Hell Creek Formation Of Montana Research Articles

Two new species of small‐bodied pachycephalosaurine (Dinosauria, Marginocephalia) from the uppermost Cretaceous of North America suggest hidden diversity in well‐sampled formations

AbstractHere we report two new small‐bodied pachycephalosaurines: one from the Dinosaur Park Formation of Alberta and the other from the Hell Creek Formation of Montana, each represented by an isolated squamosal. These two new specimens are approximately the same size as squamosals of Sphaerotholus buchholtzae, and possess several overlapping morphologies with the genus (such as a strongly posteroventrally projecting and laterally tapering parietosquamosal bar). Therefore, these two new specimens are identified as belonging to the genus Sphaerotholus. However, a suite of characters and combinations differentiate these two new specimens from the other three Sphaerotholus species. Most notably, they possess multiple posterior and lateral node rows on the squamosal, and lack a parietosquamosal node (a single row and a parietosquamosal node define these other species in the genus). Specifically, the Alberta specimen possesses two posterior and two lateral node rows, whereas the Montana specimen has three of each. This analysis provides phylogenetic support for the evolutionary hypothesis that a single‐rowed and parietosquamosal node‐less taxon underwent a cladogenetic event that gave rise to a single‐rowed and parietosquamosal node‐bearing lineage and a multi‐rowed, parietosquamosal node‐less lineage (the two taxa described herein). In turn, these two Sphaerotholus lineages possibly underwent anagenetic change throughout the latest Maastrichtian. Additionally, these new taxa suggest that while large‐bodied taxa from well sampled formations are to be found with decreasing frequency, the inverse holds true for small‐bodied taxa and that pachycephalosaurines were more diverse than previously recognized, and continued to diversify up until the very end of the Cretaceous.

Stratigraphic range extension of the turtle Boremys pulchra (Testudinata, Baenidae) through at least the uppermost Cretaceous

New material of the derived baenid turtle Boremys pulchra from the Hell Creek Formation of Montana extends the stratigraphic range of the taxon through at minimum the latest Maastrichtian. Previously, the species was constrained to the Campanian of Montana and Alberta, so this extension constitutes at least 5 million years. Due to fossil reworking at the Bug Creek Anthills assemblage, where Maastrichtian and Paleocene deposits are mixed, a definitive extension for B. pulchra cannot currently include Paleocene strata. However, the presence of B. pulchra in latest Cretaceous strata, previous identification of Paleocene Boremys sp. and the general success of baenid taxa across the K–Pg boundary, make it quite plausible that B. pulchra survived the extinction event and that previously described Maastrichtian and Paleocene Boremys sp. material probably represents a new taxon. A stratigraphic extension beyond the Campanian indicates that B. pulchra survived the paleoenvironmental conditions of the latest Cretaceous, where adaptation to locally heterogeneous aquatic habitats and paleotemperature fluctuations may have facilitated latest Cretaceous and K–Pg survivorship. Additionally, ectoparasitic bore marks on the Boremys pulchra specimen described here can be attributed to the ichnotaxon Karethraichnus lakkos.

The promise of taphonomy as a nomothetic discipline: taphonomic bias in two dinosaur-bearing faunas in North America1

The fossil record of dinosaurs is a rich, if biased, one with nearly complete skeletons, partial skeletons, and isolated parts found in diverse, well-studied faunal assemblages around the world. Among the recognized biases are the preferential preservation of large dinosaurs and the systematic underrepresentation of small dinosaurs. Such biases have been quantitatively described in the Upper Cretaceous (Campanian) Dinosaur Park Formation of Alberta, where large, nearly complete dinosaurs were found and described early in collecting history and small, very incomplete dinosaurs were found and described later. This pattern, apparently replicated in the Maastrichtian Hell Creek Formation of Montana, is so striking that it begs the question of whether this is a nomothetic principle for the preservation of dinosaur faunas elsewhere. We tested this hypothesis by analyzing the very well-studied dinosaur fauna of the Upper Jurassic (Kimmeridgian) Morrison Formation of the western United States. The Morrison Formation fails to show any correlation between body size and completeness, order of discovery, or order of description. Both large and small dinosaurs of the Morrison include highly complete as well as highly incomplete taxa, and both large and small dinosaurs were discovered and described early in collecting history as well as more recently. The differences in preservation between the Dinosaur Park Formation and the Morrison Formation are so striking that we posit a Dinosaur Park model of dinosaur fossil preservation and a Morrison model. Future study will show whether either or both represent durable nomothetic models for dinosaur fossil preservation.

