Edmontosaurus Regalis Research Articles

Craniomandibular anatomy of a juvenile specimen of Edmontosaurus regalis Lambe, 1917 clarifies issues in ontogeny and biogeography

ABSTRACT Edmontosaurus regalis is a hadrosaurid represented by abundant fossil material from the late Campanian–early Maastrichtian of what is now Alberta, Canada. Despite its prevalence, juvenile craniomandibular elements of E. regalis are poorly known. A newly collected partial skeleton from the Horseshoe Canyon Formation, UALVP 60425, represents a small saurolophine with a partial skull (skull length ∼500 mm). It can be referred to Edmontosaurini based on the presence of a postorbital fossa and anteroposteriorly long frontals that lack a dorsally oriented nasal contact and contribute significantly to the orbital margin. Despite its small size and presumed immaturity, UALVP 60425 is assigned to E. regalis based on a horizontal shelf on the postorbital process of the jugal. The deep postorbital fossa that, in part, diagnoses E. regalis is acquired during growth. UALVP 60425 shares many features with the Alaskan Prince Creek saurolophine (Ugrunaaluk kuukpikensis); both show consistent anatomical differences in the lacrimal, jugal, and postorbitals from E. annectens throughout their growth series. New data from UALVP 6025 supports referring the Alaskan saurolophine material to Edmontosaurus cf. regalis in the absence of large-sized cranial material from the Prince Creek Formation.

Re-examination of the cranial osteology of the Arctic Alaskan hadrosaurine with implications for its taxonomic status.

Hadrosaurid fossils from the Liscomb Bonebed (Prince Creek Formation, North Slope, Alaska) were the first dinosaur bones discovered from the Arctic. While the Prince Creek Formation hadrosaurids were long identified as Edmontosaurus, a member of the sub-clade Hadrosaurinae, they were recently assigned to a newly-erected taxon, Ugrunaaluk kuukpikensis. However, taxonomic status of the new taxon is ambiguous largely due to the immature nature of the specimens upon which it was based. Here we reexamine cranial elements of the Prince Creek Formation hadrosaurine in order to solve its taxonomic uncertainties. The Prince Creek Formation hadrosaurine possesses a short dorsolateral process of the laterosphenoid, one of the diagnostic characters of Edmontosaurus. The Prince Creek Formation hadrosaurine also shows affinity to Edmontosaurus regalis in the presence of a horizontal shelf of the jugal. Our morphological comparisons with other North American Edmontosaurus specimens and our phylogenetic analyses demonstrate that the Prince Creek Formation hadrosaurine should be re-assigned to Edmontosaurus. Because the Prince Creek Formation Edmontosaurus shows differences with lower latitude Edmontosaurus in a dorsoventrally short maxilla, presence of a secondary ridge on the dentary teeth, and the absence of the transverse ridge between basipterygoid processes of the basisphenoid, we consider that the Prince Creek Formation Edmontosaurus should be regarded as Edmontosaurus sp. until further discoveries of mature hadrosaurines from the Prince Creek Formation Bonebed and/or equivalently juvenile Edmontosaurus specimens from the lower latitudes allow direct comparisons. The retention of the Prince Creek Formation hadrosaurine as Edmontosaurus re-establishes a significant latitudinal distribution for this taxon. Despite the large latitudinal distribution of the taxon, the morphological disparity of Edmontosaurus is small within Hadrosaurinae. The small morphological disparity may be related to the relatively low latitudinal temperature gradient during the latest Cretaceous compared to present day, a gradient which might not have imposed significant pressure for much morphological adaptations across a broad latitudinal range.

A taphonomic analysis of a multitaxic bonebed from the St. Mary River Formation (uppermost Campanian to lowermost Maastrichtian) of Alberta, dominated by cf. Edmontosaurus regalis (Ornithischia: Hadrosauridae), with significant remains of Pachyrhinosaurus canadensis (Ornithischia: Ceratopsidae)

