A riparian plant community from the upper Maastrichtian of the Pyrenees (Catalonia,NE Spain)

Angiosperms began to colonise riparian habitats very soon in their evolutionary history, probably already in the Aptian, but it is still poorly known when flowering plants finally dominated entirely these kind of communities as they do in the present. A new fossil plant locality (Molí del Baró-1) from the upper Maastrichtian of the Southern Pyrenees is described in which meandering river facies represent one of the first riparian communities formed only by angiosperms. The fossil assemblage consists of abundant leaves, seeds, logs and sporomorphs. Angiosperms remains dominate in all these cases and the leaf sample is mostly composed of a new eudicot willow-like species, Saliciphyllum gaetei sp. nov., the palm Sabalites longirhachis and an helophytic monocot. Pollen remains suggest that the later belonged to Typhaceae. Most of these plant remains were parautochthonous and deposited in a pond formed in the accretional part of a meander loop. The locality of Molí del Baró-1 represents an unique plant fossil assemblage in the uppermost Cretaceous of southern Europe. It clearly differs from those reported in other Maastrichtian localities of the Pyrenees (Fumanya and South Isona) and from the Campanian-Maastrichtian of Austria and Romania. In addition, it reflects a surprisingly modern physiognomy for a Late Cretaceous riverine plant assemblage that was built up with willow-like plants, palms and reeds.     

A new Brachiosauridae Sauropod dinosaur from the lower Cretaceous of Europe (Soria Province,Spain)

A new dinosaur sauropod from the Golmayo Formation (upper Hauterivian-lower Barremian) in Soria province (Spain) is described as Soriatitan golmayensis gen. et sp. nov. The new material consists of one tooth, three dorsal vertebrae with ribs, a partial sacrum, five caudal centra, two caudal vertebral spines, one chevron, a humerus, an ulna, a radius, two partial ilia, two ischia, a fragment of pubis, and a partial femur. Cladistic analysis identified the material as belonging to Brachiosaruidae within Titanosauriformes. Sauropod material from the Golmayo Formation shares a diagnostic feature with Abydosaurus, Cedarosaurus, Tastavinsaurus and Venenosaurus including anteriorly deflected anterior–middle caudal neural spines and with Cedarosaurus a proximodistally straight lateral margin between the proximal head and the shaft of the humerus. Eight characters are potential autapomorphies in the sauropod from Spain, including the presence of a large rectangular ventral ridge below the preacetabular process of the ilium, the same length of the pubic and ischiadic blades in the ischium, and two lateral ridges in the lateral area of the deltopectoral crest of the humerus. The presence of Early Cretaceous brachiosarids in both, North America and Europe, give support to the hypothesis of a connection between the tectonic plates of these continents at some point during the Early Cretaceous.     

Spheroolithid eggshells in the Lower Cretaceous of Europe. Implications for eggshell evolution in ornithischian dinosaurs

Early Cretaceous spheroolithid eggshells have always been extremely scarce, unlike their putative egg layers, ornithopod dinosaurs, which are ubiquitous in Early Cretaceous ecosystems. In this paper, we re-evaluate eggshells previously assigned to the oofamily Elongatoolithidae, an oofamily related to derived theropod dinosaurs (including maniraptorans and oviraptorids), and reassign them to the oofamily Spheroolithidae, on the basis of the lack of a squamatic ultrastructure, and the presence of a prolatocanaliculate pore system and undulating extinction, typical of spheroolithid eggshells. We erect a new oogenus, Guegoolithus to include material previously reported as Macroolithus turolensis Amo-Sanjuan, Canudo and Cuenca-Bescós 2000, and new material from several lower Barremian localities of the Maestrazgo Basin in Spain. Guegoolithus eggshells exhibit an acicular radial ultrastructure in the lower part of the shell units, which develops into a tabular radial ultrastructure in the upper part, giving an appearance of a two-layered eggshell in thin sections that is not seen in SEM pictures. These features fill the gap between the two-layered prismatic eggshell of ceratopsians and the prolatospherulitic ornithopod eggshell of derived hadrosaurs.     

