Campanian

The Campanian is, in the ICS' geologic timescale, the fifth of six ages of the Late Cretaceous epoch (or, in chronostratigraphy: the fifth of six stages in the Upper Cretaceous series). The Campanian spans the time from 83.6 ± 0.7 Ma to 72.1 ± 0.6 Ma (million years ago). It is preceded by the Santonian and it is followed by the Maastrichtian.^[1]

Stratigraphic definition

The Campanian was introduced in scientific literature by Henri Coquand in 1857. It is named after the French village of Champagne in the département Charente-Maritime. The original type locality was an outcrop near the village of Aubeterre-sur-Dronne in the same region. Due to changes of the stratigraphic definitions, this section is now part of the Maastrichtian stage.

The base of the Campanian stage is laid at the extinction of crinoid species Marsupites testudinarius. A GSSP had not yet been ratified in 2009. One possible candidate is in a section near a dam at Waxahachie, Texas.

The top of the Campanian is defined as the place in the stratigraphic column were the ammonite Pachydiscus neubergicus first appears.

Subdivision

The Campanian is sometimes subdivided into Lower, Middle and Upper subages. In the Tethys domain, the Campanian encompasses six ammonite biozones. They are, from young to old:

• zone of Nostoceras hyatti
• zone of Didymoceras chayennense
• zone of Bostrychoceras polyplocum
• zone of Hoplitoplacenticeras marroti/Hoplitoplacenticeras vari
• zone of Delawarella delawarensis
• zone of Placenticeras bidorsatum

Paleontology

During the Campanian age, a radiation among dinosaur species occurred. In North America, for example, the number of known dinosaur genera rises from 4 at the base of the Campanian to 48 in the upper part. This development is sometimes referred to as the "Campanian Explosion". However, it is not yet clear if the event is artificial, i.e. the low number of genera in the lower Campanian can be caused by a lower preservation chance for fossils in deposits of that age. The generally warm climates and large continental area covered in shallow sea during the Campanian probably favoured the dinosaurs. In the following Maastrichtian stage, the number of North American dinosaur genera found is 30% less than in the upper Campanian.^[2]

Animals that lived in the Campanian include:

†Ankylosaurs

Ankylosaurs of the Campanian
Taxa	Presence	Location	Description	Images
Aletopelta		Point Loma Formation, California, USA	A medium-sized ankylosaurid, estimated to be around 6 m (20 ft) long.	Edmontonia Euoplocephalus Pinacosaurus
Antarctopelta		Santa Marta Formation, James Ross Island, Antarctica	A stocky ankylosaur protected by armor plates embedded in the skin. Although a complete skeleton has not been found, the species is estimated to have reached a maximum length of 4 meters (13 ft). Displays characteristics of both ankylosaurids and nodosaurids.
Edmontonia	Campanian to Maastrichtian	Horseshoe Canyon Formation, Alberta, Canada	A bulky nodosaurid at roughly 6.6 m (22 ft) long. It had small, ridged bony plates on its back and many sharp spikes along its body sides. The four largest spikes jutted out from the shoulders on each side, two of which were split into subspines in some specimens. Its skull had a pear-like shape when viewed from above.
Euoplocephalus
Niobrarasaurus
Nodocephalosaurus
Panoplosaurus		Judith River Formation, Alberta, Canada; Montana, USA	A 5.5–7 m long nodosaurid.
Pinacosaurus
Saichania
Shanxia
Struthiosaurus
Tarchia
Tianzhenosaurus

Birds (avian theropods)

Birds of the Campanian
Taxa	Presence	Location	Description	Images
Hesperornis				Hesperornis
Neogaeornis wetzeli			A marine bird from Chile. It had the midfeet of a foot-propelled diving bird, but its relationships are enigmatic. The only known species is from the Campanian-Maastrichtian boundary.
Ichthyornis

Bony Fish

Bony fish of the Campanian
Taxa	Presence	Location	Description	Images
Xiphactinus				Xiphactinus

Cartilaginous Fish

Cartilaginous fish of the Campanian
Taxa	Presence	Location	Description	Images
Chlamydoselachus				Chlamydoselachus
†Schizorhiza

