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.     

A new abelisaurid dinosaur from the Late Cretaceous of southern France: Palaeobiogeographical implications

The Abelisauridae are a family of mainly Cretaceous theropod dinosaurs with a wide distribution across the Gondwanan land masses. Although their presence in Europe was reported twenty-five years ago, it has often been considered as controversial largely because of the incompleteness of the available specimens. We report here the discovery of well-preserved abelisaurid material, including a highly diagnostic braincase, at a Late Cretaceous (late Campanian) locality in the Aix-en-Provence Basin, near the eponym city in south-eastern France. A new abelisaurid taxon is erected, Arcovenator escotae gen. nov., sp. nov., on the basis of cranial and postcranial material. A phylogenetic analysis reveals that the new Abelisauridae from Provence is more closely related to taxa from India and Madagascar than to South American forms. Moreover, Genusaurus, Tarascosaurus and the previous Late Cretaceous discoveries are identified as basal abelisaurids. Contrary to previously proposed palaeobiogeographical models of abelisaurid evolution, the presence of the new taxon in Europe suggests that Europe and Africa may have played a major role in abelisaurid dispersal, which apparently involved crossing marine barriers.     

A new troodontid theropod, Talos sampsoni gen. et sp. nov., from the Upper Cretaceous Western Interior Basin of North America

Background

Troodontids are a predominantly small-bodied group of feathered theropod dinosaurs notable for their close evolutionary relationship with Avialae. Despite a diverse Asian representation with remarkable growth in recent years, the North American record of the clade remains poor, with only one controversial species—Troodon formosus—presently known from substantial skeletal remains.

Methodology/Principal Findings

Here we report a gracile new troodontid theropod—Talos sampsoni gen. et sp. nov.—from the Upper Cretaceous Kaiparowits Formation, Utah, USA, representing one of the most complete troodontid skeletons described from North America to date. Histological assessment of the holotype specimen indicates that the adult body size of Talos was notably smaller than that of the contemporary genus Troodon. Phylogenetic analysis recovers Talos as a member of a derived, latest Cretaceous subclade, minimally containing Troodon, Saurornithoides, and Zanabazar. MicroCT scans reveal extreme pathological remodeling on pedal phalanx II-1 of the holotype specimen likely resulting from physical trauma and subsequent infectious processes.

Conclusion/Significance

Talos sampsoni adds to the singularity of the Kaiparowits Formation dinosaur fauna, which is represented by at least 10 previously unrecognized species including the recently named ceratopsids Utahceratops and Kosmoceratops, the hadrosaurine Gryposaurus monumentensis, the tyrannosaurid Teratophoneus, and the oviraptorosaurian Hagryphus. The presence of a distinct troodontid taxon in the Kaiparowits Formation supports the hypothesis that late Campanian dinosaurs of the Western Interior Basin exhibited restricted geographic ranges and suggests that the taxonomic diversity of Late Cretaceous troodontids from North America is currently underestimated. An apparent traumatic injury to the foot of Talos with evidence of subsequent healing sheds new light on the paleobiology of deinonychosaurians by bolstering functional interpretations of prey grappling and/or intraspecific combat for the second pedal digit, and supporting trackway evidence indicating a minimal role in weight bearing.     

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.     

山东莱阳晚白垩世恐龙与恐龙蛋研究历史和新进展

简要回顾了莱阳恐龙和恐龙蛋化石群的研究历史和以杨钟健为代表的老一代地质古生物学者对莱阳恐龙和恐龙蛋研究发现的杰出贡献,并介绍了莱阳恐龙和恐龙蛋的最新发现和若干研究进展.在近年来对莱阳周边地层进行的大规模考察中,发现了一系列发育在平原上的恐龙峡谷群,以及其中蕴含的十几个恐龙和恐龙蛋新地点和新层位.2010年开始,在对莱阳金岗口村附近的2个化石地点的发掘中,发现了以鸭嘴龙科为主的大量脊椎动物化石和蛋化石,包括一类新的栉龙亚科成员,一些大型兽脚类恐龙化石,以及一新的龟鳖类蛋化石等.2号地点化石富集层具有典型的泥石流沉积特征和骨骼埋藏特征.对棘鼻青岛龙的特殊头饰进行了CT扫描和三维重建,发现其头饰是实心结构,但其他骨骼特征证明棘鼻青岛龙属于具有头饰的赖氏龙亚科是确定无疑的,所以目前发现的头饰应不是其真实状态或根本不属于其头部骨骼.此外,对谭氏龙1属3种的重新观察研究得出以下结论:中国谭氏龙和金刚口谭氏龙应是有效属种,但部分骨骼还有疑问,还需要进一步研究,而莱阳谭氏龙为无效属种.     

