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.     

Osteology of Viavenator exxoni (Abelisauridae; Furileusauria) from the Bajo de la Carpa Formation,NW Patagonia,Argentina

The complete osteology of the abelisaurid Viavenator exxoni, from the Bajo de la Carpa Formation (Santonian, Upper Cretaceous) is described. Viavenator is characterized by a series of autapomorphies, including: transversely compressed parietal depressions on both sides of the supraoccipital crest; ventral edges of the paraoccipital processes located above the level of the dorsal edge of the occipital condyle; basioccipital-opisthotic complex about two and a half times the width and almost twice the height of the occipital condyle, in posterior view; well-developed crest below the occipital condyle, diverging ventrally and defining the subcondylar recess; deeply excavated and sub-circular basisphenoidal recess, with its major axis transversely oriented; basipterygoid processes horizontally placed with respect to the cranial roof and located slightly dorsally to the basal tubera; mid and posterior cervical centra with slightly convex lateral and ventral surfaces; hyposphene-hypanthrum articular complex present from dorsal 2 onward; presence of an interspinous accessory articular system in middle and posterior dorsal vertebrae; presence of a pair of pneumatic foramina within the prespinal fossa in anterior caudal vertebrae; distal end of the scapular blade posteriorly curved. Particularly, Viavenator shows plesiomorphic cranial characters, i.e. flat frontals lacking domes or horns, combined with derived postcranial characters, e.g. the interspinous accessory joint system of dorsal vertebrae. This combination between plesiomorphic and derived traits suggests that Viavenator is a transitional form, which is an idea supported by its intermediate stratigraphic and phylogenetic placement between the basal and older (e.g. Skorpiovenator, Ilokelesia) and derived and younger members of the clade (e.g. Aucasaurus, Carnotaurus).     

Vertebrate remains from the Upper Cretaceous (Santonian) Ajka Coal Formation,western Hungary

Vertebrate remains from the Upper Cretaceous (Santonian) Ajka Coal Formation (Bakony Mountains, western Hungary) are described. Macro- and microfossils collected from two boreholes and from isolated chunks of sediment/matrix dumped on spoil heaps of the Jókai Mine represent pycnodontiform and lepisosteiform fishes, bothremydid turtles, the mosasauroid Pannoniasaurus inexpectatus, the crocodyliforms cf. Theriosuchus, Iharkutosuchus makadii and cf. Allodaposuchus, as well as ankylosaurian and theropod dinosaurs. This unit was deposited in a swampy lacustrine environment, in contrast with the neighbouring and contemporaneous floodplain deposit of the vertebrate-bearing Csehbánya Formation at Iharkút. Despite significant environmental differences, the faunal composition of the Ajka Coal Formation assemblage completely overlaps with that of the Csehbánya Formation, suggesting the occurrence of the same semi-aquatic and terrestrial species in both settings. The ankylosaurian remains further strengthen the previous view that ankylosaurus preferred wetland habitats such as fluvial systems and coastal regions.     

Small-sized abelisaurid (Theropoda: Ceratosauria) remains from the Upper Cretaceous of northwest Patagonia,Argentina

A new abelisaurid theropod from the Cenomanian of Candeleros Formation (Neuquén Province, NW Patagonia, Argentina) is described. It includes an isolated frontal, partial pelvis, two fused sacral centra, femoral shaft, rib fragments, partial metatarsal II and shaft of metatarsal III. A paleohistological analysis shows that MMCh-PV 69 was somatically mature at time of death. The prevalence of relatively ordered intrinsic fibers in the primary bone indicates a relatively lower growth rate of MMCh-PV 69 compared with other abelisauroids, such as Aucasaurus garridoi. Phylogenetic analysis found MMCh-PV 69 as a basal abelisaurid, sister group of the node formed by Majungasaurinae and Brachyrostra. We estimated the body mass of MMCh-PV 69 in 240 kg, which makes it one of the smallest abelisaurids recorded. This new form adds to the extensive theropod fauna of the Candeleros Formation, probably the most diverse association of meat-eating dinosaurs recorded in a Cretaceous Formation from South America.     

