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We report a new specimen of the plesiosaur Cardiocorax mukulu that includes the most complete plesiosaur skull from sub-Saharan Africa. The well-preserved three-dimensional nature of the skull offers rare insight into the cranial anatomy of elasmosaurid plesiosaurians. The new specimen of Cardiocorax mukulu was recovered from Bentiaba, Namibe Province in Angola, approximately three meters above the holotype. The new specimen also includes an atlas-axis complex, seventeen postaxial cervical vertebrae, partial ribs, a femur, and limb elements. It is identified as Cardiocorax mukulu based on an apomorphy shared with the holotype where the cervical neural spine is approximately as long anteroposteriorly as the centrum and exhibits a sinusoidal anterior margin. The new specimen is nearly identical to the holotype and previously referred material in all other aspects. Cardiocorax mukulu is returned in an early-branching or intermediate position in Elasmosauridae in four out of the six of our phylogenetic analyses. Cardiocorax mukulu lacks the elongated cervical vertebrae that is characteristic of the extremely long-necked elasmosaurines, and the broad skull with and a high number of maxillary teeth (28–40) which is characteristic of Aristonectinae. Currently, the most parsimonious explanation concerning elasmosaurid evolutionary relationships, is that Cardiocorax mukulu represents an older lineage of elasmosaurids in the Maastrichtian.

ABSTRACTThe fossil record of post-Paleozoic lungfishes in Greenland is currently restricted to a few brief reports of isolated and undetermined tooth plates coming from the uppermost Fleming Fjord Formation (late Norian) in Jameson Land, central East Greenland. Here, we describe Ceratodus tunuensis, sp. nov., a new dipnoan from a thin bed of calcareous lake mudstone from the ?rsted Dal Member of the Fleming Fjord Formation. The Ceratodus fossil record indicates that during the Late Triassic, this genus was restricted to the middle latitudes of the Northern Hemisphere. This record matches previous paleobiogeographical analyses and indicates that terrestrial biota during the Late Triassic was strongly influenced by paleolatitude.Citation for this article: Agnolin, F. L., O. Mateus, J. Milàn, M. Marzola, O. Wings, J. Schulz Adolfssen, and L. B. Clemmensen. 2018. Ceratodus tunuensis, sp. nov., a new lungfish (Sarcopterygii, Dipnoi) from the Upper Triassic of central East Greenland. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2018.1439834.

ABSTRACT Diplodocid sauropods had a unique skull morphology, with posteriorly retracted nares, an elongated snout, and anteriorly restricted, peglike teeth. Because of the lack of extant analogs in skull structure and tooth morphology, understanding their feeding strategy and diet has been difficult. Furthermore, the general rarity of sauropod skulls and the fragility of their facial elements resulted in a restricted knowledge of cranial anatomy, in particular regarding the internal surface of the facial skull. Here, we describe in detail a well-preserved diplodocid skull visible in medial view. Diagnostic features recognized in other skulls observable in lateral view, such as the extended contribution of the jugal to the antorbital fenestra, are obliterated in medial view due to extensive overlapping joints between the maxilla, jugal, quadratojugal, and the lacrimal. These overlapping joints permitted limited anterior sliding movement of the snout, which likely served as a kind of ?shock-absorbing? mechanism during feeding. Diplodocid skulls therefore seem to have evolved to alleviate stresses inflicted on the snout during backward movements of the head, as would be expected during branch-stripping or raking.ABSTRACT Diplodocid sauropods had a unique skull morphology, with posteriorly retracted nares, an elongated snout, and anteriorly restricted, peglike teeth. Because of the lack of extant analogs in skull structure and tooth morphology, understanding their feeding strategy and diet has been difficult. Furthermore, the general rarity of sauropod skulls and the fragility of their facial elements resulted in a restricted knowledge of cranial anatomy, in particular regarding the internal surface of the facial skull. Here, we describe in detail a well-preserved diplodocid skull visible in medial view. Diagnostic features recognized in other skulls observable in lateral view, such as the extended contribution of the jugal to the antorbital fenestra, are obliterated in medial view due to extensive overlapping joints between the maxilla, jugal, quadratojugal, and the lacrimal. These overlapping joints permitted limited anterior sliding movement of the snout, which likely served as a kind of ?shock-absorbing? mechanism during feeding. Diplodocid skulls therefore seem to have evolved to alleviate stresses inflicted on the snout during backward movements of the head, as would be expected during branch-stripping or raking.

