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110-567: Acanthopholididae Nopcsa , 1902 Acanthopholidae Nopcsa, 1917 ? Hylaeosauridae Nopcsa, 1902 Palaeoscincidae Nopcsa, 1918 Panoplosauridae Nopcsa, 1929 Struthiosauridae Kuhn, 1966 Nodosauridae is a family of ankylosaurian dinosaurs known from the Late Jurassic to the Late Cretaceous periods in what is now Asia, Europe, North America, and possibly South America. While traditionally regarded as

220-794: A monophyletic clade as the sister taxon to the Ankylosauridae , some analyses recover it as a paraphyletic grade leading to the ankylosaurids. Nodosaurids, like their sister group the ankylosaurids, were heavily armored dinosaurs adorned with rows of bony armor nodules and spines ( osteoderms ), which were covered in keratin sheaths. Nodosaurids, like other ankylosaurians, were small- to large-sized, heavily built, quadrupedal , herbivorous dinosaurs , possessing small, leaf-shaped teeth. Unlike ankylosaurids, nodosaurids lacked mace-like tail clubs, instead having more flexible tail tips. Many nodosaurids had spikes projecting outward from their shoulders. One particularly well-preserved nodosaurid "mummy",

330-555: A definition followed by Vickaryous, Teresa Maryańska , and Weishampel in 2004. Vickaryous et al. considered two genera of nodosaurids to be of uncertain placement ( incertae sedis ): Struthiosaurus and Animantarx , and considered the most primitive member of the Nodosauridae to be Cedarpelta . Following the publication of the PhyloCode , Nodosauridae needed to be formally defined following certain parameters, including that

440-530: A diet regularly consisting of almost 85% fern material, along with 3.7% cycad matter, trace elements of conifers , and 11.4% undiagnostic plant remains. Gastroliths may have been found with Panoplosaurus as well, but their identification is uncertain as they were not originally mentioned among the material found as part of the specimen. While nodosaurids were traditionally thought to have had simply sinuses, lacking complicated cavities and paranasal sinuses. While this can be seen in some taxa like Edmontonia ,

550-427: A distinct Chassternbergia , and Panoplosaurinae including only Panoplosaurus . The first phylogenetic analysis to include Panoplosaurus was that of Yuong-Nam Lee in 1996 , where Panoplosaurus mirus , including both CMN 2759 and ROM 1215 nested next to Edmontonia , which included both E. longiceps and E. rugosidens , to the exclusion of all other nodosaurids. A similar union of Panoplosaurus and Edmontonia

660-433: A distinct shape, the osteoderms are keeled, with the strength of the keel dependent on location. Lambe identified 7 different categories of osteoderms in his 1919 description of Panoplosaurus . The first kind was large, paired elements with a low keel, which formed bands around the neck leading from the head. On the sides of the neck to the back was a second kind, individual elements that were slightly smaller, suboval, and had

770-668: A general placement in Nodosauridae . Edmontonia was for a time considered the same taxon as Panoplosaurus , making it the only nodosaur from the Campanian of North America, but this was quickly disputed and they are now considered separate. Following consistent placements in phylogenetic analyses close to Edmontonia and the American taxon Animantarx , Panoplosaurus was placed into the clade Panoplosaurini , related but not close to Nodosaurus or Struthiosaurus , which it

880-468: A generic method of feeding due to their simple teeth, stiff skulls, and unspecialized musculature, comparable to modern iguanids . However, unlike iguanids ankylosaurs frequently have significant tooth wear, sometimes all the way down to the base of the crown. Nodosaurids normally have oblique wear facets on teeth, in Panoplosaurus having a single wear facet per tooth which changes angulation across

990-580: A keel were identified as a sixth form, and were suggested to have been from the limbs, though this was not definitive. The final form of scute were small ossicles, which occurred all over the animal filling in gaps between the larger osteoderms. The scutes along the top and sides of the neck in Panoplosaurus are the most distinct form, differing significantly from the corresponding elements in Edmontonia . Three bands of cervical osteoderms were present in both genera, consisting of rounder plates that united on

1100-439: A large variety of flora and fauna . Periodic areas of standing water would occur in the low-lying plains. Where there was dense vegetation, conifers were the dominant plants, with an understory composed of ferns and tree ferns , and flowering herbs and shrubs. The forest floor was covered by decaying vegetative matter, small ferns, lycopods and angiosperms , mosses , lichens , and fungi , with plentiful algae where there

1210-551: A large variety of animals. The herbivorous fauna of the Dinosaur Park Formation can be divided into two Megafaunal Assemblage Zones, defined as the lowest 28 m (92 ft) directly above the Oldman Formation, and the deposits above those, where the common ceratopsid Centrosaurus and hadrosaurid Parasaurolophus are absent. Panoplosaurus , which was found 24.9 m (82 ft) above

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1320-532: A motorcycle journey together with his long-standing Albanian secretary and lover Bajazid Elmaz Doda to study fossils . He later returned to Vienna where he ran into financial difficulties again and was distracted in his work. To cover his debts, he sold his fossil collection to the Natural History Museum in London. Nopcsa struggled with illness, to the extent that he had to give a lecture in

1430-441: A possible evolutionary response to a tougher, fibrous diet. However, microwear on the teeth of ankylosaurids and Panoplosaurus instead show that their diets were not significantly different, with regular pitting and abrasion showing a diet consisting occasionally of fruit as well as more abundant foliage. Stomach contents of the closely related taxon Borealopelta were identified amongst gastroliths , showing that at least it had

1540-423: A reigning European monarch, I would have no difficulty coming up with the further funds needed by marrying a wealthy American heiress aspiring to royalty, a step which under other circumstances I would have been loath to take.' During his lifetime Nopcsa wrote a memoir based on diaries and notes from 1897 to 1917. Even though he finished the memoir around 1929, it was never published during his lifetime. Only in 2001

