Paleobiota of the Hell Creek Formation

This is an overview of the fossil flora and fauna of the Maastrichtian-Danian Hell Creek Formation.

Invertebrates

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Insects

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Insects from the groups Diptera, Zygoptera, and possibly Hemiphlebiidae have been unearthed in Hell Creek in amber.[1][2] Fossils found in the Hell Creek Formation and the Fort Union Formation of these insects went extinct during the K-T Event.[3][4][5]

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.
Genus Species Location Stratigraphic position Material Notes Images

Cephaloleichnites

Cephaloleichnites strongi

Possible hispine beetle herbivory on Zingiberopsis attenuta

?Hemiphlebiidae[2]

Indeterminate

Possible Hemiphlebiid damselflies

Brachycera[1]

Indeterminate

Undescribed brachyceran flies

Nematocera[1]

Indeterminate

Undescribed nematoceran flies

Aeschnidiidae[6]

Indeterminate

Indeterminate aeschnidiid dragonflies

Molluscs

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Genus Species Location Stratigraphic position Material Notes Images
Plesielliptio

P. postbiplicatus

Freshwater Unionid river mussels

P. gibbosoides

Freshwater Unionid river mussels

P. whitfieldi

Freshwater Unionid river mussels

Rhabdotophorus R. aldrichi Freshwater mussel of family Unionidae.[7]
Pleurobema P. cryptorhynchus Freshwater mussel of family Unionidae.[7]
Plethobasus P. aesopiformis Freshwater mussel of family Unionidae.[7]
P. biesopoides
Quadrula Q. cylindricoides Freshwater mussel of family Unionidae.[7]
Proparreysia P. verrucosiformis Freshwater mussel of family Unionidae.[7]
P. holmesiana
P. barnumi
P. percorrugata
P. pyramidatoides
P. letsoni
P. retusoides
P. corbiculoides
P. paucinodosa
?Obovaria ?O. pyramidella
Corbicula C. cf. subelliptica
C. sp South Dakota From a marine facies ("tongue"). Modern members of the genus live in fresh water
Sphaerium S. beckmani "Pill clam". "Nut clam". "Fingernail clam". "Pea clam". Family Sphaeriidae.
Pleiodon Indeterminate
Campeloma C. sp Freshwater snail
Anomia A. gryphorhyncha Bivalve. Family Anomiidae. From a marine facies ("tongue") in South Dakota.
Crassostrea C. subtrigonalis Oyster. Family Ostreidae. Collected from a marine facies ("tongue") in South Dakota.
Granocardium G. sp Bivalve. Family Cardiidae (cockle). Collected from a marine facies ("tongue") in South Dakota.
?Hiatella ?H. sp Bivalve. Present members of this genus are rock borers. Collected from a marine facies ("tongue") in South Dakota.
Leptosolen indeterminate Bivalve. Family Cultellidae. Collected from a marine facies ("tongue") in South Dakota.
Sphenodiscus S. lenticularis Ammonite. From a marine facies ("tongue") in South Dakota.
Discoscaphites D. rossi Microconch of an ammonite. From a marine facies ("tongue") in South Dakota.
Scaphitidae indeterminate Ammonite. From a marine facies ("tongue") in South Dakota. Other attributes: specimen has hooks on its shell.

Amphibians

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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.
Genus Species Location Stratigraphic position Material Notes Images

Anura (frog)[8]

indeterminate[8]

  • North Dakota[9]
  • South Dakota[9]

Middle to upper Hell Creek Formation[10]

3 unassigned specimens[11]

Anura indet. consists of material not currently assigned to any genus of frog.[8]

Barbourula[12]

Indeterminate[13]

Caudata (salamander)[8]

indeterminate[8]

  • North Dakota[9]
  • South Dakota[9]

Lower to uppermost Hell Creek Formation[10]

149 unassigned specimens[11]

Material of Caudata indet. is not currently assigned to any genus.[8]

Eopelobates[15]

Indeterminate[13]

Habrosaurus[8][13]

H. dilatus[8][13]

Middle to upper Hell Creek Formation[10]

6 specimens[11]

Habrosaurus is a sirenid amphibian.[8]

Lisserpeton[13]

L. bairdi[13]

Opisthotriton[8][13]

O. kayi[8][13]

Lower to upper Hell Creek Formation[10]

22 specimens[11]

Opisthotriton is classified as a Batrachosauroididae.[8]

Paranecturus[16]

P. garbanii[16]

A member of Proteidae.[16]

Proamphiuma[13]

P. cretacica[13]

Prodesmodon[13]

P. copei[13]

Scapherpeton[8][13]

S. tectum[8][13]

Lower to upper Hell Creek Formation[10]

144 specimens[11]

Scapherpeton is a scapherpetonid that is very common in the Hell Creek Formation.[8]

Scotiophryne[13]

S. pustulosa[13]

A small frog

Fish

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Bony fish

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Bony fishes
Genus Species Location Stratigraphic position Material Notes Images
Acipenser[8][17][18] A. amnisinferos[18] A sturgeon
A. eruciferus[17]
A. praeparatorum[18]
  • North Dakota (Tanis)[18]
A. cf. A. amnisinferos[18]
  • North Dakota (Tanis)[18]

cf. A. sp.[8]

  • North Dakota[9]
  • South Dakota[9]

Lower to upper Hell Creek Formation[10]

18 specimens are tentatively assigned to Acipenser sp.[11]

Acipenser sp. is tentatively referred to the genus.[8]

Anchiacipencer[20] A. acanthaspis[20] A sturgeon, originally considered as indeterminate material[20]
Belonostomus[8][17] B. longirostris[8][17] Lower to upper Hell Creek Formation[10] 28 specimens[11] A long-snouted slender fish classified as an aspidorhynchid.[8]
Coriops[21] C. amnicolus[21]

Cyclurus[17]

C. fragosus[17]

A small amiid fish (ubiquitous). Previously known as Amia fragosa or Kindleia fragosa.[23] 2610 specimens have been assigned to Kindleia, making it an extremely common genus.[11]

 
Cyclurus kehreri from Messel Pit, species within same genus as C. fragosus
"Lepisosteus"[8][17] "L. occidentalis"[8][17] Lower to uppermost Hell Creek Formation[10] 938 specimens are assigned to Lepidosteus[11] A lepidosteid that is very common in the Hell Creek Formation.[8] Nomen dubium.[24]  
Melvius[8] M. thomasi[8]
  • North Dakota[9]
  • South Dakota[9]
Lower to upper Hell Creek Formation[10] 6 specimens are assigned to Melvius[11] A large amiid fish.[8]
Phyllodus P. paulkatoi Fish with columnar teeth
Palaeolabrus[17] P. montanensis[17]
Paleopsephurus[17] P. wilsoni[17] A paddlefish
Paralbula[25] P. casei[25]
Parapsephurus[26] P. willybemisi[26]
  • North Dakota (Tanis)[26]
A paddlefish  
Platacodon[21] P. nanus[21] Small teleost fish
Protamia[17] Indeterminate[17]
Pachyrhizodontoidei Indeterminate Fish
Polyodontidae[27] Indeterminate[27] Paddlefish
Protoscaphirhynchus[17] P. squamosus[17] A sturgeon
Pugiopsephurus[26] P. inundatus[26]
  • North Dakota (Tanis)[26]
A paddlefish

Cartilaginous fish

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Chondrichthyes reported from the Hell Creek Formation
Genus Species Location Stratigraphic position Material Notes Images

Chiloscyllium[28]

C. sp.[28]

A member of Hemiscylliidae.[28]

 

Galagadon[29]

G. nordquistae

  • South Dakota

Isolated teeth

A carpet shark

Lonchidion[30]

L. selachos[30]

Lower to upper Hell Creek Formation[10]

40 specimens[11]

A genus of prehistoric sharks in the family Hybodontidae. It makes up 0.4% of the remains of the vertebrates of the Hell Creek Formation.[8]

Myledaphus[28]

M. pustulosus[28]

Lower to upper Hell Creek Formation.[10]

1677 specimens[11] previously assigned to M. bipartitus.[28]

Is an anacoracid batoid[8] rajiform related to guitarfishes.[28] Described on the basis of teeth formerly assigned to the species M. bipartitus.[28] The material assigned to Myledaphus bipartitus and makes up 16.5% of the vertebrate remains.[8]

Protoginglymostoma[28]

P. estesi[28]

A member of Ginglymostomatidae.[28] Formerly assigned to the genus Brachaelurus.