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The Spheroolithidae oospecies Guegoolithus turolensis, putatively attributed to non-hadrosauroid styracosterns was first described in the Barremian of the Iberian Basin, and later reported in the Valanginian–Hauterivian of the Cameros Basin, with both occurrences separated by a few hundred kilometres but by over 10 million years.

ABSTRACTCyclotosaurus naraserluki, sp. nov., is a new Late Triassic capitosaurid amphibian from lacustrine deposits in the Fleming Fjord Formation of the Jameson Land Basin in Greenland. It is based on a fairly complete and well-preserved skull associated with two vertebral intercentra. Previously reported as Cyclotosaurus cf. posthumus, C. naraserluki is unique among cyclotosaurs for having the postorbitals embaying the supratemporals posteromedially. The anterior palatal vacuity presents an autapomorphic complete subdivision by a wide medial premaxillary-vomerine bony connection. The parasphenoid projects between the pterygoids and the exoccipitals, preventing a suture between the two, a primitive condition shared with Rhinesuchidae, Eryosuchus, and Kupferzellia. Within Cyclotosaurus, the Greenlandic skull has a distinctive combination of circular choanae (shared with C. ebrachensis, C. posthumus, and C. robustus) and a convex posteromedial margin of the tabulars (also present in C. ebrachensis and C. intermedius). A phylogenetic analysis indicates that C. naraserluki is the sister taxon of the middle Norian C. mordax from southern Germany, with which it shares a pair of premaxillary foramina. Cyclotosaurus is one of the most successful and diverse genera of Late Triassic temnospondyls, with at least eight species reported from middle Carnian to late Norian. Cyclotosaurus naraserluki is the largest amphibian ever reported from Greenland and one of the Late Triassic vertebrates with the highest northern paleolatitude currently known.http://zoobank.org/urn:lsid:zoobank.org:pub:43AAA541-031C-4EE1-B819-4846EBBD1BBBSUPPLEMENTAL DATA?Supplemental materials are available for this article for free at www.tandfonline.com/UJVPCitation for this article: Marzola, M., O. Mateus, N. H. Shubin, and L. B. Clemmensen. 2017. Cyclotosaurus naraserluki, sp. nov., a new Late Triassic cyclotosaurid (Amphibia, Temnospondyli) from the Fleming Fjord Formation of the Jameson Land Basin (East Greenland). Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1303501.ABSTRACTCyclotosaurus naraserluki, sp. nov., is a new Late Triassic capitosaurid amphibian from lacustrine deposits in the Fleming Fjord Formation of the Jameson Land Basin in Greenland. It is based on a fairly complete and well-preserved skull associated with two vertebral intercentra. Previously reported as Cyclotosaurus cf. posthumus, C. naraserluki is unique among cyclotosaurs for having the postorbitals embaying the supratemporals posteromedially. The anterior palatal vacuity presents an autapomorphic complete subdivision by a wide medial premaxillary-vomerine bony connection. The parasphenoid projects between the pterygoids and the exoccipitals, preventing a suture between the two, a primitive condition shared with Rhinesuchidae, Eryosuchus, and Kupferzellia. Within Cyclotosaurus, the Greenlandic skull has a distinctive combination of circular choanae (shared with C. ebrachensis, C. posthumus, and C. robustus) and a convex posteromedial margin of the tabulars (also present in C. ebrachensis and C. intermedius). A phylogenetic analysis indicates that C. naraserluki is the sister taxon of the middle Norian C. mordax from southern Germany, with which it shares a pair of premaxillary foramina. Cyclotosaurus is one of the most successful and diverse genera of Late Triassic temnospondyls, with at least eight species reported from middle Carnian to late Norian. Cyclotosaurus naraserluki is the largest amphibian ever reported from Greenland and one of the Late Triassic vertebrates with the highest northern paleolatitude currently known.http://zoobank.org/urn:lsid:zoobank.org:pub:43AAA541-031C-4EE1-B819-4846EBBD1BBBSUPPLEMENTAL DATA?Supplemental materials are available for this article for free at www.tandfonline.com/UJVPCitation for this article: Marzola, M., O. Mateus, N. H. Shubin, and L. B. Clemmensen. 2017. Cyclotosaurus naraserluki, sp. nov., a new Late Triassic cyclotosaurid (Amphibia, Temnospondyli) from the Fleming Fjord Formation of the Jameson Land Basin (East Greenland). Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1303501.