1650-407: A result of this consistent support, Daniel Madzia and colleagues decided to name the clade uniting all taxa closer to Panoplosaurus than Nodosaurus or Struthiosaurus , giving it the formal name Panoplosaurini , modifying the suffix -inae from previous uses as it was continually nested within the clade Nodosaurinae . The reference phylogeny for Panoplosaurini designated by Madzia and colleagues

1760-409: A short snout, with a very domed surface, and bony plates directly covering the cheek. The neck had circular groups of plates arranged around the top surface, both the forelimb and hindlimb were about the same length, and the hand may have only included three fingers. Almost the entire surface of the body was covered in plates, osteoderms and scutes of varying sizes, ranging from large elements along

1870-412: A single one to analyse both Nodosauridae and Ankylosauridae. Panoplosaurus was resolved next to Edmontonia , deep within an unresolved polytomy of all nodosaurids more derived than Animantarx , which included Niobrarasaurus , Nodosaurus , Pawpawsaurus , Sauropelta , Silvisaurus , Stegopelta , and Texasetes . In 2016 , the phylogenetic analysis of Arbour and Currie initially meant to test

1980-414: A strong keel. Small, keeled scutes with a thick base were identified as the fourth osteoderm type, occurring on the underside of the base of the neck forwards to the chin. A fifth kind of osteoderms was identified as small, polygonal elements that fit together along the underside of Panoplosaurus , slowly grading into the larger rectangular elements of category two on the sides. Small irregular scutes lacking

2090-490: A tapering snout, lumpy cranial armour, a swollen vomer bone in the palate , tall neural arches and neural spines in the vertebrae, a small and round coracoid fused to the scapula, and a hand that may have only had three digits. There is also a prominent armour plate covering the cheek in the type specimen of Panoplosaurus , which may be unique feature of the taxon, or individual, depending on what additional skulls are referred to P. mirus . The skull of Panoplosaurus

2200-464: A taphonomic bias is present in the formation towards the better preservation of large-bodied animals like Panoplosaurus , which is 63% complete. Six small lizards are known, representing five different families, with the teiids Socognathus and Glyptogenys , the xenosaurid Exostinus , the helodermatid Labrodioctes , the necrosaurid Parasaniwa , and the varanid Palaeosaniwa . No snakes are known, which, while it contrasts with

2310-566: A theory about the dinosaurs' sexual dimorphism , which he published in 1926. Among others, he thought that hadrosaurid species with the cranial crests were males and those without them were females. He paired Kritosaurus with Parasaurolophus , Prosaurolophus with Saurolophus and others. His examples were not proved to be true, but his opinion that sexual dimorphism was present among hadrosaurid dinosaurs has gained acceptance, see for example Lambeosaurus . Nopcsa discovered and named several species in his lifetime. In 1899 he named

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2420-660: A wheelchair in 1928. Soon Nopcsa became depressed. Finally, in 1933, he fatally shot first his partner, Bayazid Elmas Doda, after having slipped sleeping powder into his tea. He then wrote a suicide note, where he states the reason for his actions a nervous breakdown, and shot himself. He was cremated at Feuerhalle Simmering in Vienna, and his ashes buried there (Section 3, Ring 3, Group 8, No. 44). In his suicide note, he describes his reasons for killing his partner: The reason that I shot my longtime friend and secretary, Mr Bayazid Elmas Doda, in his sleep without his suspecting at all

2530-397: Is a robust, 430 mm (17 in) long bone, with the shape and projection of the head suggesting the upper arm of Panoplosaurus was held in a flexed position in life. The only other portions of the forelimb known are three well-preserved, articulated digits of the hand, which may represent the complete hand suggesting Panoplosaurus had fewer fingers than its relatives. The manus

2640-448: Is broad and depressed, narrowing towards the end of the blunt snout to form a triangular shape. At a total length of 335 mm (13.2 in) in a straight line, the skull is curved in a way that across the skull roof the same length is 530 mm (21 in). Behind the orbits , the skull reaches its maximum width of 295 mm (11.6 in), giving it a very short and broad appearance. As other nodosaurid specimens from

2750-421: Is characteristic of the formation and similar deposits, and lived alongside the less common shark Hybodus montanensis , intermediate paddlefish and sturgeons , the long, slender fish Belonostomus , the gar Lepisosteus , bowfins , and an abundance of small teleosts including Paratarpon and Cretophareodus . At least nine forms of amphibians were present in the Dinosaur Park Formation, including

2860-621: Is defined in the PhyloCode as "the largest clade containing Panoplosaurus mirus , but not Nodosaurus textilis and Struthiosaurus austriacus " while Struthiosaurini has a similar definition of "the largest clade containing Struthiosaurus austriacus , but not Nodosaurus textilis and Panoplosaurus mirus ". Topology A below demonstrates these relationships, following the phylogenetic analyses of Rivera-Sylva and colleagues (2018), with clade names added by definition from Madzia et al. (2021). However, in 2023, Raven and colleagues proposed an alternate phylogeny for nodosaurids; instead of

2970-461: Is from a different age and location from other specimens, may instead represent the ankylosaurid taxon Nodocephalosaurus , although this is uncertain due to lack of overlapping material. 18 other specimens in the ROM, CMN, American Museum of Natural History , and Princeton University were referred to the genus by Walter P. Coombs in 1978 , but these were referred on the assumption that Edmontonia

3080-401: Is now shared by much of the scientific community. Nopcsa studied Transylvanian dinosaurs intensively, even though they were smaller than their "cousins" elsewhere in the world. For example, he unearthed six-meter-long sauropods , a group of dinosaurs which elsewhere commonly grew to 30 meters or more, which he named Magyarosaurus . Nopcsa deduced that the area where the remains were found