Perinates of a new species of Iguanodon (Ornithischia: Ornithopoda) from the lower Barremian of Galve (Teruel,Spain)

Although Iguanodon is one of the most abundant and well-known of Europe's dinosaur genera, fossils of young specimens are very rare. Indeed, the fossil record contains very few examples of the young of any non-hadrosaurid iguanodontian. Here we report the discovery of 13 Iguanodon perinates from the Lower Cretaceous of Galve (Teruel, Spain). The characteristics of an adult and juvenile found nearby show these perinates to belong to a new species: Iguanodon galvensis sp. nov. The histological and osteological features of these young animals indicate them to have been in their first year of life. The taphonomic features of their remains, plus the finding of clearly embryonic vertebrae alongside them, suggest the perinates of this species remained in the vicinity of their nests for some time, possibly congregating in nursery areas.     

A review of the Upper Campanian and Maastrichtian belemnite biostratigraphy of Europe

The conventional belemnite zonation of northwest Europe includes three Upper CampanianBelemnitellazones, two Lower MaastrichtianBelemnellazones, in addition to the lower Upper MaastrichtianBelemnitella juniorZone and upper Upper MaastrichtianBelemnella casimirovensisZone. These zones are critically assessed. Recent belemnite subdivisions of the Upper Campanian and Lower Maastrichtian are reviewed. The Upper Campanian of Norfolk has been subdivided into seven informalBelemnitellazones and subzones, and the Lower Maastrichtian of northwest Germany into six formalBelemnellazones. The timespan of the Early Maastrichtian zones is estimated and the duration varies from 0.12–0.60 Ma. It is shown that the base of the basal MaastrichtianBelemnella lanceolataZone is slightly diachronous, and the base of theB. casimirovensisZone is highly diachronous across Europe.     

A mosasaur,cf. Plotosaurus,from the upper Maastrichtian Quiriquina Formation in Central Chile

Here we describe a mosasaurid from the upper Maastrichtian Quiriquina Formation near the town of Cocholgüe about 30 km north of Concepción, central Chile. The specimen comprises a concretion preserving the rostral part of a pair of mandibular rami. The bulbous base of the teeth allows referral to cf. Plotosaurus sp. This is southernmost record of this piscivorous and pelagic taxon. The southern dispersal of mosasaurs during the late Maastrichtian may not only have depended from warm south-bound Equatorial currents but mainly from the nutrient-rich oceanic regime along an upwelling zone of cold abyssal water that mixed with the warm surface water, resulting in an enormous primary production. Apparently mosasaur dispersal depended more on food than on suitable temperatures or currents.     

Controls on upper Campanian–Maastrichtian chalk deposition in the eastern Danish Basin

Detailed facies analysis of a 350 m long core of upper Campanian–Maastrichtian chalk at Stevns Peninsula, eastern Denmark, shows that four mudstone and wackestone chalk facies account for close to 95% of the succession, and that bioturbated mudstone chalk alone accounts for nearly 55% of the sediment. Sedimentation took place in deep water, below the photic zone and storm‐wave base, and is characterized by decimetre to metre‐scale variations in facies and trace fossil assemblages indicating repeated shifts in depositional environment. Integration of facies with published data on sea‐surface temperature and accumulation rates suggests that sea‐surface temperature is the most important parameter in controlling stratification of the water column and thereby, indirectly, the observed variations in depositional facies. However, bioturbated mudstone chalk occurs in all stratigraphic levels independent of accumulation rates and sea temperatures and is interpreted to represent a very broad set of deep water environmental conditions with an ample supply of calcareous nannofossil debris and intense bioturbation. Longer term shifts in deposition are best expressed by distribution of clay, flint and bioturbated micro‐wackestone, bioturbated wackestone and laminated mudstone chalk facies, whereas the trace fossil assemblages appear less useful. The data set indicates overall shallowing over time with two distinctive events of clay influx to the basin during the late Campanian–earliest Maastrichtian and late Maastrichtian.     

Unusual forms of orbitoidal foraminifera in the Maastrichtian of Turkey

Maastrichtian orbitoidal foraminifera in Turkey show a variety of unusual features. They include remarkable specimens of their reproductive life cycle, lateral chambers within the embryo, teratological individuals (polyvalence), microboring activity within tests, equatorial chamber growth, and conical forms. These features provide evidence for Maastrichtian orbitoidal foraminiferal systematics, biological life cycle, palaeoecological conditions and fossilization. The growth of lateral and equatorial chambers is thought to have been controlled largely by genetic characteristics. Microboring activity is the result of their fossilization and life relationships with other micro-organisms such as Talpinella conicularia. Conical forms and polyvalence are thought to be related to sedimentological characteristics of the palaeoenvironment, such as currents. Abnormal growth of the equatorial chambers is accepted as one of the main distinguishing features of Orbitoides gruenbachensis.     

