The skull of living birds is greatly modified from the condition found in their dinosaurian antecedents. Bird skulls have an enlarged, toothless premaxillary beak and an intricate kinetic system that includes a mobile palate and jaw suspensorium. The expanded avian neurocranium protects an enlarged brain and is flanked by reduced jaw adductor muscles. However, the order of appearance of these features and the nature of their earliest manifestations remain unknown. The Late Cretaceous toothed bird Ichthyornis dispar sits in a pivotal phylogenetic position outside living groups: it is close to the extant avian radiation but retains numerous ancestral characters. Although its evolutionary importance continues to be affirmed, no substantial new cranial material of I. dispar has been described beyond incomplete remains recovered in the 1870s. Jurassic and Cretaceous Lagerstätten have yielded important avialan fossils, but their skulls are typically crushed and distorted . Here we report four three-dimensionally preserved specimens of I. dispar-including an unusually complete skull-as well as two previously overlooked elements from the Yale Peabody Museum holotype, YPM 1450. We used these specimens to generate a nearly complete three-dimensional reconstruction of the I. dispar skull using high-resolution computed tomography. Our study reveals that I. dispar had a transitional beak-small, lacking a palatal shelf and restricted to the tips of the jaws-coupled with a kinetic system similar to that of living birds. The feeding apparatus of extant birds therefore evolved earlier than previously thought and its components were functionally and developmentally coordinated. The brain was relatively modern, but the temporal region was unexpectedly dinosaurian: it retained a large adductor chamber bounded dorsally by substantial bony remnants of the ancestral reptilian upper temporal fenestra. This combination of features documents that important attributes of the avian brain and palate evolved before the reduction of jaw musculature and the full transformation of the beak.
The cranial and postcranial anatomy of the basal hadrosaurid dinosaur Eotrachodon orientalis, from the uppermost Santonian of southern Appalachia (southeastern U.S.A.), is described in detail. This animal is the only known pre-Campanian non-lambeosaurine hadrosaurid, and the most complete hadrosauroid known from Appalachia. E. orientalis possesses a mosaic of plesiomorphic and derived characters in the context of Hadrosauroidea. Characters shared with basal hadrosauroids include a short and sloping maxillary ectopterygoid shelf, caudally prominent maxillary jugal process, one functional tooth per alveolus on the maxillary occlusal plane, a jugal rostral process with a shallow caudodorsal margin and medioventrally facing articular facet, a vertical dentary coronoid process with a poorly expanded apex, and tooth crowns with accessory ridges. Derived characters shared with other hadrosaurids include a circumnarial depression compartmented into three fossae (as in brachylophosaurins and Edmontosaurus), a thin everted premaxillary oral margin (as in Gryposaurus, Prosaurolophus, and Saurolophus), and a maxilla with a deep and rostrocaudally extensive rostrodorsal region with a steeply sloping premaxillary margin (as in Gryposaurus). Eotrachodon orientalis differs primarily from the other hadrosauroid from the Mooreville Chalk of Alabama, Lophorhothon atopus, in having a slender and crestless nasal whose caudodorsal margin is not invaded by the circumnarial depression. Hadrosaurus foulkii, the only other known hadrosaurid from Appalachia, is distinct from E. orientalis in having dentary teeth lacking accessory ridges and a dorsally curved shaft of the ischium. A histological section of the tibia of the E. orientalis holotype (MSC 7949) suggests that this individual was actively growing at the time of death and, thus, had the potential to become a larger animal later in development.
