The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock that is found in the western United States, which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltstone and limestone and is light grey, greenish gray, or red. Most of the fossils occur in the green siltstone beds and lower sandstones, relics of the rivers and floodplains of the Jurassic period.
The fauna of Morrison Formation is similar to one in the coeval rocks of Tendaguru Beds (in Tanzania) and Lourinhã Formation in Portugal,[1] mostly with the second. Some genera are shared in Morrison and Lourinhã, such as Torvosaurus,[2] Ceratosaurus,[3] Stegosaurus, Dryosaurus,[4] and Allosaurus.[5] In sum, Morrison Fm has 37 valid genera of dinosaurs.
The herbivorous ornithischian dinosaurs were diverse but not as common as sauropods in the Morrison. Fruitadens, previously known as the "Fruita Echinodon", was found to be a heterodontosaurid. Plate-backed stegosaurids included Hesperosaurus mjosi, Stegosaurus armatus, S. ungulatus, S. stenops, and Alcovasaurus longispinus. Armored dinosaurs that weren't stegosaurs were unknown in the formation until the 1990s. Two have been named: Gargoyleosaurus parkpinorum and Mymoorapelta maysi. Ornithopods, bipedal herbivores, came in several types. Small "hypsilophodonts" included Drinker nisti, Laosaurus celer, "L." gracilis, Nanosaurus agilis, Othnielia rex, and Othnielosaurus consors (all of which are now synonymous with Nanosaurus). Larger but similar-looking dryosaurids were represented by two species of Dryosaurus (D. altus and D. elderae) and the camptosaurid Uteodon aphanoecetes, which is currently known only from Dinosaur National Monument. Still larger was the more common Camptosaurus dispar. Dryosaurids and camptosaurids were early iguanodonts, a group that would later spawn the duck-billed dinosaurs.
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Anomoepus | Colorado[6] | Morrison Anomoepus lack the handprint impressions found associated with earlier instances of the ichnogenus in New England. | |||
Camptosaurus[7] | C. amplus[8] | Wyoming | Now known to be based on a theropod foot.[9] | ||
C. aphanoecetes[10] | Utah, Brushy Basin member | Reclassified as Uteodon. | |||
C. browni[11] | Junior synonym of C. dispar.[12] | ||||
C. dispar[13] [14] | Colorado, Oklahoma, Utah and Wyoming | "[Twenty-five to thirty] disarticulated skull elements, some with associated postcrania, approximately [ten] partial, articulated skeletons, juvenile to adult."[15] | A member of Camptosauridae. Junior synonyms include Camptosaurus browni, C. medius, and C. nanus. | ||
C. medius | Junior synonym of C. dispar. | ||||
C. nanus | Junior synonym of C. dispar. | ||||
Dinehichnus | Utah, Saltwash member | Multiple Dinehichnus trackways have been discovered. The tracks run parallel to one another, indicating that the trackmaker was at least somewhat of a social animal. | Dinehichnus are attributed to dryosaurids. The tracks preserve feet characterized by widely splayed toes and that are rotated somewhat toward the midline of the trackmaker's body. Each track is accompanied by "distinct ... heel impressions". | ||
Drinker[16] | D. nisti | Wyoming[17] | "Partial skull and postcranial skeleton."[18] | A basal neornithischian about 2 m long, slightly smaller than Othnielosaurus. Junior synonym of Nanosaurus agilis[19] | |
Dryosaurus | D. altus | Colorado and Wyoming, Brushy Basin member[20] [21] | The remains of many individuals have been uncovered, with some sites containing hundreds of bones from Dryosaurus of multiple age groups.[22] | A large dryosaurid iguanodont up to 2.4 m (7.9 feet) long and 114 kg (251 lbs) in weight. It was physically similar to Othnielosaurus, although larger and with more derived teeth. | |
D. elderae | Utah, Brushy Basin member | A 2 partial skeletons, one a juvenile, and 2 additional fragmentary specimens. | A large dryosaurid iguanodont up to 2.4 m (7.9 feet) long and 114 kg (251 lbs) in weight. It was physically similar to Othnielosaurus, although larger and with more derived teeth. | ||
"Laosaurus" | L. altus | Now known as Dryosaurus altus | |||
L. celer | Wyoming | Considered dubious due to fragmentary remains. | |||
L. consors | Now known as Othnielosaurus consors | ||||
"L. gracilis" | Considered dubious due to fragmentary remains. | ||||
Nanosaurus | N. agilis | Colorado, Brushy Basin member[23] | A small basal Neornithischian known from dozens of individuals. The holotype, YPM VP 1913, is a fragmentary skeleton from a juvenile. | ||
Othnielia | O. rex | Colorado, Brushy Basin member | Based on an isolated femur. | A small neornithischian 2 m in length. Dubious; many remains historically assigned to this taxon are now place within Nanosaurus. | |
Othnielosaurus | O. consors | Utah and Wyoming | A basal neornithischian about 2 m long. Junior synonym of Nanosaurus agilis | ||