3190-424: Is possible that the function of these complicated sinuses was to warm incoming air as it passed through the skull. Inflowing air in Panoplosaurus was simulated to undergo a raise of between 17.9 and 18.2 °C (64.2 and 64.8 °F), primarily in the elongate nasal passage, while saturating the air with moisture. This is less heat efficient than the more complicated sinuses of Euoplocephalus , but still shows that

3300-455: Is present among the fauna of the Dinosaur Park Formation, representing an extinct group of semi-aquatic animals with crocodilian features that spanned the globe for much of the Mesozoic and Cenozoic . Mosasaurs and plesiosaurs are both known from the Dinosaur Park Formation, though mosasaurs are only represented by the teeth of Plioplatecarpus from the upper deposits, and apart from

3410-578: Is that I did not wish to leave him behind sick, in misery and without a penny, because he would have suffered too much. Nopcsa left behind a considerable quantity of scientific publications and private diaries. The diaries paint a picture of a complex man with great intuition, but without the ability to understand the motives of others. His devotion to the cause of the Albanians was in contrast to his sociopathic insensitivity. In his diaries he nonchalantly wrote about his bid to become king of Albania : Once

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3520-409: The synsacrum , but the synsacrum, as in other nodosaurids, is partially complete. Sternberg identified that the ilium of Panoplosaurus was supported by at least six vertebrae in 1921, which he identified as a dorsosacral, four true sacral vertebrae , and a caudosacral. The identification of four sacrals was questioned by Carpenter in 1990, as Edmontonia only has three true sacrals, but he

3630-538: The First World War , Nopcsa was on another mission as a spy for Austria-Hungary , working undercover as a shepherd in Transylvania. He also led a group of Albanian wartime volunteers. Nopcsa was the first to hijack  an aircraft. His motive for aircraft hijacking was to flee the nascent and ultimately short-lived Hungarian Soviet Republic in 1919 to Vienna. With the defeat of Austria-Hungary at

3740-681: The Geological Survey of Canada discovered a complete skull and significant amount of the skeleton of an armoured dinosaur in the sandstone layers of Quarry 69 of the Belly River Group , 64 m (210 ft) above sea level. The specimen, designated by the Canadian Museum of Nature accession number CMN 2759, and excavated 4.43 km (2.75 mi) south of the mouth of the Little Sandhill tributary of

3850-708: The Late Cretaceous of Alberta , Canada . Few specimens of the genus are known, all from the middle Campanian of the Dinosaur Park Formation , roughly 76 to 75 million years ago. It was first discovered in 1917, and named in 1919 by Lawrence Lambe , named for its extensive armour, meaning "well-armoured lizard". Panoplosaurus has at times been considered the proper name for material otherwise referred to as Edmontonia , complicating its phylogenetic and ecological interpretations, at one point being considered to have existed across Alberta, New Mexico and Texas , with specimens in institutions from Canada and

3960-535: The Red Deer River in Alberta , includes a nearly complete skull in articulation, most or all of the cervical vertebrae and the front dorsal vertebrae , and armour plates covering them, a majority of the disarticulated forelimb and three articulated fingers , a fragment of the pelvis and partial sacrum , a few bones of the foot , and multiple hundreds of osteoderms and dermal ossicles. This material

4070-469: The Turks . However, afterwards the newly liberated states immediately plunged into internal conflicts. During these Balkan Wars, Nopcsa spied for Austria-Hungary . Out of these conflicts, Albania arose as an independent state, which needed a king. Nopcsa volunteered, suggesting he would use money he would gain from marrying a rich American girl to fund the war efforts, however, to no avail. Later, during

4180-481: The United States . The skull and skeleton of Panoplosaurus are similar to its relatives, but have a few significant differences, such as the lumpy form of the skull osteoderms , a completely fused shoulder blade , and regularly shaped plates on its neck and body lacking prominent spines. It was a quadrupedal animal, roughly 5 m (16 ft) long and 1,600 kg (3,500 lb) in weight. The skull has

4290-419: The olfactory bulb and cerebral hemisphere being 44.0, 46.2 and 54.0 respectively. However, the size of the region of the brain devoted to the sense of smell is smaller in Panoplosaurus than expected for an animal of its size. The Dinosaur Park Formation deposits are a 70 m (230 ft) layer of sediments exposed in the badlands of Alberta, lying on top of the older Oldman Formation and underneath

4400-420: The physiology and living behavior of the dinosaurs he was studying. Nopcsa was the first to suggest that these archosaurs cared for their young and exhibited complex social behavior, an idea that did not take off until the 1980s. Because he was one of the first people to study the biology of dinosaurs, he is known as the 'father of paleobiology', even though he himself coined the term " paleophysiology " for

4510-459: The salamander -like Albanerpeton , two unnamed forms of frog , and salamanders from the genera Scapherpeton , Lisserpeton , Opisthotriton , Habrosaurus , and two or three unnamed forms. Turtles are commonly well-preserved in the formation, with many taxa known from many specimens. Three genera in the primitive turtle family Baenidae are known, Plesiobaena , Boremys and Neurankylus . The low-domed macrobaenid Judithemys

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4620-410: The 385 mm (15.2 in) tibia being noticeably longer than the 310 mm (12 in) fibula. As in ankylosaurids, the tibia of Panoplosaurus was shorter than the humerus, which contrasts with Stegosaurus and basal taxa like Scelidosaurus . This suggests that the forelimb of Panoplosaurus was only slightly shorter than the forelimbs. Both the metatarsal and phalanges of the foot resemble