A large collection of vertebrate coprolites from black lacustrine shales in the Late Triassic (Rhaetian–Sinemurian) Kap Stewart Formation, East Greenland is examined with regard to internal and external morphology, prey inclusions, and possible relationships to the contemporary vertebrate fauna. A number of the coprolites were mineralogically examined by X-ray diffraction (XRD), showing the primary mineral composition to be apatite, clay minerals, carbonates and, occasionally, quartz in the form of secondary mineral grains. The coprolite assemblage shows multiple sizes and morphotypes of coprolites, and different types of prey inclusions, demonstrating that the coprolite assemblage originates from a variety of different producers.Supplementary material: A description of the size, shape, structure, texture, contents and preservation of the 328 specimens is available at https://doi.org/10.6084/m9.figshare.c.2134335

The purpose of this application, under Articles 78.1 and 81.1 of the Code, is to replace Diplodocus longus Marsh, 1878 as the type species of the sauropod dinosaur genus Diplodocus by the much better represented D. carnegii Hatcher, 1901, due to the undiagnosable state of the holotype of D. longus (YPM 1920, a partial tail and a chevron). The holotype of D. carnegii, CM 84, is a well-preserved and mostly articulated specimen. Casts of it are on display in various museums around the world, and the species has generally been used as the main reference for studies of comparative anatomy or phylogeny of the genus. Both species are known from the Upper Jurassic Morrison Formation of the western United States. The genus Diplodocus is the basis for the family-level taxa diplodocinae Marsh, 1884, diplodocidae Marsh, 1884, diplodocimorpha Marsh, 1884 (Calvo & Salgado, 1995) and diplodocoidea Marsh, 1884 (Upchurch, 1995). It is also a specifier of at least 10 phylogenetic clades. With the replacement of D. longus by D. carnegii as type species, Diplodocus could be preserved as a taxonomic name with generally accepted content. Taxonomic stability of the entire clade diplodocoidea, and the proposed definitions of several clades within Sauropoda, could be maintained.

A large collection of vertebrate coprolites from black lacustrine shales in the Late Triassic (Rhaetian–Sinemurian) Kap Stewart Formation, East Greenland is examined with regard to internal and external morphology, prey inclusions, and possible relationships to the contemporary vertebrate fauna. A number of the coprolites were mineralogically examined by X-ray diffraction (XRD), showing the primary mineral composition to be apatite, clay minerals, carbonates and, occasionally, quartz in the form of secondary mineral grains. The coprolite assemblage shows multiple sizes and morphotypes of coprolites, and different types of prey inclusions, demonstrating that the coprolite assemblage originates from a variety of different producers.Supplementary material: A description of the size, shape, structure, texture, contents and preservation of the 328 specimens is available at https://doi.org/10.6084/m9.figshare.c.2134335

The amniote eggshell functions as a respiratory structure adapted for the optimal transmission of respiratory gasses to and from the embryo according to its physiological requirements. Therefore amniotes with higher oxygen requirements, such as those that sustain higher metabolic rates, can be expected to have eggshells that can maintain a greater gas flux to and from the egg. Studies of extant amniotes have found that eggshells of reduced porosity impose a limit on the metabolic rate of the offspring. Here we show a highly significant relationship between metabolic rates and eggshell porosity in extant amniotes that predicts highly endothermic metabolic rates in dinosaurs. This study finds the eggshell porosity of extant endotherms to be significantly higher than that of extant ectotherms. Eggshell porosity values of dinosaurs are found to be significantly higherthan that of extant ectotherms, but not extant endotherms. Dinosaur eggshells are commonly preserved in the fossil record, and porosity may be readily identified and measured. This provides a simple tool to identify metabolic rates in extinct egg-laying tetrapods whose eggs possessed a mineralized shell

We present the δ13C and paleomagnetic stratigraphy for marine strata at the coast of southern Angola, anchored by an intercalated basalt with a whole rock 40Ar/39Ar radiometric age of 84.6 ± 1.5 Ma, being consistent with both invertebrate and vertebrate biostratigraphy. This is the first African stable carbon isotope record correlated to significant events in the global carbon cycle spanning the Late Cenomanian to Early Maastrichtian. A positive ∼ 3‰ excursion seen in bivalve shells below the basalt indicates the Cenomanian-Turonian Boundary Event at 93.9 Ma, during Oceanic Anoxic Event 2. Additional excursions above the basalt are correlated to patterns globally, including a negative ∼ 3‰ excursion near the top of the section interpreted as part of the Campanian-Maastrichtian Boundary Events. The age of the basalt ties the studied Bentiaba section to a pulse of Late Cretaceous magmatic activity around the South Atlantic and significant tectonic activity, including rotation, of the African continent.

