i ()4 Anchiceratops longirostris • a Genus Arrhinoceratops • 106 Arrhinoceratops brachyops 108 Genus Pentaceratops Pentaceratops sternbergii Pentaceratops fenestratus ^ Genus Triceratops • Triceratops horridus Triceratops prorsus Triceratops brevicornus Triceratops serratus 11 + — CONTENTS x Systematic description of genera and species Genus Triceratops —Cont "a Cont 8e 121 Triceratops flabellatus **•*• Triceratops elatus Triceratops calicornis 123 Triceratops obtusus 125 Triceratops (Diceratops) "Inadequate" hatcheri species Triceratops alticorms Triceratops sulcatus • Triceratops maximus 126 12° 1AO l ^° ?Q7 * Of) Genus Torosaurus Torosaurus latus 1J " 09 Torosaurus gladius Bibliography Description of Plates (even pages only) Index 133 140-170 I PLATE 161 XII PLATE A Skull of TRICERATOPS HORRIDUS, Natural History, 1/12 natural Courtesy of the Field B Skull of XIII Museum 1/12 natural 12,003 Field Museum of of Natural History TRICERATOPS HORRIDUS, of Mines, No size No P271 South Dakota size Courtesy of the South Dakota State School of Mines 162 State School PLATE Mil PLATE XIV A Mounted skeleton of TRICERATOPS Natural History, 1/38 natural Courtesy of the American B Mounted skeletons of Canada; Museum cf at American Museum of Museum of size of Natural History CHASMOSAURUS left, plesiotype, ELATUS, No 2245 ; BELLI, right, Courtesy of the Geological Survey of Canada 164 at the National No 2280 PLATE XIV 165 PLATE XV TOROSAURUS LATUS Type, No 1830 Y.P.M., restored by R A Lateral view of skull B Anterior view of skull About 1/16 natural S Lull size Yale Peabody Museum 166 PLATE XV 167 PLATE XVI TOROSAURUS GLADIUS Type, No 1831 Y.P.M., restored by Hugh Gibb and R A Lateral view of skull B Oblique anterior view of About 1/16 natural skull size Yale Peabody Museum 168 S Lull PLATE XVI 169 PLATE XVII FLESH RESTORATION OF CHASMOSAURUS BELLI A Left lateral view B Anterior view C Right lateral view Restored by Mr L S Russell of the Geological Survey of Canada Published with the permission of the Director, Geological Survey of Canada, Dept of Mines, Ottawa, Canada 170 PLATE Mil 171 INDEX Synonyms species or genera arc discussed are in bold-face type Pages on which are in italics, Asterisks signify illustrations 96 Agath.ium.is sylvestris, Ceratops alticornis, Ceratopsian species from, 9, 78, 83, 86, 87, 91, 92, 94, 95, 103, 104, 106 albertensis (Styracosaurus), 28,* 87, 88, VIII Ceratofs horriilus, 96, 116 Ceratops beds, Alphabetical List of Genera and Species, 19 Ceratopsian Albert.) 128 alticornis (Triceratops), 28,* 103, 104, 105, 106, 108, X ornatus, 71, 103, 107, 110, XI andrewsi (Protoceratops), 74, 75,* 76, 77,* 78, 80,* IV Angular, 37 apertus (Centrosaurus), 84 Arrhinoceratops, 74, 81, 98, 103, 106, 107, 108, 125, 126 brachyops, 27, 28,* 29, 106, 110, 127, 129, XII Atlas, coalesced, 39,* 40 74, 77, 85, 88, 89, 91, 92, 93, 98, 99, 102, 104, 106, 107, 113, 124, 130, 132 belli, 1, 27, 28,* 62, 66, 67,* 92, 93, VI, XIV brevirostris, 94, 127, VII kaiseni, 26, 27, 28,* 68, 85, 94, 95,* 96, 97, 110, 1, Ceratopsian species in, 8, 10, 26, 27, 29, 81, 83, 84, 86, 87, 91, 92, 94, 95, 109, 110 Basin, Bison alticornis, Black Buttes, XVII Chclonia, 25 (Chasmosaurus), 28,* 67,* 92, 93, VI, XIV Horn 11 of, Chalk Butte region, Montana, 14 Chameleon, 25 Chasmosaurus, 24-28,* 39, 51, 54, 57, 58, 66, 68, 70, restoration of, 39 Belly River formation, Big map V, VII 37 Bearpaw formation, belli localities, Cervical vertebrae, 39, 40, 41,* 42.