Tectonic control of accommodation space and sediment supply within the Mata Amarilla Formation (lower Upper Cretaceous) Patagonia,Argentina

The Mata Amarilla Formation dates from the early Upper Cretaceous and was deposited during a transition in tectonic regime from the extensional Rocas Verdes Basin to the Austral Foreland Basin. Detailed sedimentological logs and architectural parameters were used to define 13 facies associations. The distribution of facies associations and associated variations in fluvial architecture have enabled large‐scale changes in accommodation space/sediment supply ratios (A/S ratio) to be defined for the three component sections of the Mata Amarilla Formation. The lower and upper sections are characterized by a high A/S ratio, whereas the middle section corresponds to a low A/S ratio. In the western part of the study area, small‐scale variations in the A/S ratio were recognized in the middle section. The strong west to east trend in evolution of the fluvial systems coincides with the direction of propagation of the Patagonian fold and thrust belt, which is located to the west of the study area. Intervals of high A/S ratio (i.e. lower and upper sections) are interpreted to have developed during periods of increased loading by the fold and thrust belt caused by tectonic uplift. In contrast, intervals of low A/S ratio (i.e. middle section) were developed during periods of tectonic quiescence. This article suggests that the large‐scale variations in A/S ratios are related to different rates of migration and growth of the Patagonian fold and thrust belt, whereas the small‐scale variation occurred in response to specific periods of thrusting and folding in the Patagonian fold and thrust belt (i.e. local loads). This field example of the effects of different scales of variation in A/S ratios across the Austral Foreland Basin could be used to recognize similar tectonically forced variations in stratigraphic architecture in other foreland basins throughout the world, as well as to understand the response of fluvial systems to such changes.     

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.     

New species Dalembia jiayinensis (Magnoliopsida) from the Upper Cretaceous Yong'ancun Formation,Heilongjiang, northern China

Leaves of Dalembia jiayinensis sp. nov. are reported from the Santonian Yong'ancun Formation exposed along the Heilongjiang (Amur) River in Jiayin County, Northeast China. The leaves are compound, odd-pinnate, consisting of five leaflets. Leaflets are pinnately-lobed, trilobate or unlobed, elliptic, oblong, elliptic-rhomboid or ovate in outline, with blunt or rounded apex and the leaflet base is variable being cuneate, slightly decurrent, truncate or slightly cordate and sometimes asymmetric. Margin is entire or lobed. Lobes are short, simple, decreasing upwards, with wide and rounded sinuses. Apices of the lobes are usually rounded, sometimes notched. Venation is pinnate or palmately-pinnate, craspedodromous or semi-craspedodromous. The new species D. jiayinensis extends both the geographical and stratigraphic ranges of the genus Dalembia. This is the first occurrence of Dalembia in China and the most southern and youngest (Santonian) occurrence of this genus.     

Linderochelys rinconensis (Testudines: Pan-Chelidae) from the Upper Cretaceous of northern Patagonia: New insights from shell bone histology,morphology and diagenetic implications