We here present fossil Crocodylomorpha eggshells from the Upper Jurassic Lourinhã Formation of Portugal, recovered from five sites: one nest from Cambelas with 13 eggs, and three partial eggs and various fragments from, Paimogo N (I), Paimogo S (II), Casal da Rola, and Peralta. All specimens but the nest were found in association with dinosaur egg material. Our research reveals that on a micro- and ultrastructural analysis, all samples present the typical characters consistent with crocodiloid eggshell morphotype, such as the shell unit shape, the organization of the eggshell layers, and the triangular blocky extinction observed with crossed nicols. We assign the material from the Lourinhã Formation to the oofamily Krokolithidae, making it the oldest crocodylomorph eggs known so far, as well as the best record for eggs of non- crocodylian crocodylomorphs. Furthermore, our study indicates that the basic structure of crocodiloid eggshells has remained stable since at least the Upper Jurassic.

We here present fossil Crocodylomorpha eggshells from the Upper Jurassic Lourinhã Formation of Portugal, recovered from five sites: one nest from Cambelas with 13 eggs, and three partial eggs and various fragments from, Paimogo N (I), Paimogo S (II), Casal da Rola, and Peralta. All specimens but the nest were found in association with dinosaur egg material. Our research reveals that on a micro- and ultrastructural analysis, all samples present the typical characters consistent with crocodiloid eggshell morphotype, such as the shell unit shape, the organization of the eggshell layers, and the triangular blocky extinction observed with crossed nicols. We assign the material from the Lourinhã Formation to the oofamily Krokolithidae, making it the oldest crocodylomorph eggs known so far, as well as the best record for eggs of non- crocodylian crocodylomorphs. Furthermore, our study indicates that the basic structure of crocodiloid eggshells has remained stable since at least the Upper Jurassic.

In the early 20thcentury, Belgian naturalists reported Paleocene and Eocene sharks, the bothremydid pleurodiran turtleTaphosphrys(formerlyBantuchelys), and a neosuchian and the dyrosaurid crocodyliformCongosaurusfrom coastal outcrops near Landana in the northern province of Cabinda, Angola. In 1935, rare and fragmentary mammals were reported from strata at Malembo Point, south of Landana, and originally considered to be Miocene in age. Subsequent re-identification of the mammal fossils suggested that the Malembo deposits are Lower Oligocene based on the presence of an arsinoithere and hyracoids. An anthropoid canine, originally identified as a carnivore incisor, was also recognized. In four recent expeditions to Cabinda, Projecto PaleoAngola has traced and measured the stratigraphy between Landana and Malembo Point and sampled these deposits for pollen, stable isotopes, detrital zircons, paleomagnetic stratigraphy, and for U/Pb dating of bones, teeth, and coprolites. Although the faunas from all Cabinda localities are dominated by sharks and rays, new discoveries from Landana include a complete cheloniid cryptodire turtle skull, a small snake vertebra, and a bird bone. Discoveries from the Malembo level include a narrow-snouted crocodyliform similar toCongosaurusandEuthecodon, an arsinoithere anterior tooth, an upper molar similar to that of the ptolemaiidanKelba, an unidentified mammalian incisor, and a large primate-like premolar. Recent biostratigraphic advances in East Africa and the new fossil discoveries in Cabinda suggest similarity to late Oligocene faunas in Ethiopia and Kenya, although the large primate-like premolar is unique. To this day, the fossil localities of Malembo provide the only coastal, low latitude, low elevation record of West African Cenozoic terrestrial mammals. https://gsa.confex.com/gsa/2013AM/webprogram/Paper232115.html

Ossified gastralia, clavicles and sternal ribs are known in a variety of reptilians, including dinosaurs. In sauropods, however, the identity of these bones is controversial. The peculiar shapes of these bones complicate their identification, which led to various differing interpretations in the past. Here we describe different elements from the chest region of diplodocids, found near Shell, Wyoming, USA. Five morphotypes are easily distinguishable: (A) elongated, relatively stout, curved elements with a spatulate and a bifurcate end resemble much the previously reported sauropod clavicles, but might actually represent interclavicles; (B) short, L-shaped elements, mostly preserved as a symmetrical pair, probably are the real clavicles, as indicated by new findings in diplodocids; (C) slender, rod-like bones with rugose ends are highly similar to elements identified as sauropod sternal ribs; (D) curved bones with wide, probably medial ends constitute the fourth morphotype, herein interpreted as gastralia; and (E) irregularly shaped elements, often with extended rugosities, are included into the fifth morphotype, tentatively identified as sternal ribs and/or intercostal elements. To our knowledge, the bones previously interpreted as sauropod clavicles were always found as single bones, which sheds doubt on the validity of their identification. Various lines of evidence presented herein suggest they might actually be interclavicles – which are single elements. This would be the first definitive evidence of interclavicles in dinosauromorphs. Previously supposed interclavicles in the early sauropodomorph Massospondylus or the theropods Oviraptor and Velociraptor were later reinterpreted as clavicles or furculae. Independent from their identification, the existence of the reported bones has both phylogenetic and functional significance. Their presence in non-neosauropod Eusauropoda and Flagellicaudata and probable absence in rebbachisaurs and Titanosauriformes shows a clear character polarity. This implicates that the ossification of these bones can be considered plesiomorphic for Sauropoda. The proposed presence of interclavicles in sauropods may give further support to a recent study, which finds a homology of the avian furcula with the interclavicle to be equally parsimonious to the traditional theory that furcula were formed by the fusion of the clavicles. Functional implications are the stabilizing of the chest region, which coincides with the development of elongated cervical and caudal vertebral columns or the use of the tail as defensive weapon. The loss of ossified chest bones coincides with more widely spaced limbs, and the evolution of a wide-gauge locomotor style.

