Sympatry of two ankylosaurs (Hungarosaurus and cf. Struthiosaurus) in the Santonian of Hungary

A complete and well-preserved right ankylosaurian humerus from the Upper Cretaceous (Santonian) Csehbánya Formation of Iharkút, western Hungary is described here. Based on its osteological features and 21.5 cm adult length, the new specimen is markedly different from the slender humerus of Hungarosaurus, the previously known ankylosaur from the locality, and more similar to that of Struthiosaurus. Thus, the new Hungarian specimen is tentatively assigned here to cf. Struthiosaurus thereby dating back the first occurrence of this genus to the Santonian. The new fossil demonstrates the sympatric co-existence of two different nodosaurid ankylosaurs (a smaller, robust form with 2–2.5 m total body length and a larger, cursorial form with 4–4.5 m body length) in the Iharkút fauna. This also suggests that the pattern of the European ankylosaur diversity was more complex than previously thought.     

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.     

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.     

Notes on fossil remains from the Early Cretaceous Lohan Cura Formation, Neuquén Province, Argentina

In this contribution, we describe an association of fossil remains collected in the locality of Cerro de Los Leones, approximately 8 km southwest of the town of Picún Leufú, Neuquén Province, Argentina, from different levels of the Albian Cullín Grande Member, Lohan Cura Formation. The fossils include pelecypodan (Neocorbicula dinosauriorum, Neocorbicula pehuenchensis, and genus indet.) and gastropodan (Physa wichmanni) molluscs, a mesoeucrocodylian, possible pterosaurs, theropods, and sauropods. The sedimentary environment at Cerro de los Leones is characterized by fluvial deposits corresponding to a sandy meandering fluvial system. The most fossiliferous levels (located in the lower part of the exposed sequence) were developed in a distal flood plain and flood basin areas. Sedimentological and paleopedological features suggest semi-arid conditions during the deposition of the Cullín Grande Member. Despite the fragmentary nature of the specimens described here, the new information provided from the Albian assemblages of the Lohan Cura Formation at the Cerro de Los Leones locality provides a broader panorama of the composition of terrestrial faunas during the Early Cretaceous in Patagonia.     

A new titanosaur (Dinosauria,Sauropoda) from the Upper Cretaceous of Lo Hueco (Cuenca,Spain)

The upper Campanian-lower Maastrichtian site of Lo Hueco (Cuenca, Spain) has provided a set of well-preserved partial skeletons in anatomical connection or with a low dispersion of their skeletal elements. One partial skeleton is herein described and a new titanosaurian sauropod is established, Lohuecotitan pandafilandi. This titanosaur is diagnosed by eight autapomorphic features: dorsally and ventrally widened or bifurcated posterior centrodiapophyseal lamina in anterior and middle dorsal vertebrae; short postspinal lamina with a transversely expanded distal end represented by smooth scars in the dorsal vertebrae; anteriormost caudals with the medial spinoprezygapophyseal and medial spinopostzygapophyseal laminae ventrally connected with the prespinal and postspinal laminae, respectively; anterior caudal neural spines with a dorsal projection of the prespinal and postspinal laminae; anterior caudal neural spines bears a “greek-cross”-like cross-section; middle caudal centra having two round and rough structures in the dorsal edge of the posterior articulation, which extends to the dorsal surface of the centrum; the articular ends of the rami of the haemal arches are divided in two articular surfaces; and tuberosity between the anterior and the lateral trochanter of the fibula. The herein performed phylogenetic analysis considered L. pandafilandi as a member of Lithostrotia more derived than Malawisaurus. The known palaeodiversity of the Late Cretaceous Ibero-Armorican titanosaurs is increasing, and further analyses focused on this group will be necessary to better understand the evolutionary history of European titanosaurs and to clarify their relationships within Titanosauria.     

