Paleobiota of the Morrison Formation

Short description: none

The distinctive banding of the Morrison Formation, a group of rock layers that occur throughout Dinosaur National Monument and the source of fossils like those found at the Dinosaur Quarry.

The Morrison Formation is a distinctive sequence of Late Jurassic sedimentary rock that is found in the western United States, which has a wide assortment of taxa represented in its fossil record, including 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.

## Contents

* 1 Plants
* 1.1 Conifers
* 1.2 Ginkgos
* 1.3 Cycadophyta and relatives
* 1.4 Gnetales
* 2 Invertebrates
* 2.1 Arthropods
* 3 Vertebrates
* 3.1 Fish
* 3.2 Amphibians
* 3.3 Sphenodonts
* 3.4 Squamates
* 3.5 Turtles
* 3.6 Choristoderes
* 3.7 Crurotarsans
* 3.8 Pterosaurs
* 3.9 Dinosaurs
* 3.10 Mammaliaforms
* 3.10.1 Tinodontids
* 3.10.2 Eutriconodonts
* 3.10.3 Multituberculates
* 3.10.4 Others
* 3.10.5 Dryolestoids
* 4 See also
* 5 Footnotes
* 5.1 Bibliography

## Plants

The Morrison Formation is devoid of flowering plants; during the Jurassic, the Earth was dominated by ferns, seed ferns, and a wealth of gymnosperms—such as cycads, ginkgos, and conifers. Vegetation varied from river-lining forests of conifers, tree ferns, and ferns (gallery forests), to fern savannas with occasional trees such as the Araucaria-like conifer Brachyphyllum.[1][2]

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.

### Conifers

Genus  | Species  | Locality  | Material  | Notes

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Mesembrioxylon  |

M. carterii

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Protocupressinoxylon  |

P. medlynii

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Brachyphyllum | B. rechtenii

B. sp

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Sequoia | S.sp  |  |  |
cf. Araucaria[3] | A. sp  |  |  |

### Ginkgos

Genus  | Species  | Locality  | Material  | Notes

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Ginkgo |

G. sp

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### Cycadophyta and relatives

Genus  | Species  | Locality  | Material  | Notes

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Nilssonia |

N. compta

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Zamites |

Z. arcticus

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Cycadolepis  |

C. sp

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### Gnetales

Genus  | Species  | Locality  | Material  | Notes

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Bassitheca[4]

B. hoodiorum

Colorado and Utah (Brushy Basin Member)

More than 300 specimens, three dimensional calcitic casts of a two-seeded compound cone

Tentatively assigned as a gnetale, with some attributes of the cones pointing towards a close relation with the extant genus Ephedra.

Dayvaultia[5]

D. tetragona

Henry Mountains of Utah (Brushy Basin Member)

"Three-dimensional casts and partially permineralized small cones".

A gnetale with close affinities to Cretaceous and modern genera.

## Invertebrates

### Arthropods

Trace fossils reported from the Morrison Formation are indicative of greater diversity of arthropods (especially insects) than recorded by body fossils.[6] Bone modifications likely produced by dermestid beetles were reported in vertebrate fossils from Wyoming and Colorado.[7][8]

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes

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Cambaridae[9] | Indeterminate  |  |  | A crayfish
Coleoptera[10] | Indeterminate  | Utah  | Isolated elytra.  |
Decapoda[10] |  | Utah  | A specimen preserved with structures similar to eyes on stalks, antennae and a cheliped.  | Possibly a freshwater hermit crab.

Eopolis

E. ekdalei

Utah, Brushy Basin member

Nests produced by social insects,[11] possibly termites.[12]

Morrisonnepa[13] | M. jurassica |

Utah

|  | A hemipteran belonging to the group Nepomorpha.
Parapleurites[14] | P. morrisonensis |

Colorado

| a forewing.  | A Locustopsid orthopteran
Spinicaudata[10] |  | Utah  | At least 10 specimens preserved as shells.  |

Tektonargus

T. kollaspilus

Colorado, Brushy Basin member

Five specimens were reported in the original description of the ichnogenus.

## Vertebrates

### Fish

Although the paleoclimate of the Morrison formation was semiarid with only seasonal rainfall, there were enough bodies of water to support a diverse ichthyofauna.[15] Although abundant, fish remains are constrained to only certain locations within the formation.[15] Microvertebrate sites in Wyoming are dominated by fish remains.[15] Indeterminate ray-finned fish remains have been recovered from Ninemile Hill and a microvertebrate site in the Black Hills.[15] Found in stratigraphic zones 2, 4, and 5.[16] Morrison actinopterygians generally have no close modern relatives.[15] The Wyoming microvertebrate remains are extracted from the sediment by screenwashing.[15] Paleoniscoid remains are geographically present in the western part of Colorado, where remains have been recovered from "a level above the Mygatt-Moore Quarry."[15] Largely complete remains of small individuals have been consistently recovered for over 15 years.[15] Single tooth fossil of pycnodont fish is represented from Dinosaur National Monument in Utah,[17] found in stratigraphic zone 4.[16] Dipnoan remains found at a fossil site not far from Cañon City, Colorado.[15] Remains usually in a state of rather complete preservation.[15] Halecostome remains are geographically present in the western part of Colorado, where remains have been recovered from "a level above the Mygatt-Moore Quarry."[15] Largely complete remains of small individuals have been consistently recovered for over 15 years.[15] Amiid remains found in stratigraphic zones 2, 3, and 4.[16] Found at a fossil site not far from Cañon City, Colorado.[15] Remains usually in a state of rather complete preservation.[15]

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes  | Images

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Ceratodus[18]

C. fossanovum[15]

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A lungfish genus whose members ranged from 1 to 2 m in length and weights of up to 79 pounds, with most Morrison lungfish being on the smaller end of that range.[15] These species are believed to have had similar diets to extant lungfish like the physically similar modern genus Neoceratodus.[15]

|  Ceratodus
"Hulettia" hawesi
Morrolepis schaefferi
A teleost, cf. Leptolepis

C. ?frazieri[15]

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C. guentheri[15]

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C. robustus[15]

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Indeterminate.

