Lancian
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The Lancian was a North American
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 conventi ...
of 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 ...
. 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 Judithian to the Lancian are generally regressive throughout the entire sequence, so the preserved changes in fossil communities represent not only phylogenetic changes but ecological zones from the submontane habitats to near-sea level coastal habitats.


Paleobiogeography

By the Lancian, the crested hadrosaurs are no longer the dominant inhabitant of any province of western North America; the only remaining species was ''
Hypacrosaurus ''Hypacrosaurus'' (meaning "near the highest lizard" reek υπο-, ''hypo-'' = less + ακρος, ''akros'', high because it was almost but not quite as large as ''Tyrannosaurus'') was a genus of duckbill dinosaur similar in appearance to ''Co ...
''. Lehman records three surviving chasmosaurs, ''
Triceratops ''Triceratops'' ( ; ) is a genus of herbivore, herbivorous Chasmosaurinae, chasmosaurine Ceratopsidae, ceratopsid dinosaur that first appeared during the late Maastrichtian stage of the Late Cretaceous Period (geology), period, about 68 m ...
,'' ''
Torosaurus ''Torosaurus'' ("perforated lizard", in reference to the large openings in its frill) is a genus of herbivorous ceratopsid dinosaur that lived during the late Maastrichtian stage of the Cretaceous period, between 68 and 66 million years ago, tho ...
'' and ''
Nedoceratops ''Nedoceratops'' (meaning "insufficient horned face") is a controversial genus of ceratopsid dinosaur from the Late Cretaceous period Lance Formation of North America. It is known only from a single skull discovered in Wyoming. Its status is the ...
'', with the possibility of the recently discovered ''
Ojoceratops ''Ojoceratops'' ( meaning "Ojo Alamo horned face") is a genus of ceratopsian dinosaur which lived in what is now New Mexico, United States. ''Ojoceratops'' fossils have been recovered from strata of the Ojo Alamo Formation (Naashoibito Member), d ...
,
Regaliceratops ''Regaliceratops'' (meaning "Royal horned face") is a monospecific genus of chasmosaurinae, chasmosaurine ceratopsidae, ceratopsid dinosaur from Alberta, Canada that lived during the Late Cretaceous (middle Maastrichtian stage, 68.5 to 67.5 Ma) i ...
and'' ''
Bravoceratops ''Bravoceratops'' is a genus of large chasmosaurine ceratopsid dinosaur that lived approximately 70 million years ago, and is known from the Late Cretaceous Javelina Formation in what is now Texas, United States. Discovery and naming ''Bravocer ...
''. It has recently been suggested that ''Triceratops'' and ''Torosaurus'' may be synonymous, though this is still up for debate.
Saurolophine Saurolophinae is a subfamily (biology), subfamily of hadrosaurid dinosaurs. It has since the mid-20th century generally been called the Hadrosaurinae, a group of largely non-crested hadrosaurs related to the crested sub-family Lambeosaurinae. How ...
dinosaurs like ''
Edmontosaurus ''Edmontosaurus'' ( ) (meaning "lizard from Edmonton") is a genus of hadrosaurid (duck-billed) dinosaur. It contains two known species: ''Edmontosaurus regalis'' and ''Edmontosaurus annectens''. Fossils of ''E. regalis'' have been found in rocks ...
'', ''
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 ...
'', ''
Saurolophus ''Saurolophus'' (; meaning "lizard crest") is a genus of large hadrosaurid dinosaur from the Late Cretaceous period of Asia and North America, that lived in what is now the Horseshoe Canyon and Nemegt formations about 70 million to 68 million ...
'' as well as the recently discovered ''
Augustynolophus ''Augustynolophus'' is an extinct genus of herbivorous saurolophine hadrosaur dinosaur which was discovered in the Moreno Formation in California, dating to the late Maastrichtian age, making it one of the last dinosaurs known from the fossil re ...
