Constraining the age of the last marine sediments in the late Cretaceous of central south Pyrenees (NE Spain): Insights from larger benthic foraminifera and strontium isotope stratigraphy

The uppermost Cretaceous (upper Campanian–Maastrichtian) marine deposits of the central south Pyrenees host a rich larger benthic foraminiferal fauna and several rudist-rich levels. These marine deposits are directly overlain by the continental facies of the Arén and Tremp Formations, which are famous for their fossil dinosaur remains. Larger benthic foraminiferal distribution documents an important faunal turnover in all the carbonate platform environments within the photic zone, from open marine to littoral areas. Biostratigraphy indicates that this turnover occurred close to the Campanian-Maastrichtian boundary. This is also confirmed by strontium isotope stratigraphy which indicates an earliest Maastrichtian age for the appearance of the larger benthic foraminiferal assemblage constituted by Lepidorbitoides socialis, Clypeorbis mammillata, Wannierina cataluniensis, Orbitoides gruenbachensis, Siderolites aff. calcitrapoides, Fascispira colomi, Omphalocyclus macroporus and Laffiteina mengaudi. In particular, a numerical age of 71 Ma is obtained for the Hippurites radiosus level, just a few meters below the first continental deposits of the Arén sensu stricto Formation. The youngest marine sediments of the central south Pyrenees are early Maastrichtian in age. This is also an important constraint for the age of the end-Cretaceous dinosaur fossil localities of the Tremp basin.     

Palynology of Upper Cretaceous (uppermost Campanian–Maastrichtian) deposits in the South Yellow Sea Basin, offshore Korea

Forty-seven samples from Upper Cretaceous sections penetrated by the Kachi-1 and Inga-1 wells in the South Yellow Sea Basin have been analysed for their spore and pollen content. Thirty-five species of 18 spore genera and 54 species of 28 pollen genera are documented. One new monotypic genus, Diporocolpopollenites, and its type species, D. kachiensis sp. nov., are erected, and Dilwynites Harris, 1965, and its type species, D. granulatus Harris, 1965, are emended. There are also three new combinations: Ephedripites eocaenicus (Selling, 1944), E. praeclarus (Chlonova, 1961), and Retitricolpites anguloluminosus (Anderson, 1960). Two palynological zones are erected: anAquilapollenites attenuatus Assemblage Zone, which encompasses deposits that are considered to be latest Campanian–Early Maastrichtian in age, and an Aquilapollenites eurypteronus Assemblage Zone for sections that have been dated as Late Maastrichtian. The assemblages are typical of the Yenisey-Amur Subprovince of the Aquilapollenites (floral) Province. Lowland floodplain to shallow, commonly mesotrophic, lacustrine environments of deposition are indicated. The climate was probably wet subtropical, with rainfall being somewhat higher during the Late Maastrichtian than through the latest Campanian–Early Maastrichtian.     

Morphometric analysis of the genus Omphalocyclus from the Late Cretaceous of Turkey: new data on its stratigraphic distribution in Mediterranean Tethys and description of two new taxa

Omphalocyclus is an orbitoidal taxon poorly known from the Late Cretaceous benthonic foraminiferal inventory of the Tethyan realm. Apart from its debatable diagnosis only in the late Maastrichtian of western Tethys, the genus has also been discovered in much older beds in association with Orbitoides and Lepidorbitoides having rather primitive developmental stages. The morphometric analysis of A-forms in successive assemblages, ranging in age from (late) Campanian to terminal Maastrichtian, enables the documentation of phylogenetic changes for the first time. The most conspicuous change is found to be the general increase in the size of the embryon, which on average doubles by the end of the Maastrichtian. This trend is followed by the increase in the number of epi-embryonic chamberlets, which is however, not as significant as the former parameter. Omphalocyclus in the stratigraphically lowermost populations has mainly three to four primary epi-embryonic and no accessory epi-embryonic chamberlets. With the introduction of radial stolons, which seems to have taken place in horizons referable to the Gansserina gansseri Zone, only several accessory epi-embryonic chamberlets arise from the tritoconch. Instead, epi-embryonic chamberlets become rather larger in size and they also cover a wider portion of the embryon along its thick outer wall. Considering the suitable change in embryon size, and also some other morphologic features in successive populations, two new species, O. anatoliensis sp. nov. and O. cideensis sp. nov. are erected in late Campanian and late Campanian-early Maastrichtian populations respectively. The most advanced specimens in the late Maastrichtian are attributed to the long-known species O. macroporus [Lamarck, J.B., 1816. Histoire naturelle des animaux sans vertèbres, vol. 2. Verdière, Paris, pp. 1–568]. Thus, a tentative subdivision scheme comprising three successive species, O. anatoliensis – O. cideensis – O. macroporus, is proposed.     