Preprismatoolithus[24] | P. coloradensis | Colorado, Salt Wash member | Eggshell present in great abundance at the so-called "Young Egg Locality" which seems to have been a dinosaur nesting ground. Congeneric eggshell fossils are found at additional Colorado sites including the Fruita Paleontological Area, the Uravan Locality and Garden Park. | P. coloradensis is described by John Foster as being "of the prismatic basic type," with subspherical eggs about 10 cm (4 inches) in diameter.[25] This oospecies has been attributed to "hypsilophodontid" dinosaurs, although a lack of associated embryo material currently makes confirming the egg-layer's identity impossible. | |
Tichosteus | T. aequifacies | Colorado | 2 partial dorsal centra of a juvenile. | ||
T. lucasanus | Colorado | 2 partial dorsal centra. | |||
Uteodon | U. aphanoecetes | Utah, Brushy Basin member | An articulated postcranial skeleton. | Sometimes synonymized with Camptosaurus. | |
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Gargoyleosaurus | G. parkpinorum[26] | Wyoming | "Skull [and] partial postcranium."[27] | A polacanthine nodosaur known from reasonably complete fossil remains. Its skull measures 29 centimeters (11 in) in length, and its total body length is an estimated 3 to 4 meters (9.8 to 13.1 ft.). It may have weighed as much as 1 tonne (2,200 lb.). | |
Mymoorapelta | M. maysi | Colorado and Utah, Brushy Basin member [28] | "Skull fragments, portions of [three] skeletons, [and] other postcrania."[29] | Both the first ankylosaur discovered in the formation and the first known North American Jurassic ankylosaur. It probably weighed 500 kg (1,102 lbs) in life.[30] | |
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Alcovasaurus | A. longispinus | Wyoming, Brushy Basin member | "Fragmentary postcranial skeleton, adult."[31] | A dacentrurinae stegosaurid physically similar to Stegosaurus stenops but with much larger tail spines.[32] It is also similar to Kentrosaurus in having long dermal spikes on the caudal region.[33] The thighbone length was determined at 1082 millimeters. The longest spike was 86 centimeters long. Its point was broken and it is estimated the original length of the bone core at 985 millimeters. In 2019, the genus Alcovasaurus was considered a junior synonym of Miragaia[34] | |
Hesperosaurus | H. mjosi[35] | Wyoming | "Complete skeleton with skull, subadult."[36] Known only from a single specimen. | A stegosaurine stegosaurid that was slightly smaller and more primitive than Stegosaurus itself. H. mjosi had a broader skull and longer, lower plates. Considered by some to be a species of Stegosaurus[37] | |
Stegopodus | Utah | Stegopodus represent only a portion of the Morrison's stegosaur tracks, which are already rare and generally only preserve the animal's hind feet. | Stegosaur tracks which record front feet with five digits and hind feet with three weight-bearing digits. The general morphology of the tracks fit scientific predictions made eight years in advance of the erection of Stegopodus. | ||
Stegosaurus | S. armatus | Colorado, Brushy Basin | Several caudal vertebrae and assorted fragmentary postcranial elements. | S. armatus is both the first Stegosaurus to be discovered and the type species.[38] Its type specimen is poorly preserved, incomplete, and lacks diagnostic features. It has been considered dubious, with S. stenops as the neotype species for the genus. | |
S. stenops | Colorado, Utah and Wyoming, Brushy Basin member[39] | "[Two] complete skeletons with skulls, [four] braincases, at least [fifty] partial postcrania, juvenile to adult." | The best known Stegosaurus species, it has shorter limbs and larger plates than S. ungulatus.[40] | ||
S. sulcatus | Wyoming[41] | Several postcranial elements, including a possible shoulder spike. | Often considered synonymous with S. stenops,[42] it may be distinct. Potentially has a shoulder spike, otherwise unknown in Stegosaurus, despite presence in relatives.[43] | ||
S. ungulatus | Wyoming and Utah | Several partial skeletons, including a partial braincase. | S. ungulatus had longer limbs and comparatively smaller plates than the better known S. stenops.[44] Although formerly portrayed with eight tail spikes, it is now known to have had the typical four.[45] Possibly synonymous with S. stenops. | ||
Indeterminate. | New Mexico & Oklahoma | ||||
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Anomoepus | |||||
Fruitadens[46] | F. haagarorum | Colorado, Brushy Basin member[47] | A partial skull and mandible with several postcranial elements.[48] | A heterodontosaur. | |
Sauropods, the giant long-necked long-tailed four-legged herbivorous dinosaurs, are among the most common and famous Morrison fossils. A few have uncertain relationships, like "Apatosaurus" minimus (possibly a basal titanosauriform) and Haplocanthosaurus. Sauropods including Haplocanthosaurus priscus, H. delfsi, and the diplodocid Eobrontosaurus appeared in the early stages of the Morrison. The middle stages were dominated by familiar forms such as the Giraffe-like Brachiosaurus altithorax, which were uncommon, but related camarasaurids, like Camarasaurus supremus, C. grandis, C. lentus, and Cathetosaurus, were very common. Also common were long, low diplodocids, like Apatosaurus ajax, A. louisae, Brontosaurus excelsus, B. parvus, Barosaurus lentus, Diplodocus longus, D. carnegii, Galeamopus and Dyslocosaurus polyonychius.
