Turiasauria-like teeth from the Upper Jurassic of the Lusitanian Basin,Portugal

Turiasauria is a clade of eusauropods with a wide stratigraphic range that could extend from the Bathonian to the lower Aptian including Turiasaurus, Losillasaurus, Zby and putatively, Galveosaurus, Atlasaurus and isolated remains from Middle Jurassic-to-Lower Cretaceous. Some are characterised by the presence of heart-shaped teeth. Several tooth occurrences from the Portuguese Upper Jurassic with this type of morphology (SI: 1.1–1.8) are reported and discussed. If this morphology is regarded as synapomorphic of Turiasauria, the teeth will be tentatively related to this clade. From a sample of 43 teeth, three main morphotypes are described. Three hypotheses might explain the morphological variation: (1) the range of tooth morphologies indicates variation in the jaw, (2) the range of tooth morphologies indicates taxonomic variation or (3) a combination of both. The general wear pattern in morphotypes I and II starts with a distal facet, then the appearance of mesial/apical facet and finally a ‘V’-shaped facet. In morphotype III, the wear begins with a mesial facet. The variability observed for Portuguese Upper Jurassic specimens is congruent with the morphological variability along the tooth row shown by other sauropods with spatulate/spoon-shaped teeth and it is considered the most parsimonious hypothesis to explain it.     

Ammonite succession at the Bajocian/Bathonian boundary in the Cabo Mondego region (Portugal)

The Cabo Mondego outcrops exposed along the cliffs, on the western margin of the Iberian Plate, show an expanded stratigraphic section of Lower Bathonian deposits containing abundant ammonoids. Upper Bajocian deposits correspond to similar facies, of muddy limestones alternating with marlstones, although ammonoids are scarce. A detailed succession of ammonites across the Bajocian/Bathonian boundary has been recognized at Cabo Mondego, which can form a useful bio‐ and chronostratigraphic standard for the Lusitanian Basin. The revision of previous collections from the classical section and new field samplings of two other separate sections allow the recognition through up to twenty metres of thickness, the highest zone of Bajocian (Parkinsoni Zone) and the lowest zone of Bathonian (Zigzag Zone). The Parkinsoni and the Zigzag zones established for NW European areas and belonging to the Northwest European Province, can be identified in the Lusitanian Basin, although the ammonite fossil assemblages are composed of Submediterranean taxa. However, a subdivision of the Parkinsoni Zone is not possible, due to the scarcity of well preserved ammonoids. The Zigzag Zone can be recognized and characterized as composed of two subunits (Parvum and Macrescens subzones) as represented in diverse European basins of the Submediterranean Province. Ammonite fossil assemblages of the Parvum Subzone may be grouped into two successive horizons, which are biochronostratigraphically equivalent to the subdivisions of the Convergens Subzone distinguished in the Digne‐Barrême area (SE France). New biochronostratigraphic data on the Bigotitinae, youngest members of Leptosphinctinae and oldest members of Zigzagiceratinae are relevant in understanding the evolution and faunal turnover of the West Tethyan Perisphinctidae during earliest Bathonian. The ammonite succession at the Bajocian/Bathonian boundary in the Cabo Mondego region (Portugal) represents one of the most complete biostratigraphic records so far recognized on the Iberian Plate.     

New data on Late Jurassic sauropods of central and northern sectors of the Bombarral Sub-basin (Lusitanian Basin,Portugal)

The Upper Jurassic’s central and northern sectors of the Bombarral Sub-basin are relatively poor in sauropod material, highlighting the specimens (mainly teeth) found in the Guimarota mine (Leiria) and the Andrés (Pombal) fossil site. The study of published and the unpublished sauropod material allows for a revision of the present state of sauropod diversity of the Bombarral Sub-basin. These new specimens come from Pombal, Leiria, Batalha, Porto de Mós, Alcobaça and Caldas da Rainha, and include an almost complete posterior or middle dorsal neural spine and a partial caudal series. The systematic re-evaluation of the sauropod record of this sector indicates the presence of turiasaurs, diplodocines, titanosauriforms and an indeterminate eusauropod form. During the last part of the twentieth century, the discovery of fossil vertebrates has increased significantly in the sediments cropping out in the central and northern sectors of the Bombarral Sub-basin (Alcobaça and Bombarral Formations), improving our understanding of the Late Jurassic faunas of the Lusitanian Basin.     

