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Marshosaurus
Marshosaurus
Marshosaurus
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Marshosaurus
Temporal range: Late Jurassic, 155–152 Ma
↓
Skull cast
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family: †Piatnitzkysauridae
Genus: †Marshosaurus
Madsen, 1976
Species:
†M. bicentesimus
Binomial name
†Marshosaurus bicentesimus
Madsen, 1976

Marshosaurus is a genus of carnivorous theropod dinosaur, belonging to the family Piatnitzkysauridae, from the Late Jurassic Morrison Formation of Utah and possibly Colorado.[1]

Discovery and naming

[edit]
Reconstructed skull of Marshosaurus at the Carnegie Museum of Natural History, based on referred material

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 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.

In 1976 the second was by Madsen named as the type species Marshosaurus bicentesimus. The generic name honoured the nineteenth century paleontologist Professor Othniel Charles Marsh, who described many dinosaur fossils during the Bone Wars. The specific name was chosen "in honor of the bicentennial of the United States of America".[1]

The holotype, UMNH VP 6373, was found in a layer of the Brushy Basin Member of the Morrison Formation dating from the late Kimmeridgian, approximately 155 - 152 mya. It is a left ilium, or upper pelvis bone. The paratypes consisted of three bones: the ischia UMNH VP 6379 and UMNH VP 380 and the pubic bone 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.[2] 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.[3] This specimen was also the subject of a 1997 SVP abstract.[4]

Description

[edit]
Size of Marshosaurus compared to a human

Marshosaurus was medium-sized for a theropod. In 2010, Gregory S. Paul estimated its length at 4.5 meters (15 ft) and its weight at 200 kilograms (440 lb).[5] The holotype ilium has a length of 37.5 centimeters (14.8 in). If the cranial material is correctly referred, the skull was about 60 centimeters (24 in) long.

In 2012, Matthew Carrano established one autapomorphy, a unique derived trait of the holotype: the suture between the pubic peduncle and the pubic bone is convex, curving upwards, at the front and concave at the rear.[6]

Classification

[edit]
Ischium

Madsen originally was unsure about the phylogenetic position of Marshosaurus, placing it as Theropoda incertae sedis. Some later analyses showed Marshosaurus to be a member of Avetheropoda, a group of more bird-like theropods including Tyrannosaurus, Velociraptor and Allosaurus. However, Roger Benson (2010)[7] found it to be a megalosauroid in a phylogenetic analysis of Megalosaurus and 40 other theropods.

The position of Marshosaurus in the evolutionary tree, as a possible member of the Piatnitzkysauridae, is shown by the cladogram below.[7]

Megalosauroidea

Paleobiology

[edit]
Speculative life restoration with hypothetical feathers.

Pathology

[edit]

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.[8] In a 2001 study conducted by Bruce Rothschild and other paleontologists, five foot bones referred to Marshosaurus were examined for signs of stress fracture, but none were found.[9]

Paleoecology

[edit]

Habitat

[edit]

The Morrison 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,[10] to 146.8 million years old at the top,[11] which places it in the late Oxfordian, Kimmeridgian, and early Tithonian stages of the Late Jurassic period. This 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.[12] This formation is similar in age to the Solnhofen Limestone Formation in Germany and the Tendaguru Formation in Tanzania. 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.

Paleofauna

[edit]

The Morrison Formation records an environment and time dominated by gigantic sauropod dinosaurs such as Camarasaurus, Brachiosaurus, Barosaurus, Diplodocus, and Apatosaurus. Dinosaurs that lived alongside Marshosaurus included the herbivorous ornithischians Camptosaurus, Dryosaurus, Stegosaurus and Othnielosaurus. Predators in this paleoenvironment included the theropods Saurophaganax, Torvosaurus, Ceratosaurus, Stokesosaurus, Ornitholestes and[13] Allosaurus, which accounted for 70 to 75% of theropod specimens and was at the top trophic level of the Morrison food web.[14] Stegosaurus is commonly found at the same sites as Allosaurus, Apatosaurus, Camarasaurus, and Diplodocus.[15] 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.[16]

