Michael J. Ryan's research while affiliated with Carleton University and other places

Publications (100)

Article
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In this study, we use an exceptional skeleton of the pachycephalosaur Stegoceras validum (UALVP 2) to inform a comprehensive appendicular muscle reconstruction of the animal, with the goal of better understanding the functional morphology of the pachycephalosaur postcranial skeleton. We find that S . validum possessed a conservative forelimb muscul...
Article
The early centrosaurine ceratopsid, Wendiceratops pinhornensis, was discovered in Alberta, Canada in a medium density monodominant bonebed from the Oldman Formation (mid-Campanian, ~79 Ma). The bonebed contains abundant, well-preserved, adult-sized and some juvenile-sized postcranial material, allowing for the first description of a number of eleme...
Article
A monodominant Gryposaurus sp. bonebed in the lower unit of the Campanian Oldman Formation of southern Alberta is the oldest hadrosauroid bonebed documented in the province and the first described from the formation. The sedimentology of the locality and the taphonomy of the hadrosaurid material indicates that the bonebed represents an assemblage o...
Article
The pre-Campanian trionychid fossil record in North America is composed of highly fragmentary specimens, which are often not identifiable beyond Pan-Trionychidae. Here, we describe a new species of plastomenid soft-shelled turtle based on a partial shell (ROM 56647) from the Santonian Milk River Formation of southern Alberta, estimated at approxima...
Article
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Goniopholididae is a group of basal neosuchian crocodyliforms closely related to Paralligatoridae and Eusuchia that lived during the Jurassic and Early Cretaceous. Goniopholidids have the long, flat snout and secondary palate of modern crocodylians, the acquisition of which is regarded as a key feature in the early evolution of crocodylian body pla...
Article
Hadrosaurids are a diverse and widely distributed group of ornithischian dinosaurs that are particularly well represented in the upper Campanian Dinosaur Park Formation of the Belly River Group of Alberta. However, the origin of this hadrosaurid diversity in Alberta is poorly understood, as the lower Campanian terrestrial deposits of the underlying...
Article
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A new articulated postcranial specimen of an indeterminate ankylosaurid dinosaur from the Upper Cretaceous (middle-upper Campanian) Baruungoyot Formation from Hermiin Tsav, southern Gobi Desert, Mongolia includes twelve dorsal vertebrae, ribs, pectoral girdles, forelimbs, pelvic girdles, hind limbs, and free osteoderms. The new specimen shows that...
Article
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Elasmosaurid plesiosaurian remains have been documented from non-marine to paralic (fluvial to estuarine) sediments of the upper Campanian Dinosaur Park Formation (DPF) of southern Alberta since 1898. Despite this long collection history, this material has received relatively little research attention, largely due to the highly fragmentary nature o...
Article
We describe a new partial skull with braincase of a maiasaurin hadrosaurid from the Milk River Ridge Reservoir near Warner, southern Alberta, as the first diagnostic occurrence of Maiasaura in Canada. This material was collected in the Oldman Formation, at approximately the same stratigraphic level as a nearby bonebed of the ceratopsid Coronosaurus...
Article
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Ceratopsids are among the most ubiquitous dinosaur taxa from the Late Cretaceous terrestrial formations of the Western Interior of North America, comprising two subfamilies, Chasmosaurinae and Centrosaurinae. The Two Medicine Formation of northwestern Montana has produced numerous remains of centrosaurine dinosaurs, which represent three taxa previ...
Article
Until now, the pre-Miocene fossil record of mustelids in North America has been restricted to specimens attributable to oligobunine taxa and isolated remains tentatively allocated to the genus Plesictis. In the present study, we report on a nearly complete cranium and a referred dentary of a new genus and species of mustelid. The specimens were rec...
Article
Until now, the pre-Miocene fossil record of mustelids in North America has been restricted to specimens attributable to oligobunine taxa and isolated remains tentatively allocated to the genus Plesictis. In the present study, we report on a nearly complete cranium and a referred dentary of a new genus and species of mustelid. The specimens were rec...
Article
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 din...
Article
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Ontogenetic sequences are relatively rare among dinosaurs, with Ceratopsia being one of the better represented clades, and especially among geologically earlier forms, such as Psittacosaurus . Psittacosaurus is a small, bipedal basal ceratopsian abundant in the Lower Cretaceous deposits of Asia, whose cranial and endocranial morphology has been wel...
Article
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The Dinosaur Park Formation (DPF) has a diverse assemblage of chasmosaurines currently represented by Chasmosaurus belli, C. russelli, Vagaceratops irvinensis, and Mercuriceratops gemini, and may also include remains possibly referable to Spiclypeus shipporum. Two skulls, YPM 2016 and AMNH 5402, previously referred to C. belli, both have a straight...
