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.     

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 new basal ornithopod dinosaur (Frenchman Formation,Saskatchewan, Canada), and implications for late Maastrichtian ornithischian diversity in North America

A small, articulated basal ornithopod skeleton from the Frenchman Formation (late Maastrichtian) of Saskatchewan (RSM P 1225.1), previously referred to the taxon Thescelosaurus, differs from both recognized species of this taxon (Thescelosaurus neglectus and Thescelosaurus garbanii). The differences are taxonomically informative and we recognize this specimen as the holotype of a new species, Thescelosaurus assiniboiensis sp. nov. , diagnosed by the presence of two autapomorphies, and displaying plesiomorphic traits more similar to those of Parksosaurus, than to those of the other Thescelosaurus species. The Frenchman Formation also harbours an intriguing faunal assemblage in which Thescelosaurus represents one of the most abundant dinosaur taxa, and preserves a relatively high proportion of small (putatively juvenile and subadult) specimens of many dinosaur taxa. Further work that increases the faunal sample from this formation, and that permits quantitative comparisons with contemporary formations, will determine whether or not these differences are well supported, and will determine their ultimate palaeobiological significance. Identification of a third species of Thescelosaurus from the late Maastrichtian of North America suggests that this taxon was more diverse than previously recognized, and shows an increase in diversity from the Campanian through the late Maastrichtian, contrasting the trends seen in most other ornithischian clades. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1157–1198.     

The youngest sauropod evidence in Europe

Titanosaur sauropods constituted one of the major dinosaur faunal components in Europe along the latest Cretaceous. Although they were abundant in the late Campanian–early Maastrichtian, titanosaurs apparently demised in terms of diversity and abundance in the late Maastrichtian Ibero-Armorican domain. Thus, any finding in this crucial period provides new clues to understand the biodiversity trend of the group prior to the K–Pg boundary. Here, we report the occurrence of a vertebral element attributed to a titanosaur in fluvial deposits dated as uppermost Maastrichtian (C29r chrone). The specimen corresponds to the right posterior part of a posterior cervical vertebra. CT-scan and histological analyses revealed high pneumatisation with laminar structures composed of compact bone, strongly remodelled with at least three generations of secondary osteons. These evidences suggest a relatively advanced age for the individual at death. Given that the partial vertebra was recovered from the uppermost portion of the late Maastrichtian, it might represent the youngest evidence for titanosaur sauropods in Europe discovered so far. Further, alongside to other remains (i.e. eggshells and footprints), the studied partial vertebra provides evidence for the persistence of this taxonomic group at the very end of the Cretaceous in southwestern Europe.     

The extinction of the dinosaurs

Non‐avian dinosaurs went extinct 66 million years ago, geologically coincident with the impact of a large bolide (comet or asteroid) during an interval of massive volcanic eruptions and changes in temperature and sea level. There has long been fervent debate about how these events affected dinosaurs. We review a wealth of new data accumulated over the past two decades, provide updated and novel analyses of long‐term dinosaur diversity trends during the latest Cretaceous, and discuss an emerging consensus on the extinction's tempo and causes. Little support exists for a global, long‐term decline across non‐avian dinosaur diversity prior to their extinction at the end of the Cretaceous. However, restructuring of latest Cretaceous dinosaur faunas in North America led to reduced diversity of large‐bodied herbivores, perhaps making communities more susceptible to cascading extinctions. The abruptness of the dinosaur extinction suggests a key role for the bolide impact, although the coarseness of the fossil record makes testing the effects of Deccan volcanism difficult.     

On the earliest record of Cretaceous tyrannosauroids in western North America: implications for an Early Cretaceous Laurasian interchange event

The sudden appearance of Asian dinosaur clades within Lower Cretaceous strata of western North America has long been recognised as a biotic dispersion event related to initial establishment of a Beringian land bridge. To date, uncertainty exists regarding the timing of the Early Cretaceous Laurasian interchange event (EKLInE) and the pattern of associated biotic dispersal. Here, we report a tyrannosauroid premaxillary tooth (FMNH PR 2750) from the Cloverly Formation, Wyoming, USA, that pushes back the earliest Cretaceous record of the clade in North America. Although fragmentary, the tooth is consistent with mounting evidence for a pre-108 Ma initiation of EKLInE and earliest Albian emplacement of Beringia. Previous authors have considered the Aptian/Albian of western North America a depauperate dinosaur fauna, characterised by regional extinction and diversity decline. Documentation of Albian tyrannosauroids in the region indicates a more dynamic ecosystem than previously appreciated and marks an early start to faunal mixing between immigrant and endemic dinosaur clades. Finally, we find that the enamel microstructure of FMNH PR 2750 conforms to the morphotype of tyrannosaurids, yet exhibits poor columnar differentiation. This morphology bolsters prior interpretations on the phylogenetic utility of enamel microstructure and suggests a trend of increasing enamel complexity within Tyrannosauroidea.     