†Ceratopsians

Ceratopsians of the Campanian
Taxa	Presence	Location	Description	Images
Achelousaurus Achelousaurus horneri	74.2 million years ago			Achelousaurus Agujaceratops Albertaceratops Anchiceratops Avaceratops Bagaceratops Brachyceratops Centrosaurus Cerasinops Chasmosaurus Coahuilaceratops Diabloceratops Einiosaurus Graciliceratops Kosmoceratops Medusaceratops Pentaceratops Prenoceratops Protoceratops Rubeosaurus Spinops Styracosaurus Titanoceratops Udanoceratops Utahceratops Vagaceratops
Agujaceratops Agujaceratops mariscalensis	77 million years ago
Albertaceratops Albertaceratops nesmoi
Anchiceratops Anchiceratops longirostris
Avaceratops Avaceratops lammersi
Bagaceratops Bagaceratops rozhdestvenskyi
Bainoceratops Bainoceratops efremovi
Brachyceratops Brachyceratops montanensis
Breviceratops Breviceratops kozlowskii
Centrosaurus Centrosaurus apertus
Cerasinops Cerasinops hodgskissi
Ceratops Ceratops montanus
Chasmosaurus Chasmosaurus russelli Chasmosaurus belli
Coahuilaceratops Coahuilaceratops magnacuerna
Coronosaurus Coronosaurus brinkmani
Diabloceratops Diabloceratops eatoni
?Dysganus Dysganus encaustus Dysganus bicarinatus Dysganus peiganus
Einiosaurus Einiosaurus procurvicornis
Graciliceratops Graciliceratops mongoliensis
Gryphoceratops Gryphoceratops morrisoni
Judiceratops Judiceratops tigris
Kosmoceratops Kosmoceratops richardsoni
Lamaceratops Lamaceratops tereschenkoi
Magnirostris Magnirostris dodsoni
Medusaceratops Medusaceratops lokii
Monoclonius Monoclonius crassus
Nasutoceratops Nasutoceratops titusi
Pachyrhinosaurus Pachyrhinosaurus canadensis Pachyrhinosaurus lakustai Pachyrhinosaurus perotorum
Pentaceratops Pentaceratops sternbergii
Prenoceratops Prenoceratops pieganensis
Platyceratops Platyceratops tatarinovi
Protoceratops Protoceratops andrewsi Protoceratops hellenikorhinus
Rubeosaurus Rubeosaurus ovatus
Spinops Spinops sternbergorum
Styracosaurus Styracosaurus albertensis
Titanoceratops Titanoceratops ouranos
Udanoceratops Udanoceratops tschizhovi
Unescoceratops Unescoceratops koppelhusae
Utahceratops Utahceratops gettyi
Vagaceratops Vagaceratops irvinensis
Xenoceratops Xenoceratops foremostensis

