Showing posts with label Ornithischian Dinosaurs. Show all posts
Showing posts with label Ornithischian Dinosaurs. Show all posts

Sunday, 18 June 2023

Vectipelta barretti: A new species of Ankylosaurian Dinosaur from the Early Cretaceous of the Isle of Wight.

The Ankylosaurs are Ornithischian Dinosaurs noted for their broad body, short limbs and extensive covering of dermal armour on the head, body and tail. They are a diverse group, found on all continents, but reaching their greatest diversity on the continents of the former Laurasian landmass, i.e. North America, Europe, and Asia. They split from their sister group, the Stegosaurs, in the Early or early Middle Jurassic, but appear to have remained a minor component of most ecosystems until the Early Cretaceous, when they underwent a significant radiation, remaining a significant member of the herbevore community until their demise in the End Cretaceous extinction. 

A number of Ankylosaurs have been described from the UK, from the Middle Jurassic onwards, although the group reach their maximum abundance in the UK in the Early Cretaceous Wealden Group of southern England. For a long time, all Ankylosaurs from the Wealden Group were referred to two species, Polacanthus foxii and Hylaeosaurus armatus, but this is now thought likely to hide considerable diversity, as the Wealden Group is thought to cover at least 8 million years of time, including all of the Barremian Stage and part of the Aptian.

Traditionally, all Ankylosaur material from the Isle of Wight was referred to Polacanthus foxii, but a recent study of Ankylosaur diversity within the Wealden Group, led by Thomas Raven of the Fossil Reptiles, Amphibians and Birds Section at the Natural History Museum, has suggested that this name should be restricted to the holotype specimen, NHMUKPV R175, and other specimens should be considered taxonomically uncertain until re-examined. One notable specimen examined in Raven et al.'s study was IWCMS 1996.153, known as the Spearpoint Ankylosaur, which was shown to be particularly different to the Polacanthus foxii holotype, and not likely to be closely related.

In a paper published in the Journal of Systematic Palaeontology on 15 June 2023, Stuart Pond, also of the Fossil Reptiles, Amphibians and Birds Section at the Natural History Museum, Sarah-Jane Strachan of the Department of Earth Sciences at University College London, Thomas Raven (again), Martin Simpson of Lansdowne on the Isle of Wight, Kirsty Morgan of Fayetteville in Arkansas, and Susannah Maidment, again of the Fossil Reptiles, Amphibians and Birds Section at the Natural History Museum, formally describe the Spearpoint Ankylosaur as the holotype of a new species.

The Wessex Group consists of a series of terrestrial, lacustrine, flu-vial and lagoonal deposits which outcrop in the Wessex Sub-basin of the Isle of Wight and the Weald Sub-basin of south-east England. On the Isle of Wight the group comprises two formations, the Wessex Formation, considered Barremian in age, and the Vectis Formation, thought to be late Barremian to early Aptian. The Wessex Formation is a sequence about 180 m thick, consisting of variegated mudstones and interbedded sandstones deposited as river channel, floodplain and point bar deposits by a high-sinuosity river system flowing west to east. The overlying Vectis Formation is thought to have been laid down in a shallow lagoon. The largest exposures of both formations are on the southwestern coast of the Isle of Wight, with smaller exposures at Yaverland on the south-east coast.

The Wessex Formation is thought to have been laid down in a Mediterranean climate, with distinctive dry and wet seasons, the latter associated with periodic flood events in which debris flows including large trees and Dinosaur remains were swept down from the (now lost) highlands from which the river descended, and were deposited on the flood plain. This formation is rich in fossils, producing numerous Plant (particularly Gymnosperms), Invertebrates, trace fossils, and a diverse Vertebrate assemblage, including Bony Fish, Turtles, Crocodilians, Pterosaurs and Dinosaurs.

The Spearpoint Ankylosaur was discovered within the Compton Chine to Steephill Cove Site of Special  Scientific Interest, about 50 m to the west of Chilton Chine. Chines on the Isle of Wight, are gullies which cut into the overlying chalk grassland by seasonal streams, and emerging from the cliffs as waterfalls. These features never erode down to beach level, as the cliffs are eroded backwards by the action of the sea faster than the chines are eroded downwards by the streams, and form a unique environment with their own distinctive flora and fauna, including many rare Plants and Insects.

In November 1993 fossil hunter Gavin Leng found several Ankylosaur vertebrae on beach, which he later donated to the Dinosaur Isle Museum. The following spring another fossil hunter Lin Spearpoint, discovered the remainder of the skeleton in the cliffs above, on land belonging to dairy farmer Richard Fisk. Fisk gave permission for Lin Spearpoint, Dick Spearpoint, and Martin Simpson, to excavate the skeleton, which was then prepared by Lin Spearpoint, housed at the University of Southampton, where it was the subject of several MSci, MRes and PhD projects, some of which were carried out by Stuart Pond, Sarah-Jane Strachan, and Kirsty Morgan, temporarily displayed at the SeaCity  Museum,  Southampton, and eventually acquired by the Dinosaur Isle Museum. 

Pond et al. formally describe the Spearpoint Ankylosaur as the holotype of a new species, Vectipelta barretti, where 'Vectipelta' means 'Vectis-shield' (Vectis was the name used by the Romans for the Isle of Wight), and 'barretti' honours Paul Barrett of the Natural History Museum, in recognition of his major and ongoing contributions to Dinosaurian Vertebrate palaeontology, and his importance to Stuart Pond, Thomas Raven and Susannah Maidment as a mentor, supervisor, colleague and friend.