Scabby Butte is an isolated exposure of Upper Cretaceous (uppermost Campanian to lowermost Maastrichtian) strata of the St. Mary River Formation. These rocks have produced a diverse assemblage of both terrestrial and aquatic vertebrates, although hadrosaurid and ceratopsid dinosaurs represent the largest component by volume. Almost all of these dinosaur remains were collected from a single bonebed (Site 2) at Scabby Butte and have been referred to the hadrosaurid Edmontosaurus regalis Lambe, 1917 and the ceratopsid Pachyrhinosaurus canadensis Sternberg, 1950. This study presents a quantitative taphonomic reanalysis of the originally published data, providing new information previously overlooked and important information about the age-class structure of the dinosaur fauna preserved at the site. Site 2 is a lag deposit with a minimum number of 11 individuals (two ceratopsid, nine hadrosaurid), with three-quarters of the material being adult-sized based on size-class analysis. Most elements have undergone moderate to severe breakage and abrasion, and are completely disarticulated, suggesting that they were transported from where they died; post-mortem scavenging is also a possibility, as evidenced by the presence of tooth marks and trample marks on several elements. Burial took place soon after scavenging, as there is little evidence of subaerial weathering.

High-precision U–Pb CA–ID–TIMS dating and chronostratigraphy of the dinosaur-rich Horseshoe Canyon Formation (Upper Cretaceous, Campanian–Maastrichtian), Red Deer River valley, Alberta, Canada

The non-marine Horseshoe Canyon Formation (HCFm, southern Alberta) yields taxonomically diverse, late Campanian to middle Maastrichtian dinosaur assemblages that play a central role in documenting dinosaur evolution, paleoecology, and paleobiogeography leading up to the end-Cretaceous extinction. Here, we present high-precision U–Pb CA–ID–TIMS ages and the first calibrated chronostratigraphy for the HCFm using zircon grains from (1) four HCFm bentonites distributed through 129 m of section, (2) one bentonite from the underlying Bearpaw Formation, and (3) a bentonite from the overlying Battle Formation that we dated previously. In its type area, the HCFm ranges in age from 73.1–68.0 Ma. Significant paleoenvironmental and climatic changes are recorded in the formation, including (1) a transition from a warm-and-wet deltaic setting to a cooler, seasonally wet-dry coastal plain at 71.5 Ma, (2) maximum transgression of the Drumheller Marine Tongue at 70.896 ± 0.048 Ma, and (3) transition to a warm-wet alluvial plain at 69.6 Ma. The HCFm’s three mega-herbivore dinosaur assemblage zones track these changes and are calibrated as follows: Edmontosaurus regalis – Pachyrhinosaurus canadensis zone, 73.1–71.5 Ma; Hypacrosaurus altispinus – Saurolophus osborni zone, 71.5–69.6 Ma; and Eotriceratops xerinsularis zone, 69.6–68.2 Ma. The Albertosaurus Bonebed — a monodominant assemblage of tyrannosaurids in the Tolman Member — is assessed an age of 70.1 Ma. The unusual triceratopsin, Eotriceratops xerinsularis, from the Carbon Member, is assessed an age of 68.8 Ma. This chronostratigraphy is useful for refining correlations with dinosaur-bearing upper Campanian–middle Maastrichtian units in Alberta and elsewhere in North America.

Supplementary cranial description of the types of Edmontosaurus regalis (Ornithischia: Hadrosauridae), with comments on the phylogenetics and biogeography of Hadrosaurinae.

The cranial anatomy of the flat-skulled hadrosaurine Edmontosaurus regalis (Ornithischia: Hadrosauridae) is extensively described here, based on the holotype and paratype collected from the middle part of the Horseshoe Canyon Formation in southern Alberta. Focus is given to previously undocumented features of ontogenetic and phylogenetic importance. This description facilitates overall osteological comparisons between E. regalis and other hadrosaurids (especially E. annectens), and revises the diagnosis of E. regalis, to which a new autapomorphy (the dorsal half of the jugal anterior process bearing a sharp posterolateral projection into the orbit) is added. We consider the recently named Ugrunaaluk kuukpikensis from the upper Campanian/lower Maastrichtian of Alaska a nomen dubium, and conservatively regard the Alaskan material as belonging to Edmontosaurus sp.. A phylogenetic analysis of Hadrosauroidea using maximum parsimony further corroborates the sister-taxon relationship between E. regalis and E. annectens. In the strict consensus tree, Hadrosaurus foulkii occurs firmly within the clade comprising all non-lambeosaurine hadrosaurids, supporting the taxonomic scheme that divides Hadrosauridae into Hadrosaurinae and Lambeosaurinae. Within Edmontosaurini, Kerberosaurus is posited as the sister taxon to the clade of Shantungosaurus + Edmontosaurus. The biogeographic reconstruction of Hadrosaurinae in light of the time-calibrated cladogram and probability calculation of ancestral areas for all internal nodes reveals a significantly high probability for the North American origin of the clade. However, the Laramidia–Appalachia dispersals around the Santonian–Campanian boundary, inferred from the biogeographic scenario for the North American origin of Hadrosaurinae, are in conflict with currently accepted paleogeographic models. By contrast, the Asian origin of Hadrosaurinae with its relatively low probability resulting from the biogeographic analysis is worth seriously considering, despite the lack of fossil material from the Santonian and lower Campanian of Asia. Extra fossil collecting in appropriate geographic locations and stratigraphic intervals of Asia and Europe will help to clarify the biogeographic dynamics of hadrosaurine dinosaurs in the near future.