A systematic reappraisal and quantitative study of the nonmarine teleost fishes from the late Maastrichtian of the Western Interior of North America: evidence from vertebrate microfossil localities1

The diversity and distribution of nonmarine teleost fishes in the Western Interior of North America during the late Maastrichtian is documented based on isolated elements from vertebrate microfossil localities in the Hell Creek Formation of Montana, the Lance Formation of Wyoming, and the Scollard Formation of Alberta. A minimum of 20 taxa are recognized based on >1900 abdominal centra and tooth-bearing elements. These include two elopomorphs, six osteoglossomorphs, three ostariophysans, one esocid, six acanthomorphs, and two taxa of unknown relationships. These assemblages differ from late Campanian assemblages in the absence of the Clupeomorpha and the presence of the Perciformes. Within the Hell Creek Formation, we record patterns in the relative abundances of the most abundant taxa leading up to the Cretaceous–Paleogene (K–Pg) boundary. Most notably, acanthomorphs increased in abundance upsection, whereas a group of osteoglossomorphs, represented by Coriops and (or) Lopadichthys, concurrently decreased in abundance. Conversely, some teleosts exhibited more stable or slightly fluctuating relative abundances throughout the formation (Wilsonichthyidae, Esocidae). These late Maastrichtian teleost assemblages are of higher diversity than an early Eocene assemblage from Wyoming preserved under similar taphonomic conditions. This pattern suggests either that lower Cenozoic deposits in the Western Interior are insufficiently sampled or that the K–Pg mass extinction event adversely affected nonmarine teleosts.

Tooth replacement in a specimen of Tyrannosaurus rex (Dinosauria, Theropoda) from the Hell Creek Formation (Maastrichtian), Montana

ABSTRACT A well-preserved skull of Tyrannosaurus rex from the Late Cretaceous (Maastrichtian) Hell Creek Formation of Montana is used to document its tooth replacement with the help of high-resolution medical CT scanning. A pattern of retaining only one single replacement tooth before replacing the functional tooth is reconstructed for most alveoli of the jaws, with exception of the second alveolus of the left maxilla, and the fourth alveolus of the right dentary, where another, second replacement tooth can be seen. The youngest replacement teeth have only a thin layer of enamel around the tooth crown, the larger replacement teeth have thicker enamel, fortified at the apex as in the functional teeth. The Zahnreihen approach allows to reconstruct a z-spacing of 2 in the upper jaws, whereas the right dentary displays a z-spacing of 1 in the rostral and distal third of the tooth row and a z-spacing of 2 in its medial part. In MB.R.91216, teeth of the upper jaws and the medial part of the dentary were probably replaced in a regular alternating pattern between odd and even teeth, whereas the mesial and distal teeth of the dentary were replaced simultaneously.

First high-precision U–Pb CA–ID–TIMS age for the Battle Formation (Upper Cretaceous), Red Deer River valley, Alberta, Canada: implications for ages, correlations, and dinosaur biostratigraphy of the Scollard, Frenchman, and Hell Creek formations

The Battle Formation (BFm) is a widespread Upper Cretaceous marker horizon in western Canada that records a time of low sediment input and marks the boundary between the Edmontonian and Lancian land-vertebrate ages. Here, we present the first high-precision U–Pb CA–ID–TIMS age of 66.936 ± 0.047/0.060/0.140 Ma for the Battle bentonite, an altered vitric ash in the upper portion of the BFm at Knudsen’s Farm in the Red Deer River valley of Alberta. This age supersedes those previously reported, confirms that rates of sediment accumulation for the formation were very low (∼1.40 cm/ka), and allows us to interpolate an age range of ∼66.88–67.20 Ma for the BFm. Our data also provide a maximum age of ∼66.88 Ma for the base of the overlying Scollard Formation, a dinosaur-rich unit. We combine our age data with calibrated magneto- and palynostratigraphic data to assess chronostratigraphic correlations among the Scollard Formation of Alberta, the Frenchman Formation of Saskatchewan, and the Hell Creek Formation in eastern Montana. Whereas the combined data support previous interpretations that equate the age ranges of the Scollard Formation, Frenchman Formation, and the upper one third of the Hell Creek Formation in eastern Montana, they also indicate that all of the lower one third (L3) and part of the middle one third (M3) of the Hell Creek Formation in Montana are chronostratigraphically equivalent to all or part of the sub-BFm unconformity and the BFm in Alberta. Accordingly, a minimum age of ∼67.20 Ma is assessed for the base of the Hell Creek Formation in its type area.