Formation binning: a new method for increased temporal resolution in regional studies,applied to the Late Cretaceous dinosaur fossil record of North America

The advent of palaeontological occurrence databases has allowed for detailed reconstruction and analyses of species richness through deep time. While a substantial literature has evolved ensuring that taxa are fairly counted within and between different time periods, how time itself is divided has received less attention. Stage-level or equal-interval age bins have frequently been used for regional and global studies in vertebrate palaeontology. However, when assessing diversity at a regional scale, these resolutions can prove inappropriate with the available data. Herein, we propose a new method of binning geological time for regional studies that intrinsically incorporates the chronostratigraphic heterogeneity of different rock formations to generate unique stratigraphic bins. We use this method to investigate the diversity dynamics of dinosaurs from the Late Cretaceous of the Western Interior of North America prior to the Cretaceous–Palaeogene mass extinction. Increased resolution through formation binning pinpoints the Maastrichtian diversity decline to between 68 and 66 Ma, coinciding with the retreat of the Western Interior Seaway. Diversity curves are shown to exhibit volatile patterns using different binning methods, supporting claims that heterogeneous biases in this time-frame affect the pre-extinction palaeobiological record. We also show that the apparent high endemicity of dinosaurs in the Campanian is a result of non-contemporaneous geological units within large time bins. This study helps to illustrate the utility of high-resolution, regional studies to supplement our understanding of factors governing global diversity in deep time and ultimately how geology is inherently tied to our understanding of past changes in species richness.     

Latirhinus uitstlani,a ‘broad-nosed’ saurolophine hadrosaurid (Dinosauria,Ornithopoda) from the late Campanian (Cretaceous) of northern Mexico

We describe a new genus and species of hadrosaurid dinosaur, Latirhinus uitstlani, from upper Campanian strata of the Cerro del Pueblo Formation in northern Mexico. The new taxon differs from other hadrosaurids, except Gryposaurus, in possessing a deep and arcuate dorsal process of the nasal; it differs from Gryposaurus, as well as from all other saurolophines, in having a broader narial foramen, and an anterodorsally curved deltoid crest of the scapula (convergent with Lambeosaurinae within the context of Hadrosauridae). L. uitstlani appears to be allied to ‘kritosaurs’, particularly to species of Gryposaurus, and it represents one of the southernmost saurolophines of North America. Likewise, L. uitstlani adds to the diversity of saurolophines in the southern Western Interior Basin and its occurrence in northern Mexico is consistent with the apparent predominance of ‘kritosaurs’ in southern Laramidia during late Campanian times.     

A re-evaluation of the ‘mid-Cretaceous sauropod hiatus’ and the impact of uneven sampling of the fossil record on patterns of regional dinosaur extinction

The mid-Cretaceous of North America and Europe has long been noted for the absence of sauropod dinosaurs, leading several authors to suggest that this depauperate interval is a consequence of an end-Albian sauropod extinction. This time period has become known as the ‘mid-Cretaceous sauropod hiatus’, with the subsequent presence of titanosaurian sauropods in the latest Cretaceous of North America and Europe interpreted as the result of dispersal of taxa from South America and Africa, respectively. However, several lines of evidence indicate that this hiatus is probably a sampling artefact. New fossil and trackway discoveries have considerably shortened the hiatus, reducing it to the Turonian–early Campanian in North America, and to just two short intervals in the late Cenomanian–early Turonian and late Coniacian–Santonian of Europe. Palaeoenvironmental analyses of sauropods demonstrate an inland terrestrial preference for titanosaurs, the dominant Late Cretaceous sauropods; however, during the hiatus there was a decline in inland deposits and increase in coastal sediments in Europe and North America, which would have greatly reduced the probability of preserving titanosaurs. Neither the decline in inland deposits, nor the ‘sauropod hiatus’, occurred elsewhere in the world. Statistical comparisons also demonstrate a significant positive correlation between fluctuations in inland deposits and sauropod occurrences during the mid–Late Cretaceous in Europe and North and South America. Lastly, cladistic analyses do not place latest Cretaceous North American and European titanosaurs within South American and African clades, contradicting the predictions of the ‘austral immigrant’ hypothesis. The latter hypothesis also receives little support from biogeographical analysis of dispersal among titanosaurs. Thus, the ‘sauropod hiatus’ of North America and Europe is most plausibly interpreted as the product of a sampling bias pertaining to the rarity of inland sediments and dominance of coastal deposits preserved in these two regions during the mid-Cretaceous. The presence of titanosaurs in these areas during the latest Cretaceous can be explained by dispersal from Southern Hemisphere continents, but this is no more probable than descent from Early Cretaceous incumbent faunas or dispersal from Asia.     

