Paleocene resurrection of a crocodylomorph taxon: Biotic crises,climatic and sea level fluctuations

Postcranial material of a crocodylomorph from the Danian of Oulad Abdoun Basin (Morocco) is described. Several characters, in particular the shape of its dorsal osteoderms, allows its attribution to Pholidosauridae. Up till now, the latest known pholidosaurid was Terminonaris, from the Early Turonian of North America, so, the Moroccan pholidosaurid extends the stratigraphic range of the group to more than 20 million years younger, and shows that the pholidosaurids survived the K-Pg crisis. The reevaluation and the phylogenetic analysis of Dakotasuchus kingi, Woodbinesuchus byersmauricei, and Sabinosuchus coahuilensis, previously considered as goniopholidids and dyrosaurid, respectively, revealed rather that they are pholidosaurids. This analysis also suggests that at least two independent pholidosaurid lineages reached the Maastrichtian, among which one crossed the K-Pg boundary. This study proposes the first analysis of tethysuchian diversity from the Late Jurassic to the Early Paleogene. Two diversity peaks are observed during the Oxfordian and Cenomanian, two stages of high paleotemperatures evaluated with the δ¹⁸O. The tethysuchian diversity strongly decreases after the Cenomanian, a decline that may be correlated with the “Ocean Anoxic Event” (OAE 2), which caused the strong marine faunal turnover during the Cenomanian-Turonian time interval. The large Turonian-Coniacian oceanic regression could also be a factor in the tethysuchian decline. After that, tethysuchian diversity remains low until the Maastrichtian-lower Paleocene marine tethysuchian dyrosaurid diversification, correlated with the Late Cretaceous-Paleocene transgression. At least one pholidosaurid lineage crossed the K-Pg boundary, confirming the weak impact of this crisis on the crocodylomorphs. Pholidosaurids seem to go extinct at the beginning of the Paleocene, possibly due to the strong radiation of dyrosaurids in marine environments and crocodylians in fresh-water during this time period.     

Middle Eocene vertebrates from the sabkha of Gueran,Atlantic coastal basin,Saharan Morocco,and their peri-African correlations

The sabkha of Gueran in the Southwest Moroccan Sahara has yielded a rich and diverse fauna of late middle Eocene vertebrates that include the world's richest Bartonian age archaeocete assemblage. Archeocete remains were described previously and here we report on the rest of the vertebrate fauna. The Gueran fauna includes abundant chondrichthyan species belonging to Lamniformes, Carcharhiniformes and Rhinopristiformes, and actinopterygian assemblage consisting of Cylindracanthus, of a siluriform, and of Perciformes. Turtles are represented by at least two marine taxa referred to as Cheloniidae and Dermochelyidae. Crocodylian remains belong to at least two longirostrine species, including gavialoid remains. Snakes are represented by Pterosphenus cf. schweinfurthi (Palaeophiidae). Seabirds are represented by a pseudo-toothed bird (Pelagornithidae). The avian fossil belonged to a gigantic soaring bird and constitutes the earliest occurrence of the genus Pelagornis. The presence of proboscideans is attested by dental fragments. This fossil assemblage from Gueran shows affinities with those of the Eocene beds of Egypt and Libya. The numerous shared taxa support a close biogeographical connection between faunas from southeastern and southwestern coasts of the Mediterranean Sea.     

Lower Cretaceous Mesoeucrocodylians from Scandinavia (Denmark and Sweden)

The crocodyliform faunas of the lowermost Cretaceous Rabekke and Jydegård Formations on the Baltic island of Bornholm, Denmark, and the Annero Formation of Skåne, southernmost Sweden, are represented by isolated teeth, osteoderms, and vertebrae. The rich Berriasian assemblage of the Rabekke Formation includes at least three distinctive taxa: Bernissartia sp., Theriosuchus sp., and Goniopholis sp., an association that is also known from several other contemporaneous European vertebrate localities. In contrast to this fauna, the Jydegård and Annero Formations have yielded only rare mesoeucrocodylian remains, which are assigned to Theriosuchus sp. and an undetermined mesoeucrocodylian taxon, possibly Pholidosaurus. Geographically, the Scandinavian localities represent the easternmost and northernmost distribution of typical continental Jurassic-Cretaceous crocodyliform communities in Europe.     