Geological setting and stratigraphy. (A) Map of Isle of Wight with the extent of Wessex Sub-basin marked in orange, and the site of the Vectipelta barretti (IWCMS 1996.153 and IWCMS 2021.75) excavation indicated by the black arrow. (B) Schematic stratigraphical log of the outcrop between Sudmore Point and Ship Ledge showing the approximate stratigraphical location of Vectipelta barretti (IWCMS 1996.153 and IWCMS 2021.75). Pond et al. (2023).

The new species is described from two specimens, both believed to have come from the same individual and therefore treated as one. WCMS1996.153 comprises portions of four cervical vertebrae and six  free dorsal vertebrae discovered on the foreshore by Gavin Leng in 1993, while IWCMS 2021.75 comprises five fused dorsal vertebrae, forming the sacral rod, fragments of several dorsal ribs, a fused sacrum made up of four and a half fused vertebrae, thirteen caudal vertebrae, and fragments of four more, the proximal part of the left scapula, the proximal and distal ends of the left humerus, a possible ulna fragment, multiple pieces of the iliosacral block, including parts both the left and right ilia, both pubes, fused to the anteromedial margins of the acetabula, a more-or-less complete left ischium, some possible fibula fragments, and second metatarsal, the sacral shield, a large number of osteoderms, and three recurved spines. Specimen IWCMS 2021.75 was found lying on its back, with its (missing) head pointing towards the cliff face, leading to the assumption that the cervical and upper dorsal vertebrae of WCMS1996.153 came from the same individual. Pond et al, note that several years prior to the discovery of WCMS1996.153, a beach-rolled Ankylosaur braincase was found at the same location by fossil hunter David Cooper, and acquired by the University of Cambridge's Sedgwick Museum, and a partial femur and posterior part of an ilium were found nearby by David Richards in the 1990s and donated to the Natural History Museum. These specimens may also relate to the Spearpoint Ankylosaur, but all are heavily eroded, making it impossible to assess them with any confidence, so they were excluded from the study.

Holotype material of Vectipelta barretti, IWCMS 1996.153 and IWCMS 2021.75 in (A) dorsal view; (B) left lateral view; (C) dorsal view of the osteoderms only; (D) complete skeletal reconstruction, in left lateral view. Scale baris 1m. Pond et al. (2023).

The recurved spines of the Spearpoint Ankylosaur are notable, as NHMUKPV R175, the holotype of Polacanthus foxii, lacks any such structure, supporting the idea that the two are quite different Animals. Some spines assigned to Polacanthus sp. have been described from Early Cretaceous of Soria, Spain, but these are also unlikely to derive from that genus. Recurved spines are known in a wide range of Ankylosaurs, including the other Wealden Group species, Hylaeosaurus armatus, but the form of the Spearpoint Ankylosaur's spines is different to any of these, supporting the idea that it should be described as a new species.

Vectipelta barretti, IWCMS 2021.75. Examples of recurved spines. Orientation presumed: (A)–(C) proximal; (D)–(F) distal; (G)–(I) left lateral; (J)–(L) right lateral; (M)–(O) anterior; (P)–(R) posterior. Scale bar is 10 cm. Pond et al. (2023).

Understanding the diversity of Early Cretaceous Ankylosaurs is not just important because we are interested in Dinosaurs; it has implications for our understanding of Mesozoic ecosystems and geology as a whole. The Jurassic is believed to have ended with a mass extinction event, although the nature of this is poorly understood. Terrestrial deposits with good fossil preservation are rare from the Early Cretaceous, leading to the apparent emergence of a quite new ecosystem in the Middle Cretaceous. However, the clustering of multiple specimens from diverse groups under a few poorly constrained species names in Early Cretaceous deposits, as has happened with Wealden Ankylosaurs (and, it has also been suggested, Iguanodontids), could be hiding a more gradual faunal turnover during this interval.

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Saturday, 10 June 2023

Iani Smithi: A new species of Rhabdodontomorph Iguanodontian Dinosaur from the Cretaceous Cedar Mountain Formation of Utah.

The Cenomanian (Earliest Late Cretaceous) Mussentuchit Member of the Cedar Mountain Formation, which outcrops in central Utah, is one of the most abundant Late Cretaceous bonebeds known from anywhere in the world, with more than 100 species described to date, including Tyrannosaurids, Dromaeosaurs, Velociraptors, Troodontids, Therizinosaurs, Titanosauromorphs, Thescelosaurids, Pachycephalosaurids, Neoceratopsians, Nodosaurids, Hadrosaurids, and non-Hadrosaurid Iguanodontians, as well as a number of less well defined specimens.

In a paper published in the journal PLoS One on 7 June 2023, Lindsay ZannoTerry Gates, and Haviv Avrahami of Paleontology Research at the North Carolina Museum of Natural Sciences, and the Department of Biological Sciences at North Carolina State UniversityRyan Tucker of the Department of Earth Sciences at Stellenbosch University, and Peter Makovicky of the Department of Earth and Environmental Sciences at the University of Minnesota, describe a new species of Rhabdodontomorph Iguanodontian Dinosaur from the Cedar Mountain Formation.

The new species is described upon the basis of a single partial skeleton (specimen NCSM 29373) comprising a largely complete, disarticulated skull; cervical, dorsal, sacral, and caudal vertebrae; associated ribs and haemal arches; and portions of the right and left pectoral girdle, left pelvic girdle, right forelimb, and right hindlimb. The elements of the briancase are incompletely fused, leading Zanno et al. to conclude that the specimen is immature. The specimen was recovered from an exposure of the Mussentuchit Member in Emery County, Utah, 1 m above the boundary with the Ruby Ranch Member, and 7 m below Mussentuchit Ash Zones 1, which has been dated to 99.490 million years before the present. The new species is named Iani smithi, where 'Iani' refers to the Roman god Ianus (or Janus), who presides over boundaries, in reference to the fact that the mid-Cretaceous was a time of biological transition in western North America, and 'smithi' honours Joshua Aaron Smith for his contributions to the discovery and conservation of paleontological resources in the region, particularly early explorations by the North Carolina Museum of Natural Sciences.