Article
Full-text available
Chasmosaurine ceratopsids are well documented from the Upper Cretaceous (Campanian) Dinosaur Park Formation (DPF) of southern Alberta and Saskatchewan, and include Chasmosaurus belli , Chasmosaurus russelli , Mercuriceratops gemini , Vagaceratops irvinensis , and material possibly referable to Spiclypeus shipporum. In this study, we describe three...
Article
Full-text available
Positional variation is documented in ornithomimid pedal unguals from the Dinosaur Park and Horseshoe Canyon Formations of Alberta, Canada, and characters for identifying the position of isolated ornithomimid pedal unguals are discussed. Ungual morphology has been used recently to argue for the coexistence of two distinct ornithomimosaurs, a basal...
Preprint
Vertebrate microfossil assemblages contain abundant fossil material of small and large bodied animals recruited from the local paleocommunity that can provide important information for reconstructing regional paleoecology. The Foremost Formation is the oldest unit of the Belly River Group, and records the transition from the fully marine shales of...
Article
Medusaceratops lokii Ryan, Russell, and Hartman, 2010 is an enigmatic taxon of ceratopsid represented by partial parietals from the Mansfield bonebed in the Campanian Judith River Formation, Montana. Originally, all ceratopsid material collected from this bonebed was referred to the centrosaurine ceratopsid Albertaceratops , but subsequently two pa...
Article
To date, only three pachycephalosaur specimens have been reported from the Nemegt Formation, two of which are holotypes (Homalocephale calathocercos and Prenocephale prenes) and all of which were collected from the classic Nemegt locality. Here, we describe several new cranial specimens from this formation, but which originate from Bügiin Tsav, Gur...
Article
Ornithomimid material from the Belly River Group (Campanian) of Alberta, Canada is described as sharing characters with Qiupalong henanensis from the Qiupa Formation of Henan Province, China. Derived characters and character combinations of the pubis and astragalocalcaneum were previously used to diagnose Q. henanensis and support the referral of t...
Article
The placoderm Titanichthys from the Late Devonian (Famennian) is based on incomplete and fragmentary specimens that have hindered understanding of its overall anatomy and phylogenetic relationships. A new, nearly complete, articulated specimen from the Upper Devonian Cleveland Shale provides new information about the previously undescribed rostral,...
Article
A partial skull (CMN 8804) of a ceratopsid from the upper unit of the Campanian Oldman Formation of Alberta is the first Canadian example of the newly established Nasutoceratopsini, a new subclade of Centrosaurinae defined as the stem-based clade of centrosaurine ceratopsids more closely related to Nasutoceratops titusi than to Centrosaurus apertus...
Poster
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Poster on Dunkleosteus ecology presented at the Society of Vertebrate Paleontology Annual Meeting in Salt Lake City, Utah 2016.
Article
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A monodominant bonebed of Avimimus from the Nemegt Formation of Mongolia is the first oviraptorosaur bonebed described and the only recorded maniraptoran bonebed from the Late Cretaceous. Cranial elements recovered from the bonebed provide insights on the anatomy of the facial region, which was formerly unknown in Avimimus. Both adult and subadult...
Article
A partial ornithomimid skeleton, ROM 1790, from the lower Dinosaur Park Formation (upper Campanian) of Alberta was previously referred to Struthiomimus altus, but lacks diagnostic characters of that species. It is here described as the holotype of a new species, Rativates evadens, gen. et sp. nov., diagnosed by the form of the maxilla-jugal contact...
Conference Paper
Ornithomimid dinosaurs are best known from Upper Cretaceous deposits in the western interior of Canada and the United States, and the Gobi of northern China and Mongolia. Two recently named taxa extend the geographic range of definitive ornithomimid occurrences: Qiupalong henanensis from central China (Xu et al. 2011), and Tototlmimus packardensis...
Article
An articulated, 1.5 m long skeleton of a juvenile Chasmosaurus, lacking only the front limbs, pectoral girdles, and terminal caudal vertebrae, was collected from the Dinosaur Park Formation in Dinosaur Provincial Park, Alberta. The short, tall skull has a narrow frill that lacks a posterior embayment. Many of the cranial bones are co-ossified in sp...