A new hadrosaurid dentary from the latest Maastrichtian of the Pyrenees (north Spain) and the high diversity of the duck-billed dinosaurs of the Ibero-Armorican Realm at the very end of the Cretaceous

In the latest Maastrichtian, the European hadrosauroid fauna was more diverse than those of North America and Asia. The European record of hadrosaurid dentaries is an example of this diversity, and most of the sites with mandibular remains are located in the Ibero-Armorican Realm. Within the Iberian Peninsula, most of the remains are located in the Tremp Basin (South Central Pyrenees). Two of the three valid hadrosaurid taxa defined in this basin are from the Blasi sites (Arén, Huesca province): Arenysaurus ardevoli (Blasi-3) and Blasisaurus canudoi (Blasi-1). A new locality in Blasi (Blasi 3.4) has provided a new dentary from an indeterminate euhadrosaurid. This dentary presents some characters intermediate between Arenysaurus and Blasisaurus, some characters similar to Pararhabdodon isonensis (from the nearby province of Lleida), and some characters of its own. Nevertheless, due to its fragmentary character, without dentition or its edentulous anterior part, it cannot be determined above the level of Euhadrosauria. It thus represents a fourth Iberian euhadrosaurian taxon in the Ibero-Armorican Realm, different from Arenysaurus, Blasisaurus and Pararhabdodon, increasing the diversity of hadrosauroids in this realm at the very end of the Cretaceous.     

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.     

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.     

Severe extinction and rapid recovery of mammals across the Cretaceous–Palaeogene boundary,and the effects of rarity on patterns of extinction and recovery

The end‐Cretaceous mass extinction ranks among the most severe extinctions of all time; however, patterns of extinction and recovery remain incompletely understood. In particular, it is unclear how severe the extinction was, how rapid the recovery was and how sampling biases might affect our understanding of these processes. To better understand terrestrial extinction and recovery and how sampling influences these patterns, we collected data on the occurrence and abundance of fossil mammals to examine mammalian diversity across the K‐Pg boundary in North America. Our data show that the extinction was more severe and the recovery more rapid than previously thought. Extinction rates are markedly higher than previously estimated: of 59 species, four survived (93% species extinction, 86% of genera). Survival is correlated with geographic range size and abundance, with widespread, common species tending to survive. This creates a sampling artefact in which rare species are both more vulnerable to extinction and less likely to be recovered, such that the fossil record is inherently biased towards the survivors. The recovery was remarkably rapid. Within 300 000 years, local diversity recovered and regional diversity rose to twice Cretaceous levels, driven by increased endemicity; morphological disparity increased above levels observed in the Cretaceous. The speed of the recovery tends to be obscured by sampling effects; faunas show increased endemicity, such that a rapid, regional increase in diversity and disparity is not seen in geographically restricted studies. Sampling biases that operate against rare taxa appear to obscure the severity of extinction and the pace of recovery across the K‐Pg boundary, and similar biases may operate during other extinction events.     

A NEW SPECIES OF ISCHYODUS (CHONDRICHTHYES: HOLOCEPHALI: CALLORHYNCHIDAE) FROM UPPER MAASTRICHTIAN SHALLOW MARINE FACIES OF THE FOX HILLS AND HELL CREEK FORMATIONS, WILLISTON BASIN, NORTH DAKOTA, USA

Abstract:  A new species of chimaeroid, Ischyodus rayhaasi sp. nov., is described based primarily upon the number and configuration of tritors on palatine and mandibular tooth plates. This new species is named in honour of Mr Raymond Haas. Fossils of I. rayhaasi have been recovered from the Upper Maastrichtian Fox Hills Formation and the Breien Member and an unnamed member of the Hell Creek Formation at sites in south-central North Dakota and north-central South Dakota, USA. Ischyodus rayhaasi inhabited shallow marine waters in the central part of the Western Interior Seaway during the latest Cretaceous. Apparently it was also present in similar habitats at that time in the Volga region of Russia. Ischyodus rayhaasi is the youngest Cretaceous species of Ischyodus known to exist before the Cretaceous/Tertiary extinction, and the species apparently did not survive that event. It was replaced by Ischyodus dolloi , which is found in the Paleocene Cannonball Formation of the Williston Basin region of North Dakota and is widely distributed elsewhere.     