The panchelid turtle Linderochelys rinconensis de la Fuente, Calvo and González Riga, 2007 was described on the basis of several postcranial elements from the Rio Neuquén Subgroup of Loma del Lindero site, near Rincon de los Sauces (Neuquén, Argentina). New preparation of the type material revealed new parts of the animal, including cervical vertebrae, parts of the carapace and a right tibia. The study of the newly recovered material herein improves our knowledge of the macroanatomy of this taxon. The first costal shows an axillary buttress similar to Australasian chelids and the cervical vertebrae morphology suggests a short-necked chelid condition. The re-examination of the type material of Linderochelys and the discovery of new material allowed new studies on microanatomy, osteohistology and diagenesis. Osteohistology suggests some similarities with Patagonian Cretaceous and Palaeocene Yaminuechelys, based on the presence of parallel-fibred bone in the external cortex and a much thinner internal cortex. Paleoecological studies on the proportions of forelimbs and microanatomy point towards aquatic or semi-aquatic habits, closely related to the condition seen in some extant South American chelids, such as Phrynops hilarii, Phrynops geoffroanus, and Mesoclemmys gibba. The fossil-diagenetic processes inferred substitution, fracturing, plastic deformation and permineralization events. Combined analyses through scanning electron microscopy with energy-dispersive X-ray spectrometry and petrographic studies revealed the substitution of hydroxyapatite by francolite. Permineralization stages include infilling of vascular canals, trabeculae and fractures with iron oxides and carbonate minerals during their burial history.     

Lower Cretaceous provenance in the northern Austral basin of Patagonia from sedimentary petrography

The northern Austral basin from Patagonia is characterized by an Early Cretaceous (Barremian–Albian) coarse-grained regressive sequence. These littoral to continental deposits conform a 150 km long basin cropping out along the Southern Patagonian Andes between 47 and 48°S. The basin fill consist of basal deltaic sandstones with interbedded shales and limestones from the Río Belgrano Formation, topped by up to 350 m of fluvial conglomerates and reworked tuffs of the Río Tarde and Kachaike formations. This continental depocenter represent a major geodynamic and paleoenviromental change from the underling marine Río Mayer Formation. In this study we analyze the tectonic setting and provenance during deposition of the coarse-grained sequence using sedimentary petrography of 37 thin sections in four stratigraphic profiles covering the northern basin. Our dataset indicates mainly a recycled orogenic sandstones provenance, in agreement with potential surrounding basement sources.     

An anilioid snake from the Upper Cretaceous of northern Patagonia

We report the first record of anilioids from southern South America, a region where they do not live today. The fossils come from the Allen Formation (Late Campanian–Early Maastrichtian) at Bajo Trapalcó and Bajo de Santa Rosa localities, Río Negro province, Argentina. The remains consist of several vertebrae, most of which belong to the mid–posterior precloacal region of the column. Comparisons to other extant and extinct anilioid taxa indicate that these remains represent a new taxon, Australophis anilioides gen. et sp. nov. Australophis is morphologically closer to Palaeocene Hoffstetterella from Brazil and extant South American Anilius than to any other snake. These taxa might be closely related, in which case they would represent a lineage distinct from that including Cylindrophis and uropeltids. The two lineages must have diverged by the Late Cretaceous.     

Revised identification of a reported Iguanodon-grade ornithopod tooth from the Scollard Formation, Alberta, Canada

An isolated tooth from the late Maastrichtian strata of the Scollard Formation of Alberta, Canada, has been identified as the only record of a non-hadrosaurid hadrosauriform from the Maastrichtian of North America. Here, we demonstrate that this identification is in error and that the tooth pertains to an indeterminate ceratopsid. In addition, we provide full documentation of the published collection of Scollard Formation fossils from which the tooth was originally derived.     

Dinosaur eggshells from the Santonian Milk River Formation of Alberta,Canada

The North American fossil record of dinosaur eggshells for the Cretaceous is primarily restricted to formations of the middle (Albian–Cenomanian) and uppermost (Campanian–Maastrichtian) stages, with a large gap in the record for intermediate stages. Here we describe a dinosaur eggshell assemblage from a formation that represents an intermediate and poorly fossiliferous stage of the Upper Cretaceous, the Santonian Milk River Formation of southern Alberta, Canada. The Milk River eggshell assemblage contains five eggshell taxa: Continuoolithus, Porituberoolithus, Prismatoolithus, Spheroolithus, and Triprismatoolithus. These ootaxa are most similar to those reported from younger Campanian–Maastrichtian formations of the northern Western Interior than they are to ootaxa reported from older middle Cretaceous formations (i.e., predominantly Macroelongatoolithus). Characteristics of the Milk River ootaxa indicate that they are ascribable to at least one ornithopod and four small theropod species. The taxonomic affinity of the eggshell assemblage is consistent with the dinosaur fauna known based on isolated teeth and fragmentary skeletal remains from the formation, although most ornithischians and large theropods are not represented by eggshell. Relative to the Milk River Formation eggshell, similar oospecies occurring in younger Cretaceous deposits tend to be somewhat thicker, which may reflect an increase in body size of various dinosaur lineages during the Late Cretaceous.     