References

[edit]
[edit]
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Marshosaurus

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from Grokipedia
Marshosaurus is a genus of medium-sized carnivorous theropod dinosaur from the Late Jurassic Morrison Formation of Utah and possibly Colorado, known primarily from fragmentary skeletal remains including parts of the pelvis, hindlimbs, and teeth. Named Marshosaurus bicentesimus in 1976 after paleontologist Othniel Charles Marsh and the U.S. bicentennial, the type specimen consists of a left ilium (UUVP 2826) collected from the Cleveland-Lloyd Dinosaur Quarry in the Brushy Basin Member, dating to approximately 150 million years ago. Estimated to have reached lengths of up to 5 meters and weights around 200 kilograms, it featured a robust build with serrated, laterally compressed teeth suited for tearing flesh, a heavy ilium with a long posterior blade, and a bowed pubis ending in a teardrop-shaped foot. Classified within the superfamily and the family , Marshosaurus represents an early diverging lineage of moderate-sized theropods, distinct from larger contemporaries like Allosaurus and Ceratosaurus due to differences in ilium structure, tooth serrations, and pelvic morphology. Recent phylogenetic analyses confirm its placement in alongside genera such as and Condorraptor, highlighting its role as a basal megalosauroid adapted for predation in a diverse of floodplains and river systems. Marshosaurus, a mid-sized theropod, contributed to the rich theropod diversity of the , though its incomplete fossil record leaves many aspects of its biology enigmatic.

Discovery

Initial Finds

The earliest known specimens attributed to Marshosaurus were collected in 1912 from the Carnegie Quarry at , spanning northeastern and northwestern , consisting of a partial that was initially misidentified as belonging to another theropod and later reidentified in the . Subsequent major discoveries occurred during excavations in the at the Cleveland-Lloyd Dinosaur Quarry in Emery County, east-central , within the Brushy Basin Member of the . These efforts yielded the specimen—a left ilium cataloged as UMNH VP 6373 (formerly UUVP 2826)—along with elements such as partial fragments and additional pelvic bones from multiple individuals. In the 1970s, paleontologists working at the Dry Mesa Quarry on the Uncompahgre Plateau in Mesa County, western , recovered fragmentary postcranial remains referable to Marshosaurus, also from the Morrison Formation's Brushy Basin Member. These materials contributed to early understandings of the taxon's distribution beyond . Additional fragmentary specimens, including the first associated cranial elements, were unearthed during field seasons in the 1970s and 1990s at , spanning northeastern and northwestern , further extending the known range of Marshosaurus within the . All these initial finds date to the late through early stages of the , approximately 155–152 million years ago, based on of the Brushy Basin Member.

Formal Description and Naming

Marshosaurus bicentesimus was formally named and described by paleontologist James H. Madsen Jr. in 1976. The generic name honors the 19th-century American paleontologist , while the specific epithet bicentesimus commemorates the bicentennial of the . The holotype specimen is a left ilium (UMNH VP 6373; originally designated UUVP 2826), collected from the Cleveland-Lloyd Dinosaur Quarry in Emery County, Utah, within the Brushy Basin Member of the Morrison Formation. Paratypes consist of a right ischium (UMNH VP 6379), a left ischium (UMNH VP 6380), and a right pubic bone (UMNH VP 6387), all recovered from the same locality. These elements formed the basis of Madsen's initial taxonomic description, in which he provisionally classified Marshosaurus as a theropod dinosaur of incertae sedis within the suborder Theropoda and order Saurischia. Subsequent referrals expanded the known material, though attributions remain provisional due to the fragmentary nature of the remains. In 1991, paleontologist Brooks B. Britt referred a series of anterior caudal vertebrae (BYUVP 5201) from the Dry Mesa Quarry in western to Marshosaurus, based on their morphological similarity to unidentified tail fragments from the Cleveland-Lloyd Quarry. In 1993, a partial including cranial elements, vertebrae, and limb bones (specimen CMNH 21704, housed at the ) from in was tentatively assigned to the genus by Daniel J. Chure, Brooks B. Britt, and James H. Madsen Jr., owing to comparable neural spine morphology with elements from the type locality.

