Campanien

Reconstruction of the skull of Spinops sternbergorum gen. et sp. nov. from the Campanian of Dinosaur Provincial Park, southern Alberta, in right lateral view. Preserved elements are stippled; missing portions are dotted and modeled after Centrosaurus apertus.

Photograph of the holotype skull of Titanoceratops ouranos, from the late Campanian aged Kirtland/Fruitland beds of New Mexico, USA. Photograph by Nick Longrich

Representative taxa from the early Campanian Muthmannsdorf fauna from the Grünbach Formation, eastern Austria. A Doratodon carcharidens (Mesoeucrocodylia) mandible (PIUW 2349/57) in dorsal view (photo by Márton Rabi) B Indeterminate azhdarchid (Pterosauria, Azhdarchidae), left humerus (PIUW 2349/102) in anterior view C ‘Megalosaurus pannoniensis’ basal tetanuran (Theropoda, Tetanurae), tooth (PIUW uncatalogued) in lateral view D Mochlodon suessi (Ornithopoda, Rhabdodontidae), right dentary (holotype, PIUW 2349/2) in medial view. Scale bars equal 2 cm in A, B and D and 1 cm in C.

Representative taxa from the latest Campanian–Maastrichtian faunas from Transylvania, western Romania. A–B Nidophis insularis (Serpentes, Madtsoiidae), articulated vertebrae (LPB (FGGUB) v.547/2) in left lateral (A) and dorsal (B) views (Densuş-Ciula Formation, Tuştea, Haţeg Basin; photo by Ştefan Vasile) C Allodaposuchus precedens (Eusuchia, ?Hylaeochampsidae), skull (PSMUBB V 438) in dorsal view (Sebeş = Şard Formation, Oarda de Jos, southwestern Transylvanian Basin; photo by Vlad Codrea/Massimo Delfino) D Theriosuchus sympiestodon (Mesoeucrocodylia, Atoposauridae), right maxilla (MCDRD 793) in lateral view (Sînpetru Formation, Sînpetru, Haţeg Basin) E–F Indeterminate titanosaur (?Magyarosaurus dacus) (Sauropoda, Titanosauria), isolated osteoderm (LPB (FGGUB) R.1410) in dorsal (E) and lateral (F) views (Sînpetru Formation, Sînpetru, Haţeg Basin) G Indeterminate ornithuran bird (Aves, Ornithurae), incomplete left tibiotarsus (LPB (FGGUB) R.1902) in anterior view (Densuş-Ciula Formation, Vălioara, Haţeg Basin) H Balaur bondoc (Theropoda, Dromaeosauridae), articulated left distal hindlimb (EME PV.313) in lateral view (Sebeş = Şard Formation, Sebeş-Glod, southewestern Transylvanian Basin; photo by Mick Ellison) I Zalmoxes robustus (Ornithopoda, Rhabdodontidae), right dentary (NHMUK R.3407) in medial view (Sînpetru Formation, Sînpetru, Haţeg Basin) J Telmatosaurus transsylvanicus (Hadrosauria), right maxilla (MFGI unnumbered) in lateral view (Sînpetru Formation, Sînpetru, Haţeg Basin) K Indeterminate nodosaurid – Struthiosaurus transylvanicus or new taxon – (Ankylosauria, Nodosauridae), isolated tooth (LPB (FGGUB) R.2182) in medial view (Sînpetru Formation, Sînpetru, Haţeg Basin) L Barbatodon transylvanicus (Multituberculata, Kogaionidae), right maxilla (LPB (FGGUB) M.1635) in medial view (Sînpetru Formation, Pui, Haţeg Basin). Scale bars equal 1 mm in A, B; 5 mm in K; 1 cm in G, L; 2 cm in D; and 5 cm in C, E, F, H, I, J.

Campanian fauna from the Oldman Formation: Wendiceratops, Corythosaurus, Scolosaurus, Chasmosaurus and "Troodon"

Brachylophosaurus canadensis (TMP 2005.000.0029), Royal Tyrrell Museum, Drumheller, Alberta, 2025-07-13; cast, Campanian, from the Oldman Formation

African locality: 5, Majunga, Maevarano Formation (Campanian): Rahonavis ostromi.

