Hypacrosaurus

Hypacrosaurus (meaning "near the highest lizard" [Greek υπο-, hypo- = less + ακρος, akros, high], because it was almost but not quite as large as Tyrannosaurus)[1][2] was a genus of duckbill dinosaur similar in appearance to Corythosaurus. Like Corythosaurus, it had a tall, hollow rounded crest, although not as large and straight. It is known from the remains of two species that spanned 75 to 67 million years ago,[3] in the Late Cretaceous of Alberta, Canada, and Montana, United States, and is the latest hollow-crested duckbill known from good remains in North America. It was an obscure genus until the discovery in the 1990s of nests, eggs, and hatchlings belonging to H. stebingeri.

Hypacrosaurus
Temporal range: Late Cretaceous, 75–67 Ma
Hypacrosaurus Royal Tyrrell
H. altispinus fossil at the Royal Tyrrell Museum
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Ornithischia
Suborder: Ornithopoda
Family: Hadrosauridae
Tribe: Lambeosaurini
Genus: Hypacrosaurus
Brown, 1913
Type species
Hypacrosaurus altispinus
Brown, 1913
Species
  • H. altispinus
    Brown, 1913
  • H. stebingeri
    Horner & Currie, 1994
Synonyms

Description

Hypracosaurus NT
Life restoration of H. altispinus

Hypacrosaurus is most easily distinguished from other hollow-crested duckbills (lambeosaurines) by its tall neural spines and the form of its crest. The neural spines, which project from the top of the vertebrae, are 5 to 7 times the height of the body of their respective vertebrae in the back,[4] which would have given it a tall back in profile. The skull's hollow crest is like that of Corythosaurus, but is more pointed along its top, not as tall, wider side to side, and has a small bony point at the rear.[4] Unlike other lambeosaurines, the passages for the airways do not form an S-curve in the crest (at least not in H. altispinus).[5] The animal is estimated to have been around 9.1 meters (30 feet) long,[4] and to have weighed up to 4.0 tonnes (4.4 tons).[6] As with most duckbills, its skeleton is otherwise not particularly remarkable, although some pelvic details are distinctive.[7] Like other duckbills, it was a bipedal/quadrupedal herbivore. The two known species, H. altispinus and H. stebingeri, are not differentiated in the typical method, of unique characteristics, as H. stebingeri was described as transitional between the earlier Lambeosaurus and later Hypacrosaurus.[8] Photographs of an adult H. stebingeri skull show an animal that looks very similar to H. altispinus.

Discovery and history

Hypacrosaurus altispinus, head
H. altispinus skull and neck

The type remains of Hypacrosaurus remains were collected in 1910 by Barnum Brown for the American Museum of Natural History.[7] The remains, a partial postcranial skeleton consisting of several vertebrae and a partial pelvis (AMNH 5204), came from along the Red Deer River near Tolman Ferry, Alberta, Canada, from rocks of what is now known as the Horseshoe Canyon Formation (early Maastrichtian, Upper Cretaceous). Brown described these remains, in combination with other postcranial bones, in 1913 as a new genus that he considered to be like Saurolophus.[1] No skull was known at this time, but two skulls were soon discovered and described.[9]

During this period, the remains of small hollow-crested duckbills were described as their own genera and species. The first of these that figure into the history of Hypacrosaurus was Cheneosaurus tolmanensis, based on a skull and assorted limb bones, vertebrae, and pelvic bones from the Horseshoe Canyon Formation.[10] Not long after, Richard Swann Lull and Nelda Wright identified an American Museum of Natural History skeleton (AMNH 5461) from the Two Medicine Formation of Montana as a specimen of Procheneosaurus.[11] These and other taxa were accepted as valid genera until the 1970s, when Peter Dodson showed that it was more likely that the "cheneosaurs" were the juveniles of other established lambeosaurines. Although he was mostly concerned with the earlier, Dinosaur Park Formation genera Corythosaurus and Lambeosaurus, he suggested that Cheneosaurus would turn out to be composed of juvenile individuals of the contemporaneous Hypacrosaurus altispinus.[12] This idea has become accepted,[6] although not formally tested. The Two Medicine Procheneosaurus, meanwhile, was not quite like the other Procheneosaurus specimens studied by Dodson, and for good reason: it was much more like a species that would not be named until 1994, H. stebingeri.[8]