Upper Campanian–Maastrichtian chronostratigraphy of the Skælskør‐1 core,Denmark: correlation at the basinal and global scale and implications for changes in sea‐surface temperatures

The lithostratigraphy, calcareous nannofossil biostratigraphy, carbon‐ and oxygen‐isotope stratigraphy and gamma‐ray profile are presented for the Skælskør‐1 core, eastern Denmark. The correlation of carbon isotopes to Gubbio (Italy) and ODP Site 762C (Indian Ocean) provides the chronostratigrahical framework of the core through a tie to magnetostratigraphy. Two new carbon‐isotope excursions are defined for the uppermost Maastrichtian of the core and prove useful for long‐distance correlation. Twenty stratigraphic tie‐points are used for correlation of the upper Campanian–Maastrichtian interval by combining carbon‐isotope and gamma‐ray variations. Significant dissimilarities in the gamma‐ray profiles of the Danish Basin cores preclude the sole use of this tool for basin‐scale correlations. Bulk oxygen‐isotopes and semi‐quantitative abundance changes in the warm‐water calcareous nannofossil Watznaueria barnesiae and the cool‐water Kamptnerius magnificus highlight the following past changes in sea‐surface temperatures (SSTs): relatively warm late Campanian SSTs, cooling across the Campanian–Maastrichtian boundary and through the early Maastrichtian, warming across the early–late Maastrichtian transition, cooling in the late Maastrichtian, intense warming in the latest Maastrichtian chron C29r, followed by a very short episode of cooling immediately before the Cretaceous–Palaeogene boundary. The late Campanian–Maastrichtian evolution in sea water temperatures inferred from the Danish Basin is similar to that delineated at tropical latitude oceanic sites.     

Species extinction across the cretaceous‐tertiary boundary: Observed patterns versus predicted sampling effects,stepwise or otherwise?

Stepwise extinction and origination patterns have been reported around the Cenomanian‐Turonian, Cretaceous‐Tertiary, and the Eocene‐Oligocene boundaries. Some of these observed stepwise patterns may result from only sampling effects. It is possible to test whether or not the observed stepwise patterns result from sampling effects if the appropriate information is known.

An example of such a test involves the molluscan species from the Maastrichtian Prairie Bluff Chalk at Bragg, Alabama. Species abundances are known and sample sizes are 2144, 558, and 680 specimens from the lower, middle, and upper portions of the Prairie Bluff at Braggs. The observed ranges are not significantly different from the predicted values.

These molluscs are representative of the Campanian‐Maastrichtian temperate fauna distributed from northern Mexico, through the U.S. coastal plain, to eastern Europe. This test suggests that this species‐rich and widely distributed fauna persisted unchanged until the latest Cretaceous.     

“South American” Marsupials from the Late Cretaceous of North America and the Origin of Marsupial Cohorts

Newly described marsupial specimens of Judithian (late Campanian) and Lancian (Maastrichtian) age in the western interior of North America (Wyoming to Alberta) have dental morphologies consistent with those expected in comparably aged sediments in South America (yet to be found). Three new Lancian species are referable to the didelphimorphian Herpetotheriidae, which suggests that the ameridelphian radiation was well under way by this time. The presence of a polydolopimorphian from Lancian deposits with a relatively plesiomorphic dental morphology and an additional polydolopimorphian taxon from Judithian deposits with a more derived molar form indicate that this lineage of typically South American marsupials was diversifying in the Late Cretaceous of North America. This study indicates that typical South American lineages (e.g. didelphimorphians and polydolopimorphians) are not the result of North American peradectian progenitors dispersing into South America at the end of the Cretaceous (Lancian), or at the beginning of the Paleocene (Puercan), and giving rise to the ameridelphian marsupials. Instead, these lineages, and predictably others as well, had their origins in North America (probably in more southerly latitudes) and then dispersed into South America by the end of the Cretaceous. Geophysical evidence concerning the connections between North and South America in the Late Cretaceous is summarized as to the potential for overland mammalian dispersal between these places at those times. Paleoclimatic reconstructions are considered, as is the dispersal history of hadrosaurine dinosaurs and boid snakes, as to their contribution to an appraisal of mammalian dispersals in the Late Cretaceous. In addition, we present a revision of the South American component of the Marsupialia. One major outcome of this process is that the Polydolopimorphia is placed as Supercohort Marsupialia incertae sedis because no characteristics currently known from this clade securely place it within one of the three named marsupial cohorts.     