Restesia[28]

R. americana[28]

Middle Hell Creek Formation[10]

5 specimens previously assigned to Squatirhina[11]

A wobbegong-like shark.[28] Formerly assigned to Squatirhina. The remains consist of 0.05% of the vertebrates.[11] Also known from the Lance Formation.[28]

Carcharhinidae indet.[29]

Indeterminate

  • South Dakota

An isolated tooth.

Scapanorhynchus[32]

.sp.

  • North Dakota

Dinosaurs

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A paleo-population study is one of the most difficult of analyses to conduct in field paleontology. Here is the most recent estimate of the proportions of the eight most common dinosaurian families in the Hell Creek Formation, based on detailed field studies by White, Fastovsky and Sheehan.[33]

  • Ceratopsidae 61%
  • Hadrosauridae 23%
  • Ornithomimidae 5%
  • Tyrannosauridae 4%
  • Hypsilophodontidae 3%
  • Dromaeosauridae 2%
  • Pachycephalosauridae 1%
  • Troodontidae 1% (represented only by teeth)
 
Pie chart of the time averaged census for large-bodied dinosaurs from the entire Hell Creek Formation in the study area.

Outcrops sampled by the Hell Creek Project were divided into three sections: lower, middle and upper slices. The top and bottom sections were the focus of the PLoS One report, and within each portion many remains of Triceratops, Edmontosaurus, and Tyrannosaurus were found. Triceratops was the most common in each section, but, surprisingly, Tyrannosaurus was just as common, if not slightly more common, than the hadrosaur Edmontosaurus. In the upper Hell Creek section, for example, the census included twenty two Triceratops, five Tyrannosaurus, and five Edmontosaurus.

The dinosaurs Thescelosaurus, Ornithomimus, Pachycephalosaurus and Ankylosaurus were also included in the breakdown, but were relatively rare. Other dinosaurs, such as Sphaerotholus, Denversaurus, Torosaurus, Struthiomimus, Acheroraptor, Dakotaraptor, Pectinodon, Richardoestesia, Paronychodon, Anzu, Leptorhynchos and Troodon (more likely Pectinodon), were reported as being rare and are not included in the breakdown.

The dinosaur collections made over the past decade during the Hell Creek Project yielded new information from an improved genus-level collecting schema and robust data set that revealed relative dinosaur abundances that were unexpected, and ontogenetic age classes previously considered rare. We recognize a much higher percentage of Tyrannosaurus than previous surveys. Tyrannosaurus equals Edmontosaurus in U3 and in L3 comprises a greater percentage of the large dinosaur fauna as the second-most abundant taxon after Triceratops, followed by Edmontosaurus. This is surprisingly consistent in (1) the two major lag deposits (MOR loc. HC-530 and HC-312) in the Apex sandstone and Jen-rex sand where individual bones were counted and (2) in two thirds of the formation reflected in L3 and U3 records of dinosaur skeletons only.

Triceratops is by far the most common dinosaur at 40% (n = 72), Tyrannosaurus is second at 24% (n = 44), Edmontosaurus is third at 20% (n = 36), followed by Thescelosaurus at 8% (n = 15), Ornithomimus at 5% (n = 9), and Pachycephalosaurus and Ankylosaurus both at 1% (n = 2) are relatively rare.

Fossil footprints of dinosaurs from the Hell Creek Formation are very rare. As of 2017, there is only one find of a possible Tyrannosaurus rex footprint, dating from 2007 and described a year later.[34] A trackway made by mid-sized theropod, possibly a small tyrannosaurid individual, was discovered in South Dakota in 1997, and in 2014 these footprints were named Wakinyantanka styxi.[35]

Ornithischians

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Ankylosaurs

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Indeterminate nodosaur remains have been unearthed in the Hell Creek Formation and other nearby areas.[36][37]

Ankylosauria reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Ankylosaurus[38]

A. magniventris[38]

Upper A partial skull, teeth, cervical vertebrae, dorsal vertebrae, caudal vertebrae, right scapulocoracoid, otic capsule, maxilla fragment, right jugal, left jugal and quadratojugal, sacral centra, additional fragments of the cervical half rings and a partial tail club handle.

An ankylosaur. Also found in the Lance, Frenchman, Ferris and Scollard Formations.

 

Denversaurus[38]

D. schlessmani[38]

Skull roof, pelvis and osteoderms.

A nodosaurid ankylosaur whose remains have been found in the Lance and Laramie Formation.[36]

 

Pachycephalosaurs

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An undescribed and unnamed pachycephalosaur is present in North Dakota.[40] Pachycephalosaur remains have been unearthed in Montana as in the case of Platytholus and the now invalid genus Stenotholus kohleri, which is now a junior synonym of Pachycephalosaurus.[41]

Pachycephalosaurs reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Dracorex[42]

D. hogwartsia[42]

A pachycephalosaur, possibly synonymous with Pachycephalosaurus.

 

Pachycephalosaurus[38]

P. wyomingensis[38]

A pachycephalosaur. Also found in the Lance Formation.

 

Platytholus[43] P. clemensi[43] A partial skull[43] A medium-sized pachycephalosaurine closely related to Acrotholus and Prenocephale

 

Sphaerotholus[38]

S. buchholtzae[38]

"Skull material."[46]

A pachycephalosaur whose remains have also been found in the Frenchman Formation. Genus also known from the Kirtland Formation, Dinosaur Park Formation and the Horseshoe Canyon Formation.

 

S. triregnum[45]

A left squamosal.[45]

A pachycephalosaur distinguished from S. buchholtzae by its more ornamented squamosal.[45]

Stygimoloch[38]

S. spinifer[38]

  • Montana[38]
  • South Dakota
  • North Dakota

A pachycephalosaur, possibly synonymous with Pachycephalosaurus. Also found in the Ferris Formation and the Lance Formation.

 

Ceratopsians

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Indeterminate ceratopsid teeth and some identifiable bones from Triceratops can be extremely common.[47][48][49][50][51] 8.31% of all vertebrate remains from the Hell Creek Formation are unassigned ceratopsids.[11] In 2012, a new unidentified species of chasmosaur ceratopsian with noticeable differences from Triceratops was unearthed in South Dakota by a fossil hunter named John Carter.[52][53][54]

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.
Ceratopsians reported from the Hell Creek Formation
Genus Species Synonyms State Stratigraphic position Material Notes Images

Leptoceratops[38]

L. gracilis[38]

A small primitive-looking ceratopsian. Fossils have also been found in the Lance Formation in Wyoming.[55]

 

Tatankaceratops[56]

T. sacrisonorum[56]

A controversial ceratopsian possibly synonymous with Triceratops[57]

 

Torosaurus[8][38]

T. latus[8][38]

Upper Hell Creek Formation[10]

A ceratopsian that was once proposed to be synonymous with Triceratops,[58] but is now regarded as a valid and distinct ceratopsian.[59] A rare ceratopsid.[10] Fossils have been in the Lance Formation, Javelina Formation, North Horn Formation, Laramie Formation, El Picacho Formation, Frenchman Formation and Scollard Formation.

 

Triceratops[8][38]

T. horridus[8][38]

Lowermost to the middle Hell Creek Formation[61]

Four specimens are assigned to Triceratops horridus from the Hell Creek Formation.[11] Isolated, shed ceratopsid teeth are incredibly common in the Hell Creek and Lance Formations, being by most collectors, with some being nothing more than worn down fragments up to superb teeth containing complete, preserved roots. Because the teeth of different ceratopsians are so similar to one another, its hard to differentiate between genera/species, but based on the abundance of identifiable bones belonging to Triceratops in Lancian-aged North American formations, especially the Hell Creek, isolated ceratopsid teeth from the lower and middle Hell Creek Formation have a high likelihood of originating from T. horridus.

A ceratopsian.[8] Also found in the Evanston, Frenchman, Lance, Laramie, and Scollard Formations.

 

T. prorsus[38]

  • Montana[38]
  • South Dakota
Upper 1/3 of the Hell Creek Formation[61]

Very common.[citation needed] Because the teeth of different ceratopsians are so similar to one another, its hard to differentiate between genera/species, but based on the abundance of identifiable bones belonging to Triceratops in Lancian-aged North American formations, especially the Hell Creek, isolated ceratopsid teeth from the upper 1/3 of the Hell Creek Formation have a high likelihood of originating from T. prorsus.

Also found in the Frenchman and Lance Formations.

 

Ornithopods and relatives

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Indeterminate hadrosaurid remains are very common in the Hell Creek Formation.[8]

Ornithopods and Thescelosaurs reported from the Hell Creek Formation
Genus Species Synonyms State Stratigraphic position Material Notes Images

Edmontosaurus

E. annectens

  • Anatosaurus annectens[62]
  • Anatotitan copei[62]

Very common.