Species

Hypacrosaurus stebingeri holotype
H. stebingeri holotype skull

H. altispinus, the type species, is known from 5 to 10 articulated skulls with some associated skeletal remains, from juvenile to adult individuals found in the Horseshoe Canyon Formation. H. stebingeri is known from an unknown but substantial number of individuals, with an age range of embryos to adults.[6] The hypothesis that H. altispinus and H. stebingeri form a natural group excluding other known hadrosaur species may be incorrect, as noted in Suzuki et al.'s 2004 redescription of Nipponosaurus; their phylogenetic analysis found that Nipposaurus was more closely related to H. altispinus than H. stebingeri was to H. altispinus.[13] This was rejected by Evans and Reisz (2007), though.[14]

The new species Hypacrosaurus stebingeri was named for a variety of remains, including hatchlings with associated eggs and nests, found near the top of the late Campanian (Upper Cretaceous) Two Medicine Formation in Glacier County, Montana, and across the border in Alberta. These represent "the largest collection of baby skeletal material of any single species of hadrosaur known".[8]

Classification

Hypacrosaurus was a lambeosaurine hadrosaurid, and has been recognized as such since the description of its skull.[9] Within the Lambeosaurinae, it is closest to Lambeosaurus and Corythosaurus,[6] with Jack Horner and Phil Currie (1994) suggesting that H. stebingeri is transitional between Lambeosaurus and H. altispinus,[8] and Michael K. Brett-Surman (1989) suggesting that Hypacrosaurus and Corythosaurus are the same genus.[15] These genera, particularly Corythosaurus and Hypacrosaurus, are regarded as the "helmeted" or "hooded" branch of the lambeosaurines, and the clade they form is sometimes informally designated Lambeosaurini. Although Suzuki et al.'s 2004 redescription of Nipponosaurus found a close relationship between Nipponosaurus and Hypacrosaurus stebingeri, indicating that Hypacrosaurus may be paraphyletic,[13] this was rejected in a later, more comprehensive reanalysis of lambeosaurines, which found the two species of Hypacrosaurus to form a clade without Nipponosaurus, with Corythosaurus and Olorotitan being the closest relatives.[14]

Hypacrosaurus Size
Size comparison between the two species and a human

The following cladogram illustrating the relationships of Hypacrosaurus and its close relatives was recovered in a 2012 phylogenetic analysis by Albert Prieto-Márquez, Luis M. Chiappe and Shantanu H. Joshi.[16]

Lambeosaurini

Magnapaulia

Velafrons

Lambeosaurus lambei

Lambeosaurus magnicristatus

Corythosaurus casuarius

Corythosaurus intermedius

"Hypacrosaurus" stebingeri

Hypacrosaurus altispinus

Olorotitan

Paleobiology

Hypacrosaurus stebingeri nest
H. stebingeri nest

As a hadrosaurid, Hypacrosaurus would have been a bipedal/quadrupedal herbivore, eating a variety of plants. Its skull permitted a grinding motion analogous to chewing, and its teeth were continually replacing and packed into dental batteries that contained hundreds of teeth, only a relative handful of which were in use at any time. Plant material would have been cropped by its broad beak, and held in the jaws by a cheek-like organ. Its feeding range would have extended from the ground to about 4 m (13 ft) above.[6]

Nests and growth

Hypacrosaurus Altispinus
Juvenile H. altispinus, American Museum of Natural History.

Hypacrosaurus stebingeri laid roughly spherical eggs of 20 by 18.5 cm (7.9 by 7.3 in), with embryos 60 cm (24 in) long. Hatchlings were around 1.7 m (5.6 ft) long. Studies of lines of growth (i.e. lines of von Ebner) in the teeth of embryonic H. stebingeri suggests plesiomorphically long incubation times, with a minimum incubation time of 171.4 days for H. stebingeri.[17] Young and embryonic individuals had deep skulls with only slight expansion in the bones that would one day form the crest.[8] Growth was faster than that of an alligator and comparable to ratite growth, for several years, based on the amount of bone growth seen between lines of arrested growth (analogous to growth rings in trees).[18] Research by Lisa Cooper and colleagues on H. stebingeri indicates that this animal may have reached reproductive maturity at the age of 2 to 3 years, and reached full size at about 10 to 12 years old. The circumference of the thigh bone at postulated reproductive maturity was about 40% that of its circumference at full size. The postulated growth rate of H. stebingeri outpaces those of tyrannosaurids (predators of hypacrosaurs) such as Albertosaurus and Tyrannosaurus; rapidly growing hypacrosaurs would have had a better chance to reach a size large enough to be of defensive value, and beginning reproduction at an early age would also have been advantageous to a prey animal.[19] Secondary cartilage has been found in the skull of a hatchling specimen of H. stebingeri.[20]

