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The Judithian was a
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
n
faunal stage In chronostratigraphy, a stage is a succession of rock strata laid down in a single age on the geologic timescale, which usually represents millions of years of deposition. A given stage of rock and the corresponding age of time will by convent ...
lasting from 83.5 to 70.6 million years ago. It overlaps with the
Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...
global stage.


Fauna

Dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...
fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is '' funga''. Flora, fauna, funga and other forms of life are collectively referred to as '' biota''. Zoo ...
s of the Judithian age may represent the peak of dinosaur evolution in North America.
Hadrosaurs Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which incl ...
were universally the dominant
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
of the period and comprised more than half of "a typical assemblage." This was also the period of greatest genetic diversity among large herbivorous dinosaurs. Just in
Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...
and
Southern Alberta Southern Alberta is a region located in the Canadian province of Alberta. In 2004, the region's population was approximately 272,017.genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
of
ceratopsians Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. ...
and ten genera of hadrosaurs.


Paleobiogeography

Thomas M. Lehman has observed that ''
Corythosaurus ''Corythosaurus'' (; ) is a genus of hadrosaurid "duck-billed" dinosaur from the Upper Cretaceous Period (geology), Period, about 77–75.7 million years ago. It lived in what is now North America. Its name means "helmet lizard", derived fr ...
'' and ''
Centrosaurus ''Centrosaurus'' ( ; ) is a genus of herbivorous ceratopsian dinosaur from the Late Cretaceous of Canada. Their remains have been found in the Dinosaur Park Formation, dating from 76.5 to 75.5 million years ago. Discovery and naming The firs ...
'' haven't been discovered outside of southern Alberta even though they are the most abundant Judithian dinosaurs in the region. Large herbivores like the ceratopsians and hadrosaurs living in North America during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
had "remarkably small geographic ranges" despite their large body size and high mobility. This restricted distribution strongly contrasts with modern
mammalian Mammals () are a group of vertebrate animals constituting the class (biology), class Mammalia (), characterized by the presence of mammary glands which in Female#Mammalian female, females produce milk for feeding (nursing) their young, a ...
faunas whose large herbivores' ranges "typical y... span much of a continent." Another example is ''
Pentaceratops ''Pentaceratops'' ("five-horned face") is a genus of herbivorous ceratopsid dinosaur from the late Cretaceous Period of what is now North America. Fossils of this animal were first discovered in 1921, but the genus was named in 1923 when its ty ...
'', the only known Judithian ceratopsian from
New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe , LargestCity = Albuquerque , LargestMetro = Tiguex , OfficialLang = None , Languages = English, Spanish ( New Mexican), Navajo, Ker ...
. Only the rarer species among modern mammal communities would be able to distinguish different latitudinal zones, and some of these
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
are likely too rare to fossilize. This lack of provinciality exists despite the strong temperature gradient. Restrictions in herbivorous dinosaur distribution may be due to foliage preferences, narrow tolerance for variation in climate or other environmental factors. The restrictions on herbivorous dinosaur distribution must have been due to ecological factors rather than physical barriers because carnivorous dinosaurs tended to have wider distributions, especially smaller forms. As of his 2001 paper, restrictions in dinosaur occurrences based on distance from the paleo-shoreline had already been well documented. Vaguely distinguished inland-versus-coastal dinosaurs had been discussed previously in the
scientific literature : ''For a broader class of literature, see Academic publishing.'' Scientific literature comprises scholarly publications that report original empirical and theoretical work in the natural and social sciences. Within an academic field, scient ...
. Terrestrial
sedimentary Sedimentary rocks are types of rock (geology), rock that are formed by the accumulation or deposition of mineral or organic matter, organic particles at Earth#Surface, Earth's surface, followed by cementation (geology), cementation. Sedimentati ...
strata In geology and related fields, a stratum ( : strata) is a layer of rock or sediment characterized by certain lithologic properties or attributes that distinguish it from adjacent layers from which it is separated by visible surfaces known as ei ...
from the Judithian to the
Lancian The Lancian was a North American faunal stage of the Late Cretaceous. It was the final stage of the Cretaceous period in North America, lasting from approximately 70.6 to 66 million years ago. Geology Terrestrial sedimentary strata from the Judit ...
are generally regressive throughout the entire sequence the preserved changes in fossil communities represent not only
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
changes but ecological zones from the submontane habitats to near-sea level coastal habitats. Modern life at high elevations in lower latitudes resembles life at low elevation in higher latitudes. There may be parallels to this phenomenon in Cretaceous
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s, for instance, ''
Pachyrhinosaurus ''Pachyrhinosaurus'' (meaning in Greek "thick-nosed lizard", from ' (), thick; ' (), nose; and (), lizard) is an extinct genus of centrosaurine ceratopsid dinosaur from the Late Cretaceous period of North America. The first examples were discove ...
'' is found in both
Alaska Alaska ( ; russian: Аляска, Alyaska; ale, Alax̂sxax̂; ; ems, Alas'kaaq; Yup'ik: ''Alaskaq''; tli, Anáaski) is a state located in the Western United States on the northwest extremity of North America. A semi-exclave of the U.S., ...
and upland environments in southern Alberta. Northern and Southern animal
biome A biome () is a biogeographical unit consisting of a biological community that has formed in response to the physical environment in which they are found and a shared regional climate. Biomes may span more than one continent. Biome is a broader ...
s approximately correspond respectively with the ''
Aquillapollenites ''Aquillapollenites'' is an extinct morphogenus of Late Cretaceous angiosperm pollen grain. ''Aquillapollenites'' was a very large group containing something like 80 total species, but all of them were typified by their triprojectate structure: th ...
'' and ''
Normapolles ''Normapolles'' is an extinct angiosperm pollen group that arose in the Cenomanian (Late Cretaceous). ''Normapolles'' persisted until the early Oligiocene Epoch (Tertiary Period) but were reduced in diversity by the early Eocene Epoch. ''Normapo ...
'' palynofloral provinces.