A new twisted-wing parasitoid from mid-Cretaceous amber of Myanmar (Strepsiptera)

A second Mesozoic twisted-wing parasite (Strepsiptera) is described and figured based on an exceptionally well-preserved male in mid-Cretaceous (Cenomanian) amber from northern Myanmar. Phthanoxenos nervosus Engel and Huang, gen. et sp. nov., is distinguished from other strepsipteran lineages, particularly the contemporaneous Cretostylops engeli Grimaldi and Kathirithamby, also in Burmese amber, and Protoxenos janzeni Pohl et al. in mid-Eocene Baltic amber, and assigned to a new family, Phthanoxenidae Engel and Huang, fam. nov. Phthanoxenos exhibit features indicative of a more primitive phylogenetic position than Cretostylopidae but still more derived than Protoxenidae. Brief remarks are made on the geological history of the Strepsiptera.     

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.     

The earliest record of Asian Eusuchia from the Lower Cretaceous Khok Kruat Formation of northeastern Thailand

We describe remains of a new crocodyliform found from the Lower Cretaceous (Aptian) Khok Kruat Formation, northeastern Thailand. Remains consist of two caudal ends of mandibles, two rostral symphyseal parts of right rami of mandibles, a dorsal part of postorbital, a cranial end of squamosal and one osteoderm. Phylogenetic analyses supported inclusion of this crocodyliform into the Eusuchia as it shares several morphological characters with other eusuchians, such as a dorsocaudally oriented retroarticular process, smooth lateral surface of the caudoventral region of mandible, and a craniocaudally oriented ridge on the dorsal surface of retroarticular process. The shape of symphyseal region showed this crocodyliform had a longirostrine snout shape, which is uncommon in early eusuchians. Finding of this crocodyliform draws back the oldest record of Asian eusuchians, which was Tadzhikosuchus, approximately 30 million years and it is the only Mesozoic eusuchian found in East and Southeast Asia.     

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.     

Tail anatomy of Buitreraptor gonzalezorum (Theropoda,Unenlagiidae) and comparisons with other basal paravians

Tail anatomy of unenlagiid theropods remains poorly known. The most complete and informative taxon of this family is Buitreraptor gonzalezorum from the Upper Cretaceous of Rio Negro province, Argentina. The aim of the present contribution is to carry out an analysis of the tail anatomy of Buitreraptor based on its holotype and a newly collected specimen as well. Similarities shared by Buitreraptor, Rahonavis, Anchiornis and Archaeopteryx include: mid-caudal vertebrae with postzygapophyses longer than prezygapophyses, and mid-caudal centra with a system of lateral laminae and concavities. Preservation of paravian skeletons, as well as muscular reconstruction, indicate the presence of two different functional sections of the tail. This contribution sheds light on paravian tail evolution and provides new data on tail changes that occurred along the theropod line towards modern birds.     

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 trematochampsid crocodile from the Early Cretaceous of north-western Patagonia,Argentina and its palaeobiogeographical and phylogenetic implications

Amargasuchus minor is described from the Early Cretaceous (Neocomian, of the La Amarga Formation of northwest Patagonia (Argentina). This species possesses a moderately high and narrow snout, strong festooning, a relatively large number of maxillary teeth and a well developed antorbital fenestra, and it lacks hypertrophied teeth, a combination of characters which suggests that it belongs to the crocodile family Trematochampsidae. Amargasuchus differs, however, from Trematochampsa in several respects. These include an almost straight lateral edge of the maxilla in dorsal aspect, a different distribution of the largest teeth, a longer snout, and laterally compressed alveoli. The new material supports a Gondwanan origin of the Trematochampsidae and agrees with a new phylogenetic hypothesis which considers trematochampsids to be the ancestral group of certain lineages of ziphodont mesosuschian crocodiles.     

