Some aspects of Cretaceous ostracod biostratigraphy of South Africa and relationships with other Gondwanide localities

One hundred and thirty-six species, representing 67 genera have been recorded from the late Jurassic-Maastrichtian marine sediments of South Africa. The faunas show a major dichotomy across a regionally-developed late Cenomanian-early Coniacian hiatus with the Portlandian-Cenomanian Cytheruridae/Progonocytheridae/Schizocytheridae dominated faunas being replaced in the Coniacian by Trachyleberididae/Brachycytheridae/Schizocytheridae dominated faunas. Comparison with other Gondwanide localities shows that the two South African basins from which ostracods have been described (Outeniqua and Natal/Zululand) formed part of a Callovian-Cenomanian South Gondwana ostracod province that stretched from the Neuquen Basin of Argentina to Madagascar/Tanzania/Kutch and west Australia. The most characteristic and cosmopolitan forms within this province belong to the Majungaella/Amicytheridea/Progonocythere group, along with Arculicythere in the Aptian-Cenomanian.In Tanzania, (the only locality of the old South Gondwana province where the succession is complete) these assemblages are replaced in the Turonian by the influx of Brachycythere, and Cythereis and various other trachyleberids. Changes of a similar nature are seen whenever marine sedimentation resumed after the local “mid” Cretaceous hiatus (South Africa, India, Argentina). Argentina differs in not having Brachycythere, whose rapid appearance in the West Indian Ocean basin soon after its earliest record in Brazil, is attributed to the destruction of the barrier at the eastern end of the Walvis Ridge/Rio Grande Rise in late Cenomanian or early Turonian times. Despite this common element with Brazil and West Africa, the South African Coniacian to Maastrichtian faunas are closer to those of Tanzania and Australia than they are to either Argentina or Brazil/West Africa. In Zululand they show evidence of a steady increase in water depth, leading to the establishment of progressively more diverse cytheracean populations, with a particularly large increase across the Santonian/Campanian boundary.     

Theropod remains from the uppermost Cretaceous of Colombia and their implications for the palaeozoogeography of western Gondwana

Dinosaur remains from Upper Cretaceous outcrops of northern Gondwana are extremely rare, in contrast with the much richer sample of coeval beds from southern Gondwana. Dinosaur remains from the uppermost Cretaceous Ortega locality of the Upper Magdalena Basin (Maastrichtian) of the Department of Tolima, Colombia, provides new information on northern Gondwanan faunas of this time. A revision of dinosaur material from this outcrop, consisting of three theropod shed teeth, reveals the presence of two morphotypes. One of them is referred to Abelisauridae based on the presence of crowns with mesial margin with a strong curvature beginning at about the second-third of the crown height and straight to slightly concave distal margin. The second morphotype exhibits un-serrated mesial and distal margins without carinae and no constriction at the base of the crown, a combination of features only observed in unenlagiine dromaeosaurids within Theropoda. Members of these clades are also present in coeval beds of southern and central South America, Madagascar, northern Africa, and India, indicating a cosmopolitan distribution in western and central Gondwana during the Late Cretaceous. Regarding South America, abelisaurid and probably dromaeosaurid theropods are recorded across a large latitudinal area, from the Palaeo-Equator to considerably high palaeo-latitudes in Patagonia, and probably spanning quite different environmental conditions.     

Biogeographical network analysis of Cretaceous Australian dinosaurs

The biogeography of Cretaceous Australian dinosaur fauna has been characterized variously as endemic, cosmopolitan or closely related to other Gondwanan faunas. Over the past decade, a large number of new Australian dinosaur taxa have been described and included in phylogenetic analyses, allowing for new insights into their biogeographical affinities to be obtained. Here, I combine the latest phylogenies of Australian dinosaurs into a supertree that includes as many Australian taxa as possible and apply it to the construction of biogeographical networks of Cretaceous dinosaurs. The results show that the Cretaceous dinosaurs of Australia have a strong connection with South American dinosaurs and to those of other Gondwanan continents via South America. Community detection algorithms indicate that the Gondwanan continents form a community within Cretaceous dinosaur biogeographical networks. The biogeographical affinities between Australia and Laurasia detected in previous studies were largely due to the scarce dinosaur fossil record of Gondwana at that time and to the unstable phylogenetic position of Australian dinosaurs because of the fragmentary nature of their fossils. Continuing discoveries of new Australian taxa and refinement of resulting phylogenetic analyses can further deepen our understanding of the biogeographical history of Australia.     

