New insights into skeletal morphology of the oldest known silicoflagellates: Variramus, Cornua and Gleserocha gen. nov.

Two of the oldest known silicoflagellate-bearing sediments, lower Albian at Ocean Drilling Program Site 693 in the Weddell Sea of Antarctica and Santonian in the Devon Island sequence of the Canadian Archipelago, are re-examined with a focus on silicoflagellate genera Variramus, Cornua, and Gleserocha that lack basal rings, a feature appearing during late Santonian/early Campanian time within the genus Corbisema. The extraordinary variability of Variramus aculeifera is studied, and a new genus Gleserocha is proposed for taxa with apical structures made of three struts, but without pikes. This new genus includes previously described Variramus wisei and Cornua tapiae and new species G. harrisonii, and is here proposed as a genus transitional between Variramus and Cornua. Two unusual new species of Cornua, C. deflandrei and C. witkowskiana, are also described and a revised terminology is proposed for the skeletal components of Cornua and closely related genera.     

Benthic foraminiferal paleoenvironmental reconstruction from the Upper Coniacian-Lower Campanian succession in north-western Tunisia

Quantitative analysis of benthic foraminifera is used to characterize the paleoenvironments of the Upper Coniacian-Lower Campanian succession in the Jbil section of north-western Tunisia. Foraminiferal parameters and benthic foraminiferal assemblages show that the studied section includes four distinct paleoenvironmental phases. From oldest to youngest, these are as follows: (1) an interval with a Praebulimina reussi assemblage with infaunal ratios as high as 96.1%. High abundances of P. reussi, reflecting an increase in organic matter flux to the seafloor (meso-to eutrophic) under oxygenated bottom-water conditions. (2) An interval characterized by a Gavelinella costulata assemblage with mixed infaunal/epifaunal foraminifera with higher Fisher's alpha values (ranging from 4 to 15.2), reflecting mesotrophic conditions in an outer shelf environment. (3) An interval with a Gaudryina laevigata assemblage indicative of a middle to outer shelf environment; there is a considerable increase in infaunal agglutinated foraminifera, as well as a relatively abundant and moderately diversified oxic/suboxic foraminifera. (4) The final interval occurs in the lower Campanian (the Globotruncana ventricosa Zone) and includes a Bolivinoides decoratus assemblage reflecting an outer shelf to upper bathyal environment. It contains a higher planktonic percentage and biodiversity with a slight increase in dysoxic species; the mixed infaunal/epifaunal content (57.6 to 73.3%) reflects mesotrophic conditions. Four well-recognized major sea-level falls are matched by the dual signatures of eustatic sea-level changes. These are coincident with the results of this study, which represent the first documentation of these events in Tunisian faunal and paleoenvironmental changes, at the following boundaries: Coniacian/Santonian, intra-Santonian, Santonian/Campanian, and intra-early Campanian.     

Integrated biostratigraphy for the Santonian/Campanian boundary interval,the Kurdistan Region,northeast Iraq

The first high-resolution integrated biostratigraphic study for Santonian/Campanian sediments of the Tabin section in the Kurdistan Region, northeast Iraq is provided. The study, based on 28 closely spaced samples, combines data from planktic foraminifers (25 species), calcareous nannofossils (32 species) and two ammonite genera in the Kometan Formation, marking the Santonian/Campanian boundary (S/C boundary) in the Kurdistan Region. In the absence of the crinoid Marsupites testudinarius, the proposed boundary marker, secondary markers such as calcareous nannofossils, planktic foraminifers and ammonites, have been used to establish a multi-stratigraphic biozonation for the late Santonian–early Campanian duration. Based on the occurrences of calcareous nannofossils, three biozones are identified — Lucianorhabdus cayeuxii (late Santonian), Calculites obscurus (latest Santonian–earliest Campanian), and Broinsonia parca parca (early Campanian). Seven calcareous nannofossil bioevents and three planktic foraminiferal bioevents are also identified. The Santonian/Campanian boundary is marked by: (a) the LO (Last Occurrence) of the planktic foraminifera D. asymetrica, (b) the FOs (First Occurrence) of the calcareous nannofossil species B. parca parca and B. parca constricta, (c) the extinction of several planktic foraminiferal species of Dicarinella and Marginotruncana, (d) the abundance and diversification of the planktic foraminifera genera, Globotruncana and Globotruncanita at the beginning of the Campanian, and (e) the disappearance of the ammonite genus Texanites, 0.5 m below (i.e., at 19 m) the disappearance of all Dicarinella and Marginotruncana species in the study section. Similar to several other Tethyan sections, the FO of B. parca parca is above the LOs of D. concavata and D. asymetrica; the LO of D. asymetrica is used here to mark the S/C boundary     