Hadrosaurid (Edmontosaurus) bonebeds from the Horseshoe Canyon Formation (Horsethief Member) at Drumheller, Alberta, Canada: geology, preliminary taphonomy, and significance

Three monodominant hadrosaurid bonebeds in the Horsethief Member of the Horseshoe Canyon Formation (uppermost Campanian) in southern Alberta, Canada, are documented. Each bonebed is hosted by a decimetre-thick deposit of poorly sorted and graded organic-fragment-rich mudstone. These fossil deposits are interpreted as having been carried and deposited by debris flows or hyperconcentrated mass sediment flows initiated by overbank flooding from local channels. Each bonebed is dominated (>50% of identifiable elements) by the disarticulated to occasionally associated remains of hadrosaurine hadrosaurids, inferred to be Edmontosaurus regalis. The majority of hadrosaurid elements at two of the sites (Bleriot Ferry and Prehistoric Park) are from large, presumably adult-sized individuals, whereas the majority of elements from the Fox Coulee site are from subadults and juveniles. Fossil elements from all the sites exhibit similar taphonomic signatures suggestive of a high degree of biostratinomic modification including: (i) thorough disarticulation of carcasses, (ii) a large amount of breakage, (iii) modest amounts of size sorting, and (iv) minimum to modest occurrences of abrasion, and scratch and tooth marks. These signatures indicate that carcasses were exposed for significant amounts of time on the floodplain, where they rotted, were scavenged and trampled, and were exposed to moving water prior to final burial. The size of each bonebed together with the density of bones suggest that the biocoenoses comprised large groups of hadrosaurids, and bone size distributions suggest the possibility of age-segregated populations. The monodominant nature of the assemblages combined with homogenous taphonomic signatures within and between sites suggests that these bonebed assemblages are best interpreted as the result of mass kills rather than attrition, with recurring tropical storm-induced coastal-plain flooding postulated as a likely mechanism for what killed and eventually buried these dinosaurs.

Introduction to the Special Issue commemorating the 30th anniversary of the Royal Tyrrell Museum of Palaeontology, with a summary of the museum’s early history and its research contributions

