Wapiti Formation Research Articles

Enigmatic vertebrate swimming trace fossils from the Wapiti Formation, Alberta, Canada, and their implications for paleoenvironmental reconstruction

Vertebrate ichnofossil research, when compared with invertebrate ichnological analyses, tends to put greater emphasis on identifying the trace makers rather than associating the trace fossils with animal behaviors to interpret local paleoenvironmental conditions. In 2011, several sandstone blocks bearing unusual deformational structures were collected from the Wapiti Formation exposed near Red Willow Falls, Alberta, Canada. The sandstone blocks were analyzed for their sedimentary characteristics and the morphology of the enigmatic structures in order to identify their origins and their significance for paleoenvironmental interpretation. The deformational features are interpreted to be the swim tracks of small vertebrates attempting to escape a flash flood event. The sedimentology of the sandstone suggests a rapid increase in water flow in an overbank setting, and the orientation of the traces indicate that the trace maker was initially moving against, and then moving perpendicular to, the current. This is likely the first case of a trace fossil exhibiting clear association between vertebrates and flash flood events. As such, these traces provide an excellent example of the utility of vertebrate trace fossils for assessing organism response to changing environmental conditions, thus facilitating interpretation of local geological events.

New Lizard Specimens from the Campanian Wapiti Formation of Alberta, Canada

Reported lizard material from the Wapiti Formation (central-western Alberta, Canada) is limited to fragmentary remains of Kleskunsaurus grandeprairiensis and Socognathus unicuspis, a partial dentary attributed to Chamops cf. C. segnis, and a vertebra reportedly comparable to those of the much larger lizard Palaeosaniwa canadensis. P. canadensis is a Late Cretaceous North American member of Monstersauria, a Mesozoic and Cenozoic anguimorph group represented today by five species of Heloderma. Here, we document new squamate material from the DC Bonebed locality (Wapiti Unit 3; Campanian), including a right frontal identified as cf. P. canadensis and a taxonomically indeterminate squamate astragalocalcaneum. A partial skeleton from the Two Medicine Formation of Montana provisionally attributed to P. canadensis has a frontal resembling the corresponding element from the DC Bonebed in overall shape, in having narrowly separated facets for the prefrontal and postorbitofrontal, and in bearing osteoderms similar to the DC specimen’s in ornamentation and configuration. The Two Medicine and DC specimens differ from a roughly contemporaneous frontal from southern Alberta referred to the monstersaur Labrodioctes montanensis. The DC specimen confirms the presence of monstersaurian squamates in the Wapiti Formation, representing the northernmost record of any definitive Late Cretaceous monstersaur to date.

Filling the Bearpaw gap: Evidence for palaeoenvironment-driven taxon distribution in a diverse, non-marine ecosystem from the late Campanian of west-Central Alberta, Canada

Patterns of Late Cretaceous terrestrial vertebrate diversity across North America have been interpreted primarily in terms of biogeographic provincialism driven by latitude or coastal-inland habitat gradients. A major difficulty in determining the influence of these two gradients is the existence of some large gaps in the terrestrial fossil record, notably the ‘Bearpaw gap’ caused by a transgression of the inland Bearpaw Seaway during the latter part of the Campanian. In this context, the terrestrial fauna preserved in the Campanian deposits of the Wapiti Formation (west-central Alberta, Canada) is crucial for addressing the information deficit. Deposited at the edge of the palaeo-circumpolar region, Unit 3 of the strictly terrestrial Wapiti Formation (WU3) is coeval with the ‘Bearpaw gap’, a period when the terrestrial record from better-sampled areas elsewhere in Canada and the U.S.A. gives way to marine sediments. Here we show, based largely on evidence from the recently discovered DC (Dinosaur-Chelonian) Bonebed locality, that the diverse WU3 vertebrate fauna shares similarities with lowland to marginal marine ecosystems in the Oldman and Dinosaur Park formations which were deposited in southern Alberta prior to the Bearpaw gap. In addition, a major change in faunal composition demarcates the upper boundary of WU3, related to the disappearance of the Bearpaw Sea in Canada. Data presented here help, first and foremost, to bridge an ~1.2-million-year gap in the North American record of Campanian terrestrial vertebrates. Resemblances between the WU3 vertebrate fauna and slightly older assemblages from southern Alberta underscore the importance of determining the spatiotemporal changes in environmental factors (e.g., coastal proximity). The occurrence of one seemingly endemic lizard, together with differences in relative taxon abundance, suggest additional latitude-correlated factors, implicating both latitudinal and coastal-inland habitat gradients in driving the taxonomic composition of Late Cretaceous terrestrial faunas. • We introduce a new microvertebrate site from the late Campanian of western Canada. • A diverse non-marine fauna coeval with the Bearpaw Formation is discussed. • Vertebrate community is compared with others from the Campanian of Alberta. • Data help filling the information deficit related to the ‘Bearpaw Gap’.