Les continents péri-atlantiques au crétacé supérieur: Migrations des faunes continentales et problèmes paléogéographiques

Terrestrial vertebrates offer possibilities of reconstructing the migrations by land-routes followed during the late Cretaceous on the peri-Atlantic continents (North America, South America, Europe, Africa). South America and Africa were not separated before the Aptian. Later, migrations could still have occurred between Africa and South America during the late Cretaceous by a land-route (probably discontinuous) situated on the Rio Grande Rise-Walvis Ridge barrier; it is not impossible that some amphibia used this route. In Laurasia, two provinces were largely separated during the early part of the late Cretaceous: Euramerica and Asiamerica in the terminology of Cox (1974), (that is, Europe plus eastern North America and Asia plus western North America). During the latest Cretaceous, western North America became connected with Euramerica, but probably separated from Asia. During the latest Cretaceous, a route, probably terrestrial, permitted important faunal exchanges between South America and Laurasia; this connection was situated in the Caribbean region, perhaps where Central America is today. Limited faunal exchange occurred between Euramerica and North Africa.     

Zurupleuropholis gen. nov. (Teleostei,Albian, Argentina), first pleuropholids from the Cretaceous of South America

Although the Cretaceous is characterized by a rich fish diversity, Cretaceous continental fishes from Gondwana are poorly known and comparatively scarce. Among these fishes, the family Pleuropholidae is only known by a few species relatively poorly preserved, from the Middle Jurassic to Lower Cretaceous of Europe, Africa, North America, and South America. In this paper, two new species of the pleuropholid new genus Zurupleuropholis are described, Z. quijadensis gen. et sp. nov. and Z. decollavi gen. et sp. nov. The new fishes were recovered in the Lower Cretaceous lacustrine Lagarcito Formation of central-west Argentina. This taxon constitutes a relevant finding considering that the representation of the family Pleuropholidae is rare worldwide. Zurupleuropholis gen. nov. appears to be the youngest known member of Pleuropholidae, and it represents the second record of the family in South America and the first record in the Cretaceous of the continent.     

A large-clawed theropod (Dinosauria: Tetanurae) from the Lower Cretaceous of Australia and the Gondwanan origin of megaraptorid theropods

Megaraptoridae comprises a clade of enigmatic Gondwanan theropods with characteristic hypertrophied claws on the first and second manual digits. The majority of megaraptorids are known from South America, although a single genus (Australovenator) plus additional indeterminate material is also known from Australia. This clade has a controversial placement among theropods, and recently has been interpreted alternatively as a carcharodontosaurian or a tyrannosauroid lineage. We describe new fragmentary but associated postcranial remains from the opal fields of Lightning Ridge (middle-Albian, Griman Creek Formation) in north-central New South Wales. The new unnamed taxon exhibits a number of unusual features that suggest the presence of a hitherto unrecognised Australian megaraptorid. From an Australian perspective, the Lightning Ridge taxon predates Australovenator by ca. 10 Ma and is minimally coeval with megaraptoran material reported from the Eumeralla Formation of Victoria (but potentially 6.1–9.5 Ma younger). It is also notable as the largest predatory dinosaur yet identified from Australia and is only the second theropod known from more than a single element. A Bayesian phylogenetic approach integrating morphological, stratigraphic and palaeogeographic information tested both the carcharodontosaurian and tyrannosauroid placements for Megaraptora. Regardless of the preferred placement among Tetanurae, rigorous palaeobiogeographic analyses support an Asian origin of Megaraptora in the latest Jurassic (about 150–135 Ma), an Early Cretaceous (about 130–121 Ma) divergence of the Gondwanan lineage leading to Megaraptoridae, and an Australian root for megaraptorid radiation. These results indicate that Australia's Cretaceous dinosaur fauna did not comprise simply of immigrant taxa but was a source for complex two-way interchange between Australia–Antarctica–South America leading to the evolution of at least one group of apex predatory dinosaurs in Gondwana.     