Brushy Basin and Saltwash members

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Represented by tooth plates.

"Hulettia"[18]

"H. hawesi"[19]

Colorado

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A small fish, originally considered as a member of Halecostomi about 7.6 cm in length and 5g of live mass which probably preferred quiet water. Its fossils prominently preserve its thick interlocking scales. Later study question its assignment to genus Hulettia, and suggested that it would be a semionotid instead.[20]

cf. Leptolepis[18]

N/A

Colorado[19]

Known only from a single nearly complete skeleton found at Rabbit Valley.[21] Found in stratigraphic zone 5.[16]

A 13 cm (5 inch) fish that was deeper bodied than its co-occurring contemporaries Morrolepis and "Hulettia".[21] The Morrison cf. Leptolepis probably had a live mass of about 37g.[21] It is the only teleost fish known from the formation and was morphologically more highly derived than other Morrison fish.[21] It is believed to have fed on contemporary fish and small invertebrates.[21]

Morrolepis[18]

M. schaefferi[22]

Colorado

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A coccolepidid "palaeoniscoid" with forward-set eyes positioned past the front end of the lower jaw. It had a tall dorsal fin set far back on the body and an asymmetrical caudal fin.[22] Adult specimens would reach about 20 cm in length and 113 g (4 oz) in mass.[22]

Potamoceratodus

P. guentheri

Colorado

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Once thought to be a species of Ceratodus.

Purgotybaris

P. fosteri

Utah

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A regurgitalite containing frog and salamander bones, probably produced by an amioid fish.[23]

A hybodont?

Indeterminate

Utah

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Known from a dorsal spine.[24]

### Amphibians

A modern frog from the same family as Rhadinosteus parvus, the rhinophrynidae.

Frogs are known from several sites in the Morrison Formation, but are not particularly well represented.[25] The history of Morrison anuran discoveries began with the recovery of remains from Quarry 9 near Como Bluff, Wyoming. The new genus Eobatrachus was erected for some of these remains by O. C. Marsh, but the material was later considered non-diagnostic. Decades later another dubious anuran genus, Comobatrachus, was erected based on additional fragmentary remains. Despite the erection of multiple new names, only two frog species are currently recognised from the Morrison: Enneabatrachus hechti[26] and Rhadinosteus parvus.[27]

In addition to formally named taxa, indeterminate anuran remains have been recovered from Morrison strata in Colorado, Wyoming, and Utah, with the best specimens found in Dinosaur National Monument and Quarry 9.[25] Stratigraphically speaking, indeterminate anurans have been found in stratigraphic zones 2 and 4.[16] Indeterminate anurans with remains diagnostic down to the family level have also been reported from the Morrison, with pelobatids being represented by the ilium of an unnamed and indeterminate species, which was recovered from Quarry 9.[28] Pelobatids are present in stratigraphic zones 5 and 6.[16]

Indeterminate salamander remains are present in stratigraphic zones 2, 4, and 5.[16] A distinctive type of salamander known only as Caudata B is present in stratigraphic zone 6.[16]

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes

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Comobatrachus[25]

C. aenigmaticus

Wyoming, Brushy Basin member[25]

| A partial right humerus.[29] |

A dubious genus of prehistoric frog erected by O. C. Marsh to house fragmentary remains recovered from Reed's Quarry 9 near Como Bluff Wyoming.[25] Along with Eobatrachus it was among the earliest frog remains from the formation, although the two dubious genera were erected decades apart.[25]

Comonecturoides[30]

C. marshi[30]

Wyoming, Brushy Basin member[30]

Represented by a single femur.[30]

Considered a nomen dubium because the name is based on non-distinctive remains which cannot be classified in detail.[30]

Enneabatrachus[18]

E. hechti[26]

Utah and Wyoming[31][26]

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A small discoglossid frog whose live weight would have only been a few grams.[26]

Eobatrachus[25]

E. agilis

Wyoming, Brushy Basin member[25]

| A partial right humerus.[29] |

A dubious genus of prehistoric frog erected by O. C. Marsh to house fragmentary remains recovered from Reed's Quarry 9 near Como Bluff Wyoming.[25] Along with Comobatrachus it was among the earliest frog remains from the formation, although the two dubious genera were erected decades apart.[25]

Iridotriton[18]

I. hechti

Utah, Brushy Basin[32]

| A partial skull and incomplete postcranial skeleton.[32] |

A basal salamandroid closely related to today's advanced salamanders.

Rhadinosteus[18]

R. parvus[27]

Utah

Known from several slabs of rock which contain multiple partial specimens in association.[27]

A pipoid and possible rhinophrynid, Rhadinosteus parvus was only about 42 mm (1.6 inches) long in life.[27]

### Sphenodonts

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes  | Images

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Eilenodon[18]

E. robustus

Colorado

Several fragmentary skull, mandible, and tooth elements.

A herbivorous eilenodontine of relatively large size.

Opisthias[18]

O. rarus

Colorado, Utah and Wyoming, Brushy Basin member

Several skulls and partial skeletons.

An opisthodontian sphenodontian

Opisthiamimus[33]

O. gregori

Northern Wyoming, possibly west-central Colorado. Brushy Basin Member

Most of the skull and postcranium

A small-bodied eusphenodontian, and one of the most complete rhynchocephalian taxa yet known from North America.

Theretairus[18]

T. antiquus

Wyoming, Brushy Basin member

A mandible.

A small sphenodontian.