A primitive tube-bearing antillocaprinid rudist bivalve,Parasarcolites sohli sp. nov,from Jamaica and Puerto Rico,West Indies

The new antillocaprinid rudist bivalve Parasarcolites sohli sp. nov. is described from the Maastrichtian rocks of Puerto Rico (El Rayo Formation) and Jamaica (Sunderland Inlier). The species shows that all tubes, one of the diagnostic features of the genus, arise by the development of flanges on costae to roof over the intercostal space between adjacent costae. This is the first time that this feature has been unequivocally shown in an antillocaprinid rudist and suggests that tubes in other antillocaprinid rudists arose in a similar way. The species occurs in early Maastrichtian deposits of Puerto Rico and Jamaica, and represents a potential marker species in future biostratigraphic studies.     

Biostratigraphic and biogeographic implications of a hadrosaurid (Ornithopoda: Dinosauria) from the Upper Cretaceous Almond Formation of Wyoming,USA

The results of Barnum Brown's 1937 expedition to the Almond Formation of Wyoming consisted of two unidentified ceratopsian skulls and a partial hadrosaurid specimen (AMNH 3651). The hadrosaurid is here attributed to the Maastrichtian genus Saurolophus, verifying previous biostratigraphic correlations of this formation using ammonite zones. Fossiliferous lower Maastrichtian formations occurring latitudinally between those of Alberta, Canada, and southwestern Texas, USA, such as the Almond Formation, are essential for testing the effects and duration of apparent hadrosaurid faunal segregation earlier in the Campanian, and indirectly aiding in the placement of faunal boundaries that are currently unknown for the late Campanian. The discovery of Saurolophus in Wyoming, a close relative of the Campanian genus Prosaurolophus, affirms that the segregation of hadrosaurid faunas established in the late Campanian (～75 Ma) continued for at least 3 million years. Combining occurrences of Saurolophus from Mongolia and the Moreno Formation of California with those of Alberta, Canada, this genus appears to have had one of the largest geographic ranges of any equivalent clade of hadrosaurid dinosaur, although species level distributions are still uncertain.     

New Stigmaphronidae and Megaspilidae (Hymenoptera: Ceraphronoidea) from Canadian Cretaceous amber

Three new species within the stigmaphronid genus Tagsmiphron Engel and Grimaldi, 2009, and one new species within the megaspilid genus Conostigmus Dahlbom, 1858 are described from Upper Cretaceous (Campanian) amber originating at the Grassy Lake locality in Alberta, Canada. New taxa include Tagsmiphron spiculum sp. nov., Tagsmiphron leucki sp. nov., Tagsmiphron exitorum sp. nov., and Conostigmus cavannus sp. nov. The new Conostigmus species is a rare discovery. It is the third megaspilid species to be found in Cretaceous amber, with the two specimens described herein effectively doubling the number of known Mesozoic exemplars for the family. We provide the first comprehensive report of known Ceraphronoidea within Canadian amber, and contrast this against other Cretaceous amber assemblages, discussing the potential palaeobiogeographic and palaeoenvironmental implications of the Canadian amber assemblage.     

New Spanish Late Cretaceous eusuchian reveals the synchronic and sympatric presence of two allodaposuchids

The recently described clade Allodaposuchidae includes European eusuchian crocodyliforms restricted to the Late Cretaceous (Campanian and Maastrichtian). A new allodaposuchid crocodyliform is here described based on two specimens from the upper Campanian–lower Maastrichtian fossil site of Lo Hueco (Cuenca, Spain). This new taxon, Agaresuchus fontisensis gen. et sp. nov., is described by two complete skulls and a lower jaw associated with one of them. This new species can be distinguished unambiguously from Lohuecosuchus megadontos, the other allodaposuchid known from the same fossil site. The presence of two allodaposuchid crocodyliforms in Lo Hueco allows the recognition of the synchronic and sympatric existence of two representatives of this clade for the first time. The new genus Agaresuchus, comprises a previously described Iberian allodaposuchid species, “Allodaposuchus” subjuniperus, as Agaresuchus subjuniperus, new combination.     