A new brachyrostran with hypertrophied axial structures reveals an unexpected radiation of latest Cretaceous abelisaurids

A well preserved skeleton of a new abelisaurid is reported here. The holotype of Viavenator exxoni was found in the outcrops of the Bajo de la Carpa Formation (Santonian, Upper Cretaceous), northwestern Patagonia, Argentina. This new taxon belongs to the South American clade of abelisaurids, the brachyrostrans. The current phylogenetic analysis places it as the basalmost member of a new clade of derived brachyrostrans, named Furileusauria, characterized by distinctive cranial, axial and appendicular anatomical features. The Santonian age of Viavenator allows filling the stratigraphic gap exhibited between the basal brachyrostrans of Cenomanian–Turonian age, and the derived forms from the Campanian-Maastrichtian. The evolution of abelisaurids during the Late Cretaceous, faunal replacements, and the adaptive radiation that occurred during that period of time in South America are discussed.     

Tetrapods from the Upper Cretaceous (Turonian–Maastrichtian) Bauru Group of Brazil: a reappraisal

An updated, annotated list of all tetrapods from the Adamantina, Uberaba and Marília formations (Bauru Group), which constitute some of the best studied Upper Cretaceous units in Brazil, is presented. Tetrapod diversity in the Bauru Group is remarkable, including an admixture of typically austral Gondwanan taxa (e.g., abelisaurids, notosuchians) and boreal Gondwanan forms (e.g., carcharodontosaurids). Of note is the absence of Laurasian taxa in the upper portion of the Bauru Group. With the exception of some turtles, an anuran, mesoeucrocodylians and one titanosaur, most taxa from the Bauru Group are based on fragmentary and isolated bones, and as such many specimens can be identified only to a higher taxonomic level. Fishes, turtles, anurans, mesoeucrocodylians, dinosaurs, birds and mammals from the Adamantina and Marília formations resemble the latest Late Cretaceous vertebrate faunas from southern South America, except for the absence of ornithischian dinosaurs.     

白垩纪四足动物足印的生物地层学、生物年代学与遗迹相

从全球范围来看,白垩纪四足动物的足印多数是非鸟恐龙与鸟类留下的痕迹;少量足印来自翼龙、鳄鱼、龟、哺乳动物和其他四足动物。白垩纪的足迹化石以东亚(尤其是中国和朝鲜)和北美西部的最为人所知。南美(主要是阿根廷和巴西)也有一定数量广泛分布的足迹化石,欧洲、非洲与澳大利亚的白垩纪足迹组合则鲜为人知。以白垩纪四足动物的足印记录为基础,我们对两个全球足印生物年代重新进行了检查。早白垩世生物年代以蜥脚类与鸟脚类的足迹为特征。晚白垩世生物年代中的蜥脚类足迹较少,但是鸭嘴龙、暴龙和角龙的足迹增多了。另外,白垩纪足印化石的记录中记载了许多重要的生物地层学信息,如北美白垩纪中期蜥脚类恐龙的消失,以及白垩纪末恐龙的绝灭。越来越多来自东亚的白垩纪足印记录使我们对更精细的地方性白垩纪足印生物年代学有了初步印象。因此,以地方性四足恐龙(包括鸟类)遗迹属的地层分布为基础,可以识别出三个或四个足印生物年代。种类丰富并具有地方性特色的东亚的白垩纪鸟类动物的遗迹群,可能指示白垩纪时东亚存在着一个独特而繁盛的鸟类动物群。以足印化石为基础的这一假说有待进一步的验证。     

Diversity and faunal changes in the latest Cretaceous dinosaur communities of southwestern Europe