* Anchiceratops, 33, 19, 51, 56, 57, 65, 71, 72, 73, 74, 76, 81, 98, 100, 102, 103, 108, 110, 113 Articular, Ceratops paucidens, 93 Ceratopsoidea, 6, 81 Alveolar grooves, 33 longirostris, 3, Wyoming, 15 Climate of Dinosaur Environment, 20 Colorado, Ceratopsian species in, 16, 128 Combat, methods of, 22-23 Converse County, see Niobrara Co., Wyo., Brachyceratops, 5, 39, 74, 78, 79, 84, 97, 98, 99, 100, 102, 110 montanensis, 32, 97, 98, 99, IX brachyops (Arrhinoceratops), 28,* 106, XII Brain, 71, 72,* 73 Coracoid, 50,* Coronoid, 37 crassus (Monoclonius), 82 Crest, 24, 34 Crocodilia, 65 cutleri (Centrosaurus), 87, III brevicornus (Triceratops), 28,* 119, 120 dawsoni (Monoclonius), 83, 84 (Chasmosaurus), 94, VII British Columbia, 18, 64 Dendy, A, Brontosaurus, 20, 52 Denver formation, 3, 128 Denver and Arapahoe formations, brevirostris Brown, 102, 103, 129 calicornis (Triceratops), 123, 124, 125 Camptosaurus, 38, 81 B., 1, 52, 72, 73, 86, Dentary, 37 Diceratops, 3, 6, 107, 108, 126, 127 canadensis (Eoceratops), 91 Diplodocus, 52 Discovery, history Caudal vertebrae, 49-50 Distribution, geographic, 9, map, 11 Centrosaurus, 1, 24, 26, 29, 30, 33, 34, 39, 40, 51, 56, 65, 66, 68, 69, 74, 77, 78, 81, 83, 84, 85, 88, 89, 92, 100, 101, 102, 104, 106, 113, 122, 130 apertus, 5, 38, 40, 82, 84, 85, 86 cutleri, 47, 87, 48, 53, 58, 60, 61, 63, 64, 66, 69, 85, 49,* 50,* 52, 53,* 54,* 55,* 57, 58,* 59,* 60, 61,* 62,* 63, 64, 66, 69, 81, 82, 85, 86, 87, 98, II, III nasicornus, 30, 32, 42, 43,* 45,* 47, 48, 51-57, 60, 61, 63, 64, 85, 86, 87, 90, 91, 113, IX Ceratopsidae, 24, 75, 78, 81 1, 5, 74, 81, 94, 96 montanus, 26, 96, 97 of, geologic, Ditmars, R L., 93 Dja-doch-ta formation, 8, 74 Ceratopsian species in, 8, 74 Dorsal vertebrae, 42,* 43,* 44,* 46.* Edmonton III nexus, 28,* 30, 31,* 32-36,* 37, 41,* 43,* 44,* 46,* Ceratops, Cope, E D., 81 Wyoming, Classification, 74 formation, 1, Ceratopsian species in, 8, 9, 26, 27, 78, 103, 106, 109, 110 XIV Environment of Ceratopsia, 20-21 elatus (Triceratops), 122, 123, Eoceratops, 74, 91, 96, 108, 110 canadensis, 84, 91, 94, 95, 97, 98, 102 cpoccipitals, 34 Evolutionary Changes, Progressive, 24 Feeding habits, 21-22 104, INDEX 174 Manus, 56,* 57 Marsh, O C, 1, 114, 116, 117, 120, 121, 122, 123, 128, Femur, 59,* 60 Fenestra, parietal, 24, 34 130, 132 fenestratus (Pentaceratops), 111, 112.* Matthew, W D., 20 Fibula, 61.