Theropod fauna from southern Australia indicates high polar diversity and climate-driven dinosaur provinciality

The Early Cretaceous fauna of Victoria, Australia, provides unique data on the composition of high latitude southern hemisphere dinosaurs. We describe and review theropod dinosaur postcranial remains from the Aptian-Albian Otway and Strzelecki groups, based on at least 37 isolated bones, and more than 90 teeth from the Flat Rocks locality. Several specimens of medium- and large-bodied individuals (estimated up to ~8.5 metres long) represent allosauroids. Tyrannosauroids are represented by elements indicating medium body sizes (~3 metres long), likely including the holotype femur of Timimus hermani, and a single cervical vertebra represents a juvenile spinosaurid. Single specimens representing medium- and small-bodied theropods may be referrable to Ceratosauria, Ornithomimosauria, a basal coelurosaur, and at least three taxa within Maniraptora. Thus, nine theropod taxa may have been present. Alternatively, four distinct dorsal vertebrae indicate a minimum of four taxa. However, because most taxa are known from single bones, it is likely that small-bodied theropod diversity remains underestimated. The high abundance of allosauroids and basal coelurosaurs (including tyrannosauroids and possibly ornithomimosaurs), and the relative rarity of ceratosaurs, is strikingly dissimilar to penecontemporaneous dinosaur faunas of Africa and South America, which represent an arid, lower-latitude biome. Similarities between dinosaur faunas of Victoria and the northern continents concern the proportional representatation of higher clades, and may result from the prevailing temperate-polar climate of Australia, especially at high latitudes in Victoria, which is similar to the predominant warm-temperate climate of Laurasia, but distinct from the arid climate zone that covered extensive areas of Gondwana. Most dinosaur groups probably attained a near-cosmopolitan distribution in the Jurassic, prior to fragmentation of the Pangaean supercontinent, and some aspects of the hallmark 'Gondwanan' fauna of South America and Africa may therefore reflect climate-driven provinciality, not vicariant evolution driven by continental fragmentation. However, vicariance may still be detected at lower phylogenetic levels.     

New Horned Dinosaurs from Utah Provide Evidence for Intracontinental Dinosaur Endemism

Background

During much of the Late Cretaceous, a shallow, epeiric sea divided North America into eastern and western landmasses. The western landmass, known as Laramidia, although diminutive in size, witnessed a major evolutionary radiation of dinosaurs. Other than hadrosaurs (duck-billed dinosaurs), the most common dinosaurs were ceratopsids (large-bodied horned dinosaurs), currently known only from Laramidia and Asia. Remarkably, previous studies have postulated the occurrence of latitudinally arrayed dinosaur “provinces,” or “biomes,” on Laramidia. Yet this hypothesis has been challenged on multiple fronts and has remained poorly tested.

Methodology/Principal Findings

Here we describe two new, co-occurring ceratopsids from the Upper Cretaceous Kaiparowits Formation of Utah that provide the strongest support to date for the dinosaur provincialism hypothesis. Both pertain to the clade of ceratopsids known as Chasmosaurinae, dramatically increasing representation of this group from the southern portion of the Western Interior Basin of North America. Utahceratops gettyi gen. et sp. nov.—characterized by short, rounded, laterally projecting supraorbital horncores and an elongate frill with a deep median embayment—is recovered as the sister taxon to Pentaceratops sternbergii from the late Campanian of New Mexico. Kosmoceratops richardsoni gen. et sp. nov.—characterized by elongate, laterally projecting supraorbital horncores and a short, broad frill adorned with ten well developed hooks—has the most ornate skull of any known dinosaur and is closely allied to Chasmosaurus irvinensis from the late Campanian of Alberta.

Conclusions/Significance

Considered in unison, the phylogenetic, stratigraphic, and biogeographic evidence documents distinct, co-occurring chasmosaurine taxa north and south on the diminutive landmass of Laramidia. The famous Triceratops and all other, more nested chasmosaurines are postulated as descendants of forms previously restricted to the southern portion of Laramidia. Results further suggest the presence of latitudinally arrayed evolutionary centers of endemism within chasmosaurine ceratopsids during the late Campanian, the first documented occurrence of intracontinental endemism within dinosaurs.     