Sphenodontia[33] | Indeterminate  | Dinosaur National Monument  | A crushed partial skull (DINO 16454)  | The specimen is undescribed. Although previously considered to have belonged to Opisthias or Theretairus, recent studies have doubted this referral, thus placing it as an indeterminate sphenodont.[33] |

### Squamates

Numerous squamate remains have been found in the sediments of the Morrison Formation, most commonly at sites in Utah, Colorado and Wyoming. A number of taxa have been described, coming from three different groups: anguimorph and scincomorph lizards and early snakes.[34][35][36] The first squamates to be reported from the Morrison Formation were Paramacellodus and Dorsetisaurus, which were described from Wyoming's Quarry 9 by Don Prothero and Richard Estes.[34] Later remains would include Diablophis, originally described as a species of Parviraptor by Susan Evans in 1996[37] but subsequently moved to the new genus Diablophis by Michael Caldwell et al. in 2015, with extra material also being reported from Utah's Cisco Mammal Quarry, and Schillerosaurus, originally described as "Schilleria" and reported from Dinosaur National Monument by Evans and Dan Chure in 1999.[38] Two later additions to the Morrison's squamate assemblages are Eoscincus and Microteras, two scincomorph lizards found at Dinosaur National Monument and Como Bluff's Quarry 9, respectively. They were described by Chase Brownstein et al. in 2022.[35] Indeterminate squamate remains have currently been described from Dinosaur National Monument.[38]

The majority of modern-day scincomorph lizards are small insectivores that feed on a range of invertebrates.[39] It is thought that their counterparts from the Morrison Formation would have occupied a similar niche due to their morphological similarities. Anguimorph lizards most likely hunted small vertebrates, and Diablophis is thought to have done so too. Prey items would have included the other squamates from the formation as well as its large diversity of small mammals. All squamates might have been prey for the larger predators of the Morrison Formation, including the abundant theropod dinosaurs and crocodilians.

Name  | Species  | Locality  | Material  | Notes

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Diablophis[36]

D. gilmorei

Fruita Palaeontological Area, Colorado and Cisco Mammal Quarry, Utah (Brushy Basin Member)

Broken skull material including a right maxilla, mandible and dentary. Broken axis vertebrae, precloacal vertebrae, one caudal vertebra and a possible sacral vertebra have also been found.[36]

A basal snake. Originally described as a species of Parviraptor, it was subsequently moved to its own genus.[18][36]

Dorsetisaurus[18]

D. sp.

Quarry 9 at Como Bluff, Wyoming, Utah and Colorado (Brushy Basin Member)

Multiple dentaries.[34][38]

An anguimorph lizard.

Eoscincus[35]

E. ornatus

Site 412 at Dinosaur National Monument, Utah (Brushy Basin Member)

A partial skull including a complete rostrum and palate, a partial skull roof, and both mandibles.

A scincomorph lizard.

Helioscopos[40] | H. dickersonae | Site 317 at Dinosaur National Monument, Utah (Brushy Basin Member)  | A nearly complete skull consisting of a partial cranium and mandibles, lacking the anterior snout.  | A stem-gecko.

Microteras[35]

M. borealis

Quarry 9 at Como Bluff, Wyoming (Brushy Basin Member)

A partial skull consisting of an associated maxilla and braincase.

A scincomorph lizard.

Paramacellodus[18]

P. sp.

Fruita Palaeontological Area & Rainbow Park, Colorado and Quarry 9 at Como Bluff, Wyoming (Brushy Basin Member)

Numerous specimens including jaws, skulls, osteoscutes, trunk vertebrae and hindlimbs. Possibly a left prefrontal and postcranial material consisting of a scapulocoracoid, trunk vertebra and multiple articulated bones consisting of a partial pelvis, seven caudal vertebrae and a nearly complete left hind limb.[34][38]

A small scincomorph lizard with blunt teeth.

Saurillodon[18]

S. sp

Fruita Palaeontological Area, Colorado (Brushy Basin Member)

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A scincomorph lizard whose remains have been found in Middle Jurassic strata in England and Scotland as well as Late Jurassic strata in Portugal in addition to the Morrison formation remains.[41]

Schillerosaurus[18][42]

S. utahensis [43][38]

Dinosaur National Monument, Utah (Brushy Basin Member)

Part and counterpart of a partial dissociated skeleton and a few limb bone fragments.[38]

A small scincomorph lizard of otherwise uncertain evolutionary affinities.[43]

Squamata[38]

Indeterminate

Dinosaur National Monument, Utah (Brushy Basin Member)

Multiple specimens including fragmentary and disarticulated skeletons.[44]

Squamates of uncertain affinities. Noted as distinct from Paramacellodus and Dorsetisaurus, of which remains have been found in the same quarry.[38]

### Turtles

Turtles (Testudines) are very common fossils in the Morrison, due to their bony shells. The most common were Glyptops plicatus (very common) and Dinochelys whitei (also common, but not as common as Glyptops). Also present were Dorsetochelys buzzops and Uluops uluops.

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes

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Chelonipus

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Colorado and Utah, Salt Wash member

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Compsemys

C. plicatulus

Colorado

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Dinochelys[18]

D. whitei[45]

Colorado, Utah and Wyoming, Brushy Basin member[45]

Several shells and some postcranial material.[45]

Dorsetochelys[18]

D. buzzops

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Glyptops[18]

G. plicatulus[45]

Colorado and Wyoming, Brushy Basin member[45]

Several shells, skulls, and partial skeletons.[45]

G. ornatus[46]

Wyoming, Brushy Basin member[46][45]

A skull.[45][46]

Synonym of Glyptops plicatulus.[45]

G. utahensis[47]

Utah, Brushy Basin member[47][45]

A complete shell.[47]

Synonym of Glyptops plicatulus.[45]

Uluops[18]

U. uluops

Wyoming, Brushy Basin member[48]

A cranium.[48]

Most basal Pleurosternid from the Morrison Formation.[48]

### Choristoderes

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes

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Cteniogenys[18]

C. antiquus

Colorado, Oklahoma, South Dakota, Utah and Wyoming

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A choristoderan about 25 to 50 cm in length.

### Crurotarsans

Crocodiles of a variety of sizes and habitats were common Morrison animals. Cursorial mesosuchians, or small terrestrial running crocs, included Hallopus victor and Fruitachampsa callisoni. More derived crocodilians included Diplosaurus ferox, Amphicotylus, Hoplosuchus kayi, and Macelognathus vagans.

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes  | Images

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Amphicotylus[49]

A. gilmorei

Wyoming

|  |  |  Hallopus Hoplosuchus

A. lucasii

Colorado

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A. milesi

Wyoming, Brushy Basin member[50]

A nearly complete skeleton.[50]

A. stovalli

Oklahoma

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Diplosaurus[51]

D. felix

Colorado

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Eutretauranosuchus[18]

E. delfsi

Colorado and Wyoming

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Fruitachampsa[18]

F. callisoni

Colorado, Brushy Basin and Saltwash members

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Hallopus[18]

H. victor

Colorado, Brushy Basin member[52]

A partial skeleton including a fragmentary skull roof.[52]

Hatcherichnus

H. sanjuanensis

Colorado and Utah

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H. sp.