Description

Known Remains

The known remains of Marshosaurus bicentesimus consist of fragmentary skeletal elements, collectively representing a small portion of the , primarily the pelvic girdle, hindlimbs, and some axial elements, with only fragmentary forelimbs and partial cranial material preserved, but no complete . The (UMNH VP 6373, formerly UUVP 2826) is a left ilium from the Cleveland-Lloyd Dinosaur in the Brushy Basin Member of the , ; this element measures 375 mm in length but is incomplete, lacking much of the anterior blade. Paratypes are restricted to additional pelvic girdle bones from the same site, including two (UMNH VP 6379 and UMNH VP 380) and a pubis (UMNH VP 6387), which preserve articulation details such as the convex-concave suture on the pubic peduncle of the ilium. Referred specimens include three middle caudal vertebrae (DMNH 460) from the Fruita Paleontological Area, , attributed in 1991 based on shared morphology with unidentified caudal fragments from Cleveland-Lloyd. A partial skeleton (CMNH 21704, formerly DINO 16455b) from , , was referred in 1993 and includes posterior skull elements, posterior , atlas, axis, , several dorsal vertebrae, partial , a , , and fragmentary limb bones, matched via neural spine characteristics to indeterminate material from the type locality. Provisionally referred cranial fragments, such as isolated premaxillae, maxillae, and dentaries from Cleveland-Lloyd, have been suggested but lack overlapping elements with the and remain uncertain in attribution. These remains are characteristically disarticulated and weathered, derived from mixed bonebeds at sites like Cleveland-Lloyd that contain fossils from multiple individuals, which poses challenges for distinguishing intra- or inter-specific variation. Preservation issues, including surface and matrix adhesion, further limit and analysis of finer details.

Morphological Characteristics

Marshosaurus possessed a slender, body plan indicative of an agile predator, with limb proportions comparable to those of megalosaurids and a generalized tetanuran morphology that emphasized efficient locomotion over specialized extremes. The pelvic girdle featured an elongated ilium approximately 375 mm long, with a straight dorsal margin and low posterior blade that provided robust support for hindlimb ; the pubis was slender and anteriorly bowed, terminating in a boot-like distal expansion for enhanced stability. A diagnostic autapomorphy of the is the uniquely convex-concave articular surface between the pubis and ilium, formed by a small anterior bulge on the pubis creating a peg-and-socket reversed from the ceratosaur condition. Referred cranial material, if correctly attributed, suggests a skull up to 60 cm long with a low, elongate profile (length-to-height ratio exceeding 3:1) and minimal ornamentation, including sharp, slender teeth typical of theropods, though such dental elements are not directly preserved in the holotype. Scaling from the preserved ilium and overall body length of about 4.5–6 m implies hindlimb elements such as the femur reached approximately 1 m, underscoring its adaptation as a medium-sized, fast-moving carnivore.

Classification

Historical Interpretations

Following its formal description, Marshosaurus was initially classified as by James H. Madsen Jr., who noted the limited and fragmentary nature of the available material, consisting primarily of pelvic and cranial elements that precluded a more precise placement within theropod clades. Madsen loosely compared the ilium and other bones to those of fragilis, highlighting superficial resemblances in size and general proportions but emphasizing diagnostic differences, such as the shallower preacetabular wing of the ilium in Marshosaurus. This conservative assignment reflected the incompleteness of the hypodigm, which included only disarticulated remains from the , raising early concerns about the taxon's validity as a distinct genus. In the late 1970s and 1980s, subsequent studies built on Madsen's work by referring additional isolated elements from the to Marshosaurus, including comparisons of pubic morphology that distinguished it from contemporaneous theropods like tanneri. Galton and Jensen (1979) specifically discussed the pubis attributed to Marshosaurus (UUVP 2869), noting its straighter shaft and reduced distal expansion compared to , while reinforcing the risks of status due to the overall scarcity of diagnostic material. By the 1990s and into the early 2000s, some researchers tentatively assigned Marshosaurus to higher-level groups like or specifically , based on shared pelvic features such as the elongate preacetabular process of the ilium and robust ischial morphology. Holtz et al. (2004) exemplified this approach, placing it as a basal tetanuran due to these similarities with allosaurids, though acknowledging the ongoing debates over fragmentary evidence that limited robust phylogenetic resolution. By the early 2000s, interpretations shifted toward viewing Marshosaurus as a megalosaurid-like theropod, with its morphology suggesting affinities to basal members of that group rather than allosaurids. Benson (2008) proposed a position as a basal tetanuran in a new phylogenetic analysis focused on early theropod relationships, incorporating the limited material into a that highlighted its primitive features amid persistent concerns about the taxon's completeness and potential synonymy with better-known Morrison theropods. These early classifications, drawn from key works like Madsen's description and the comparative referrals in Galton and Jensen (1979), consistently underscored the challenges posed by the incomplete fossil record, often debating whether Marshosaurus warranted generic distinction or risked being relegated to a . Later phylogenetic matrices have occasionally revisited these tentative placements but generally affirm the basal tetanuran hypothesis pending more complete specimens.