Located just west of Drumheller, in southern Alberta, Canada, the Royal Tyrrell Museum of Palaeontology (RTMP) is nestled among outcrops of the Upper Cretaceous Horseshoe Canyon Formation exposed along the Red Deer River valley (Fig. 1). Since it officially opened on 25 September 19851, the museum has become one of Alberta’s must-see tourist attractions, a significant contributor to the local economy, and earned an international reputation for the quality of its palaeontological collections and the research conducted by its scientists. The RTMP is unique within Canada in being the only government-supported museum dedicated to palaeontology, and it is one of the few such institutions in the world. Fig. 1. The Royal Tyrrell Museum of Palaeontology. (A) Front of museum and main entrance (circa 2010), looking westwards and with horizontal beds of the Upper Cretaceous Horseshoe Canyon Formation exposed in the walls of the Red Deer River valley in the distance. (B) Aerial photograph, oblique angle and looking to the north-northeast. The museum sits on top of the number 7 coal seam (sensu Gibson 1977) and at the base of the Horsethief Member of the Horseshoe Canyon Formation (D.A. Eberth, pers. comm., 2015). That stratigraphic position correlates with an age of about 71.5 Ma (i.e., late Campanian) and lies within the upper part of the Edmontosaurus regalis – Pachyrhinosaurus canadensis dinosaur assemblage zone (see Eberth et al. 2013, fig. 6). (C) The iconic “running theropod” logo that until recently graced the exterior of the museum, below a life-sized tyrannosaurid sculpture by Brian Cooley (one of many such sculptures by Cooley that have been featured at the museum since its opening in 1985) and with the education wing (opened in 2003 as the ATCO Tyrrell Learning Centre) in the background. (D) The Cretaceous Alberta life-sized diorama (opened in 2007), an example of a RTMP exhibit that relies heavily on discoveries and interpretations by museum researchers, in this case work done in the Albertosaurus bonebed and at other localities throughout the Horseshoe Canyon Formation. All photographs courtesy of the Royal Tyrrell Museum of Palaeontology. This Special Issue of the Canadian Journal of Earth Sciences presents a collection of papers commemorating the RTMP’s 30th or pearl anniversary. Our rationale for doing a 30th anniversary volume was two-fold. First, it is an opportunity to showcase the breadth of research conducted by current and former museum scientists as well as by some colleagues having a history of collaboration with the RTMP. The 13 invited papers included in this volume are a mix of review and original research papers, all thematically focused on the geology and palaeontology of Alberta. To varying degrees, each contribution is the result of work supported by the RTMP and relies on specimens housed in the museum’s collections. Second, the 30th anniversary is an opportunity to reflect on the RTMP’s history and its research program. As is typical for most institutions, the RTMP has done little to formally document its own history. This year likely will be the last major milestone for which the original generation of museum staff are still working and available to share their recollections about the museum, fieldwork, and research. Already, deaths and departures have winnowed away part of the original …

Stratigraphy of the Danek Bonebed (Upper Cretaceous Horseshoe Canyon Formation, central Alberta) and correlations with strata in the Drumheller and Grande Prairie regions

Although considerable work has been conducted on the stratigraphy and dinosaur assemblages of the Horseshoe Canyon Formation of southern Alberta, equivalent strata and assemblages in central Alberta remain poorly understood. Data from the Danek Bonebed (Edmonton, Alberta) are beginning to fill this gap. The bonebed occurs 4 m above the #9 Big Island Coal Seam. This stratigraphic position lies just below the middle of the Horseshoe Canyon Formation in the Edmonton region, and also lies below a thick, stratigraphically significant non-coaly interval that is expressed throughout central and southern Alberta. The stratigraphic position of the Danek Bonebed equates best with the uppermost Horsethief Member of the Horseshoe Canyon Formation in the Drumheller region and the upper part of Unit 4 of the Wapiti Formation in the Grande Prairie region. In both Drumheller and Grande Prairie, the correlated position of the bonebed underlies a zone of marine transgression (Drumheller Marine Tongue), which, in turn, includes the Campanian–Maastrichtian boundary. In the context of Geologic Time Scale 2004, we infer a late Campanian age of 71.0–71.3 Ma for the bonebed. The Danek’s dinosaurian assemblage is limited taxonomically, but compares well with the Edmontosaurus regalis – Pachyrhinosaurus canadensis fossil assemblage zone in the Drumheller region. We propose that a mega-herbivore faunal assemblage, characterized by Edmontosaurus and Pachyrhinosaurus, extended continuously across the climatically wet coastal plain of latest Campanian southern and central Alberta, and likely extended northwest into the North Slope of Alaska, where it persisted into the early Maastrichtian.

Enhancing bonebed mapping with GIS technology using the Danek Bonebed (Upper Cretaceous Horseshoe Canyon Formation, Edmonton, Alberta, Canada) as a case study

The Danek Bonebed is a monodominant Edmontosaurus bonebed preserving predominantly disarticulated material from the Upper Campanian Horseshoe Canyon Formation within the city limits of Edmonton, Alberta, Canada. To date, at least six dinosaur taxa have been identified at the site: Albertosaurus sarcophagus, Chasmosaurinae indet., Dromaeosauridae indet., Edmontosaurus regalis, Ornithomimidae indet., and Troodontidae indet. This bonebed has been used as a case study for creating a digital, searchable bonebed map using a geographic information system (GIS) platform. The original quarry maps produced on site are refined when digitized with new anatomical information gathered during preparation of collected specimens. Each specimen is labeled with the known specimen identification, quarry coordinates, and catalogue number. Creating a digital map of the bonebed allows easier interpretation of data and the ability to share maps to compare specific elements within the bonebed.

