User:Montanoceratops/sandbox

Pachyrhinosaurus is a genus of Centrosaurine Ceratopsian that lived in North America during the Late Cretaceous

History
In 1945-1946 several skulls are discovered from the Horseshoe Canyon Formation. In 1947 Charles Sternberg published the abstract on a new Ceratopsian dinosaur representing a new family. In 1950 Sternberg named this species Pachyrhinosaurus canadensis. In 1975 Langston, W. Jr. described more material of P. canadensis from the St. Mary River Formation's Scabby Butte. Al Lakusta (1974) found a large bonebed along Pipestone Creek, Alberta, being excavated between 1986 and 1989 by staff and volunteers of the Royal Tyrrell. They found a very large, dense selection of material - up to 100 bones/square meter, totaling to 3500 bones and 14 skulls. In the 1980s a young Philip Currie and Darren Tanke were looking through the specimens in the TMP's collection when stumbling on the remains of a Ceratopsian from Al Lakusta's excavations, representing material from, "numerous individuals, with virtually every bone in the skeleton represented." In 2008 Currie, Langston & Tanke formally described the bones as Pachyrhinosaurus lakustai. In the 1998 a multitaxic bonebed in the Prince Creek Formation was discovered. Out of this a multitude of Pachyrhinosaurus skeletons were perserved. In 2012 Fiorillo & Tykoski described the remains from the 2006 expedition as belonging to a new species, Pachyrhinosaurus perotorum. In 2015 Kruk published a thesis on Pachyrhinosaurus ontogeny and histology. In that thesis she named a fourth species of Pachyrhinosaurus, P. "youngi". This specimen was collected in 2010. She says that, "This chapter will be submitted to PLoS ONE for publication." So far an official description hasn't been published, making P. "youngi" a nomen nudum. Finally, TMP 2002.76.1 was discovered in the Iddesleigh of the Dinosaur Park Formation in 1996, and was deposited and cataloged in the TMP in 2002. It was described by ''Ryan et. al''. (2010) as a new Ceratopsian, but wasn't given a name.

Description
Pachyrhinosaurus can be distinguished from Non-Pachyrostran Centrosaurs by the presence of a nasal boss instead of a nasal horn. Further, it can be distinguished from Achelousaurus, the other known Pachyrostran, by the more medially projected, curved P3 epiparietals, more pronounced P1 epiparietals, a shorter, rounder frill, by, "a relatively broader, longer, and typically thicker boss.", "the nasal rugosity extends posteriorly onto the frontals", and "Supraorbital boss is stouter and less rugose than in Achelousaurus."

P. canadensis
P. canadensis is known from the Horsehoe Canyon Formation and the laterally equivalent St. Mary River Formation. It was estimated by to be 6 meters long, 2 meters tall, and weigh 3 metric tones. P. canadensis can be distinguished from the other species of Pachyrhinosaurus by: the lack of a small rostral comb dorsal to the rostral (shared with P. "youngi"); nasal boss and supraorbital bosses that contact each other on the skull roof (shared with P. perotorum and P. "youngi"); presence of a supranarial ossification on the lateral surface of the rostrum (shared with P. perotorum, not perserved in P. "youngi"); nasal boss and other anterior facial ornamentations (including "fist-sized knobs") are more hypertrophied than P. lakustai, but less so than P. perotorum and P. "youngi"; lack of a single transverse shelf of bone on the medial side of the premaxilla (shared with P. perotorum and P. "youngi")' "long slender, finger-like premaxillary process that extends forward and then curves 90° medially"; "irregularly developed and vertical fluting", a flat parietal bar in lateral view"; "lateral parietal spike twists dorsolaterally"; and the presence of a "Supranasal Boss" (not shared with any other species). In 1967/1968 a skull from Drumheller was discovered by the Drumheller Dinosaur and Fossil Museum, which is now the Badlands Historical Centre. A taxon number wasn't given to this skull, but Currie, Langston, and Tanke dub this specimen the "Drumheller Skull". It is a large Pachyrhinosaurus canadensis.

P. lakustai
P. lakustai is known from the Wapiti Formation. It was around 5m long, and weigh 2 metric tons. P. lakustai can be distinguished from other known Pachyrhinosaurus species by: the "presence of a small rostral comb dorsal to the rostral" (shared with P. perotorum); the "presence of a supranarial ossification on the lateral surface of the rostrum" (shared with P. canadensis); nasal boss and other anterior facial ornamentations (including "fist-sized knobs") less hypertrophied than both P. canadensis, P. perotorum, and P. "youngi"; a single transverse shelf of bone on the medial side of the premaxilla (not shared by any other species); the "short robust premaxillary process projects straight forward into nares"; the parietal bar curves slightly upwards in lateral (possibly shared with P. "youngi"); "lateral parietal spike twists anteroventrally" (shared with P. "youngi" and to a lesser extent P. perotorum [more horizontal]); the lack of a "Supranasal Boss" (shared with P. perotorum and P. "youngi").