Location of holotype locality for Iani smithi (NCSM 29373). (A) Global map showing location of Mussentuchit Member outcrop in central Utah, western North America, and a stratigraphic section at the quarry with dated ash horizons; and (B) graphical representation of preserved skeletal elements of the holotype specimen (NCSM 29373). Preserved elements are coloured on the left facing skeletal whether they derive from the right or left side of the body. Exact positions of chevrons and ribs unknown due to poor preservation. See text and figures for specific positioning and completeness of elements. Abbreviations: MAZ1–4, Mussentuchit Ash Zones 1–4. Zanno et al. (2023).

Most of the skull is present, but the missing elements include the nasal and maxilla bones, so that the length and shape of the face of Iani smithi can only be estimated from the length of the mandible. This mandible comprises fragmentary predentary, complete pair of dentaries, right surangular, partial splenial, and several isolated dentary teeth. The rear part of the premaxila is thickened and shows the alveoli (sockets) of three premaxilary teeth. Premaxilary teeth are unusual in Ornithopod Dinosaurs, with most groups having lost them early in their history, but they are known in several other Rhabdodontomorphs.

Three-dimensional reconstruction of the skull of Iani smithi (NCSM 29373). (A) right lateral and (B) medial views of skull with left facial bones removed; skull reconstruction with all preserved elements (some mirrored) in (C) caudal, (D) dorsal, (E) ventral, and (F) rostral views. Scale bar 5 cm. Zanno et al. (2023).

The dentition of Iani smithi appears similar the other Rhabdodontomorphs, and non-Rhabdodontomorph early-branching Ornithopods such as Tenontosaurus and Qantassaurus. All of the teeth on the left dentary are still present, although the crowns of the middle teeth are damaged. Five teeth are still present in the left dentary, and thirteen loose teeth were found with the skeleton, interpretted as eight maxillary teeth, five dentary teeth, and a tooth of indeterminate origin. A single predentary tooth is present.

Predentary, dentary, and surangular of Iani smithi (NCSM 29373). Predentary in (A) right lateral, (B), rostral (right half mirrored and combined to show approximate shape of complete element), (C) ventral, and (D) dorsal views. Right dentary in (E) lateral view; left (pathological) dentary in (F) medial, (G) lateral, (H) dorsal, and (I) ventral views; right surangular in (J), lateral and (K), medial views. Abbreviations: a.co, estimated coronoid articulation; ad, accessory denticle; a.pd, predentary articulation; a.sr, surangular articulation; de, dentary; dp, dorsal process; dpa, dentary parapet; f, foramen; lg, lateral groove; mkc, Meckelian canal; ml, midline; path, pathological bone; pd, primary denticle; pathological bone; pr?, prearticular?; prd, peripheral denticle; sb, surangular boss; te, teeth; tr, exposed tooth root; vp, ventral process. Colour annotation: white, depressions/fossae/grooves; green, articular surfaces; blue circles, foramina; light blue lines, marginal contours; peach dashed lines, ridges/internal contours; light purple, dentition. Scale bar 5 cm. Zanno et al. (2023).

Portions of at least seven cervical (neck) vertebrae are present, although the atlas vertebra (which connects the spine to the skull) is absent. The axis (second vertebra) is present, and is compressed dorsoventrally, somewhat eroded ventrally, and approximately 150% longer than wide. At least eight unfused neural arches from the dorsal part of the spine are preserved, along with two partially fused dorsal vertebrae are preserved in variable states of damage and distortion. The dorsal vertabrae appear to be shorter and more gracile towards the head. Six sacral neural arches and four isolated sacral centra are present, showing the sacrum to have been unfused. The first seven caudal (tail) vertebrae are also present. 

Sacral vertebrae of Iani smithi (NCSM 29373). Neural arch of S1 in (A) dorsal and (D) cranial views; (B) sacral neural arches 1–6 in left lateral view; (C) neural arch of S6 in caudal view; (E) neural arch of S2 in cranial view; (F) neural arch of S4 in cranial view; (G) neural arch of S5 in cranial view; (H) neural arch of S4 in caudal view; Sacral centrum S1? in (I) left lateral, (J) caudal, and (L) cranial views; Sacral centra S2?–5? in (K) left lateral and (N) ventral views; (M) sacral centrum S2? In cranial view; (O) sacral centrum S5? in caudal view. Abbreviations: ipodf, infrapostdiapophyseal fossae; ns, neural spine; pop, postzygapophysis; prz, prezygapophysis; vk, ventral keel. Colour annotation: white, depressions/ fossae/grooves; green, articular surfaces; blue circles, foramina; light blue lines, marginal contours; peach dashed lines, ridges/internal contours. Scale bar 5 cm. Zanno et al. (2023).

At least fourteen dorsal ribs are present, these being more robust and strongly bowed than in other Rhabdodontomorphs. Both scapulae are preserved, these being stout and robust, but within the range of other Rhabdodontomorph species. An incomplete right humerus is present, with most of the proximal portion preserved, with the exception of most of the deltopectoral crest, but the distal portion is missing. The right ulna is well preserved and undistorted. A portion of a radius is also present, though this is highly eroded and lacks both ends, making it unclear which side it came from. Four elements of a manus (hand) are present; these are poorly preserved and somewhat compressed, making identification difficult, although they may be two metacarpals and penultimate and ultimate (ungual) phalanges of digits 1, 2, or 3. Few early-branching Ornithopods have well preserved manus, making comparison between these elements difficult. The shaft and distal end of the left ischium are preserved; the shaft is generally straight, but has a twist towards the distal end. The right femur is preserved, generally in good condition, but with some erosion in the region between the greater and lesser trochanters. 