By the late Morrison, gigantic diplodocids (or likely diplodocids) had appeared, including Diplodocus hallorum (formerly Seismosaurus), Supersaurus vivianae, Amphicoelias altus, and M. fragilimus. Smaller sauropods, such as Suuwassea emiliae from Montana, tend to be found in the northern reaches of the Morrison, near the shores of the ancient Sundance Sea, suggesting ecological niches favoring smaller body size there compared with the giants found further south.[49]
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Haplocanthosaurus | H. delfsi | Colorado, Brushy Basin member | Partial skeleton lacking a skull.[50] | Small haplocanthosaurs of indeterminate classification, ranging about 14 m (46 ft) long. | |
H. priscus | Colorado and Wyoming, Brushy Basin member | Two skulless partial skeletons. | |||
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Dyslocosaurus | D. polyonychius | Wyoming | A fragmentary forelimb and partial hindlimbs. | Phylogenetic placement is uncertain. | |
Smitanosaurus[52] | S. agilis | Colorado, Brushy Basin member | "Partial skull and cervicals." | A dicraeosaurid sauropod. Originally described as "Morosaurus" agilis. | |
Suuwassea | S. emilieae | Montana, Brushy Basin member[53] | A partial skull and some postcrania. | A dicraeosaurid about 15m in length. | |
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Amphicoelias | A. altus | Colorado, Brushy Basin member | 2 dorsal vertebrae, femur, and a pubis.[54] | Large diplodocids about 25 m (82 ft) in length.[55] | |
Apatosaurus | A. ajax | Colorado, Brushy Basin member | A partial postcranial skeleton and posterior skull. | Robust and abundantly widespread apatosaurine diplodocids reaching lengths of up to 25 m (82 ft).[56] "A." minimus likely belongs to a separate genus. | |
A. louisae | Colorado and Utah, Brushy Basin member | Four partial specimens, one of them including a skull. Three of the specimens are from DNM, one is a mid cervical from Como Bluff. | |||
"A." minimus | Wyoming | "Sacrum and pelvis."[57] | |||
Atlantosaurus | A. montanus | Colorado, Brushy Basin member[58] | A partial sacrum that cannot be distinguished from Camarasaurus or Apatosaurus.[59] | Dubious neosauropod. | |
Brontosaurus | B. excelsus | Wyoming, Brushy Basin member[60] | Two postcranial skeletons. | Previously considered a species of Apatosaurus as per Riggs (1903). | |
B. parvus | Utah and Wyoming, Salt Wash and Brushy Basin members | Three headless skeletons. | Originally called Elosaurus. | ||
B. yahnahpin[61] | Wyoming, Brushy Basin member | A partial postcranial skeleton. | An apatosaurine diplodocid slightly more primitive than Apatosaurus. Formerly placed in separate genus Eobrontosaurus. | ||
Barosaurus | B. lentus | South Dakota and Utah | 2 partial postcranial skeletons, with possibly more specimens assignable. | A diplodocid about 24 m (79 ft) in length, similar in appearance to Diplodocus.It was the rarest sauropod in the Morrison Formation.[62] | |
B. sp. | Wyoming, Utah and Oklahoma | ||||
Diplodocus | D. carnegii | Wyoming, Brushy Basin Member[63] | Known from two skulls, five partial skeletons that lack skulls and manus, and hundreds of isolated postcranial remains.[64] | Large diplodocids reaching lengths of up to 28 m (92 ft). It was one of the most abundant sauropods in the area.[65] | |