Crocodylomorphs

Mammals

†Ornithopods

Ornithopods of the Campanian
Taxa	Presence	Location	Description	Images
Aralosaurus	85.8 mya	Asia	Aralosaurus was about the size of an elephant. Although very little is known about Aralosaurus (only one near complete skull has been found); it was identified by a beak with nearly 1,000 small teeth in 30 rows. These teeth were used for breaking up plant matter by chewing, a feature common in herbivorous dinosaurs, but unusual for reptiles.The back of an Aralosaurus skull was wide, a feature suggestive of large jaw muscles used to power its chewing apparatus.	Edmontosaurus Gasparinisaura Hypacrosaurus Maiasaura Mochlodon Nipponosaurus Prosaurolophus Shantungosaurus Velafrons
Brachylophosaurus	76.5 mya	Montana, USA; Alberta, Canada	Brachylophosaurus was a typical hadrosaur which reached an adult length of 9 meters (30 ft).
Corythosaurus	77-76.5 mya	Alberta, Canada	Corythosaurus weighed in at 4 tonnes and measured roughly 10 metres (33 ft) from nose to tail. Like other hadrosaurs it had a toothless beak, the back of the jaws contained a dental battery composed of hundreds of small, interlocking teeth. These were used to crush and grind plant matter and were continually replaced as they wore away.
Diclonius	75 mya	Montana, USA
Edmontosaurus	73.0-76.5 mya	Canada	Edmontosaurus included some of the largest hadrosaurid species, measuring up to 12 metres (39 ft) long and weighing around 4.0 metric tons (4.4 short tons).
Gasparinisaura	85 mya	Argentina	Gasparinisaura was a small bipedal herbivore. In 2010 Gregory S. Paul estimated the length at 1.7 metres, the weight at thirteen kilogrammes.
Gilmoreosaurus	70 mya	Mongolia
Gryposaurus	83-75.5 mya	Alberta, Canada	Gryposaurus was a hadrosaurid of typical size and shape.
Hadrosaurus	79.5 mya	New Jersey, USA	It was likely bipedal for the purposes of running, but could use its forelegs to support itself while feeding.
Hypacrosaurus	75-67 mya	Alberta, Canada	Hypacrosaurus is most easily distinguished from other hollow-crested duckbills 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
Hypsibema		North Carolina and Missouri, USA
Kritosaurus	73 mya	North America	The type specimen of Kritosaurus navajovius is only represented by a partial skull and lower jaws, and associated postcranial remains.
Lambeosaurus	76-75 mya	Alberta, Canada
Lophorhothon	80 mya	Alabama, USA
Maiasaura	76.7 mya	Montana, USA	Maiasaura was large, attaining an adult length of about 9 metres (30 ft) and had the typical hadrosaurid flat beak and a thick nose. It had a small, spiky crest in front of its eyes. The crest may have been used in headbutting contests between males during the breeding season.
Mandschurosaurus		Asia
Microhadrosaurus		China
Mochlodon		Austria	A rhabdodontid.
Naashoibitosaurus	73 mya	New Mexico, USA	Naashoibitosaurus, based as it is on a single partial skeleton, is not well known in terms of anatomy. Its skull, the most thoroughly described portion, has a low nasal crest that peaks in front of the eyes, but does not strongly arch as in Gryposaurus.
Nipponosaurus		Russia
Orodromeus	76.7 mya	Montana, USA	Orodromeus was a small fast bipedal herbivore that probably coexisted with dinosaurs such as Daspletosaurus and Einiosaurus. Its length was estimated by Horner & Weishampel at 2.5 metres.
Parasaurolophus	76.5-73 mya	Alberta, Canada; New Mexico and Utah, USA
Prosaurolophus	76-75 mya	Alberta, Canada	Prosaurolophus was a large-headed duckbill; the most complete described specimen has a skull around 0.9 meters long (2.9 ft) on a ~8.5 meter long skeleton (~28 ft).[2] It had a small, stout, triangular crest in front of the eyes; the sides of this crest were concave, forming depressions. The upper arm was relatively short.
Pteropelyx		Montana, USA
Rhabdodon	70 mya	France; Spain; Haţeg Island, Romania	It is unclear whether it was an iguanodont or a hypsilophodont, and may be a "missing link" between the two. Current evidence indicates it is an iguanodont similar to Tenontosaurus.
Saurolophus	69.5-68.5 mya	North America, Asia	Saurolophus is known from material including nearly complete skeletons, giving researchers a clear picture of its bony anatomy. S. osborni, the rarer Albertan species, was around 9.8 meters long (32 feet), with its skull a meter long (3.3 feet). Its weight is estimated at 1.9 tonnes (2.1 tons). S. angustirostris, the Mongolian species, was larger; the type skeleton is roughly 12 meters long (39.4 ft), and larger remains are reported.
Shantungosaurus	70 mya	China	It is one of the longest and largest known hadrosaurids; the composite skeleton of a medium-sized individual mounted at the Geological Institute of China in Beijing measures 14.72 metres (48.3 ft) in length.
Tanius		China
Trachodon	77 mya	Montana, USA
Tsintaosaurus	70 mya	Southern China
Velafrons	72 mya	Mexico

†Pachycephalosaurs

Pachycephalosaurs of the Campanian
Taxa	Presence	Location	Description	Images
Alaskacephale		Prince Creek Formation, Alaska, USA		Alaskacephale Homalocephale
Colepiocephale		Alberta, Canada	The oldest known pachycephalosaurid.
Goyocephale		Mongolia
Gravitholus
Hanssuesia		Alberta, Canada; Montana, USA	Distinguished from other pachycephalosaurs by having a depressed parietal region, wide frontoparietal dome, broad nasal characteristics, reduced prefontal lobes, and a reduced parietosquamosal shelf.
?Heishansaurus
Homalocephale		Mongolia	Sporting a flat, wedge-shaped skull roof, Homalocephale was different from other pachycephalosaurs.
?Micropachycephalosaurus
Ornatotholus
Prenocephale
Sphaerotholus
Stegoceras
Tylocephale
Wannanosaurus

†Plesiosaurs

Plesiosaurs of the Campanian
Taxa	Presence	Location	Description	Images
†Elasmosaurus	80.5 mya	Pierre Shale, Kansas, USA	Elasmosaurus is a genus of plesiosaur with an extremely long neck.	Elasmosaurus Styxosaurus
†Styxosaurus	83.5-80.5 mya	Logan County, Kansas	Styxosaurus is an Elasmosaurid plesiosaur.