Crest functions

Hypacrosaurus altispinus - AMNH - DSC06304
H. altispinus skull, AMNH

The hollow crest of Hypacrosaurus most likely had social functions, such as a visual signal allowing individuals to identify sex or species, and providing a resonating chamber for making noises.[6] The crest and its associated nasal passages have also figured in the debate about dinosaur endothermy, specifically in discussions about nasal turbinates.

Turbinates are thin bones or cartilages that come in two types, with two functions. Nasal olfactory turbinates are found in all living tetrapods and function in smell. Respiratory turbinates function to prevent water loss through evaporation and are found only in birds and mammals, modern endotherms (warm-blooded animals) who could lose a great deal of water while breathing because they breathe more often than comparably sized ectotherms (cold-blooded animals) to support their higher metabolism.[21] Ruben and others in 1996 concluded that respiratory turbinates were probably not present in Nanotyrannus, Ornithomimus or Hypacrosaurus based on CT scanning, thus there was no evidence that those animals were warm-blooded.[22]

Paleopathology

The discovery of tooth marks in the fibula of a Hypacrosaurus specimen inflicted by a bite from the teeth of a tyrannosaurid indicated that this, and other hadrosaurids were either preyed upon or scavenged by large theropod dinosaurs during the Late Cretaceous period.

Thermoregulation

Examining the oxygen-isotope ratio from the bones from different parts of an extinct animal's body should indicate which thermoregulation mode an animal used during its lifetime. An endothermic (warm-blooded) animal should maintain a very similar body temperature throughout its entire body (which is called homeothermy) and therefore there should be little variation in the oxygen-isotope ratio when measured in different bones. Alternatively, the oxygen-isotope ratio differs considerably when measured throughout the body of an organism with an ectothermic (cold-blooded) physiology.[23] Oxygen-isotope ratios calculated for Hypacrosaurus suggesting that the ratios varied little, indicating that Hypacrosaurus was a homeotherm, and likely was endothermic.[24] This is in contrast to the Ruben et al. (1996) finding that Hypacrosaurus was not warm-blooded, which was based on the absence of nasal turbinates (see Crest functions subsection, above).

Taphonomy

Taphonomy is the study of the processes an organism’s body undergoes after it dies, which includes a study of preservation, the cause of death, and the circumstances of burial. The large, monospecific assemblage of Hypacrosaurus stebingeri in Montana was interpreted as a group of dinosaurs that was killed by a volcanic ashfall.[23] This assemblage is considered autochthonous, meaning that the remains are thought to have been buried on or near the same spot where the individuals died. The variety of ages in this group supports that this was a biocoenosis- an actual life assemblage of animals.[25] The cause of death in a volcanic ashfall is suffocation by the ash and by the gases released from volcanic eruptions. The preservation of this diverse group of dinosaurs provides researchers with a growth series, which is a sequence of growth stages from juvenile to adult.

Paleoecology

Hypacrosaurus skeleton
Hypacrosaurus sp. skeleton exhibited in the National Museum of Nature and Science, Tokyo, Japan.

H. altispinus shared the Horseshoe Canyon Formation with fellow hadrosaurids Edmontosaurus and Saurolophus, hypsilophodont Parksosaurus, ankylosaurid Anodontosaurus, nodosaurid Edmontonia, horned dinosaurs Montanoceratops, Anchiceratops, Arrhinoceratops, and Pachyrhinosaurus, pachycephalosaurid Stegoceras, ostrich-mimics Ornithomimus and Struthiomimus, a variety of poorly known small theropods including troodontids and dromaeosaurids, and the tyrannosaurs Albertosaurus and Daspletosaurus.[26] The dinosaurs from this formation are sometimes known as Edmontonian, after a land mammal age, and are distinct from those in the formations above and below.[27] The Horseshoe Canyon Formation is interpreted as having a significant marine influence, due to an encroaching Western Interior Seaway, the shallow sea that covered the midsection of North America through much of the Cretaceous.[27] H. altispinus may have preferred to stay more landward.[6]