New evidence of Ceratosaurus (Dinosauria: Theropoda) from the Late Jurassic of the Lusitanian Basin,Portugal

A theropod assigned to Ceratosaurus was previously reported from the Portuguese Lusitanian Basin based on a limited number of elements of a single individual. Here, we describe newly discovered elements that likely pertain to same, earlier described, specimen. The new elements provide additional evidence that the range of Ceratosaurus spanned from what is now North America into Europe. Previously, some differences were noted between the Portuguese specimens and the North American Ceratosaurus. We consider these differences to be trivial and attribute them to individual variation and/or ontogeny. The following set of features (lesser trochanter positioned low on the femur; crista tibiofibularis obliquely oriented with respect to the axis of the femoral shaft; infrapopliteal ridge present posteriorly on the femur; large cnemial crest; and medial condyle of the tibia continuous with proximal end) indicate that the Portuguese specimen is assignable to Ceratosaurus. This record constitutes one of the scarce evidence of basal ceratosaurian theropods in the Late Jurassic of Europe. Despite the abundance, diversity and wide geographical distribution of ceratosaurs during the Late Cretaceous, its early evolutionary history remains poorly understood. The Portuguese specimens constitute an important evidence for the knowledge of the paleobiogeographic evolution of the clade during the Late Jurassic.     

The Early Jurassic palynostratigraphy of the Lusitanian Basin,western Portugal

A comprehensive investigation of the Early Jurassic stratigraphical palynology of the Lusitanian Basin in western Portugal was undertaken, with most emphasis placed on dinoflagellate cysts. A total of 214 samples from an upper Sinemurian to upper Toarcian composite section based on six successions were examined. The Sinemurian material examined was barren of dinoflagellate cysts; however, the Pliensbachian and Toarcian successions are characterised by relatively low diversities where Luehndea spinosa, Mancodinium semitabulatum, Mendicodinium microscabratum, Nannoceratopsis gracilis, Nannoceratopsis senex, and Scriniocassis priscus are relatively common and biostratigraphically significant. Luehndea spinosa dominates the lowermost Toarcian (Dactylioceras polymorphum ammonite Biozone), and is an index species. At the base of the Hildaites levisoni ammonite Biozone, the effects of the Toarcian Oceanic Anoxic Event (T-OAE) caused Luehndea spinosa to become extinct. At the same time, dinoflagellate cyst abundance and diversity markedly decreased. After the T-OAE, during the middle and late Toarcian, phytoplankton recovery was prolonged and slow in the Lusitanian Basin. The Luehndea spinosa and Mendicodinium microscabratum dinoflagellate cyst biozones are defined, both of which are subdivided into two dinoflagellate cyst subbiozones.     

Ammonite-benthic Foraminifera turnovers across the Lower-Middle Jurassic transition in the Lusitanian Basin (Portugal)

This paper describes and characterises the co-occurrence of ammonite and benthic foraminiferal assemblages across the São Gião outcrop (Central Portugal), a reference section for the Lower-Middle Jurassic boundary in the Lusitanian Basin. The upper Toarcian-lower Aalenian marls and marly-limestones in this section provide a precise and detailed ammonite-based biostratigraphic zonation, with a mixed assemblage of northwest European and Mediterranean faunal elements, associated with benthic foraminifera assemblages with northern hemisphere affinities, both correlatable with the Aalenian GSSP at the Fuentelsaz section (Iberian Cordillera, Spain). A total of 447 well-preserved ammonite specimens and 13.116 foraminifera have been studied; no evidence was detected of any taphonomic processes that could have changed the original assemblages. From a biostratigraphic point of view, the ammonite record has enabled four biostratigraphic units to be recognised (the Mactra and Aalensis subzones of the Aalensis Biozone in the upper Toarcian, and the Opalinum and Comptum subzones of the Opalinum Biozone in the lower Aalenian). With regard to the benthic foraminifera, the taxa identified have enabled the Astacolus dorbignyi Zone and 11 bioevents to be identified, most of which representing local biostratigraphic proxies. However, the increase in the relative abundance of Lenticulina exgaleata Dieni from the upper part of the Opalinum Subzone to the lower part of the Comptum Subzone has a regional value. The constant and continuous ammonite record of northwest European taxa, together with typical Mediterranean taxa – namely Grammoceratinae – throughout the section, the high relative abundance of Miliolina representatives – generally interpreted as foraminifers typical of shallow waters – and the absence of foraminiferal forms typical of cool waters, do not support the inference of cool seawater temperatures attributed to the Early Aalenian, or the global character of the “Comptum cooling event”, at least with reference to the Lusitanian Basin.     