A hadrosaur. Also found in the Denver, Frenchman, Lance, Laramie, and Scollard Formations.[65][66] Hatchlings have also been unearthed.[66]

 

Thescelosaurus[39]

T. garbanii[67][68]

  • Bugenasaura garbanii[68]

T. neglectus[8][39]

Lower to upper Hell Creek Formation[10]

50 specimens[11]

A small thescelosaurine. Also found in the Frenchman, Lance, Laramie, and Scollard Formations.[70] Two species are known from Hell Creek; T. neglectus and T. garbanii.[71]

 

Theropods

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Theropod tracks have been found in South Dakota.[39] A trackway from South Dakota, named Wakinyantanka, was made by a mid-sized theropod with three slender toes, possibly a small tyrannosaurid.[35] A second footprint that may have been made by a specimen of Tyrannosaurus was first reported in 2007 by British paleontologist Phil Manning, from the Hell Creek Formation of Montana.[72] This second track measures 72 centimeters (28 in) long, shorter than the track described by Lockley and Hunt. Whether or not the track was made by Tyrannosaurus is unclear, though Tyrannosaurus is the only large theropod known to have existed in the Hell Creek Formation, though in past albertosaurine remains have described here but its most likely that they are the remains of Tyrannosaurus rex.[73][74] Theropod remains are very common in Hell Creek, some of which belong to indeterminate species on maniraptorans.[75]

Alvarezsaurs

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Alvarezsaurs reported from the Hell Creek Formation
Genus Species Synonyms State Stratigraphic position Material Notes Images
"Ornithomimus" "O." minutus

Trierarchuncus[76]

T. prairiensis[76]

upper Hell Creek Formation.

An alvarezsaur known from a partial post-cranial skeleton.[77]

Tyrannosaurids

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Tyrannosaurids reported from the Hell Creek Formation
Genus Species Synonyms State Stratigraphic position Material Notes Images

Tyrannosaurus[8][38]

T. rex[8][38]

Lower to upper Hell Creek Formation.

A tyrannosaur, known from several specimens including a juvenile nicknamed "Jane".[78] Also found in the Denver, Frenchman, Hill Creek South, Javelina, Lance, Ferris, Livingston, McRae, North Horn, Scollard, Willow Creek Formation, and also found in Lomas Coloradas Formations. Isolated teeth in the Hell Creek are common enough to be dug commercially by collectors, but rare enough that they are often sold for very high prices with fragmentary teeth usually beginning at least in the hundreds of USD, and complete teeth in the thousands of USD. Perhaps the best known iconic dinosaur.

 

Nanotyrannus?[8][38]

N.lancensis[8][38]

  • Stygivenator?
  • Deinodon lancensis
  • Albertosaurus lancensis[38]
  • Tyrannosaurus lancensis?
  • Montana
  • Wyoming

Lower to upper Hell Creek Formation.

A few specimens are known

Invalid genus, now recognised as juvenile T. rex.

 

Ornithomimosaurs

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Ornithomimid remains are not uncommon in the Hell Creek Formation.[8] Fifteen specimens from the Hell Creek Formation are undetermined ornithomimids[11]

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.
Ornithomimids reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

"Orcomimus"

unnamed

One partial skeleton.

An ornithomimid; nomen nudum.

Struthiomimus[39]

S. sedens[80]

  • Montana
  • AMNH 975, a foot claw
  • UCMP 154569, a partial skeleton

A large ornithomimid similar to Gallimimus in size. Also found in the Lance Formation.[80]

 

Ornithomimus

O. velox[80]

  • Montana
  • South Dakota
  • North Dakota

Fragmentary specimens

An ornithomimid which was also found in the Denver Formation.

 

Oviraptorosaurs

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Oviraptorosaur fossils have been found at the Hell Creek Formation for many years, most notably from isolated elements until the discovery of Anzu. In the past, oviraptorosaur fossils found were thought to have belonged to Caenagnathus, Chirostenotes, and Elmisaurus.[38][81][82][83][84] In 2016, an undescribed large-bodied caenagnathid was unearthed in Montana.[85]

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.
Oviraptorosaurs reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Anzu[38][81]

A. wyliei[81]

Lower to upper Hell Creek Formation[11]

12 well-preserved specimens[10]

One of the largest known oviraptorosaurs, and the largest known from North America. Material previously assigned to Caenagnathidae indet. is now placed in the genus Anzu.[81]

 

Caenagnathidae

Indeterminate

  • Montana

Similar to Citipes and Elmisaurus.

Eoneophron E. infernalis A partial right hindlimb[86] Closely related to Citipes and Elmisaurus.[86]  

Eumaniraptorans

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Historically, numerous teeth have been attributed to various dromaeosaurid and troodontid taxa with known body fossils from only older formations, including Saurornithoides, Zapsalis, Dromaeosaurus, Saurornitholestes, and Troodon. However, in a 2013 study, Evans et al. concluded that there is little evidence for more than a single dromaeosaurid taxon, Acheroraptor, in the Hell Creek-Lance assemblages, which would render these taxa invalid for this formation. This was disproved in a 2015 study, DePalma et al., when they described the new genus Dakotaraptor, a large species of dromaeosaur.[87] Fossilized teeth of various troodontids and coelurosaurs are common throughout the Hell Creek Formation; the best known examples include Paronychodon, Pectinodon and Richardoestesia, respectively. Teeth belonging to possible intermediate species of Dromaeosaurus[88] and Saurornitholestes[89] have been unearthed at the Hell Creek Formation and the nearby Lance Formation.

Eumaniraptorans reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Acheroraptor[87]

A. temertyorum [87]

Lower? to upper Hell Creek Formation[8]

  • ROM 63777, a maxilla and tooth[87]
  • ROM 63778, a partial dentary[87]
  • isolated teeth[87]

A velociraptorine dromaeosaurid. Teeth previously referred to various Campanian dromaeosaurids Saurornitholestes and Dromaeosaurus, frequently found throughout the formation, probably belong to this one species. Evans et al. conclude that there is little evidence for the former two taxa being present in the Hell Creek-Lance assemblages.[87]

 

Avisaurus[8]

A. archibaldi[8][38]

Middle Hell Creek Formation[10]

  • UCMP 117600, holotype, a tarsometatarsus
  • PU 17324, a tarsometatarsus

An avisaurid.[8]

 

cf. A. archibaldi[90]

Uppermost Hell Creek Formation[90]

  • YPM 57235, a coracoid

An avisaurid tentatively referred to A. archibaldi based on its size.[90]

A. darwini[91]
  • Montana
  • DDM 1577.730, a right tarsometatarsus

A. sp.[92]

  • distal tarsals, metatarsus (juvenile)[92]
  • MOR3070, a partial right tarsometatarsus

Brodavis[93]

B. baileyi[93]

  • UNSM 50665, a left tarsometatarsus missing proximal end, trochleae II and III.[93]

A primitive hesperornithiform.[93]

Dakotaraptor[94]

D. steini [94]

Upper Hell Creek Formation[94]

  • PBMNH.P.10.113.T, a partial skeleton.[94]
  • PBMNH.P.10.115.T, a tibia.[94]
  • PBMNH.P.10.118.T, an astragalocalcaneum.[94]
  • isolated teeth.[94]

A large dromaeosaurid.[87]

 

Magnusavis[91] M. ekalakaensis
  • Montana
  • CCM V2019.5.1, partial right tarsometatarsus and toe bone
A large enantiornithine closely related to avisaurids.

Paronychodon[95]

P. caperatus[95]

  • North Dakota, South Dakota, and Montana.
  • YPM 10624, a fossilized tooth. Teeth of this genus have been found too.

A troodontid theropod who is known from fossil teeth. Fossils have also been found in the Lance Formation in Wyoming.

 

Pectinodon[96][95]

P. bakkeri[96][95]

  • Montana
  • UCM 38445, a fossilized tooth. Teeth of this genus have been found too.
  • A troodontid theropod who is known from fossil teeth. Fossils have also been found in the Lance Formation in Wyoming.

 

Potamornis[93]

P. skutchi [97]

  • UCMP 117605, a tarsometatarsus

A hesperornithiform also found in the Lance Formation.[97]

Richardoestesia[98][99]

R. sp.[98][99]

  • Montana.
  • Teeth.

A coelurosaur that is known from teeth and from two species Richardoestesia gilmorei and Richardestesia isosceles, which have also been unearthed in the Lance Formation in Wyoming.