Phylogenetic Analyses

In 2010, Roger B. J. Benson conducted a cladistic of basal tetanurans, incorporating Marshosaurus into a matrix of 41 taxa scored for 196 characters, primarily focusing on postcranial morphology. This recovered Marshosaurus as a basal member of , positioned outside Allosauria, supported by ilium features such as a reduced brevis fossa and a straight postacetabular process. The phylogenetic position showed moderate support, with Marshosaurus as the sister taxon to in the resulting trees. Building on this, Matthew T. Carrano and colleagues in 2012 expanded the dataset to 111 taxa and 406 characters in their comprehensive phylogeny of , confirming Marshosaurus within and formally establishing the family to include it alongside and Condorraptor. Shared synapomorphies for the family included a reduced olecranon process on the (based on referred material) and a laterally compressed manual ungual with a prominent flexor , though the fragmentary nature of Marshosaurus limited some scorings. This placement reinforced its as an early-diverging megalosauroid, distinct from more derived carcharodontosaurids. Recent phylogenetic studies from 2019 onward have introduced debates regarding the precise position of Marshosaurus and Piatnitzkysauridae, with some analyses shifting them to basal Allosauroidea. For instance, Oliver W. M. Rauhut and Diego Pol's 2019 matrix of 59 taxa and 289 characters, focused on Jurassic tetanurans, recovered Piatnitzkysauridae as the sister group to a clade comprising Carcharodontosauridae and Neovenatoridae, emphasizing transitional pelvic and femoral traits. A 2023 macroevolutionary analysis of theropod locomotor systems, using a modified matrix from prior studies, upheld the traditional Megalosauroidea assignment but noted provisional status due to Marshosaurus's incomplete skeleton, with consensus favoring Piatnitzkysauridae as an early Jurassic radiation of medium-sized predators. More recent work, such as Hendrickx et al. (2025), confirms Piatnitzkysauridae as the earliest branching clade within Allosauroidea in a revised phylogenetic analysis of tetanurans. These debates highlight ongoing uncertainties, as larger matrices continue to refine theropod interrelationships.

Paleoecology

Depositional Environment

The , dating to the Upper epochs of the and stages (approximately 156.3 to 146.8 million years ago), represents a vast that extended across present-day states including , , and in the . Fossils of Marshosaurus are primarily known from the Brushy Basin Member, the uppermost unit of this formation, which consists of variegated mudstones, sandstones, and occasional limestones indicative of fluvial and lacustrine deposition. This member formed in a semiarid environment characterized by meandering rivers, seasonal lakes, and episodic flooding on an extensive , with such as cross-bedded sandstones and overbank mudstones reflecting low-energy depositional settings interspersed with channel migrations. in these habitats included conifer-dominated woodlands and forests along watercourses, interspersed with open fern prairies, as evidenced by wood, leaves, and root traces preserved in paleosols. The presence of calcretes and minerals further supports a with periodic and under fluctuating water tables. Paleoclimate reconstructions indicate a warm, subtropical with strong seasonal wet-dry cycles, possibly influenced by monsoon-like patterns, as inferred from sedimentological features and stable analyses of pedogenic carbonates. Oxygen isotope data from authigenic minerals suggest mean annual temperatures ranging from 25°C to 35°C, fostering diverse fluvial ecosystems despite the overall . Taphonomic evidence from Marshosaurus-bearing sites in the Brushy Basin Member points to attritional accumulation of remains in low-energy marginal aquatic environments, such as margins or backwaters, where bones were concentrated through hydraulic sorting and minimal transport. Bonebeds in this context, including those with theropod elements, likely formed via drought-induced mortality traps followed by redistribution, preserving disarticulated skeletons in fine-grained sediments without significant or scavenging.