* Maxillary, 33 121, 122 flabellatus (Triceratops), 41,* 43,* 44,* 46,* 49,* 50,* 53,* 54,* 55,* 58,* 59,* 61,* 62,* 86, II, flexus (Centrosaurus), 28,* 31,* 36,* III Flora of dinosaur environment, 20 Footprints, 64 Fossa, preorbital, 32 Fossa, supratemporal, 34 Fox Hills formation, maximus (Triceratops), 129, 130 Mexico, 18 Mivart, St George, 65 Mongolia, 5, 18, 74 Monoclonius, 1, 5, 26, 29, 74, 81, 85, 86, 88, 89, 91, 92, 96-101, 103, 108, 130 crassus, 3, 30, 40-45, 47, 48, 49, 51, 54, 58, 59, 60, 61, 81,82, 83, 96, 102 dawsoni, 30, 83, 84, 91, 102 Frill, see crest Fruitland formation, Ceratopsian species Genera and species, Alphabetical List of, Gilmore, C W., 1, recurvicornis, 3, 81, 83, 97 26, 27, 108 Geological sequence of Ceratopsia, gladius (Torosaurus) 3, fissus, in, 8, sphenocerus, 72, 90, 95, 97, 99 28,* 132, XVI 80,* 81 Granger, W., 1, 74, 76 Gregory, W K., 74 Gregory, W K and Mook, C C, 75, 76, 77 Hatcher, J B., 1, 62, 73, 82, 119, 126, 127 Moodie, R hatcheri (Diceratops), 28,* 126 Nasal, 32 Hay, O P., 25 Hayden, F V., Nasal septum, 32 13, 78, 82, 90, 96, 97, L., 2, 131 Morosaurus, 52 Morphology, general, 30-73 Mounted specimens, 64, 65, 66 nasicornus (Centrosaurus), 43,* 45,* 86, IX New 3, Ceratopsian species in, 14, 29, 111, 117, 119, 123, Mexico, Ceratopsian species in, Niobrara County, Wyo., 129 Horn, Horn, in, montanensis (Brachyceratops), 97, 98, 99, IX montanus (Ceratops), 96 Gorgosaurus, 23 Hell Creek beds, 30, 37, 81, 82, 83, 90, 91 117, 119, 123, 129 , gracilis (Leptoceratops), 78, 79, 3, Monoclonius belli, 92 Montana, ceratopsian species Ceratopsian species Nopcsa, F., 58, 60 nasal, 30 supraorbital, 30 16, 108, 111 3, in, 15, 29, 116 horridus (Triceratops), 28,* 116, XIII North Dakota, ceratopsian Huene, F v., 18, 32, 99 Humerus, 53,* 54 Hyoid, 37, 38.* obtusus (Triceratops), 28,* 125, 126.* Ilium, 47,* 49,* ornatus (Anchiceratops), 103 Ojo Alamo Ceratopsian species formation, Osborn, H F., 1,71, 108, 109, Integument, 66, 69, 70 Ischium, 58.* Jaw, lower, 37 Judith River formation, C, O'Harra, C species in, 14 10 ovatus (Styracosaurus), 90 Palatal aspect of skull, 35, 36,* 67.* 3, Parasaurolophus, 82 in, 8, 26, 29, 82, 83, 96, 109 Parietal, 34 W 38, 106, 107, 108 Jugal, 33 Parks, kaiseni (Chasmosaurus), 28,* 94, 95,* V, VII Pathologic bone, 107, 111, 121, 127, 131,* VII Kindle, E M., Kirtland formation, Ceratopsian species Lacrymal, 33 Pelvis, 48 117, 119, 120, fenestratus, in, 8, 121, H H., 123, 125, 126, 28,* 130, 131,* 132, XV Leptoceratops, 24, 26, 76, 78, 98 gracilis, 78, 79, 80,* 81 longirostris (Anchiceratops), 28,* 104, 105, 106, X Lower Ravenscrag formation, 13 Lull, R S., 126 110, 111, 112,* 127 Pes, 62,* 63 latus (Torosaurus), 6, 39, 66, 74, 77, 94, 95, 98, sternbergii, 27, 28,* 108, 109,* 110, V 26, 27, 28, 109, 110, 116, 122, 5, 103, 108, 130 111 130, 132 Lane, 1, Pentaceratops, 4, in, 8, Lacrymal foramen, see preorbital fossa Lambe, L M., 83, 84, 87, 91, 92 Lance formation, Ceratopsian species A., 128, Phrynosoma, Phylogeny, 24, 28.* Polyonax mortuarius, Postfrontal fontanelle, 25-26, Predentary, 37 Premaxillary, 32 prorsus (Triceratops), 28,* 117, 118, 119 Protoceratopsidae, 24, 74, 75 Protoceratops, 2, 5, 24, 26, 39, 74, 79, 100, 110 andrewsi, 74, 75,* 76, 77,* 78, 80,* 81, IV restoration of, frontispiece 101, INDEX 175 ToroMurus, Protorosaurus, 92 ins, Pubis, 57, 5S.* 3, 6, 26, 27, 29, 66, 68, 73, 74, 92, 107, no, iii9, 14, 127, 130 Radius 55,* 56 gladius, 28,* 88, 107, 114, 128, 130, 131, 132, Rattlesnake beds, 17 l tus, Red Deer River, Alberta, Romer, A S., Rostral 31, 65 Sacrum, 46, +7,* 4S, 49.