The slightly older Two Medicine Formation, home to H. stebingeri, was also populated by another well-known nesting hadrosaur, Maiasaura, as well as the troodontid Troodon, which is also known from nesting traces. The tyrannosaurid Daspletosaurus, caenagnathid Chirostenotes, dromaeosaurids Bambiraptor and Saurornitholestes, armored dinosaurs Edmontonia, Oohkotokia, and Scolosaurus, hypsilophodont Orodromeus, hadrosaur Prosaurolophus, and horned dinosaurs Achelousaurus, Brachyceratops, Einiosaurus, and Rubeosaurus were also present.[26] This formation was more distant from the Western Interior Seaway, higher and drier, with a more terrestrial influence.[28]

See also

References

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  2. ^ Creisler, Benjamin S. (2007). "Deciphering duckbills". In Carpenter Kenneth (ed.). Horns and Beaks: Ceratopsian and Ornithopod Dinosaurs. Bloomington and Indianapolis: Indiana University Press. pp. 185–210. ISBN 0-253-34817-X.
  3. ^ Arbour, V.M.; Burns, M. E.; Sissons, R. L. (2009). "A redescription of the ankylosaurid dinosaur Dyoplosaurus acutosquameus Parks, 1924 (Ornithischia: Ankylosauria) and a revision of the genus". Journal of Vertebrate Paleontology. 29 (4): 1117–1135. doi:10.1671/039.029.0405.
  4. ^ a b c Lull, Richard Swann; Wright, Nelda E. (1942). Hadrosaurian Dinosaurs of North America. Geological Society of America Special Paper 40. Geological Society of America. pp. 206–208.
  5. ^ Weishampel, David B. (1981). "The nasal cavity of lambeosaurine hadrosaurids (Reptilia:Ornithischia): comparative anatomy and homologies". Journal of Paleontology. 55 (5): 1046–1057.
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  9. ^ a b Gilmore, Charles Whitney (1924). "On the genus Stephanosaurus, with a description of the type specimen of Lambeosaurus lambei, Parks". Canada Department of Mines Geological Survey Bulletin (Geological Series). 38 (43): 29–48.
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  15. ^ Brett-Surman, Michael K. (1989). A revision of the Hadrosauridae (Reptilia:Ornithischia) and their evolution during the Campanian and Maastrichtian. Ph.D. dissertation. Washington, D.C.: Graduate School of Arts and Sciences of The George Washington University. pp. 1–272.
  16. ^ Prieto-Márquez, A.; Chiappe, L. M.; Joshi, S. H. (2012). Dodson, Peter (ed.). "The lambeosaurine dinosaur Magnapaulia laticaudus from the Late Cretaceous of Baja California, Northwestern Mexico". PLoS ONE. 7 (6): e38207. doi:10.1371/journal.pone.0038207. PMC 3373519. PMID 22719869.
  17. ^ Erickson, G.M.; Zelenitsky, D.K.; Kay, D.I.; Norrell, M.A. (2017). "Dinosaur incubation periods directly determined from growth-line counts in embryonic teeth show reptilian-grade development" (PDF). Proceedings of the National Academy of Sciences. doi:10.1073/pnas.1613716114. PMC 5255600.
  18. ^ Cooper, Lisa N.; Horner, John R. (1999). "Growth rate of Hypacrosaurus stebingeri as hypothesized from lines of arrested growth and whole femur circumference". Journal of Vertebrate Paleontology. 19 (3, Suppl.): 35A. doi:10.1080/02724634.1999.10011202.
  19. ^ Cooper, Lisa N.; Lee, Andrew H.; Taper, Mark L.; Horner, John R. (2008). "Relative growth rates of predator and prey dinosaurs reflect effects of predation". Proceedings of the Royal Society B. 275 (1651): 2609–2615. doi:10.1098/rspb.2008.0912. PMC 2605812. PMID 18682367.
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  25. ^ Behrensmeyer, A. K. 1991. Terrestrial vertebrate accumulations. In Allison P. A. and Briggs D. E. G. (Eds), Taphonomy: Releasing the Data Locked in the Fossil Record. New York: Plenum Press.
  26. ^ a b Weishampel, David B.; Barrett, Paul M.; Coria, Rodolfo A.; Le Loueff, Jean; Xu Xing; Zhao Xijin; Sahni, Ashok; Gomani, Elizabeth M.P.; Noto, Christopher N. (2004). "Dinosaur distribution". In Weishampel, David B.; Dodson, Peter; Osmólska Halszka (eds.). The Dinosauria (2nd ed.). Berkeley: University of California Press. pp. 517–606. ISBN 0-520-24209-2.
  27. ^ a b Dodson, Peter (1996). The Horned Dinosaurs: A Natural History. Princeton: Princeton University Press. pp. 14–15. ISBN 0-691-05900-4.
  28. ^ Rogers, Raymond R. (1990). "Taphonomy of three dinosaur bone beds in the Upper Cretaceous Two Medicine Formation of northwestern Montana: evidence for drought-related mortality". PALAIOS. 5 (5): 394–413. doi:10.2307/3514834. JSTOR 3514834.

