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.     

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.     

A new ornithopod (Dinosauria,Ornithischia) from the Upper Cretaceous of Antarctica and its palaeobiogeographical implications

A new ornithopod dinosaur from the Upper Cretaceous (Maastrichtian) Snow Hill Island Formation, at James Ross Island, Antarctica is here described. This new taxon, named as Morrosaurus antarcticus gen. et sp. nov., is represented by a fragmentary right hind limb belonging to a medium-sized individual. Our phylogenetic analysis nests the new taxon in a monophyletic clade of Southern Hemisphere ornithopods that includes most Patagonian and Antarctic ornithopods. Several members of this group share a slender and bunched foot with narrow metatarsal IV, expanded chevrons, and bowed humerus without deltopectoral crest. Several features indicate that these ornithopods exhibit adaptations for a specialized cursorial mode of life. The recognition of Patagonian and Antarctic Ornithopoda belonging to a monophyletic clade reinforces palaeobiogeographical signals indicating that Patagonia, Antarctica and Australia shared a common Late Cretaceous terrestrial fauna.     

Anurans from the Candeleros Formation (?Cenomanian-Turonian) of west-central Argentina: new evidence for pipoid evolution

The fluvio-lacustrine deposits of the ?Cenomanian-Turonian Candeleros Formation in the northwestern fringe of Patagonia have yielded numerous remains of vertebrates, including anurans. A new, partially articulated immature specimen of a small pipimorph pipoid from the upper part of this unit is described herein. Although incompletely preserved, the morphology and proportions of the skeleton are strongly reminiscent of corresponding elements of the holotype of Avitabatrachus uliana from the lower section of the same formation at a nearby locality, thus suggesting that these specimens might represent the same, or a closely related, taxon. The new specimen is a metamorphosing individual: it has a zygapophyseal articulation between the sacrum and the neural arch of a postsacral vertebra that bears conspicuous transverse processes, and four pairs of ribs. These traits are not present in the available material of A. uliana, but they might have occurred within the normal range of variation of a single species. Although the generic and specific allocation of the new material is uncertain, a parsimony analysis performed to clarify the relationships of A. uliana supports a phylogenetic placement of this taxon outside the crown Pipidae. The evidence provided by the Candeleros anurans confirms the presence of stem pipids in Patagonia in the early Late Cretaceous.     

Early Cretaceous palm pollen tetrads from Patagonia,Argentina

Pollen tetrads and monads of spiny pollen grains with close affinities to palms have been found in several localities from the Lower Cretaceous (Barremian-Albian) of the Austral Basin (Magallanes), Patagonia (Argentina). When found dispersed, spiny and zonasulcate pollen grains, are commonly referred to the fossil genus Spinizonocolpites Muller, with close affinities to the extant palm Nypa. The Patagonian specimens were compared with fossil and extant members of the Arecaceae, showing close similarities in shape and sculpture with the primitive members of the subfamily Calamoideae. Nypa produces tetragonal tetrads different from the tetrahedral tetrads of the Spinizonocolpites-type recovered from Patagonia. The specimens were studied with LM and SEM. The polarity, aperture orientation and tetrad type allow segregating the Patagonian grains from the Nypoideae and relating it to the Calamoideae subfamily. These records suggest an antique origin of monocots and a probably initial diversification of calamoid palms during the Early Cretaceous in high latitudes of Gondwana. The presence of palmae during the Early Cretaceous in southern South America suggests a warm and humid climate, similar to that found in present days at tropics.     

Frullania pinnata spec. nov. (Frullaniaceae,Porellales), a new leafy liverwort in mid-Cretaceous Burmese amber from Myanmar

An inclusion in Cretaceous amber from Myanmar is described as a new fossil species in the extant liverwort genus Frullania. The name Frullania pinnata is proposed for the taxon that is characterized by entire underleaves paired with a conspicuous pinnate branching pattern, two distinctive and stable morphological features. Entire underleaves are known in several extant Frullania species but had not previously been documented in Frullania in amber. The combination of morphological characters in this new fossil species is unknown in any crown group lineage of Frullania, and may very well represent a stem lineage element of the genus. This discovery is important because it expands our understanding of the diversity of Frullaniaceae in the Burmese amber forest as well as important ramifications for the phylogenetic reconstruction of extant Frullania lineages.     