D. hallorum | New Mexico, Wyoming and Utah, Brushy Basin and Salt Wash members | At least 4 partial postcranial skeletons | Once classified as Seismosaurus.[66] [67] [68] [69] | ||
D. lacustris | Colorado, Brushy Basin member | Known from teeth and skull remains, the latter now referred to Camarasaurus. Teeth cannot be referred beyond Flagellicaudata. | |||
D. longus | Colorado, Brushy Basin member | Several caudal vertebrae.[70] | |||
Galeamopus | G. hayi | Wyoming, Brushy Basin member | Known from a partial skeleton and braincase. | ||
G. pabsti | Colorado and Wyoming, Brushy Basin member | A skull and partial postcranial skeleton, another individual known from partial skull. | |||
Kaatedocus[71] | K. siberi | Wyoming, Salt Wash member | Skull and cervical vertebrae. | ||
Parabrontopodus | |||||
Supersaurus | S. vivianae | Colorado and Wyoming, Brushy Basin member | Known from several partial postcranial skeletons. | A large diplodocid about 33–34 m 108–111 ft) in length | |
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Brachiosaurus | B. altithorax | Colorado, Brushy Basin and Salt Wash members | Several partial skeletons and a partial skull. | A large brachiosaurid about long. | |
Brontopodus | |||||
Camarasaurus | C. annae | Utah, Brushy Basin member | A dorsal vertebra. | Camarasaurs reached an adult size of about 18 m (60 ft) in length.[72] C. annae junior synonym of C. lentus. C. lewisi was originally described as Cathetosarus lewisi and was later sunk into Camarasaurus, until being considered valid once again in 2013. | |
C. grandis | Colorado, Montana and Wyoming[73] | "At least 6 partial skeletons including 2 skulls, hundreds of postcranial elements." | |||
C. lentus | Utah and Wyoming | "5 skeletons with skulls, hundreds of postcranial elements" | |||
C. lewisi | Colorado, Brushy Basin member | A nearly complete postcranial skeleton. | |||
C. supremus | Colorado and Oklahoma(?), Brushy Basin member[74] | At least 4 partial skeletons including partial skulls. Many possible postcranial remains from Oklahoma. | |||
Indeterminate. | Oklahoma, South Dakota and Texas[75] | ||||
Dystrophaeus | D. viaemalae[76] | Utah, Tidwell member | Multiple vertebrae, teeth and incomplete forelimb material. | Previously recovered as a diplodocid, now recovered as a macronarian Type material fragmentary, but recent rediscovery of type locality has discovered more material.[77] | |
Theropod dinosaurs, the carnivorous dinosaurs, came in several different types. The less derived types, the ceratosaurs and megalosaurids, included Ceratosaurus nasicornis, C. dentisulcatus, C. magnicornis, Elaphrosaurus sp., and the megalosaur Torvosaurus tanneri (including Edmarka rex). Allosaurids included the common Allosaurus fragilis (including Epanterias amplexus), Allosaurus new species, A. lucasi, and giant Saurophaganax maximus (potentially included in Allosaurus?).
Indeterminate theropod remains have been recovered in Utah. Indeterminate theropod tracks have been recovered from both Utah and Arizona.