Utah and Wyoming, Salt Wash member

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Hoplosuchus[18]

H. kayi

Arizona and Utah

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Macelognathus[18]

M. vagans

Colorado and Wyoming, Brushy Basin member[53]

A partial left mandible (type) and several fragmentary referred fossils.[53]

Theriosuchus[54]

T. morrisonensis

Wyoming

A nearly complete left mandible missing teeth.

### Pterosaurs

Pterosaurs are very uncommon fossils in the Morrison, because the fragility of their thin walled bones often prevented their remains from being preserved.[55] Despite being uncommon they are geographically widespread;[56] indeterminate pterosaur remains have been found in stratigraphic zones 2 and 4-6.[16] In addition to indeterminate remains, several species have been identified from both the rhamphorhynchoids (long-tailed pterosaurs) and pterodactyloids (short-tailed pterosaurs).[55] Since the 1970s and 80s, pterosaur finds have become more common, but are still rare.[55] Most Morrison pterosaurs have been found in marine and shoreline deposits.[55] Pterosaur tracks have been found in both the Tidwell and Saltwash members.[55] Morrison pterosaurs probably lived on fish, insects and scavenged dinosaur carcasses, or even foraged for prey, and actively hunted;[55] they are fairly ecologically diverse, ranging from small hawking insectivore Mesadactylus to the raptorial Harpactognathus. While relatively few pterosaur genera are named from the Morrison Formation, fragmentary material that is not referrable to the genus level suggests the presence of dsungaripteroids, ctenochasmatids, dimorphodontids, and more tentatively wukongopterids and pteranodontians.[57][58][59][60]

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Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

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| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.
Name  | Species  | Locality  | Material  | Notes  | Images

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Comodactylus[18]

C. ostromi

Wyoming, Brushy Basin member

A metacarpal.

Nomen dubium

Dermodactylus[18]

D. montanus

Wyoming, Brushy Basin member

A metacarpal.

Nomen dubium

Harpactognathus[18]

H. gentryii

Wyoming, Brushy Basin member

A partial snout.

A large rhamphorhynchid with a wingspan of about 2.5 m and live mass of about 1.5 kg (3.3 lbs).[61] Harpactognathus was related to the Solnhofen genus Scaphognathus.[61]

Kepodactylus[18]

K. insperatus

Colorado, Brushy Basin member

A partial postcranial skeleton.

A large pterodactyloid with a 2.5 m (8 foot) wingspan and a live weight of about 1.5 kg (3 lbs).[62] Kepodactylus may be related to the Asian dsungaripteroid pterosaurs.[62]

Laopteryx[56]

L. priscus

Wyoming, Brushy Basin member

A braincase.

Nomen dubium initially misidentified as a bird.

Mesadactylus[18]

M. ornithosphyos

Colorado, Brushy Basin member

| A synsacrum.  | Several specimens have been incorrectly referred to Mesadactylus.[58][59]

Pteraichnus[63]

P. saltwashensis*

Arizona and Oklahoma, Saltwash member

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Utahdactylus[18]

U. kateae

Utah, Tidwell member

A fragmentary skeleton.

Previously thought to be an indeterminite diapsid, newer material suggests an affinity with ctenochasmatids.[60]

### Dinosaurs

### Mammaliaforms

Many types of mammaliaform cynodonts, mostly early mammals, are known from the Morrison; almost all of them were small sized animals, though occupying a very large variety of ecological niches, from the more rodent-like multituberculates to the carnivorous eutriconodonts (including the possibly volant Triconolestes) to the anteater-like Fruitafossor. Unclassified types include the digger Fruitafossor windscheffelia. Docodonts included the common genus Docodon, represented by D. victor, D. striatus, and D. superbus, and Peraiocynodon sp. Multituberculates, a common type of early mammal, were represented by Ctenacodon serratus, C. laticeps, C. scindens, Glirodon grandis, Morrisonodon brentbaatar, Psalodon fortis, ?P. marshi, P. potens, and Zofiabaatar pulcher. Triconodonts present included Amphidon superstes, Aploconodon comoensis, Comodon gidleyi, Priacodon ferox, P. fruitaensis, P. gradaevus, P. lulli, P. robustus, Triconolestes curvicuspis, and Trioracodon bisulcus.

Tinodontids were represented by Eurylambia aequicrurius (probably Tinodon), and Tinodon bellus (including T. lepidus). Finally, two families of Dryolestoidea were present: Paurodontidae, including Comotherium richi, Euthlastus cordiformis, Paurodon valens, and Tathiodon agilis; and Dryolestidae, including Amblotherium gracilis, Dryolestes obtusus (common genus), D. priscus, D. vorax, Laolestes eminens, L. grandis, and Miccylotyrans minimus.

In 2009, a study by J. R. Foster was published which estimated the body masses of mammals from the Morrison Formation by using the ratio of dentary length to body mass of modern marsupials as a reference. Foster concludes that Docodon was the most massive mammaliaform genus of the formation at 141g and Fruitafossor was the least massive at 6g. The average Morrison mammal had a mass of 48.5g. A graph of the body mass distribution of Morrison mammal genera produced a right-skewed curve, meaning that there were more low-mass genera.[64]

|  |

|
|

Color key
| Taxon | Reclassified taxon | Taxon falsely reported as present | Dubious taxon or junior synonym | Ichnotaxon | Ootaxon | Morphotaxon

|
|
|
|
|
|

| Notes
Uncertain or tentative taxa are in small text; ~~crossed out~~ taxa are discredited.

#### Tinodontids

Name  | Species  | Locality  | Material  | Notes

|
|
|
|

Eurylambia

E. aequicrurius

Wyoming

|  |

A tinodontid similar in appearance to Tinodon.

Tinodon[18]

T. bellus

Wyoming

|  |

Tinodontids.

