Exceptionally preserved shark fossils from Mexico elucidate the long-standing enigma of the Cretaceous elasmobranch Ptychodus.

The fossil fish Ptychodus Agassiz, 1834, characterized by a highly distinctive grinding dentition and an estimated gigantic body size (up to around 10 m), has remained one of the most enigmatic extinct elasmobranchs (i.e. sharks, skates and rays) for nearly two centuries. This widespread Cretaceous taxon is common in Albian to Campanian deposits from almost all continents. However, specimens mostly consist of isolated teeth or more or less complete dentitions, whereas cranial and post-cranial skeletal elements are very rare. Here we describe newly discovered material from the early Late Cretaceous of Mexico, including complete articulated specimens with preserved body outline, which reveals crucial information on the anatomy and systematic position of Ptychodus. Our phylogenetic and ecomorphological analyses indicate that ptychodontids were high-speed (tachypelagic) durophagous lamniforms (mackerel sharks), which occupied a specialized predatory niche previously unknown in fossil and extant elasmobranchs. Our results support the view that lamniforms were ecomorphologically highly diverse and represented the dominant group of sharks in Cretaceous marine ecosystems. Ptychodus may have fed predominantly on nektonic hard-shelled prey items such as ammonites and sea turtles rather than on benthic invertebrates, and its extinction during the Campanian, well before the end-Cretaceous crisis, might have been related to competition with emerging blunt-toothed globidensine and prognathodontine mosasaurs.

Manta-like planktivorous sharks in Late Cretaceous oceans.

The ecomorphological diversity of extinct elasmobranchs is incompletely known. Here, we describe Aquilolamna milarcae, a bizarre probable planktivorous shark from early Late Cretaceous open marine deposits in Mexico. Aquilolamna, tentatively assigned to Lamniformes, is characterized by hypertrophied, slender pectoral fins. This previously unknown body plan represents an unexpected evolutionary experimentation with underwater flight among sharks, more than 30 million years before the rise of manta and devil rays (Mobulidae), and shows that winglike pectoral fins have evolved independently in two distantly related clades of filter-feeding elasmobranchs. This newly described group of highly specialized long-winged sharks (Aquilolamnidae) displays an aquilopelagic-like ecomorphotype and may have occupied, in late Mesozoic seas, the ecological niche filled by mobulids and other batoids after the Cretaceous-Paleogene boundary.

The first Gondwanan borioteiioid lizard and the mid-Cretaceous dispersal event between North America and Africa.

Borioteiioidea are an enigmatic group of Cretaceous lizards widely distributed in northern continents (Laurasia). Here, we describe the first borioteiioid lizard from Gondwana, represented by a new species of the polyglyphanodontine genus Bicuspidon, B. hogreli sp. nov., from the Cenomanian Kem Kem beds of Morocco. The discovery of Bicuspidon hogreli sp. nov., which is one of the oldest known member of Polyglyphanodontini, challenges previous assumptions on the center of origin and dispersal of the group. In addition, the known distribution of Bicuspidon (Cenomanian of Utah and Morocco, and Santonian-Maastrichtian of eastern Europe) suggests a complex palaeobiogeographical history for the genus. The existence of a terrestrial dispersal route persisting during the whole Early Cretaceous between North America and Africa is hypothesized to explain some similarities observed between the Cenomanian squamate assemblages of these two continents. Alternatively, dispersal between the two landmasses may have occurred by transatlantic rafting. During the Cenomanian-Santonian interval, Bicuspidon colonized the European archipelago probably from North Africa, like many "Eurogondwanan" taxa.

A unique Cretaceous-Paleogene lineage of piranha-jawed pycnodont fishes.

The extinct group of the Pycnodontiformes is one of the most characteristic components of the Mesozoic and early Cenozoic fish faunas. These ray-finned fishes, which underwent an explosive morphological diversification during the Late Cretaceous, are generally regarded as typical shell-crushers. Here we report unusual cutting-type dentitions from the Paleogene of Morocco which are assigned to a new genus of highly specialized pycnodont fish. This peculiar taxon represents the last member of a new, previously undetected 40-million-year lineage (Serrasalmimidae fam. nov., including two other new genera and Polygyrodus White, 1927) ranging back to the early Late Cretaceous and leading to exclusively carnivorous predatory forms, unique and unexpected among pycnodonts. Our discovery indicates that latest Cretaceous-earliest Paleogene pycnodonts occupied more diverse trophic niches than previously thought, taking advantage of the apparition of new prey types in the changing marine ecosystems of this time interval. The evolutionary sequence of trophic specialization characterizing this new group of pycnodontiforms is strikingly similar to that observed within serrasalmid characiforms, from seed- and fruit-eating pacus to flesh-eating piranhas.

Denticle-embedded ampullary organs in a Cretaceous shark provide unique insight into the evolution of elasmobranch electroreceptors.