Associations

An association between ''Centrosaurus'' and ''Corythosaurus'' is characteristic of southern Alberta. Earlier research had found that lambeosaurines are less common in contemporary Montanan strata and with different centrosaurs as ''
Monoclonius ''Monoclonius'' (meaning "single sprout") is a dubious genus of herbivorous ceratopsian dinosaur found in the Late Cretaceous layers of the Judith River Formation in Montana, United States, and the uppermost rock layers of the Dinosaur Park Forma ...
'' takes the place of ''Centrosaurus''. Inland environments also differed, with the contemporary
Two Medicine Formation The Two Medicine Formation is a geological formation, or rock body, in northwestern Montana and southern Alberta that was deposited between and (million years ago), during Campanian (Late Cretaceous) time. It crops out to the east of the Rocky M ...
preserving an inland fauna characterized by ''
Maiasaura ''Maiasaura'' (from the Greek ''μαῖα'', meaning "good mother" and ''σαύρα'', the feminine form of ''saurus'', meaning "reptile") is a large herbivorous saurolophine hadrosaurid ("duck-billed") dinosaur genus that lived in the area curre ...
'' and the early pachyrhinosaur ''
Einiosaurus ''Einiosaurus'' is a genus of herbivorous centrosaurine ceratopsian dinosaur from the Upper Cretaceous (Campanian stage) of northwestern Montana. The name means 'buffalo lizard', in a combination of Blackfeet Indian ''eini'' and Latinized Ancient ...
''. Farther south was characterized by lower taxonomic diversity in communities where lambeosaurine were less common and centrosaurs were completely lacking. There ''
Kritosaurus ''Kritosaurus'' is an incompletely known genus of hadrosaurid (duck-billed) dinosaur. It lived about 74.5-66 million years ago, in the Late Cretaceous of North America. The name means "separated lizard" (referring to the arrangement of the cheek ...
'', ''
Parasaurolophus ''Parasaurolophus'' (; meaning "near crested lizard" in reference to '' Saurolophus)'' is a genus of herbivorous hadrosaurid ornithopod dinosaur that lived in what is now North America and possibly Asia during the Late Cretaceous Period, abou ...
'' and ''
Pentaceratops ''Pentaceratops'' ("five-horned face") is a genus of herbivorous ceratopsid dinosaur from the late Cretaceous Period of what is now North America. Fossils of this animal were first discovered in 1921, but the genus was named in 1923 when its ty ...
'' are the dominant fauna. The giant eusuchian ''
Deinosuchus ''Deinosuchus'' () is an extinct genus of alligatoroid crocodilian, related to modern alligators and caimans, that lived 82 to 73 million years ago (Ma), during the late Cretaceous period. The name translates as "terrible crocodile" and i ...
'' is also "conspicuous" in the southern biome. Farther south, in Texas, ''Kritosaurus'' predominates. The biomes of the Eastern US may have resembled those of Texas except completely lacking in ceratopsians. ''Parasaurolophus'' and ''Kritosaurus'' are also present in northern latitudes, so evidently exchange between them occurred, but both are uncommon outside of the southern biome.


References

* * *Lehman, T. M., 2001, Late Cretaceous dinosaur provinciality: In: Mesozoic Vertebrate Life, edited by Tanke, D. H., and Carpenter, K., Indiana University Press, pp. 310–328. * Reprinted in {{cite book, last1=Halstead, first1=L.B., title=Dinosaur studies : commemorating the 150th anniversary of Richard Owen's Dinosauria., date=1991, publisher=Gordon and Breach, location=Philadelphia, isbn=9782881248337 Cretaceous geochronology