Characterization of macroecological patterns for latest Cretaceous dinosaur communities is essential to understand how those faunas were changing during the run-up to the Cretaceous-Paleogene extinction event, and thus the cause of the extinction. Outside of the well-studied latest Cretaceous dinosaurs of North America, southwestern Europe (France, Spain and Portugal) preserves one of the richest end-Cretaceous dinosaur fossil records, as it has produced hundreds of dinosaur fossil localities. We compiled a comprehensive database of all dinosaur fossil occurrences from the uppermost Cretaceous of the Ibero-Armorican region and analyze it statistically, providing the first numerical study of the ecological and taxonomic diversities of these communities. Our study corroborates previous work that has identified a major faunal change in the latest Cretaceous terrestrial vertebrate assemblages, and places this event around the C31r-C31n reversal, in the early late Maastrichtian (c. 69 Ma). Significant differences in ecological diversity metrics (dominance, Shannon and Simpson) characterize the pre- and post-turnover assemblages. The turnover event, therefore, did not only lead to a taxonomic replacement but also important reorganizations in the structure of dinosaur communities. Herbivorous dinosaurs suffered the most dramatic alterations across the turnover, in terms of relative dominances, by shifting their contributions within the communities (hadrosauroids replacing titanosaurids as the dominant taxon in the medium-to large-bodied herbivore niche) or even disappearing (rhabdodontids and nodosaurids). The carnivores apparently maintained similar relative abundances before and after the turnover, and the relative proportions between carnivorous and herbivorous taxa remained static through time. Further improvement of the present database might allow for the identification of new ecological patterns, and higher-resolution comparison with the North American records.     

A ceratopsian dinosaur from the Late Cretaceous of eastern North America,and implications for dinosaur biogeography

Tyrannosaurs and hadrosaurs from the Late Cretaceous of eastern North America (Appalachia) are distinct from those found in western North America (Laramidia), suggesting that eastern North America was isolated during the Late Cretaceous. However, the Late Cretaceous fauna of Appalachia remains poorly known. Here, a partial maxilla from the Campanian Tar Heel Formation (Black Creek Group) of North Carolina is shown to represent the first ceratopsian from the Late Cretaceous of eastern North America. The specimen has short alveolar slots, a ventrally projected toothrow, a long dentigerous process overlapped by the ectopterygoid, and a toothrow that curves laterally, a combination of characters unique to the Leptoceratopsidae. The maxilla has a uniquely long, slender and downcurved posterior dentigerous process, suggesting a specialized feeding strategy. The presence of a highly specialized ceratopsian in eastern North America supports the hypothesis that Appalachia underwent an extended period of isolation during the Late Cretaceous, leading the evolution of a distinct dinosaur fauna dominated by basal tyrannosauroids, basal hadrosaurs, ornithimimosaurs, nodosaurs, and leptoceratopsids. Appalachian vertebrate communities are most similar to those of Laramidia. However some taxa-including leptoceratopsids-are also shared with western Europe, raising the possibility of a Late Cretaceous dispersal route connecting Appalachia and Europe.     

Intracontinental movements in Western Gondwanaland: a palaeomagnetic test

Early Cretaceous intracontinental movements within Africa and/or South America, to account for misfit problem in the pre-drift reconstruction of Western Gondwana, have been tested using palaeomagnetic poles from both continents. Each continent has been considered as comprising separate subplates according to the boundaries proposed by Pindell and Dewey (1982) and Curie (1983): i.e., northern Africa ( ), southern Africa ( ), northern South America (nsa) and southern South America ( ). Visual and statistical distribution of Late Permian to Jurassic poles from the African subplates indicate two distinct groups before rotating relative to , while after the rotation they become indistinguishable. The distribution and sparsity of data from South America limit the study in this continent as the data are statistically indistinguishable before and after rotation of with respect to . When the test is extended to the reconstruction of Western Gondwana, the tightest grouping of the poles occurs when an intracontinental movement is assumed within Africa. This suggests that the misfit between the two continents is probably due to intracontinental deformation within Africa and not in South America. If confirmed, this may have profound implications in the kinematics of the processes related to the opening of the South and Central Atlantic Oceans and also the origin of the West African Rift System.     