4730-541: The Albanian mountains, he was held hostage by the bandit Mustafa Lita, together with Bajazid Doda. Lita demanded ten thousand Turkish pounds for his release. In his memoirs Nopcsa described his elaborate plan to get out of this situation, which involved being taken to Prizren as a spy. He was eventually rescued by Doda's father, who had brought 'ten armed retainers'. In 1912 the Balkan states joined forces to drive out

4840-720: The addition of another internal specifier was deemed unnecessary. Nodosauridae is traditionally composed of the basal clade Polacanthinae (sometimes recovered outside of the Nodosauridae), as well as the Panoplosaurini and Struthiosaurini within the Nodosaurinae . Nodosaurinae is defined in the PhyloCode as "the largest clade containing Nodosaurus textilis , but not Hylaeosaurus armatus , Mymoorapelta maysi , and Polacanthus foxii ". Panoplosaurini

4950-547: The armour on the torso and tail of Panoplosaurus is unknown, as no elements were found in articulation or association with this region of the skeleton. Panoplosaurus was originally named simply as a genus of armoured dinosaur by Lambe in 1919, within the group Stegosauria , though it was subsequently specified by Sternberg in 1921 that within Stegosauria it was closest to taxa like Ankylosaurus and Euoplocephalus , and so within Ankylosauridae . The description of

5060-461: The bone separation being a slight thickening along what would be the suture. The scapular region of the bone is relatively short, but is concave following the curvature of the body and curves down towards its distal end. Because of the curvature of the blade, the coracoid in Panoplosaurus sat at the front of the chest, as in Stegosaurus , Triceratops and Hadrosaurus . The top margin of

5170-475: The bones with Professor Eduard Suess , who encouraged him to study them. Following the professor's advice, he started studying geology at the University of Vienna in 1897, where he quickly advanced in his studies. He gave his first academic lecture in 1899 at the age of twenty-two. He acquired a PhD in geology in 1903 from the university; his doctorate focused on geologically mapping the area surrounding

5280-599: The classification of Panoplosaurus and Edmontonia to Nodosauridae. In 1998 , Sereno modified his earlier position, using the name Nodosaurinae for the group of derived nodosaurids including Panoplosaurus , and informally defining the subfamily as all taxa closer to Panoplosaurus than Sarcolestes or Hylaeosaurus . Tracy Ford published a novel classification of Panoplosaurus in 2000 , using osteoderm anatomy to divide Nodosauridae into Nodosaurinae including only Nodosaurus , Sauropeltinae including Sauropelta and Silvisaurus , Edmontoniinae including Edmontonia and

5390-526: The classifications of Ankylosauria in 1978, which he used as the group to encompass all heavily armoured ornithischians in a similar usage to Nodosauridae of Nopcsa. Within this suborder, he divided all taxa into two families, Ankylosauridae and Nodosauridae, distinguished by cranial and postcranial anatomy. Within Nodosauridae, which he considered a senior synonym of Acanthopholinae, Panoplosaurinae, Edmontoniinae, and other equivalent names, Coombs synonymized Panoplosaurus with Edmontonia , making Panoplosaurus

5500-445: The contact, is from layers just younger than those bearing Edmontonia , but still corresponding to the upper portion of the older MAZ-1, and would have coexisted with the ankylosaurid Euoplocephalus , ceratopsids Centrosaurus and Chasmosaurus belli , and hadrosaurids Corythosaurus intermedius , Lambeosaurus lambei , Lambeosaurus clavinitialis , and Parasaurolophus . As well as herbivorous megafauna, unnamed ornithopods ,

5610-457: The corresponding elements of the hand, but are slightly larger and more robust. Though the tibia and fibula of Panoplosaurus are crushed, they appear to be straighter than the elements in Edmontonia , which are curved along their length. Armour from the neck and trunk of Panoplosaurus , some of it still in articulation with bones, is known. A gradient of sizes exist from plate-like paired elements through to indistinct ossicles. Where they have

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5720-624: The end of the war, Nopcsa's native Transylvania was ceded to Romania . As a consequence, the Baron of Felső-Szilvás lost his estates and other possessions in 1920. Compelled to find paid employment, he landed a job as the head of the Hungarian Geological Institute in 1925. But Nopcsa's tenure in the Geological Institute was short-lived, he soon became bored of the sedentary job. He went to Europe on

5830-515: The family estate. On 20 November 1906 Nopcsa met the then eighteen-year-old Bajazid Elmaz Doda in Bucharest and hired him as his secretary. Nopcsa later recounted this meeting in his memoir: [H]e has been the only person who has truly loved me and in whom I had full confidence, never doubting for a moment that he would misuse my trust. Additionally, Nopcsa was interested in Albania , which

5940-624: The formation show that the alluvial channels were flowing to the east or southeast, towards the Bearpaw Sea of the Western Interior Seaway, creating meandering paths as the flow of water changed down and up during tidal cycles across the floodplain was balancing out. The climate of the environment in the Cretaceous was much different than in present day, being warmer and more temperate, with wet and dry seasons allowing for

6050-412: The front and rear edges of the crown, though there are more on the front than rear edge of the tooth. There are not significant differences between the teeth of Panoplosaurus and those of other armoured dinosaurs. The number of vertebrae in Panoplosaurus is unknown, as the dorsal and caudal series' are incomplete, and the cervical vertebrae are obscured by armour. The first three cervicals after

6160-426: The genus to it, the holotype CMN 2759, and specimen AMNH 3072, all from the upper Dinosaur Park Formation. Arbour and Philip J. Currie further restricted Panoplosaurus to just the holotype in 2016 , as a result of uncertainties surrounding the referrals of various specimens between it, Edmontonia rugosidens and Edmontonia longiceps , all from the Dinosaur Park Formation and similar deposits. Panoplosaurus

