Temporal range: Late Jurassic, 155–145 Ma
|D. altus skeleton in Japan|
|Species:||† D. altus|
(Marsh, 1878 [originally Laosaurus altus])
Dryosaurus (// DRY-o-SAWR-əs; meaning 'tree lizard', Greek δρυς/drys meaning 'tree, oak' and σαυρος/sauros meaning 'lizard'; the name reflects the forested habitat, not a vague oak-leaf shape of its cheek teeth as is sometimes assumed) is a genus of an ornithopod dinosaur that lived in the Late Jurassic period. It was an iguanodont (formerly classified as a hypsilophodont). Fossils have been found in the western United States, and were first discovered in the late 19th century. Valdosaurus canaliculatus and Dysalotosaurus lettowvorbecki were both formerly considered to represent species of Dryosaurus.
Discovery and naming
In 1876, Samuel Wendell Williston in Albany County, Wyoming discovered the remains of small euornithopods. In 1878, Professor Othniel Charles Marsh named these as a new species of Laosaurus, Laosaurus altus. The specific name altus, meaning "tall" in Latin, refers to it being larger than Laosaurus celer. In 1894, Marsh made the taxon a separate genus, Dryosaurus. The generic name is derived from the Greek δρῦς, drys, "tree, oak", referring to a presumed forest-dwelling life mode. Later it was often assumed to have been named after an oak-leaf shape of its cheek teeth, which, however, is absent. The type species remains Laosaurus altus, the combinatio nova is Dryosaurus altus.
The holotype, YPM 1876, was found in a layer of the Upper Brushy Basin Member of the Morrison Formation, dating from the Tithonian. It consists of a partial skeleton including a rather complete skull and lower jaws. Several other fossils from Wyoming have been referred to Dryosaurus altus. They include specimens YPM 1884: the rear half of a skeleton; AMNH 834: a partial skeleton lacking the skull from the Bone Cabin Quarry; and CM 1949: a rear half of a skeleton dug up in 1905 by William H. Utterback. From 1922 onwards in Utah, Earl Douglass discovered Dryosaurus remains at the Dinosaur National Monument. These include CM 11340: the front half of a skeleton of a very young individual; CM 3392: a skeleton with skull but lacking the tail; CM 11337: a fragmentary skeleton of a juvenile; and DNM 1016: a left ilium dug up by technician Jim Adams. Other fossils were found in Colorado. In Lily Park, Moffat County, James Leroy Kay and Albert C. Lloyd in 1955 recovered CM 21786, a skeleton lacking skull and neck. From Sheetz’ Quarry 1, at Uravan, Montrose County, in 1973 Peter Malcolm Galton and James Alvin Jensen described specimen BYU ESM-171R found by Rodney Dwayne Scheetz and consisting of some vertebrae, a left lower jaw, a left forelimb and two hindlimbs. Gregory S. Paul in 2010 suggested that the Utah material represented a separate species.
Apart from Dryosaurus altus, several other species have been named in the genus. The first of these was created accidentally when in 1903 Giuseppe de Stefano renamed Crocodilus phosphaticus Thomas 1893 into Dryosaurus phosphaticus; he had intended to call it Dyrosaurus phosphaticus. This was only emended by Éric Buffetaut in 1981.
Dryosaurus had a long neck, long, slender legs and a long, stiff tail. Its arms, however, with five fingers on each hand, were short. Known specimens were about 8 to 14 feet (2.4 to 4.3 m) long and weighed 170 to 200 pounds (77 to 91 kg). However, the adult size is unknown, as no known adult specimens of the genus have been found.
Dryosaurus had a horny beak and cheek teeth and, like other ornithopods, was a herbivore. Some scientists suggest that it had cheek-like structures to prevent the loss of food while the animal processed it in the mouth.
Diet and dentition
The teeth of Dryosaurus were, according to museum curator John Foster, characterized by "a strong median ridge on the lateral surface."Dryosaurus subsisted primarily on low growing vegetation in ancient floodplains.
Growth and development
A Dryosaurus hatchling found at Dinosaur National Monument in Utah confirmed that Dryosaurus followed similar patterns of craniofacial development to other vertebrates; the eyes were proportionally large while young and the muzzle proportionally short. As the animal grew, its eyes became proportionally smaller and its snout proportionally longer.
Paleobiogeography and fossil distribution
The Dryosaurus holotype specimen YPM 1876 was discovered in Reed’s YPM Quarry 5, in the Upper Brushy Basin Member, of the Morrison Formation. In the Late Jurassic Morrison formation of Western North America, Dryosaurus remains have been recovered from stratigraphic zones 2-6. A spectacular digsite near Uravan, Colorado held hundreds of D. altus fossils which represented multiple stages of the animal's life cycle. This formation is a sequence of shallow marine and alluvial sediments which, according to radiometric dating, ranges between 156.3 million years old (Ma) at its base, to 146.8 million years old at the top, which places it in the late Oxfordian, Kimmeridgian, and early Tithonian stages of the Late Jurassic period. In 1877 this formation became the center of the Bone Wars, a fossil-collecting rivalry between early paleontologists Othniel Charles Marsh and Edward Drinker Cope. The Morrison Formation is interpreted as a semiarid environment with distinct wet and dry seasons. The Morrison Basin where dinosaurs lived, stretched from New Mexico to Alberta and Saskatchewan, and was formed when the precursors to the Front Range of the Rocky Mountains started pushing up to the west. The deposits from their east-facing drainage basins were carried by streams and rivers and deposited in swampy lowlands, lakes, river channels and floodplains. This formation is similar in age to the Solnhofen Limestone Formation in Germany and the Tendaguru Formation in Tanzania.