Locality map: Grand Staircase-Escalante National Monument, southern Utah. Map showing the Nipple Butte area (indicated by yellow star) of Grand Staircase-Escalante National Monument (GSENM). GSENM is bounded by the red rectangle and silhouetted in dark gray on the inset of Utah and surrounding states (modified from [1]). The original map has been modified to show the Nipple Butte area instead of the Machairoceratops fossil locality as in the original source. New location based on map in A New Macrovertebrate Assemblage from the Late Cretaceous (Campanian) of Southern Utah, page 601.

Kaiparowits Formation (Campanian; Upper Cretaceous) at The Blues, near Powell Point (peak to the left) composed of the Claron Formation.

Map of Grand Staircase-Escalante National Monument (GSENM), southern Utah (A), and generalized stratigraphic section of the upper Campanian Kaiprowits Formation (B). The approximate stratigraphic position of Akainacephalus johnsoni is located near the base of the middle unit within the Kaiparowits Formation. The map highlights the GSENM boundary (dashed line), showing the geological distribution and outcrops of the Cretaceous Kaiparowits, Wahweap, and Straight Cliffs formations. The red star indicates the Horse Mountain area, from which Akainacephalus johnsoni was recorded. Map and stratigraphic column modified from Roberts (2005). Radioisotopic dates used from Roberts, Deino & Chan (2005) and Roberts et al. (2013), respectively.

Holotype of Khunnuchelys lophorhothon sp. nov. from Baybishe, northeastern Aral Sea region, Kazakhstan, Bostobe Formation, Santonian–early Campanian, Late Cretaceous, ZIN PH 5/55, a partial skull; in dorsal (A), ventral (B), lateral (C), and anterior (D) views. Photographs (A1–D1), explanatory drawings (A2–D2).

World map showing Late Cretaceous metatherian locales. Europe 1. Font-de-Benon quarry, Archingeay-Les Nouillers (Cenomanian, Late Cretaceous), Charente-Maritime, southwestern France (Vullo et al. 2009) 2. Valkenburg Member, Maastricht Formation (late Maastrichtian, Late Cretaceous), southern Limburg, The Netherlands (Martin et al. 2005) Asia 3. Yixian Formation, China (Barremian, Early Cretaceous) 4. Bissekty Formation, Kyzylkum Desert, Uzbekistan (Turonian, Late Cretaceous) 5. Darbasa Formation, southern Kazakhstan (Campanian, Late Cretaceous) 6. Grey Mesa locality (Averianov 1997) 7. Barun Goyot Formation, Umuni Gobi, Mongolia (Campanian, Late Cretaceous) 8. Nemegt Formation, Omnogov, Mongolia (Maastrichtian, Late Cretaceous) 9. Djadokhta Formation, Mongolia (Campanian, Late Cretaceous) North America Alaska 9. Prince Creek Formation, Alaska (early Maastrichtian, Late Cretaceous) Alberta and Saskatchewan, Canada 10. Milk River Formation, southern Alberta, Canada (late Santonian, Late Cretaceous) 11. Oldman Formation, southern Alberta, Canada (Campanian, Late Cretaceous) 12. Dinosaur Park Formation, southern Alberta, Canada (late Campanian, Late Cretaceous) 13. Horseshoe Canyon Formation, southern Alberta, Canada (early Maastrichtian, Late Cretaceous) 14. St. Mary River Formation, Alberta and northwestern Montana (early Maastrichtian, Late Cretaceous) 15. Scollard Formation, Alberta (late Maastrichtian, Late Cretaceous) 16. Frenchman Formation, Saskatchewan (late Maastrichtian, Late Cretaceous) Montana and North Dakota 17. Judith River Formation (late Campanian, Late Cretaceous) 18. Two Medicine Formation (late Campanian, Late Cretaceous) 19. Hell Creek Formation, Montana and North Dakota (late Maastrichtian, Late Cretaceous) South Dakota 20. Fox Hills Formation, South Dakota (late Maastrichtian, Late Cretaceous) 21. Hell Creek Formation, South Dakota (late Maastrichtian, Late Cretaceous) Wyoming 22. “Mesa Verde Formation” (late Campanian, Late Cretaceous) 23. Lance Formation, Wyoming (late Maastrichtian, Late Cretaceous) 24. Ferris Formation, Wyoming (late Maastrichtian, Late Cretaceous) Utah 25. Cedar Mountain Formation (Albian-Cenomanian) 26. Dakota Formation fauna (late Cenomanian, Late Cretaceous) 27. Smoky Hollow Member, Straight Cliffs Formation (Turonian, Late Cretaceous) 28. John Henry Member, Straight Cliffs Formation (Coniacian-Santonian, Late Cretaceous) 29. Wahweap Formation (early-middle Campanian, Late Cretaceous) 30. Kaiparowits Formation (late Campanian, Late Cretaceous) 31. Iron Springs Formation fauna, southern Utah (Turonian – Santonian, Late Cretaceous) 32. North Horn Formation, Utah (late Maastrichtian, Late Cretaceous) Colorado 33. Williams Fork Formation, Colorado (late Campanian-early Maastrichtian) 34. Laramie Formation, northeastern Colorado (late Maastrichtian, Late Cretaceous) Baja California Del Norte, Mexico 35. El Gallo Formation (late Campanian, Late Cretaceous) (Clemens 1980; Lillegraven 1972; Lillegraven 1976) New Mexico 36. Fruitland and lower Kirtland Formation, San Juan Basin (late Campanian, Late Cretaceous) 37. Naashoibito Member, Kirtland Formation, New Mexico (late Maastrichtian, Late Cretaceous) Oklahoma 38. Antlers Formation, Texas and Oklahoma (Aptian-Albian, Early Cretaceous) Texas 39. Aguja Formation, West Texas (late Campanian, Late Cretaceous) New Jersey 40. Marshalltown Formation, New Jersey (Campanian, Late Cretaceous) Szalay 1994)