Spatial distribution of Late Pliensbachian (Early Jurassic) calcareous nannofossils within the Lusitanian basin (Portugal)

Quantitative analyses of Pliensbachian calcareous nannofossils have been carried out on a proximal-distal transect in the Lusitanian Basin (Portugal). The studied sections Vale Venteiro near Tomar and Peniche represent proximal and distal environments with respect to the emerged land of the Iberian Meseta (to the East). The upper portion of the Vale das Fontes Fm (Davoei and Margaritatus ammonite Zones) is studied in both sections and correlated by means of ammonite and nannofossil integrated biostratigraphy. A careful analysis of the preservation state of nannofossils is performed. Preservation state is moderate to good in the two settings; changes in nannofossil assemblages are therefore considered as primary. Samples were analysed for nannofossil absolute and relative abundances, species diversity and wt%CaCO₃. The analysis of calcareous nannofossils (absolute abundance, percentage, average abundance) shows changes in the community structure in space (i.e., from proximal to distal), and vertically within the water column. This pattern suggests a partitioning of habitats within the photic zone, and with respect to emerged lands. Proximal environments within the Lusitanian Basin were probably more eutrophic, because of the proximity of emerged lands to the East (Iberian Meseta). This was the source area for nutrients delivered into the basin by river transport. Distal environments were likely characterized by a greater water depth and by a deeper light penetration leading to a relatively more expanded photic zone. The assemblages recorded in the proximal setting exhibit high mean relative abundance of placolith-bearing coccolithophorids (Lotharingius spp., Biscutum spp. and Similiscutum spp.) and of Schizosphaerella spp., while distal assemblages are dominated by Crepidolithus crassus and Schizospharella spp. Crepidolithus crassus is interpreted as a deep-dweller coccolithophorid, inhabiting preferentially distal and deeper settings in the Lusitanian Basin. Placolith-bearing coccolithophorids were more abundant in proximal settings with respect to emerged lands, under relatively elevated trophic conditions. The probable calcareous dinocyst Schizospharella spp. proliferated in surface waters of both proximal and distal environments.     

First evidence of stegosaurian Deltapodus footprints in North Africa (Iouaridène Formation, Upper Jurassic, Morocco)

Abstract:  New findings of dinosaur footprints are described from the Upper Jurassic Iouaridène ichnosite of Morocco. On the top of two surfaces, stratigraphically close to that bearing the famous Breviparopus taghbaloutensis trackways, two footprints were excavated and assigned to the ichnogenus Deltapodus . This ichnogenus is well known from the Middle Jurassic of Yorkshire and also occurs in Upper Jurassic deposits from Iberia and the United States. This finding represents the first record of Deltapodus from Africa. These footprints, probably produced by stegosaurian dinosaurs, add new data on the distribution of this type of dinosaur and on the connection between the northern and southern margin of Tethys. 3D models have been generated to allow more detailed studies and to record these unique footprints.     

Hoof-Like Unguals,Skin, and Foot Movements Deduced from Deltapodus Casts of the Galve Basin (Upper Jurassic-Lower Cretaceous,Teruel, Spain)

Galve (Teruel, Spain) is a town in the interior of a synclinal fold with Upper Jurassic marine limestones along its flanks, and, in its core, Upper Jurassic–Lower Cretaceous continental and shoreline sediments crop out. The core sediments cover an area about 8 km², and contain a concentration of sites with footprints, bones, and eggshells of dinosaurs. The footprints are both shafts and natural casts. Some casts are attributed to stegosaurs (Deltapodus). The Deltapodus casts are characterized by features that allow us to make direct observations on the skin formed by polygonal scales, and ellipsoidal “hooves,” as well as deductions on the movement of the limbs during walking. According to the opinion of some authors, dinosaur footprints are indicators of the motion of their limbs and sometimes of the whole body. So far, results have been deduced from theropod, ornithopod, and sauropod footprints. This article shows the results obtained from analis of the aforementioned Deltapodus casts, i.e., forelimb movement similar to that of the forelimbs of sauropods, and the rigid structure of the autopodial part of the hind limb.     