A juvenile Edmontosaurus from the late Maastrichtian (Cretaceous) of North America: Implications for ontogeny and phylogenetic inference in saurolophine dinosaurs

The anatomy of an articulated juvenile specimen of the saurolophine hadrosaurid dinosaur Edmontosaurus annectens, LACM 23504, is described in detail. This individual consists of a partial skull and nearly complete articulated postcranium, collected from upper Maastrichtian strata of the Hell Creek Formation of Montana, USA. This study aims to help understand the ontogenetic morphological changes occurring in the cranium and postcranium of saurolophine hadrosaurids using E. annectens as a case study. Emphasis is placed on those morphological attributes that are ontogenetically variable via comparative osteology between the juvenile and the available adult specimens. It is observed that much of the cranial ontogenetic variation relates to the elongation of the skull and mandible. In the postcranium, most of the ontogenetic variation concentrates in the pectoral and pelvic girdles and the stylopodia. Not all the identified patterns of ontogenetic variation may be generalized to all hadrosaurids. The impact of ontogenetic variation on phylogenetically informative characters of saurolophine hadrosaurids is evaluated. It is concluded that, at least for Edmontosaurus annectens (and perhaps other saurolophine hadrosaurids), most characters used in phylogenetic inference of these animals are not affected by ontogeny. Thus, juvenile specimens are still a source of substantial character data suitable for use in phylogenetic analyses of saurolophine relationships. Nevertheless, it is recommended that ontogenetically variable characters are left as missing data in a character–taxon matrix when only juvenile material is available for a given saurolophine taxon. Scoring those characters based solely on juveniles is likely to decrease the accuracy of the phylogenetic inference.     

Past diversity of Proteaceae on subantarctic Campbell Island, a remote outpost of Gondwana

Twelve fossil species of Proteaceous pollen, predominantly attributable to Proteacidites and Beaupreaidites, were recovered from the Maastrichtian-Paleocene sedimentary succession of the Garden Cove Formation on Campbell Island, the southernmost landmass of the Zealandia continent. Among these are two new species, Proteacidites campbellensis and Proteacidites hortisinus. The high diversity of Proteaceae pollen in the sediments encompassing the Cretaceous-Paleogene boundary on Campbell Island is consistent with the fossil record from neighbouring landmasses but strongly contrasts with the impoverished record of the family in the extant New Zealand flora. Examples of Beauprea- and Knightia-like pollen in the Campbell Island assemblages confirm the presence of these lineages on Zealandia by the end of the Cretaceous and suggest that their present endemism in New Caledonia and New Zealand can be explained in terms of relictual vicariant distributions, perhaps modified by northward tracking of warmer climates on Zealandia through the Cenozoic.     

Reworked marine pythonomorph (Reptilia,Squamata) remains in Late Pleistocene cave deposits in central Spain

Three sites at the Calvero de la Higuera complex (Camino Cave, Navalmaíllo Rock Shelter, and Buena Pinta Cave), near the village of Pinilla del Valle (Madrid, Spain), are known for their record of Neanderthals and other Late Pleistocene mammals. Occasionally, they also yield much more ancient reworked remains, which come from the Upper Cretaceous dolomites and carbonatic sandstones in which these caves and shelters are developed. These are mostly teeth of sharks and rays and vertebrae and teeth of bony fishes, but several reptile vertebrae and teeth have also been found. These reptile remains, which we describe here, likely belong to the pythonomorph incertae sedis Carentonosaurus cf. mineaui. This taxon is known from several outcrops in the southwest of France but is rare on the Iberian Peninsula; indeed, only a few remains possibly related to the genus have been found (in the Cabaña Formation, Asturias, Spain). The pythonomorph remains discussed here are the first fossils of marine reptiles from the Madrid region. Should the assignment to Carentonosaurus be confirmed, the teeth would provide novel data on the characteristics of this rather poorly known taxon, and might help clarify its phylogenetic relationship within Pythonomorpha.     

