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.     

Parataxonomy and palaeobiogeographic significance of dinosaur eggshell fragments from the Upper Cretaceous strata of the Cauvery Basin,South India

Abstract

In the past, fossilised dinosaur eggshells have been extensively documented from the Upper Cretaceous Lameta Formation of Central India and as many as nine oospecies are known at present from this formation. Compared to this, only one dinosaur oospecies has been described from the Cretaceous succession of the Cauvery Basin. However, the first fossil egg from India, identified as a chelonian egg, was documented from the Aptian – Albian Karai Formation of the Cauvery Basin in 1957. Following this, a solitary titanosaurid dinosaur egg was described from the Upper Cretaceous (Lower Maastrichtian) Kallankuruchhi Formation, Cauvery Basin in 1996. More recently, we have recovered isolated eggshell fragments from the marine part of the Upper Cretaceous (Late Maastrichtian) Kallamedu Formation. Based on eggshell morphology, microstructure and ultrastructure, these eggshell fragments are assigned to the oospecies Fusioolithus baghensis. The new find from the Cauvery Basin is important from palaeobiogeographic point of view as the oofamily Fusioolithidae is found in the Upper Cretaceous strata of India, France, Argentina and Morocco. Based on the common occurrence of similar oospecies in South America, Africa, Europe and India, a Late Cretaceous palaeobiogeographic connection between India and South America as well as Europe via Africa is suggested.     

Latest Cretaceous hadrosauroid (Dinosauria: Ornithopoda) remains from Bulgaria

Disarticulated dinosaur bones have been discovered in a fossiliferous lens in the Labirinta Cave, southwest of the town of Cherven Bryag, in NW Bulgaria. This cave is formed within marine limestones belonging to the Kajlâka Formation of Latest Cretaceous age. Associated fossils and Sr isotopy suggest that the fossiliferous sediments belong to the uppermost part of the Upper Maastrichtian. The dinosaur bones discovered in this lens include the distal portion of a left femur, a right tibia, the proximal part of a right fibula, a left metatarsal II, the second or third phalanx of a left pedal digit IV, the proximal end of a second metacarpal, and a caudal centrum. All the bones undoubtedly belong to ornithopod dinosaurs and more accurately to representatives of the hadrosauroid clade. All belong to small-sized individuals, although it cannot be assessed whether they belong to juveniles or small-sized adults, pending histological analyses. Hadrosauroid remains have already been discovered in Late Maastrichtian marine sediments from western, central and eastern Europe, reflecting the abundance of these dinosaurs in correlative continental deposits. Indeed, hadrosauroids were apparently the dominating herbivorous dinosaurs in Eurasia by Late Maastrichtian time.     

Late Cretaceous sharks from Cuba,first record of Serratolamna serrata (Agassiz) (Lamniformes,Serratolamnidae)

Only one species of Elasmobranchii, Ptychodus cyclodontis Mutter, Iturralde-Vinent and Carmona (2005), has been reported so far from the Late Cretaceous of Cuba. Herein we describe the first record of a Maastrichtian Serratolamna serrata (Agassiz, 1843) as well as non-diagnostic remains which include a tooth referred to a lamniform shark and an isolated vertebra of an indeterminate elasmobranch. These fossils expand the temporal distribution of Cretaceous fossil sharks known from Cuba and increase our understanding of the group’s fossil diversity.     