Small theropod teeth from the Upper Jurassic coal mine of Guimarota (Portugal)

Isolated teeth of small theropod dinosaurs from the Upper Jurassic lignite coal mine of Guimarota (near Leiria, Portugal) are described and illustrated. The well known Upper Jurassic theropods from Europe,Archaeopteryx andCompsognathus, are the most common taxa in the Guimarota assemblage. One morphotype is closely related to an allosaurid theropod. Six further morphotypes of theropod teeth are also described, which are closely related to Cretaceous theropods such as dromaeosaurids, troodontids, tyrannosaurids,Richardoestesia andParonychodon. A Late Jurassic origin of these groups of theropods, which is very often postulated, is discussed.     

Additional dinosaur teeth from the Cenomanian (Late Cretaceous) of Charentes,southwestern France

Some isolated teeth of theropod and sauropod dinosaurs from the Cenomanian (Late Cretaceous) of Charentes are described. Two new teeth of Troodontidae confirm the presence of this theropod family, previously based on a single specimen. New dental morphotypes are recognized within Dromaeosauridae and Brachiosauridae in comparison with those already known from Charentes. Lastly, a very small tooth is tentatively assigned to an embryonic or neonatal sauropod. The palaeobiogeographical history of European hadrosauroids is briefly discussed. This history was probably more complex than it appears, involving exchanges with both North America and Asia as early as the mid-Cretaceous (Albian–Cenomanian).     

Dinosaurs of Romania

The dinosaurs of Romania are exclusively Cretaceous. Lowermost Cretaceous dinosaurs come from a bauxite mine in the Bihor county (northwest Romania) that has yielded thousands of disarticulated bones. Uppermost Cretaceous dinosaurs have been known from the Haţeg Basin (south Transylvania) since the end of the 19th century, mostly as bone concentrations (‘fossiliferous pockets’); more recently, nests with dinosaur eggs, including hatchlings, have been found in Haţeg. Although separated by a ca 60 Myr gap, the two dinosaur faunas from Romania share some common features: predominance of ornithopods, absence of large theropods (substituted in the case of the Maastrichtian Haţeg assemblage by several small theropods), and, in general, the small size of the individuals (dwarfism). These aspects seem to be explained by the isolated island habitat of both assemblages. To cite this article: D. Grigorescu, C. R. Palevol 2 (2003) 97–101.     

A New Large-Bodied Oviraptorosaurian Theropod Dinosaur from the Latest Cretaceous of Western North America

The oviraptorosaurian theropod dinosaur clade Caenagnathidae has long been enigmatic due to the incomplete nature of nearly all described fossils. Here we describe Anzu wyliei gen. et sp. nov., a new taxon of large-bodied caenagnathid based primarily on three well-preserved partial skeletons. The specimens were recovered from the uppermost Cretaceous (upper Maastrichtian) Hell Creek Formation of North and South Dakota, and are therefore among the stratigraphically youngest known oviraptorosaurian remains. Collectively, the fossils include elements from most regions of the skeleton, providing a wealth of information on the osteology and evolutionary relationships of Caenagnathidae. Phylogenetic analysis reaffirms caenagnathid monophyly, and indicates that Anzu is most closely related to Caenagnathus collinsi, a taxon that is definitively known only from a mandible from the Campanian Dinosaur Park Formation of Alberta. The problematic oviraptorosaurs Microvenator and Gigantoraptor are recovered as basal caenagnathids, as has previously been suggested. Anzu and other caenagnathids may have favored well-watered floodplain settings over channel margins, and were probably ecological generalists that fed upon vegetation, small animals, and perhaps eggs.     

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.     

Tyrant Dinosaur Evolution Tracks the Rise and Fall of Late Cretaceous Oceans

The Late Cretaceous (∼95–66 million years ago) western North American landmass of Laramidia displayed heightened non-marine vertebrate diversity and intracontinental regionalism relative to other latest Cretaceous Laurasian ecosystems. Processes generating these patterns during this interval remain poorly understood despite their presumed role in the diversification of many clades. Tyrannosauridae, a clade of large-bodied theropod dinosaurs restricted to the Late Cretaceous of Laramidia and Asia, represents an ideal group for investigating Laramidian patterns of evolution. We use new tyrannosaurid discoveries from Utah—including a new taxon which represents the geologically oldest member of the clade—to investigate the evolution and biogeography of Tyrannosauridae. These data suggest a Laramidian origin for Tyrannosauridae, and implicate sea-level related controls in the isolation, diversification, and dispersal of this and many other Late Cretaceous vertebrate clades.     

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.     

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.     