The dinosaur tracks of Tyrants Aisle: An Upper Cretaceous ichnofauna from Unit 4 of the Wapiti Formation (upper Campanian), Alberta, Canada.

The Wapiti Formation of northwest Alberta and northeast British Columbia, Canada, preserves an Upper Cretaceous terrestrial vertebrate fauna that is latitudinally situated between those documented further north in Alaska and those from southern Alberta and the contiguous U.S.A. Therefore, the Wapiti Formation is important for identifying broad patterns in vertebrate ecology, diversity, and distribution across Laramidia during the latest Cretaceous. Tracksites are especially useful as they provide a range of palaeoecological, palaeoenvironmental, and behavioural data that are complementary to the skeletal record. Here, we describe the Tyrants Aisle locality, the largest in-situ tracksite known from the Wapiti Formation. The site occurs in the lower part of Unit 4 of the formation (~72.5 Ma, upper Campanian), exposed along the southern bank of the Redwillow River. More than 100 tracks are documented across at least three distinct track-bearing layers, which were deposited on an alluvial floodplain. Hadrosaurid tracks are most abundant, and are referable to Hadrosauropodus based on track width exceeding track length, broad digits, and rounded or bilobed heel margins. We suggest the hadrosaurid trackmaker was Edmontosaurus regalis based on stratigraphic context. Tyrannosaurids, probable troodontids, possible ornithomimids, and possible azhdarchid pterosaurs represent minor but notable elements of the ichnofauna, as the latter is unknown from skeletal remains within the Wapiti Formation, and all others are poorly represented. Possible social behaviour is inferred for some of the hadrosaurid and small theropod-like trackmakers based on trackway alignment, suitable spacing and consistent preservation. On a broad taxonomic level (i.e., family or above), ichnofaunal compositions indicate that hadrosaurids were palaeoecologically dominant across Laramidia during the late Campanian within both high-and low-latitude deposits, although the role of depositional environment requires further testing.

Taphonomy and taxonomy of a juvenile lambeosaurine (Ornithischia: Hadrosauridae) bonebed from the late Campanian Wapiti Formation of northwestern Alberta, Canada.

Hadrosaurid (duck-billed) dinosaur bonebeds are exceedingly prevalent in upper Cretaceous (Campanian–Maastrichtian) strata from the Midwest of North America (especially Alberta, Canada, and Montana, U.S.A) but are less frequently documented from more northern regions. The Wapiti Formation (Campanian–Maastrichtian) of northwestern Alberta is a largely untapped resource of terrestrial palaeontological information missing from southern Alberta due to the deposition of the marine Bearpaw Formation. In 2018, the Boreal Alberta Dinosaur Project rediscovered the Spring Creek Bonebed, which had been lost since 2002, along the northern bank of the Wapiti River, southwest of Grande Prairie. Earlier excavations and observations of the Spring Creek Bonebed suggested that the site yielded young hadrosaurines. Continued work in 2018 and 2019 recovered ~300 specimens that included a minimum of eight individuals, based on the number of right humeri. The morphology of several recovered cranial elements unequivocally supports lambeosaurine affinities, making the Spring Creek sample the first documented occurrence of lambeosaurines in the Wapiti Formation. The overall size range and histology of the bones found at the site indicate that these animals were uniformly late juveniles, suggesting that age segregation was a life history strategy among hadrosaurids. Given the considerable size attained by the Spring Creek lambeosaurines, they were probably segregated from the breeding population during nesting or caring for young, rather than due to different diet and locomotory requirements. Dynamic aspects of life history, such as age segregation, may well have contributed to the highly diverse and cosmopolitan nature of Late Cretaceous hadrosaurids.