External links

1913 in paleontology

Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 1913.

Amurosaurus

Amurosaurus (; "Amur lizard") is a genus of lambeosaurine hadrosaurid dinosaur found in the latest Cretaceous period (66 million years ago) of eastern Asia. Like most lambeosaurs, it would have been a primarily bipedal herbivore with a "duckbill" shaped snout and a hollow crest on top of its head, although such a crest has not been found. Fossil bones of adults are rare, but an adult would most likely have been at least 6 metres (20 ft) long. According to Gregory S. Paul, it was about 8 metres (26 ft) long and weighed about 3,000 kilograms (6,600 lb).

Ankylopollexia

Ankylopollexia is an extinct clade of ornithischian dinosaurs that lived from the Late Jurassic to the Late Cretaceous. It is a derived clade of iguanodontian ornithopods and contains the subgroup Styracosterna.

The name stems from the Greek word, “ankylos”, mistakenly taken to mean stiff, fused (in fact the adjective means bent or curved; used of fingers, it can mean hooked), and the Latin word, “pollex”, meaning thumb. Originally described in 1986 by Sereno, this most likely synapomorphic feature of a conical thumb spine defines the clade.First appearing around 156 million years ago, in the Jurassic, Ankylopollexia became an extremely successful and widespread clade during the Cretaceous, and were found around the world. The group died out at the end of the Maastrichtian. Even though they grew to be quite large, comparable to some carnivorous dinosaurs, they were universally herbivorous. Most ankylopollexians were bipedal.

Aralosaurini

Aralosaurini is a tribe of basal lambeosaurine hadrosaurs endemic to Eurasia. It currently contains Aralosaurus (from the Aral sea of Kazakhstan) and Canardia (from Toulouse, Southern France).

Barsboldia

Barsboldia (meaning "of Barsbold", a well-known Mongolian paleontologist) was a genus of large hadrosaurid dinosaur from the early Maastrichtian Nemegt Formation of Ömnogöv', Mongolia. It is known from a partial vertebral column, partial pelvis, and some ribs.

Blasisaurus

Blasisaurus is a genus of lambeosaurine hadrosaurid dinosaur from the Late Cretaceous. It is known from a partial skull and skeleton found in late Maastrichtian-age rocks of Spain. The type species is Blasisaurus canudoi, described in 2010 by Penélope Cruzado-Caballero, Xabier Pereda-Suberbiola and José Ignacio Ruiz-Omeñaca, a group of researchers from Spain.

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 spans the time from 83.6 (± 0.7) to 72.1 (± 0.6) million years ago. It is preceded by the Santonian and it is followed by the Maastrichtian.The Campanian was an age when a worldwide sea level rise covered many coastal areas. The morphology of some of these areas has been preserved: it is an unconformity beneath a cover of marine sedimentary rocks.

Canardia

Canardia is an extinct genus of aralosaurin lambeosaurine dinosaur known from the Late Cretaceous Marnes d’Auzas Formation (late Maastrichtian stage) of Toulouse, Haute-Garonne Department, southern France. The type species Canardia garonnensis was first described and named by Albert Prieto-Márquez, Fabio M. Dalla Vecchia, Rodrigo Gaete and Àngel Galobart in 2013.

