Othnielosaurus is a genus of ornithischian dinosaur that lived about 155 to 148 million years ago, during the Late Jurassic-age Morrison Formation of the western United States. It is named in honor of famed paleontologist Othniel Charles Marsh, and was formerly assigned to the genus Laosaurus. This genus was coined to hold fossils formerly included in Othnielia, which is based on remains that may be too sparse to hold a name. O.C. Marsh named several species and genera in the late 19th century that have come to be recognized as hypsilophodonts or hypsilophodont-like animals, including Nanosaurus agilis, "N." rex (Othnielia), Laosaurus celer, L. consors, and L. gracilis. This taxonomy has become very complicated, with numerous attempts at revision in the years since; Othnielosaurus is part of decades of research to untangle the taxonomy left behind by Marsh and his rival Edward Drinker Cope from the Bone Wars. Othnielosaurus has usually been classified as a hypsilophodont, a type of generalized small bipedal herbivore or omnivore, although recent research has called this and the existence of a distinct group of hypsilophodonts into question.
|Fossil, Paleontology Museum of Zurich|
Othnielosaurus is known from material from all parts of the body, including two good skeletons, although the skull is still poorly known (note that earlier references use a multitude of names for this material, with most of them since 1977 using Othnielia rex). Othnielosaurus was a small animal, 2 meters (6.6 ft) or less in length and 10 kilograms (22 lb) or less in weight.
It was a bipedal dinosaur with short forelimbs and long hindlimbs with large processes for muscle attachments. The hands were short and broad with short fingers. From the partial type skull and the skull on the possible specimen "Barbara", the head was small. It had small leaf-shaped cheek teeth (triangular and with small ridges and denticles lining the front and back edges), and premaxillary teeth with less ornamentation. Like several related dinosaurs, such as Hypsilophodon, Thescelosaurus, and Talenkauen, Othnielosaurus had thin plates lying along the ribs. Called intercostal plates, these structures were cartilaginous in origin.
In 1877, Marsh named two species of Nanosaurus in separate publications, based on partial remains from the Morrison Formation of Garden Park, Colorado. One paper described N. agilis, based on YPM 1913, with remains including impressions of a dentary, and postcranial bits including an ilium, thigh bones, shin bones, and a fibula. The other paper named N. rex, a second species which Marsh based on YPM 1915 (also called 1925 in Galton, 2007), a complete thigh bone. He regarded both species as small ("fox-sized") animals. He assigned this genus to the now-abandoned family Nanosauridae.
The next year, he named the new genus Laosaurus on material collected by Samuel Wendell Williston from Como Bluff, Wyoming. Two species were named: the type species L. celer, based on parts of eleven vertebrae (YPM 1875); and the "smaller" L. gracilis, originally based on a back vertebra's centrum, a caudal centrum, and part of an ulna (review by Peter Galton in 1983 finds the specimen to now consist of thirteen back and eight caudal centra, and portions of both hindlimbs).
A third species, L. consors, was established by Marsh in 1894 for YPM 1882, which consists of most of one articulated skeleton and part of at least one other individual. The skull was only partially preserved, and the fact that the vertebrae were represented only by centra suggests a partially grown individual. Galton (1983) notes that much of the current mounted skeleton was restored in plaster, or had paint applied.
These animals attracted little professional attention until the 1970s and 1980s, when Peter Galton reviewed many the "hypsilophodonts" in a series of papers. In 1973, he and Jim Jensen described a partial skeleton (BYU ESM 163 as of Galton, 2007) missing the head, hands, and tail as Nanosaurus (?) rex, which had been damaged by other collectors prior to description. By 1977, he had determined that Nanosaurus agilis was quite different from N. rex and the new skeleton, and coined Othnielia for N. rex. The 1977 reference, somewhat buried in a paper concerning the transcontinental species of Dryosaurus, did not elaborate, but did assign Laosaurus consors and L. gracilis to the new genus, and considered L. celer a nomen nudum. The publication of Drinker further complicated matters.