A temperate palaeodiversity peak in Mesozoic dinosaurs and evidence for Late Cretaceous geographical partitioning

Aim Modern biodiversity peaks in the tropics and declines poleward, a pattern that is potentially driven by climate. Although this latitudinal biodiversity gradient (LBG) also characterizes the marine invertebrate fossil record, distributions of ancient terrestrial faunas are poorly understood. This study utilizes data on the dinosaur fossil record to examine spatial patterns in terrestrial biodiversity throughout the Mesozoic. Location We compiled data on fossil occurrences across the globe. Methods We compiled a comprehensive dataset of Mesozoic dinosaur genera (738), including birds. Following the utilization of sampling standardization techniques to mediate for the uneven sampling of the fossil record, we constructed latitudinal patterns of biodiversity from this dataset. Results The dominant group of Mesozoic terrestrial vertebrates did not conform to the modern LBG. Instead, dinosaur diversity was highest at temperate palaeolatitudes throughout the 160 million year span of dinosaurian evolutionary history. Latitudinal diversity correlates strongly with the distribution of land area. Late Cretaceous sauropods and ornithischians exhibit disparate LBGs. Main conclusions The continuity of the palaeotemperate peak in dinosaur diversity indicates a diminished role for climate on the Mesozoic LBG; instead, dinosaur diversity may have been driven by the amount of land area among latitudinal belts. There is no evidence that the tropics acted as a cradle for dinosaur diversity. Geographical partitioning among major clades of herbivorous dinosaurs in the Late Cretaceous may result from the advanced stages of continental fragmentation and/or differing responses to increasing latitudinal climatic zonation. Our results suggest that the modern‐day LBG on land was only established 30 million years ago, following a significant post‐Eocene recalibration, potentially related to increased seasonality.     

Small Theropod Teeth from the Late Cretaceous of the San Juan Basin,Northwestern New Mexico and Their Implications for Understanding Latest Cretaceous Dinosaur Evolution

Studying the evolution and biogeographic distribution of dinosaurs during the latest Cretaceous is critical for better understanding the end-Cretaceous extinction event that killed off all non-avian dinosaurs. Western North America contains among the best records of Late Cretaceous terrestrial vertebrates in the world, but is biased against small-bodied dinosaurs. Isolated teeth are the primary evidence for understanding the diversity and evolution of small-bodied theropod dinosaurs during the Late Cretaceous, but few such specimens have been well documented from outside of the northern Rockies, making it difficult to assess Late Cretaceous dinosaur diversity and biogeographic patterns. We describe small theropod teeth from the San Juan Basin of northwestern New Mexico. These specimens were collected from strata spanning Santonian – Maastrichtian. We grouped isolated theropod teeth into several morphotypes, which we assigned to higher-level theropod clades based on possession of phylogenetic synapomorphies. We then used principal components analysis and discriminant function analyses to gauge whether the San Juan Basin teeth overlap with, or are quantitatively distinct from, similar tooth morphotypes from other geographic areas. The San Juan Basin contains a diverse record of small theropods. Late Campanian assemblages differ from approximately co-eval assemblages of the northern Rockies in being less diverse with only rare representatives of troodontids and a Dromaeosaurus-like taxon. We also provide evidence that erect and recurved morphs of a Richardoestesia-like taxon represent a single heterodont species. A late Maastrichtian assemblage is dominated by a distinct troodontid. The differences between northern and southern faunas based on isolated theropod teeth provide evidence for provinciality in the late Campanian and the late Maastrichtian of North America. However, there is no indication that major components of small-bodied theropod diversity were lost during the Maastrichtian in New Mexico. The same pattern seen in northern faunas, which may provide evidence for an abrupt dinosaur extinction.     

Dinosaurs and fluctuating sea levels during the mesozoic

The very different frequency of dinosaurs during the Mesozoic can be allied to the correlation between global sea level cyclicity and fossilization. This is based upon the sedimentary situation in the inner shelf, the area of predominant fossil record of dinosaurs, and sea level fluctuations. A rich fossil record is found in times of high sea level, and vice versa. Due to natural laws acting on sea level stands, the fossil record of dinosaurs and other terrestrial tetrapods is incomplete. This is causally explainable in the sequence stratigraphy. Among causes of global sea level fluctuations, the change from warm to cold times has been accorded greatest probability even in the Mesozoic. Consequently, the problem of dinosaur evolution and distribution should not be confused with the pattern of their fossil record. The latter, however, is so far nearly always used for all interpretations. The context presented here results in basic modifications.