Histologic growth dynamic study of Edmontosaurus regalis (Dinosauria: Hadrosauridae) from a bonebed assemblage of the Upper Cretaceous Horseshoe Canyon Formation, Edmonton, Alberta, Canada

The Danek Bonebed (Edmonton, Alberta, Canada) is a monodominant Edmontosaurus regalis assemblage of the upper Campanian (Upper Cretaceous) Horseshoe Canyon Formation. Bone histology of humeri and femora are used in this paper to test hypotheses about the growth dynamics and palaeobiology of Edmontosaurus. The high number of elements collected from the Danek Bonebed allow for an expansion of the multi-element histological record for hadrosaurs. Results indicate that Edmontosaurus had a growth trajectory similar to other large-bodied dinosaurs and reached the onset of somatic maturity at about 10–15 years of age; however, even the largest elements to preserve lines of arrested growth do not have external fundamental systems. This timing of the onset of somatic maturity agrees with the estimated body size of Edmontosaurus relative to other dinosaurs for which life-history data are available. Vascularity patterns support the hypothesis that edmontosaurs preserved at the Danek Bonebed were not subject to the same extreme seasonal environmental shifts as congenerics preserved at higher latitudes, further supporting overwintering behaviour in the latter.

Taphonomy of the Danek Bonebed: a monodominant Edmontosaurus (Hadrosauridae) bonebed from the Horseshoe Canyon Formation, Alberta

The Danek Bonebed (Horsethief Member, Horseshoe Canyon Formation, Late Campanian) is dominated by the remains of at least 12 Edmontosaurus regalis. Skeletal remains of a tyrannosaurid and ceratopsid are also known. The predominantly disarticulated remains were interred on a periodically inundated floodplain and, although the cause of death is unknown, a sudden, catastrophic death explains the demographic spread, faunal diversity, rare greenstick fractures, and homogeneous weathering/abrasion categories of the assemblage. The Danek Bonebed shares a similar taphonomic signature to the Liscomb Bonebed (Prince Creek Formation, Alaska), but it is unique among all other described hadrosaurid bonebeds in the unusually high proportion of bite-marked bones (∼30%), suggesting scavenging played a major role in the reworking of the assemblage. The highest frequency of bite marks is found on small, often unidentifiable (and commonly ignored) bone fragments, underscoring the role that such fragments can play in taphonomic interpretation. Finally, the recognition of E. regalis from central Alberta is an important datum linking contemporaneous occurrences in southern Alberta with slightly older records of this species from the Wapiti Formation in northwestern Alberta.

Quantitative diagenetic analyses of Edmontosaurus regalis (Dinosauria: Hadrosauridae) postcranial elements from the Danek Bonebed, Upper Cretaceous Horseshoe Canyon Formation, Edmonton, Alberta, Canada: implications for allometric studies of fossil organisms

For fossil assemblages, quantitative size and shape studies are often complicated by diagenetic distortion. Different vertebrate elements, although subjected to similar burial stresses, exhibit deformations based upon their original shapes; this hypothesis is tested here by quantitatively comparing deformed humeri and femora from the Danek Bonebed (a monodominant Edmontosaurus regalis bonebed from the upper Campanian Horseshoe Canyon Formation in Edmonton, Alberta, Canada) with samples of undeformed humeri and femora from modern and fossil assemblages. Analyses suggest that at the Danek Bonebed a strong relationship exists between element length and circumference despite being distorted by crushing deformation. Major and minor axes of the midshaft cross section, however, were not uniformly distorted. Although their anatomical position did not change, the major axis became longer relative to the minor axis in distorted specimens. A regression based on the undeformed humeri was not able to accurately predict circumference in the Danek humeri. Further study might quantify the deformation of other bones in the Danek Bonebed and could be extended to other assemblages and genera. Caution should be taken when conducting studies in which diagenetic crushing may have altered morphological features of fossil vertebrate remains.