6270-664: The genus, including Royal Ontario Museum (ROM) 1215 and Royal Tyrrell Museum of Paleontology (RTMP) 83.25.2 from the Dinosaur Park Formation of Alberta, and an unnumbered Oklahoma Museum of Natural History (OMNH) scapulocoracoid from the Aguja Formation of Texas . A scapula from the Naashoibito member of the Kirtland Formation in New Mexico was referred to Panoplosaurus in 1981 , but as it

6380-472: The ground, which they would flap as they made a jump. The scales on its forearms would develop into feathers to aid this, and eventually allowing for flight. This theory found favor in the 1960s and later gained wide acceptance, though later fossil finds of tree-living feathered dinosaurs suggest the development of flight may have been more complex than Nopcsa envisioned. Additionally, Nopcsa's conclusion that at least some Mesozoic era reptiles were warm-blooded

6490-521: The holotype of Borealopelta markmitchelli , preserves a nearly complete set of armor in life position, as well as the keratin covering and mineralized remains of the underlying skin, which indicate reddish pigments in a countershading pattern. The family Nodosauridae was erected by Othniel Charles Marsh in 1890, and anchored on the genus Nodosaurus . The clade Nodosauridae was first informally defined by Paul Sereno in 1998 as "all ankylosaurs closer to Panoplosaurus than to Ankylosaurus ,"

6600-492: The latter from many specimens such as teeth throughout the entire formation, and troodontids from the MAZ-1 are limited to Stenonychosaurus . Three genera of oviraptorosaurs are known, all of which likely coexisted in the Dinosaur Park Formation, Caenagnathus , Chirostenotes and Citipes . It is possible that a therizinosaur was present based on a single frontal bone , and three ornithomimosaurs are preserved in

6710-408: The lower portion of the snout, are somewhat horseshoe-shaped, form a sharp beak that fits within the overhang of the premaxillaries in the upper jaw. Both the predentaries and their premaxillary counterparts lack any teeth, a derived feature among nodosaurids where premaxillary teeth are sometimes present. The teeth of the mandible are hidden by armour on the right side, and on the left side of

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6820-410: The midline of the animal, and one narrower element on each side with a sharp keel. In Panoplosaurus both the first and second bands of neck osteoderms had a third pair, lower on the side of the animal, again possessing a sharper keel than the elements on the top of the neck. While Edmontonia possesses lateral spines on the rear neck and shoulders, these are absent in Panoplosaurus . The arrangement of

6930-511: The modern diversity of the group, does correspond with the rarity of the group during the Cretaceous. Two pterosaurs have been found, the azhdarchid Cryodrakon and an unnamed pterodactyloid . All three major types of mammal present in the Cretaceous have been found in the Dinosaur Park Formation, Multituberculata , Marsupialia and Placentalia . The named multituberculates include Cimexomys , Cimolodon , Cimolomys , Meniscoessus , and Mesodma . Marsupials are represented by

7040-458: The nasal system of Panoplosaurus is far more complex than previously thought. The complete nasal passage of Panoplosaurus undergoes two complete 360 degree loops in different planes along its length, before entering the olfactory recess for scent processing. However, while the shape of the nasal passages is more complicated, Panoplosaurus does lack the additional parallel sinus tracts that can be found in ankylosaurids like Euoplocephalus . It

7150-562: The new genus Edmontonia in 1928 by Sternberg identified it as a very close relative of Panoplosaurus , almost close enough to be considered the same genus, both being closely related to Palaeoscincus . Following further description of the armoured taxon Struthiosaurus by Franz Nopcsa in 1929 , a different classification war proposed. Thyreophora replaced Stegosauria as the clade encompassing multiple families, with Panoplosaurus falling alongside Edmontonia , Dyoplosaurus , Hierosaurus , Scolosaurus and Stegopelta within

7260-423: The new subfamily Edmontoniinae by Loris S. Russell in 1940 , along with Edmontonia and Palaeoscincus , an equivalent classification to Gilmore's where Panoplosaurinae was used. Disagreeing with the classifications based on the work of Nopcsa, Evgeny Maleev moved Panoplosaurus into the family Ankylosauridae, which he considered to contain all ankylosaurs except Syrmosaurus . Coombs reviewed and revised

7370-405: The new subfamily Panoplosaurinae , which formed the family Nodosauridae with Nodosaurinae , Ankylosaurinae and Acanthopholinae . Nopcsa's classification of Panoplosaurinae was amended slightly by Charles W. Gilmore in 1930 , who moved Palaeoscincus into the subfamily and removed Dyoplosaurus , which was discovered to have the skull of an ankylosaurine. Panoplosaurus was then placed in

7480-527: The new subgenus Chassternbergia , remove a specimen from Edmontonia longiceps making it the new genus Denversaurus , and identify a former specimen of Panoplosaurus mirus as P. sp.. To distinguish the Panoplosaurus species from the Edmontonia complex, Bakker placed Panoplosaurus in Panoplosaurinae, and Edmontonia , Chassternbergia and Denversaurus in Edmontoniinae, uniting

7590-496: The only Late Cretaceous nodosaurid from North America. Panoplosaurinae was then revived in 1986 by Paul Sereno , who used it to unite Panoplosaurus , Sauropelta , Silvisaurus and Struthiosaurus within Nodosauridae, excluding Hylaeosaurus and Polacanthus . Robert Bakker then redescribed specimens from the Late Cretaceous of North America in 1988 , prompting him to separate Edmontonia rugosidens into

7700-412: The primitive ceratopsian Unescoceratops and the pachycephalosaurians Stegoceras validum , Stegoceras sternbergi and Stegoceras brevis are known from the Dinosaur Park Formation, though their stratigraphic correlations are uncertain. Among theropods , the dromaeosaurids Dromaeosaurus and Saurornitholestes are known, the former from a single specimen of uncertain provenance, and