During the phases of reduced to missing fossil record (low sea level, cold times), dinosaurs existed at least in circumequatorial regions in high diversity. Highly diverse faunas recorded exceptionally in the Upper Jurassic, Middle and Late Cretaceous, were each time the result of a long previous evolution and not the result of short term radiations at these times. Phases of sea level highstand and warm times caused an increased fossil record and poleward distribution. Cretaceous dinosaurs in paleolatitudes of 70° to 80° N and S are no proof for endothermy, but are only the effect of favorable climatic conditions at limited times. Any endothermy of the dinosaurs is not coincident with the supposedly uniformly warm equable climate of the Mesozoic, but with the opposite. Cold times did not hamper the existence of dinosaurs, but led in extreme cases (Aalenian and Valanginian) to the global lack of their fossil record. The situation at the Cretaceous‐Tertiary boundary is also explainable in this context. According to the sea level cyclicity, no extreme sea level fall and no globablly cold time were present in the critical time segment. The regression in the late Maastrichtian is found to belong to a sequence of third‐order cycles beginning in the Campanian. Every one of the cycle boundaries with regression and transgression produced apparent extinction effects which in reality are only gaps in the fossil record. After the late Maastrichtian regression the dinosaurs persisted with six lineages. The so far youngest dinosaur fauna in the Puercan (basal Paleocene) lies in a phase of sea level highstand of minor amplitude and duration with comparatively minor chances for a fossil record. The occurrences in the Puercan are governed by natural law, and, thus, dinosaurs are untied from the short term problems of the Cretaceous‐Tertiary boundary. Why dinosaurs are then missing at the next highstand, remains an open question. Anyhow, mechanisms which control fossil record, diversification and distribution, including global cold periods, do not belong to the direct causes of extinction, because identical occurrences happened many times during the Mesozoic without inducing extinction.     

First record of Megaraptora (Theropoda,Neovenatoridae) from Brazil

An isolated caudal vertebral centrum of a theropod dinosaur was discovered in the Bauru Basin (Late Cretaceous) of Brazil, in the Maastrichtian São José do Rio Preto Formation. The vertebral centrum has pneumatic features that are similar to those in the megaraptoran theropods Aerosteon, Megaraptor, and Orkoraptor. For example, all these taxa share with the caudal centrum here described the presence of true pleurocoels or pneumatic foramina, immersed within a depression or fossa. Thus, the specimen is considered the first record of Megaraptora in Brazil. The present analysis provides new information on the vertebral caudal anatomy of this clade of bizarre Cretaceous theropods.     

A new crocodylian from the late Maastrichtian of Spain: implications for the initial radiation of crocodyloids

Background

The earliest crocodylians are known primarily from the Late Cretaceous of North America and Europe. The representatives of Gavialoidea and Alligatoroidea are known in the Late Cretaceous of both continents, yet the biogeographic origins of Crocodyloidea are poorly understood. Up to now, only one representative of this clade has been known from the Late Cretaceous, the basal crocodyloid Prodiplocynodon from the Maastrichtian of North America.

Methodology/Principal Findings

The fossil studied is a skull collected from sandstones in the lower part of the Tremp Formation, in Chron C30n, dated at −67.6 to 65.5 Ma (late Maastrichtian), in Arén (Huesca, Spain). It is located in a continuous section that contains the K/P boundary, in which the dinosaur faunas closest to the K/P boundary in Europe have been described, including Arenysaurus ardevoli and Blasisaurus canudoi. Phylogenetic analysis places the new taxon, Arenysuchus gascabadiolorum, at the base of Crocodyloidea.

Conclusions/Significance

The new taxon is the oldest crocodyloid representative in Eurasia. Crocodyloidea had previously only been known from the Palaeogene onwards in this part of Laurasia. Phylogenetically, Arenysuchus gascabadiolorum is situated at the base of the first radiation of crocodyloids that occurred in the late Maastrichtian, shedding light on this part of the cladogram. The presence of basal crocodyloids at the end of the Cretaceous both in North America and Europe provides new evidence of the faunal exchange via the Thulean Land Bridge during the Maastrichtian.     