Modeling cranial biomechanics in archosaurs using 3D computational methods (17.5)

Accurately modeling cranial function in vertebrates remains a challenging yet important tool for ecologists, functional morphologists, and paleobiologists. Three‐dimensional computational methods now offer new approaches for reconstructing jaw muscle anatomy and the cranial biomechanical environment so that evolutionary and functional hypotheses can be tested. Here we present several examples that reconstruct muscle forces, joint moments and bite forces in extant and extinct archosaurs using 3D modeling techniques. 1) We compared modeled bite forces in an ontogenetic sample of Alligator mississippiensis with published in vivo bite forces to validate whether the method is applicable to diapsid vertebrates. 2) We modeled bite forces and joint moments within the skulls of the non‐avian dinosaurs Tyrannosaurus rex and Edmontosaurus regalis to test the sensitivity of input parameters with force estimates. 3) Using these latter taxa, we compared moments about joints implicated in cranial kinesis to identify functional patterns. These models will enable us to test for coevolution between bite force and dietary correlates in fossil archosaurs, estimate loading patterns within cranial joints, and determine patterns in jaw muscle anatomy and function among tetrapods.

A Mummified Duck-Billed Dinosaur with a Soft-Tissue Cock’s Comb

Among living vertebrates, soft tissues are responsible for labile appendages (combs, wattles, proboscides) that are critical for activities ranging from locomotion to sexual display [1]. However, soft tissues rarely fossilize, and such soft-tissue appendages are unknown for many extinct taxa, including dinosaurs. Here we report a remarkable "mummified" specimen of the hadrosaurid dinosaur Edmontosaurus regalis from the latest Cretaceous Wapiti Formation, Alberta, Canada, that preserves a three-dimensional cranial crest (or "comb") composed entirely of soft tissue. Previously, crest function has centered on the hypertrophied nasal passages of lambeosaurine hadrosaurids, which acted as resonance chambers during vocalization [2-4]. The fleshy comb in Edmontosaurus necessitates an alternative explanation most likely related to either social signaling or sexual selection [5-7]. This discovery provides the first view of bizarre, soft-tissue signaling structures in a dinosaur and provides additional evidence for social behavior. Crest evolution within Hadrosaurinae apparently culminated in the secondary loss of the bony crest at the terminal Cretaceous; however, the new specimen indicates that cranial ornamentation was in fact not lost but substituted in Edmontosaurus by a fleshy display structure. It also implies that visual display played a key role in the evolution of hadrosaurine crests and raises the possibility of similar soft-tissue structures among other dinosaurs.

Dinosaur biostratigraphy of the Edmonton Group (Upper Cretaceous), Alberta, Canada: evidence for climate influence

A high-resolution biostratigraphic analysis of 287 dinosaurian macrofossils and 138 bonebeds in the Edmonton Group (Upper Cretaceous) of southern Alberta provides evidence for at least three dinosaurian assemblage zones in the Horseshoe Canyon Formation (HCFm). From bottom to top the zones comprise unique assemblages of ornithischians and are named as follows: (1) Edmontosaurus regalis – Pachyrhinosaurus canadensis (lower zone); (2) Hypacrosaurus altispinus – Saurolophus osborni (middle zone); and (3) Eotriceratops xerinsularis (upper zone). Whereas the lower and middle zones are well defined and based on abundant specimens, the validity of the uppermost zone (E. xerinsularis) is tentative because it is based on a single specimen and the absence of dinosaur taxa from lower in section. The transition from the lower to the middle zone coincides with the replacement of a warm-and-wet saturated deltaic setting by a cooler, coastal-plain landscape, characterized by seasonal rainfall and better-drained substrates. Whereas changes in rainfall and substrate drainage appear to have influenced the faunal change, changes in mean annual temperature and proximity to shoreline appear to have had little influence on faunal change. We speculate that the faunal change between the middle and upper zones also resulted from a change in climate, with ornithischian dinosaurs responding to the re-establishment of wetter-and-warmer climates and poorly-drained substrates. Compared with the shorter-duration and climatically-consistent dinosaurian assemblage zones in the older Dinosaur Park Formation of southern Alberta, HCFm assemblage zones record long-term morphological stasis in dinosaurs. Furthermore, the coincidence of faunal and paleoenvironmental changes in the HCFm suggest climate-change-driven dinosaur migrations into and out of the region.