7810-448: The read of the skull, where they curve inwards at the jaw articulation, and towards the front they bend inwards slightly where the predentary articulates with the dentary , with the thin predentaries meeting at the midline of the jaw. The mandibles are deepest near the rear of the skull, approximately maintaining their depth along the tooth row before narrowing sharply at the front where the predentaries are. Predentaries, which make up

7920-453: The relationships of Ankylosauridae was expanded to include many of the nodosaurids known at the time, with Panoplosaurus limited to the holotype due to a lack of consensus about referred specimens. Following further modifications and expansions, Panoplosaurus was continually found within a group including Edmontonia and also at times Animantarx , Texasetes , Denversaurus (if considered separate from Edmontonia ) and ‘’Patagopelta’’. As

8030-539: The relatively complete elasmosaurid Fluvionectes , plesiosaurs are only known from isolated and incomplete elements. Two or three true crocodilians are known, including the named genera Leidyosuchus and Albertochampsa . A rich and diverse vertebrate assemblage is known from the Dinosaur Park Formation, with the lower region, excluding the Lethbridge Coal Zone, being formed by terrestrial and coastal deposits. Microfossil sites are common, but

8140-407: The same location that may be referred to Panoplosaurus lack the distinctively short snout of the holotype, it may be that its unique appearance is due to it being a different age or sex than other individuals. The orbits of Panoplosaurus are very small, and placed far from the snout, which is very squared and bearing laterally open nostrils . Bony plates are present across the entire exterior of

8250-425: The scapula forms a shelf as it approaches the coracoid, terminating in a prominent acromion process that projects outwards from the animal, and directly overhangs a large rugose attachment area for the deltoid muscle . The scapula itself is 410 mm (16 in) long, and the coracoid is 255 mm (10.0 in). The complete fusion of the scapulocoracoid is unique to Panoplosaurus . The humerus

8360-404: The sinus cavities of Panoplosaurus increased the recovery of lost heat and moisture by over 60%. The brain of Panoplosaurus takes up 33% of the length of the skull, similar to the nodosaurid Pawpawsaurus where the value is 30%, a higher value than in ankylosaurids. Panoplosaurus had a similar sense of smell to both Pawpawsaurus and Euoplocephalus , with the ratio between the length of

8470-446: The skull and neck, to smaller, round bones underneath the chin and body, to small ossicles that filled in the spaces between other, larger osteoderms. Panoplosaurus was originally classified as a stegosaur related to the similarly armoured form Ankylosaurus , a group that was later divided with ankylosaurs becoming their own group. It was then considered close to Edmontonia in the subfamily Panoplosaurinae , but then moved into

8580-422: The skull are fused together into one single ossification, a feature unique to Panoplosaurus and Edmontonia among ankylosaurs. Following the fused atlas-axis-third cervical complex, the next three cervicals are all identifiable, but very morphologically conservative, without significant differences along the series, all vertebrae being amphicoelous . Dorsal vertebrae are poorly preserved where present, but

8690-477: The skull where the cheek plate is not in place, the mandibular teeth are hidden by the teeth of the maxilla in the upper jaw. Eight maxillary teeth are preserved, and though it is not certain that was the full tooth count there is not room for many more in the jaw. All the teeth are similar to those of Edmontonia and Palaeoscincus , with a mild expansion of the crown above the root ( cingulum ), and denticles formed by prominent ridges on both

8800-406: The skull, including large osteoderms on all the sides of the skull and mandible , small scutes within the nostrils, and underneath the head in the form of an aggregation of small ossicles covering the surface between the two lower jaws. The occipital , where the skull articulates with the neck, is short and thick, facing nearly directly downwards, which would have meant the head was held with

8910-452: The snout down in life, about 20 degrees below the horizontal. Unlike in Edmontonia , the groove separating cranial osteoderms in Panoplosaurus never disappear, which show that there is a unique narrow scute across the entire rear of the skull. Due to fusion and the covering of osteoderms, the only individual bone of the 310 mm (12 in) long mandible than can be identified is the predentary . The rami diverge strongly towards

9020-504: The species Mochlodon robustus , which he later renamed to Rhabdodon robustum in 1915. He also named Struthiosaurus transylvanicus , which he described in 1915. In 1928 he named the Teinurosaurus (meaning "extended tail lizard"). He named the turtle species Kallokibotion bajazidi , which literally means ' beautiful box of Bajazid '. The reason for this name was that the shell reminded him of Bajazid's arse. Nopcsa

9130-453: The study of the evolution of physiology and biology. Some of his works in this field exploited paleohistological data. Another of Nopcsa's theories that was ahead of its time was that birds evolved from ground-dwelling dinosaurs, which is the theory of cursorial origin of flight. He theorized that the Proavis , a theorized predecessor of birds, was running animal with forearms lifted off

9240-439: The taxa Alphadon , Eodelphis , Pediomys , and Turgidodon , and placentals by Cimolestes , Gypsonictops , and Paranyctoides , though all the material represents isolated partial skull or skeletal bones, or isolated teeth. While no other dinosaurs were recovered at the same sites as any Panoplosaurus specimens, correlation and comparisons of localities throughout the Dinosaur Park Formation show it coexisted with

9350-488: The tooth row, going from sub-vertical at the snout to nearly horizontal at the back of the mouth. This differs from ankylosaurids, but may be due to the alignment of teeth between the upper and lower jaws, instead of a difference in form of mastication . While the shape of teeth in Panoplosaurus and other ankylosaurs suggests a simple, soft food diet, their variability implies a less restrictive feeding strategy. Nodosaurid teeth are more blade-shaped than those of ankylosaurids,