Dinosaur eggshell assemblage from Japan reveals unknown diversity of small theropods

The Lower Cretaceous (Albian) Sasayama Group in the Hyogo Prefecture of southwestern Japan has yielded various vertebrate fossils, including skeletal remains of dinosaurs, anurans, lizards, and mammals, and recently eggshell fragments. Here we report on numerous fossil eggshells from the bone-bearing Kamitaki locality in Tamba City, which represents a diverse dinosaur eggshell assemblage. Of the more than 90 eggshell fragments recovered, five different types were identified, including eggshells that likely belong to a variety of theropods (Nipponoolithus ramosus oogen. et oosp. nov., Elongatoolithus sp., Prismatoolithus sp., and Prismatoolithidae indet.) and at least one ornithopod (Spheroolithus sp.). All eggshells are relatively thin, and a new derived estimation method correlating egg mass with eggshell thickness indicates that they are among the smallest (28–135 g) theropod eggs known, likely laid by small bodied forms. The eggshell assemblage from this locality suggests that a diverse small dinosaur fauna, consisting primarily of theropods, nested in the region, a diversity yet to be evidenced from skeletal remains in Japan.     

The first lizard fossil (Reptilia: Squamata) from the Mesozoic of South Korea

Upper Cretaceous deposits in Mongolia, Chinese Inner Mongolia, and, more recently, southern China, have yielded individually rich and taxonomically diverser lizard assemblages. Here we describe the remains of a new terrestrial lizard, Asprosaurus bibongriensis gen. et sp. nov., from the Upper Cretaceous of South Korea. It represents the first record of a Mesozoic lizard from the Korean Peninsula and, although incomplete, is exceptional in its very large size. Characters of the mandible support attribution to crown-group Anguimorpha, with the closest similarities being to monstersaurs, the group represented today by the venomous North American Beaded lizard and Gila monster, genus Heloderma. This group is well-represented in the Upper Cretaceous fossil record in of eastern Asia, and the remains of large monstersaurs have been recovered from several dinosaur egg localities, suggesting dietary preferences similar to those of the living genus. The new Korean lizard, recovered from the Boseong Bibong-ri Dinosaur Egg Site, fits the same pattern.     

A saltasaurine titanosaur (Sauropoda: Titanosauriformes) from the Angostura Colorada Formation (upper Campanian,Cretaceous) of northwestern Patagonia,Argentina

Saltasaurine titanosaurs are characterized by their relatively small size compared to other sauropods, extreme postcranial pneumaticity, and dermal armour covering the body. This group has been reported in the Upper Cretaceous of the Lecho, Allen, and Anacleto formations of Argentina. We describe here a new saltasaurine specimen (MACN-Pv RN 233) from the Campanian of the Angostura Colorada Formation (Río Negro Province) that is represented by eight caudal vertebrae and six osteoderms. This specimen is described in detail and its phylogenetic relationships with the other three known saltasaurines, as well as its implications to the knowledge of caudal vertebra and osteoderm anatomy, are discussed. Our results place MACN-Pv RN 233 more closely related to Saltasaurus loricatus and Rocasaurus muniozi than to Neuquensaurus australis. MACN-Pv RN 233 possesses a combination of features that differ from other saltasaurines, but because of the fragmentary nature of the specimen we decided for the sake of taxonomic stability to not erect a new taxon. This specimen shows the first unambiguous evidence of chevron pneumatisation for a sauropodomorph, implying a broader osteological invasion of the diverticula from the abdominal air sac than previously thought for this group of dinosaurs. MACN-Pv RN 233 preserves two osteoderm morphotypes, one similar to those reported for Neuquensaurus australis and Saltasaurus loricatus. This new specimen expands the distribution of the group to a new geological unit and increases the dinosaur diversity known for the Angostura Colorada Formation.     

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.