 

"Styginetta"[100] "S. lofgreni."
  • Montana.
A Presbyornithid, it is notable for being one of the few birds known to have survived the Cretaceous–Paleogene extinction event.

"Unnamed enantiornithine B"[90]

Unnamed

  • YPM 57823, a partial coracoid[90]

An unnamed enantiornithean.[90]

"Unnamed hesperornithiform A"[90]

Unnamed

  • Montana
  • UCMP 13355, a tarsometatarsus

A primitive hesperornithiform.[93] The Hell Creek specimen was referred to the same unnamed taxon as RSM P 2315.1 from the Canadian Frenchman Formation.[90] RSM P 2315.1 was later made the holotype of Brodavis americanus.[93] May be a synonym of Potamornis.[90]

"Unnamed ornithurine B"[90]

Unnamed[101]

  • UCMP 129143, a partial coracoid[90]

An ornithurine possibly similar to Cimolopteryx[90][101]

"Unnamed ornithurine C"[90]

Unnamed

  • SDSM 64281A, a partial coracoid[90]
  • SDSM 64281B, a partial coracoid[90]
  • UCMP 175251, a partial coracoid[90]
  • MOR 2918, a partial coracoid[90]

An ornithurine, also present in the Lance Formation and Fort Union Formation, one of the few individual bird species known to have survived the Cretaceous–Paleogene extinction[90]

"Unnamed ornithurine D"[90]

Unnamed

  • UCMP 187207, a partial coracoid[90]

An ichthyornithean also present in the Frenchman Formation[90]

Pterosaurs

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Undescribed pterosaur remains were reported from North Dakota.[103] A specimen of an azhdarchid pterosaur from Montana likely belongs to Quetzalcoatlus, though it is not diagnostic to the species level.[103]

Pterosaurs of the Hell Creek Formation
Taxa Species State Stratigraphic location Material Notes Images

cf. Quetzalcoatlus[103]

Indeterminate

Montana[103] A single azhdarchid neck bone which may belong to the genus Quetzalcoatlus.[103]
 

Crocodylomorphs

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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.
Crocodylomorphs reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Borealosuchus[104]

  • Montana
  • North Dakota
  • South Dakota

Extinct genus of crocodylians that lived from the Late Cretaceous to the Eocene in North America.

   

Brachychampsa[104]

  • Montana
  • South Dakota
  • North Dakota

Extinct genus of alligatoroid.

   

Thoracosaurus[104]

  • Montana

Extinct genus of gavialoid crocodilian which existed during the Late Cretaceous and early Paleocene.

   

Turtles

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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.
Turtles reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Adocus[105]

Indeterminate[105]

Extinct genus of aquatic turtles belonging to the family Adocidae.

 

Axestemys[107]

A. infernalis[107]

  • Montana
  • North Dakota
  • South Dakota
  • Wyoming

A turtle belonging to the family Trionychidae. Its fossils from the Hell Creek Formation were formerly assigned to the late Campanian species Axestemys splendidus.[108][107]

Compsemys[105]

C. victa[105]

A relative of Dermatemydidae.

Peckemys

P. brinkman

  • Montana
  • North Dakota

A relative of Baenidae.

Emarginachelys

E. cretacea

  • Montana

A relative of chelydrids.[109]

Eubaena[105]

E. cephalica[105]

Baenid turtle

Gamerabaena

G. sonsalla

  • North Dakota

Extinct genus of baenid turtle.

Palatobaena

P. cohen

  • North Dakota

A relative of extinct family of cryptodiran turtles.

Cedrobaena

C. putorius

  • South Dakota
  • North Dakota

A relative of Baenidae.

Gilmoremys

G. lancensis

  • Montana
  • North Dakota

Trionychidae related to the softshell turtle.

Hoplochelys[109]

H. clark[109]

A kinosternoid related to the Central American river turtle.[109]

Hutchemys[110] H. walkerorum[110] A shell[110] A Plastomeninae related to the softshell turtle.[110]

Plastomenus

P. sp

Trionychidae turtle.

Basilemys

B. sinuosa

Largest dermatemydid land tortoise.

Trionyx[105]

Indeterminate[105]

A genus of softshell turtles belonging to the family Trionychidae.

 

Aspideretoides

A. foveatus

Trionychidae turtle.

 

Helopanoplia

H. distincta

Trionychidae turtle.

Judithemys

J. backmani

Thin-shelled macrobaenid turtle.

Plesiobaena

P. antiqua

Baenid turtle.

Stygiochelys

S. estesi

Baenid turtle.

Neurankylus

N. eximius

Largest baenid turtle in Hell Creek Formation.

 

Saxochelys[111]

S. gilberti

  • North Dakota
  • A population of over 30 individual skeletons

A member of the family Baenidae.

Thescelus

T. insiliens

Baenid turtle.

Chelydridae

Indeterminate

Chelydrids-like turtle.

Squamata

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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.
Squamates reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Cemeterius[112][113]

C. monstrosus[112][113]

A platynotan lizard of uncertain phylogenetic placement, also known from the Lance Formation.[112]

Cerberophis[112][113]

C. robustus[112][113]

An alethinophidian snake of uncertain phylogenetic placement.[112]

Obamadon[112][113]

O. gracilis[112][113]

A polyglyphanodontian lizard of uncertain phylogenetic placement. Also known from the Lance Formation.[112]

Peneteius[112]

P. aquilonius[112]

A chamopsiid polyglyphanodontian lizard.[112]

Haptosphenus

H. placodon

Teiidae lizard.

 

Leptochamops

L. denticulatus

Small Teiidae lizard.

Chamops

C. segnis

Largest Teiidae lizard in Hell Creek Formation

Contogenys

C. sloani

Scincidae? lizard.

Exostinus

E. lancensis

xenosaurid lizard.

 

Proxestops

P. jepseni

Anguidae lizard.

Parasaniwa

P. wyomingensis

Necrosaurid lizard.

Paraderma

P. bogerti

Helodermatidae? lizard.

Palaeosaniwa

P. canadensis

A large Monstersauria lizard, closely related to today's varanid lizards. It was the largest lizard in the Hell Creek formation.

Boidae

Indeterminate

Snake. Earliest-known boid.

 

Mosasauridae

Indeterminate

Indeterminate mosasaur remains have been unearthed in North Dakota; they may belong to a mosasaur measuring 11 m (36 ft) in length.[114][115]

 

Choristoderans

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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.
Choristoderans reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Champsosaurus[104]

C. sp.[104]

  • Montana

champsosaur.

 
 

Mammals

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Multituberculates

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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.
Multituberculates reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Cimexomys[116]

C. minor[116]

A multituberculate of uncertain phylogenetic placement.

 

Cimolodon[116]

C. nitidus

  • Montana
  • North Dakota[116]

A cimolodontid multituberculate.

C. cf. nitidus[116]

A cimolodontid multituberculate.

C. sp.[116]

A cimolodontid multituberculate.

Cimolomys[116]

C. gracilis

A cimolomyid multituberculate.

 

A cimolomyid multituberculate.

Essonodon[116]

E. browni[116]

A cimolomyid multituberculate.

Meniscoessus[116]

M. conquistus

A cimolomyid multituberculate.

 

M. robustus

  • Montana
  • North Dakota
  • South Dakota[116]

A cimolomyid multituberculate.

 

M. cf. robustus

A cimolomyid multituberculate.

M. sp.

A cimolomyid multituberculate.

?M. sp.[116]

A cimolomyid multituberculate.

Mesodma[116]

M. formosa

  • Montana
  • South Dakota[116]

A neoplagiaulacid multituberculate.

M. cf. formosa

A neoplagiaulacid multituberculate.

M. hensleighi

  • Montana
  • South Dakota[116]

A neoplagiaulacid multituberculate.

M. cf. hensleighi

A neoplagiaulacid multituberculate.

M. thompsoni

  • Montana
  • North Dakota
  • South Dakota[116]

A neoplagiaulacid multituberculate.

M. cf. thompsoni

A neoplagiaulacid multituberculate.

M sp.[116]

A neoplagiaulacid multituberculate.

?M sp.[116]

A neoplagiaulacid multituberculate.

?Neoplagiaulax[116]

?N. burgessi[116]

A neoplagiaulacid multituberculate.

Paracimexomys[116]

P. priscus[116]

A multituberculate of uncertain phylogenetic placement.

Paressonodon[117]

P. nelsoni[117]

A cimolomyid multituberculate.