T. lepidus

Wyoming

#### Eutriconodonts

Name  | Species  | Locality  | Material  | Notes

|
|
|
|

Amphidon[18]

A. superstes

Wyoming

|  |

A small amphidontid.

Aploconodon[18]

A. comoensis

Wyoming

|  |

An amphilestid eutriconodont.

Comodon[18]

C. gidleyi

Wyoming, Brushy Basin member

| A mandible.  |

An amphilestid eutriconodont slightly larger in size than Aploconodon.

Phascalodon

P. gidleyi

Wyoming

|  |

Triconolestes[18]

T. curvicuspis

Utah

|  |

A volaticotherian eutriconodont.

Trioracodon[18]

T. bisulcus

Wyoming

|  |

A triconodontid eutriconodont similar to Priacodon.

#### Multituberculates

Name  | Species  | Locality  | Material  | Notes

|
|
|
|

Ctenacodon[18]

C. laticeps

Wyoming

|  |

C. scindens

|  |  |

C. serratus

Wyoming

|  |

Glirodon[18]

G. grandis

Colorado and Utah

|  |

Morrisonodon

M. brentbaatar

Wyoming

|  |

Priacodon[18]

P. ferox

Wyoming

|  |

P. fruitaensis

Colorado

P. grandaevus

Wyoming

P. lulli

Wyoming

P. robustus

Wyoming

Psalodon[18]

P. fortis

|  |  |

P. marshi

|  |

P. potens

|  |

Zofiabaatar[18]

Z. pulcher

Wyoming

|  |

#### Others

Name  | Species  | Locality  | Material  | Notes  | Images

|
|
|
|
|

Cifellilestes[65]

C. ciscoensis[65]

Utah, Brushy Basin Member

|  |  | Fruitafossor

Docodon[18]

D. victor[66][67]

|  |  |

Fruitafossor[18]

F. windscheffeli

| Colorado  |  |

#### Dryolestoids

Name  | Species  | Locality  | Material  | Notes

|
|
|
|

Amblotherium[18]

A. gracilis

Colorado

|  |

A small Dryolestid dryolestoid.

A. megistodon[68]

Wyoming

Araeodon[18]

A. intermissus

Utah and Wyoming

|  |

A paurodontid dryolestoid, somewhat smaller than Archaeotrigon and Paurodon. Considered to be a junior synonym of Paurodon valens by Averianov and Martin (2015).[69]

Archaeotrigon[18]

A. brevimaxillus

Wyoming

|  |

Paurodontid dryolestoids similar in appearance to Paurodon. Both species were considered to be junior synonyms of Paurodon valens by Averianov and Martin (2015).[69]

A. distagmus

|  |

Comotherium[18]

C. richi

Wyoming

|  |

A paurodontid dryolestoid.

Dryolestes[18]

D. obtusus

|  |  |

Dryolestid dryolestoids.

D. priscus

Wyoming

D. tenax

|  |

Euthlastus[18]

E. cordiformis

Wyoming

|  |

A paurodontid dryolestoid.

Foxraptor[18]

F. atrox

Wyoming

|  |

A paurodontid dryolestoid similar in size to Paurodon. Considered to be a junior synonym of Paurodon valens by Averianov and Martin (2015).[69]

Herpetairus

H. sp.

|  |  |

Kepolestes

K. coloradensis

Colorado

|  |

Laolestes[18]

L. eminens

|  |  |

Common Dryolestid dryolestoids.

L. grandis

|  |

Malthacolestes

M. sp.

|  |  |

Melanodon

M. goodrichi

M. hodsoni

M. oweni

|  |  |

Miccylotyrans

M. minimus

|  |  |

A Dryolestid dryolestoid.

Paurodon[18]

P. valens

Wyoming

|  |

A paurodontid dryolestoid.

Pelicopsis

P. dubius

Wyoming

|  |

A paurodontid dryolestoid. Considered to be a junior synonym of Paurodon valens by Averianov and Martin (2015).[69]

Tathiodon[18]

T. agilis

Wyoming

|  |

A paurodontid dryolestoid.

## See also

* List of dinosaur-bearing rock formations
* Molluscs
* Synthesis
* lacustrine carbonates