Gondwanan dinosaur evolution and biogeographic analysis

Gondwanan dinosaurs, though less well-known than their Laurasian counterparts,are being discovered and described at an ever accelerating pace. Dinosaurs are known from every major Gondwanan landmass, including South America, Africa, Antarctica, Australia, India, Madagascar and New Zealand. Much of the Gondwanan dinosaur literature includes speculation on vicariant and dispersal events relating to the fragmentation of Gondwana during the Jurassic and Cretaceous. These prolific biogeographic hypotheses are often inconclusive, speculative in nature, and untestable with the data at hand. To formulate well-supported biogeographic hypotheses, resolved dinosaur phylogenies and sound hypotheses of Earth history are necessary. Other factors, such as taphonomy, depositional setting, and missing taxonomic, temporal and geographic data, also contribute to understanding dinosaur faunal compositions. Additional phylogenetic information, a continuation of the current exploration and discovery, and knowledge of recent advances in tectonic plate reconstructions are paramount to developing a well-supported view of Gondwanan dinosaur biogeography.     

Neuroanatomy of the abelisaurid theropod Viavenator: The most complete reconstruction of a cranial endocast and inner ear for a South American representative of the clade

The type braincase of Viavenator exxoni (MAU-Pv-LI-530) was recovered complete and isolated from most of the other skull bones. Although the braincase is crossed by numerous fractures, using CT scans allowed the generation of 3D renderings of the endocranial cavity enclosing the brain, cranial nerves, and blood vessels, as well as the labyrinth of the inner ear. Within the abelisaurids, the only taxon with a complete braincase and known endocranial morphology is Majungasaurus crenatissimus, from Madagascar. In turn, in Argentina, partial endocranial morphology is known for another two Cretaceous forms: Abelisaurus comahuensis and Aucasaurus garridoi. Here, we present the most complete reconstruction of the neuroanatomy for a representative of the clade in South America. These findings add knowledge to the field of theropod paleoneuroanatomy in general, and abelisaurid diversity in particular. Comparisons of Viavenator with other abelisaurids indicate greater similarity with Aucasaurus than with Majungasaurus, suggesting that South American forms shared the same neurosensorial capabilities, which include larger flocculus of the cerebellum and larger olfactory ratios than the form from Madagascar.     

全球显生宙碳酸盐岩时空分布规律及其控制因素

世界上约60%的油气产自碳酸盐岩。全球哪些地区、哪些时代碳酸盐岩发育？其时空分布受哪些因素控制？弄清这些问题,不仅对我国海外油气勘探战略选区有指导作用,而且可为了解全球古地理环境演化提供重要信息。根据全球179个碳酸盐岩盆地的数据统计分析,对全球显生宙碳酸盐岩时空分布规律进行了研究,并探讨了其控制因素。在显生宙的各个地质时期,碳酸盐岩均有分布,但不同时期,碳酸盐岩发育程度不同。在泥盆纪、白垩纪和古近纪,碳酸盐岩分布广泛,而在志留纪、二叠纪、三叠纪和侏罗纪,分布局限。不同时期,碳酸盐岩发育地区不同。寒武纪-奥陶纪,碳酸盐岩主要分布于俄罗斯、中国、北美洲、澳大利亚;三叠纪以后,碳酸盐岩发育区域转移至中东、北欧、北非、南美洲;至古近纪和新近纪,碳酸盐岩发育区主要分布于中东、北非、南亚地区。研究表明,全球碳酸盐岩时空分布受大陆漂移和全球海平面变化控制。古生代,古劳亚大陆、西伯利亚、中国华南地区、澳大利亚均位于赤道附近温暖浅海地带,碳酸盐岩发育,上述地区是这一时期碳酸盐岩分布主要区域;冈瓦纳大陆在古生代位于高纬度区,碳酸盐岩少。中生代,古劳亚大陆漂移至高纬度区,碳酸盐岩减少;冈瓦纳大陆解体为南美板块、非洲板块并漂移至低纬度区,发育碳酸盐岩。新生代,碳酸盐岩在南亚地区的增多,这也和板块的位置相印证。另外,当全球海平面上升时,海侵形成广阔的陆表海,碳酸盐岩广泛发育;当全球海平面下降时,海退形成陆缘海,碳酸盐岩发育面积减小。     

Mawsoniid remains (Sarcopterygii: Actinistia) from the lacustrine Missão Velha Formation (Lower Cretaceous) of the Araripe Basin,North-East Brazil

Mawsoniids are a lineage of extinct fresh/brackish water coelacanth fishes, common in Cretaceous Godwanan deposits of South America, North and West Africa and Madagascar. Here we formally describe mawsoniid remains from the fluvio-lacustrine Missão Velha Formation (Lower Cretaceous) of the Araripe Basin, North-East Brazil. The examples from the Missão Velha Formation are here described as Mawsonia cf. gigas, based mainly on elements of the lower jaw and opercular series. The occurrence of M. cf. gigas in the Missão Velha Formation expands the stratigraphic and geographical ranges of the type species.     