Phylogeny,Histology and Inferred Body Size Evolution in a New Rhabdodontid Dinosaur from the Late Cretaceous of Hungary

Background

Rhabdodontid ornithopod dinosaurs are characteristic elements of Late Cretaceous European vertebrate faunas and were previously collected from lower Campanian to Maastrichtian continental deposits. Phylogenetic analyses have placed rhabdodontids among basal ornithopods as the sister taxon to the clade consisting of Tenontosaurus, Dryosaurus, Camptosaurus, and Iguanodon. Recent studies considered Zalmoxes, the best known representative of the clade, to be significantly smaller than closely related ornithopods such as Tenontosaurus, Camptosaurus, or Rhabdodon, and concluded that it was probably an island dwarf that inhabited the Maastrichtian Haţeg Island.

Methodology/Principal Findings

Rhabdodontid remains from the Santonian of western Hungary provide evidence for a new, small-bodied form, which we assign to Mochlodon vorosi n. sp. The new species is most similar to the early Campanian M. suessi from Austria, and the close affinities of the two species is further supported by the results of a global phylogenetic analysis of ornithischian dinosaurs. Bone histological studies of representatives of all rhabdodontids indicate a similar adult body length of 1.6–1.8 m in the Hungarian and Austrian species, 2.4–2.5 m in the subadults of both Zalmoxes robustus and Z. shqiperorum and a much larger, 5–6 m adult body length in Rhabdodon. Phylogenetic mapping of femoral lengths onto the results of the phylogenetic analysis suggests a femoral length of around 340 mm as the ancestral state for Rhabdodontidae, close to the adult femoral lengths known for Zalmoxes (320–333 mm).

Conclusions/Significance

Our analysis of body size evolution does not support the hypothesis of autapomorhic nanism for Zalmoxes. However, Rhabdodon is reconstructed as having undergone autapomorphic giantism and the reconstructed small femoral length (245 mm) of Mochlodon is consistent with a reduction in size relative to the ancestral rhabdodontid condition. Our results imply a pre-Santonian divergence between western and eastern rhabdodontid lineages within the western Tethyan archipelago.     

A New Hadrosauroid Dinosaur from the Early Late Cretaceous of Shanxi Province,China

Background

The origin of hadrosaurid dinosaurs is far from clear, mainly due to the paucity of their early Late Cretaceous close relatives. Compared to numerous Early Cretaceous basal hadrosauroids, which are mainly from Eastern Asia, only six early Late Cretaceous (pre-Campanian) basal hadrosauroids have been found: three from Asia and three from North America.

Methodology/Principal Findings

Here we describe a new hadrosauroid dinosaur, Yunganglong datongensis gen. et sp. nov., from the early Late Cretaceous Zhumapu Formation of Shanxi Province in northern China. The new taxon is represented by an associated but disarticulated partial adult skeleton including the caudodorsal part of the skull. Cladistic analysis and comparative studies show that Yunganglong represents one of the most basal Late Cretaceous hadrosauroids and is diagnosed by a unique combination of features in its skull and femur.

Conclusions/Significance

The discovery of Yunganglong adds another record of basal Hadrosauroidea in the early Late Cretaceous, and helps to elucidate the origin and evolution of Hadrosauridae.     

Dinosaurs and the Cretaceous Terrestrial Revolution

The observed diversity of dinosaurs reached its highest peak during the mid- and Late Cretaceous, the 50 Myr that preceded their extinction, and yet this explosion of dinosaur diversity may be explained largely by sampling bias. It has long been debated whether dinosaurs were part of the Cretaceous Terrestrial Revolution (KTR), from 125-80 Myr ago, when flowering plants, herbivorous and social insects, squamates, birds and mammals all underwent a rapid expansion. Although an apparent explosion of dinosaur diversity occurred in the mid-Cretaceous, coinciding with the emergence of new groups (e.g. neoceratopsians, ankylosaurid ankylosaurs, hadrosaurids and pachycephalosaurs), results from the first quantitative study of diversification applied to a new supertree of dinosaurs show that this apparent burst in dinosaurian diversity in the last 18 Myr of the Cretaceous is a sampling artefact. Indeed, major diversification shifts occurred largely in the first one-third of the group's history. Despite the appearance of new clades of medium to large herbivores and carnivores later in dinosaur history, these new originations do not correspond to significant diversification shifts. Instead, the overall geometry of the Cretaceous part of the dinosaur tree does not depart from the null hypothesis of an equal rates model of lineage branching. Furthermore, we conclude that dinosaurs did not experience a progressive decline at the end of the Cretaceous, nor was their evolution driven directly by the KTR.