P. perotorum
P. perotorum is known from the Prince Creek Formation. It was around 5m long, and weigh 2 metric tons P. perotorum can be distinguished from other species of Pachyrhinosaurus by: "rostrum anterior tip upturned with anteroventrally directed premaxillary oral margin, resulting in a rostral bone shifted to a level dorsal to the maxillary tooth row"; "rostral bone lacks the acute, ventrally downturned, parrot-like beak common to other ceratopsians"; "nasal boss dorsal surface with median ridge near posterior end of boss" (none of these feature is shared with another species of Centrosaur). Both "epiparietal 2 (P2) posterior margin excavated by enhanced, trough-like, mediolaterally oriented sulcus" and "epiparietal 2 (P2) horn base penetrated by canal passing dorsoventrally through body of horn." are tentative autapomorphies of P. perotorum. Further, these features can be used to better distinguish P. perotorum from its relatives. "small rostral comb dorsal to the rostral" (shared with P. lakustai); the "nasal boss and supraorbital bosses that contact each other on the skull roof" (shared with P. canadensis and P. "youngi"); "presence of a supranarial ossification on the lateral surface of the rostrum" (shared with P. canadensis and not perserved in P. "youngi"); nasal boss and other anterior facial ornamentations (including "fist-sized knobs") are more hypertrophied than P. canadensis, P. lakustai, and P. "youngi"; the lateral parietal spike twists anteroventrally (similar, but less pronounced than P. lakustai and P. "youngi"; the lack of a "supranasal boss (shared with P. lakustai and P. "youngi".

P. "youngi"
P. "youngi" was found in the Wapiti Formation. Kruk didn't provide estimates on size of P. "youngi", however it can be assumed to be around 5-6m long and weighed 2-3 tons due to it's phylogenetic position. P. "youngi" can be differentiated from other Pachyrhinosaurus species by: the lack of a "small rostral comb dorsal to the rostral" (shared with P. canadensis); the "nasal boss and supraorbital bosses that contact each other on the skull roof" (shared with P. canadensis and P. perotorum); "dorsolaterally curved parietal horns"; a "distinctive tall separation between supraorbital bosses"; an "anteriorly protruding “pommel"" (all 3 unique to P. "youngi"); lack of a "single transverse shelf of bone on the medial side of the premaxilla" (shared with P. canadensis and P. perotorum); the lack of "Supranasal Boss" (shared with P. perotorum and P. lakustai.)

P. sp. nov.
TMP 2002.76.1 is known from the Dinosaur Park Formation. This specimen is known from a mostly complete skull missing the back of the frill, both radii and ulnae of the front feet, the right humerus, both scapulae, both scapulacoracoids, 3 complete feet, 2 tibiae (one partial one complete), a fibula, around 5 cervical vertebrae, around 14 dorsal vertebrae, and around 11 caudal vertebrae, and around 12 pairs of ribs, as well as a highly incomplete sacrum. If this taxon is Pachyrhinosaurus it would be the oldest species of the genus. Both Kruk (2015) and Tykoski et. al. (2019) recover this taxon as a taxon closer to Pachyrhinosaurus rather than Achelousaurus. However, Savhannah Carpenter's unpublished phylogeny from 2021 (labeled as Achelousaurus [Iddesleigh]) as well as ''Ryan et. al. (2010) places TMP 2002.76.1 in a polytomy with Achelousaurus and Pachyrhinosaurus''. The reason why it's position within Pachyrostra hasn't been consistent is due to the preservation of TMP 2002.76.1. TMP 2002.76.1 is missing the top of its frill, which is a vital character that distinguished Ceratopsids in general and Pachyrhinosaurus in particular. Until further expeditions to the Dinosaur Park Formation uncover another specimen of this species, it is unlikely to receive a name (despite several characters that distinguish it from Achelousaurus horneri and the other known Pachyrhinosaurus species ) due to the uncertainty of whether it is a new genus, a new species of Achelousaurus, or a new species of Pachyrhinosaurus.

In October 2013 a Darla Zelenitsky and her team discovered the skull of a "gargantuan pachyrhinosaur" within the limits of Drumheller, Alberta. The Pachyrhinosaur was unveiled at the University of Calgary in February of 2014. It was the largest skull of a Pachyrhinosaur discovered, over 6 meters in length. Zelenitsky estimates that the skull is 75-80% complete. She and her team planned on returning to the site in spring/summer of 2014 to determine if more of the "gargantuan pachyrhinosaur" is perserved, and determine "if this specimen represents a new species.” So far, no more news of the Pachyrhinosaur has been publicized since the original unveiling.