## Footnotes

1. ↑ Carpenter, Kenneth (2006). "Biggest of the big: a critical re-evaluation of the mega-sauropod Amphicoelias fragillimus". in Foster, John R.. 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.
2. ↑ Peterson, Fred. SUMMARY LIST OF LATE JURASSIC PLANTS, WESTERN INTERIOR U.S. & SW CANADA.
3. ↑ Bader, K (2003). "The local flora and fauna of a site in the upper Morrison Formation (Upper Jurassic) of northeastern Wyoming.". Journal of Vertebrate Paleontology 23 (3 suppl).
4. ↑ Steven R. Manchester; Xiaoqing Zhang; Carol L. Hotton; Scott Wing; Peter R. Crane (2022). "Two-seeded cones of probable gnetalean affinity from the Morrison Formation (Late Jurassic) of Utah and Colorado, USA". Acta Palaeobotanica 62 (2): 77–92. doi:10.35535/acpa-2022-0006.
5. ↑ Steven R. Manchester; Xiaoqing Zhang; Carol L. Hotton; Scott Wing; Peter R. Crane (2021). "Distinctive quadrangular seed-bearing structures of gnetalean affinity from the Late Jurassic Morrison Formation of Utah, USA". Journal of Systematic Palaeontology 19 (4): 1–18. doi:10.1080/14772019.2021.1968522. Bibcode: 2021JSPal..19..743M.
6. ↑ Stephen T. Hasiotis (2004). "Reconnaissance of Upper Jurassic Morrison Formation ichnofossils, Rocky Mountain Region, USA: paleoenvironmental, stratigraphic, and paleoclimatic significance of terrestrial and freshwater ichnocoenoses". Sedimentary Geology 167 (3–4): 177–268. doi:10.1016/j.sedgeo.2004.01.006.
7. ↑ Brooks B. Britt; Rodney D. Scheetz; Anne Dangerfield (2008). "A Suite of Dermestid Beetle Traces on Dinosaur Bone from the Upper Jurassic Morrison Formation, Wyoming, USA". Ichnos 15 (2): 59–71. doi:10.1080/10420940701193284.
8. ↑ Julia B. McHugh; Stephanie K. Drumheller; Anja Riedel; Miriam Kane (2020). "Decomposition of dinosaurian remains inferred by invertebrate traces on vertebrate bone reveal new insights into Late Jurassic ecology, decay, and climate in western Colorado". PeerJ 8. doi:10.7717/peerj.9510. PMID 32742794.
9. ↑ Audo, Denis; Hasiotis, Stephen T; Kawai, Tadashi (2023-12-01). "Diversity and evolutionary history of fossil crayfishes" (in en). Journal of Crustacean Biology 43 (4). doi:10.1093/jcbiol/ruad079. ISSN 0278-0372. https://academic.oup.com/jcb/article/doi/10.1093/jcbiol/ruad079/7504443.
10. ↑ 10.0 10.1 10.2 María B. Lara; John R. Foster; Thomas F. Howells; James I. Kirkland (2023). "14th Symposium on Mesozoic Terrestrial Ecosystems and Biota. Arthropoda body fossils from the Jurassic Salad Bar site in the Upper Jurassic Morrison Formation in southeastern Utah, USA". The Anatomical Record 306 (S1): 163–165. doi:10.1002/ar.25219.
11. ↑ Elliott Armour Smith; Mark A. Loewen; James I. Kirkland (2020). "New social insect nests from the Upper Jurassic Morrison Formation of Utah". Geology of the Intermountain West 7: 281–299. doi:10.31711/giw.v7.pp281-299. https://giw.utahgeology.org/giw/index.php/GIW/article/view/84.
12. ↑ Alayna Rea; Edward L. Simpson; Michael C. Wizevich (2026). "A FESEM examination of Eopolis ekdalei, an eusocial ichnofossil, Brushy Basin Member, Late Jurassic Morrison Formation: evidence for preserved fungus in a fungal garden". New Mexico Museum of Natural History and Science Bulletin 102: 347–358. https://www.researchgate.net/publication/400053442.
13. ↑ María B. Lara; John R. Foster; James I. Kirkland; Thomas F. Howells (2020). "First fossil true water bugs (Heteroptera, Nepomorpha) from Upper Jurassic strata of North America (Morrison Formation, southeastern Utah)". Historical Biology: An International Journal of Paleobiology 33 (10): 1996–2004. doi:10.1080/08912963.2020.1755283.
14. ↑ D. M. Smith, M. A. Gorman, J. D. Pardo and B. J. Small. 2011. First fossil Orthoptera from the Jurassic of North America. Journal of Paleontology 85(1):102-105
15. ↑ 15.00 15.01 15.02 15.03 15.04 15.05 15.06 15.07 15.08 15.09 15.10 15.11 15.12 15.13 15.14 15.15 15.16 15.17 15.18 15.19 Foster, J. (2007). "The Forgotten Aquatic Denizens: The Fish." pp. 129-131.
16. ↑ 16.0 16.1 16.2 16.3 16.4 16.5 16.6 16.7 16.8 Foster, J. (2007). "Appendix." Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. pp. 327-329.
17. ↑ Foster, J. (2007). "Pycnodontoidea." Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. p. 135.
18. ↑ 18.00 18.01 18.02 18.03 18.04 18.05 18.06 18.07 18.08 18.09 18.10 18.11 18.12 18.13 18.14 18.15 18.16 18.17 18.18 18.19 18.20 18.21 18.22 18.23 18.24 18.25 18.26 18.27 18.28 18.29 18.30 18.31 18.32 18.33 18.34 18.35 18.36 18.37 18.38 18.39 18.40 18.41 18.42 18.43 18.44 18.45 18.46 18.47 18.48 18.49 18.50 18.51 18.52 18.53 Foster, J. (2007). "Table 2.1: Fossil Vertebrates of the Morrison Formation." pp. 58-59.
19. ↑ 19.0 19.1 Foster, J. (2007). "Hulettia hawesi." p. 132-134.
20. ↑ Foster, John; Hunt-Foster, ReBecca; Gorman, Mark; Trujillo, Kelli; Suarez, Celina; McHugh, Julia; Peterson, Joseph; Warnock, Jonathan et al. (2018-04-19). "Paleontology, taphonomy, and sedimentology of the Mygatt-Moore Quarry, a large dinosaur bonebed in the Morrison Formation, western Colorado—Implications for Upper Jurassic dinosaur preservation modes" (in en). Geology of the Intermountain West 5: 23–93. doi:10.31711/giw.v5.pp23-93. ISSN 2380-7601. https://giw.utahgeology.org/giw/index.php/GIW/article/view/24.