A New European Marsupial Indicates a Late Cretaceous High-Latitude Transatlantic Dispersal Route

The first record of an undoubted opossum-like marsupial from the Mesozoic of Europe indicates an invasion from North America at the end of Late Cretaceous (Maastrichtian). The new 66.1 million-year-old marsupial, Maastrichtidelphys meurismeti n. gen., n. sp., represented by a right upper molar, comes from the type Maastrichtian of The Netherlands. The Maastricht marsupial exhibits affinities with earlier (early Maastrichtian) North American herpetotheriids providing definitive evidence of a high-latitude North Atlantic dispersal route between North America and Europe during the latest Cretaceous. Previously, the first major interchange for marsupials was thought to have occurred nearly 10 million years later in the Eocene. The occurrence of this new marsupial in Europe implies that at some time during the latest Cretaceous, sea level and climatic conditions must have been sufficiently favorable to allow for such a high-latitude dispersal. The fragmentary remains of hadrosaurid and theropod dinosaurs, as well as boid snakes from northwestern Europe which have affinities with North American taxa help substantiate assumptions made by the occurrence of the herpetotheriid marsupial in Maastricht.     

A preliminary phylogeny of the Pterasteridae (Echinodermata,Asteroidea) and the first fossil record: Late Cretaceous of Germany and Belgium

The Pterasteridae comprises a diversified group of extant largely deep-sea starfishes. Generic diagnoses have been based classically on soft tissue characters and skeletal architecture. A preliminary phylogeny of sixteen extant species is here worked out by cladistic analysis. The resulting tree suggests monophyly of extant genera and the validity of dissociated plates for identification of genera. Fossil remains of Pterasteridae are here described for the first time. By comparison with extant species, all the skeletal remains from the lower Upper Campanian of Belgium and from the lower Maastrichtian of Germany are tentatively assigned to the genusPteraster. The fossil record of starfishes is poor, but the present Late Cretaceous pterasterids provide one more piece of evidence of the high diversity of starfishes during the Mesozoic. Known Late Cretaceous and Paleogene fossils are broadly similar, which suggests the end-Cretaceous extinction event did not cause major turnover in asteroid faunal composition. As suggested for other starfish groups, both the fossil record of deep-sea Pterasteridae in shelf settings and tree topology imply an onshore-offshore evolutionary trend.      

Diverse Dinosaur-Dominated Ichnofaunas from the Potomac Group (Lower Cretaceous) Maryland

Until recently fossil footprints were virtually unknown from the Cretaceous of the eastern United States. The discovery of about 300 footprints in iron-rich siliciclastic facies of the Patuxent Formation (Potomac Group) of Aptian age is undoubtedly one of the most significant Early Cretaceous track discoveries since the Paluxy track discoveries in Texas in the 1930s. The Patuxent tracks include theropod, sauropod, ankylosaur and ornithopod dinosaur footprints, pterosaur tracks, and miscellaneous mammal and other vertebrate ichnites that collectively suggest a diversity of about 14 morphotypes. This is about twice the previous maximum estimate for any known Early Cretaceous vertebrate ichnofauna. Among the more distinctive forms are excellent examples of hypsilophodontid tracks and a surprisingly large mammal footprint. A remarkable feature of the Patuxent track assemblage is the high proportion of small tracks indicative of hatchlings, independently verified by the discovery of a hatchling-sized dinosaur. Such evidence suggests the proximity of nest sites. The preservation of such small tracks is very rare in the Cretaceous track record, and indeed throughout most of the Mesozoic.