Pelvic girdle and hind limb of Iani smithi (NCSM 29373). Right ischium in (A) dorsal, (B) ventral, and (C) medial views. Right femur in (D) medial, (E) lateral, (F) proximal, and (G), distal views. Abbreviations: fh, femoral head; ft, fourth trochanter; lc, lateral condyle; mc, medial condyle; op, obturator process. Colour annotation: white, depressions/fossae/grooves; green, articular surfaces; blue circles, foramina; light blue lines, marginal contours; peach dashed lines, ridges/internal contours. Scale bar 10 cm. Zanno et al. (2023).

Zanno et al. carried out several phylogenetic analyses, using a number of different matrices, which consistantly found that Iani smithi is closely related to the Rhabdodontomorphs and the non-Rhabdodontomorph Tenontosaurus, with all analyses bar one placing Iani smithi within the Rhabdodontomorphs.

Hypothesized evolutionary relationships of Iani smithi. Strict consensus tree of the Poole Matrix using (A) maximum parsimony optimality, and (B) Bayesian inference, and strict consensus tree of the Dieudonné et al. Matrix using (C) maximum parsimony optimality, and (D) Bayesian inference. Rhabdodontomorph taxa in green. Bremer support values (left) and posterior probabilities (right) shown. Grey names reflect taxa outside the illustrated clade for a particular analysis, or in the case of Dryosauridae (an ad hoc combined Operational Taxonomic Units with variable composition using different analytical approaches). Zanno et al. (2023).

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Wednesday, 1 June 2022

A Thyreophoran osteoderm from the Early Jurassic of northeastern Germany.

Early Jurassic Dinosaur fossils are rare in Europe, with only a few specimens known from Dorset in England, central Portugal, and several sites in Germany. One of the German sites which produces Dinosaur material is the Grimmen Clay Pit in Mecklenburg-Western Pomerania, northeastern Germany. This site was formerly worked for clays for the aggregates industry, and comprises a detached block of Lower Toarcian clay within a Pleistocene glacial sequence. This block has been badly distorted, making it hard to establish an internal stratigraphy, but contains fossiliferous concretions, which have produced numerous Ammonites (which have been used to establish its age), as well as marine, and more rarely terrestrial, Vertebrate material. 

In a paper published in the journal Paläontologische Zeitschrift on 19 January 2022, Marco Schade of the Institute of Geography and Geology and Zoological Institute and Museum at the University of Greifswald, and the Department of Earth and Environmental Sciences at  Ludwig-Maximilians-Universität, and Jörg Ansorge, also of the Institute of Geography and Geology at the University of Greifswald, describe an osteoderm from a Thyreophoran Dinosaur from the Grimmen Clay Pit.

The osteoderm, given the catalogue number GG 504, was discovered in several pieces by Jörg Ansorge in 2017, and subsequently prepared by him. The internal structure of the bone examined by micro-computed tomography at the Department of Cytology and Evolutionary Biology at the University of Greifswald.

GG 504, osteoderm of an unknown Thyreophoran Dinosaur. (a) Top view of the slightly convex side; (b) view on the acute edge, showing the inclination of the osteoderm and its base towards the less convex side; (c) cross section of the proximodistal mid-length; d, intertrabecular spaces filled with aragonite; (e) magnified part of a cross section. Abbreviations: ae, acute edge; al, aragonitic layers; be, blunt edge; co, cortex. Schade & Ansorge (2022).

The specimen comprises five separate fragments, which can be assembled to form a single piece with a maximum dimension of 15.5 cm. From the overall shape of this piece, Schade and Ansorge calculate that it represents about 75% of the original bone. Distinct striations can be seen on both surfaces of the specimen, appearing wrinkled under a light microscope, though these are rougher and more irregular on the more convex side. The osteoderm is thickest in the middle of what Schade and Ansorge assess to be its forward edge, where it is about 2 cm thick, tapering to the rear and either side. Both sides of the bone are convex, although one more than the other, with the specimen being slightly bent and the more convex side forming the inner part of the bend. The osteoderm appears to have originally been sub-parallel along much of its length, tapering towards the rear.

GG 504, osteoderm of an unknown Thyreophoran Dinosaur. (a) Top view of the pronounced convex side; (b) base in proximal view; (c) close-up of striated cortex shown in (a); (d) magnified cortex; (e) close-up of section close to the base shown in (a). Abbreviations: co, cortex; ob, offset base; sd, slight depression. Schade & Ansorge (2022).

The bulk of the osteoderm appears to be made up of Haversian bone (bone made up of a series of parallel tubes, which in life would have held blood vessels), surrounded by a thin, laminated cortex. These laminae are separated by layers of aragonite in places, and appear to show lines of arrested growth.

GG 504, osteoderm of an unknown Thyreophoran Dinosaur. (a) Magnified section of the distal-most fragment, arrow points towards former apex; (b) close-up of cortex from the pronounced convex side shown in (a), arrows without lines hint towards potential lines of arrested growth; (c) close-up of cortex from the slightly convex side shown in (a). Abbreviations: ?ShF, potential Sharpey’s fibres; ?so, potential secondary osteons; vc, vascular cavities. Schade & Ansorge (2022).