Here, we report a novel type of dermal denticle (or placoid scale), unknown among both living and fossil chondrichthyan fishes, in a Cretaceous lamniform shark. By their morphology and location, these dermal denticles, grouped into clusters in the cephalic region, appear to have been directly associated with the electrosensory ampullary system. These denticles have a relatively enlarged (∼350 µm in diameter), ornamented crown with a small (∼100 µm) asterisk- or cross-shaped central perforation connected to a multi-alveolate internal cavity. The formation of such a complex structure can be explained by the annular coalescence and fusion, around an ampullary vesicle, of several developmental units still at papillary stage (i.e. before mineralization), leading to a single denticle embedding an alveolar ampulla devoid of canal. This differs from larger typical ampullae of Lorenzini with a well-developed canal opening in a pore of the skin and may represent another adaptive response to low skin resistance. Since it has been recently demonstrated that ampullary organs arise from lateral line placodes in chondrichthyans, this highly specialized type of dermal denticle (most likely non-deciduous) may be derived from the modified placoid scales covering the superficial neuromasts (pit organs) of the mechanosensory lateral line system of many modern sharks.

A Cretaceous eutriconodont and integument evolution in early mammals.

The Mesozoic era (252-66 million years ago), known as the domain of dinosaurs, witnessed a remarkable ecomorphological diversity of early mammals. The key mammalian characteristics originated during this period and were prerequisite for their evolutionary success after extinction of the non-avian dinosaurs 66 million years ago. Many ecomorphotypes familiar to modern mammal fauna evolved independently early in mammalian evolutionary history. Here we report a 125-million-year-old eutriconodontan mammal from Spain with extraordinary preservation of skin and pelage that extends the record of key mammalian integumentary features into the Mesozoic era. The new mammalian specimen exhibits such typical mammalian features as pelage, mane, pinna, and a variety of skin structures: keratinous dermal scutes, protospines composed of hair-like tubules, and compound follicles with primary and secondary hairs. The skin structures of this new Mesozoic mammal encompass the same combination of integumentary features as those evolved independently in other crown Mammalia, with similarly broad structural variations as in extant mammals. Soft tissues in the thorax and abdomen (alveolar lungs and liver) suggest the presence of a muscular diaphragm. The eutriconodont has molariform tooth replacement, ossified Meckel's cartilage of the middle ear, and specialized xenarthrous articulations of posterior dorsal vertebrae, convergent with extant xenarthran mammals, which strengthened the vertebral column for locomotion.

Taxonomic composition and trophic structure of the continental bony fish assemblage from the early late cretaceous of Southeastern Morocco.

The mid-Cretaceous vertebrate assemblage from south-eastern Morocco is one of the most diversified continental vertebrate assemblages of this time worldwide. The bony fish component (coelacanths, lungfishes and ray-finned fishes) is represented by relatively complete specimens and, mostly, by fragmentary elements scattered along 250 kilometres of outcrops. Here we revisit the bony fish assemblage by studying both isolated remains collected during several fieldtrips and more complete material kept in public collections. The assemblage comprises several lungfish taxa, with the first mention of the occurrence of Arganodus tiguidiensis, and possibly two mawsoniid coelacanths. A large bichir cf. Bawitius, is recorded and corresponds to cranial elements initially referred to 'Stromerichthys' from coeval deposits in Egypt. The ginglymodians were diversified with a large 'Lepidotes' plus two obaichthyids and a gar. We confirm here that this gar belongs to a genus distinctive from Recent gars, contrary to what was suggested recently. Teleosteans comprise a poorly known ichthyodectiform, a notopterid, a probable osteoglossomorph and a large tselfatiiform, whose cranial anatomy is detailed. The body size and trophic level for each taxon are estimated on the basis of comparison with extant closely related taxa. We plotted the average body size versus average trophic level for the Kem Kem assemblage, together with extant marine and freshwater assemblages. The Kem Kem assemblage is characterized by taxa of proportionally large body size, and by a higher average trophic level than the trophic level of the extant compared freshwater ecosystems, but lower than for the extant marine ecosystems. These results should be regarded with caution because they rest on a reconstructed assemblage known mostly by fragmentary remains. They reinforce, however, the ecological oddities already noticed for this mid-Cretaceous vertebrate ecosystem in North Africa.

A new crested pterosaur from the Early Cretaceous of Spain: the first European tapejarid (Pterodactyloidea: Azhdarchoidea).