†Pterosaurs

Pterosaurs of the Campanian
Taxa	Presence	Location	Description	Images
Geosternbergia		USA, North America	Geosternbergia was originally a species of Pteranodon and is famous for its oddly shaped crest.
Nyctosaurus		mid-western United States	Nyctosaurus is a genus of pterodactyloid pterosaur.	Pteranodon Quetzalcoatlus
Pteranodon		Kansas, USA, North America	Pteranodon is a genus of pterosaurs which included some of the largest known flying reptiles, with wingspans over 6 metres
Quetzalcoatlus		Texas, USA	Quetzalcoatlus was a pterodactyloid pterosaur known from the Late Cretaceous of North America and one of the largest known flying animals of all time.
Piksi		Montana, USA	Piksi is a genus of pterosaurs containing the single species Piksi barbarulna.
Navajodactylus		New Mexico, USA, and Alberta, Canada	Known primarily from forearm elements; tentatively assigned to Azhdarchidae, though most likely not part of it.
Montanazhdarcho		Montana, USA	Small azhdarchid pterosaur.
Volgadraco		Saratov, Russia	Azhdarchid pterosaur.

†Sauropods

Sauropods of the Campanian
Taxa	Presence	Location	Description	Images
Alamosaurus		Southwestern United States	Alamosaurus is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now North America. It was a large quadrupedal herbivore.	Alamosaurus Saltasaurus
Andesaurus		Neuquén Province, Argentina	Andesaurus is a genus of basal titanosaurian sauropod dinosaur.
Dreadnoughtus		Cerro Fortaleza Formation, Argentina	Dreadnoughtus is one of the largest titanosaurs known.
Huabeisaurus		North East, China	A member of the Euhelopodidae sauropods.
Overosaurus		Neuquén Province, Argentina	Small-sized titanosaur.
Quaesitosaurus		Shar Tsav, Mongolia	Quaesitosaurus is a genus of titanosaurian sauropod.
Saltasaurus		north-west Argentina; Uruguay	Saltasaurus is a genus of titanosaurid sauropod dinosaur. An estimated length of 12 metres (39 feet) and a mass of 7 tonnes (8 tons).

Squamates

Squamates of the Campanian
Taxa	Presence	Location	Description	Images
Halisaurus				Mosasaurus Taniwhasaurus
Mosasaurus
Plotosaurus
Taniwhasaurus		New Zealand, Japan, Antarctica
Tylosaurus

Testudines

Testudines of the Campanian
Taxa	Presence	Location	Description	Images
†Archelon				Reconstruction of Archelon

†Theropods (non-avian)

David J. Varrichio observes that during the late Campanian Alberta and Montana had very similar theropods despite significant differences in the types of herbivorous dinosaur faunas.^[3]

Non-avian theropods of the Campanian
Taxa	Presence	Location	Description	Images
Abelisaurus		Allen Formation?, Anacleto Formation?, Argentina	Bipedal carnivore that probably reached 7 to 9 meters in length; known from only one partial skull.	Abelisaurus Bambiraptor Citipati Portrait of Saurornithoides Velociraptor
Albertosaurus
Appalachiosaurus
Archaeornithomimus
Bambiraptor
Byronosaurus
Citipati
Carnotaurus
Chirostenotes
Daspletosaurus
Dromaeosaurus
Dromiceiomimus
Dryptosaurus
Gorgosaurus
Harpymimus
Khaan
Lythronax		Wahweap Formation, Utah	A 7-meter tyrannosaurid known from a partially complete skull, some vertebrae and a complete pubis
Luanchuanraptor
Mononykus
Noasaurus
Ornithomimus
Oviraptor		Mongolia
Pyroraptor		Var, France
Saurornithoides
Saurornitholestes
Shuvuuia
Struthiomimus
Troodon
Variraptor		Var, France
Velociraptor		Mongolia and China
Xenotarsosaurus
Zhuchengtyrannus		Wangshi Group, Zhucheng, China	One of the largest tyrannosaurids at between 10–12 meters. Known from a lower jaw and maxilla slightly smaller than those of the later Tyrannosaurus.

Footnotes

References

• Gradstein, F.M.; Ogg, J.G. & Smith, A.G.; 2004: A Geologic Time Scale 2004, Cambridge University Press.
• Varricchio, D. J. 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from Montana. pp. 42–57 in D. H. Tanke and K. Carpenter (eds.), Mesozoic Vertebrate Life. Indiana University Press, Indianapolis, Indiana.
• Weishampel, D.B.; Barrett, P.M.; Coria, R.A.; Le Loueff, J.; Xu, X.; Zhao, X.; Sahni, A.; Gomani, E.M.P. & Noto, C.N.; 2004: Dinosaur distribution, in: Weishampel, D.B.; Dodson, P. & Osmólska, H. (eds.): The Dinosauria, University of California Press, Berkeley (2nd ed.), ISBN 0-520-24209-2, pp 517–606.

External links

• GeoWhen Database - Campanian
• Late Cretaceous timescale, at the website of the subcommission for stratigraphic information of the ICS
• Stratigraphic chart of the Late Cretaceous, at the website of Norges Network of offshore records of geology and stratigraphy
• Campanian Microfossils: 75+ images of Foraminifera