21. ↑ 21.0 21.1 21.2 21.3 21.4 Foster, J. (2007). "cf. Leptolepis." p. 135.
22. ↑ 22.0 22.1 22.2 Foster, J. (2007). "Morrolepis schaefferi." pp. 131-132.
23. ↑ Hunt, A. P.; Lucas, S. G. (2025). "The ichnotaxonomy of regurgitalites with description of new ichnogenera". New Mexico Museum of Natural History and Science Bulletin 101: 153–162. https://www.researchgate.net/publication/396523005.
24. ↑ Chure, Daniel J.; Engelmann, George F. (1989), Flynn, John J.; McKenna, Malcolm C.; Chure, Daniel J. et al., eds., "The fauna of the Morrison Formation in Dinosaur National Monument" (in en), Mesozoic/Cenozoic Vertebrate Paleontology: Classic Localities, Contemporary Approaches. Salt Lake City, Utah to Billings, Montana, July 19–27, 1989 (Washington, D. C.: American Geophysical Union): pp. 8–14, doi:10.1029/ft322p0008, ISBN 978-0-87590-608-9, http://doi.wiley.com/10.1029/FT322p0008, retrieved 2024-09-01
25. ↑ 25.00 25.01 25.02 25.03 25.04 25.05 25.06 25.07 25.08 25.09 Foster, J. (2007). "Anura (Frogs)." pp. 135-136.
26. ↑ 26.0 26.1 26.2 26.3 Foster, J. (2007). "Enneabatrachus hechti" p. 137.
27. ↑ 27.0 27.1 27.2 27.3 Foster, J. (2007). "Rhadinosteus parvus." p. 137.
28. ↑ Foster, J. (2007). "Pelobatidae indet." p. 137.
29. ↑ 29.0 29.1 Evans, S. E. and Milner, A. R. (1993). Frogs and salamanders from the Upper Jurassic Morrison Formation (Quarry Nine, Como Bluff) of North America. Journal of Vertebrate Paleontology 13(1):24-30
30. ↑ 30.0 30.1 30.2 30.3 30.4 Foster, J. (2007). "Caudata (Salamanders)" p. 138.
31. ↑ Foster, J. (2007). "Enneabatrachus hechti" p. 137. Note that Dinosaur National Monument is in Utah, see ibid. pg. 6.
32. ↑ 32.0 32.1 Evans, S. E., Lally, C., Chure, D. C., Elder, A., & Maisano, J. A. (2005). A late Jurassic salamander (Amphibia: Caudata) from the Morrison formation of north America. Zoological Journal of the Linnean Society, 143(4), 599-616.
33. ↑ 33.0 33.1 33.2 DeMar Jr, D. G.; Jones, M. E. H.; Carrano, M. T. (2022). "A nearly complete skeleton of a new eusphenodontian from the Upper Jurassic Morrison Formation, Wyoming, USA, provides insight into the evolution and diversity of Rhynchocephalia (Reptilia: Lepidosauria)". Journal of Systematic Palaeontology 20 (1): 779–786. doi:10.1080/14772019.2022.2093139. PMID 2093139.
34. ↑ 34.0 34.1 34.2 34.3 Prothero, D. R.; Estes, R. (1980). "Late Jurassic lizards from Como Bluff, Wyoming and their palaeobiogeographic significance". Nature 286 (5772): 484–486. doi:10.1038/286484a0. Bibcode: 1980Natur.286..484P.
35. ↑ 35.0 35.1 35.2 35.3 Brownstein, C. D.; Meyer, D. L.; Fabbri, M.; Bhullar, B. S.; Gauthier, J. A. (2022). "Evolutionary origins of the prolonged extant squamate radiation". Nature Communications 13 (7087): 1–11. doi:10.1038/s41467-022-34217-5. PMID 36446761. Bibcode: 2022NatCo..13.7087B.
36. ↑ 36.0 36.1 36.2 36.3 Caldwell, M. W.; Nydam, R. L.; Palci, A.; Apesteguía, S. N. (2015). "The oldest known snakes from the Middle Jurassic-Lower Cretaceous provide insights on snake evolution". Nature Communications 6: 1–11. doi:10.1038/ncomms6996. PMID 25625704. Bibcode: 2015NatCo...6.5996C.
37. ↑ Evans, S.E. (1996). "Parviraptor (Squamata: Anguimorpha) and Other Lizards from the Morrison Formation at Fruita, Colorado". The Continental Jurassic 60: 243–248.
38. ↑ 38.0 38.1 38.2 38.3 38.4 38.5 38.6 38.7 Evans, S. E.; Chure, D. J. (1999). "Upper Jurassic lizards from the Morrison Formation of Dinosaur National Monument, Utah". Vertebrate Paleontology in Utah. Miscellaneous Publications of the Utah Geological Survey 99 (1): 151–159.
39. ↑ Benton, M. J. (2015). Vertebrate palaeontology (4th ed.). Chichester, West Sussex. ISBN 978-1-118-40764-6. OCLC 867852756.
40. ↑ Meyer, Dalton; Brownstein, Chase D.; Jenkins, Kelsey M.; Gauthier, Jacques A. (2023-11-29). "A Morrison stem gekkotan reveals gecko evolution and Jurassic biogeography" (in en). Proceedings of the Royal Society B: Biological Sciences 290 (2011). doi:10.1098/rspb.2023.2284. ISSN 0962-8452.
41. ↑ Foster, J. (2007). "Saurillodon sp." p. 145.
42. ↑ Randall L. Lydam, Daniel J. Chure and Susan E. Evans (2013). "Schillerosaurus gen. nov., a replacement name for the lizard genus Schilleria Evans and Chure, 1999 a junior homonym of Schilleria Dahl, 1907". Zootaxa 3734 (1): 99–100. doi:10.11646/zootaxa.3736.1.6. PMID 25112616. http://www.mapress.com/zootaxa/2013/f/zt03736p100.pdf.
43. ↑ 43.0 43.1 Foster, J. (2007). "Schilleria utahensis" p. 145.
44. ↑ Evans, Susan E.; Chure, Daniel J. (1998). "Morrison Lizards: structure, relationships and biogeography". Modern Geology 23: 35–48. https://doc.rero.ch/record/14349.
45. ↑ 45.00 45.01 45.02 45.03 45.04 45.05 45.06 45.07 45.08 45.09 45.10 Gaffney, E. S. (1979). The Jurassic turtles of North America. Bulletin of the AMNH; v. 162, article 3.
46. ↑ 46.0 46.1 46.2 Marsh, Othniel Charles (1890-08-01). "Notice of some extinct Testudinata" (in en). American Journal of Science s3-40 (236): 177–179. doi:10.2475/ajs.s3-40.236.177. ISSN 0002-9599. Bibcode: 1890AmJS...40..177M. https://www.ajsonline.org/content/s3-40/236/177.
47. ↑ 47.0 47.1 47.2 Gilmore, C. (1912). Description of a new species of tortoise from the Jurassic of Utah. Annals of the Carnegie Museum 10:7–12.