BACKGROUND: The Tapejaridae is a group of unusual toothless pterosaurs characterized by bizarre cranial crests. From a paleoecological point of view, frugivorous feeding habits have often been suggested for one of its included clades, the Tapejarinae. So far, the presence of these intriguing flying reptiles has been unambiguously documented from Early Cretaceous sites in China and Brazil, where pterosaur fossils are less rare and fragmentary than in similarly-aged European strata. METHODOLOGY/PRINCIPAL FINDINGS: Europejara olcadesorum gen. et sp. nov. is diagnosed by a unique combination of characters including an unusual caudally recurved dentary crest. It represents the oldest known member of Tapejaridae and the oldest known toothless pterosaur. The new taxon documents the earliest stage of the acquisition of this anatomical feature during the evolutionary history of the Pterodactyloidea. This innovation may have been linked to the development of new feeding strategies. CONCLUSION/SIGNIFICANCE: The discovery of Europejara in the Barremian of the Iberian Peninsula reveals an earlier and broader global distribution of tapejarids, suggesting a Eurasian origin of this group. It adds to the poorly known pterosaur fauna of the Las Hoyas locality and contributes to a better understanding of the paleoecology of this Konservat-Lagerstätte. Finally, the significance of a probable contribution of tapejarine tapejarids to the early angiosperm dispersal is discussed.

Direct evidence of hybodont shark predation on Late Jurassic ammonites.

Sharks are known to have been ammonoid predators, as indicated by analysis of bite marks or coprolite contents. However, body fossil associations attesting to this predator-prey relationship have never been described so far. Here, I report a unique finding from the Late Jurassic of western France: a complete specimen of the Kimmeridgian ammonite Orthaspidoceras bearing one tooth of the hybodont shark Planohybodus. Some possible tooth puncture marks are also observed. This is the first direct evidence of such a trophic link between these two major Mesozoic groups, allowing an accurate identification of both organisms. Although Planohybodus displays a tearing-type dentition generally assumed to have been especially adapted for large unshelled prey, our discovery clearly shows that this shark was also able to attack robust ammonites such as aspidoceratids. The direct evidence presented here provides new insights into the Mesozoic marine ecosystem food webs.

Mammalian hairs in Early Cretaceous amber.

Two mammalian hairs have been found in association with an empty puparium in a approximately 100-million-year-old amber (Early Cretaceous) from France. Although hair is known to be an ancestral, ubiquitous feature in the crown Mammalia, the structure of Mesozoic hair has never been described. In contrast to fur and hair of some Jurassic and Cretaceous mammals preserved as carbonized filaments, the exceptional preservation of the fossils described here allows for the study of the cuticular structure. Results show the oldest direct evidence of hair with a modern scale pattern. This discovery implies that the morphology of hair cuticula may have remained unchanged throughout most of mammalian evolution. The association of these hairs with a possible fly puparium provides paleoecological information and indicates peculiar taphonomic conditions.

The oldest modern therian mammal from Europe and its bearing on stem marsupial paleobiogeography.

We report the discovery of mammalian tribosphenic teeth from the basal Cenomanian of southwestern France that we refer to a new primitive marsupial-like form identified as a basal taxon of Marsupialiformes, a new clade recognized here to include the crown group Marsupialia and primitive stem lineages more closely related to Marsupialia than to Deltatheroida. Arcantiodelphys marchandi gen et sp nov. shares several significant marsupial-like features (s.l.) with marsupialiform taxa known from the North American Mid-Cretaceous. Among marsupialiforms, it shows a closer resemblance to Dakotadens. This resemblance, which is plesiomorphic within "tribotherians," makes Arcantiodelphys one of the most archaic known Marsupialiformes. Moreover, Arcantiodelphys is characterized by an original and precocious crushing specialization. Both the plesiomorphic and autapomorphic characteristics of Arcantiodelphys among Marsupialiformes might be explained by an Eastern origin from Asian stem metatherians, with some in situ European evolution. In addition, the presence of a mammal with North American affinities in western Europe during the early Late Cretaceous provides further evidence of a large Euramerican biogeographical province at this age or slightly before. Concerning the paleobiogeographical history of the first stem marsupialiforms during the Albian-Cenomanian interval, 2 possible dispersal routes from an Asian metatherian ancestry can be proposed: Asia to Europe via North America and Asia to North America via Europe. The main significance of the Archingeay-Les Nouillers mammal discovery is that it indicates that the beginning of the stem marsupialiforms history involved not only North America but also Europe, and that this early history in Europe remains virtually unknown.

The early evolution of feathers: fossil evidence from Cretaceous amber of France.

The developmental stages of feathers are of major importance in the evolution of body covering and the origin of avian flight. Until now, there were significant gaps in knowledge of early morphologies in theoretical stages of feathers as well as in palaeontological material. Here we report fossil evidence of an intermediate and critical stage in the incremental evolution of feathers which has been predicted by developmental theories but hitherto undocumented by evidence from both the recent and the fossil records. Seven feathers have been found in an Early Cretaceous (Late Albian, ca 100 Myr) amber of western France, which display a flattened shaft composed by the still distinct and incompletely fused bases of the barbs forming two irregular vanes. Considering their remarkably primitive features, and since recent discoveries have yielded feathers of modern type in some derived theropod dinosaurs, the Albian feathers from France might have been derived either from an early bird or from a non-avian dinosaur.