Specimen GG 504 resembles a previously described specimen, GG 85/1 23, which has been assigned to Emausaurus, an early diverging Thyreophoran Dinosaur (that is to say, an early Thyreophoran Dinosaur that cannot be confidently assigned to either of the main Thyreophoran groups, the Stegosaurs and Ankylosaurs, and which presumably lived before those groups diverged), and which was also recovered from the Grimmen Clay Pit. However, GG 504 is significantly larger than GG 85/1 23 (15.5 cm vs 4.5 cm), and GG85/1 23 is the largest known osteoderm associated with Emausaurus. Furthermore, GG 85/1 23 has a rougher, more vascularised base than GG 504, and a less symmetrical shape.

GG 504 also shows some resemblances to osteoderms associated with Scutellosaurus lawleri, another early diverging Thyreophoran from the Early Jurassic of Arizona, although again it is significantly larger than any osteoderm associated with this species (Scutellosaurus lawleri is only thought to have reached about 120 cm in total length).

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Monday, 21 March 2022

Torosaurus in Canada.

The Ceratopsids, an iconic group of Dinosaurs which were an important part of the faunas of Late Cretaceous North America and Asia, were large, quapruped herbivores with distinctive neck frills and facial horns over their eyes and noses. One of the largest Ceratopsids was Torosaurus, known from the End Cretaceous of North America, which is thought to have weighed up to 6500 kg, and achieved skull lengths of about 3 m. However, Torosaurus, is similar to Triceratops, another large End Cretaceous Ceratopsid, which has led to an active debate in recent years, as to whether the two genera are in fact the same, and, therefore, whether or not Torosaurus is a valid name at all (in taxonimy, where a group has been named twice, the older name is considered to have priority). This argument derives from the fact that all known specimens of Torosaurus are as large as, or larger than, the largest specimens of Triceratops, while the horns and crest of Torosaurus are essentially a more exagerated variant in the pattern seen in Triceratops, which might imply that Torosaurus fossils represent more mature specimens of Triceratops, rather than a separate species.

To date, there have been two reports of Torosauraus specimens being found in Canada, both of fragments of frills that have not fully been described. The first, EM P16.1, was recovered from the Frenchman Formation of southern Saskatchewan in 1947 by Charles Morey, upon whose land it was found, and local amateur palaeontologist Harold ‘Corky’ Jones. This specimen was briefly described and illustrated by palaeontologist Tim Tokaryk in 1986, who attributed the specimen to Torosauraus, but this diagnosis has since been disputed, and it has been suggested that the specimen might in fact belong to Arrhinoceratops, which can be hard to differentiate from Torosaurus. The second specimen, UALVP 1646, was excavated from the lower Scollard Formation in the Red Deer River Valley of southern Alberta by Charles Stelck in 1964, and has been mentioned as an example of Torosaurus in a number of publications, but never formally described or illustrated.

In a paper published in the Zoological Journal of the Linnean Society on 1 March 2022, Jordan Mallon of the Beaty Centre for Species Discovery and Palaeobiology Section at the Canadian Museum of Nature, and the Ottawa-Carleton Geoscience Centre and Department of Earth Sciences at Carleton University, Robert Holmes, also of the Beaty Centre for Species Discovery and Palaeobiology Section at the Canadian Museum of Nature, and of the Department of Biological Sciences at the University of Alberta, Emily Bamforth of the T. rex Discovery Centre at the Royal Saskatchewan Museum, and Dirk Shuman of Fibics Incorporated, formally describe specimens EM P16.1 and UALVP 1646, and discuss their attribution to Torosaurus and the implications of this to the ongoing Torosaurus/Triceratops debate.

The site where specimen EM P16.1 was collected is no longer accessible, but is believed to be an outcrop of the upper-half of Frenchman Formation along the Frenchman River, making it uppermost Maastrichtian in origin (i.e. from the very end of the Cretaceous). Shortly after it's discovery, Harold Jones wrote a letter to Charles Sternberg of the Geological Survey of Canada in which he described, in addition to the crest fragment, the associated discovery of two large bones whicj he believed to be the Animal's femurs, which were in good condition and each measured 46" in length and 9" in diameter at the thinnest part (117 cm long and 23 cm in diameter). The subsequent fate of these bones has not been recorded, however, the Eastend Historical Museum, which houses EM P16.1, also hosts some partial, poorly preserved limb material bearing the field number B9, which is also the field number assigned to EM P16.1, suggesting that these are the femurs mentioned by Jones. Much of Jones' collection is known to have been damaged in 1952, when the basement of an old schoolhouse in which it was housed was flooded, so the disparity in condition between Jones' description and that of the Eastend Historical Museum material is not surprising.

 
Femoral material pertaining to Torosaurus cf. Torosaurus latus (EM P16.1). (A) Proximal portion of right femur, medial view; (B) distal portion of left femur, cranial view; (C) proximal portion of left femur, cranial view; (D) proximal diaphysis of left femur, transverse view; (E) proximal portion of left femur, caudal view. Red arrows indicate where left femur was sampled for osteohistology. Mallon et al. (2022).

The second specimen, UALVP 1646, was uncovered in a quarry at a quarry in the Red Deer River Valley in 1964, with its current assignment to the lower Scollard Formation being based upon a description that it was found approximately 8 m below the Nevis coal seam.

 
Locality information pertinent to Canadian Torosaurus specimens. (A) Map showing locations of EM P16.1 and UALVP 1646 (stars). (B) stratigraphic context of EM P16.1 and UALVP 1646, with distribution of Hell Creek Formation Torosaurus for context. Note that the precise placement of EM P16.1 within the Frenchman Formation is unknown. The precise location of MPM VP6841 within the upper Hell Creek Formation is likewise uncertain. Abbreviations: Fm, Formation; K-Pg, Cretaceous-Palaeogene boundary; Mtn, Mountain. Mallon et al. (2022).