The Tallahatta Formation, Lisbon Formation, and Gosport Sand are the three lithostratigraphic units that make up the lower-to-middle Eocene Claiborne Group. In Alabama, these marine units are among the most fossiliferous in the state and a long history of scattered reports have attempted to document their fossil diversity. In this study, we examined 20931 elasmobranch and bony fish elements, including otoliths, derived from Claiborne Group units in Alabama and identified 115 unequivocal taxa. Among the taxa identified, one new species is described, Carcharhinus mancinae sp. nov., and Pseudabdounia gen. nov. is a new genus erected to include two species formerly placed within Abdounia Capatta, 1980. New taxonomic combinations proposed include Pseudabdounia claibornensis (White, 1956) gen. et comb. nov., Pseudabdounia recticona (Winkler, 1874) gen. et comb. nov., Physogaleus alabamensis (Leriche, 1942) comb. nov., and Eutrichiurides plicidens (Arambourg, 1952) comb. nov. We also report the first North American paleobiogeographic occurrences of Aturobatis aff. A. aquensis Adnet, 2006, Brachycarcharias atlasi (Arambourg, 1952), Eutrichiurides plicidens comb. nov., Galeorhinus louisi Adnet & Cappetta, 2008, Ginglymostoma maroccanum Noubhani & Cappetta, 1997, Gymnosarda sp., Mennerotodus sp., Rhizoprionodon ganntourensis (Arambourg, 1952), Stenoscyllium aff. S. priemi Noubhani & Cappetta, 1997, Trichiurus oshosunensis White, 1926, and the first North American occurrence for a fossil member of the Balistidae Risso, 1810. Our sample also included 26 taxa that represented first paleobiogeographic occurrences for Alabama, including Abdounia beaugei (Arambourg, 1935), Albula eppsi White, 1931, Ariosoma nonsector Nolf & Stringer, 2003, Anisotremus? sp., Anomotodon sp., Brachycarcharias twiggsensis (Case, 1981), Burnhamia daviesi (Woodward, 1889), Eoplinthicus yazooensis Capetta & Stringer, 2002, Galeorhinus ypresiensis (Casier, 1946), Gnathophis meridies (Frizzell & Lamber, 1962), Haemulon? obliquus (Müller, 1999), Hypolophodon sylvestris (White, 1931), Malacanthus? sulcatus (Koken, 1888), Meridiania cf. M. convexa Case, 1994, Palaeocybium proosti (Storms, 1897), Paraconger sector (Koken, 1888), Paralbula aff. P. marylandica Blake, 1940, Phyllodus toliapicus Agassiz, 1844, Propristis schweinfurthi Dames, 1883, Pycnodus sp., Pythonichthys colei (Müller, 1999), Scomberomorus stormsi (Leriche, 1905), Signata stenzeli Frizzell & Dante, 1965, and Signata nicoli Frizzell & Dante, 1965, and the first Paleogene occurrences in Alabama of a member of the Gobiidae Cuvier, 1816. A biostratigraphic analysis of our sample showed stratigraphic range extensions for several taxa, including the first Bartonian occurrences of Eoplinthicus yazooensis, Jacquhermania duponti (Winkler, 1876), Meridiania cf. M. convexa, Phyllodus toliapicus, and “Rhinobatos” bruxelliensis (Jaekel, 1894), range extensions into the late Ypresian and Bartonian for Tethylamna dunni Cappetta & Case, 2016 and Scoliodon conecuhensis Cappetta & Case, 2016, the first late Ypresian records of Galeorhinus louisi, the first Lutetian occurrence of Gymnosarda Gill, 1862, and a range extension for Fisherichthys aff. F. folmeri Weems, 1999 into the middle Bartonian. Larger biostratigraphic and evolutionary trends are also documented, such as the acquisition of serrations in Otodus spp., possible population increases for the Rhinopterinae Jordan & Evermann, 1896 and Carcharhiniformes Compagno, 1973 in the Bartonian, and the apparent diversification of the Tetraodontiformes Berg, 1940 during the same stage. This study helps better our understanding of early-to-middle Eocene elasmobranch and bony fish diversity, paleobiogeography, and biostratigraphy in the Gulf Coastal Plain of North America.
Resolving the phylogeny of sea turtles is uniquely challenging given the high potential for the unification of convergent lineages due to systematic homoplasy. Equivocal reconstructions of marine turtle evolution subsequently inhibit efforts to establish fossil calibrations for molecular divergence estimates and prevent the accurate reconciliation of biogeographic or palaeoclimatic data with phylogenetic hypotheses. Here we describe a new genus and species of marine turtle, Asmodochelys parhami, from the Upper Campanian Demopolis Chalk of Alabama and Mississippi, USA represented by three partial shells. Phylogenetic analysis shows that A. parhami belongs to the ctenochelyids, an extinct group that shares characteristics with both pan-chelonioids and pan-cheloniids. In addition to supporting Ctenochelyidae as a sister taxon of Chelonioidea, our analysis places Protostegidae outside of the Chelonioidea crown group and recovers Allopleuron hofmanni as a stem dermochelyid. Gap excess ratio (GER) results indicate a strong stratigraphic congruence of our phylogenetic hypothesis; however, the highest GER value is associated with the phylogenetic hypothesis of marine turtles which excludes Protostegidae from the Cryptodira crown group. Ancestral range estimations derived from our phylogeny imply a European or North American origin of Chelonioidea in the middle-to-late Campanian, approximately 20 Myr earlier than current molecular divergence studies suggest.
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