Corythosaurus

Corythosaurus is a genus of hadrosaurid "duck-billed" dinosaur from the Upper Cretaceous Period, about 77–75.7 million years ago. It lived in what is now North America. Its name means "helmet lizard", derived from Greek κόρυς. It was named and described in 1914 by Barnum Brown. Corythosaurus is now thought to be a lambeosaurine, related to Nipponosaurus, Velafrons, Hypacrosaurus, and Olorotitan. Corythosaurus has an estimated length of 9 metres (30 ft), and has a skull, including the crest, that is 70.8 centimetres (27.9 in) tall.

Corythosaurus is known from many complete specimens, including the nearly complete holotype found by Brown in 1911. The holotype skeleton is only missing the last section of the tail, and part of the forelimbs, but was preserved with impressions of polygonal scales. Corythosaurus is known from many skulls with tall crests. The crests resemble the crests of the cassowary and a Corinthian helmet. The most likely function of the crest is thought to be vocalization. As in a trombone, sound waves would travel through many chambers in the crest, and then get amplified when Corythosaurus exhaled. A Corythosaurus specimen has been preserved with its last meal in its chest cavity. Inside the cavity were remains of conifer needles, seeds, twigs, and fruits: Corythosaurus probably fed on all of these.

The two species of Corythosaurus are both present in slightly different levels of the Dinosaur Park Formation. Both still co-existed with theropods and other ornithischians, like Daspletosaurus, Brachylophosaurus, Parasaurolophus, Scolosaurus, and Chasmosaurus.

Devil's Coulee Dinosaur Heritage Museum

Devil's Coulee Dinosaur Heritage Museum, located in Warner, Alberta, Canada, is a key historic site in southern Alberta. In 1997, ten fossilized dinosaur eggs, believed to have come from a Hadrosaur, specifically a Hypacrosaurus were found at Devil's Coulee site. These were not the first fossils to be found in what was often called the Fossil Coulee region of the province and as a result the town of Warner established the museum to help interpret the story.

Horseshoe Canyon Formation

The Horseshoe Canyon Formation is a stratigraphic unit of the Western Canada Sedimentary Basin in southwestern Alberta. It takes its name from Horseshoe Canyon, an area of badlands near Drumheller.

The Horseshoe Canyon Formation is part of the Edmonton Group and is up to 230 metres (750 ft) thick. It is of Late Cretaceous age, Campanian to early Maastrichtian stage (Edmontonian Land-Mammal Age), and is composed of mudstone, sandstone, carbonaceous shales, and coal seams. A variety of depositional environments are represented in the succession, including floodplains, estuarine channels, and coal swamps, which have yielded a diversity of fossil material. Tidally-influenced estuarine point bar deposits are easily recognizable as Inclined Heterolithic Stratification (IHS). Brackish-water trace fossil assemblages occur within these bar deposits and demonstrate periodic incursion of marine waters into the estuaries.

The Horseshoe Canyon Formation crops out extensively in the area around Drumheller, as well as farther north along the Red Deer River near Trochu and along the North Saskatchewan River in Edmonton. It is overlain by the Battle, Whitemud, and Scollard formations. The Drumheller Coal Zone, located in the lower part of the Horseshoe Canyon Formation, was mined for sub-bituminous coal in the Drumheller area from 1911 to 1979, and the Atlas Coal Mine in Drumheller has been preserved as a National Historic Site. In more recent times, the Horseshoe Canyon Formation has become a major target for coalbed methane (CBM) production.

Dinosaurs found in the Horseshoe Canyon Formation include Albertavenator, Albertosaurus, Anchiceratops, Anodontosaurus, Arrhinoceratops, Atrociraptor, Epichirostenotes, Edmontonia, Edmontosaurus, Hypacrosaurus, Ornithomimus, Pachyrhinosaurus, Parksosaurus, Saurolophus, and Struthiomimus. Other finds have included mammals such as Didelphodon coyi, non-dinosaur reptiles, amphibians, fish, marine and terrestrial invertebrates and plant fossils. Reptiles such as turtles and crocodilians are rare in the Horseshoe Canyon Formation, and this was thought to reflect the relatively cool climate which prevailed at the time. A study by Quinney et al. (2013) however, showed that the decline in turtle diversity, which was previously attributed to climate, coincided instead with changes in soil drainage conditions, and was limited by aridity, landscape instability, and migratory barriers.

Lambeosaurinae

Lambeosaurinae is a group of crested hadrosaurid dinosaurs.