Galton (2007), in his discussion of the teeth of Morrison ornithischians, concluded that the holotype femur of Othniela rex is not diagnostic, and reassigned the BYU skeleton to Laosaurus consors, which is based on more diagnostic material. As the genus Laosaurus is also based on nondiagnostic material, he gave the species L. consors its own genus, Othnielosaurus. As a result, in practical terms, what had been thought of as Othnielia is now known as Othnielosaurus consors. In a 2018 paper, however, Carpenter and Galton (2018) treated Othnielosaurus, along with Othnielia, as junior synonyms of Nanosaurus (they acknowledged possible disagreement with their taxonomic action).
Othnielosaurus (previously under the names Laosaurus, Nanosaurus, and Othnielia) has typically been regarded as a hypsilophodont ornithopod, a member of a nebulous and poorly defined group of small, running herbivorous dinosaurs. This was challenged by Robert Bakker et al. in 1990. In their description of the new taxon Drinker nisti, they split Othnielia into two species (O. rex and O. consors) and placed "othnieliids" as more basal than hypsilophodontids. With recent analyses suggesting a paraphyletic Hypsilophodontidae, the general idea of "othnielids" as basal to other hypsilophodonts has been supported, although Drinker has been controversial because virtually nothing new has been published on it since its description. Other basal ornithopods have sometimes been linked to Othnielosaurus, particularly Hexinlusaurus, considered by at least one author to be a species of "Othnielia", O. multidens. New studies concur with the hypothesis that Othnielosaurus is more basal than other traditional hypsilophodonts, but go even farther and remove the genus from Ornithopoda and the larger group Cerapoda, which also includes Ceratopsians and pachycephalosaurs.
Othnielosaurus was one of the smaller members of the diverse Morrison Formation dinosaur fauna, diminutive in comparison to the giant sauropods. The Morrison Formation is interpreted as a semiarid environment with distinct wet and dry seasons, and flat floodplains. Vegetation varied from river-lining gallery forests of conifers, tree ferns, and ferns, to fern savannas with rare trees. It has been a rich fossil hunting ground, holding fossils of green algae, fungi, mosses, horsetails, ferns, cycads, ginkgoes, and several families of conifers. Other fossils discovered include bivalves, snails, ray-finned fishes, frogs, salamanders, turtles, sphenodonts, lizards, terrestrial and aquatic crocodylomorphans, several species of pterosaur, numerous dinosaur species, and early mammals such as docodonts, multituberculates, symmetrodonts, and triconodonts. Such dinosaurs as the theropods Ceratosaurus, Allosaurus, Ornitholestes, and Torvosaurus, the sauropods Apatosaurus, Brachiosaurus, Camarasaurus, and Diplodocus, and the ornithischians Camptosaurus, Dryosaurus, and Stegosaurus are known from the Morrison. Othnielosaurus is present in stratigraphic zones 2-5.
Typically, Othnielosaurus has been interpreted like other hypsilophodonts as a small, swift herbivore, although Bakker (1986) interpreted the possibly related Nanosaurus as an omnivore. This idea has had some unofficial support, but little in the formal literature; description of more complete skull remains will be needed to test this hypothesis.
Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.Avetheropoda
Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs.Cerapoda
Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.Dinosauriformes
Dinosauriformes is a clade of archosaurian reptiles that include the dinosaurs and their most immediate relatives. All dinosauriformes are distinguished by several features, such as shortened forelimbs and a partially to fully perforated acetabulum, the hole in the hip socket traditionally used to define dinosaurs. The oldest known member is Asilisaurus, dating to about 245 million years ago in the Anisian age of the middle Triassic period.Drinker nisti
Drinker (named after the palaentologist Edward Drinker Cope) is a genus of neornithischian dinosaur from the late Jurassic period of North America. Although based on good remains, it remains obscure due to a lack of post-naming publications.Jeholosauridae
Jeholosaurids were herbivorous neornithischian dinosaurs from the Cretaceous Period (Aptian - Santonian, with a possible Campanian record) of Asia. The family was first proposed by Han et al. in 2012. The jeholosaurids were defined as those ornithischians more closely related to Jeholosaurus shangyuanensis than to Hypsilophodon foxii, Iguanodon bernissartensis, Protoceratops andrewsi, Pachycephalosaurus wyomingensis, or Thescelosaurus neglectus. The Jeholosauridae includes the type genus Jeholosaurus and Yueosaurus.Jingshanosaurus
Jingshanosaurus (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period.Kimmeridgian
In the geologic timescale, the Kimmeridgian is an age or stage in the Late or Upper Jurassic epoch or series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxfordian and precedes the Tithonian.Laosaurus
Laosaurus (meaning "stone or fossil lizard") is a genus of neornithischian dinosaur. The type species is Laosaurus celer, first described by O.C. Marsh in 1878 from remains from the Oxfordian-Tithonian-age Upper Jurassic Morrison Formation of Wyoming. The validity of this genus is doubtful because it is based on fragmentary fossils. A second species from the Morrison Formation, L. gracilis, and a species from the late Cretaceous of Alberta, Laosaurus minimus, are also considered dubious.Melanorosauridae
The Melanorosauridae were a family of sauropodomorph dinosaurs which lived during the Late Triassic and Early Jurassic. The name Melanorosauridae was first coined by Friedrich von Huene in 1929. Huene assigned several families of dinosaurs to the infraorder "Prosauropoda": the Anchisauridae, the Plateosauridae, the Thecodontosauridae, and the Melanorosauridae. Since then, these families have undergone numerous revisions. Galton and Upchurch (2004) considered Camelotia, Lessemsaurus, and Melanorosaurus members of the family Melanorosauridae. A more recent study by Yates (2007) indicates that the melanorosaurids were instead early sauropods.Nanosaurus
Nanosaurus ("small or dwarf lizard") is the name given to a genus of dinosaur from the Late Jurassic. Described by Othniel Charles Marsh in 1877, it is a poorly known ornithischian of uncertain affinities. Its fossils are known from the Morrison Formation of Colorado and possibly Wyoming. It has often been illustrated in the popular literature (as a "tiny dinosaur"), leaving the impression that more is known about it than actually is. Most representations are based on remains referred to Othnielosaurus.Neornithischia
Neornithischia ("new ornithischians") is a clade of the dinosaur order Ornithischia. They are the sister group of the Thyreophora within the clade Genasauria. Neornithischians are united by having a thicker layer of asymmetrical enamel on the inside of their lower teeth. The teeth wore unevenly with chewing and developed sharp ridges that allowed neornithischians to break down tougher plant food than other dinosaurs. Neornithischians include a variety of basal forms historically known as "hypsilophodonts", including the Parksosauridae; in addition, there are derived forms classified in the groups Marginocephalia and Ornithopoda. The former includes clades Pachycephalosauria and Ceratopsia, while the latter typically includes Hypsilophodon and the more derived Iguanodontia.Neotheropoda
Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and the only group of theropods who survived the Triassic–Jurassic extinction event. Yet all of the neotheropods became extinct during the early Jurassic period except for Averostra.Orionides
Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.Orodrominae
Orodrominae is a subfamily of parksosaurid dinosaurs from the Cretaceous of North America and Asia.Othnielia
Othnielia is a dubious genus of ornithischian dinosaur, named after its original describer, Professor Othniel Charles Marsh, an American paleontologist of the 19th century.Riojasauridae
Riojasauridae is a family of sauropod-like dinosaurs from the Upper Triassic. It is known primarily from the genera Riojasaurus and Eucnemesaurus. Sites containing Riojasauridae include the Lower Elliot Formation of Orange Free State, South Africa (where fossils of Eucnemesaurus have been found), and Ischigualasto, in La Rioja Province, Argentina ( where fossils of Riojasaurus have been recovered).Tithonian
In the geological timescale, the Tithonian is the latest age of the Late Jurassic epoch or the uppermost stage of the Upper Jurassic series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the Kimmeridgian and followed by the Berriasian stage (part of the Cretaceous).Xixiposaurus
Xixiposaurus is a genus of prosauropod dinosaur which existed in what is now Lower Lufeng Formation, China during the lower Jurassic period. It was first named by Sekiya Toru in 2010 and the type species is Xixiposaurus suni.