Palaeoceanographic and palaeoclimatic inferences from Late Cretaceous planktonic foraminiferal assemblages from the Exmouth Plateau (ODP Sites 762 and 763, eastern Indian Ocean)

The evolution of planktonic foraminifera during the Late Cretaceous is marked in the Santonian by the disappearance of complex morphotypes (the marginotruncanids), and the contemporary increasing importance and diversification of another group of complex taxa, the globotruncanids. Upper Turonian to lower Campanian planktonic foraminiferal assemblages from Holes 762C and 763B (Ocean Drilling Program, Leg 122, Exmouth Plateau, 47°S palaeolatitude) were studied in detail to evaluate the compositional variations at the genus and species level based on the assumption that, in the Cretaceous oceans as in the modern, any faunal change was associated with changes in the characteristics and the degree of stability of the oceanic surface waters. Three major groups were recognised based on gross morphology, and following the assumption that Cretaceous planktonic foraminifera, although extinct, had life-history strategies comparable to those of modern planktonics: 1 – r-selected opportunists; 2 – k-selected specialists; 3 – r/k intermediate morphotypes which include all genera that display a range of trophic strategies in-between opportunist and specialist taxa. Although planktonic foraminiferal assemblages are characterised by a progressive appearance of complex taxa, this trend is discontinuous. Variation in number of species and specimens within genera has allowed recognition of five discrete intervals each of them reflecting different oceanic conditions based on fluctuations in diversity and abundance of the major morphotypes. Planktonic forms show cyclical fluctuations in diversity and abundance of cold (r-strategists) and warm taxa (k-strategists), perhaps representing alternating phases of unstable conditions (suggesting a weakly stratified upper water column in a mesotrophic environment), and well-stratified surface and near-surface waters (indicating a more oligotrophic environment). Interval 1, middle Turonian to early Coniacian in age, is dominated by the r/k intermediate morphotypes which alternate with r-strategists. These cyclical alternations are used to identify three additional sub-intervals. Interval 2, aged middle to late Coniacian, is characterised by the increasing number of species and relative abundance of k-strategists. After this maximum diversification the k-strategists show a progressive decrease reaching a minimum value in Interval 3 (early to late Santonian), which corresponds to the extinction of the genus Marginotruncana. In the Interval 4, latest Santonian in age, the k-strategists, represented mainly by the genera Globotruncana, increase again in diversity and abundance. The last Interval 5 (early Campanian) is dominated by juvenile globotruncanids and r-strategists which fluctuate in opposite phase. The positive peak (Interval 2) related to the maximum diversification of warm taxa (k-strategists) in the Coniacian seems to correspond to a warmer episode. It is followed by a marked decrease in the relative abundance of warm taxa (k-strategists crisis) with a minimum in the late Santonian (Interval 3), reflecting a decrease in temperature. Detailed analysis of faunal variations allows the Santonian faunal turnover to be ascribed to a cooling event strong enough to cause the extinction of the marginotruncanids.     

Global historical biogeography of hadrosaurid dinosaurs

Hadrosaurids were the most derived ornithopods and amongst the most diverse herbivore dinosaurs during the Late Cretaceous of Europe, Asia, and the two Americas. Here, their biogeographical history is reconstructed using dispersal‐vicariance analysis (DIVA). The results showed that Hadrosauridae originated in North America and soon after dispersed to Asia no later than the Late Santonian. The most recent common ancestor of Saurolophidae (= Saurolophinae + Lambeosaurinae) is inferred to have been widespread in North America and Asia. The split between saurolophines and lambeosaurines occurred in response to vicariance no later than the Late Santonian: the former clade originated in North America, whereas the latter did so in Asia. Saurolophine biogeographical history included a minimum of five dispersal events followed by vicariance. Four of these dispersals were inferred to have occurred from North America to Asia during the Campanian and Early Maastrichtian, whereas a fifth event represented a southward dispersal from North to South America no later than the Late Campanian. The historical biogeography of lambeosaurines was characterized by an early evolution in Asia, with a Campanian dispersal to the European archipelago followed by vicariance. Reconstruction of the ancestral areas for the deepest nodes uniting the more derived lambeosaurines clades (‘hypacrosaurs’, ‘corythosaurs’, and ‘parasaurolophs’) is ambiguous. The split between North American and Asian clades of ‘hypacrosaurs’ and ‘parasaurolophs’ occurred in response to vicariance during the Campanian. The evolutionary history of North American ‘hypacrosaurs’ and ‘parasaurolophs’ was characterized by duplication events. The latter also characterized the Late Campanian ‘corythosaurs’, which remained restricted to North America. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 159 , 503–525.