Stygimys

S. kuszmauli

  • Montana

It was a member of the extinct order Multituberculata.

Metatherians

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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.
Metatherians reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Alphadon[116]

A. marshi

  • Montana
  • North Dakota[116]

An alphadontid. genus of small, primitive mammal that was a member of the Metatheria, a group of mammals that includes modern-day marsupials.

A. cf. marshi

  • Montana
  • South Dakota[116]

An alphadontid.

A. wilsoni

An alphadontid.

A. cf. wilsoni

An alphadontid.

A. sp.[116]

  • North Dakota
  • South Dakota[116]

An alphadontid.

 

Didelphodon[116]

D. padanicus

A stagodontid.

D. vorax

  • Montana
  • North Dakota[116]

A stagodontid. genus of Stagodontidae marsupials from the Late Cretaceous of North America.

 
 

D. cf. vorax

A stagodontid.

 

D. sp.

A stagodontid.

cf. D. sp.[116]

  • North Dakota
  • South Dakota[116]

A stagodontid.

Glasbius[116]

G. twitchelli

A glasbiid.

G. cf. twitchelli[116]

A glasbiid.

Leptalestes[118]

L. cooki

A pediomyid.

L. krejcii[118]

  • Montana
  • South Dakota[116]

A pediomyid.

Nanocuris[117]

N. improvida[117]

A deltatheridiid.

Nortedelphys

N. jasoni (= N. intermedius)[117][119]

  • Montana
  • South Dakota

A herpetotheriid.

Pediomys[116]

P. elegans[116]

  • Montana
  • South Dakota[116]

A pediomyid.

Protalphadon[116]

P. foxi

An alphadontid.

P. lulli[116]

An alphadontid.

Protolambda[118][120]

P. florencae[118][120]

  • Montana
  • North Dakota
  • South Dakota[116]

A pediomyid.

P. hatcheri[118][120]
  • Montana
  • South Dakota[116]

A pediomyid.

P. mcgilli[121]

A pediomyid.

Turgidodon[116]

T. rhaister[116]

An alphadontid.

Eutherians

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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.
Eutherians reported from the Hell Creek Formation
Genus Species State Stratigraphic position Material Notes Images

Altacreodus

A. magnus

  • Montana
  • North Dakota

a possible creodont, formerly a species of Cimolestes[122]

Alostera[116]

A. saskatchewanensis[116]

A eutherian of uncertain phylogenetic placement.

Ambilestes

A. cerberoides

  • Montana

A eutherian of uncertain classification, formally a species of Cimolestes

Batodon[116]

B. tenuis[116]

A cimolestid eutherian.

Cimolestes[116]

C. incisus

A cimolestid eutherian.

C. stirtoni[116]

A cimolestid eutherian.

Gypsonictops[116]

G. hypoconus

  • Montana
  • South Dakota[116]

A gypsonictopsid eutherian.

G. illuminatus

  • Montana
  • North Dakota[116]

A gypsonictopsid eutherian.

G. cf. illuminatus

A gypsonictopsid eutherian.

G. sp.[116]

A gypsonictopsid eutherian.

cf. Paranyctoides[116]

cf. Paranyctoides sp.[116]

A nyctitheriid eutherian.

Protungulatum[118]

P. coombsi[118]

A stem-placental.

 

Purgatorius

P. ceratops

  • Montana

A genus with four species believed to be either stem-placentals or stem-primates.

 

Scollardius

S. propalaeoryctes

  • Montana

A eutherian of uncertain classification, formally a species of Cimolestes

Flora

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Detail of Hell Creek diorama at Milwaukee Public Museum is represented by angiosperm-dominated riparian forest.

The Hell Creek Formation was a low floodplain at the time before the sea retreated, and in the wet ground of the dense woodland, the diversity of angiosperms and conifers were present. An endless diversity of herbaceous flowering plants, ferns and moss grew in the forest understory. On the exposed point bars of large river systems, there were shrubs and vines. The evidence of the forested environment is overwhelmingly supported by petrified wood, rooted gley paleosols,[123] and ubiquitous tree leaves. The presence of the simple and lobed leaves, combined with an extremely high dicot diversity, extinct cycadeoid Nilssoniocladus, Ginkgo, many types of monocots, and several types of conifers is different from any modern plant community. There are numerous types of leaves, seeds, flowers and other structures from Angiosperms, or flowering plants. The Hell Creek Formation of this layer contains over 300 tablets, of which angiosperms are by far the most diverse and dominant flora of the entire population, about 90 percent, followed by about 5% of conifers, 4% of ferns, and others. Compared to today Hell Creek's flora which is prairie, then Hell Creek's flora was hardwood forest mixed with deciduous and evergreen forest. In sharp contrast to the Great Plains today, the presence of many thermophilous taxa such as palm trees and gingers meant the climate was warmer and wetter then.

 
Fossil fruits from the Hell Creek Formation of Spinifructus antiquus of the palm family (Arecaceae), closely related to the genus Astrocaryum.

The plants of the Hell Creek Formation generally represent angiosperm-dominated riparian forests of variable diversity, depending on stratigraphic position and sedimentary environment. There appears to be floral transitions visible on a stratigraphic range from the lower to the upper Hell Creek Formation. For this reason, Kirk Johnson and Leo Hickey divided it into five zones and described them as HCIa, HCIb, HCIIa, HCIIb, and HCIII as a reflection of floral change through time.[124] For example, the HCIa zone is dominated by "Dryophyllum" subfalcatum, Leepierceia preartocarpoides, "Vitis" stantonii, and "Celastrus" taurenensis, and is located 55 to 105 meters below the K-Pg boundary layer. Although the HCIb zone is a very thin layer, about 5 meters of rock, it bears unusually high diversity of herbaceous and shrubby plants, including Urticaceae, Ranunculaceae, Rosaceae, and Cannabaceae.[125][126]

 
Bisonia niemii is one of the common fossil leaf species in the HCIII zone.

There is evidence of transitional floras in the middle of the Hell Creek Formation as shown by HCII and HCIII zones. The HCII flora represents a transitional period where taxa from the lower Hell Creek are replaced by the HCIII flora. The diversity of the HCIII zone is very high, and its composition is more uniform than that of HCII, many of which were rare or absent from the zones below, and some others that used to be common below became rarer in the HCIII zone. These forms include Elatides longifolia, "Dryophyllum" tennessensis, Liriodendrites bradacii, and many members of the Laurales including Bisonia niemii, "Ficus" planicostata, and Marmarthia trivialis, while "Celastrus" taurenensis, Leepierceia preartocarpoides, and many cupressaceous conifers became rarer. This phenomenon suggests that the global temperature was warming during the last 300,000-500,000 years of the Cretaceous period.[125][126][127][128]

Johnson claims that there are no grasses, oaks, maples, beeches, figs, or willows in the Hell Creek Formation. There is no evidence of fern prairie either.[129] However, there was an extremely high angiosperm diversity — common plane trees, "Dryophyllum" subfalcatum, Leepierceia preartocarpoides, and palm trees — along with extinct cycadeoid Nilssoniocladus, Ginkgo, araucariaceous, podocarpaceous, and cupressaceous conifers. This represents the mixed deciduous and evergreen broad-leaved forest as the Hell Creek landscape. The nature of these forests is uncertain because Johnson found that the majority of the angiosperm and conifer genera are now extinct. He also believes that, very roughly 80% of the terrestrial plant taxa died out in what is now Great Plains at the K-Pg boundary. On other hand, there is a great increase in the abundance of fossil fern spores in the two centimeters of rock that directly overlies the impact fallout layer (the famous K-Pg boundary layer). This increase in fern spore abundance is commonly referred as "the fern spike" (meaning that if the abundance of spores as a function of stratigraphic position were plotted out, the graph would show a spike just above the impact fallout layer).

Many of the modern plant affinities in the Hell Creek Formation (e.g., those with the prefix "aff." or with quotes around the genus name) may not in reality belong to these genera; instead they could be entirely different plants that resemble modern genera. Therefore, there is some question regarding whether the modern Ficus or Juglans, as two examples, actually lived in the Late Cretaceous.

Compared to the rich Hell Creek Formation fossil plant localities of the Dakotas, relatively few plant specimens have been collected from Montana. A few taxa were collected at Brownie Butte Montana by Shoemaker, but most plants were collected from North Dakota (Slope County) and from South Dakota. Among the localities, the Mud Buttes, located in Bowman County, North Dakota, is probably the richest megaflora assemblage known and the most diverse leaf quarry from the Hell Creek Formation.[126] "TYPE" after the binomial means that it is represented by a type specimen found in the Yale-Peabody Museum collections. "YPM" is the prefix for the Yale-Peabody Museum specimen number; "DMNH" is for the Denver Museum of Nature & Science; "USNM" is for Smithsonian National Museum of Natural History; and so on. The majority of Hell Creek megafloral specimens are collected at the Denver Museum of Nature & Science.