Probable deinonychosaur tracks from the Upper Cretaceous Wapiti Formation (upper Campanian) of Alberta, Canada

AbstractLate Cretaceous tracks attributable to deinonychosaurs in North America are rare, with only one occurrence ofMenglongipusfrom Alaska and two possible, but indeterminate, occurrences reported from Mexico. Here we describe the first probable deinonychosaur tracks from Canada: a possible trackway and one isolated track on a single horizon from the Upper Cretaceous Wapiti Formation (upper Campanian) near Grande Prairie in Alberta. The presence of a relatively short digit IV differentiates these from argued dromaeosaurid tracks, suggesting the trackmaker was more likely a troodontid. Other noted characteristics of the Wapiti specimens include a rounded heel margin, the absence of a digit II proximal pad impression, and a broad, elliptical digit III. Monodactyl tracks occur in association with the didactyl tracks, mirroring similar discoveries from the Early Cretaceous Epoch of China, providing additional support for their interpretation as deinonychosaurian traces. Although we refrain from assigning the new Wapiti specimens to any ichnotaxon because of their relatively poor undertrack preservation, this discovery is an important addition to the deinonychosaur track record; it helps to fill a poorly represented geographic and temporal window in their known distribution, and demonstrates the presence of a greater North American deinonychosaur ichnodiversity than has previously been recognized.

Exploring possible ontogenetic trajectories in tyrannosaurids using tracks from the Wapiti Formation (upper Campanian) of Alberta, Canada

ABSTRACT Fossil tracks should theoretically capture differences in pedal anatomy between growth stages of the same taxon, particularly those related to the soft tissue of the foot, providing a more realistic view of pedal ontogeny than skeletal material alone. However, recognizing these ontogenetic trajectories is complicated by the influence of preservation and kinematics on track morphology, as well as the inherent difficulty of referring different tracks to a single taxon. Here, we explore differences in track morphology from a collection of tracks attributed to tyrannosaurids from Unit 4 of the Wapiti Formation (upper Campanian) in western Canada. Along with morphology, close geographic and stratigraphic associations suggest that the tracks pertain to similar tyrannosaurid trackmakers. A geometric morphometric analysis of the track outlines reveals size-dependent increase in relative track robusticity, driven primarily by an increase in ‘heel’ breadth and surface area. This relationship is lost when the dataset is expanded to include tyrannosaurid tracks globally, which we attribute to increased stratigraphic and taxonomic ‘noise’ within the global dataset that masks the tightly constrained patterns obtained from the Wapiti Formation tracks. Although there is some substrate and kinematic influence on certain aspects of track morphology, we hypothesize that the observed size-dependent relationship reflects genuine expansion in the breadth of the heel soft tissues and probably their overall surface area associated with growth. Increased pedal robusticity likely assisted with weight bearing and locomotor stability as body mass increased over ontogeny, supporting previous hypotheses that some tyrannosaurids underwent a growth-related reduction in relative agility and/or cursorial performance.

Bonebed amber as a new source of paleontological data: The case of the Pipestone Creek deposit (Upper Cretaceous), Alberta, Canada

