Marshosaurus Bicentesimus
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''Marshosaurus'' is a genus of medium-sized carnivorous
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
, belonging to the
Megalosauroidea Megalosauroidea (meaning 'great/big lizard forms') is a superfamily (or clade) of tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period. The group is defined as '' Megalosaurus bucklandii'' and all taxa s ...
, from the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
Morrison Formation of
Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...
and possibly
Colorado Colorado (, other variants) is a state in the Mountain states, Mountain West subregion of the Western United States. It encompasses most of the Southern Rocky Mountains, as well as the northeastern portion of the Colorado Plateau and the wes ...
.


Description

''Marshosaurus'' was medium-sized for a theropod. In 2010,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
estimated its length at and its weight at . The holotype ilium has a length of . If the cranial material is correctly referred, the skull was about long. In 2012, Matthew Carrano established one
autapomorphy In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...
, a unique derived trait of the holotype: the suture between the pubic peduncle and the
pubic bone In vertebrates, the pubic region ( la, pubis) is the most forward-facing ( ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior ...
is convex, curving upwards, at the front and concave at the rear.


Discovery and naming

During the 1960s, over fourteen thousand fossil bones were uncovered at the Cleveland-Lloyd Quarry in central Utah. The majority of these belonged to ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'' but some were of at least two theropods new to science. In 1974 one of these was named by James Henry Madsen Jr. as the genus ''
Stokesosaurus ''Stokesosaurus'' (meaning "Stokes' lizard") is a genus of small (around in length), carnivorous early tyrannosauroid theropod dinosaurs from the late Jurassic period of Utah, United States. History From 1960 onwards Utah geologist William ...
''. In 1976 the second was by Madsen named as the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
''Marshosaurus bicentesimus''. The generic name honoured the nineteenth century paleontologist Professor
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among ...
, who described many dinosaur fossils during the
Bone Wars The Bone Wars, also known as the Great Dinosaur Rush, was a period of intense and ruthlessly competitive fossil hunting and discovery during the Gilded Age of American history, marked by a heated rivalry between Edward Drinker Cope (of the Ac ...
. The specific name was chosen "in honor of the bicentennial of the United States of America". The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
, UMNH VP 6373, was found in a layer of the
Brushy Basin Member The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock 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, ...
of the Morrison Formation dating from the late
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
, approximately 155 - 152 mya. It is a left ilium, or upper pelvis bone. The
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). O ...
s consisted of three bones: the ischia UMNH VP 6379 and UMNH VP 380 and the
pubic bone In vertebrates, the pubic region ( la, pubis) is the most forward-facing ( ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior ...
UMNH VP 6387. Three ilia and six jaw fragments were provisionally referred. The material represents at least three individuals. In 1991, Brooks Britt referred tail vertebrae from Colorado, because they resembled non-identified tail vertebrae fragments from the Cleveland-Lloyd Dinosaur Quarry. In 1993 a partial skeleton, CMNH 21704, from the Dinosaur National Monument was referred because its dorsal neural spines resembled non-identified spines from the Cleveland-Lloyd Dinosaur Quarry. This specimen was also the subject of a 1997 SVP abstract.


Classification

Madsen originally was unsure about the phylogenetic position of ''Marshosaurus'', placing it as Theropoda ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
''. Some later analyses showed ''Marshosaurus'' to be a member of
Avetheropoda Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs. Definition Avetheropoda was named by Gregory S. Paul in 1988, and was first defined as a clade by Currie and Padia ...
, a group of more bird-like theropods including ''
Tyrannosaurus ''Tyrannosaurus'' is a genus of large theropod dinosaur. The species ''Tyrannosaurus rex'' (''rex'' meaning "king" in Latin), often called ''T. rex'' or colloquially ''T-Rex'', is one of the best represented theropods. ''Tyrannosaurus'' live ...
'', ''
Velociraptor ''Velociraptor'' (; ) is a genus of small dromaeosaurid dinosaur that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago. Two species are currently recognized, although others have been assigned in the p ...
'' and ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
''. However,
Roger Benson Roger is a given name, usually masculine, and a surname. The given name is derived from the Old French personal names ' and '. These names are of Germanic origin, derived from the elements ', ''χrōþi'' ("fame", "renown", "honour") and ', ' ( ...
(2010) found it to be a
megalosauroid Megalosauroidea (meaning 'great/big lizard forms') is a Taxonomic rank, superfamily (or clade) of Tetanurae, tetanuran theropod dinosaurs that lived from the Middle Jurassic to the Late Cretaceous period. The group is defined as ''Megalosaurus, M ...
in a phylogenetic analysis of ''
Megalosaurus ''Megalosaurus'' (meaning "great lizard", from Greek , ', meaning 'big', 'tall' or 'great' and , ', meaning 'lizard') is an extinct genus of large carnivorous theropod dinosaurs of the Middle Jurassic period (Bathonian stage, 166 million years ...
'' and 40 other theropods. The position of ''Marshosaurus'' in the evolutionary tree, as a possible member of the
Piatnitzkysauridae Piatnitzkysauridae is an extinct family of megalosauroid or basal Allosauroid dinosaurs. It only consists of three known dinosaur genera: '' Condorraptor'', ''Marshosaurus'', and '' Piatnitzkysaurus.'' The most complete and well known member of ...
, is shown by the cladogram below.