Data
Phylogenetic character-taxon matrix (NEXUS file). (NEX)
Data
Ontogenetic character-specimen matrix (TNT file). (TNT)
Article
Full-text available
Background: The chasmosaurine ceratopsid Chasmosaurus is known from the Upper Cretaceous (Campanian) Dinosaur Park Formation of southern Alberta and Saskatchewan. Two valid species, Chasmosaurus belli and C. russelli, have been diagnosed by differences in cranial ornamentation. Their validity has been supported, in part, by the reported stratigrap...
Data
Ontogenetic character-specimen matrix (NEXUS file). (NEX)
Data
Phylogenetic character-taxon matrix (TNT file). (TNT)
Data
Ontogenetically variable phylogenetic characters. (DOC)
Conference Paper
The Cleveland Museum of Natural History has a deep history of collecting in the Cleveland Member of the Ohio Shale and related rocks of northeastern Ohio. In the mid-1960s, staff, volunteers, and local citizens rallied to collect significant vertebrate, invertebrate and plant fossils during the construction of the regional interstate, I-71, that wi...
Conference Paper
Histological evidence derived from tibiae from a monodominant Gryposaurus sp. bonebed in the Oldman Formation (Campanian) indicates that the individuals represent two growth stages, early and late juveniles. Bone microstructure data indicates that the gryposaurs experienced rapid growth and achieved approximately 70% of adult size before the end of...
Conference Paper
Ceratopsid dinosaurs are diagnosed primarily on cranial ornamentation characters. Some taxa, such as the valid species of Chasmosaurus, C. belli and C. russelli, have very similar cranial ornamentations that have been traditionally difficult to quantify. The validity of these taxa have been supported, in part, by their reported stratigraphic segreg...
Article
The horned dinosaur Centrosaurus apertus from the Belly River Group (Campanian) is represented by multiple articulated skulls and skeletons, and is particularly notable for its occurrence in dozens of large-scale monodominant bonebeds, which have been found in the Dinosaur Park Formation across southern Alberta and Saskatchewan. Here we present a d...
Article
Full-text available
Previously reported and new specimens of marine turtles from the late Campanian of Alberta, Canada, provide additional information on the diversity and distribution of chelonioid turtles at this time. An articulated carapace from the Bearpaw Formation previously interpreted as Lophochelys niobrarae is considered to be a juvenile of a specifically i...
Article
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 deposit...
Article
Disentangling ontogenetic from interspecific variation is key to understanding biodiversity in the fossil record, yet information on growth in the ceratopsid subfamily Chasmosaurinae is sparse. Here, we describe the partial skull of a juvenile chasmosaurine, attributed to Arrhinoceratops brachyops, within the context of more mature specimens of thi...
Article
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The fossil record of ceratopsid dinosaurs between the occurrence of their proximate sister taxa in the Turonian and the beginning of their well-documented radiation from the late Campanian of North America onwards (approximately 90 and 77 Ma) is poor, with only seven taxa described from this early period in their evolution. We describe a new taxon...
Article
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Background Osteohistological examinations of fossil vertebrates have utilized a number of proxies, such as counts and spacing of lines of arrested growth (LAGs) and osteocyte lacunar densities (OLD), in order to make inferences related to skeletochronology and mass-specific growth rates. However, many of these studies rely on samplings of isolated...
Article
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A new taxon of chasmosaurine ceratopsid demonstrates unexpected disparity in parietosquamosal frill shape among ceratopsid dinosaurs early in their evolutionary radiation. The new taxon is described based on two apomorphic squamosals collected from approximately time equivalent (approximately 77 million years old) sections of the upper Judith River...
Article
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Ceratopsidae represents one of the last and best-known radiations of non-avian dinosaurs. Interspecific variation is well documented qualitatively with linear measurements, but little has been done to quantify shape differences in the frill that may indicate functional or evolutionary signals. In order to investigate shape change in the squamosal a...
Chapter
A new species of Neurankylus (N. lithographicus sp. nov.) is described on the basis of skull and shell material from the Santonian-aged Milk River Formation, Alberta, Canada. The genus Neurankylus is also rediagnosed on the basis of the Milk River material and on new material pertaining to the type species N. eximius. N. eximius previously was cons...
Article
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An isolated centrum collected ex situ from marine shales of the Lower Cretaceous (Albian) Arctic Red Formation along the Road River represents the first documented occurrence of a plesiosaurian from Yukon. This centrum represents the northernmost occurrence of plesiosaurians in the Western Interior Sea of North America prior to the establishment of...
Article
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The original hypodigm of the controversial mid-Cretaceous Moroccan dinosaur Sigilmassasaurus brevicollis is redescribed, and the diagnosis of the taxon is revised. Unambiguously referred material is restricted to cervical and anterior dorsal vertebrae sharing apomorphies with the holotype. A newly recognized diagnostic character of Sigilmassasaurus...