New Spanish Late Cretaceous eusuchian reveals the synchronic and sympatric presence of two allodaposuchids

The recently described clade Allodaposuchidae includes European eusuchian crocodyliforms restricted to the Late Cretaceous (Campanian and Maastrichtian). A new allodaposuchid crocodyliform is here described based on two specimens from the upper Campanian–lower Maastrichtian fossil site of Lo Hueco (Cuenca, Spain). This new taxon, Agaresuchus fontisensis gen. et sp. nov., is described by two complete skulls and a lower jaw associated with one of them. This new species can be distinguished unambiguously from Lohuecosuchus megadontos, the other allodaposuchid known from the same fossil site. The presence of two allodaposuchid crocodyliforms in Lo Hueco allows the recognition of the synchronic and sympatric existence of two representatives of this clade for the first time. The new genus Agaresuchus, comprises a previously described Iberian allodaposuchid species, “Allodaposuchus” subjuniperus, as Agaresuchus subjuniperus, new combination.     

Two new species of mid-Cretaceous webspinners in amber from northern Myanmar (Embiodea: Clothodidae,Oligotomidae)

Two new genera and species of Cretaceous webspinners (Embiodea) are described and figured, both preserved in amber from northern Myanmar. Atmetoclothoda orthotenes Engel and Huang, gen. et sp. nov., is the first fossil representative of the putatively primitive family Clothodidae, and is segregated into its own subfamily, Atmetoclothodinae Engel and Huang, subfam. nov., owing to its primitive retention of a distinct mentum, a quadrate submentum with straight borders, a postocciptal suture that meets the hypostomal sulcus at the posterior tentorial pit, and subgenae that do not meet medially (thus a ventral bridge or gula is lacking), completely symmetrical terminalia, and unsegmented cerci that apically bear a small inner lamellar projection. Litoclostes delicatus Engel and Huang, gen. et sp. nov., is the first definitive fossil of Oligotomidae (not including a subfossil species in Pleistocene copal and which is likely synonymous with an extant taxon), and differs from modern genera in a combination of head, tarsal, and terminalic morphology. The new fossil species double the known Cretaceous fossils and add a further two families to the Mesozoic record for the order.     

A new Late Cretaceous crocodyliform from the western margin of Gondwana (La Rioja Province,Argentina)

The Cretaceous witnessed the greatest diversity of Gondwanan notosuchian crocodyliforms, which displayed high levels of diversity and a notable array of specialized forms that developed in different ecological niches. Among this diversity, the advanced notosuchians are a clade of mid body sized forms which displays a remarkable abundance but is restricted to two lithological units from the Late Cretaceous of South America, the Adamantina Formation in southeastern Brazil and the Bajo de la Carpa Formation in Patagonia (Argentina). The only exceptions known so far were the Late Cretaceous Labidiosuchus from the Marilia Formation in Brazil and Yacarerani from the Cajones Formation in Bolivia. Herein we report a new Cretaceous crocodyliform, Llanosuchus tamaensis gen. nov. et sp. nov., found in the Los Llanos Formation (Campanian?) in northwestern Argentina (La Rioja Province). The small specimen includes well preserved fragments of the cranium and mandible with an estimated skull size of about 9 cm lengths. This crocodyliform shares several derived characters with Notosuchus terrestris from the Bajo de la Carpa Formation (Patagonia) and it was found in a weakly developed sandy paleosol profile formed in wet local conditions but in a region characterized by semi-arid climate and active eolian sedimentation. The presence of a new advanced notosuchian in the Cretaceous of western Gondwana, and its intermediate geographical region between the known South American species (Brazil, Argentina and Bolivia), has interesting implications, and adds another record of an advanced notosuchian in deposits with inferred warm climates and semi-arid paleoenvironments. Finally, Llanosuchus tamaensis supports a Late Cretaceous age for Los Llanos Formation with important geological implications for La Rioja Province.     