This unusual preservation not only provides us with a window into a diverse Early Cretaceous ecosystem, but it also suggests the potential of such facies to provide ichnological bonanzas. A remarkable feature of the assemblage is that it consists largely of reworked nodules and clasts that may have previously been reworked within the Patuxent Formation. Such unusual contexts of preservation should provide intriguing research opportunities for sedimentologists interested in the diagenesis and taphonomy of a unique track-bearing facies.     

The fossil record of Limopsis (Bivalvia: Limopsidae) in Antarctica and the southern high latitudes

Abstract: Limopsis is one of the most speciose and widespread bivalve genera in the Southern Ocean at the present day. However, the fossil record of the genus is poorly known from the southern high latitudes. Here, we review the fossil record in this region to help understand the evolutionary origins of the genus. Limopsis infericola sp. nov. and additional specimens of a previously described species are added to the fossil record of Antarctica. The globally distributed limopsid clade had its earliest occurrences in the Early Cretaceous of Europe and New Zealand, then radiated during the Late Cretaceous (Maastrichtian, 70.6–65.5 Ma). Fossil evidence shows that the genus underwent a second, Cenozoic, radiation related to the isolation of Antarctica and the onset of cooling in the southern hemisphere. The genus has persisted in Antarctica for the last 50 myr, adapting to extreme changes in the environmental conditions, including surviving the last glacial maximum in marine refugia.     

First Record of Theropod and Ornithopod Tracks and Detailed Description of Sauropod Trackways from the Tacuarembó Formation (Late Jurassic-?Early Cretaceous) of Uruguay

The record of dinosaur tracks and trackways from the Tacuarembó Formation (Late Jurassic-?Early Cretaceous) is described in detail: two trackways and four isolated tracks. Ichnites are preserved in sandstones of interdune facies of the Batoví (lower) Member. The interpretation herein presented indicates a sauropod affinity of the trackways, while the isolated prints are considered to have been made by an ornithopod and small theropods. The fossil record for this formation was primarily integrated by fresh-water organisms. The presence of new groups (sauropods and ornithopods) has specially contributed to the reconstruction of the terrestrial fauna, represented up until this time only by theropods.     

Fused mid‐caudal vertebrae in the titanosaur Uberabatitan ribeiroi from the Late Cretaceous of Brazil and other bone lesions

Bone pathology in the titanosaur dinosaur Uberabatitan ribeiroi from the Marília Formation (Bauru Group, Late Cretaceous) of Uberaba city (Minas Gerais State, Brazil) is analysed here. They include two fused procoelous mid‐caudal vertebrae (CPPLIP‐1020) and a haemal arch (CPPLIP‐1006) of the middle section of the tail with a healing fracture callus. The analyses of the caudal vertebrae CPPLIP‐1020 of Uberabatitan permit us to recognize the following signs, based on CT scan and external macroscopic observations: (1) ossified longitudinal tendons; (2) likely ossified intervertebral disc, producing fused vertebral bodies; (3) fused right zygapophyseal process with a laterally developed osteophyte affecting this joint; (4) osteophytes and exostoses at different portions of the vertebrae; (5) cloacae, bone erosion and likely internal infection. According to all the processes observed in these caudals, we could not discard at least two possibilities for the diagnosis of the fused vertebrae. It could be the result of a spondyloarthropathy process (considering most of the observed signs) or possibly have been associated with an infection (e.g. discospondylitis/infections spondylitis or septic arthritis). The bone lesion record in Uberabatitan ribeiroi from the Late Cretaceous of Brazil increases the range of study of titanosaur dinosaurs, which although have a large fossil record, have few pathological studies.     