First ornithopod remains from the Bajo de la Carpa Formation (Santonian,Upper Cretaceous), northern Patagonia,Argentina

In the last decades, the Argentinian ornithopod record has been increased with new and diverse bone remains found along all the Upper Cretaceous. Most of them are very incomplete and represent taxa of different size. As result, the studies about the palaeobiodiversity of the Ornithopoda clade in South America are complex. In this paper, new postcranial remains of an indeterminate medium-sized ornithopod from the Santonian Bajo de la Carpa Formation (Rincón de los Sauces, Neuquén province) are presented. They present diagnostic features of the Ornithopoda clade, and several characters that relate them with other Argentinian ornithopods, especially with the medium-sized members of the Elasmaria clade sensu Calvo et al. (2007) (Macrogryphosaurus and Talenkauen). The postcranial material allows to identificate at least three different ontogenetic stages: adult, subadult more immature and subadult. These bones are the first record of Ornithopoda for the Bajo de la Carpa Formation and one of the very scarce Santonian records of this clade in South America. The diversity of the Late Cretaceous South American ornithopods presents two clear distributions: the Cenomanian-Santonian was characterized by small and medium euiguanodonts and elasmarian; and the Campanian–Maastrichtian by the medium sized elasmarian and large sized hadrosaurids.     

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.     

The controversial Les Labadous eggshells: A new and peculiar dromaeosaurid (Dinosauria: Theropoda) ootype from the Upper Cretaceous of Europe

The fossil record of small-sized theropods in south-western Europe is scarce and fragmentary. In absence of more direct evidence, the oological record arises as an alternative source of information to infer biodiversity. In this regard, the controversial dinosaur eggshells from the Les Labadous locality in southern France are herein re-evaluated in the light of new material and current parataxonomic information. New data and analyses reveal a new eggshell type, Montanoolithus labadousensis oosp. nov., characterized by a unique combination of characters. Additionally, the phylogenetic analysis reinforces the placement of the new oospecies as the sister taxon of the North American Montanoolithus strongorum, refusing the previous attribution of the Les Labadous remains to Elongatoolithidae. The combination of microstructural and phylogenetic results suggests that Montanoolithus labadousensis was likely produced by a dromaeosaurine dromaeosaurid, a taxonomic attribution that is consistent with the biodiversity of small-sized theropods at the latest Cretaceous of south-western Europe.     

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.     

豫西栾川地区晚白垩世秋扒组——新的驰龙类化石(英文)

根据以下特征,建立驰龙类恐龙一新属新种——河南栾川盗龙(Luanchuanraptor henanensisgen.etsp.nov):牙齿后缘每5mm有24个小锯齿;前部背椎椎体具有小的椎体侧孔;后部脉弧的前后突短且为三分支;最后部的尾椎体的椎体前突拉长且包裹与其相邻的前部椎体,两椎体前突之间夹有低的刀片状的神经棘;乌喙骨的内面强烈凹陷;肱骨骨干直,肱骨三角脊几乎占肱骨长度的前部一半。河南栾川盗龙代表除在亚洲戈壁地区及中国东北地区之外发现的第一件驰龙类化石标本。     

豫西栾川地区晚白垩世秋扒组一新的驰龙类化石

根据以下特征，建立驰龙类恐龙一新属新种——河南栾川盗龙（Luanchuanraptor henartens／sgen．et sp．nov）：牙齿后缘每5mm有24个小锯齿；前部背椎椎体具有小的椎体侧孔；后部脉弧的前后突短且为三分支；最后部的尾椎体的椎体前突拉长且包裹与其相邻的前部椎体，两椎体前突之间夹有低的刀片状的神经棘；乌喙骨的内面强烈凹陷；肱骨骨干直，肱骨三角脊几乎占肱骨长度的前部一半。河南栾川盗龙代表除在亚洲戈壁地区及中国东北地区之外发现的第一件驰龙类化石标本。     

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.     

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.     