Overview (from Johnson, 2002)

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302 plant morphotypes based on leaf only, including:

  • 1 bryophyte (mosses and liverworts)
  • 11 ferns
  • 1 sphenopsid
  • 10 conifers
  • 1 ginkgo (uncommon)
  • 278 angiosperms (roughly 92% of all taxa found)

Paleoflora

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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.

Liverworts

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Genus Species Location Stratigraphic position Abundance Notes Images

Marchantia

M. pealii

Only known liverwort in Hell Creek.

 

Ferns

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Genus Species Location Stratigraphic position Abundance Notes Images

Equisetum

E. sp

Rare in the Hell Creek Formation.

 

Polypodiaceae?

indeterminate

Salvinia

S. sp

Floating aquatic plant.

 

Cycadophytes

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Genus Species Location Stratigraphic position Abundance Notes Images
Nilssoniocladus

N. comtula

Unlike N. yukonensis, its leaves are pinnatisect pinnules. Common.

N. yukonensis

The only Hell Creek Formation cycadophyte. A simple leaf. Common.

 

Ginkgoales

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Genus Species Location Stratigraphic position Abundance Notes Images

Ginkgo

G. adiantoides

The only ginkgoalean in the Hell Creek Formation; uncommon

 

Conifers

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Genus Species Location Stratigraphic position Abundance Notes Images

Araucaria?

indeterminate[126]

Casts of Monkey-puzzle leaves.

Cupressinocladus

C. interruptus

Casts of cupressaceae foliage. Uncommon

Dammarites

D. sp

A conifer seed cone. It's likely to belong to Elatides longifolia.

Ditaxocladus

D. catenulata

A cupressaceous conifer closely related to Cupressinocladus. Common.

 

Elatocladus

Elatocladus sp,[126]

A taxodioid leaf morphogenus[126]

Elatides

E. longifolia

Araucaria-like or Cunninghamia-like conifer.
Less common in the lower 2/3 but more common in the upper 1/3 of the Hell Creek Formation.[126]

 
Glyptostrobus

G. europaeus

Uncommon conifer.
 

Metasequoia

M. occidentalis

M. occidentalis seed cones are known from the Hell Creek.

Platyspiroxylon

Platyspiroxylon sp.[126]

A cupressaceaeous wood morphogenus

Podocarpoxylon

Podocarpoxylon sp.[126]

A possibly podocarpaceous wood morphogenus

Sequoiaxylon

Sequoiaxylon sp.[126]

A cupressaceaeous wood morphogenus

Taxodioxylon

Taxodioxylon sp.[126]

A cupressaceaeous wood morphogenus

Taxodium

T. olrikii

Related to today's bald cypress.

Unidentified

Unidentified[126]

Unspecified cheirolepidiacous fossils[126]

Angiosperms

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Genus Species Location Stratigraphic position Abundance Notes Images

"Artocarpus"

"A." lessigiana

Abundant at Brownie Butte, Montana.

 

Annona?

A?. robusta

Abundant at Brownie Butte, Montana.

 

Araliaephyllum

A. polevoi

A lobed leaf. Closely relating to Bisonia. Fairly common.
Averrhoites cf. A. affinis An uncommon taxon with compound leaves.
 
Bisonia

B. niemi

A broad leaf, probably in the Laurales.
A common taxon. Type specimen was found in South Dakota.

 
Browniea B. serrata In the Nyssaceae, closely relating to extant Camptotheca. Less common.
Cannabaceae?

indeterminate

Incertae sedis. It's not likely in the Cannabaceae.[130]

 

Carpites C. ulmiformis Though this fossil fruit is abundant in the Early Paleocene, it's also found in Hell Creek. It may belong to Apiaceae.[131]

"Celastrus"

"C." taurenensis

Incertae sedis. It's common in the lowermost to the middle Hell Creek Formation, but less common in the upper 1/3 Hell Creek Formation.[126]

 

"Cinnamomum"

"C." lineafolia

Included in Ficus affinis by L. Hickey.
Belongs in Rhamnaceae (modern buckthorns and Ceanothus).[citation needed] Some other specimens referred to Cinnamomum sezanensis(?) sp.), a real cinnamon bush. Its affinity is questionable.

 

Cissites cf. C. acerifolia This morphotype was first described from the Cenomanian Dakota Group.[125]

C. insignis

This form represents of the group of Cenomanian leaves from Dakota Formation.[125]

C. lobata

A lobed leaf with half-naked basal lateral veins. Common in the upper 1/3 of the Hell Creek Formation.

C. puilasokensis

A palmately lobed leaf with 5 primary veins. Common in the upper 1/3 of the Hell Creek Formation.
 

Cobbania

C. corrugata

A prehistoric species of water lettuce, previously assigned to the genus Pistia.

 

C. hickeyi Another Cobbania species from pond sediments known as "Licking Leaves."[132]
Cornophyllum C. newberryi The majority of leaves are entire-margined but some may develop a few teeth.[125]
"Cypercites" "C." sp A reed-type plant.
 
Dryophyllum

D. subfalcatum

This taxon is extremely common in the Hell Creek Formation, but is rare in Paleocene sediments.
It is believed to be in Sabiaceae, closely related to the living Meliosma.[125][133]

 

aff. "Dryophyllum" subfalcatum

see above.

"D." tenneseensis

This taxon is similar to D. subfalcatum]] but with extremely high L/W ratio and craspedodromous venation.[125]
Erlingdorfia

E. montana

Johnson, 1996. In the Platanaceae (related to today's Sycamore). A very common taxon.

 

"Ficus"

"F." planicostata

Despite the genus name, it's in Lauraceae.

 

Grewiopsis

G. saportana

Another generic Platanaceae.

 

Harmsia

H. hydrocotyloidea

Incertae sedis. May be related to lotus. An uncommon taxon.

Humulus?

aff. Humulus sp.

May be related to the extant genus Humulus.
 

Hydropteris

H. pinnata

Floating aquatic fern

 
Limnobiophyllum

L. scutatum

Floating aquatic monocot, closely related to Pistia.
 

Liriodendrites

L. bradacii

Johnson, 1996. In the Magnoliidae: a common taxon.

 

"Liriodendron"

"L." laramiense

Unlobed leaf. May be related to today's tulip tree (yellow poplar). An uncommon taxon.
 

Liriodendron

L. sp

Four-lobed leaf. May be related to today's tulip tree (yellow poplar). An uncommon taxon.

 

Leepierceia

L. preartocarpoides

Incertae sedis but possibly in Proteales. Johnson, 1996.

 
Laurophyllum

L. wardiana

Large leaves. Closely related to "Dryophyllum" subfalcatum.
"Magnolia"

M. pulchra

Occurs in southern the Wyoming flora,
Leo Hickey claims it is found further north in Montana and the Dakotas.[citation needed]

 

Marmarthia

M. johnsonii

A new Marmarthia species described in Peppe et al. 2007.[134]

M. pearsonii

Johnson, 1996. In the Lauraceae: a very common taxon.

 

M. trivialis

Johnson, 1996. In the Lauraceae: a very common taxon.

"Myrica"

"Myrica" torreyi

Incertae sedis. Not actually a bayberry.

 

Nelumbo

N. sp

One of the most common aquatic plants in Hell Creek.
 
Nelumbium

N. montanum

An aquatic angiosperm, closely related to lotus. Uncommon.
Nordenskioldia

N. borealis

A fossil fruit likely to belonging to Zizyphoides flabella.[135]

Paranymphaea

P. hastata

Despite the name, it's not related to extant genus Nymphaea.
Palaeoaster

P. porosia

A papaveraceous with fruits and seeds visually similar to Romneya though it's dubious.[citation needed]

 

Penosphyllum

P. cordatum

May be related to Sterculioideae. A common taxon.

Platanites

P. marginata

Johnson, 1996. In the Platanaceae. A common taxon.

 

P. raynoldsii

An uncommon taxon.

Porosia

P. verrucosa

A fossil fruit that may belong to Rutaceae.[136]

Rhamnica

R. cleburnii

Incertae sedis. A Tetracera look-alike leaf.[125]

 

Sabalites

S. sp

Coryphoid palm tree. Very common.