Amber deposits and dinosaur bonebeds provide some of the most detailed sources of information on terrestrial ecosystems, but these sources have rarely been studied in tandem. The Pipestone Creek bonebed from the Campanian Wapiti Formation of Alberta, Canada, provides an opportunity to explore both data sources in the same deposit for the first time. The site has yielded an exceptional fauna dominated by abundant remains of the centrosaurine ceratopsian Pachyrhinosaurus. The initial campaigns of excavation in the 1980s led to the collateral discovery of amber, and the site became the first published source of insect-bearing amber in a dinosaur bonebed. Here we describe amber inclusions in detail and analyze the composition and geochemistry of the amber (FTIR spectroscopy and stable isotope studies) in order to draw paleoecological and paleoenvironmental inferences. We describe a feather fragment from an aquatic bird, as well as new species of Psocodea and Hymenoptera (Mymarommatidae) with ecological constraints. Indeterminate aphid and spider inclusions were also found, but preservation was not sufficient for formal description. FTIR spectroscopy suggests a botanical source among the Cupressaceae, while stable isotope data indicate that the resin-producing forest did not receive precipitation directly from the Western Interior Seaway. These initial results show that amber is a valuable source of additional information about the habitat surrounding the vertebrates found in bonebed deposits. Given the abundance and range of inclusions encountered so far, Pipestone Creek amber also appears to be a promising new source for a relatively diverse Campanian amber assemblage.

Detrital zircon geochronology of the Aycross Formation (Eocene) near Togwotee Pass, western Wind River Basin, Wyoming

The Aycross Formation is the basal unit of the Absaroka Volcanic Supergroup in the southern Absaroka Range and consists of volcanic sandstone, mudstone, breccia, tuff and conglomerate. The Aycross was deposited during the waning stages of the Laramide Orogeny and the earliest phases of volcanism in the Absaroka Range. U-Pb geo-chronology using laser ablation multicollector inductively coupled plasma mass spectrometry LA-ICP-MS was performed on detrital zircons collected from an Aycross sandstone bed at Falls Campground east of Togwotee Pass. The detrital zircon age spectrum ranged fom ca 47 to 2856 Ma. Peak ages, as indicated by the zircon age probability density plot are ca. 51, 61, and 72 Ma. Tertiary zircons were the most numerous (n = 32), accounting for 42% of the zircon ages spectrum. Of these 19 are Eocene, and 13 are Paleocene, which are unusual ages in the Wyoming-Idaho-Montana area. Mesozoic zircons (n = 21) comprise 27% of the age spectrum and range in age from 68–126 Ma; all but one being late Cretaceous in age. No Paleozoic zircons are present. Proterozoic zircons range in age from 1196–2483 Ma, and also consist of 27% of the age spectrum. The maximum depositional age of the Aycross Formation is estimated to be 50.05 +/− 0.65 Ma based on weighted mean of the eight youngest grains. The Aycross Formation detrital zircon age spectrum is distinct from that of other 49–50 Ma rocks in northwest Wyoming, which include the Hominy Peak and Wapiti Formations and Crandall Conglomerate. The Aycross must have been derived largely from distal westerly source areas, which include the late Cretaceous and Paleocene Bitteroot Lobe of the Idaho Batholith. In contrast, the middle Eocene units further to the north must have been derived from erosion of the Archean basement-cored uplift of the Laramide Foreland in southwest Montana.

A high-latitude dromaeosaurid, Boreonykus certekorum, gen. et sp. nov. (Theropoda), from the upper Campanian Wapiti Formation, west-central Alberta

ABSTRACTDromaeosaurids were rare components of most Late Cretaceous terrestrial ecosystems and are poorly known from high palaeolatitudes. New dromaeosaurid material, including a frontal and associated postcranial elements, is described from a dense monodominant ceratopsid bonebed on Pipestone Creek, near the city of Grande Prairie (Unit 3, Wapiti Formation, upper Campanian), central-western Alberta, Canada. This stratigraphic interval is significant because it records a period of terrestrial deposition at a time when much of the western interior of Canada and the United States was inundated by the Bearpaw Sea. A phylogenetic analysis recovers Boreonykus certekorum, gen. et sp. nov., as a derived eudromaeosaur, possibly within Velociraptorinae. The identification of a new dromaeosaurid from the Wapiti Formation simultaneously helps fill an important gap in the record of late Campanian dromaeosaurids, bolsters support for a partly endemic fauna within the Wapiti Formation, and potentially adds to the North American record of a predominantly Asian Velociraptorinae.http://zoobank.org/urn:lsid:zoobank.org:pub:8A64DF33-58C3-4C43-830E-78F725094A13SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVPCitation for this article: Bell, P. R., and P. J. Currie. 2015. A high-latitude dromaeosaurid, Boreonykus certekorum, gen. et sp. nov. (Theropoda), from the upper Campanian Wapiti Formation, west-central Alberta. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2015.1034359.