* St Mary Trachodon, 8, Triceratops, 1, formation, 78 San Juan County, New Mexico, Ceratopsian species in, 16-17 Saskatchewan, localities, 13 brevicornus, 27, 28,* 35, 102, 108, 109, C, elatus, 20 107, 114, 115, 118, 119, 120, 122, 123, 124, 125, 128, 129, XIV 72, 98, 114, 115, 118, 120, 121, 122 serratus (Triceratops), 38,* 120, 121 flabellatus, 71, Skeleton, appendicular, 50-63 galeus, 115, 128 (Diceratops) hatchcri, 28,* 114, 115, 121, 126 Skeleton, axial, 30-50 Monoclonius (Centrosaurus) flexus, 30, 31,* 32- horridus, 3, 28,* 71, 114, 115, 116, 117, 118, 119, 125, 126, 130, XIII "ingens," 29, 117, 130 maximus, 115, 129, 130 35, 36,* 37-38 Chasmosaurus, 66, 67,* 68, 95,* V, VI, VII South America, South Dakota, ceratopsian species in, 14, 117 sphenocerus (Monoclonius), 83 obtusus, 28,* 107, 115, 120, 122, 125, 126,* 127, 128, 129 prorsus, 25, 28,* 37, 40, 60, 71, 76, 107, 109, 110, Sphenodon, 71 Splenial, 115, 116, 117, 118, 119, 120, 121, 123, 124 35, 38,* 71, 78, 98, 109, 114, 115, 116, 37 Squamosal, 33, 34 Stanton, T W., serratus, Stegosauria, 24 sulcatus, 71, 117, 120, 121, 122 114, 115, 128, 129 64, 104 sternbergii (Pentaceratops), 28,* 108, 109,* 110, V Two Sternum, 51, 52,* Tyrannosaurus, 23, 71 Ulna, 54,* 55 Vertebrae, cervicals, 39,* 40, 41,* 42.* Vertebrae, dorsals, 42,* 43,* 44,* 45,* 46.* Vertebral column, of Monoclonius (Centrosaurus) Sternberg, C M., 1, Stratigraphic sequence, table of, Stvracosaurus, 4, 25-27, 33, 68, 74, 82, 87, 89, 91 albertensis, 122, 123, 124, 125 Sense organs, 73 Skull, of 114, 115, 117, 118, 119, 120, 121, 123-125, 129 calicornis, 29, 114, 115, 116, 118, 120, 121, Scapula, 50,* 51 Schuchert, 72 6, 26, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 47, 48, 49, 51, 52, 53, 54, 55, 57, 58, 62, 64, 71, 72,* 73, 74, 77, 82, 89, 91, 96, 98, 99, 100, 101, 103, 106, 114, 127, 128, 131, 132 alticornis, 3, 27, 115, 128 35 Russell, L S XV 132, recurvicomis (Monoclonius), 83 XVI 114, 127, 129, 130, 131,* 22, 25, 28,* 111, 28,* 83, 87, 88, 90, 91, 104, VIII ovatus, 5, 90, 104 sulcatus (Triceratops), 128, 129 Medicine formation, Ceratopsian species in, 5, 8, 84, 90, 97, 109, 110 flexus, 39-50 of Chasmosaurus, 68, 69 Surangular, 37 Sweetwater County, Wyoming, 16 Tait, J and Brown, B., 73 Vertebral formula, 39 Tapirus, 79 Williston, S Teeth, 35 Tetrapodosaurus Texas, Tibia, 60, 61.* Wiman, C, 111, 112, 113 Wyoming, ceratopsian species borealis, 64 Vomer, W., 74, 113 in, 15, 117, 119, 122, 123, 125, 126, 128, 130, 132 120, 121, ... Triceratofs flabellatus Triceratofs galeus Triceratofs (Diceratofs) hatcheri Triceratofs horridus Triceratofs Two Medicine Formation maximus Triceratofs obtusus Brachy ceratofs montanensis Triceratofs... Styracosaurus ovatus Triceratofs serratus Triceratofs sulcatus Edmonton Formation Anchiceratofs ornatus Anchiceratofs longirostris Arrhinoceratofs brachyofs Leftoceratofs gracilis Dja-doch-ta Formation,... species of the types of most of or partial ceratopsian skulls and other skeletal material, including the now either in the United Triceratops, two of Torosaurus, and one of Diceratops All of these