Species

 * P. canadensis (Sternberg, 1950)
 * P. lakustai (Currie, Langston & Tanke, 2008)
 * P. perotorum (Fiorillo & Tykoski, 2012)
 * P. "youngi" (Kruk, 2015, Thesis)
 * P. sp. nov.? TMP 2002.76.1, possibly the U of C "Gargantuan Pachyrhinosaur" See text

Phylogeny
This cladogram was recovered by Kruk (2015) in her thesis on P. "youngi".

Horseshoe Canyon Formation/ St. Mary River Formation
Pachyrhinosaurus canadensis is known from both the Horseshoe Canyon Formation and the laterally equivalent St. Mary River Formation. It shared it's environment with the Leptoceratopsid Montanoceratops, The Chasmosaurs Regaliceratops , Anchiceratops, and Arrhinoceratops. It would have also coexisted with hadrosaur Edmontosaurus regalis, The Ornithomimosaurs Ornithomimus and Struthiomimus, the Tyrannosaur Albertosaurus sarcophagus, The Ankylosaurs Anodontosaurus and Edmontonia longiceps. It would have also coexisted with the Troodontid Albertavenator, the Dromeosaur Atrociraptor , and the Caenagnathid Ovoraptorosaurs Apatoraptor and Epichirostenotes . Among Non-Dinosaurians the mammal Didelphodon coyi from Drumheller.  Stangerochampsa is from Horseshoe Canyon isn't clear in the article's description, but probably belongs to the Drumheller unit, making it coocurring. Basilemys is known from the Horseshoe Canyon Formation, but isn't given a locality, based on other formations it probably cooccurs with P. canadensis. The Choristoderan Champsosaurus albertensis and the plesiosaur Leurospondylus were found in Horseshoe Canyon, but no further locality information can be gathered.

Wapiti Formation
Pachyrhinosaurus lakustai and P. "youngi" are known from the Wapiti Formation. It would have shared its environment with A large Tyrannosaur, possibly Albertosaurus (represented by teeth and Bellatoripes tracks ), the Velociraptorine Boreonykus certekorum , the Hadrosaur Edmontosaurus, and Saurornitholestes sp. . "Troodon" and the morphotaxon Richardoestesia. are also present.

Prince Creek Formation
Pachyrhinosaurus perotorum is known from the Prince Creek Formation. It would have shared it's environment with an indeterminate Leptoceratopsid, The Hadrosaur ''Edmontosaurus sp. (previously within the junior synonym "Ugrunaaluk")'', an indeterminate Lambeosaurine represented by the supraoccipital DMNH 2014-12-266. The Pachycephalosaur Alaskacephale gangloffi is also present. Nanuqsaurus was also present within this formation. ''Dromaeosaurus cf. albertensis, Saurornitholestes cf. langstoni'', indeterminate Thescelosaurinae and indeterminate "Hypsilophodontid" teeth have been found within the formation. An indeterminate genus of saurornitholestinae, represented by the dentary fragment DMNH 21183 is present. A Distal metatarsal IV from an indeterminate Ornithomimosaur and the tracks of ichnogenus bird Gruipeda have also been reported. Of all the remains, only one non-dinosaurian faunal taxon is known, the metatherian mammal Unnuakomys.

Dinosaur Park Formation
TMP 2002.76.1 is known from the Dinosaur Park Formation, specifically the Lethbridge Coal. Very little has been reported from the Lethbridge Coal, among the known remains from the Dinosaur Park Formation: Centrosaurus, Chasmosaurus belli and C. russeli are Ceratopsids it would have coexisted with. The Hadrosaurids Corythosaurus, Gryposaurus, and Parasaurolophus walkeri. The Tyrannosaur Gorgosaurus would have lived at the same time and possibly hunted this taxon.

Notable Specimens

 * CMN 8867: The holotype of Pachyrhinosaurus canadensis and by proxy Pachyrhinosaurus as a whole.
 * CMN 8866: The paratype of Pachyrhinosaurus canadensis.
 * TMP 1986.55.258: The holotype of Pachyrhinosaurus lakustai.
 * DMNH 21200: The holotype of Pachyrhinosaurus perotorum.
 * DMNH 22558: Paratype of Pachyrhinosaurus perotorum.
 * DMNH 21201: Paratype of Pachyrhinosaurus perotorum.
 * UALVP 53300: Holotype of P. "youngi".
 * UALVP 54165: Paratype of P. "youngi"
 * UALVP 50652: Paratype of P. "youngi"
 * UALVP 55805 : Paratype of P. "youngi"
 * TMP 2002.76.1: A new species that possibly belongs to Pachyrhinosaurus, if so, it is the oldest Pachyrhinosaurus known.
 * "University of Calgary's Gargantuan Pachyrhinosaur": a taxon of pachyrhinosaur, possibly belonging to the new species represented by TMP 2002.76.1. It is the largest Pachyrhinosaur discovered.