Charophytes from the Cretaceous – Paleogene transition in the Pingyi Basin (Eastern China) and their Eurasian correlation

Charophytes from uppermost Campanian to Paleocene deposits in the Pingyi Basin (Shandong Province, Eastern China) are studied from the perspectives of taxonomy, paleoecology, biogeography, and biostratigraphy. The taxonomy of charophytes used by previous authors is revisited based on a study of intraspecific gyrogonite polymorphism, facilitating comparisons between China and Europe. A number of synonymies are proposed. Gobichara deserta is confirmed as a younger synonym of Microchara cristata. The genus Euaclistochara Z. Wang is shown to be a younger synonym of Lamprothamnium J. Groves. Charophyte assemblages from the Pingyi Basin were generally species-poor but showed a high degree of variation depending on the paleoenvironment. During the latest Campanian–Maastrichtian, brackish water assemblages were monospecific, formed by Feistiella anluensis. Permanent lakes were dominated by Microchara cristata and Peckichara praecursoria. Lamprothamnium ellipticum and Mesochara voluta inhabited the overbank ponds near braided rivers. The Paleocene was much more homogeneous and was dominated by Peckichara varians in permanent lakes. A new biozonation is proposed which encompasses two biozones based on species with broad paleoecological requirements and a Eurasiatic distribution. These are the Microchara cristata biozone starting in the latest Campanian and lasting at least until the earliest Danian and the Peckichara varians biozone encompassing the late Danian–earliest Eocene. These biozones allow direct correlation between Chinese and European basins.     

Clypeorbis? ultima n. sp. from the uppermost Maastrichtian of Austria: The youngest representative of the Clypeorbinae Sigal, 1952 (calcareous benthic foraminifera)?

A second possible species of the genus Clypeorbis Douvillé [type-species C. mammillata (Schlumberger)] is described from the topmost Maastrichtian of the Kambühel Formation of the Northern Calcareous Alps, Austria. This comparatively small-sized, asymmetric calcareous benthic foraminifer is characterized by its rather large biloculine embryonic apparatus and a reduced ventral umbo. It occurs in a mixed calcilithic–bioclastic littoral facies together with orbitoids, rotaliids, Siderolites, and other mainly calcareous benthic foraminifera as well as red algae. The stratigraphy is constrained by means of associated planktonic foraminifera, indicating the latest Maastrichtian age (CF-3 hariaensis zone). Clypeorbis? ultima n. sp. might represent the youngest representative of the Clypeorbinae Sigal that became extinct at the Cretaceous–Paleogene boundary.     

Integrated biostratigraphy and palaeoenvironmental interpretation of the Upper Cretaceous to Paleocene succession in the northern Moldavidian Domain (Eastern Carpathians,Romania)

This study of the upper Maastrichtian to Danian sedimentary succession from the northern part of the Romanian Eastern Carpathians (Varniţa section) aims to establish an integrated biostratigraphy based on calcareous nannofossils, organic-walled dinoflagellate cysts (dinocysts) and foraminiferal assemblages, and to reconstruct the depositional environments of the interval. The stratigraphic record across the studied section is incomplete, considering that an approximately 16 m thick strata interval from the top of the Maastrichtian to lowermost Danian cannot be analyzed due to a landslide covering the outcrop. The upper Maastrichtian is marked by a succession of biostratigraphic events, such as the First Appearance Datum (FAD) of the nannoplankton taxon Nephrolithus frequens and FAD of the dinocyst species Deflandrea galeata and Disphaerogena carposphaeropsis, and the Last Appearance Datum (LAD) of Isabelidinium cooksoniae in the lower part of the section. These bioevents are followed by the LAD of the Dinogymnium spp. and Palynodinium grallator dinocyst markers in the top of the Maastrichtian deposits analyzed. In terms of foraminiferal biostratigraphy, the upper Maastrichtian Abathomphalus mayaroensis Zone is documented in the lower part of the studied section. Some bioevents, such as the bloom of the calcareous dinoflagellate genus Thoracosphaera and the FAD of the organic-walled dinocysts Damassadinium californicum, Senoniasphaera inornata, Xenicodinium lubricum and X. reticulatum suggest an early Danian age for the middle part of the section. From the Danian deposits in the Varniţa section, we describe a new organic-walled dinocyst species, Pentadinium darmirae sp. nov., which is until now the only species of the Pentadinium genus discovered in the Paleocene. The occurrence of the global Danian dinocyst marker Senoniasphaera inornata in the top of the section, suggests an age not younger than middle Danian (62.6 Ma) for the analyzed deposits.The palynofacies constituents, as well as the agglutinated foraminiferal morphogroups, used to reconstruct the depositional environments, show that the late Maastrichtian sediments were deposited in an outer shelf to distal (bathyal) environment, followed by a marine transgression during the Danian.     