Genus | Species | Locality | Material | Notes | Images | |
---|---|---|---|---|---|---|
Allosaurus | A. fragilis | Colorado, New Mexico, Oklahoma, South Dakota, Utah and Wyoming, Brushy Basin member[78] [79] [80] [81] | "At least [three] complete skulls, many partial skulls and skull elements, many partial and complete skeletons representing at least 60 individuals."[82] It was the most common large carnivore in the area.[83] [84] [85] | |||
A. jimmadseni[86] | Wyoming and Utah, Brushy Basin and Salt Wash members | At least 15 specimens, including several skulls. | ||||
Antrodemus | A. valens | Colorado[87] | Half of a proximal caudal centrum | Considered dubious due to fragmentary remains.[88] | ||
Creosaurus | C. atrox | Wyoming, Brushy Basin member[89] | A fragmentary skull and assorted postcrania. | Considered a junior synonym of Allosaurus. | ||
Epanterias | E. amplexus | Colorado, Brushy Basin member[90] | Several fragmentary postcranial elements of 2 individuals,[91] possible additional remains known from Wyoming.[92] | Considered dubious due to fragmentary remains. | ||
Labrosaurus | L. lucaris | Wyoming, Brushy Basin member | A fragmentary skull and several postcranial elements. | Junior synonym of A. fragilis. | ||
L. ferox | Wyoming, Brushy Basin member | A partial skull, dentaries, and fragmentary postcrania.[93] | Known from one specimen with several pathologies. Today considered a junior synonym of A. fragilis. | |||
Saurophaganax | S. maximus | Oklahoma, Brushy Basin member[94] | Several partial postcranial skeletons and fragmentary skull material. | Generally considered to be its own genus. Sometimes considered to be a species of Allosaurus.[95] [96] |
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Ceratosaurus | C. dentisulcatus | Utah | "Partial skull, vertebrae, [and] limb elements." | Large ceratosaurs grew to lengths of about 6–7 meters (20–23 ft.) in length with large nasal horns on their snouts as well as two smaller horns above the eyes. | |
C. magnicornis | Colorado[97] | "Skull [and] assorted postcrania." | |||
C. nasicornis | Colorado and Utah, Brushy Basin member | Remains of "5 individuals, including [a] nearly complete adult skeleton and subadult skeleton."[98] | |||
Indeterminate. | Wyoming[99] | ||||
Ceratosauria | Indeterminate. | Colorado | Previously referred to Elaphrosaurus,[100] [101] [102] these remains are probably not referable to that genus and are best considered indeterminate beyond Ceratosauria.[103] | ||
Elaphrosaurus | Intermediate | Colorado | Now thought to be indeterminate beyond Ceratosauria. | ||
Fosterovenator[104] | F. churei | Wyoming | Tibia, astragalus, fibula. | A fragmentary theropod which may be a ceratosaurid. | |
Coelurosaurs, the group of theropods most closely related to and including birds, included Coelurus fragilis, Ornitholestes hermanni, Tanycolagreus topwilsoni, the possible troodontid Koparion douglassi, the definite troodontid Hesperornithoides, and the early tyrannosauroid Stokesosaurus clevelandi.
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Coelurus | C. fragilis | Utah and Wyoming, Brushy Basin member | A partial skeleton in several syntypes and several fragmentary referred specimens.[105] | A basal coelurosaurian about 2.3 m (7.5 ft) long. | |
Indeterminate. | Colorado | ||||
Hesperornithoides[106] | H. miessleri | Wyoming | An early troodontid. | ||
Koparion | K. douglassi | Utah | A small theropod thought to be one of the oldest known troodontids. | ||
Palaeopteryx | P. thomsoni | Colorado, Brushy Basin member | A distal radius | ||
Ornitholestes | O. hermanni | Wyoming | "Skull and associated postcranial skeleton." | A small basal coelurosaurian about 2 m (6.5 ft) long. | |
Stokesosaurus | S. clevelandi | Utah | "Illium, associated elements and pelvic cranial material. [sic]"[107] | A possible early tyrannosauroid about 4 m (13 ft) in length. | |
Tanycolagreus | T. topwilsoni | Wyoming and Utah | A basal coelurosaurian about 3.4 m (11.3 ft) long, similar in appearance to Coelurus. | ||
Genus | Species | Locality | Material | Notes | Images |
---|---|---|---|---|---|
Edmarka | E. rex |
| Possibly a junior synonym of Torvosaurus tanneri. | ||
Marshosaurus | M. bicentesimus | Colorado and Utah | Partial skeleton, including part of a skull.[108] | A medium-sized piatnitzkysaurid megalosauroid about 6 m (20 ft) in length. | |
Torvosaurus | T. tanneri | Colorado, Utah and Wyoming, Brushy Basin member | "Partial skeletons of at least [three] individuals."[109] | A large, robust megalosaurid reaching lengths of up to .[110] One of the largest carnivores of the formation. | |
Dinosaur eggs have been found in Utah.
Morrison ornithopod trace fossils are represented by three toed tracks which are generally small. The toes of Morrison ornithopod tracks are usually more widely splayed than the theropod tracks preserved in the formation.
Stegosaur tracks were first recognized in 1996 from a hindprint-only trackway discovered at the Cleveland-Lloyd quarry, which is located near Price, Utah.[111] Two years later, a new ichnogenus called Stegopodus was erected for another set of stegosaur tracks which were found near Arches National Park, also in Utah. Unlike the first, this trackway preserved traces of the forefeet. Fossil remains indicate that stegosaurs have five digits on the forefeet and three weight-bearing digits on the hind feet. From this, scientists were able to successfully predict the appearance of stegosaur tracks in 1990, six years in advance of the first actual discovery of Morrison stegosaur tracks. Since the erection of Stegopodus, more trackways have been found, however none have preserved traces of the front feet, and stegosaur traces remain rare.
Indeterminate theropod tracks have been recovered from both Utah and Arizona.[112]