A juvenile cf. Edmontosaurus annectens (Ornithischia, Hadrosauridae) femur documents a poorly represented growth stage for this taxon

A nearly complete, but isolated, femur of a small hadrosaurid from the Hell Creek Formation of Montana is tentatively referred to Edmontosaurus annectens. At 28 cm long, the element can be classified as that from an “early juvenile” individual, approximately 24 percent of the maximum known femur length for this species. Specimens from this size range and age class have not been described previously for E. annectens. Notable trends with increasing body size include increasingly distinct separation of the femoral head and greater trochanter, relative increase in the size of the cranial trochanter, a slight reduction in the relative breadth of the fourth trochanter, and a relative increase in the prominence of the cranial intercondylar groove. The gross profile of the femoral shaft is fairly consistent between the smallest and largest individuals. Although an ontogenetic change from relatively symmetrical to an asymmetrical shape in the fourth trochanter has been suggested previously, the new juvenile specimen shows an asymmetric fourth trochanter. Thus, there may not be a consistent ontogenetic pattern in trochanteric morphology. An isometric relationship between femoral circumference and femoral length is confirmed for Edmontosaurus. Overall ontogenetic trends in the femur of Edmontosaurus are concordant with patterns seen in other Hadrosauridae, supporting a fairly conserved pattern of development for this element within the clade.

The shell morphology of the latest Cretaceous (Maastrichtian) trionychid turtle Helopanoplia distincta.

BackgroundHelopanoplia distincta is an extinct soft-shelled turtle (Pan-Trionychidae) for which the type specimen is a fragmentary costal and the inguinal notch portion of the left hypoplastron from the Late Cretaceous (Maastrichtian) Lance Formation of Wyoming, USA that bear a distinct surface sculpture pattern consisting of raised tubercles. Over the course of the past few decades, a number of additional, fragmentary specimens from the Late Cretaceous (Maastrichtian) Hell Creek Formation of Montana and North Dakota have been referred to this taxon based on the presence of these tubercles, but a more complete understanding of the anatomy and phylogenetic relationships of this distinctive soft-shelled turtle is still outstanding.MethodsWe here figure and describe shell remains of eight fossils referable to Helopanoplia distincta from the Hell Creek Formation of Montana and North Dakota that, in combination, document nearly all aspects of the shell morphology of this taxon. We furthermore explore the relationships of this fossil turtle by inserting it into a modified phylogenetic analysis of pan-trionychid relationships.ResultsThe new fossil material thoroughly supports the validity of Helopanoplia distincta. In addition to its unique surface sculpture pattern, this turtle can be diagnosed relative to all other named pan-trionychids by the presence of a distinct corner along the margin of costals II, the complete covering of costal ribs I–VI by metaplastic bone, midline contact of the main plastral elements, hyoplastral shoulder, presence of a lateral, upturned margin on the hyo/hypoplastron that is covered dorsally and laterally by sculptured metaplastic bone, a single, lateral hyoplastral process, and the apomorphic presence of fine scallops along the margin of costals VIII, formation of a laterally embraced, rounded nuchal, anteriorly rounded costals I, distally expanded costals II, and narrow costals VII. A phylogenetic analysis places Helopanoplia distincta as sister to the clade formed by Plastomenus thomasii and Hutchemys spp., thereby confirming its identity as a plastomenid. The vast majority of Helopanoplia distincta material has been recovered from fine-grained overbank deposits, thereby supporting the hypothesis that this turtle favored ponded waters.