'' were some of the known surviving hadrosaurs. All lacked the elaborate ornamentation of their predecessors, the lambeosaurs. The ankylosaurs had been reduced to down to a handful of species, with ankylosaurs like ''
Anodontosaurus ''Anodontosaurus'' is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the entire span of the Late Cretaceous Horseshoe Canyon Formation (mid Late Campanian to "middle" Maastrichtian stage, about 72 ...
'' and ''
Ankylosaurus ''Ankylosaurus'' is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of the n ...
'', as well as nodosaurs like ''
Denversaurus ''Denversaurus'' (meaning "Denver lizard") is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous (late Maastrichtian) of western North America. Although at one point treated as a junior synonym of ''Edmontonia'' by some taxon ...
'', ''
Edmontonia ''Edmontonia'' is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous Period. It is part of the Nodosauridae, a family within Ankylosauria. It is named after the Edmonton Formation (now the Horseshoe Canyon Formation in Canada) ...
'' and ''
Glyptodontopelta ''Glyptodontopelta'' (meaning "''Glyptodon'' shield") is a monospecific genus of nodosaurid dinosaur from New Mexico that lived during the Late Cretaceous (lower to upper Maastrichtian, 69 to 66 Ma) in what is now the Naashoibito member of the O ...
'' being the only known survivors. In the south the transition to the Lancian is even more dramatic, which Lehman describes as "the abrupt reemergence of a fauna with a superficially "Jurassic" aspect." These faunas are dominated by ''
Alamosaurus ''Alamosaurus'' (; meaning "Ojo Alamo lizard") is a genus of opisthocoelicaudiine titanosaurian sauropod dinosaurs, containing a single known species, ''Alamosaurus sanjuanensis'', from the late Cretaceous Period of what is now southern North Am ...
'' and feature abundant ''
Quetzalcoatlus ''Quetzalcoatlus'' is a genus of pterosaur known from the Late Cretaceous period of North America (Maastrichtian stage); its members were among the largest known flying animals of all time. ''Quetzalcoatlus'' is a member of the Azhdarchidae, ...
, Bravoceratops'' and ''Ojoceratops'' in Texas. The extreme changes occurring in the make-up of herbivore communities during the faunal turnover suggests that a change in the ecosystems' flora was "the most immediate cause...though perhaps not the ultimate one." The rapid expansion of land and drying of inland climate accompanying a drop in sea level could explain some of the environmental changes occurring Late Cretaceous western North America. The wetland habitat enjoyed by many dinosaurs would have shrunk and fragmented. Since many species had very limited geographic ranges its plausible that some of the fragments would be smaller than the area needed to support the species. However, there's no direct evidence for the shrinking of wetland environments. Lehman contends that the actual area of coastal lowlands within 150m of the shoreline must have actually increased significantly. Further, dinosaurs that inhabited inland or arid environments were among the most prevalent in the Lancian. The ''Alamosaurus''-''Quetzalcoatlus'' association probably represent semi-arid inland plains. In previous research Jack Horner speculated that a rise in sea level during the
Bearpaw Transgression Bearpaw or Bear Paw may refer to: *Bearpaw (brand), a brand of footwear, including sheepskin boots, slippers and casual shoes *Bearpaw Formation, a rock formation in North America * Bear-paw poppies, the genus of the poppy family Papaveraceae * Mik ...
created selective pressure as coastal lowlands were swallowed up the sea, resulting in
anagenesis Anagenesis is the gradual evolution of a species that continues to exist as an interbreeding population. This contrasts with cladogenesis, which occurs when there is branching or splitting, leading to two or more lineages and resulting in separate ...
. If the geographic range of some dinosaur species were truly as limited as the fossil record suggests, then a rapid rise and in sea level could cause intense pressure even the event was local. Additionally, a rapid drop in sea level could allow for "rapid colonization by a few dinosaur generalists." The appearance or reappearance of basal
neoceratopsian 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. ...
s could be explained by immigration from Asia. Dinosaurs like ''
Nodocephalosaurus ''Nodocephalosaurus'' (meaning "knob headed lizard") is a monospecific genus of ankylosaurid dinosaur from New Mexico that lived during the Late Cretaceous (late Campanian to early Maastrichtian stage, 73.49 to 73.04 Ma) in what is now the De-na- ...