 

Spinifructus

S. antiquus

A fruit seed that may belong to the palm family (Arecaceae), closely related to the genus Astrocaryum.[137]

 

Trapago

T. angulata

A water caltrop look-alike.

 

Trochodendroides

T. arctica

A fossil fruit that may belong to Trochodendroides nebrascensis'.
T. ellipticum A katsura look-alike. An uncommon taxon.
T. genetrix A katsura look-alike. A common taxon.

T. nebrascensis

A very common taxon.
 

"Vitis"

"V." stantonii

Possibly a member of the Platanaceae rather than Vitaceae[126][138]

 

"Ziziphus"

"Z." fibrillosus

A common taxon.

 

Zingiberopsis

Z. attenuata

Related to today's ginger plant. Its closest living relative is the Asian genus Alpinia.
Some Hell Creek Formation specimens show damage from hispine beetles ("leaf beetles" (Wilf et al., 2000)).

 

Z. magnifolia Another Zinigberopsis species, previously assigned to Canna? magnifolia.

Zizyphoides

Z. flabella

An uncommon taxon.

Palynology

edit
Genus Species Location Stratigraphic position Abundance Notes Images
Abietineaepollenites

Abietineaepollenites foveoreticulatus[139]

Montana

A conifer palynomorph

Abietineaepollenites microalatus[139]

Montana

A conifer palynomorph

Abietineaepollenites varius[139]

Montana

A conifer palynomorph

Acanthotriletes

Acanthotriletes levidensis[139]

Montana

Alnipollenites

Alnipollenites verus[139]

Montana

Appendicisporites

Appendicisporites tricornitatus[139]

Montana

Aquilapollenites

Aquilapollenites amplus[139]

Montana

Aquilapollenites attenuatus

Aquilapollenites collaris

Aquilapollenites conatus[139]

Montana

Aquilapollenites delicatus[139]

Montana

Aquilapollenites marmarthensis

Aquilapollenites polaris[139]

Montana

Aquilapollenites pulvinus[139]

Montana

Aquilapollenites pyriformis[139]

Montana

Aquilapollenites quadricretaeus

Aquilapollenites quadrilobus

Aquilapollenites reductus[139]

Montana

Aquilapollenites reticulatus[139]

Montana

Aquilapollenites senonicus

Aquilapollenites turbidus

Aquilapollenites striatus

 

Azolla

Azolla cretacea[139]

Montana

A mosquito fern palynomorph

Balmeisporites

Balmeisporites sp.

Calamospora

Calamospora mesozoica[139]

Montana

Camarozonosporites

Camarozonosporites heskemensis[139]

Montana

Cicatricosisporites

Cicatricosisporites carlylensis[139]

Montana

Cicatricosisporites dorogensis[139]

Montana

Cingulatisporites

Cingulatisporites dakotaensis[139]

Montana

Cingulatisporites scabratus[139]

Montana

Clavatricolpites

Clavatricolpites prolatus[139]

Montana

Concavisporites

Concavisporites rugulatus[139]

Montana

Concavisporites rugulatus[139]

Montana

Converrucosisporites

Converrucosisporites sp.[139]

Montana

Corylus

Corylus granilabratus[139]

Montana

A hazelnut palynomorph

Cupanieidites

Cupanieidites major[139]

Montana

Cyathidites

Cyathidites foveolatus[139]

Montana

Cyathidites minor[139]

Montana

Cycadopites

Cycadopites scabratus[139]

Montana

Deltoidospora

Deltoidospora diaphana[139]

Montana

Dicotetradites

Dicotetradites granulatus[139]

Montana

Ephedripites

Ephedripites ovatus[139]

Montana

Ephedripites undulatus[139]

Montana

Erdtmanipollis

Erdtmanipollis cretaceus[139]

Montana
South Dakota

Gleicheniidites

Gleicheniidites excelsus[139]

Montana

Gleicheniidites senonicus[139]

Montana

Gnetaceaepollenites

Gnetaceaepollenites eocenipites[139]

Montana

Haloragacidites

Haloragacidites quadratus[139]

Montana

Hamulatisporis

Hamulatisporis hamulatis[139]

Montana

Hymenophyllumsporites

Hymenophyllumsporites parvus[139]

Montana

A fern spore palynomorph

Hymenophyllumsporites pseudomaximus[139]

Montana

A fern spore palynomorph

Ilexpollenites

Ilexpollenites compactus

Inaperturopollenites

Inaperturopollenites rugulatus[139]

Montana

Interpollis

Interpollis cf. I. supplingensis

Kurtzipites

Kurtzipites trispissatus[139]

Montana

Kurtzipites trispissatus[139]

Montana

Laevigatosporites

Laevigatosporites anomalus[139]

Montana

Laevigatosporites discordatus[139]

Montana

Laevigatosporites gracilis[139]

Montana

Laevigatosporites ovatus[139]

Montana

Liliacidites

Liliacidites variegatus[139]

Montana

Liliacidites sp.[139]

Montana

Lycopodiumsporites

Lycopodiumsporites austroclavatidites[139]

Montana

Momipites

Momipites circularis[139]

Montana

Momipites parvus[139]

Montana

Monosulcites

Monosulcites carpentieri[139]

Montana

Monosulcites crescentus[139]

Montana

Monosulcites latus[139]

Montana

Monosulcites tectatus[139]

Montana

Monosulcites sp.[139]

Montana

Myrtipites

Myrtipites granulatus[139]

Montana

Myrtipites scabratus[139]

Montana

Nyssapollenites

Nyssapollenites analepticus[139]

Montana

Nyssapollenites pseudocruciatus[139]

Montana

Osmundacidites

Osmundacidites wellmanii[139]

Montana

Pachysandra

Pachysandra cretaceae[139]

Montana
South Dakota

Palmidites

Palmidites maximus[139]

Montana

Peromonolites

Peromonolites granulatus[139]

Montana

Phyllocladidites

Phyllocladidites mawsonii[139]

Montana

Phyllocladidites ruei[139]

Montana

Podocarpidites

Podocarpidites otagoensis[139]

Montana

Polyadopollenites

Polyadopollenites psilatus[139]

Montana

Polycolpites

Polycolpites granulatus[139]

Montana

Polypodiidites

Polypodiidites inangahuensis[139]

Montana

Proteacidites

Proteacidites retusus[139]

Montana

Proteacidites retusus[139]

Montana

Proteacidites thalmannii[139]

Montana

Psilatricolporites

Psilatricolporites prolatus[139]

Montana

Pterocaryapollenites

Pterocaryapollenites stellatus[139]

Montana

Reticuloidosporites

Reticuloidosporites dentatus[139]

Montana

Schizosporis

Schizosporis complexus[139]

Montana

Schizosporis parvus[139]

Montana

Spheripollenites

Spheripollenites subgranulatus[139]

Montana

Spinamonoporites

Spinamonoporites typicus[139]

Montana

Stereisporites

Stereisporites antiquasporites[139]

Montana

Stereisporites psilatus[139]

Montana

Striainaperturites

Striainaperturites ovatus[139]

Montana

Styx

Styx major[139]

Montana

Styx minor[139]

Montana

Taxodiaceaepollenites

Taxodiaceaepollenites hiatus[139]

Montana

Triatriopollenites

Triatriopollenites granilabratus[139]

Montana

Tricolpites

Tricolpites bacustriatus[139]

Montana

Tricolpites delicatulus[139]

Montana

Tricolpites foveolatus[139]

Montana

Tricolpites interangulus

Tricolpites parvistriatus[139]

Montana

Tricolpites psilascabratus[139]

Montana

Tricolpites reticulatus[139]

Montana

Tricolpites striatus[139]

Montana

Tricolpopollenites

Tricolpopollenites clavireticulatus[139]

Montana

Tricolpopollenites megaexactus[139]

Montana

Tricolpopollenites microreticulatus[139]

Montana

Tricolpopollenites microscabratus[139]

Montana

Tricolpopollenites sp1, sp2[139]

Montana

Tricolporopollenites

Tricolporopollenites elongatus[139]

Montana

Tricolporopollenites foveotectatus[139]

Montana

Tricolporopollenites granustriatus[139]

Montana

Tricolporopollenites megaexactus[139]

Montana

Tricolporopollenites prolatus[139]

Montana

Tricolporopollenites striatus[139]

Montana

Triplanosporites

Triplanosporites sinuosus[139]

Montana

Triporopollenites

Triporopollenites rugatus[139]

Montana

Ulmipollenites

Ulmipollenites undulosus[139]

Montana

Ulmipollenites verrucatus[139]