Paleopathology

One right ilium of a ''Marshosaurus bicentesimus'' is deformed by "an undescribed pathology" which probably originated as a consequence of injury. Another specimen has a pathological rib. In a 2001 study conducted by Bruce Rothschild and other paleontologists, five foot bones referred to ''Marshosaurus'' were examined for signs of
stress fracture A stress fracture is a fatigue-induced bone fracture caused by repeated stress over time. Instead of resulting from a single severe impact, stress fractures are the result of accumulated injury from repeated submaximal loading, such as running or ...
, but none were found.


Paleoecology


Habitat

The Morrison Formation is a sequence of shallow marine and alluvial sediments which, according to
radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares ...
, 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 In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxford ...
, and early Tithonian
stages Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * S ...
of the Late Jurassic period. This formation is interpreted as a
semiarid A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of semi- ...
environment with distinct wet and
dry season The dry season is a yearly period of low rainfall, especially in the tropics. The weather in the tropics is dominated by the tropical rain belt, which moves from the northern to the southern tropics and back over the course of the year. The te ...
s. The Morrison Basin where dinosaurs lived, stretched from New Mexico to Alberta and Saskatchewan, and was formed when the precursors to the
Front Range The Front Range is a mountain range of the Southern Rocky Mountains of North America located in the central portion of the U.S. State of Colorado, and southeastern portion of the U.S. State of Wyoming. It is the first mountain range encountered ...
of the Rocky Mountains started pushing up to the west. The deposits from their east-facing
drainage basin A drainage basin is an area of land where all flowing surface water converges to a single point, such as a river mouth, or flows into another body of water, such as a lake or ocean. A basin is separated from adjacent basins by a perimeter, ...
s were carried by streams and
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of w ...
s and deposited in swampy lowlands, lakes, river channels and
floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river which stretches from the banks of its channel to the base of the enclosing valley walls, and which experiences flooding during periods of high discharge.Goudi ...
s. This formation is similar in age to the Solnhofen Limestone Formation in
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
and the
Tendaguru Formation The Tendaguru Formation, or Tendaguru Beds are a highly fossiliferous formation and Lagerstätte located in the Lindi Region of southeastern Tanzania. The formation represents the oldest sedimentary unit of the Mandawa Basin, overlying Neoproter ...
in
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands ...
. In 1877 this formation became the center of the Bone Wars, a fossil-collecting rivalry between early paleontologists
Othniel Charles Marsh Othniel Charles Marsh (October 29, 1831 – March 18, 1899) was an American professor of Paleontology in Yale College and President of the National Academy of Sciences. He was one of the preeminent scientists in the field of paleontology. Among ...
and Edward Drinker Cope.