Mallon et al. identified the three remaining fragments of limb bone associated with specimen EM P16.1 as the proximal portion of the right femur, the distal end of the left femur missing the condyles, and the proximal metaphysis and diaphysis of the left femur (excluding the fourth trochanter, which is missing). This third piece appeared to be the least damaged, although it shows signs of having been hastily repaired in places with plaster and shellac, and from this specimen Mallon et al. took a sample from the caudolateral margin of the metaphysis and caudal left femoral diaphysis for osteohistological sampling.

The frill of specimen EM P16.1 was reconstructed with plaster and clay shortly after being extracted, and previous descriptions of the specimen have been based upon that reconstruction, even though it was known to be inaccurate. The specimen was re-prepared by experts from the Royal Saskatchewan Museum in the early 2000s, and Mallon et al. present a new description of the specimen based upon that preparation.

The frill of EM P16.1 is mostly complete, although the left squamosal is heavily reconstructed with plaster, and the median parietal bar is missing entirely. The frill is of moderate dimensions, being somewhat larger than that of Arrhinoceratops brachyops, but smaller than the largest Torosaurus frills, particularly in the transverse dimensions. The frill is nearly square in outline, being just 1.1 times wider than long, as in the holotype of Arrhinoceratops brachyops. The frill is notably more triangular in the holotype of Torosaurus latus (YPM 1830, excavated from the Lance Formation of southern Wyoming by John Bell Hatcher in 1891), which is 1.79 times wider caudally than rostrally (although the parietal of this specimen is heavily reconstructed). The width of the transverse parietal bar of EM P16.1 is among the smallest known for Torosaurus, and most nearly approximates that of ANSP 15192 (which is 1225 mm). The frill is flat in the transverse plane, not saddle-shaped as in Triceratops, although this flattening may be exaggerated by taphonomic compression.

 
Parietosquamosal frill of Torosaurus cf. Torosaurus latus (EM P16.1.). (A) Dorsal surface; (B) interpretive reconstruction of dorsal surface; (C) schematic transverse cross-section of squamosal near caudal edge, based on sketch by Harold Jones; (D) schematic of frill, showing paired fossae and channels on ventral surface of squamosals, based on sketch by H. Jones. Abbreviations: p, parietal; pf, parietal fenestra; sq, squamosal; sqaf, accessory fenestra of squamosal; sqb, squamosal bar. Mallon et al. (2022).

Specimen UALVP 1646 consists of a partial parietal, including portions of the median and caudal bars. As preserved, the maximum dimensions of the specimen are 513 mm rostrocaudally by 568 mm transversely. The parietal is weakly bowed dorsally, indicating that the frill was originally saddle-shaped, albeit not to the extent seen in Triceratops. The partial midline parietal bar is thickened (20 mm dorsoventrally) and bears three sagittally aligned bumps dorsally, similar to those reported in other Torosaurus. The anterior-most of these is most prominent; the remaining two become progressively broader and lower toward the caudal margin of the parietal. Portions of the finished margin of the left parietal fenestra are preserved. The fenestra seems to have been small and probably had a maximum diameter of not much more than 15 cm. The bone surrounding the fenestra is thin (8 mm), thickening outwardly in all directions. The caudal margin of the parietal is 21–25 mm thick (compared to 10–19 mm thick in YPM 1831) and lacks a median emargination. The parietal bears three epiparietals co-ossified to the frill. The interstitial sutures are visible. These epiparietals are interpreted as the left and right P1s and the left P2. There is no epiossification straddling the midline.

 
Parietal of Torosaurus cf. Torosaurus latus (UALVP 1646). (A) Dorsal surface; (B) interpretive reconstruction of dorsal surface; (C) ventral surface; (D) interpretive reconstruction ventral surface; (E) detail of left parietal fenestra. Arrows in (B) and (D) indicate epiparietal loci. Arrows in (E) indicate unbroken margin of parietal fenestra. Abbreviations: mb, median bump; pf, parietal fenestra; pfo, parietal fossa. Mallon et al. (2022).

Triceratops is known to have ranged into Canada, but the question of whether-or-not the closely related Torosaurus also did so has until now remained open; two specimens found in Canada have previously been referred to Torosaurus, but only one of these has previously been (briefly) described, and both have had their reference to the genus questioned. Mallon et al.'s redescription of EM P16.1 rules out the exclusion of that specimen from Torosaurus, on the basis that all of the features used to reject that hypothesis were ether artefacts of the way in which the specimen had been preserved, or were also present in the holotype of Torosaurus, if not in every specimen attributed to that taxon. Specimen UALVP 1646 is less complete than specimen EM P16.1, but again has no features which preclude its inclusion within the genus Torosaurus. Mallon et al. note that specimens assigned to Torodsaurus do seem to be somewhat variable in some features, and note that Triceratops, which is closely related, is considered to have undergone some species turnover during the 200 000 years represented by the Hell Creek Formation, and it would not be unreasonable to expect a similar level of turnover in Torosaurus.

Mallon et al. also note that the frill of specimen EM P16.1 is considerably smaller than seen in many other Torosaurus specimens, although the associated femora were of comparable size to the largest known Triceratops specimens. They note that size is not always a reliable indicator of maturity in Dinosaurs, but that EM P16.1 shows a number of morphological and osteological features which lead them to conclude that it was not a mature specimen. The existence of a sub-adult Torosaurus specimen of comparable size to the largest Triceratops specimens leads them to conclude that it is unlikely that Triceratops represents an immature form of Torosaurus.

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Saturday, 5 March 2022

Bashanosaurus primitivus: A new species of Stegosaur from the Middle Jurassic of Chongqing Municipality, China.