The Late Pleistocene distribution of vicuñas (Vicugna vicugna) and the “extinction” of the gracile llama (“Lama gracilis”): New molecular data

We present the results of a palaeogenetic analysis of two Late Pleistocene camelids originating in southern Chile. Our analysis of two mitochondrial DNA fragments (control region and cytochrome b gene) reveals that these specimens do not belong to an extinct taxon, but rather to extant vicuña (Vicugna vicugna), whose modern distribution is restricted to the extreme elevations of the Andes, more than 3500 km to the north of where these specimens originated. Our results also suggest fossil specimens from Patagonia that are currently assigned to the extinct taxon Lama gracilis, may actually belong to V. vicugna, implying a continuous distribution of the latter from the southern tip of South America to the Andes during the Final Pleistocene. The haplotypes of both specimens are not present in modern populations, suggesting a loss of genetic diversity concomitant with the contraction of the vicuña geographical distribution during the Final Pleistocene or early Holocene.     

The first water measurers from the Lower Cretaceous amber of Spain (Heteroptera,Hydrometridae, Heterocleptinae)

Two fossils belonging to a new genus and species of water measurer (Gerromorpha, Hydrometridae), Alavametra popovi Sánchez-García and Nel gen. n., sp. n., are described as first definitive record of the family in Lower Cretaceous (upper Albian) amber from the Utrillas Group (Peñacerrada I site) in Spain. Although several parts of the specimens are obscured due to preservation, a sufficient number of taxonomical characters are visible to consider adequate placement within Heterocleptinae, including the very long posterior pair of cephalic trichobothria inserted on tubercles and the preapical articulation between the first and second antennal segments. The new fossil taxon is included into a cladistic analysis with extinct and extant hydrometrids, and it results putatively basal among the subfamily Heterocleptinae, suggesting that this clade was already present 105 Ma ago.     

New abelisaurid remains from the Anacleto Formation (Upper Cretaceous), Patagonia,Argentina

New theropod remains with abelisaurid affinities from the Upper Cretaceous (Anacleto Formation, lower Campanian), NW Patagonia, Argentina, are here described. The specimen (MPCN-PV 69) consists of a partial premaxilla, fragmentary vertebrae, proximal portion of both humeri, distal portion of the pubis, and an incomplete pedal ungual. Characters linking with Abelisauridae are a premaxilla with a subquadrangular body, externally ornamented, and paradental plates with a striated surface; and humerus with bulbous proximal head, conical internal tuberosity, and reduced greater tubercle. The humerus is similar to those of Carnotaurus and Aucasaurus, due to the presence of a bulbous head and a discontinuity between the head and the internal tuberosity, but also differs from both taxa in the more distal location of the greater tubercle with respect to the internal tuberosity. Aucasaurus also comes from Anacleto Formation, but differences in the humeri suggest that MPCN-PV 69 is a different taxon. The phylogenetic analysis performed supports the affiliation to Abelisauridae, but fails to determinate a more precise relationship with others abelisaurids. However, a majority rule consensus of the analysis shows a position within Brachyrostra. Despite being fragmentary, MPCN-PV 69 probably represents a new abelisaurid from the Anacleto Formation, thus increasing the knowledge and diversity of Late Cretaceous South American abelisaurids.     