The Morrison Formation records an environment and time dominated by gigantic sauropod dinosaurs such as Camarasaurus, Barosaurus, Diplodocus, Apatosaurus and Brachiosaurus. Dinosaurs that lived alongside Dryosaurus included the herbivorous ornithischians Camptosaurus, Stegosaurus and Othnielosaurus. Predators in this paleoenvironment included the theropods Saurophaganax, Torvosaurus, Ceratosaurus, Marshosaurus, Stokesosaurus, Ornitholestes and Allosaurus, which accounting for 70 to 75% of theropod specimens and was at the top trophic level of the Morrison food web. Other vertebrates that shared this paleoenvironment included bivalves, snails, ray-finned fishes, frogs, salamanders, turtles, sphenodonts, lizards, terrestrial and aquatic crocodylomorphans, and several species of pterosaur. Early mammals were present in this region, such as docodonts, multituberculates, symmetrodonts, and triconodonts. The flora of the period has been revealed by fossils of green algae, fungi, mosses, horsetails, cycads, ginkgoes, and several families of conifers. Vegetation varied from river-lining forests of tree ferns, and ferns (gallery forests), to fern savannas with occasional trees such as the Araucaria-like conifer Brachyphyllum.
- Tom R. Hübner and Oliver W. M. Rauhut (2010). "A juvenile skull of Dysalotosaurus lettowvorbecki (Ornithischia: Iguanodontia), and implications for cranial ontogeny, phylogeny, and taxonomy in ornithopod dinosaurs". Zoological Journal of the Linnean Society 160 (2): 366–396. doi:10.1111/j.1096-3642.2010.00620.x.
- McDonald AT, Kirkland JI, DeBlieux DD, Madsen SK, Cavin J; et al. (2010). Farke, Andrew Allen, ed. "New Basal Iguanodonts from the Cedar Mountain Formation of Utah and the Evolution of Thumb-Spiked Dinosaurs". PLoS ONE 5 (11): e14075. doi:10.1371/journal.pone.0014075. PMC 2989904. PMID 21124919.
- Galton, P.M., 1977. "The Upper Jurassic dinosaur Dryosaurus and a Laurasia-Gondwana connection in the Upper Jurassic", Nature 268(5617): 230-232
- O.C. Marsh, 1878, "Principal characters of American Jurassic dinosaurs. Part I", American Journal of Science and Arts 16: 411-416
- O.C. Marsh, 1894, "The typical Ornithopoda of the American Jurassic", American Journal of Science, series 3 48: 85-90
- Gilmore C.W., 1925, "Osteology of ornithopodous dinosaurs from the Dinosaur National Monument, Utah. Camptosaurus medius, Dryosaurus altus, Laosaurus gracilis", Memoirs of the Carnegie Museum 10: 385-409
- Galton, P.M. & Jensen, J.A., 1973, "Small bones of the hypsilophodontid dinosaur Dryoraurus altus from the Upper Jurassic of Colorado", Great Basin Nature, 33: 129-132
- Paul, G.S., 2010, The Princeton Field Guide to Dinosaurs, Princeton University Press p. 281
- G. de Stefano, 1903, "Nuovi rettili degli strati a fosfato della Tunisia", Bollettino della Societa Geologica Italiana 22: 51-80
- R.T.J. Moody and E. Buffetaut, 1981, "Notes on the systematics and palaeoecology of the crocodiles and turtles of the Metaloui Phosphates (Eocene) of southern Tunisia", Tertiary Research 3(3): 125-140
- Horner, John R.; de Ricqlés, Armand; Padian, Kevin; and Scheetz, Rodney D. (2009). "Comparative long bone histology and growth of the "hysilophodontid" dinosaurs Orodromeus makelai, Dryosaurus altus, and Tenontosaurus tillettii (Ornithischia: Euornithopoda)". Journal of Vertebrate Paleontology 29 (3): 734–747. doi:10.1671/039.029.0312.
- Marshall (1999) pp. 138-139
- "Dryosaurus altus," Foster (2007) pp. 218-219.
- "Appendix," Foster (2007) pp. 327-329.
- Trujillo, K.C.; Chamberlain, K.R.; Strickland, A. (2006). "Oxfordian U/Pb ages from SHRIMP analysis for the Upper Jurassic Morrison Formation of southeastern Wyoming with implications for biostratigraphic correlations". Geological Society of America Abstracts with Programs 38 (6): 7.
- Bilbey, S.A. (1998). "Cleveland-Lloyd Dinosaur Quarry - age, stratigraphy and depositional environments". In Carpenter, K.; Chure, D.; and Kirkland, J.I. (eds.). The Morrison Formation: An Interdisciplinary Study. Modern Geology 22. Taylor and Francis Group. pp. 87–120. ISSN 0026-7775.
- Russell, Dale A. (1989). An Odyssey in Time: Dinosaurs of North America. Minocqua, Wisconsin: NorthWord Press. pp. 64–70. ISBN 978-1-55971-038-1.
- Foster, J. (2007). "Appendix." Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 327-329.
- Foster, John R. (2003). Paleoecological Analysis of the Vertebrate Fauna of the Morrison Formation (Upper Jurassic), Rocky Mountain Region, U.S.A. New Mexico Museum of Natural History and Science Bulletin, 23. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science. p. 29.
- Carpenter, Kenneth (2006). "Biggest of the big: a critical re-evaluation of the mega-sauropod Amphicoelias fragillimus". In Foster, John R.; and Lucas, Spencer G. (eds.). Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin, 36. Albuquerque, New Mexico: New Mexico Museum of Natural History and Science. pp. 131–138.
- Foster, J. (2007). Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. 389pp.
- Palmer, D., ed. (1999). The Marshall Illustrated Encyclopedia of Dinosaurs and Prehistoric Animals. London: Marshall Editions. pp. 138–139. ISBN 1-84028-152-9.