The Stegosaurs were a distinctive group of Ornithischian Dinosaurs whose fossils are known from the Middle Jurassic to the Early Cretaceous, and which are distinctive for the presence of a double row of plates or spines which run along the back from the neck to the tip of the spine. The earliest known Stegosaurs appear in the Middle Jurassic in Europe, China, Argentina, and Morocco, a distribution which implies the group emerged some time before the oldest fossils we know.

In a paper published in the Journal of Vertebrate Paleontology on 3 March 2022, Dai Hui and Li Ning of the Chongqing Laboratory of Geoheritage Protection and Research at the Chongqing Bureau of Geological and Mineral Resource Exploration and Development, Susannah Maidment of the Department of Earth Sciences at the Natural History Museum, Wei Guangbiao  of the Chongqing Institute of Geological Survey, Zhou Yuxuan, Hu Xufeng, Ma Qingyu, Wang Xunqian, and Hu Haiqian, also of the Chongqing Laboratory of Geoheritage Protection and Research, and Peng Guangzhao of the Zigong Dinosaur Museum, describe a new species of Stegosaur from the Middle Jurassic Lower Member of the Shaximiao Formation of Chongqing Municipality, China. 

The new species is described upon the basis of material from the Dinosaur Quarry at Laojun Village, a site discovered in 2017, and which has been excavated systematically to form a wall of Dinosaur fossils 150 m long and 8 m high. The Lower Member of the Shaximiao Formation comprises grayish-yellow massive fine-grained lithic feldspathic sandstones and purplish red silty mudstones in multiple rhythmic layers of uneven thickness; the siltstones contain calcareous nodules. This 'redbed' facies is interpretted to indicate a semi-arid, terrestrial paleoenvironment. Various attempts have been made to date this bed radiometrically, yielding dates of between 158 and 172 million years old.

 
Maps showing the location where Bashanosaurus primitivus was recovered. (A) Sketch map of China showing the location of Chongqing Municipality. (B) sketch map of Yunyang County. (C) Sketch map of the fossil locality. (D) Stratigraphic chart with Lower Member of the Shaximiao Formation in Yunyang County, Chongqing Municipality, China. Dai et al. (2022).

The new species is named Bashanosaurus primitivus, where 'Bashanosaurus' is a combination of 'Bashan-', an ancient name for Chongqing, and '-saurus', meaning 'Lizard', a comon suffix for Dinosaur names, and 'primitivus' refers to the fact that a phylogenetic analysis revealed the species to be the most primitive member of the Stegosauria (that it is to say it is the earliest branching member of the group, with all other Stegosaurs being more closely related to one-another than they are to Bashanosaurus primitivus). The new species is described from three specimens, CLGPR V00006-1, V00006-2, and V00006-3, all from the 'wall' of Dinosaur specimens from the Dinosaur Quarry at Laojun Village.

 
(A) Photograph (top) and interpretative sketch (bottom) of Bashanosaurus primitivus (CLGPR V00006-1) on the west side of the Dinosaur Fossil Wall. (B) Photograph (top) and interpretative sketch (bottom) of Bashanosaurus primitivus (CLGPR V00006-2) in the middle of the Dinosaur Fossil Wall. Abbreviations: cav, caudal vertebra; co, coracoid; dv, dorsal vertebra; f, femur; fl, fibula; mt, metatarsal; pl, plate; r, rib; sc, scapula; sp, spine; t, tibia. Dai et al. (2022).

Specimen CLGPR V00006-1 is designated the holotype of the species (that is to say, any other specimen which is judged to be the same species as CLGPR V00006-1 is therefore considered to be a specimen of Bashanosaurus primitivus). This specimen is preserved on the west side of the wall of Dinosaur fossils and comprises one dorsal vertebra, two caudal vertebrae, right scapula, right coracoid, left femur, left tibia, left fibula, one metatarsal, three pieces of dermal armor (1 plate and 2 spines) and some ribs. Specimen CLGPR V00006-2 is preserved in the middle of the wall of Dinosaur fossils, and comprises five dorsal vertebrae, right tibia, right fibula, one piece of dermal plate, and some ribs. CLGPRV00006-3 is a dorsal vertebra from the east side of the wall of Dinosaur fossils.

 
(A)–(F) Dorsal vertebrae referred to Bashanosaurus primitivus (CLGPR V00006-3) from the Middle Jurassic of China. (A) Anterior; (B) left lateral; (C) dorsal; (D) posterior; (E) right lateral; (F) ventral; (G) posterior views of dorsal vertebrae referred to Bashanosaurus primitivus (CLGPR V00006-1) from the Middle Jurassic of China. (H) Posterior view of dorsal vertebrae referred to Bashanosaurus primitivus (CLGPR V00006-2) from the Middle Jurassic of China. I, anterior view of dorsal vertebrae referred to Bashanosaurus primitivus (CLGPR V00006-2) from the Middle Jurassic of China. (J)–(O) anterior caudal vertebrae referred to Bashanosaurus primitivus (CLGPR V00006-1) from the Middle Jurassic of China. (J) anterior; (K) left lateral; (L) dorsal; (M) posterior; (N) right lateral; (O) ventral. (P)–(R) middle caudal vertebrae referred to Bashanosaurus primitivus (CLGPR V00006-1) from the Middle Jurassic of China. (P) right lateral; (Q) posterior; (R) ventral. Abbreviations: acpl, anterior centroparapophyseal lamina; dia, diapophysis; ipzs, intraprezygapophyseal shelf; nc, neural canal; ns, neural spine; para, parapophysis; podl, postzygodiapophyseal lamina; pozyg, postzygapophysis; ppdl, paradiapophyseal lamina; przyg, prezygapophysis. Dai et al. (2022).