The chronology of dinosaur oospecies in south-western Europe: Refinements from the Maastrichtian succession of the eastern Pyrenees

The Late Cretaceous deposits of the continental Tremp Formation in the Vallcebre Syncline (South-eastern Pyrenees) provide an extensive egg record of dinosaurs. The parataxonomical study and analysis of multiple eggshell samples, the precise stratigraphical control of several sections and the time calibration of the abundant egg levels enable the establishment of a robust oospecies succession. The successive occurrence of three megaloolithid oospecies (Megaloolithus siruguei-Megaloolithus mamillare-Megaloolithus sp.) in the Early and Late Maastrichtian is well correlated with the magnetic polarity time scale throughout the chron 31. The replacement of Megaloolithus siruguei with Megaloolithus mamillare occurs around the reversal of chrons 31r-31n. A comparison with oospecies successions from Arc basin localities (France) allows the age calibration for such oospecies replacement to be confirmed. This age refinement implies that some of the stated boundaries for the proposed oospecies assemblages may change and that an in-depth revision of the age and magnetic calibration of some south European egg localities is required.     

The Last Pterosaurs: First Record from the Uppermost Maastrichtian of the Tremp Syncline (Northern Spain)

Pterosaur bones are for the first time reported from the Tremp Syncline(Spain) and the southern Pyrenees.They come from the Torrebilles-2 site(Isona,Lleida Province) in the upper Maastrichtian part of the Tremp Formation,which falls within the C29r magnetochron.Fragmentary long bones are referred to the Pterosauria based on their cortical thinness,micro-architecture,and histological features.They belong to very large to possibly giant individuals.The most complete specimen is possibly a partial right femur.The review of the Maastrichtian pterosaur record shows that only a few specimens can be referred to the C29r(which contains the K/Pg boundary).The identification of fragmentary remains could improve our knowledge about the abundance and distribution of these flying sauropsids just before the end-Cretaceous mass extinction.     

Pes shape variation in an ornithopod dinosaur trackway (Lower Cretaceous,NW Spain): New evidence of an antalgic gait in the fossil track record

Trackways can provide unique insight to animals locomotion through quantitative analysis of variation in track morphology. Long trackways additionally permit the study of trackmaker foot anatomy, providing more insight on limb kinematics. In this paper we have restudied the extensive tracksite at Barranco de La Canal-1 (Lower Cretaceous, La Rioja, NW Spain) focussing on a 25-m-long dinosaur (ornithopod) trackway that was noted by an earlier study (Casanovas et al., 1995; Pérez-Lorente, 2003) to display an irregular pace pattern. This asymmetric gait has been quantified and photogrammetric models undertaken for each track, thus revealing distinct differences between the right and the left tracks, particularly in the relative position of the lateral digits II–IV with respect to the central digit III. Given that the substrate at this site is homogenous, the consistent repetition of the collected morphological data suggests that differences recorded between the right and the left tracks can be linked to the foot anatomy, but more interestingly, to an injury or pathology on left digit II. We suggest that the abnormal condition registered in digit II impression of the left pes can be linked to the statistically significant limping behaviour of the trackmaker. Furthermore, the abnormal condition registered did not affect the dinosaur's speed.     

Marine connections in North America during the late Maastrichtian: palaeogeographic and palaeobiogeographic significance ofJeletzkytes nebrascensisZone cephalopod fauna from the Elk Butte Member of the Pierre Shale, SE South Dakota and NE Nebraska

The Elk Butte Member of the Pierre Shale of southeast South Dakota and northeast Nebraska yields a late Maastrichtian cephalopod fauna of nautiloids, belemnites and ammonites of theFeletzkytes nebrascensisZone, best known from the near-shore facies of the Fox Hills Formation. ThenebrascensisZone is the highest distinct marine assemblage that can be recognised in the Western Interior, although ammonites occur as rarities high in the Lance Formation in Wyoming. Elements of the fauna occur in the Gulf Coast and Atlantic Seaboard, and extend into the highest Maastrichtian nannofossil Subzone CC26b, ofMicula prinsii, in Texas. These occurrences point to the existence of a southerly marine route for migration into and out of the northern Interior during the late late Maastrichtian. An analysis of Maastrichtian ammonite occurrences in West Greenland reveals no evidence for a marine link to the western Interior at this time, but rather indicates an open marine link to the North Atlantic region.The presence of upper upper Maastrichtian Pierre Shale in southeast South Dakota and northeast Nebraska, deposited in water depths that are conservatively estimated at 100-200 m, suggests that marine conditions (evidence for which has been removed by post-Cretceous erosion) may have extended well to the north of the shoreline position indicated in recent palaeogeographic reconstructions.     