A review of Late Cretaceous fossil birds from Hungary

We review the previously described Late Cretaceous (Santonian) bird remains from the Csehbánya Formation in the Bakony Mountains of Hungary, augmenting initial work by Ősi ( 2008 ), and add a number of newly collected fossils. All together, the eight fossil specimens so far collected from this site are important to our understanding of avian evolution because they document a large range of taxon body sizes from at least one major lineage (Enantiornithes) and come from a critically undersampled time period in the Cretaceous. Globally, very little fossil bird material has been collected from the middle stages of the Late Cretaceous, the Coniacian and Santonian; most known taxa are either Early Cretaceous (ca. 120 Ma) in age or are from the terminal Campanian and Maastrichtian (ca. 70–65 Ma). Indeed, one of the Csehbánya Formation fossil birds is recognized as a new taxon of large enantiornithine, an avisaurid apparently similar in its largely unfused foot morphology to the Argentine Soroavisaurus and to the North American Avisaurus. The Central European records reviewed in this paper highlight the wide distribution of some Late Cretaceous fossil birds, particularly avisaurid enantiornithines, and lead us to a brief discussion of avian biogeography at the end of the Mesozoic. Copyright © 2010 John Wiley & Sons, Ltd.     

A new tyrannosaurine specimen (Theropoda: Tyrannosauroidea) with insect borings from the Upper Cretaceous Honglishan Formation of Northwestern China

A large theropod ilium was recently collected from the Upper Cretaceous Honglishan Formation in the Sangequan area of the northern Junggar Basin, Xinjiang, China, which represents the first officially reported dinosaur fossil from this formation. Several morphological features, including robust supracetabular ridge, reduced supracetabular crest, concave anterior margin of the pubic peduncle, ventral flange on the pubic peduncle, converging dorsal surface of the iliac blades, laterally visible cuppedicus shelf, and ventral flange on the posterior surface of pubic peduncle, suggest that this specimen can be referred to Tyrannosaurinae, and furthermore, a few differences between this specimen and other tyrannosaurines in particular the contemporary Asian tyrannosaurine Tarbosaurus suggest that IVPP V22757 may represent a new tyrannosaurine species. However, in the absence of extensive data that would make it possible to properly evaluate these differences, we refrain from naming a new taxon based on this specimen. Some insect borings are also identified in this specimen, and are referable to the ichnogenus Cubiculum, which is interpreted as the insect pupichnia. This new fossil documents the presence of a gigantic theropod in the Upper Cretaceous of Junggar Basin, adding new information on its poorly studied ecosystems.     

First record of a clupeomorph fish in the Neuquén Group (Portezuelo Formation), Upper Cretaceous of Patagonia, Argentina

A new genus and species of clupeomorph fish, Leufuichthys minimus, is described from the fluvial deposits of the Portezuelo Formation, Upper Cretaceous (Turonian-Coniacian) of the Neuquén Group, Patagonia, Argentina. It is a small-sized fish with an estimated body length up to 46 mm. Among other characters, the new species shows the following: abdominal scutes; abdomen moderately convex; anal fin elongate-based; three uroneurals; two epurals; caudal fin bearing very elongate rays; and cycloid scales. Leufuichthys minimus gen. et sp. nov. shows a greater similarity with Kwangoclupea dartevellei, a clupeomorph described from a marine Cenomanian deposit of the Democratic Republic of Congo (Africa), mainly due to the presence of an elongate-based anal fin, bearing more than 20 fin-rays, differing from it by the presence of a not hypertrophied abdomen. As far as known, L. minimus gen. et sp. nov. is the first clupeomorph described in the Upper Cretaceous of Patagonia and represents one fortuitous preservation of an articulated fish in fluvial deposits.     

Edentulous pterosaurs from the Cambridge Greensand (Cretaceous) of eastern England with a review of Ornithostoma Seeley, 1871