Reassessment of coelurosaurian (Dinosauria,Theropoda) remains from the Upper Cretaceous Wangshi Group of Shandong Province,China

Non-avian theropods are well represented in the Cretaceous of Asia. However, the first theropod remains ever reported from China, from the Wangshi Group of Shandong Province, have not been reassessed since they were described by Carl Wiman in 1929, despite the great strides that have been made in theropod phylogenetics in the last eight decades. The remains redescribed herein consist of four vertebrae (two cervicals, one dorsal and one caudal) from one site, and a caudal vertebra and ungual phalanx from another. The vertebrae from the first site are assigned to tyrannosauroids and ornithomimosaurs, as is the caudal vertebra from the second site, whereas the manual ungual also bears resemblances to those of non-maniraptoran coelurosaurs (specifically compsognathids and ornithomimosaurs). Theropods remain relatively rare in the Wangshi Group; nevertheless, the remains redescribed herein highlight the potential for the discovery of more theropod remains in future excavations.     

Recognition of massive Upper Cretaceous carbonate bodies as olistoliths using rudist bivalves as internal bedding indicators (Campanian Merfeg Formation,Central Tunisia)

The Merfeg Formation (upper Campanian) of Central Tunisia crops out around the southwestern periclinal termination of Jebel el Kébar, near Sidi Bouzid. At its base is a massively bedded unit of locally dolomitized, sparsely fossiliferous micritic to microbioclastic limestone that contains several discrete, plurimetric mound-like bodies (lithosomes) of micritic limestone containing locally abundant rudists and corals. The lithosomes are separated laterally from one another by megabreccias and conglomerates containing clasts of similar lithology and are overlain, with sharp contact, by onlapping argillaceous pelagic limestones, within which are intercalated at least two more, somewhat thinner rudist/coral limestone units. This complex of facies is laterally equivalent to thicker, deep platform limestones of the Abiod Formation to the north and east, and to restricted carbonate platform facies of the Berda Formation to the south and west. The lithosomes have previously been interpreted as in situ downslope mudmounds that became capped by rudist and coral formations, cemented, and then surrounded by erosively emplaced debris flows. However, our detailed studies of rudist orientations imply variable and in some cases relatively high angles of bedding within the lithosomes with respect to the regional dip of the host strata. Such steep inclinations of internal bedding are unlikely to have been primary. Accordingly, we propose an alternative interpretation that the lithosomes were platform-derived olistoliths, emplaced along with the associated debris flow deposits. Micritic beds, neighbouring the olistoliths are of variable thickness and contain rare large inoceramids and randomly oriented rudists, as well as locally developed microbioclastic beds with planar and small-scale swaley cross stratification. These micritic and microbioclastic beds are, by contrast, interpreted as primary (i.e., non-olistostromal) slope deposits. Whether the proposed catastrophic collapses of the original platform margin were induced by sea-level fall or seismically triggered (or a combination of the two) remains uncertain.     

Late Albian (Early Cretaceous) ammonites from the Provincial Formation of central Cuba

A late Albian ammonite assemblage from the Provincial Formation of Villa Clara Province, Cuba is described. The Provincial Formation is a lithostratigraphic unit of Albian-Cenomanian age extensively exposed in central Cuba and formed within a part of the Caribbean Tethys that was volcanic during the Cretaceous. The formation is mainly composed of calcareous, terrigenous marine, and volcano-sedimentary deposits characterized by a series of micritic limestones intercalated with marls, sandstones, calcareous conglomerates, ash, and tuffaceous material. A rich assemblage of ammonites recovered from the calcareous biomicrites and marls is of late Albian (Stoliczkaia dispar Zone, Mortoniceras rostratum Subzone) age. The ammonite fauna shows a strong Tethyan affinity, and only a single hoplitid ammonite species was recorded. Although scarce, the first Cuban report of this and other boreal ammonite species now allows precise correlations to be made between Cuba and Albian sediments elsewhere in the world.     