Paleofauna

The Morrison Formation records an environment and time dominated by gigantic sauropod dinosaurs such as '' Camarasaurus'',''
Brachiosaurus ''Brachiosaurus'' () is a genus of sauropod dinosaur that lived in North America during the Late Jurassic, about 154to 150million years ago. It was first described by American paleontologist Elmer S. Riggs in 1903 from fossils found in th ...
'', ''
Barosaurus ''Barosaurus'' ( ) was a giant, long-tailed, long-necked, plant-eating sauropod dinosaur closely related to the more familiar ''Diplodocus''. Remains have been found in the Morrison Formation from the Upper Jurassic Period of Utah and South Da ...
'', ''
Diplodocus ''Diplodocus'' (, , or ) was a genus of diplodocid sauropod dinosaurs, whose fossils were first discovered in 1877 by S. W. Williston. The generic name, coined by Othniel Charles Marsh in 1878, is a neo-Latin term derived from Greek δι ...
'', and ''
Apatosaurus ''Apatosaurus'' (; meaning "deceptive lizard") is a genus of herbivorous sauropod dinosaur that lived in North America during the Late Jurassic period. Othniel Charles Marsh described and named the first-known species, ''A. ajax'', in 1877, ...
''. Dinosaurs that lived alongside ''Marshosaurus'' included the herbivorous
ornithischians Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek st ...
''
Camptosaurus ''Camptosaurus'' ( ) is a genus of plant-eating, beaked ornithischian dinosaurs of the Late Jurassic period of western North America and possibly also Europe. The name means 'flexible lizard' ( Greek (') meaning 'bent' and (') meaning 'li ...
'', ''
Dryosaurus ''Dryosaurus'' ( , meaning 'tree lizard', Greek ' () meaning 'tree, oak' and () 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 ...
'', ''
Stegosaurus ''Stegosaurus'' (; ) is a genus of herbivorous, four-legged, armored dinosaur from the Late Jurassic, characterized by the distinctive kite-shaped upright plates along their backs and spikes on their tails. Fossils of the genus have been fou ...
'' and '' Othnielosaurus''. Predators in this paleoenvironment included the
theropods Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally ca ...
''
Saurophaganax ''Saurophaganax'' ("lord of lizard-eaters") is a genus of large allosaurid dinosaur from the Morrison Formation of Late Jurassic (latest Kimmeridgian age, about 151 million years ago) Oklahoma, United States.Turner, C.E. and Peterson, F., (1999) ...
'', ''
Torvosaurus ''Torvosaurus'' () is a genus of carnivorous megalosaurid theropod dinosaur that lived approximately 165 to 148 million years ago during the late Middle and Late Jurassic period (Callovian to Tithonian stages) in what is now Colorado, Portuga ...
'', ''
Ceratosaurus ''Ceratosaurus'' (from Greek κέρας/κέρατος, ' meaning "horn" and σαῦρος ' meaning "lizard") was a carnivorous theropod dinosaur in the Late Jurassic period ( Kimmeridgian to Tithonian). The genus was first described in 1 ...
'', ''
Stokesosaurus ''Stokesosaurus'' (meaning "Stokes' lizard") is a genus of small (around in length), carnivorous early tyrannosauroid theropod dinosaurs from the late Jurassic period of Utah, United States. History From 1960 onwards Utah geologist William ...
'', ''
Ornitholestes ''Ornitholestes'' (meaning "bird robber") is a small theropod dinosaur of the late Jurassic (Brushy Basin Member of the Morrison Formation, middle Kimmeridgian age, about 154 million years agoTurner, C.E. and Peterson, F., (1999). "Biostratigrap ...
'' and ''
Allosaurus ''Allosaurus'' () is a genus of large carnosaurian theropod dinosaur that lived 155 to 145 million years ago during the Late Jurassic epoch ( Kimmeridgian to late Tithonian). The name "''Allosaurus''" means "different lizard" alludin ...
'', which accounted for 70 to 75% of theropod specimens and was at the top
trophic level The trophic level of an organism is the position it occupies in a food web. A food chain is a succession of organisms that eat other organisms and may, in turn, be eaten themselves. The trophic level of an organism is the number of steps it ...
of the Morrison
food web A food web is the natural interconnection of food chains and a graphical representation of what-eats-what in an ecological community. Another name for food web is consumer-resource system. Ecologists can broadly lump all life forms into one o ...
. ''Stegosaurus'' is commonly found at the same sites as ''Allosaurus'', ''Apatosaurus'', ''Camarasaurus'', and ''Diplodocus''. Early mammals were present in this region, such as
docodonts Docodonta is an order of extinct mammaliaforms that lived during the Mesozoic, from the Middle Jurassic to Early Cretaceous. They are distinguished from other early mammaliaforms by their relatively complex molar teeth, from which the order ge ...
,
multituberculates Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, a ...
, symmetrodonts, and triconodonts. The flora of the period has been revealed by fossils of green algae,
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
, mosses,
horsetails ''Equisetum'' (; horsetail, snake grass, puzzlegrass) is the only living genus in Equisetaceae, a family of ferns, which reproduce by spores rather than seeds. ''Equisetum'' is a " living fossil", the only living genus of the entire subclass ...
,
cycad Cycads are seed plants that typically have a stout and woody (ligneous) trunk with a crown of large, hard, stiff, evergreen and (usually) pinnate leaves. The species are dioecious, that is, individual plants of a species are either male o ...
s,
ginkgo ''Ginkgo'' is a genus of non-flowering seed plants. The scientific name is also used as the English name. The order to which it belongs, Ginkgoales, first appeared in the Permian, 270 million years ago, and is now the only living genus with ...
es, and several families of conifers. Vegetation varied from
river A river is a natural flowing watercourse, usually freshwater, flowing towards an ocean, sea, lake or another river. In some cases, a river flows into the ground and becomes dry at the end of its course without reaching another body of w ...
-lining forests of
tree fern The tree ferns are arborescent (tree-like) ferns that grow with a trunk elevating the fronds above ground level, making them trees. Many extant tree ferns are members of the order Cyatheales, to which belong the families Cyatheaceae (scaly tree ...
s, and
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s (
gallery forest A gallery forest is one formed as a corridor along rivers or wetlands, projecting into landscapes that are otherwise only sparsely treed such as savannas, grasslands, or deserts. The gallery forest maintains a more temperate microclimate above th ...
s), to fern
savanna A savanna or savannah is a mixed woodland- grassland (i.e. grassy woodland) ecosystem characterised by the trees being sufficiently widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to ...
s with occasional trees such as the '' Araucaria''-like conifer ''
Brachyphyllum ''Brachyphyllum'' (meaning "short leaf") is a form genus of fossil coniferous plant foliage. Plants of the genus have been variously assigned to several different conifer groups including Araucariaceae and Cheirolepidiaceae. They are known from ...
''.


References


External links


''Marshosaurus'' at Dinosaur Mailing List
{{Taxonbar, from=Q136872 Megalosaurs Late Jurassic dinosaurs of North America Dinosaurs of the Morrison Formation Fossil taxa described in 1976 Paleontology in Utah