A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae

ABSTRACTOur understanding of caenagnathids has benefited from recent discoveries, including nearly complete skeletons from the Hell Creek Formation of Montana. However, their phylogenetic relationships remain unclear. A new specimen from the Horseshoe Canyon Formation of Alberta has implications for the phylogeny and paleobiology of these creatures. The partial skeleton is articulated and includes a mandible, a full cervical and dorsal series of vertebrae, a right pectoral girdle and arm, a sternum, gastralia, a partial ilium, and a partial hind limb. The mandible is edentulous and the articular ridge is intermediate in form between Caenagnathus collinsi and Chirostenotes pergracilis. The neck is long and composed of at least 11 well-pneumatized cervical vertebrae with fused cervical ribs. The dorsal ribs have finger-like uncinate processes dissimilar in shape to those of other oviraptorosaurs. The pectoral girdle is large and typically maniraptoran, except that the glenoid of the scapulocoracoid faces laterally instead of posteroventrally. The arm is well muscled and can be interpreted to have been a pennibrachium, as indicated by ulnar papillae on the ulna. The manus is characterized by a short first metacarpal but an elongate phalanx I-1 and oviraptorid-like phalangeal proportions in the second digit. These and other features indicate that the specimen represents a new taxon, Apatoraptor pennatus, gen. et sp. nov. Phylogenetic analysis resolves the complicated relationships of Caenagnathidae and allows the evolution of display features to be traced throughout Oviraptorosauria.http://zoobank.org/urn:lsid:zoobank.org:pub:122957D7-F65F-4A2C-86AF-386AE8AAF2C4SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVPCitation for this article: Funston, G. F., and P. J. Currie. 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1160910.

A New Specimen of the Controversial Chasmosaurine Torosaurus latus (Dinosauria: Ceratopsidae) from the Upper Cretaceous Hell Creek Formation of Montana.

Torosaurus latus is an uncommon and contentious taxon of chasmosaurine ceratopsid known from several upper Maastrichtian units in western North America. We describe a partial parietal of To. latus from the Hell Creek Formation of Montana. Although the specimen’s ontogenetic maturity means that it cannot inform the ongoing debate over whether To. latus is the old adult form of the contemporary Triceratops, the specimen is one of the best-preserved To. latus parietals and supplements previous descriptions.

A new crocodylian (Eusuchia) from the uppermost Cretaceous of Alberta, Canada

Albertosuchus knudsenii gen. et sp. nov. is described on the basis of an incomplete skeleton from the lower part (uppermost Cretaceous) of the Scollard Formation, southern Alberta, Canada. It is probably a crocodyloid, the only Canadian representative of the group and also the sole crocodylian known during the latest Cretaceous in Canada. Within the Crocodyloidea, A. knudsenii is one of the most basal forms and shares a sister-group relationship with the European genus Arenysuchus. However, it needs to be emphasized that because of its incompleteness, phylogenetic relationships proposed here for A. knudsenii may change when better-preserved specimens are found. Albertosuchus knudsenii differs from other crocodyloids in having no premaxilla-maxillary notch, a very short mandibular symphysis with the involvement of the splenial, and an extremely deep hypapophysis on the last cervical and anterior dorsal vertebrae. The discovery of A. knudsenii may expand the geographical range of the Crocodyloidea into Canada during the Cretaceous. Crocodylians in the lower part of the Scollard Formation are of low diversity compared with that of the corresponding Hell Creek Formation of Montana and Lance Formation of Wyoming, both farther to the south in the USA.

A juvenile Edmontosaurus from the late Maastrichtian (Cretaceous) of North America: Implications for ontogeny and phylogenetic inference in saurolophine dinosaurs

The anatomy of an articulated juvenile specimen of the saurolophine hadrosaurid dinosaur Edmontosaurus annectens, LACM 23504, is described in detail. This individual consists of a partial skull and nearly complete articulated postcranium, collected from upper Maastrichtian strata of the Hell Creek Formation of Montana, USA. This study aims to help understand the ontogenetic morphological changes occurring in the cranium and postcranium of saurolophine hadrosaurids using E. annectens as a case study. Emphasis is placed on those morphological attributes that are ontogenetically variable via comparative osteology between the juvenile and the available adult specimens. It is observed that much of the cranial ontogenetic variation relates to the elongation of the skull and mandible. In the postcranium, most of the ontogenetic variation concentrates in the pectoral and pelvic girdles and the stylopodia. Not all the identified patterns of ontogenetic variation may be generalized to all hadrosaurids. The impact of ontogenetic variation on phylogenetically informative characters of saurolophine hadrosaurids is evaluated. It is concluded that, at least for Edmontosaurus annectens (and perhaps other saurolophine hadrosaurids), most characters used in phylogenetic inference of these animals are not affected by ontogeny. Thus, juvenile specimens are still a source of substantial character data suitable for use in phylogenetic analyses of saurolophine relationships. Nevertheless, it is recommended that ontogenetically variable characters are left as missing data in a character–taxon matrix when only juvenile material is available for a given saurolophine taxon. Scoring those characters based solely on juveniles is likely to decrease the accuracy of the phylogenetic inference.