Taphonomy, age, and paleoecological implication of a new Pachyrhinosaurus (Dinosauria: Ceratopsidae) bonebed from the Upper Cretaceous (Campanian) Wapiti Formation of Alberta, Canada

The Grande Prairie region (Alberta, Canada) includes some of the richest Cretaceous fossil sites in North America, including the recently described bonebed of Pachyrhinosaurus lakustai at the Pipestone Creek locality. Here we describe a new multi-taxa, ceratopsian-dominated bonebed from the region, integrating taphonomic, radioisotopic, and paleoecological data. The bonebed can be traced for 107 m and has been excavated over an area of 40 m2 with an average bone density of 30–50 elements/m2. The new bonebed occurs within Unit 4 of the upper Campanian Wapiti Formation, and 40Ar/39Ar dating provides an age of 71.89 ± 0.14 Ma, thus making the site equivalent in age to the upper Drumheller Member of the lower Horseshoe Canyon Formation of central Alberta. About 88% of vertebrate remains are ceratopsian, and dromaeosaurid, hadrosaurid, troodontid, and tyrannosaurid remains have also been identified. Juvenile material, although scarce, indicates an assemblage of individuals of different ages. Specimens showed no strong preferred two-dimensional orientation but are clearly sorted vertically. Taphonomic and sedimentological interpretation support a complex pre-burial history of preserved elements as well as a depositional setting characterized by persistent waterlogged conditions as those typical of large oxbow lakes or marshy/swampy areas, as well as lacustrine settings within an alluvial plain. Being located more than 450 km inland from the paleo-coastline, the new bonebed represents one of the farthest-known inland occurrence of centrosaurines in North America, further supporting the presence of large aggregations of ceratopsian far from the coastal lowlands of the Western Interior Seaway.

The first appearance of Troodon in the Upper Cretaceous site of Danek Bonebed, and a reevaluation of troodontid quantitative tooth morphotypes

The systematic position of the small theropod dinosaur Troodon has had a confusing history and has been identified at different times as a lizard, a pachycephalosaurid, and a coelurosaurid. Troodon is now most commonly considered as a sister taxon to Dromeaosauridae, within Maniraptora. This study records the first evidence of Troodon in the Danek bonebed, an organic-rich member of the upper Campanian Horseshoe Canyon Formation. A single tooth (UALVP 55489) was recovered during the 2012 dig season, and it is exceptionally well-preserved. Its discovery prompted a reevaluation of variation within troodontid teeth in Alberta; it is compared here with 110 troodontid teeth from the Dinosaur Park, Horseshoe Canyon, and Wapiti formations. The results show that no distinctive morphotypes can be separated, in contrast to the results of other studies. This suggests that either troodontid teeth are not sufficiently different for identifying different species or that only one troodontid taxon is present in the Campanian of Alberta.

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.

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.

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.

A possible pterosaur manus track from the Late Cretaceous of Alberta

Bell, P.R., Fanti, F. & Sissons, R. 2012: A possible pterosaur manus track from the LateCretaceous of Alberta. Lethaia, DOI: 10.1111/let.12006.An isolated track from the latest Campanian–Maastrichtian Wapiti Formation (Alber-ta) is tentatively identified as a pterosaur manus print. The basic digital formula (digitI < digit II < digit III) is consistent with the plesiomorphic condition in pterosaurs.The track measures 25.5 cm in length, making it the largest putative pterosaur manusprint (estimated wingspan 7.7 m) from North America. Although precise chrono-stratigraphic data are lacking, sedimentary evidence indicates the track comes from amid- to late Campanian fluvial deposits that accumulated approximately 400 km fromthe closest shoreline within a high-latitude (65°N) setting.