Kinetic Limitations of Intracranial Joints in Brachylophosaurus canadensis and Edmontosaurus regalis (Dinosauria: Hadrosauridae), and Their Implications for the Chewing Mechanics of Hadrosaurids

The highly specialized tooth morphology and arrangement of the dental battery of hadrosaurids has led to much speculation surrounding the chewing mechanics of this successful group of herbivorous dinosaurs. Pleurokinesis, a long established hypothesis explaining the ornithopod chewing mechanism, proposes a transverse power stroke in hadrosaurids that was accommodated by vertical adduction of the mandible, lateral rotation of the maxilla at the maxilla-premaxilla joint, lateral rotation of the jugal-maxilla complex at its contact with the lacrimal, and posterolateral rotation of the quadrate at its contact with the squamosal. A secondary series of movements were also thought to have occurred as a consequence of these primary movements. In this article, the intracranial joint morphology is described for both Brachylophosaurus canadensis and Edmontosaurus regalis and their permissive kinematics are established. Based on this evidence, the movements associated with pleurokinesis are not accommodated in these hadrosaurine dinosaurs. Rather, the movements that seem most likely to have produced the observed dental wear patterns are those associated with the mandible about the jaw joint. The structure of this joint appears well-suited to have accommodated some translation as well as rotation of the mandible about the quadrate condyle. Three-dimensional modeling of the alternate mandibular movements reveals that not all the combined labiolingual width of the lingual and buccal facets of the tooth row was involved in the power stroke. Rather, limits on the degree of mandibular long axis rotation suggest that only the lingual facet and the more medial portion of the buccal facet were utilized.

Cranial Growth and Variation in Edmontosaurs (Dinosauria: Hadrosauridae): Implications for Latest Cretaceous Megaherbivore Diversity in North America

The well-sampled Late Cretaceous fossil record of North America remains the only high-resolution dataset for evaluating patterns of dinosaur diversity leading up to the terminal Cretaceous extinction event. Hadrosaurine hadrosaurids (Dinosauria: Ornithopoda) closely related to Edmontosaurus are among the most common megaherbivores in latest Campanian and Maastrichtian deposits of western North America. However, interpretations of edmontosaur species richness and biostratigraphy have been in constant flux for almost three decades, although the clade is generally thought to have undergone a radiation in the late Maastrichtian. We address the issue of edmontosaur diversity for the first time using rigorous morphometric analyses of virtually all known complete edmontosaur skulls. Results suggest only two valid species, Edmontosaurus regalis from the late Campanian, and E. annectens from the late Maastrichtian, with previously named taxa, including the controversial Anatotitan copei, erected on hypothesized transitional morphologies associated with ontogenetic size increase and allometric growth. A revision of North American hadrosaurid taxa suggests a decrease in both hadrosaurid diversity and disparity from the early to late Maastrichtian, a pattern likely also present in ceratopsid dinosaurs. A decline in the disparity of dominant megaherbivores in the latest Maastrichtian interval supports the hypothesis that dinosaur diversity decreased immediately preceding the end Cretaceous extinction event.

Evolution of Dentary Diastema in Iguanodontian Dinosaurs

Abstract:The dentary diastema of iguanodontians has been considered to be related to its unique jaw mechanism for herbivorous adaptation. The dentary and diastema lengths of iguanodontians were measured and compared to elucidate the evolution of iguanodontian diastema. A gap in ratios between most non‐hadrosaurid iguanodontians and hadrosaurids was observed, suggesting that all non‐hadrosaurid iguanodontians, expect forOuranosaurus nigeriensisandProtohadros byrdi, lack a diastema or have a short diastema, although some other taxa have been considered to have a long diastema in previous studies. In non‐hadrosaurid iguanodontians, some large‐sized forms, such asIguanodon bernissartensis, Shuangmiaosaurus gilmorei, and possiblyEolambia caroljonesa, had a short diastema through ontogeny, whereasOuranosaurus nigeriensisandProtohadros byrdideveloped a long diastema convergently. The development of a long diastema of hadrosaurine hadrosaurids may be different from that of lambeosaurine hadrosaurids. Some hadrosaurunes (Edmontosaurus annectens, Edmontosaurus regalis, andSaurolophus angustirostris) may have developed a long diastema in the subadult stage and showed little elongation of dentary diastema through ontogeny. Lambeosaurines (Corythosaurus casuariusandLambeosaurus lambei) tend to have a short diastema in the embryonic and subadult stages, and an elongated diastema from the subadult to adult stages.