Article
Relative to large-bodied dinosaurs, the diversity of small-bodied dinosaurs from the Campanian of North America is poorly understood due to a lack of well-preserved skeletons. We document the first articulated remains, as well as the first cranial bones, of non-iguanodontian ornithopods from the Belly River Group of Alberta. The geologically oldest...
Article
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Taphonomic biases dictate how organisms are represented in the fossil record, but their effect on studies of vertebrate diversity dynamics is poorly studied. In contrast to the high diversity and abundance of small-bodied animals in extant ecosystems, small-bodied dinosaurs are less common than their large-bodied counterparts, but it is unclear whe...
Article
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Background Herbivore coexistence on the Late Cretaceous island continent of Laramidia has been a topic of great interest, stemming from the paradoxically high diversity and biomass of these animals in relation to the relatively small landmass available to them. Various hypotheses have been advanced to account for these facts, of which niche partiti...
Data
Skeletal measurements of CMN 12068, 12069, and 12070. Asterisk (*) indicates fragmentary, distorted or incomplete remains. Dash (-) indicates elements that are not present. (XLS)
Data
Pedal unguals of CMN 12069, in dorsal, ventral, and lateral views. A, digit II; B, digit III; C, digit IV. (TIF)
Data
Pedal phalanx measurements used in principal component analysis. Unshared measurements removed. Abbreviations: DH, distal width; DW, distal width; L, length; PH, proximal height; PW, proximal width. (XLS)
Article
Full-text available
Bonebeds can provide a wealth of anatomical, taphonomic, and ontogenetic information about the specimens preserved within them, and can provide evidence for inferred behavior. The material described here represents the first known bonebed of ornithomimids in North America, and the fourth record of an ornithomimosaur bonebed in the world. Partial sk...
Article
Xenoceratops foremostensis gen. et. sp. nov., a new centrosaurine ceratopsid from the Foremost Formation (Campanian) of Alberta, is described based on frill material from at least three adult-sized individuals collected from a low-density bone bed. The material can be assigned to Centrosaurinae based on features of the preserved squamosal. Although...
Article
Ongoing research into the biostratigraphy of the upper Campanian Dinosaur Park Formation (DPF) of Alberta has demonstrated that megaherbivorous dinosaur taxa (ankylosaurs, ceratopsids, hadrosaurids) are not homogeneously distributed throughout the unit. This has compelled proposals of different, informal assemblage zone schemes, and the hypothesis...
Article
Full-text available
In 1916, a centrosaurine dinosaur bonebed was excavated within the Campanian-aged deposits of what is now Dinosaur Provincial Park, Alberta, Canada. Specimens from this now-lost quarry, including two parietals, a squamosal, a skull missing the frill, and an incomplete dentary, were purchased by The Natural History Museum, London. The material was r...
Article
Anchiceratops is a chasmosaurine ceratopsid from the Upper Cretaceous Horseshoe Canyon Formation (HCF) of Alberta. It is distinguished primarily by its unique parietosquamosal frill ornamentation and possibly by the presence of a ventrally flexed olfactory bulb of the brain. Although Anchiceratops is known from at least ten partial skulls, only two...
Article
The cranial roof ornamentation and the degree of neurocentral closure in the holotype of the pachycephalosaurian dinosaur Homalocephale calathocercos is described in detail for the first time in order to assess its ontogenetic status and taxonomic validity. The parietosquamosal ornamentation consists of five primary nodes along the posterior margin...
Article
Full-text available
Chasmosaurus irvinensis (sp. nov.) is distinguished from other species of this genus by the possession of a broad snout, absence of a brow horn (the position of which is occupied by a pit or rugosities suggestive of bone resorption), broadly rounded and open jugal notch, subrectangular squamosal, straight posterior parietal bar bearing 10 epoccipit...
Data
Character-taxon matrix (21 taxa, 136 characters). Data matrix is compiled by adding three new characters (char. 134–136) to a NEXUS version of the matrix of Makovicky and Norell (2006) and Makovicky
Data
Measurements of Koreaceratops hwaseongensis, gen. et sp. nov. (in millimeters). Asterisk indicates as preserved
Article
In 2008, a new basal neoceratopsian was discovered in the Tando beds (Albian) of Tando Basin in South Korea. It represents the first ceratopsian dinosaur in the Korean peninsula and is assigned to Koreaceratops hwaseongensis gen. et sp. nov. Autapomorphies of Koreaceratops include very tall neural spines over five times higher than the associated c...