World map showing Late Cretaceous metatherian locales. Europe 1. Font-de-Benon quarry, Archingeay-Les Nouillers (Cenomanian, Late Cretaceous), Charente-Maritime, southwestern France (Vullo et al. 2009) 2. Valkenburg Member, Maastricht Formation (late Maastrichtian, Late Cretaceous), southern Limburg, The Netherlands (Martin et al. 2005) Asia 3. Yixian Formation, China (Barremian, Early Cretaceous) 4. Bissekty Formation, Kyzylkum Desert, Uzbekistan (Turonian, Late Cretaceous) 5. Darbasa Formation, southern Kazakhstan (Campanian, Late Cretaceous) 6. Grey Mesa locality (Averianov 1997) 7. Barun Goyot Formation, Umuni Gobi, Mongolia (Campanian, Late Cretaceous) 8. Nemegt Formation, Omnogov, Mongolia (Maastrichtian, Late Cretaceous) 9. Djadokhta Formation, Mongolia (Campanian, Late Cretaceous) North America Alaska 9. Prince Creek Formation, Alaska (early Maastrichtian, Late Cretaceous) Alberta and Saskatchewan, Canada 10. Milk River Formation, southern Alberta, Canada (late Santonian, Late Cretaceous) 11. Oldman Formation, southern Alberta, Canada (Campanian, Late Cretaceous) 12. Dinosaur Park Formation, southern Alberta, Canada (late Campanian, Late Cretaceous) 13. Horseshoe Canyon Formation, southern Alberta, Canada (early Maastrichtian, Late Cretaceous) 14. St. Mary River Formation, Alberta and northwestern Montana (early Maastrichtian, Late Cretaceous) 15. Scollard Formation, Alberta (late Maastrichtian, Late Cretaceous) 16. Frenchman Formation, Saskatchewan (late Maastrichtian, Late Cretaceous) Montana and North Dakota 17. Judith River Formation (late Campanian, Late Cretaceous) 18. Two Medicine Formation (late Campanian, Late Cretaceous) 19. Hell Creek Formation, Montana and North Dakota (late Maastrichtian, Late Cretaceous) South Dakota 20. Fox Hills Formation, South Dakota (late Maastrichtian, Late Cretaceous) 21. Hell Creek Formation, South Dakota (late Maastrichtian, Late Cretaceous) Wyoming 22. “Mesa Verde Formation” (late Campanian, Late Cretaceous) 23. Lance Formation, Wyoming (late Maastrichtian, Late Cretaceous) 24. Ferris Formation, Wyoming (late Maastrichtian, Late Cretaceous) Utah 25. Cedar Mountain Formation (Albian-Cenomanian) 26. Dakota Formation fauna (late Cenomanian, Late Cretaceous) 27. Smoky Hollow Member, Straight Cliffs Formation (Turonian, Late Cretaceous) 28. John Henry Member, Straight Cliffs Formation (Coniacian-Santonian, Late Cretaceous) 29. Wahweap Formation (early-middle Campanian, Late Cretaceous) 30. Kaiparowits Formation (late Campanian, Late Cretaceous) 31. Iron Springs Formation fauna, southern Utah (Turonian – Santonian, Late Cretaceous) 32. North Horn Formation, Utah (late Maastrichtian, Late Cretaceous) Colorado 33. Williams Fork Formation, Colorado (late Campanian-early Maastrichtian) 34. Laramie Formation, northeastern Colorado (late Maastrichtian, Late Cretaceous) Baja California Del Norte, Mexico 35. El Gallo Formation (late Campanian, Late Cretaceous) (Clemens 1980; Lillegraven 1972; Lillegraven 1976) New Mexico 36. Fruitland and lower Kirtland Formation, San Juan Basin (late Campanian, Late Cretaceous) 37. Naashoibito Member, Kirtland Formation, New Mexico (late Maastrichtian, Late Cretaceous) Oklahoma 38. Antlers Formation, Texas and Oklahoma (Aptian-Albian, Early Cretaceous) Texas 39. Aguja Formation, West Texas (late Campanian, Late Cretaceous) New Jersey 40. Marshalltown Formation, New Jersey (Campanian, Late Cretaceous) Szalay 1994)