Contemporaneous Biota

The Morrison Formation of the Late Jurassic preserved a rich and diverse vertebrate fauna, dominated by dinosaurs but including a variety of other taxa. Theropod dinosaurs were prominent carnivores, with Allosaurus serving as the apex predator at lengths of 8–12 meters and accounting for over 70% of theropod specimens, Ceratosaurus as a smaller, horned form reaching 5–6 meters, and Marshosaurus as a mid-sized representative at about 4.5 meters long. Herbivorous dinosaurs formed the bulk of the large-bodied fauna, including abundant sauropods such as Diplodocus, Apatosaurus, and Camarasaurus, which together comprised roughly 50% of all vertebrate fossils recovered, alongside ornithischians like the plated Stegosaurus and the bipedal Dryosaurus. The assemblage also featured smaller vertebrates, such as early mammals of the clade Dryolestida (e.g., Dryolestes priscus), semiaquatic crocodilians including Goniopholis, flying reptiles like the pterosaur Harpactognathus, as well as fish, frogs, salamanders, lizards, turtles, and invertebrates; notably, no avian birds are known from the formation. High theropod diversity in the , with at least eight sympatric genera, indicates niche partitioning based on and morphology, allowing coexistence through differences in prey size, diet, and habitat use. Recent analyses of bite marks on sauropod bones further suggest community dynamics involved scavenging and predation on juvenile herbivores, supporting partitioned roles among predators.

Paleobiology

Pathological Evidence

A referred right ilium (UUVP 2742) of Marshosaurus bicentesimus displays a traumatic deformity along the posterodorsal margin grading into a lateral ridge, likely evidencing injury from physical confrontation or environmental factors. Analysis of five pedal bones referred to Marshosaurus revealed no signs of stress fractures, differing from patterns observed in larger Morrison theropods such as Allosaurus, where such injuries are common and suggest intense, repetitive hunting exertion; this absence implies comparatively lower locomotor strain in Marshosaurus. The documented pathologies in Marshosaurus specimens underscore a lifestyle fraught with injury risks from predation and competition.

Inferred Ecology and Behavior

Marshosaurus, as a medium-sized theropod estimated at 4.5 meters in length and approximately 200 kg in mass, is inferred to have been a carnivorous predator or targeting smaller vertebrates, including ornithopods, mammals, and possibly juvenile sauropods, within the . Recent dental microwear textural analysis (3D DMTA) of teeth from the Cleveland-Lloyd Dinosaur Quarry reveals statistically significant dietary differences between Marshosaurus and the larger fragilis, supporting niche partitioning where Marshosaurus likely focused on softer or more accessible prey compared to the apex predatory role of . This partitioning may have extended to nasicornis, a smaller theropod potentially specialized for different prey types, allowing coexistence among carnivores despite a high predator-to-prey ratio. Reconstruction of the pelvic girdle and musculature indicates that Marshosaurus was a biped capable of agile maneuvers, with robust and muscles such as the M. caudofemoralis brevis providing strong propulsive force for pursuits or evasion. The wider brevis fossa on the suggests enhanced force generation relative to closely related piatnitzkysaurids like , implying moderate agility suited to navigating environments rather than sustained high-speed chases. As a member of , its lightly built yet sturdy aligns with active predatory behaviors typical of basal tetanurans. Fragmentary remains, including elements from at least three individuals, limit detailed insights into growth and . Multiple specimens from sites like the Cleveland-Lloyd may reflect taphonomic aggregation due to the quarry's , though direct evidence for remains inconclusive due to the scarcity of associated assemblages.

References

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  6. https://npshistory.com/publications/paleontology/nmmnhs-79-703.pdf
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  11. https://iknowdino.com/marshosaurus-episode-76/
  12. https://www.researchgate.net/publication/272152788_A_new_large_theropod_dinosaur_from_the_Upper_Jurassic_of_Colorado
  13. https://dml.reptilis.net/2008Nov/msg00165.html
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  15. https://pubs.usgs.gov/bul/1009e/report.pdf
  16. https://npshistory.com/publications/paleontology/nrtr-93-11.pdf
  17. https://www.nps.gov/dino/learn/nature/morrison-formation.htm
  18. https://ugspub.nr.utah.gov/publications/special_studies/SS-48.pdf
  19. https://www.researchgate.net/publication/264762599_Palaeobiodiversity_of_conifer_seed_cones_in_the_Upper_Jurassic_Morrison_Formation_of_Utah_USA
  20. https://www.sciencedirect.com/science/article/abs/pii/S0037073804000119
  21. https://pubs.geoscienceworld.org/gsa/gsabulletin/article/126/7-8/1105/126022/Multiproxy-approach-reveals-evidence-of-highly
  22. https://doc.rero.ch/record/14574/files/PAL_E1788.pdf
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  33. https://onlinelibrary.wiley.com/doi/full/10.1111/joa.13983
  34. https://www.prehistoric-wildlife.com/species/marshosaurus/
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