First postcranial remains of Multituberculata (Allotheria,Mammalia) from Gondwana

Multituberculates (Allotheria) are generally regarded as the evolutionarily most successful and longest-lived (Middle Jurassic to late Eocene) clade of Mesozoic and early Paleogene mammals. Despite this “reputation” and the fact that the group is particularly well represented in both taxonomic diversity and relative abundance on Laurasian landmasses during the Cretaceous and Paleocene, multituberculates are exceedingly poorly represented on the southern supercontinent Gondwana. Previous records on Gondwanan landmasses have been based on fragmentary dental remains and all except the three most recently published (each represented by a single isolated tooth or fragment of tooth) have been disputed and allocated to either Haramiyida or Gondwanatheria. Furthermore, several previous records, disputed or not, are based on fragmentary dental remains of a type (plagiaulacoid) that has evolved independently several times in mammalian evolution.Here we place on record a multituberculate femur from the Upper Cretaceous (Maastrichtian) Maevarano Formation of the Mahajanga Basin, Madagascar. This specimen, although fragmentary as well, exhibits a number of features common to all multituberculate femora: neck cylindrical in cross section and set apart from shaft; greater trochanter prominent, extending proximally beyond head, inclined dorsally, and separated from neck by deep incisure; lesser trochanter prominent and protruding ventrally; posttrochanteric fossa present on ventral aspect, lateral to lesser trochanter; subtrochanteric tubercle present on dorsal aspect, distal to incisure between greater trochanter and neck; diaphysis straight, elliptical in cross section (slightly compressed dorsoventrally); and third trochanter absent. Three of these features (prominent, ventrally placed lesser trochanter; presence of posttrochanteric fossa; presence of subtrochanteric tubercle) are regarded as autapomorphies of Multituberculata. This specimen therefore not only independently and conclusively confirms the presence of the clade on Madagascar—previously based on a small molar fragment—but on the entire supercontinent as well.     

New fossil mesosciophilids (Diptera: Nematocera) from the Yixian Formation (Lower Cretaceous) of western Liaoning,China

A new genus with two new species, Orentalphila gravia gen. et sp. nov. and O. caloa sp. nov., and three new species in two known genera, Sinosciophila angustia sp. nov., Sinosciophila seboa sp. nov. and Similsciophila undulata sp. nov., are described from the Lower Cretaceous of Yixian Formation at Huangbanjigou, Beipiao City, western Liaoning, China. These new taxa, representing the first records of mesosciophilids from this locality, provide morphological information and diversity for Mesosciophilidae in the Early Cretaceous. An updated key to known genera of Mesosciophilidae is provided. In addition, according the generic diagnosis revised by Zhang, 2007, Mesoplecia antiqua Hao and Ren, 2009 should be transferred to Mesosciophila Rohdendorf, 1946.     

Ornithomimidae (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan

The stratigraphically oldest remains of ornithomimid theropod dinosaurs are known from the Cenomanian Khodzhakul Formation and the Turonian Bissekty Formation of Uzbekistan. The ornithomimid from the Bissekty Formation is documented by more than 800 isolated bones that represent much of the skeleton. It shows at least three unambiguous synapomorphies of Ornithomimidae: length of anterior cervical centra three to five times greater than transverse width; low and rounded fibular crest of tibia; metatarsal III pinched between metatarsals II and IV; and proximal end of metatarsal III not visible in anterior view. Phylogenetic analysis, based on a dataset with 568 morphological characters and including all known ornithomimosaurian taxa, places the ornithomimid from the Bissekty Formation near the base of the ornithomimid radiation, between Archaeornithomimus asiaticus and Sinornithomimus dongi.     

北美西部内陆海盆Niobrara组的钙质超微化石及其环境意义——Ⅱ：古环境研究

沉积于北美西部白垩纪内陆海盆的Niobrara组，是一套以白垩，白垩质灰岩，以及钙质页岩为主的海相沉积，其中自垩的主要组分为颗石藻及其它钙质超微化石。通过对Kansas州西部和South Dakota州东部四条剖面中Niobrara组的岩性，沉积物厚度，及钙质超微化石的研究，证明北美西部内陆海盆的沉积环境及水体温度在Niobrara组沉积时期因地而异。在Niobrara组沉积初期，海盆内的水体温度为北冷南暖，其后，至少在位于Kansas西部海域的海水温度呈与时逐渐降低趋势。钙质超微化石有关属种在盆地内的分布差异，进一步说明海水变冷的直接原因源自北极海域水体的缓慢南浸。