9460-1387: The traditional dichotomous split between nodosaurids and ankylosaurids, their analyses resulted in a paraphyletic grade of these taxa comprising the monophyletic clades Panoplosauridae, Polacanthidae and Struthiosauridae. These results are displayed in Topology B below. Corresponding clades are shown in matching colors for clarity, and ⊞ buttons can be clicked to expand nodes: Sauroplites Mymoorapelta Dongyangopelta Gastonia Gargoyleosaurus Hoplitosaurus Polacanthus Peloroplites Taohelong Sauropelta Acantholipan Nodosaurus Niobrarasaurus Ahshislepelta Tatankacephalus Silvisaurus CPC 273 ( Aguja Formation specimen) Hungarosaurus Europelta Pawpawsaurus Borealopelta Stegopelta Struthiosaurus languedocensis Struthiosaurus transylvanicus Struthiosaurus austriacus Animantarx Panoplosaurus Patagopelta Texasetes Denversaurus Edmontonia longiceps Edmontonia rugosidens Kunbarrasaurus Paw Paw juvenile Hylaeosaurus Liaoningosaurus Crichtonpelta Chuanqilong Cedarpelta Aletopelta Later-diverging ankylosaurids Sarcolestes Franz Nopcsa von Fels%C5%91-Szilv%C3%A1s Nopcsa

9570-471: The two subfamilies into the family Edmontoniidae , which he placed in Nodosauroidea , the family Nodosauridae of Coombs' use elevated to superfamily rank. Carpenter further reviewed Late Cretaceous North American nodosaurids in 1990, concluding that Bakker's justification for separation was lacking; placing Chassternbergia and Denversaurus as junior synonyms of Edmontonia , and only narrowing down

9680-413: The type genus Nodosaurus was required as an internal specifier. In formally defining Nodosauridae, Madzia and colleagues followed the previously established use for the clade, defining it as the largest clade including Nodosaurus textilis but not Ankylosaurus magniventris . As all phylogenies referenced included both Panoplosaurus and Nodosaurus within the same group relative to Ankylosaurus ,

9790-644: The work of the Albanian Congress of Trieste , published his notes on the congress that became of particular historical interest. He left the Albanological part of his estate along with a letter of manuscripts to be published to Norbert Jokl , a renowned specialist in Albanian studies and Nopcsa's former colleague. At that time, Nopcsa's material consisted of thousands of pages of notes, sketches, and finished text. Subsequently, this library came into possession of Mid'hat Bey Frashëri . When Frashëri

9900-511: The younger Bearpaw Formation . Panoplosaurus was found about 20 to 30 m (66 to 98 ft) above boundary with the Oldman Formation, which is 76.5 million years old, within the "sandy zone" of the formation deposited by alluvial paleochannels. All the deposits of the Dinosaur Park Formation were lain during the beginning of the last significant sea level rise of the Western Interior Seaway . Sediments deposited during

10010-584: Was a province of the Ottoman Empire contending for independence in that time. He was one of the few outsiders who ventured into the mountainous areas in the north of Albania. He soon learned the Albanian dialects and customs. Eventually, he got on good terms with the leaders of the Albanian nationalist resistance who fought against the Turks in the region. Nopcsa gave passionate speeches and smuggled in weapons. In 1907 on one of his expeditions into

10120-502: Was a rather large animal at 1,600 kg (3,500 lb), a comparable size to other ankylosaurs from the same location, and heavier than or approximately equivalent to all predators it coexisted with. Panoplosaurus was about 5 m (16 ft) long, and was armoured like most ankylosaurs, but lacked prominent spikes anywhere on its body. The characteristics of its armour are one of the features that distinguishes Panoplosaurus from its relative and contemporary Edmontonia , along with

10230-529: Was a synonym of Panoplosaurus , which was considered unjustified by Kenneth Carpenter in a 1990 review on Late Cretaceous nodosaurids . The referral of ROM 1215 to Panoplosaurus was questioned by Coombs in 1990, and Roland A. Gangloff referred it and the Alaskan specimen DPMWA 90–25 to Edmontonia in 1995 based on the anatomy of the lack of diagnostic traits, but Victoria Arbour and colleagues retained ROM 1215 in Panoplosaurus in 2009 , limiting

10340-428: Was also interested in evolutionary theory, especially on macroevolution, on which the fossil record can yield relevant data. Nopcsa was also an important geologist. Indeed, Nopcsa was one of the first scholars to study the geology of the western Balkans , particularly northern Albania. Nopcsa became fascinated with Albania during his lifetime, probably through the tales of Albania's mountain tribesmen, to which he

10450-604: Was an island, Hațeg Island (now the Haţeg or Hatzeg basin in Romania) during the Mesozoic era. He theorized that "limited resources" found on islands commonly have an effect of "reducing the size of animals" over the generations, producing a localized form of dwarfism . Nopcsa's theory of insular dwarfism —also known as the island rule —is today widely accepted. Additional pygmy sauropods, named Europasaurus , were recently discovered in northern Germany. Nopcsa also created

10560-608: Was being described by Canadian paleontologist Lawrence M. Lambe of the Geological Society, who completed the description of the skull and osteoderms prior to his death in March of 1919 . Lambe's work was published posthumously by the Royal Society of Canada, where he named the new material as Panoplosaurus mirus , and a supplemental description of the vertebrae and limb material, which Lambe had not yet gotten to,

10670-679: Was born in 1877 in Déva, Transylvania , Kingdom of Hungary (today Deva, Romania ), to the Hungarian Nopcsa aristocratic family of Romanian origin. He was the son of Elek Nopcsa  [ hu ] (1848-1918), a member of the Hungarian Parliament and his wife, Matylda Henrietta Żeleński z Żelanki (1852-1938). In 1895 Nopcsa's younger sister Ilona discovered dinosaur bones at the family estate at Szacsal (today part of Sânpetru, Sântămăria-Orlea , Romania). He shared