The diversity of fossil crocodylomorphs in Portugal is high, with occurrence as old as Mystriosaurus (=Steneosaurus) bollensis from the Lower Jurassic. The Late Jurassic forms are the better documented, and include the following taxa: Machimosaurus hugii, Lisboasaurus estesi Seiffert, 1973, Lusitanisuchus mitrocostatus Seiffert, 1975; Schwarz & Fechner 2004, Theriosuchus guimarotae Schwarz and Salisbury 2005, Cf. Alligatorium, Goniopholis baryglyphaeus, and a crocodylomorph-like eggs in dinosaur nests (Mateus et al., 1998; Ricqlès et al., 2001). From the Lower Cretaceous were reported a few dinosaurs but its record is strangely scarce in crocodylomorphs (Mateus et al., 2011). The Upper Cretaceous crocodiles show a large diversity, but it is mostly based in fragmentary material that require revision, such as “Crocodylus” blavieri? Grey from the Upper Campanian - Maastrichtian of Viso, near Aveiro (initially reported by Sauvage 1897-98), Goniopholis cf. crassidens Owen 1841 and Oweniasuchus pulchelus Jonet 1981. Moreover there is a fascinating, but poorly understood, crocodylomorph diversity in the Cenomanian of Portugal, documented by fragmentary specimens that have been doubtfully assigned to Thoracosaurus Leidy 1852 of the Middle Cenomanian of Cacém, to the nomen dubium Oweniasuchus lusitanicus Sauvage 1897-98 (interpreted as a mesosuchian goniopholid) based in a fragmentary mandible from the Campanian-Maastrichtian, and also from the Middle Cenomanian of Portugal, Buffetaut and Lauverjat (1978) report an fragmentary unidentified possible dyrosaurid from Nazaré. All this specimens are too incomplete to be compared with the specimen here described. In contrast, Cenozoic crocodiles of Portugal are often known after complete skulls and several individuals. The taxa list include Iberosuchus macrodon (Lower to Middle Eocene), Tomistoma calaritanus (Early Miocene) and T. lusitanica (Burdigalian-Helvetian), and Diplocynodon sp. (Antunes, 1961, 1987, 1994).
At least, two different morphotypes of crocodylomorph eggs from the Late Jurassic of Lourinhã Formation are also known.
A new specimen here reported of crocodile based in a partial skull and mandible (ML1818) from the Uppermost Middle Cenomanian platform carbonates of Baixo Mondego, west central Portugal (Tentúgal Fm., Callapez, 2004). The taxon is phylogenetically positioned as a basal Eusuchia, due to the choanae enclosed by the pterygoid, and closely related with stem Crocodylia and Borealosuchus. This specimen represents the only well documented and valid eusuchian species in the Cenomanian of Europe and is the oldest representative of an eusuchian crocodylomorph, with the exception for the Barremian Hylaeochampsa vectiana.

Although rich in Cretaceous vertebrate fossils, prior to 2005 the amniote fossil record of Angola was poorly known. Two horizons and localities have yielded the majority of the vertebrate fossils collected thus far; the Turonian Itombe Formation of Iembe in Bengo Province and the Maastrichtian Mocuio Formation of Bentiaba in Namibe Province. Amniotes of the Mesozoic of Angola are currently restricted to the Cretaceous and include eucryptodire turtles, plesiosaurs, mosasaurs, pterosaurs, and dinosaurs. Recent collecting efforts have greatly expanded our knowledge of the amniote fauna of Angola and most of the taxa reported here were unknown prior to 2005.

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