Map of Cretaceous-aged dinosaur fossil localities of Mongolia. Gobihadros mongoliensis was collected from Bayshin Tsav in Area C. Open squares indicate Late Cretaceous sites, solid squares represent Early Cretaceous localities. Abbreviations: A, Localities of Western Gobi Desert in Mongolia, mainly group of localities of Nemegtian age (early Maastrichtian), Late Cretaceous; B, Localities of Central Gobi Desert in Mongolia, mainly Djadokhtian age (Campanian), Late Cretaceous; C & D- Localities of Eastern Gobi Desert in Mongolia, mainly Baynshirenian age (Cenomanian-Santonian), Late Cretaceous. Figure has been modified from Tsogtbaatar et al. 2014, Figure 1 [24].

Tarsometatarsi of caenagnathids from the Upper Campanian Dinosaur Park Formation, Dinosaur Provincial Park, Alberta, Canada. A. TMP 1993.036.0181 (Chirostenotes pergracilis). B. TMP 1982.016.0006 (Citipes elegans). C. ROM 781 (Citipes elegans holotype). D. TMP 1996.012.0141 (Citipes elegans). Tarsometatarsi in proximal (A1–D1), anterior (A2–D2), and posterior (A3–D3) views.

Crâne de mosasaure d'espèce Platecarpus tympaniticus (squamates, mosasaures). Provenance : Smoky Hill Chalk, Kansas (aux Etats-Unis). Date : Crétacé supérieur, période du Campanien, 88 millions d'années avant notre ère. Collections du Muséum national d'histoire naturelle de Paris (France). Exposé à l'occasion de l'exposition "Un T-Rex à Paris" au Muséum national d'histoire naturelle de Paris (France) du 6 juin au 2 septembre 2018. Légende du fossile dans cette exposition : "Les mosasaures ne sont pas des dinosaures mais des reptiles marins, très répandus à la fin du Crétacé. Certains genres comme Globidens et Halisaurus sont connus aussi bien aux Etats-Unis qu'au Maroc. Ces animaux étaient d'excellents nageurs, capables de traverser l'Atlantique."