Upper Maastrichtian shallow marine environments and neoselachian assemblages in North Iberian palaeomargin (Castilian Ramp,Spain)

Upper Maastrichtian deposits formed in a nearshore subtidal environment within the Valdenoceda Formation (Castilian Ramp, North Iberian margin) are described together with two recently found selachian assemblages. Rare earth element concentrations (REE) have been used to assess the degree of taphonomic mixing and reworking, observing that it is minor or non-existent, and differences in degree of preservation and ecologic mixing can be explained by biostratinomic processes. The patterns of REE also helped to obtain a better understanding of the depositional environment, including the diagenetic history from burial to final degree of bone preservation.The fossil assemblages here described are close to that of the late Maastrichtian of Albaina (in the enclave of Condado de Treviño, Burgos), both in the Basque-Cantabrian Region, but their age may be slightly older (early late Maastrichtian). In total, the new assemblages consist of 17 taxa, assigned to 11 genera of shallow-water dwellers combined with individuals from the outer shelf. They represent cosmopolitan taxa (Squalicorax pristodontus, Serratolamna serrata and Rhombodus binkhorsti) together with local species (Rhinobatos echavei, Rhinobatos ibericus). Although there are not significant differences between Albaina and Quintanilla la Ojada faunas, the new assemblages add interesting taphonomic and geochemical information to the few existing uppermost Cretaceous deposits with fossil sharks in southwestern Europe.     

Planktonic foraminifera and calcareous nannofossils record in the upper Campanian-Maastrichtian pelagic deposits of the Malatya Basin in the Hekimhan area (NW Malatya,eastern Anatolia)

The uppermost Cretaceous (upper Campanian-Maastrichtian) pelagic successions from the Malatya Basin (NW Malatya, eastern Anatolia) were studied by 688 samples, which were collected from five stratigraphic sections in the Hekimhan area. The pelagic deposits conformably overlie rudist bearing shallow-water limestones and are overlain conformably by Maastrichtian dolomites and unconformably by Paleocene-Eocene deposits, respectively.The pelagic successions in the Hekimhan area comprise the Kösehasan Formation at the base and the Zorbehan Formation at the top and reach up to 1100 m in thickness. The Kösehasan Formation rests over the neritic rudist-bearing limestones of the Güzelyurt Formation along a sharp contact and consists mainly of flysch-type sandstone-mudstone alternation with complete and partial Bouma sequences. The carbonate content of abundant planktonic foraminifera and nannoplankton-bearing 980-m-thick succession increases upwards and the formation passes gradually to the clayey limestones and marlstones of the Zorbehan Formation to the top. Occurrences of nannoplankton Lithraphidites quadratus Bramlette and Martini and Micula praemurus (Bukry) in the first beds of the Kösehasan Formation indicate that the age of the Kösehasan Formation and overlying Zorbehan Formation is of late Maasthrichtian. Another late Maastrichtian taxa Cribrosphaerella daniae Perch-Nielsen and Arkhangelskiella maastrichtiana Burnett are observed from the lowermost part of the succession. Maastrichtian planktonic foraminifera such as Contusotruncana walfischensis (Todd) and Globotruncanita pettersi (Gandolfi) were recorded through the successions. Although planktonic foraminifera are diverse and abundant particularly in the Kösehesan Formation, index late Maasthrichtian species were not encountered. Campanian and Santonian-Campanian planktonic foraminifera, e.g. Radotruncana calcarata (Cushman) and Globotruncanita elevata (Brotzen), obtained particularly from the lower part of the succession and calcareous nannofossils such as Broinsonia parca parca Bukry, Reinhardtites anthophorus (Deflanre) and Eiffellithus eximius (Stover) are interpreted as reworked from older strata. Trace fossils are common throughout the succession.Rareness of planktonic foraminifera and nannoplankton in the uppermost part of the succession (Zorbehan Formation) indicates maximum shallowing of the latest Maastrichtian sea in this part of the basin. Rare echinoids, bivalves and ammonites are observed in that part of the sequence.The obtained data indicate that sediment accumulation rate of the pelagic deposits is rather high and about 27.5 cm/ky for this part of the basin. Changes in thickness of the formations along short distances in the five stratigraphic sections analysed in this study should be related to the diachroneity of the depositional and erosional events.     