Taxonomic and biostratigraphic re-evaluation ofPerisphinctes internispinosus Krantz, 1926 (Upper Jurassic, Ammonoidea)

The lectotype ofPerisphinctes internispinosus Krantz, 1926, type species ofWindhauseniceras Leanza, 1945, from the upper Middle Tithonian of Cerro Lotena (Neuquén-Mendoza Basin, Argentina) is designated. The type series, housed in the Paläontologisches Institut der Universität Bonn, was originally illustrated by hand-drawings of two of the eleven specimens. The lectotype is illustrated by a photograph for the first time. The specimen is probably a large microconch; the most likely corresponding macroconch appears to beH. aff.steinmanni (Steuer, 1897) from Cerro Lotena.Windhauseniceras internispinosum is the index and guide species of theInternispinosum Biozone, upper Middle Tithonian, most probablyPonti Zone in age. The occurrence in Arroyo del Yeso (Mendoza) below a succession ofMicracanthoceras species includingM. mirum (Leanza, 1945) which is very close toM. microcanthum, strongly supports this correlation. The species occurs throughout the Neuquén-Mendoza Basin and probably also in the Tarapacá Basin, Northern Chile.     

A new Xinjiangchelyid turtle (Testudines, Eucryptodira) from the Jurassic Qigu Formation of the southern Junggar Basin, Xinjiang, north-west China

A new eucryptodiran turtle, Xinjiangchelys qiguensis sp. nov. from the Upper Jurassic (Oxfordian — ?Kimmeridgian) Qigu Formation of the southern Junggar Basin (north‐west China) is described. The type material consists of a partial skeleton, including the complete carapace, plastron, nearly all cervical vertebrae, both scapulae, the pelvis and one ulna. It is clearly identifiable as a basal eucryptodire since it lacks the mesoplastron. It is distinguished from other species of Xinjiangchelys by several autapomorphies of the carapace and plastron, such as the first and fifth vertebrals extending on the peripherals, the plastron with three pairs of gulars, and an intergular which does not contact the hyoplastron. In the postcranium, the scapula with a long acromial and a small scapular process, the pelvis with a short ilial shaft and the elongated cervical vertebrae are characteristic. A new phylogenetic analysis of the in‐group phylogeny of the Xinjiangchelyidae is proposed and discussed, resulting in a new classification of the family. Xinjiangchelys (Toxocheloides) narynensis is regarded as a nomen dubium. Shartegemys is referred to Xinjiangchelys, whereas the holotypes of ‘Plesiochelys’chungkingensis and ‘P’. latimarginalis are excluded from the genus Xinjiangchelys but included in the Xinjiangchelyidae.     

A modern-type Kimmeridgian reef (Ota Limestone,Portugal): Implications for Jurassic reef models

Upper Jurassic reefs rich in microbial crusts generally appear in deeper (sponge—‘algal’ crust reefs) or in very shallow but protected settings (coral or coral-coralline sponge meadows with ‘algal’ crusts). Upper Jurassic high-energy reefs (coral reefs and coral-stromatoporoid reefs) normally lack major participation of microbial crusts but rather represent huge bioclastic piles with only minor framestone patches preserved. An exception to this rule is represented by the high-energy, coral-‘algal’ Ota Reef from the Kimmeridgian of the Lusitanian Basin (Portugal). The narrow Ota Reef tract rims a small intra-basinal carbonate platform exhibiting perfect facies zonation (from W to E: Reef tract, back reef sands, peritidal belt, low-energy shallow lagoon). The reef is dominated by massive corals (Thamnasteria, Microsolena, Stylina). Complete preservation of coral framework is rare: like other Upper Jurassic high-energy reefs, the Ota Reef is very rich in debris; however, this debris is largely stabilized by algal and microbial crusts, what contrasts the other examples and gives the Ota Reef the appearance of a typical modern high-energy coral-melobesioid algal reef. Further similarities to modern reefs are the likely existence of a spur-and-groove system, the perfect sheltering of inner platform areas and the occurrence of small islands, as indicated by local blackenings and early vadose and karstic features.     