Lithostratigraphy and biostratigraphy of the Campanian–Maastrichtian Toyajo Formation in Wakayama, southwestern Japan

The Upper Cretaceous Toyajo Formation is distributed around the Mt. Toyajo in the Aridagawa area, Wakayama, southwestern Japan. The formation is subdivided into three newly defined members, the Nakaibara Siltstone Member, Hasegawa Muddy Sandstone Member, and Buyo Sandstone Member, in ascending order. Close field observation elucidated the detailed biostratigraphy of the Toyajo Formation, and high-precision biostratigraphic correlation was made with the Yezo Group in Hokkaido (northern Japan) and Sakhalin and the Izumi Group in southwestern Japan.The Toyajo Formation contains diversified lower Campanian to upper Campanian heteromorph ammonoid assemblages, including Eubostrychoceras and Scaphites. Discovery of the heteromorph fauna demonstrates that scaphitid ammonoids survived until Campanian time in the northwestern Pacific region. Although Eubostrychoceras elongatum has been known in the northeastern Pacific region, the occurrence of this species in the northwestern Pacific region has been uncertain before. The rich occurrence of E. elongatum in the Aridagawa area indicates that this species was distributed widely in the northern Pacific realm.The Toyajo Formation is similar to the Izumi Group in various geologic features, and may indicate that the Toyajo Formation was deposited in a strike-slip basin along the Chichibu Belt formed by the movement along the Kurosegawa Tectonic Zone in the latest Cretaceous, like the Izumi Group, along the Median Tectonic Line.     

New evidence for the possible occurrence of Tyrannosaurus in West Texas,and discussion of Maastrichtian tyrannosaurid dinosaurs from Big Bend National Park

Speculation regarding Tyrannosaurus in West Texas has been largely based upon a sub-adult tyrannosaurid maxilla from the Javelina Formation (Late Cretaceous–Maastrichtian) of Big Bend National Park. However, a very large anterior caudal vertebra, recently collected from the Javelina Formation, exhibits a morphology that can confidently be assigned to Tyrannosauridae and, because of its size, likely pertains to an adult Tyrannosaurus. The stratigraphic position of the specimen is closely bracketed by titanosaurid remains and further supports coexistence of these taxa. The stratigraphic position of the specimen possibly records one of the earliest occurrences of Tyrannosaurus. If so, Tyrannosaurus likely existed during roughly equivalent temporal intervals in disparate paleobiomes in both northern and southern late Maastrichtian faunal realms of North America.     

Integrating outcrop data and forward computer modelling to unravel the development of a Messinian carbonate platform in SE Spain (Sorbas Basin)

Geological mapping, definition of facies distributions and reconstruction of platform‐interior growth geometries of the Messinian Cariatiz carbonate platform (Sorbas basin, South Spain), were performed to evaluate the controlling factors in platform growth and to test a 3‐D computer simulation program. For the simulation with the program REPRO, five platform‐related facies were modelled: (1) the reef crest facies by the numerical solution of a Fisher equation; (2) the lagoonal facies by a function of water depth‐dependent carbonate production; (3) the proximal and middle slope facies (breccia and block facies, calcarenite facies) by a subroutine simulating gravity‐driven particle export from the reef crest; (4) a distal slope; and (5) a basinal facies by a pelagic rain function. Development of a fan delta conglomeratic system is simulated by using a siliciclastic point source and gravity‐driven particle redistribution. A best fit between the observed platform growth geometries and modelling results is achieved by assuming that high‐frequency sea‐level changes superimposed onto a longer term sea‐level fall controlled platform growth. For the modelling, a relative sea‐level curve was reconstructed, which is based on a deep‐sea benthic foraminiferal stable oxygen isotope record at ODP Site 926 with a 45 m eustatic sea‐level fall, and a tectonic uplift component of 20 m. The consistency of 3‐D simulation results is corroborated by the coral growth rates provided by the Fisher‐equation subroutine. These rates of 2–8 mm year⁻¹ compare well to the coral growth rates in Recent fringing reefs. We propose that during the early stage of platform evolution the high‐frequency fluctuations were obliquity‐modulated precessional cycles, whereas precessional cycles control later stages of platform growth. REPRO provides a separate visualization of the different facies bodies as a function of time and space, showing the intrinsic pattern of facies distribution in the platform. This is the result of a combination of platform growth and syndepositional subaerial erosion. For example, only the youngest stages of reef framework facies in the development of the Cariatiz carbonate platform are preserved.     