An Upper Cretaceous (Campanian-Maastrichtian) actinopterygian fish assemblage from the marginal marine Adaffa Formation of Saudi Arabia

Actinopterygian remains have been recovered from Upper Cretaceous (lower Campanian to lower Maastrichtian) marginal marine deposits of the Adaffa Formation in northwestern Saudi Arabia. The fossils comprise gars (Lepisosteidae), pachycormids (cf. Protosphyraena sp.), indeterminate pycnodontiforms, enchodontid teleosts (cf. Enchodus sp.) and other indeterminate Teleostei. This assemblage is significant because it includes a novel occurrence for the Middle East (Pachycormidae) together with taxa (Lepisosteidae, Pycnodontiformes, Enchodontidae) that have been previously recorded from Late Cretaceous faunas elsewhere in the Mediterranean Tethyan region.     

Pseudorhapydionina bilottei sp. nov., an endemic foraminifera from the post-Cenomanian/Turonian boundary (Pyrenees,NE Spain)

A new species of the foraminiferal genus Pseudorhapydionina is reported from the Santonian shallow-water carbonate and mixed deposits of La Cova Unit, in the Montsec and Pedraforca thrust sheets (Southern Pyrenees, NE Spain). Pseudorhapydionina bilottei sp. nov. differs from the Cenomanian species of the genus in its larger test size and the number of chambers in its early planispiral-involute stage. The discovery of P. bilottei sp. nov in the Santonian indicates that the genus Pseudorhapydionina escaped extinction during the Cenomanian-Turonian boundary (CTB) event in the Pyrenean bioprovince. A new subfamily Pseudorhapydionininae is described.     

Hesperornithiformes (Aves: Ornithurae) from the Upper Cretaceous Pierre Shale,Southern Manitoba,Canada

A historical collection of hesperornithiform fossils from the Gammon Ferruginous, Pembina, and Millwood members of the Pierre Shale (Campanian, Upper Cretaceous) in southern Manitoba, Canada, was examined to revise their taxonomy. Only two species of Hesperornis have been recognized in previous studies on the Pierre Shale in Manitoba, but our study recognizes six species of two genera, including H. lumgairi sp. nov. H. regalis is the most common species but absent in the uppermost unit within the studied sequence. The result of this study supports the paleobiogeographic subdivision of the Campanian vertebrate fauna within the Western Interior Seaway, but not the faunal boundary that distinguishes the avian fauna of Manitoba from that of South Dakota and Kansas.     

Oldest amber species of Palaeoaphididae (Hemiptera) from Baikura (Taimyr amber)

Ambaraphis baikurensis sp. nov. (Hemiptera: Palaeoaphididae) is described from the mid-Cretaceous (Albian–lower Cenomanian) of northern Siberia (Taimyr amber, Baikura locality) on the basis of wing morphology, which is most similar to that of aphids of the genus Ambaraphis Richards, 1966, previously reported only from Campanian Canadian amber. The new species primarily differs by pterostigma shape. The composition of the Baikura fauna supports assigning it to the Albian–lower Cenomanian Ognevka Formation.     

The Late Cretaceous Pterophyllum (Bennettitales) in the North-East of Russia

The Late Cretaceous Pterophyllum species survived as relict elements in areas influenced by volcanism in the North-East of Russia. Two new species are described: P. philippoviae Gnilovskaya from the Turonian-Coniacian deposits of the Vetvinskaya Unit (Penzhina and Oklan Rivers interfluve, Magadan Region) and P. terechoviae from the Maastrichtian deposits of the Kakanaut Formation (Kakanaut River Basin, Koryak Upland). The upper boundary of Pterophyllum stratigraphic range is extended from the Turonian-Coniacian up to the K/Pg boundary. Pterophyllum terechoviae is the youngest member of the genus and probably the latest occurrence of Bennettitales in the Northern Hemisphere.     

The latest record of the extinct subfamily Eoptychopterinae (Diptera: Ptychopteridae) from Upper Cretaceous amber of North Carolina

A new genus and species of female phantom midge Neuseptychoptera carolinensis gen. et sp. nov. in the fossil subfamily Eoptychopterinae is described and illustrated; this ptychopterid is from Upper Cretaceous (Campanian) amber of North Carolina. This is the first member of Eoptychopterinae from North America and the first found in Campanian age amber. We propose to treat the extinct subfamily Eoptychopterininae Lukashevich, 1995, as a junior synonym of the subfamily Eoptychopterinae Handlirsch, 1906, syn. nov. The elongated mouthparts of the new species have broad labella, and these are functionally interpreted as feeders for honeydew, fermenting plant juices, and nectar rather than for biting.