10780-511: Was considered close to around when it was named. Panoplosaurus is from deposits slightly younger than Edmontonia rugosidens , and existed alongside hadrosaurids like Corythosaurus and Lambeosaurus , ceratopsids like Centrosaurus , and the tyrannosaurid Gorgosaurus , as well as other small dinosaurs like Stegoceras , Dromaeosaurus and Ornithomimus , and various fishes, amphibians, crocodiles and pterosaurs . In 1917 , Canadian paleontologist Charles M. Sternberg of

10890-455: Was first introduced by Louis Drašković, a man thought to be his first lover. During his lifetime Nopcsa published more than fifty scientific studies concerning Albania, covering a wide range of linguistics , folklore , ethnology , history and kanun (that is, Albanian customary law). He was one of the leading experts on Albania in his time. After Nopcsa's death, several of his important manuscripts were left unpublished. He participated in

11000-542: Was forced to flee the country, Nopcsa's materials were confiscated by the communist regime of Enver Hoxha . Eventually, Nopcsa's manuscripts, drawings, and completed writings formed the core of the Albanological section of Albania's National Library . Season 7, Episode 9 of the Bad Gays podcast covers Nopsca's life. Panoplosaurus mirus Panoplosaurus is a genus of armoured dinosaur from

11110-568: Was it published in German and it was later translated to English in 2014 as Traveler, Scholar, Political Adventurer: A Transylvanian Baron at the Birth of Albanian Independence , edited by Robert Elsie . Nopcsa's main contribution to paleontology – and hence " paleobiology " – was, that he was one of the first researchers who tried to "put flesh onto bones." At a time when paleontologists were mainly interested in assembling bones, he tried to deduce

11220-446: Was named for a nearly complete skeleton and skull from the deposits, an unnamed but new form of snapping turtle is known, Adocus and Basilemys represent the diversity of primitive trionychoids , and the softshelled turtles Apalone , Aspideretoides , and a new form, are also present. Fragmentary material from the choristodere Cteniogenys , and many well-preserved skulls and skeletons of its relative Champsosaurus ,

11330-413: Was not able to definitively identify the first sacral of Sternberg as a dorsosacral. The neural spines of all four sacrals are fused into a single plate-like process. Few caudals are known in Panoplosaurus , but where preserved they are very similar to the corresponding elements of Ankylosaurus . In Panoplosaurus the scapula and coracoid are completely fused together, with the only indication of

11440-452: Was not analysed. Expansions on the analysis of Vickaryous and colleagues resolved the same clade of Panoplosaurus and Edmontonia with the additions of the nodosaurid Hungarosaurus , the ankylosaurid Crichtonsaurus benxiensis , the nodosaurid Struthiosaurus , and the supposed ankylosaurid Tatankacephalus . The next novel analysis was that of Richard Thompson and colleagues in 2012 , combining previous ankylosaurian analyses into

11550-597: Was present during all of the duration of the formation and its over- and underlying deposits, but the mussel and brackish -water assemblages that can be seen in older and younger layers are absent in the range of 10 to 35 m (33 to 115 ft) above the Dinosaur Park Formation-Oldman Formation contact. A variety of forms of fish are present in the fluvial beds of the Dinosaur Park Formation, including chondrichthyans , teleosts and other ray-finned fishes . The ray Myledaphus

11660-523: Was published by Sternberg in 1921 . The name Panoplosaurus derived from the hoplon of Greece , translating as "well -armoured lizard". While the beds of discovery of the holotype , CMN 2759, were originally described as the Belly River beds, they are now considered part of the upper level of the Dinosaur Park Formation , pertaining to the late middle Campanian , 75 to 76 million years ago. Many additional specimens have since been referred to

11770-499: Was recovered in 1998 by Jim Kirkland and colleagues, but the 2001 analysis of Carpenter instead recovered Edmontonia closest to Animantarx , while Panoplosaurus nested next to a clade of Sauropelta and Silvisaurus . The 2003 analysis of Robert Hill and colleagues resolved a group of Edmontonia and Animantarx , with Panoplosaurus as the sister taxon, and the analysis of Matthew Vickaryous et al. recovered Panoplosaurus and Edmontonia as sisters, though Animantarx

11880-432: Was splayed, with two phalanges on the first digit, and three on the second and third. All ungual bones were hoof-shaped, lacking a point and bearing a flat bottom. The only portions of the pelvis and hindlimb that are known in Panoplosaurus are the tibia , fibula , and a few isolated bones from the foot, the remainder being unpreserved. The right tibia and fibula were found still in articulation, with

11990-472: Was standing water. The constant presence of water in the Dinosaur Park Formation led many forms of freshwater or marine animals to enter the otherwise predominantly terrestrial ecosystem. In the lower Dinosaur Park Formation, assemblages of crevasse sites show that mollusks were commonly dominated by the freshwater clam Sphaerium , which occurred with abundant gastropods of the genera Goniobasis and Lioplacodes . This sphaeriid-dominated diversity

12100-767: Was that of Rivera-Sylva et al. ( 2018 ), which is a modified version of the Arbour and Currie analysis expanded to include the Mexican taxon Acantholipan . Additional modifications to the analysis were done by Soto Acuña and colleagues in 2021 and 2024, with Panoplosaurus maintaining a position close to Edmontonia . Clades follow Madzia and colleagues. Peloroplites Sauropelta Borealopelta Tatankacephalus Nodosaurus Struthiosaurini Animantarx Panoplosaurus Texasetes Edmontonia rugosidens Edmontonia longiceps Denversaurus Ankylosaurs were traditionally viewed as having

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