A diverse, high-latitude ichnofauna from the Late Cretaceous Wapiti Formation, Alberta, Canada

The Wapiti Formation in west-central Alberta preserves one of the most diverse Late Cretaceous terrestrial track records yet identified in Canada. At least seven morphotypes are recognized and attributed to mammals, small reptiles or amphibians, tyrannosaurids, medium-sized theropods, hadrosaurids, and ankylosaurs. Most tracks occur isolated on slump blocks associated with latest Campanian (Wapiti Formation unit 4) exposures found along Pipestone Creek and Red Willow River. With the possible exception of hadrosaurids, tracks provide some of the most compelling evidence for the occurrence of such taxa within the Wapiti Formation ecosystem. The apparent absence of ceratopsian tracks is surprising considering their bones are abundantly preserved in nearby monodominant bonebeds. The overall faunal signal represented by the Wapiti Formation trackmakers is typical of and consistent with other coeval assemblages in similar environments. The Wapiti Formation tracks, combined with the known fossil bone record, provide another data point in a growing palaeobiogeographical picture of the dinosaur faunas of high-latitude northwestern North America during the Late Cretaceous.

Fossil mayfly larvae (Ephemeroptera, cf. Heptageniidae) from the Late Cretaceous Wapiti Formation, Alberta, Canada

Fossil mayfly larvae (cf. Heptageniidae) are reported for the first time in the Cretaceous of Canada. The new fossils come from the latest Campanian part of the Wapiti Formation, which crops out in west-central Alberta, near the British Columbia border. These sediments represent mixed lentic and lotic fluvial environments consistent with modern heptageniid ecology. This discovery helps fill a significant temporal gap in heptageniid evolution between the Eocene and their earliest appearance in the Turonian.

Late Maastrichtian paleovalley systems in west-central Alberta: Mapping the Battle Formation in the subsurface

Abstract The Maastrichtian Battle Formation forms an important stratigraphic marker in the nonmarine Upper Cretaceous succession across a wide area of west-central Alberta. This mudstone-dominated unit, with a maximum thickness of 18.0 m, is distinctive both at outcrop and in down-hole geophysical well logs. New stratigraphic picks for the top and base of the Battle Formation in 8935 wells, together with data-points from a further 608 wells in which the Battle Formation is absent, were used to generate a regional isopach map. In combination with closely spaced well-log cross-sections, the isopach map allows delineation of a series of paleovalley systems incised into the Battle Formation and filled by sandy fluvial deposits of the overlying Scollard Formation. Across wide areas within the paleovalleys, the Battle Formation has been completely removed and Scollard Formation strata rest directly on the underlying Horseshoe Canyon or Wapiti formations. Paleovalley trends appear to have been broadly similar to those of major present-day river systems in the region. A paleo-Athabasca River system with a main trunk valley up to 30 km across trends southwest to northeast in the area between Edson and Whitecourt. Farther south, a well-defined paleovalley system with a main valley up to 10 km wide trending northwest to southeast in the area between Red Deer and Drumheller can be considered as a precursor to the present Red Deer River drainage system. A less well-defined paleovalley trends northwest to southeast in the area of the present Bow River to the east of Calgary.

Borioteiioidean lizard skulls from Kleskun Hill (Wapiti Formation; upper Campanian), west-central Alberta, Canada

ABSTRACT New material of borioteiioidean lizards (Squamata: Scincomorpha) from west-central Alberta, Canada, represent the first and northernmost record of multiple articulated skull elements from the Cretaceous of North America. Specimens were recovered from the fluvial beds of the Wapiti Formation (Campanian) within a bentonitic paleosol exposed at the Kleskun Hill Park, east of the city of Grande Prairie. Such beds accumulated during the maximum transgression of the Bearpaw Seaway (73–74 Ma), thus providing crucial information on lizard faunas during a time interval represented in most of coeval North American deposits by marine strata. Cranial material ascribed to Socognathus unicuspis give the occasion for a revision of the taxon with respect to osteologically better-known Polyglyphanodon sternbergi from the Late Cretaceous of Utah as well as a comparison with several lizards reported from coeval strata of Mongolia. Furthermore, a new scincomorphan lizard, Kleskunsaurus grandeprairiensis, gen. et sp. nov., is described. Socognathus unicuspis is assigned to Chamopsiidae, taxon nov., which also includes Chamops, Leptochamops, and several other morphologically similar taxa from the Cretaceous of North America.