Laminated–bioturbated cycles in Maastrichtian chalk of the North Sea: oxygenation fluctuations within the Milankovitch frequency band

The Maastrichtian chalk of the southern Central Graben, Danish North Sea, is a homogeneous pure white coccolithic chalk mudstone deposited in a deep epeiric shelf sea, which covered large parts of northern Europe. The sediment displays a pronounced cyclicity marked by decimetre‐thick bioturbated beds alternating with slightly thinner non‐bioturbated, mainly laminated beds. The laminated half‐cycles consist of alternating millimetre‐thick, graded, high‐porosity laminae and non‐graded, low‐porosity laminae. The cyclicity has been interpreted previously as caused by periods of slow background sedimentation and bioturbation interrupted by periods of rapid deposition of laminated beds, with the latter reflecting random and local resedimentation processes. Based on textural and structural analysis, the millimetre‐scale, non‐graded laminae are interpreted as having been deposited directly from pelagic rain of pelleted coccoliths representing the primary production. The graded laminae were deposited from small‐volume, low‐density turbidity currents and suspension clouds. The sedimentation rates of the cyclical chalk are similar to those known elsewhere, and the lamination is interpreted as having been preserved from destruction through bioturbation by anoxic conditions at the seafloor. Bioturbated–laminated cycles are thus formed by slow sedimentation during alternating seafloor redox conditions probably on a Milankovitch scale. A direct implication of this interpretation is that the cycles are areally widespread, probably extending throughout the southern Central Graben area and may be useful for correlation and high‐resolution cyclostratigraphy in the chalk fields of the Danish North sea. If the laminated half‐cycles represent a few rapid resedimentation events, with a high sedimentation rate as suggested by most workers, then the sediment would not be truly cyclic, but would represent event sedimentation within a pelagic background represented by the bioturbated beds. In this case, the cycles would have very limited potential for correlation.     

四川盆地北缘发现白垩纪恐龙足迹化石

2020年10月,笔者在四川盆地北缘的四川省通江县下白垩统苍溪组粉砂岩层面之上,新发现5个恐龙足迹化石,其形态均与经典的兽脚类足迹Eubrontes相似（如双足动物所留,足迹宽,功能性三趾型,较短的第Ⅲ趾）。该发现表明,Eubrontes这种中大型的兽脚类足迹广泛分布在四川盆地的下白垩统。该发现对于四川盆地北缘的古气候、古地理和地层对比都具有重要意义。     

陕西安塞首次发现中侏罗世晚期恐龙足迹

在安塞地区野外地质考察过程中, 在直罗组顶部的一套砂岩地层中发现4枚恐龙足迹, 其中3枚为三趾型恐龙足迹; 另外还观察到3枚疑似恐龙足迹。根据足迹形态对比和足迹特征研究, 推断P3观察点三趾型恐龙足迹造迹者为大型肉食性兽脚类恐龙, 其他足迹的造迹者难以确认。该发现不仅扩大了鄂尔多斯盆地中侏罗世恐龙足迹的分布范围, 也为研究直罗组晚期的古环境提供了素材。     

Sr and O isotope constraints on source and crustal contamination in the high-K calc-alkaline and shoshonitic neogene volcanic rocks of SE Spain