A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America

Dromaeosaurids from the Maastrichtian of North America have a poor fossil record and are known largely from isolated teeth, which have typically been referred to taxa based on more complete material from earlier Campanian strata. An almost complete maxilla with well-preserved dentition and an associated dentary from the Hell Creek Formation of Montana are used to establish a new dromaeosaurid taxon in the latest Maastrichtian, immediately prior to the end-Cretaceous extinction event. Acheroraptor temertyorum gen. et sp. nov. is differentiated from other dromaeosaurids on the basis of a hypertrophied postantral wall that projects posteriorly into the antorbital fenestra, a maxillary fenestra positioned low in the antorbital fossa and directly posterior to the promaxillary fenestra, and distinctive dentition with marked apicobasal ridges. The new material allows a dromaeosaurid from the Maastrichtian of North America to be placed within a phylogenetic framework for the first time. Phylogenetic analysis suggests Acheroraptor is a velociraptorine that is more closely related to Asian dromaeosaurids, including Tsaagan and Velociraptor, than it is to Dromaeosaurus, Saurornitholestes, or any other taxon from North America. As part of the Lancian Tyrannosaurus-Triceratops fauna, A. temertyorum is the latest occurring dromaeosaurid. Its relationships and occurrence suggest a complex historical biogeographic scenario that involved multiple, bi-directional faunal interchanges between Asia and North America during the Late Cretaceous.

New Examples ofTyrannosaurus rexfrom the Lance Formation of Wyoming, United States

The dinosaur remains from the Upper Cretaceous Lance Formation of Wyoming, USA, have been known since 1870. Most of the fossils that are attributed to theropod dinosaurs are fragmentary skeletons consisting of cranial, axial and appendicular elements, and many isolated teeth are also known. The largest known theropod of the Lance Formation dinosaur fauna is Tyrannosaurus rex. Numerous specimens of this taxon are known from the Maastrichtian strata of the northwestern United States and southwestern Canada. In 1890 and 1892, two pedal phalanges of the left foot and several isolated teeth identified here as belonging to Tyrannosaurus were discovered near Lusk, Wyoming, by John Bell Hatcher and the prospecting party from Yale Univeristy's Peabody Museum of Natural History in New Haven, Connecticut, USA. The discovery of these fossils is significant. One of the first discoveries of Tyrannosaurus from Wyoming, it predates all other discoveries and naming of this taxon in the coeval Hell Creek Formation of Montana, North Dakota and South Dakota, USA, and the Willow Creek and Scollard formations of the province of Alberta, Canada. Additionally, a well-preserved anterior end of the right dentary with teeth discovered in 1947 of a large Tyrannosaurus from the Lance Formation of the Bighorn basin, Wyoming, is described. The discovery of this specimen is also significant, because it is the first occurrence of Tyrannosaurus in the upper coastal plain paleoenvironments located farther inland away from the coastline of the Western Interior Seaway. Previous discoveries of Tyrannosaurus fossil remains from the Lance Formation were from strata deposited near the coastline of the Western Interior Seaway. The discovery of these specimens and their lack of significant morphologic divergence from other specimens of T. rex suggest that this taxon is the only large-bodied theropod that is currently known from the Upper Cretaceous (late Maastrichtian) of northwestern North America.