Sequence stratigraphic analysis and sea-level history of the Upper Jurassic deposits (Mozduran Formation), south of Aghdarband,NE Iran

Abstract

The Kopet-Dagh Basin is a large sedimentary basin in northeastern Iran that host the giant Khangiran and Gonbadli gas fields. The Mozduran Formation with its various sedimentary facies is an important reservoir widely distributed in the basin. A sedimentological analysis of Upper Jurassic Mozduran Formation resulted in an accurate reconstruction of the sedimentary environments and the sequence stratigraphic framework south of Aghdarband. The strata consist of six different facies associations including 12 carbonate, one evaporate and two siliciclastic subfacies. On the basis of their various components, structural and textural characteristics, these facies were deposited on a homoclinal ramp in tidal flat to open marine environments ranging from supratidal to subtidal settings. Facies A1 and A2 represent open marine, B1-B4 Shoal, C1-C4 lagoonal and D1, D2, E, T1 and T2 tidal flat and Salina environments. In addition, based on detailed field and laboratory studies on the facies architecture, several large-scale (long-term) depositional sequences could be distinguished in the stratigraphic sections of the study area. These sequences are composed of LST, TST and HST that are separated by a SB1 and SB2 sequence boundaries. The paleogeography of the study area during the Late Jurassic time is reconstructed in five block diagrams.     

Microtaphonomy of bioclasts and paleoecology of microencrusters from Upper Jurassic spongiolithic limestones (External Prebetic,Southern Spain)

Microencrusters and microtaphonomic features of the Oxfordian spongiolithic limestones of the External Prebetic were studied using thin-section analysis. The spongiolithic limestone is a bioclastic-rich packstone with common echinoderm, mollusc and brachiopod remains. The bioclasts show a high fragmentation index and frequent microborings. The encrustation index (E _i) is higher for fragments of serpulids, ammonoids and bivalves, and increases with the initial grain-size of bioclasts. The main microencrusters consist of benthic microbial communities (BMC) and nubeculariids, as well as subordinate calcareous and siliceous agglutinated foraminifera, serpulids and bryozoans. BMC are usually the first colonizers, and encrusting foraminifera mainly appear on bioclasts larger than 2 mm. BMC dominate in well-developed encrustations on upward facing surfaces of larger bioclasts that are also colonized by foraminifera (nubeculariids and Subdelloidina). Bullopora, serpulids and bryozoans are more common on lower surfaces. The fact that the values of encrustation index, encrustation thickness and diversity of the microencrusters increase with the size of bioclasts is related to a higher stability and exposure time of the available bioclastic substrate. The microencruster distribution on upper and lower surfaces of large bioclasts may be related to photic control, space competition and/or predation avoidance.     

An ammonite–fish association from the Kimmeridgian (Upper Jurassic) of La Rochelle, western France

A partially disarticulated actinopterygian fish preserved in a large three-dimensional ammonite body chamber is described from the Kimmeridgian of western France. Taphonomic observations on the degree of preservation of the fish and the development of epibiont organisms on the inner wall of the shell indicate a rather long time interval before sediment totally filled the body chamber. The fish, referred to an indeterminate Macrosemiidae, probably used this empty ammonite ( Rasenioides , Aulacostephanidae) shell as a refuge, or possibly for spawning and/or brooding. It can be assumed that ammonite shells may have constituted common shelters for demersal fishes living in an open-marine shelf environment, near to a muddy bottom devoid of rocks.     

Cranial anatomy,taxonomic implications and palaeopathology of an Upper Jurassic Pliosaur (Reptilia: Sauropterygia) from Westbury,Wiltshire, UK

Abstract: Complete skulls of giant marine reptiles of the Late Jurassic are rare, and so the discovery of the 1.8‐m‐long skull of a pliosaur from the Kimmeridge Clay Formation (Kimmeridgian) of Westbury, Wiltshire, UK, is an important find. The specimen shows most of the cranial and mandibular anatomy, as well as a series of pathological conditions. It was previously referred to Pliosaurus brachyspondylus, but it can be referred reliably only to the genus Pliosaurus, because species within the genus are currently in need of review. The new specimen, together with another from the same locality, also referred to P. brachyspondylus, will be crucial in that systematic revision, and it is likely that the genus Pliosaurus will be found to include several genera. The two Westbury Pliosaurus specimens share many features, including the form of the teeth, but marked differences in the snout and parietal crest suggest sexual dimorphism; the present specimen is probably female. The large size of the animal, the extent of sutural fusion and the pathologies suggest this is an ageing individual. An erosive arthrotic condition of the articular glenoids led to prolonged jaw misalignment, generating a suite of associated bone and dental pathologies. Further damage to the jaw joint may have been the cause of death.