A re-examination of fossil material from the Late Cretaceous Cambridge Greensand Member (CGM) of the West Melbury Marly Chalk Formation revealed a number of new specimens of edentulous pterosaur jaw fragments previously identified as shark fin spines and fish jaws and accessioned under the epithet ‘cestraciontid finray’ and ‘jaws of fish’. These are now recognised as pterosaurian jaw tips and referred to Ornithostoma sedgwicki Seeley, 1891 and Azhdarchoidea indet. This material increases the diversity of edentulous pterosaurs from the CGM.The edentulous pterosaur Ornithostoma sedgwicki Seeley, 1891 from the Cretaceous Cambridge Greensand of eastern England is reviewed. The holotype specimen is confirmed as a fragment of a premaxilla/maxilla of a non-tapejarid azhdarchoid on account of the conspicuous curvature of the dorsal and occlusal margins posteriorly and the presence of small neural foramina on the lateral margins. Neural foramina are not seen on jaws of members of the Pteranodontia, a group to which O. sedgwicki was included previously. The referral of O. sedgwicki to Azhdarchoidea eliminates the single known Lower Cretaceous occurrence of Pteranodontidae, restricting the temporal range of this taxon to the Upper Cretaceous. Postcranial material referred to O. sedgwicki from the type horizon is regarded as indeterminate Pterosauria.     

The first crocodyliform from the Chubut Group (Chubut Province, Argentina) and its phylogenetic position within basal Mesoeucrocodylia

A new crocodyliform specimen is presented here found in the Cerro Castaño Member of the Cerro Barcino Formation (Chubut Group). The material consists of cranial and postcranial remains that represent a new taxon that has strong affinities with Peirosauridae, but also shares derived features present in Araripesuchus. The phylogenetic relationships of this new taxon were tested through a cladistic analysis depicting it as a member of the Peirosauridae. The inclusion of Barcinosuchus within this clade of basal mesoeucrocodylians is supported by the presence of hypapophyses up to the third or fourth dorsal vertebrae, anterolateral facing edge on postorbital, quadrate dorsal surface divided in two planes by a ridge; mandibular symphysis tapering anterirorly in ventral view, lateral surface of dentary convex anterior to mandibular fenestra, distal body of quadrate well developed, anteroposteriorly thin and lateromedially broad. The new specimen broadens the temporal and geographical distribution of Peirosauridae during the Cretaceous of Gondwana, representing the southern-most and the most ancient record of the group in Patagonia. The new material also provides insights on the postcranial anatomy of peirosaurids, a group that has been so far studied almost exclusively from cranial material.     

Prophaenognatha, a New Aclopinae Genus from the Yixian Formation, China and Its Phylogenetic Position Based on Morphological Characters (Coleoptera: Scarabaeidae)

Prophaenognatha robusta gen. et sp. nov. (Scarabaeoidea: Scarabaeidae: Aclopinae), the best-preserved aclopine fossil so far, is described and illustrated from the Upper Jurassic–Lower Cretaceous Yixian Formation of the Jehol Biota, western Liaoning Province, NE China. The key to extinct and extant genera of Aclopinae is given and the monophyly of extant and fossil Aclopinae lineages is supported by five character states. The new taxon provides evidence about the evolution of Scarabaeoidea with its phylogenetic position inferred based on 68 morphological characters.     

New materials of lithostrotian titanosaurs (Dinosauria: Sauropoda) from the Upper Cretaceous of central Patagonia

Continental Cretaceous deposits exposed in central Patagonia, Argentina, preserve a rich and important record of titanosaurian evolution that spans the entire Upper Cretaceous. Recently, a new lithostragraphic unit, the Lago Colhué Huapi Formation (Coniacian – Maastrichtian), was located in the Golfo San Jorge Basin. Here, we describe new titanosaurian sauropod material from that formation (UNPSJB-Pv 1051). The material consists of a partially articulated left hind limb and disarticulated but associated skeletal elements. They are confidently referred to Titanosauria, and within that clade, to Lithostrotia, probably occupying a derived position. We interpret the bone concentration as an assemblage of hydraulic origin deposited in a fluvial channel. This new material begins the incipient fossil record of the Lago Colhué Huapi Formation, thereby also increasing the titanosaurian diversity of the latest Cretaceous. Additionally, the new titanosaurian enhances evidence in support of strong faunal similarities among the Allen and Marilia formations, chronological equivalents in the Cretaceous of South America. Furthermore, the Lago Colhué Huapi specimen adds to the Late Cretaceous record of Titanosauria and augments our knowledge of central Patagonian terrestrial vertebrate assemblages during this interval.