Dinosaur tracks in the Kokorkom Desert,Candeleros Formation (Cenomanian,Upper Cretaceous), Patagonia Argentina: Implications for deformation structures in dune fields

The Kokorkom desert extended over an area of 826 km² in the central-west sector of Neuquén and Río Negro provinces along the area of the backbulge basin within the Andean Foreland basin in the Neuquén Basin. Its deposits constitute the middle-upper section of the Candeleros Formation (Cenomanian) and reach approximately 130 m thick. Tracks are found on wet and dry interdune and within draa slipface deposits. They constitute biogenic deformation structures characterized by folded-up and/or brecciated sandstone levels formed under dry and/or wet substrate conditions with passive filling. The degree of preservation varies, but the identification of digit impression suggests that they were produced by theropods or iguanodontians.     

Calcareous nannofossils from the Santa Marta Formation (Upper Cretaceous), northern James Ross Island,Antarctic Peninsula

This study reports on the most stratigraphically extensive nannofloras yet recovered from the Lachman Crags Member of the Santa Marta Formation, James Ross Island, Antarctic Peninsula. The productive samples are dated as early Campanian. These ages are in accord with those provided by ammonites, foraminifera, ostracods and radiolarians from the same locality. The consistent and relatively abundant presence of Gephyrobiscutum diabolum throughout the productive part of the section, a species that has previously only been documented from the Falkland Plateau, extends its geographic distribution to higher latitudes, at least to the Antarctic Peninsula area.     

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.     

New pterosaurian remains from the Late Cretaceous (Santonian) of Hungary (Iharkút, Csehbánya Formation)

The Late Cretaceous (Santonian) continental vertebrate locality of Iharkút, western Hungary has provided numerous azhdarchid pterosaur remains including the recently described, Bakonydraco galaczi. Since the first report of these fossils, additional remains have been discovered that improve considerably our knowledge of some aspects of the anatomy of azhdarchid pterosaurs. New cranial material described here indicates an edentulous, non-crested premaxillary rostrum in Bakonydraco similar to that of Quetzalcoatlus and reveals that this rostrum was considerably thinner and more lightly built than the relatively massive, pointed mandibular symphysis. In addition, the contact surface of the upper and lower jaws of Bakonydraco at least in the symphyseal region was more similar to Tapejara wellnhoferi than to other azdarchids: the contact is irregular and the premaxilla does not fit closely the rostral part of the mandible. Among the postcranial material the atlas-axis complex possessing lateral pneumatic foramina is of great importance because it further supports the notion of interspecific variability of vertebral pneumaticity among pterosaurs.     

Down‐slope facies variability within deep‐water channel systems: Insights from the Upper Cretaceous Cerro Toro Formation,southern Patagonia