First report of mammal-like tracks from the Cretaceous of North Africa (Tunisia)

This paper describes Cretaceous mammal-like tracks from southern Tunisia. The tracks, referred to the Cenomanian Kerker Member of the Zebbag Formation, are the first mammal-like footprints reported from the Cretaceous of North Africa. The good preservation of the two tracks and their distinctive morphology support their attribution to a mammalian trackmaker, although the limited available data prevents attribution to a specific ichnotaxon. Morphologically, the Tunisian tracks resemble those of modern Mustelidae, however, based on mammalian faunas in the Cretaceous of Africa, they probably have affinity with members of Multituberculate family. Theropod dinosaur and bird tracks occur on the same track-bearing layer. The sediments are interpreted as an arid tidal flat environment, suggesting that African mammals might have shared their environment with a diverse fauna of larger animals.     

Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia

Patagonia has yielded the most comprehensive fossil record of Cretaceous theropods from Gondwana, consisting of 31 nominal species belonging to singleton taxa and six families: Abelisauridae, Noasauridae, Carcharodontosauridae, Megaraptoridae nov. fam., Alvarezsauridae, and Unenlagiidae. They provide anatomical information that allows improved interpretation of theropods discovered in other regions of Gondwana. Abelisauroids are the best represented theropods in Patagonia. They underwent an evolutionary radiation documented from the Early Cretaceous through to the latest Cretaceous, and are represented by the clades Abelisauridae and Noasauridae. Patagonian carcharodontosaurids are known from three taxa (Tyrannotitan, Giganotosaurus and Mapusaurus), as well as from isolated teeth, collected from Aptian to Cenomanian beds. These allosauroids constituted the top predators during the mid-Cretaceous, during which gigantic titanosaur sauropods were the largest herbivores. Megaraptorans have become better documented in recent years with the discovery of more complete remains. Megaraptor, Aerosteon and Orkoraptor have been described from Cretaceous beds from Argentina, and these taxa exhibit close relationships with the Aptian genera Australovenator, from Australia, and Fukuiraptor, from Japan. The Gondwanan megaraptorans are gathered into the new family Megaraptoridae, and the Asiatic Fukuiraptor is recovered as the immediate sister taxon of this clade. Although megaraptorans have been recently interpreted as members of Allosauroidea, we present evidence that they are deeply nested within Coelurosauria. Moreover, anatomical information supports Megaraptora as more closely related to the Asiamerican Tyrannosauridae than thought. Megaraptorans improve our knowledge about the scarcely documented basal radiation of Gondwanan coelurosaurs and tyrannosauroids as a whole. Information at hand indicates that South America was a cradle for the evolutionary radiation for different coelurosaurian lineages, including some basal forms (e.g., Bicentenaria, Aniksosaurus), megaraptorans, alvarezsaurids less derived than those of Laurasia, and unenlagiids, revealing that Gondwanan coelurosaurs played sharply differing ecological roles, and that they were taxonomically as diverse as in the northern continents. The unenlagiids represent an endemic South American clade that has been recently found to be more closely related to birds than to dromaeosaurid theropods. Analysis of the theropod fossil record from Gondwana shows the highest peak of origination index occurred during the Aptian–Albian and a less intense one in the Campanian time spans. Additionally, peaks of extinction index are recognized for the Cenomanian and Turonian–Coniacian time spans. In comparison, the Laurasian pattern differs from that of Gondwana in the presence of an older extinction event during the Aptian–Albian time-span and a high origination rate during the Cenomanian time-bin. Both Laurasian and Gondwanan theropod records show a peak of origination rates during the Campanian.