Bashanosaurus primitivus shows a number of distinctive features not seen in any previously described Stegosaur. These include parapophyses of dorsal vertebrae, which are elevated on stalks at the base of the transverse process, the shape of the scapula blade, which is slender and flares distally, the acromial process of the scapula, which is small, triangular and poorly developed, the 4th trochanter of the femur, which is positioned below the middle of the shaft, and the bases of the dermal plates, which are strongly convex and transversely thickened, and are separated from the anterior/posterior margins of the plates by a distinct groove.

A fourth Stegosaur specimen, CLGPR V00006-4, was recovered from an exploratory trench, about 900 m away from the Dinosaur Fossil Wall. This specimen comprises a partial cranium, a dorsal vertebra, and a piece of dermal armour, which is not sufficient to assign it two species level, since two Stegosaur species have now been described from the Lower Member of the Shaximiao Formation, Bashanosaurus primitivus, and Huayangosaurus taibaii, which was described in 1984, from material from the Shaximiao Formation in of Sichuan Province. 

 
(A)–(F) Braincase referred to the Stegosauria but not to species level. (CLGPRV0005-4) from the Middle Jurassic of China. (A) Anterior; (B) left lateral; (C) right lateral; (D) posterior; (E) dorsal; (F) ventral. (G)–(L) Dorsal vertebrae referred to the Stegosauria but not to species level. (CLGPRV0005-4) from the Middle Jurassic of China. (G) Anterior; (H) left lateral; (I) dorsal; (J) posterior; (K) right lateral; (L) ventral. (M)–(R) Dermal armour referred to the Stegosauria but not to species level. (CLGPRV0005-4) from the Middle Jurassic of China. (M) Anterior; (N) posterior; (O) left lateral; (P) right lateral; (Q) dorsal; (R) ventral. Abbreviations: boc, basioccipital; bpt, basipterygoid process; bsp, basiphenoid; cc, carotid canal; c.n.I, III, V, VII, XII exit for cranial nerves; dia, diapophysis; f, frontal; fime, fissura metotica; fm, foramen magnum; fo, foramen ovale; ipzs, intraprezygapophyseal shelf; ist, integumentary structure; nc, neural canal; ns, neural spine; oc, occipital condyle; os, orbitosphenoid; p, parietal; para, parapophysis; poc, paroccipital process; pozyg, postzygapophysis; przyg, prezygapophysis; ps, parasphenoid; soc, supraoccipital; sf, supratemporal fossae; sq, squamosal. Dai et al. (2022).

A phylogenetic analysis, based upon a previous study of the group published by a team led by Sussanah Maidment in 2020, found that Bashanosaurus primitivus is the sister taxon to Chungkingosaurus jiangbeiensis, a Stegosaur from the Late Jurassic Upper Member of the Shaximiao Formation, and that together these two species form a sister group to all other Stegosaurs. 

 
A single most parsimonious tree recovered by the phylogenetic analysis. The numbers below the line are Bremer supports, the numbers above the line are GC frequencies. Ankylosauria was represented by three taxa, Gastonia, Euoplocephalus, and Sauropelta in the analysis, but these are shown here as a single terminal taxon for clarity. Dai et al. (2022).

Dai et al. also carried out a radiometric analysis of the deposits which produced Bashanosaurus primitivus, using thorium/uranium ratios in detrital zircons (uranium-thorium dating works because uranium decays to thorium at a known rate, so that the ratio of the two elements in minerals that naturally incorporate uranium but not thorium can be used to establish a date for the minerals).  This produced a range of ages, which would be expected from durable minerals in sedimentary rocks, but the youngest were dated to roughly 169 million years old, implying that the rocks of the Lower Member of the Shaximiao Formation were deposited no earlier than the Bajocian Stage of the Middle Jurassic. 

If the Lower Member of the Shaximiao Formation is Bajocian in origin, then that would make the Stegosaurs found within it, Bashanosaurus primitivus and Huayangosaurus taibaii, the oldest known Stegosaur speciemens. Since Huayangosaurus taibaii is the next branching clade within the Stegosauria after the clade that comprises Bashanosaurus primitivus and Chungkingosaurus jiangbeiensis, this strongly supports an Asian origin for the Stegosauria as a whole.

Dai et al.'s results suggest that the Lower Member of the Shaximiao Formation at Dinosaur Quarry at Laojun Village is Middle Jurassic in age. However, another recent study has suggested that the Lower Member of the Shaximiao Formation at Zigong in Sichuan Province should be dated to the Late Jurassic. Dai et al. suggest that this is because, in the Middle Jurassic, the centre of deposition within the Sichaun Basin (which contains the Shaximiao Formation) was in the northeast, shifted over time due to the rapid filling of the basin by material derived from the intracontinental subduction of the Qinling orogenic belt and Daba Mountains, with the upshot that the Shaximiao Formation may be laterally diachronous (that is to say of different ages across it's horizontal extent). If this is the case, then the rocks which produced Huayangosaurus taibaii may be younger than those which produced Bashanosaurus primitivus, even though they are part of the same member of the same formation.

Bashanosaurus primitivus is therefore the oldest known Stegosaur in Asia, and one of the oldest known Stegosaurs in the world, with geochronological data supporting a Bajocian age for the Lower Member of the Shaximiao Formation at Dinosaur Quarry. Bashanosaurus primitivus is also one of the earliest branching Stegosaurs phylogenetically, forming a clade with Chungkingosaurus jiangbeiensis that forms the sister group to all other Stegosaurs. This strongly supports the hypothesis that the Stegosauria as a whole had an Asian origin.

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