In southern Patagonia, outcrops of the Upper Cretaceous Cerro Toro Formation preserve a >150 km long deep‐water axial channel belt in the Magallanes–Austral Basin, providing a unique opportunity to investigate longitudinal variations in the depositional characteristics of a deep‐water channel system. This study documents sedimentological, stratigraphical and geochronological data from the Cerro Toro Formation in the Argentine sector of the basin. New results are integrated with previous work from the Chilean basin sector to conduct a basin‐scale comparison of the timing of deposition, provenance and lithofacies proportions. The Cerro Toro channel belt includes a nearly 1000 m thick section characterized by high‐density turbidites and mass‐wasting deposits. Two ash beds from the base of the section yield U–Pb zircon ages of 90·4 ± 2 Ma and 88·0 ± 3 Ma, indicating similar initiation ages as documented in the Chilean sector. The U–Pb detrital zircon age spectra from samples in the study area reveal similar provenance trends to samples from the Chilean basin sector, with peak age populations at 310 to 260 Ma, 160 to 135 Ma and 110 to 82 Ma. The maximum depositional age of the channel belt in the Argentine sector is 87·8 ± 1·5 Ma and all new geochronology data corroborate an 86 to 80 Ma depositional age for the main Cerro Toro channel belt. Statistical analyses of 7370 beds from nearly 8000 m of new and previously published stratigraphic sections along the entire outcrop belt suggest progressive variations in the down‐system proportion of lithofacies. In the up‐slope region, lithofacies representing mass wasting processes (for example, debris‐flow and mass‐transport deposits) account for ca 29% of the stratigraphic thickness, as opposed to 5% in the down‐slope region of the channel belt, where turbidity current deposits are more prevalent. The proportion of beds >1 m thick also decreases systematically down slope, particularly for conglomeratic turbidite deposits. This work highlights that: (i) the proportion of thick beds and distribution of lithofacies are key down‐system changes in the stratigraphic fill of this deep‐water channel belt; (ii) detrital zircon trends suggest a relatively well‐mixed longitudinal depositional system; and (iii) geochronology of the main Cerro Toro outcrop belt supports but does not necessitate the model of a single, roughly age‐equivalent, channel system. This study has implications for understanding the downslope variability in depositional processes, stratigraphic architecture and reservoir quality of submarine channel systems.     

The first record of a bryophyte in Upper Cretaceous amber from Taimyr,northern Siberia: Taimyrobryum martynoviorum gen. et sp. nov. (Bryopsida)

Cretaceous bryophyte fossils are known only from a few localities and only a few are known in ultrastructural fidelity. We describe a fossil moss gametophyte from Upper Cretaceous Taimyr amber and place it in the fossil genus Taimyrobryum martynoviorum gen. et sp. nov. (Bryopsida). Its unbranched, somewhat zig-zagged stems with distantly spaced, spreading, narrowly lanceolate leaves and the unbranched costa match the gross morphology of Dicranidae; however, the uniformly prosenchymatous leaf cells contradict an affiliation to this subclass. The Yantardakh in the Taimyr Peninsula is the fourth and northernmost Cretaceous amber locality yielding inclusions of bryophytes.     

Multiple paleopathologies in the dinosaur Bonitasaura salgadoi (Sauropoda: Titanosauria) from the Upper Cretaceous of Patagonia,Argentina

Bones in the fossil record sometimes exhibit unusual structures that can be attributed to pathologies, taphonomic alterations, or morphological variation. The goal of this work is to describe three bone abnormalities present in the type specimen of Bonitasaura salgadoi from the Cretaceous of north Patagonia, Argentina. The studied material corresponds to a left femur, a left metatarsal III, and a right prezygapophysis of a mid-caudal vertebra. Macroscopic and/or histological examinations were conducted, and a discussion concerning the origin of each abnormality is provided. The results of this study suggest that the anomalous structures are pathologies. In the femur, an osteoblastic tumor was identified by the presence of a large outgrowth of ovoid appearance with a spiculated microstructural pattern. The metatarsal III shows an enthesophyte (or bone spur) based on its location, shape and growth in parallel to the long axis of the bone element. Finally, the abnormal tissue observed in the prezygapophysis of the caudal vertebra was determined to be an infection by the presence of reactive new bone associated with a local widening of the subperiosteal margin and a sinus of drainage. This is the first report of multiple pathologies in a single specimen of a titanosaur, and it provides new insights about paleopathologies in sauropod dinosaurs.