Mountain building triggered late cretaceous north american megaherbivore dinosaur radiation

Prior studies of Mesozoic biodiversity document a diversity peak for dinosaur species in the Campanian stage of the Late Cretaceous, yet have failed to provide explicit causal mechanisms. We provide evidence that a marked increase in North American dinosaur biodiversity can be attributed to dynamic orogenic episodes within the Western Interior Basin (WIB). Detailed fossil occurrences document an association between the shift from Sevier-style, latitudinally arrayed basins to smaller Laramide-style, longitudinally arrayed basins and a well substantiated decreased geographic range/increased taxonomic diversity of megaherbivorous dinosaur species. Dispersal-vicariance analysis demonstrates that the nearly identical biogeographic histories of the megaherbivorous dinosaur clades Ceratopsidae and Hadrosauridae are attributable to rapid diversification events within restricted basins and that isolation events are contemporaneous with known tectonic activity in the region. SymmeTREE analysis indicates that megaherbivorous dinosaur clades exhibited significant variation in diversification rates throughout the Late Cretaceous. Phylogenetic divergence estimates of fossil clades offer a new lower boundary on Laramide surficial deformation that precedes estimates based on sedimentological data alone.     

The occurrence of the fossil genus Graminidites in the Maastrichtian Scollard Formation,Alberta, Canada,and its palaeoecological and palaeogeographical significance

Fossil grass pollen is common in Late Tertiary sediments and its rare occurrence in the Early Tertiary led to the belief that its presence is restricted to Tertiary sediments. A literature survey shows that rare occurrences of Graminidites spp. have been reported from Campanian–Maastrichtian strata worldwide. All reported Graminidites spp. have been studied by transmitted light microscopes, which are inadequate to resolve their fine exine sculpture. Graminidites sp. occurs in the Maastrichtian Scollard Formation of Alberta, Canada, and is studied here with light microscopy and scanning electron microscopy. A new species, G. ulkapites, occurs at the initiation of a temperate climate in the area. It occurs in post‐dinosaurian beds locally but in pre‐Cretaceous–Tertiary boundary strata, which implies that grasses could also be present in dinosaur inhabitations. Grass phytoliths, reported from dinosaurian coprolites from the Maastrichtian Deccan Intertrappean sediments of India, indicate that grasses were ingested by dinosaurs even if not included in the dinosaurian diet. A plot of worldwide Senonian occurrences of Graminidites spp. on a late Maastrichtian palaeogeographic map demonstrates that the data are inadequate to reveal the place of origin or migratory pattern. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 235–248.     

The First Dinosaur from Washington State and a Review of Pacific Coast Dinosaurs from North America

We describe the first diagnostic dinosaur fossil from Washington State. The specimen, which consists of a proximal left femur, was recovered from the shallow marine rocks of the Upper Cretaceous (Campanian) Cedar District Formation (Nanaimo Group) and is interpreted as pertaining to a large theropod on the basis of its hollow medullary cavity and proximally placed fourth trochanter. The Washington theropod represents one of the northernmost occurrences of a Mesozoic dinosaur on the west coast of the United States and one of only a handful from the Pacific coast of Laramidia during the Cretaceous. Its isolated nature and preservation in marine rocks suggest that the element was washed in from a nearby fluvial system. If the femur pertains to a tyrannosauroid, which seems likely given its size and the widespread occurrence of the group across Laramidia during Late Cretaceous times, then it would represent an earlier occurrence of large body size than previously recognized (complete femur length estimated at 1.2 meters). Uncertainty surrounding the latitude of deposition of the Nanaimo Group (i.e., the Baja-British Columbia hypothesis) precludes assigning the Washington theropod to either of the putative northern or southern biogeographic provinces of Laramidia.     

Dinosaurs of France

The French dinosaur record is one of the most extensive in Europe; it ranges stratigraphically from the Late Triassic to the Latest Cretaceous. All major clades of dinosaurs but marginocephalians are known. About 20 species are based on significant material; the theropods are the best represented. Most of these taxa have been described or revised in recent years. Important specimens have been discovered in the Late Triassic of eastern France, the Middle Jurassic of Normandy, and the Late Cretaceous of Provence and Languedoc. The ichnological record is good for the Late Triassic-Early Jurassic, and the Late Cretaceous egg sites are among the richest in the world. To cite this article: R. Allain, X.P. Suberbiola, Palevol 2 (2003) 27–44.