'' resembled Asian forms, and some like ''Saurolophus'' co-occurred in Asia as well as North America. Potential Asian immigrants were especially common in upland environments. The appearance of ''Alamosaurus'' may have represented an immigration event from South America. Some taxa may have co-occurred on both continents, including ''
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 ...
'' and ''
Avisaurus ''Avisaurus'' (meaning "bird lizard") is a genus of enantiornithine bird from the Late Cretaceous of North America. Discovery ''Avisaurus archibaldi'' was discovered in the Late Cretaceous Hell Creek Formation of North America (Maastrichtian, fr ...
''. ''Alamosaurus'' appears and achieves dominance in its environment very abruptly. Some scientists speculated that ''Alamosaurus'' was an immigrant from Asia. Inhabitants of upland environment are more likely to be endemic than coastal species, and tend to have less of an ability to cross bodies of water. Further, early Cretaceous titanosaurs were already known, so North American potential ancestors for ''Alamosaurus ''already existed. ''Quetzalcoatlus'' also had precursors in North America and its apparent range expansion may represent the expansion of its preferred habitat rather than an immigration event.
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
deposits in North America reveal that basal neoceratopsians were already present on the continent before their apparent reemergence in the Lancian, so an immigration event from Asia is unnecessary to explain their appearance. The major potential immigrants represent archaic forms that probably wouldn't have directly competed with the disappearing forms anyway. Lehman described the evidence for immigration as a driving force in Lancian dinosaur faunal turnovers as "not particularly compelling." The faunal turnover may be explained by the descent of more primitive forms existing in upland refugia characterized by
conifer Conifers are a group of conifer cone, cone-bearing Spermatophyte, seed plants, a subset of gymnosperms. Scientifically, they make up the phylum, division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single ...
-dominated flora into areas that were formerly coastal lowlands as the seas retreated and conditions became more arid. The decline of mammal diversity in Western North America from the Miocene to the present primarily effected large herbivores and occurred over roughly the same length of time as the Late Cretaceous changes, and so may be parallel. They have many commonalities, including the replacement of diverse with single species environments (
caribou Reindeer (in North American English, known as caribou if wild and ''reindeer'' if domesticated) are deer in the genus ''Rangifer''. For the last few decades, reindeer were assigned to one species, ''Rangifer tarandus'', with about 10 subspe ...
in the north,
bison Bison are large bovines in the genus ''Bison'' (Greek: "wild ox" (bison)) within the tribe Bovini. Two extant and numerous extinct species are recognised. Of the two surviving species, the American bison, ''B. bison'', found only in North Ame ...
to the south). The most spectacular and specialized forms became extinct. The mammalian turnover was preceded by an episode of immigration, and was associated with the rapid expansion of terrestrial habitat due to melting
glaciers A glacier (; ) is a persistent body of dense ice that is constantly moving under its own weight. A glacier forms where the accumulation of snow exceeds its ablation over many years, often centuries. It acquires distinguishing features, such as ...
. In the mammalian turnover, the newly emerging dominant fauna were clearly Old World immigrants, the
cervids Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...
and
bovids The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, the ...
. In the southern biome, by Lancian time sauropods had replaced both hadrosaurs and ceratopsians. In the north, both were still present although hadrosaurs were demoted to a "subordinate" role in dinosaur ecosystems. ''Edmontosaurus'' was the dominant northern hadrosaurid. At the end of the Cretaceous, most ecosystems were dominated by a single herbivore. The northern biome was dominated by ''Triceratops'' and the southern biome by ''Alamosaurus''. This faunal turnover coincides with the Laramide orogeny and the uplift of the central Rockies. Strata exhibit changes in lithology and the direction of paleocurrents, and a severe drop in relative sea level. At the very least, Lehman argues, the altitudinal life zones would shift, and a change in the distribution of vegetation utilized by herbivorous dinosaurs would have probably resulted. By the end of the Judithian, North America had 7.7 million km2 of land area, but by the end of the Lancian it had reached 17.9 million km2, nearly the modern value of 22.5 million km2.


Footnotes


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 *