Lambeosaurini

Lambeosaurini, previously known as Corythosaurini, is one of four tribes of hadrosaurid ornithopods from the family Lambeosaurinae. It is defined as all lambeosaurines closer to Lambeosaurus lambei than to Parasaurolophus walkeri, Tsintaosaurus spinorhinus, or Aralosaurus tuberiferus, which define the other three tribes. Members of this tribe possess a distinctive protruding cranial crest. Lambeosaurins walked the earth for a period of around 12 million years in the Late Cretaceous, though they were confined to regions of modern day North America and Asia.

Lambeosaurus

Lambeosaurus ( LAM-bee-o-SAWR-əs; meaning "Lambe's lizard") is a genus of hadrosaurid dinosaur that lived about 75 million years ago, in the Late Cretaceous period (Campanian) of North America. This bipedal/quadrupedal, herbivorous dinosaur is known for its distinctive hollow cranial crest, which in the best-known species resembled a hatchet. Several possible species have been named, from Canada, the United States, and Mexico, but only the two Canadian species are currently recognized as valid.

Material relevant to the genus was first named by Lawrence Lambe in 1902. Over twenty years later, the modern name was coined in 1923 by William Parks, in honour of Lambe, based on better preserved specimens. The genus has a complicated taxonomic history, in part because small-bodied crested hadrosaurids now recognized as juveniles were once thought to belong to their own genera and species. Currently, the various skulls assigned to the type species L. lambei are interpreted as showing age differences and sexual dimorphism. Lambeosaurus was closely related to the better known Corythosaurus, which is found in slightly older rocks, as well as the less well-known genera Hypacrosaurus and Olorotitan. All had unusual crests, which are now generally assumed to have served social functions like noisemaking and recognition.

Magnapaulia

Magnapaulia is a genus of herbivorous lambeosaurine hadrosaurid dinosaurs known from the Latest Cretaceous Baja California, of northwestern Mexico. It contains a single species, Magnapaulia laticaudus. Magnapaulia was first described in 1981 as a possible species of Lambeosaurus by William J. Morris, and was given its own genus in 2012 by Prieto-Márquez and colleagues.

Nipponosaurus

Nipponosaurus (meaning "Japanese lizard") is a lambeosaurine hadrosaur from Sinegorsk, Sakhalin Island in Russia, part of Japan at the time of naming. The type and only species is N. sachalinensis, known only from a single juvenile specimen discovered in 1934 and named in 1936, by Takumi Nagao, with further material of the same individual found in 1937. Since then, the taxon has been largely ignored, and its validity has been doubted, with synonymy with other Asian hadrosaurs or status as a nomen dubium being suggested. Redescriptions from 2004 and 2017, however, have supported recognition as a distinct species. Dating the only specimen has been difficult, but based on associated mollusc taxa, the species likely lived sometime in the upper Santonian or lower Campanian, around 80 million years ago.

Olorotitan

Olorotitan was a genus of lambeosaurine duckbilled dinosaur from the middle or latest Maastrichtian-age Late Cretaceous, whose remains were found in the Udurchukan Formation beds of Kundur, Amur Region, Far Eastern Russia. The type, and only species is Olorotitan arharensis.

Tsintaosaurus

Tsintaosaurus (; meaning "Qingdao lizard", after the old transliteration "Tsingtao") is a genus of hadrosaurid dinosaur from China. It was about 8.3 metres (27 ft) long and weighed 2.5 tonnes. The type species is Tsintaosaurus spinorhinus, first described by Chinese paleontologist C. C. Young in 1958.

A hadrosaur, Tsintaosaurus had a characteristic 'duck bill' snout and a battery of powerful teeth which it used to chew vegetation. It usually walked on all fours, but could rear up on its hind legs to scout for predators and flee when it spotted one. Like other hadrosaurs, Tsintaosaurus probably lived in herds.

Velafrons

Velafrons (meaning "sailed forehead") is a genus of lambeosaurine hadrosaurid dinosaur from the Late Cretaceous of Mexico. It is known from a mostly complete skull and partial skeleton of a juvenile individual, with a bony crest on the forehead. Its fossils were found in the late Campanian-age Cerro del Pueblo Formation (about 72 million years old), near Rincon Colorado, Coahuila, Mexico. The type specimen is CPC-59, and the type species is V. coahuilensis.

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