Montana

Ulmoideipites

Ulmoideipites tricostatus[139]

Montana

Wodehouseia

Wodehouseia spinata[139]

Montana

 

Zlivisporis

Zlivisporis blanensis[139]

Montana

See also

edit

References

edit
  1. ^ a b c DePalma, Robert; Cichocki, Frederich; Dierick, Manuel; Feeney, Robert (2010). "Preliminary Notes on the First Recorded Amber Insects from the Hell Creek Formation" (PDF). The Journal of Paleontological Sciences. 10: 1–7. Retrieved 16 November 2020.
  2. ^ a b Nel, André; DePalma, Robert A.; Engel, Michael S. (2010). "A possible hemiphlebiid damselfly in Late Cretaceous amber from South Dakota (Odonata: Zygoptera)". Transactions of the Kansas Academy of Science. 113 (3/4): 231–234. doi:10.1660/062.113.0312. JSTOR 41309615. S2CID 84827761.
  3. ^ Messer, A'ndrea Eluse. "Leaf-mining insects destroyed with the dinosaurs, others quickly appeared". Penn State News. Pennsylvania State University. Retrieved 20 August 2014.
  4. ^ "North Dakota site shows wreckage from same object that killed the dinosaurs". UW News. University of Washington. Retrieved 29 March 2019.
  5. ^ Peterson, Douglas. "The Day the Dinosaurs Died". The New Yorker.
  6. ^ Nel, André (2021). "Maastrichtian representatives of the dragonfly family Aeschnidiidae question the entomofaunal turnover of the early Late Cretaceous". Palaeoentomology. 4 (3). doi:10.11646/palaeoentomology.4.3.5. Retrieved 17 Feb 2024.
  7. ^ a b c d e Graf, Dan; Cummings, Kevin. "The Mussel Project". UWSP Mussel Project. The University of Wisconsin-Stevens Point. Retrieved 7 June 2021.
  8. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw Pearson et al. (2002) p. 154
  9. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj Pearson et al. (2002) pp. 145–167
  10. ^ a b c d e f g h i j k l m n o p q Pearson et al. (2002) p. 155
  11. ^ a b c d e f g h i j k l m n o p q r s Pearson et al. (2002) pp. 156
  12. ^ Listed as "cf. Barbourula sp." in "Class Amphibia," Estes and Berberian, (1970). Page 4.
  13. ^ a b c d e f g h i j k l m n o p "Class Amphibia," in Estes and Berberian, (1970). Page 4.
  14. ^ a b c d e f g h i "Class Amphibia," in Estes and Berberian, (1970). Page 4. All taxa listed occur in Montana, see page 1.
  15. ^ Listed as "Eopelobates? sp." in "Class Amphibia," Estes and Berberian, (1970). Page 4.
  16. ^ a b c d David G. Demar Jr. (2013). "A new fossil salamander (Caudata, Proteidae) from the Upper Cretaceous (Maastrichtian) Hell Creek Formation, Montana, U.S.A". Journal of Vertebrate Paleontology. 33 (3): 588–598. Bibcode:2013JVPal..33..588D. doi:10.1080/02724634.2013.734887. S2CID 128532897.
  17. ^ a b c d e f g h i j k l m n o p "Class Osteichthyes," in Estes and Berberian, (1970). Page 3.
  18. ^ a b c d e f g Hilton, Eric J.; Grande, Lance (2022-10-03). "Late Cretaceous sturgeons (Acipenseridae) from North America, with two new species from the Tanis site in the Hell Creek Formation of North Dakota". Journal of Paleontology. 97 (1): 189–217. doi:10.1017/jpa.2022.81. S2CID 252702937.
  19. ^ a b c d e f g h "Class Osteichthyes," in Estes and Berberian, (1970). Page 3. All taxa listed occur in Montana, see page 1.
  20. ^ a b c d Sato, Hiroki; Murray, Alison M.; Vernygora, Oksana; Currie, Philip J. (4 July 2018). "A rare, articulated sturgeon (Chondrostei: Acipenseriformes) from the Upper Cretaceous of Dinosaur Provincial Park, Alberta, Canada". Journal of Vertebrate Paleontology. 38 (4): (1)-(15). doi:10.1080/02724634.2018.1488137. S2CID 92574712.
  21. ^ a b c d "Class Osteichthyes," in Estes and Berberian, (1970). Page 4.
  22. ^ a b c "Class Osteichthyes," in Estes and Berberian, (1970). Page 4. All taxa listed occur in Montana, see page 1.
  23. ^ Grande, Lance; Bemis, William E. (10 April 1998). "A Comprehensive Phylogenetic Study of Amiid Fishes (Amiidae) Based on Comparative Skeletal Anatomy. an Empirical Search for Interconnected Patterns of Natural History". Journal of Vertebrate Paleontology. 18 (sup1): 1–696. Bibcode:1998JVPal..18S...1G. doi:10.1080/02724634.1998.10011114.
  24. ^ Grande, Lance (2010). "An Empirical Synthetic Pattern Study of Gars (lepisosteiformes) and Closely Related Species, Based Mostly on Skeletal Anatomy. the Resurrection of Holostei". Copeia. 2010 (2A): iii–871. JSTOR 20787269.
  25. ^ a b Listed as "cf. Paralbula casei" in "Class Osteichthyes," Estes and Berberian, (1970). Page 4.
  26. ^ a b c d e f Hilton, Eric J.; During, Melanie A.D.; Grande, Lance; Ahlberg, Per E. (May 2023). "New paddlefishes (Acipenseriformes, Polyodontidae) from the Late Cretaceous Tanis Site of the Hell Creek Formation in North Dakota, USA". Journal of Paleontology. 97 (3): 675–692. Bibcode:2023JPal...97..675H. doi:10.1017/jpa.2023.19. S2CID 258095684.
  27. ^ a b c Grande, Lance; Bemis, William E. (1991). "Osteology and Phylogenetic Relationships of Fossil and Recent Paddlefishes (Polyodontidae) with Comments on the Interrelationships of Acipenseriformes". Memoir. 1. Society of Vertebrate Paleontology: ii–121. doi:10.2307/3889328. JSTOR 3889328.
  28. ^ a b c d e f g h i j k l m n o p q r s Todd D. Cook; Michael G. Newbrey; Donald B. Brinkman; James I. Kirkland (2014). "Euselachians from the freshwater deposits of the Hell Creek Formation of Montana". Through the End of the Cretaceous in the Type Locality of the Hell Creek Formation in Montana and Adjacent Areas. Geological Society of America Special Papers. Vol. 503. pp. 229–246. doi:10.1130/2014.2503(08). ISBN 978-0-8137-2503-1.
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  30. ^ a b "Class Chondrichthyes," in Estes and Berberian, (1970). Page 3.
  31. ^ "Class Chondrichthyes," in Estes and Berberian, (1970). Page 3. All taxa listed occur in Montana, see page 1.
  32. ^ "Stratigraphy and paleontology of the Cretaceous Hell Creek Formation, Stumpf site, Morton County, North Dakota". Proceedings of the North Dakota Academy of Sciences. 48. 1994.
  33. ^ White, Paul; Fastovsky, David; Sheehan, Peter (1998). "Taphonomy and suggested structure of the dinosaurian assemblage of the Hell Creek Formation (Maastrichtian), eastern Montana and western North Dakota". PALAIOS. 13 (1): 41. Bibcode:1998Palai..13...41W. doi:10.2307/3515280. JSTOR 3515280.
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  36. ^ a b Carpenter, Kenneth; Breithaupt, Brent (2 September 1986). "Latest Cretaceous occurrence of nodosaurid ankylosaurs (Dinosauria, Ornithischia) in Western North America and the gradual extinction of the dinosaurs". Journal of Vertebrate Paleontology. 6 (3): 251–257. Bibcode:1986JVPal...6..251C. doi:10.1080/02724634.1986.10011619.
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  64. ^ Vajda, Vivi; Lyson, Tyler R.; Bercovici, Antonie; Doman, Jessman H.; Pearson, Dean A. (November 2013). "A snapshot into the terrestrial ecosystem of an exceptionally well-preserved dinosaur (Hadrosauridae) from the Upper Cretaceous of North Dakota, USA". Cretaceous Research. 46: 114–122. Bibcode:2013CrRes..46..114V. doi:10.1016/j.cretres.2013.08.010.
  65. ^ Rohrer, W. L.; Konizeski, R. (May 1960). "On the Occurrence of Edmontosaurus in the Hell Creek Formation of Montana". 34. 34 (3): 464–466. JSTOR 1300943.
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