The Neogene volcanic province of SE Spain (NVPS) is characterized by calc-alkaline (CA), high-K calc-alkaline (KCA), shoshonitic (SH), ultrapotassic (UP), and alkaline basaltic (AB) volcanic series. All these series, except the AB, have high LILE/LREE, LILE/HFSE and B/Be ratios and high but variable Sr, Pb and O isotope compositions. The KCA and SH lavas contain metapelitic xenoliths whose mineralogical and chemical composition are typical of anatectic restites. The geochemical characteristics of CA, KCA, SH and UP series suggest that they originated from the lithospheric mantle, previously contaminated by fluids derived from pelagic sediments. Additionally, the presence of restite xenoliths in the KCA and SH lavas indicates some sort of interaction between the mantle-derived magmas and the continental crust. Trace element and isotope modeling for the KCA and SH lavas and the restites, point towards the existence of two mixing stages. During the first stage, the lithospheric mantle was contaminated by 1–5% of fluids derived from pelagic sediments, which produced a fertile source heterogeneously enriched in incompatible elements (particularly LILE and LREE), as well as in ⁸⁷Sr/⁸⁶Sr, without significant modifications of the δ¹⁸O values. In the second stage, the primary melts derived from this metasomatized mantle, which inherited the enrichment in LILE, LREE and ⁸⁷Sr/⁸⁶Sr, interacted with crustal liquids from the Betic Paleozoic basement during their ascent towards the surface. This mixing process caused an increase in δ¹⁸O values and, to a lesser extent, in ⁸⁷Sr/⁸⁶Sr ratios. However, the incompatible trace elements abundances only change slightly, even for high mixing rates, due to their similar concentrations in both components. We suggest the following geodynamic scenario to account for the global evolution of this area: (1) a Late Cretaceous to Oligocene subduction scheme during which mantle metasomatism took place, shortly followed by Upper Oligocene to Lower Miocene continental collision, and (2) a Middle to Upper Miocene extensional event triggering partial melting of the previously metasomatized mantle and the extrusion of the CA and associated magmas.     

Maastrichtian to Paleocene depositional environment of the Dakhla Formation, Western Desert, Egypt: sedimentology, mineralogy, and integrated micro- and macrofossil biostratigraphies

Integrated sedimentology, mineralogy, geochemistry, and microfossil and macrofossil biostratigraphies of the Maastrichtian–early Paleocene Dakhla Formation of the Western Desert, Egypt, provide improved age resolution, information on the cyclic nature of sediment deposition, and the reconstruction of depositional environments. Age control based on integrated biostratigraphies of planktic foraminifera, calcareous nannofossils and macrofossils yields the following ages for stratigraphic and lithologic sequences. The contact between the Duwi and Dakhla formations marks the Campanian/Maastrichtian boundary (zone CF8a/b boundary) and is dated at about 71 Ma. The age of the Dakhla Formation is estimated to span from 71 Ma at the base to about 63 Ma at the top (zones CF8a–Plc). The Cretaceous/Tertiary (K/T) boundary is within the upper unit of the Kharga Shale Member and marked by a hiatus that spans from 64.5 Ma in the lower Paleocene (base Plc) to at least 65.5 Ma (base CF2, base M. prinsii zones) in the upper Maastrichtian at Gebel Gifata, the type locality of the Dakhla Formation. As a result, the Bir Abu Minqar horizon, deposited between about 64.2 and 64.5 Ma (Plc(l) zone), directly overlies the K/T boundary hiatus. Major hiatuses also span the late Maastrichtian–early Paleocene in sections to the northwest (c. 61.2–65.5 Ma at North El Qasr, c. 61.2–69 Ma at Bir Abu Minqar and c. 61.2–65.5 Ma at Farafra), and reflect increased tectonic activity.During the Maastrichtian–early Paleocene a shallow sea covered the Western Desert of Egypt and the clastic sediment source was derived primarily from tectonic activity of the Gilf El Kebir spur to the southwest of Dakhla and the Bahariya arch. Uplift in the region resulted in major hiatuses in the late Maastrichtian–early Paleocene with increased erosion to the southwest. The area was located near the palaeoequator and experienced warm, wet, tropical to subtropical conditions characterized by low seasonality contrasts and predominantly chemical weathering (high kaolinite and smectite). A change towards perennially more humid conditions with enhanced runoff (increased kaolinite) occurred towards the end of the Maastrichtian and in the early Paleocene with shallow seas fringed by Nypa palm mangroves. Sediment deposition was predominantly cyclic, consisting of alternating sandstone/shale cycles with unfossiliferous shales deposited during sea-level highstands in inner neritic to lagoonal environments characterized by euryhaline, dysaerobic or low oxygen conditions. Fossiliferous calcareous sandstone layers were deposited in well-oxygenated shallow waters during sea-level lowstand periods.