48. ↑ 48.0 48.1 48.2 Rollot, Yann; Evers, Serjoscha W.; Joyce, Walter G. (2021-10-06). "A redescription of the Late Jurassic (Tithonian) turtle Uluops uluops and a new phylogenetic hypothesis of Paracryptodira" (in en). Swiss Journal of Palaeontology 140 (1): 23. doi:10.1186/s13358-021-00234-y. ISSN 1664-2384. PMID 34721284. Bibcode: 2021SwJP..140...23R.
49. ↑ Pritchard, A. C.; Turner, A. H.; Allen, E. R.; Norell, M. A. (2013). "Osteology of a North American Goniopholidid (Eutretauranosuchus delfsi) and Palate Evolution in Neosuchia". American Museum Novitates 3783 (3783): 1. doi:10.1206/3783.2. edit
50. ↑ 50.0 50.1 Yoshida, Junki; Hori, Atsushi; Kobayashi, Yoshitsugu; Ryan, Michael J.; Takakuwa, Yuji; Hasegawa, Yoshikazu (2021). "A new goniopholidid from the Upper Jurassic Morrison Formation, USA: novel insight into aquatic adaptation toward modern crocodylians". Royal Society Open Science 8 (12). doi:10.1098/rsos.210320. PMID 34909210. Bibcode: 2021RSOS....810320Y.
51. ↑ Allen, Eric Randall (Summer 2012). "Analysis of North American goniopholidid crocodyliforms in a phylogenetic context" (pdf). Iowa Research Online. doi:10.17077/etd.317zy27t. https://iro.uiowa.edu/esploro/outputs/graduate/Analysis-of-North-American-goniopholidid-crocodyliforms/9983776735102771.
52. ↑ 52.0 52.1 A. D. Walker. A Revision of the Jurassic Reptile Hallopus victor (Marsh), with Remarks on the Classification of Crocodiles. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences Vol. 257, No. 816 (Feb. 26, 1970), pp. 323-372
53. ↑ 53.0 53.1 Göhlich, U., L.M. Chiappe, J.M. Clark, and H.-D. Sues (2005) The systematic position of the Late Jurassic alleged dinosaur Macelognathus (Crocodylomorpha: Sphenosuchia). Canadian Journal of Earth Sciences 42: 307–321.
54. ↑ Foster, J. (2018). "A new atoposaurid crocodylomorph from the Morrison Formation (Upper Jurassic) of Wyoming, USA" (in en). Geology of the Intermountain West 5: 287–295. doi:10.31711/giw.v5i0.32. ISSN 2380-7601. https://www.utahgeology.org/giw/index.php/giw/article/view/32. Retrieved 2018-12-22.
55. ↑ 55.0 55.1 55.2 55.3 55.4 55.5 Foster, J. (2007). "Soaring Overhead: The Pterosaurs." pp. 157-158.
56. ↑ 56.0 56.1 Foster, J. (2007). "Laopteryx priscus." p. 160.
57. ↑ Jensen, J. A., and Padian, K. (1989) Small pterosaurs and dinosaurs from the Uncompahgre fauna (Brushy Basin Member, Morrison Formation: ?Tithonian), late Jurassic, western Colorado. Journal of Paleontology 63:363–374.
58. ↑ 58.0 58.1 McLain, M. A., & Bakker, R. T. (2018). Pterosaur material from the uppermost Jurassic of the uppermost Morrison Formation, Breakfast Bench Facies, Como Bluff, Wyoming, including a pterosaur with pneumatized femora. Geological Society, London, Special Publications, 455(1), 105-124.
59. ↑ 59.0 59.1 Sprague, M. & McLain, M. A. (2018). Resolving the Mesadactylus Complex of Dry Mesa Quarry, Morrison Formation, Colorado. Journal of Vertebrate Paleontology, Program and Abstracts, 2018, p. 220.
60. ↑ 60.0 60.1 Stephen Czerkas; Tracy Ford (2018). "Pterosaur or diapsid? The search for the true Utahdactylus". Flugsaurier 2018: The 6th International Symposium on Pterosaurs. Los Angeles, USA. Abstracts: 35–36.
61. ↑ 61.0 61.1 Foster, J. (2007). "Harpactognathus gentryii." p. 160.
62. ↑ 62.0 62.1 Foster, J. (2007). "Kepodactylus insperatus." p. 160.
63. ↑ Lockley et al. (2008).
64. ↑ Foster, J.R. 2009. Preliminary body mass estimates for mammalian genera of the Morrison Formation (Upper Jurassic, North America). PaleoBios 28(3):114-122.
65. ↑ 65.0 65.1 Brian M. Davis; Kai R.K. Jäger; Guillermo W. Rougier; Kelli Trujillo; Kevin Chamberlain (2022). "A morganucodontan mammaliaform from the Upper Jurassic Morrison Formation, Utah, USA". Acta Palaeontologica Polonica. doi:10.4202/app.00955.2021.
66. ↑ Julia A. Schultz; Bhart-Anjan S. Bhullar; Zhe-Xi Luo (2018). "Re-examination of the Jurassic mammaliaform Docodon victor by computed tomography and occlusal functional analysis". Journal of Mammalian Evolution. in press. doi:10.1007/s10914-017-9418-5.
67. ↑ Averianov, Alexander O.; Sues, Hans‐Dieter (March 16, 2026). "A docodont ulna from the Upper Jurassic in North America and its implications for the forelimb morphology in Docodonta (Mammaliaformes)" (in en). The Anatomical Record. doi:10.1002/ar.70192. ISSN 1932-8486.
68. ↑ John R. Foster; Darrin C. Pagnac; ReBecca K. Hunt-Foster (2020). "An unusually diverse northern biota from the Morrison Formation (Upper Jurassic), Black Hills, Wyoming". Geology of the Intermountain West 7: 29–67. doi:10.31711/giw.v7.pp29-67.
69. ↑ 69.0 69.1 69.2 69.3 A.O. Averianov and T. Martin (2015). "Ontogeny and taxonomy of Paurodon valens (Mammalia, Cladotheria) from the Upper Jurassic Morrison Formation of USA". Proceedings of the Zoological Institute of the Russian Academy of Sciences 319 (3): 326–340. http://www.zin.ru/journals/trudyzin/doc/vol_319_3/TZ_319_3_Averyanov.pdf.

### Bibliography

* Foster, J. (2007). Jurassic West: The Dinosaurs of the Morrison Formation and Their World. Indiana University Press. 389pp. ISBN 978-0-253-34870-8
* Lockley, M.; Harris, J.D.; and Mitchell, L. 2008. "A global overview of pterosaur ichnology: tracksite distribution in space and time." Zitteliana. B28. p. 187-198. ISSN 1612-4138

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