Soft sheets of fibrillar bone from a fossil of the supraorbital horn of the dinosaur Triceratops horridus

Soft fibrillar bone tissues were obtained from a supraorbital horn of Triceratops horridus collected at the Hell Creek Formation in Montana, USA. Soft material was present in pre and post-decalcified bone. Horn material yielded numerous small sheets of lamellar bone matrix. This matrix possessed visible microstructures consistent with lamellar bone osteocytes. Some sheets of soft tissue had multiple layers of intact tissues with osteocyte-like structures featuring filipodial-like interconnections and secondary branching. Both oblate and stellate types of osteocyte-like cells were present in sheets of soft tissues and exhibited organelle-like microstructures. SEM analysis yielded osteocyte-like cells featuring filipodial extensions of 18–20μm in length. Filipodial extensions were delicate and showed no evidence of any permineralization or crystallization artifact and therefore were interpreted to be soft. This is the first report of sheets of soft tissues from Triceratops horn bearing layers of osteocytes, and extends the range and type of dinosaur specimens known to contain non-fossilized material in bone matrix.

Torosaurus Marsh, 1891, is Triceratops Marsh, 1889 (Ceratopsidae: Chasmosaurinae): synonymy through ontogeny

ABSTRACT Although they have been considered distinct genera for over a century, ontogenetic analyses reveal that Triceratops and “Torosaurus” actually represent growth stages of a single genus. Major changes in cranial morphology—including the opening of parietal fenestrae and the elongation of the squamosals—occur rapidly, very late in Triceratops ontogeny and result in the characteristic ‘Torosaurus’ morphology. This report presents the results of a 10-year field study of the dinosaurs of the Hell Creek Formation in Montana and is based on a collection of over 50 specimens of Triceratops, including over 30 skulls, which have been amassed in that time, in addition to specimens from numerous other North American museums. This large sample of individuals reveals the full ontogenetic spectrum of Triceratops. The synonymy of Triceratops and ‘Torosaurus’ contributes to an unfolding view of extremely reduced dinosaur diversity just before the end of the Mesozoic Era.

An Exquisitely Preserved Skeleton Representing a Primitive Sturgeon from the Upper Cretaceous Judith River Formation of Montana (Acipenseriformes: Acipenseridae: N. Gen. and Sp.)

A new species of sturgeon (Acipenseridae), †Psammorhynchus longipinnisn. gen. and sp., is described based on a single well-preserved specimen from the Upper Cretaceous Judith River Formation of Montana. Although based on only a single individual, the osteological detail we are able to describe for this species is extensive because of the completeness of the specimen, the relatively three-dimensional preservation, and the soft, loosely consolidated sandstone matrix in which it was contained (allowing some dissection of the specimen to reveal the internal bones of the head region). We prepared the specimen completely out of the matrix, and described the head region in stages of “dissection.” †Psammorhynchus longipinnisis the only fossil sturgeon to date that provides a relatively complete, reliable look at its osteology. Although there have previously been numerous and widespread reports of fossil sturgeons, the material is notoriously fragmentary and poorly preserved, consisting mostly of isolated scutes or pectoral spines (summarized here in an appendix) and it is of very limited use to phylogenetic studies of living sturgeon taxa. The only other described articulated skeleton of a fossil sturgeon is †Protoscaphirhynchus squamosusWilimovsky from the Upper Cretaceous Hell Creek Formation of Montana. That specimen (the only one known for the species) is a badly preserved, very poorly prepared skeleton showing little detail of the head region. It is interesting to note that the only known articulated fossil sturgeon skeletons (†Psammorhynchus longipinnisn. gen. and sp.; †Protoscaphirhynchus squamosusWilimovsky; and a third, yet undescribed specimen) were apparently each independently discovered in the abdominal regions of hadrosaurian dinosaurs (as was the Cretaceous paddlefish †Paleopsephurus wilsoniMacAlpin). Perhaps hadrosaur carcasses provided the right combination of taphonomic factors to protect acipenseriform carcasses from damage in high energy aquatic environments predominantly preferred by sturgeons. In any case, based on known specimens, sediments within hadrosaurs appear to have been the main preservational medium for articulated fossil sturgeon skeletons in North America.†Psammorhynchus longipinnisappears to be the sister taxon to the rest of Acipenseridae based on our preliminary analysis here. We will reevaluate this relationship elsewhere in our (Hilton, Grande, and Bemis) ongoing study of fossil and living Acipenseridae.

Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America

The tooth taxon Aublysodon mirandus was reinstated following the collection of nondenticulate tyrannosaurid premaxillary teeth from late Maastrichtian deposits in western North America. A small skull from the Hell Creek Formation of Montana (the ‘Jordan theropod’, LACM 28471), that was associated with a nondenticulate premaxillary tooth, was referred to Aublysodon and the diagnosis was revised to include cranial bones. However, the ‘premaxillary’ tooth of the specimen is actually a maxillary tooth. The small size of Aublysodon crowns, and evidence that some denticles develop late in growth in theropods, indicates that the nondenticulate condition represents immaturity. Therefore, Aublysodon is a nomen dubium. The Jordan theropod was recently designated as the type specimen of Stygivenator molnari. A tyrannosaurid from the Hell Creek Formation of Montana (LACM 23845) was first referred to Albertosaurus cf. A. lancensis and then later became the type specimen of Dinotyrannus megagracilis. On the basis of shared derived characters and a quantitative reconstruction of the growth series of Tyrannosaurus rex, the type specimens of S. molnari and D. megagracilis are juvenile and subadult specimens of T. rex, respectively. There is currently evidence for only one tyrannosaurid species in the late Maastrichtian of western North America: T. rex. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142, 479–523.

A review of nonmarine turtles from the Late Cretaceous of Alberta

The Late Cretaceous of Alberta preserves one of the most complete records of fossil turtles within a single geographic area in North America. The Cenomanian Dunvegan Formation contains the earliest record of the family Trionychidae in North America. The Santonian Milk River Formation contains a minimum of ten taxa with Adocus, a small trionychid, and a member of the Solemydidae being the most abundant. Diversity remains high in the mid-Campanian Judith River Group. The solemyidid last occurs in the basal beds of the Judith River Group. A member of the Macrobaenidae first occurs in the Dinosaur Park Formation, the uppermost formation in the Judith River Group. Turtles diversity is low in the late Campanian lower Horseshoe Canyon Formation, and they are absent in the early Maastrichtian upper Horseshoe Canyon Formation. Diversity increases in the late Maastrichtian Scollard Formation, although it is much less than in the contemporaneous Hell Creek Formation of Montana. Two of the taxa present in the Scollard Formation, Compsemys and Plastomenus, occur in late Campanian or early Maastrichtian formations in more southerly areas of North America. The changes in turtle diversity through the Campanian and Maastrichtian are interpreted as a result of shifts in a latitudinal turtle diversity gradient resulting from changes in climate. Based on this interpretation a decrease in temperature from the mid-Campanian to early Maastrichtian, followed by a rapid increase at the beginning of the late Maastrichtian is supported.

A new genus of derived pachycephalosaurian from western North America

ABSTRACT Pachycephalosaurian specimens from the Upper Cretaceous (upper Campanian) Kirtland Formation of northwestern New Mexico include a partial skull that preserves much of the basicranium. It represents a new genus and species, Sphaerotholus goodwini. A new species, S. buchholtzae, from the upper Maastrichtian Hell Creek Formation of Montana is referred to this genus. Additional New Mexico pachycephalosaurians include a partial dentary with associated skull material that is tentatively referred to S. goodwini and a partial frontoparietal that is referred to Pachycephalosauridae incertae sedis. On the basis of a hypothetical growth series of Stegoceras, we excluded characters based on dome development and suture closure from a cladistic analysis of pachycephalosaurian relationships. Ornatotholus browni, the only putative “flat-headed” pachycephalosaurian from North America, is considered a nomen dubium and may represent a juvenile Stegoceras. Gravitholus albertae is an adult Stegoceras sp.; Stegoceras edmontonense is a nomen dubium and is referred to cf. Sphaerotholus sp. Based on the results of a quantitative cladistic analysis, Stegoceras (including Stegoceras breve, S. lambei, S. stembergi, and UCMP 130051) is the sister taxon to all other domed pachycephalosaurians. Derived pachycephalosaurids consist of two principal clades: a lineage that includes Stygimoloch, Pachycephalosaurus, and Sphaerotholus and a lineage represented by the Asian taxa, Tylocephale and Prenocephale. With biogeographic occurrences mapped onto the phytogeny, a single dispersal event from Asia into North America, followed by dispersal of Prenocephale and Tylocephale into Asia prior to the late Campanian is indicated (ACCTRAN) or two independent dispersals into North America prior to the late Campanian is indicated (DELTRAN). Pachycephalosaurian phylogeny does not support Asian-North American contiguity throughout the Campanian and Maastrichtian.