New latest Coniacian to middle Campanian foraminiferal data from the lower Zongshan Formation in the Chaqiela section,Gamba, southern Tibet

A first and detailed foraminiferal biostratigraphic work on the lower part of the Zongshan Formation (Limestone I and Calcareous Marl I sequence) in the Chaqiela section, Gamba, southern Tibet, allows the recognition of three latest Coniacian to middle Campanian planktic foraminiferal biozones: Dicarinella asymetrica Total Range Zone, Globotruncanita elevata Partial-Range Zone, and Contusotruncana plummerae Interval Zone. The base and top of the Santonian Stage in the Chaqiela section were placed at the lowest occurrence (LO) of Globotruncana linneiana and the highest occurrence (HO) of Dicarinella asymetrica, respectively. The deposition of the latest Coniacian to middle Campanian sediments of the lower Zongshan Formation in the Chaqiela section seems to have been continuous or at least without any major gap based on the planktic foraminiferal biozones and events.     

Turonian–Santonian radiolarians and planktonic foraminifers of the Upper Cretaceous sequences between Trabzon and Giresun areas in the Eastern Pontides,NE Turkey

In the “Eastern Pontides” (NE Turkey), the intense volcanic activity accompanying carbonate-siliceous sedimentation during the Late Cretaceous period ensured the occurrence of thick volcano-sedimentary sequences. Seventy-one radiolarian species belonging to 26 genera were identified from five stratigraphic sections of the Upper Cretaceous sequences between Trabzon and Giresun areas in NE Turkey. These species were obtained from the lower Turonian–lower Coniacian strata of the Çe?meler Formation, the Coniacian–lower Santonian strata of the Elmal? Dere Formation and the upper Santonian strata of the Ça?layan Formation. Coniacian–Campanian fauna, consisting of planktonic foraminifera Hedbergella, Marginotruncana, Globotruncana, Whiteinella and biserial heterohelicids as well as radiolarians, was recovered from the red pelagic limestones of the Elmal? Dere Formation known as marker horizon.     

The Pugnellidae, a new stromboidean family (Gastropoda) from the Upper Cretaceous

Members of the new family Pugnellidae have long been considered to represent the earliest members of the Strombidae but have more recently been included with the Aporrhaidae. We here propose them to represent a new family since their low-spired, bulgous shells are quite distinct from the usually slender and high-spired shell of the Aporrhaidae. Eleven species are described here belonging to six genera from which one genus and five species are new. The new genusBizarrus includes large, unornamentedGymnarus-like species, and new species are described asPugnellus klitzschi, Perustrombus africanus, Perustrombus indicus, Pugnellus?bartheli andPyktes popenoei. The here described Pugnellidae are from the Coniacian/ Santonian of Mungo River in Cameroon, the Santonian Umzamba Formation in the Eastern Cape Province of South Africa, the Campanian-Maastrichtian Ariyalur Group in Tamil Nadu, southern India, the Maastrichtian Quiriquina Formation of central Chile and the Maastrichtian Ammonite Hill Formation in the Western Desert of Egypt. Their radiation and possible phylogenetic relations to other Late Cretaceous stromboids are discussed. The origin of this family appears to lie in eastern Asia from where they spread through the Indopacific, Tethys and Atlantic Oceans during the Cenomanian-Santonian before they finally reached eastern North America in the Campanian.     

Planktonic foraminiferal biostratigraphy and paleoenvironments at the Santonian-Campanian boundary of Northern Tunisia

The Santonian-Campanian boundary in Northern Tunisia has been studied in Oued el Khannga section, based on biostratigraphic and paleoecological analysis. The detailed biostratigraphic analysis allows us to define the stratigraphic distribution of diverse planktonic foraminifera and to establish a continuous biozonation. Sixty-six planktonic foraminiferal species belonging to 13 genera have been identified and two Tethyan planktonic foraminiferal zones have been organized, indicating the upper Santonian - lower Campanian age. These zones are: Dicarinella asymetrica (Total Range Zone) and Globotruncanita elevata/Globotruncana arca (Concurrent range zone). A major turn over event was recorded in this interval including the extinction of several species characterized by complex morphotypes with a double keel such as the genera of Dicarinella, Marginotruncana. In fact, the extinction of the index species D. asymetrica is the most important event which defines the Santonian-Campanian boundary. Many representative species of the genera Globotruncanita and Globotruncana occurred, for the first time, in the uppermost of the Santonian and thrived close to the Santonian-Campanian boundary. In addition some surviving species with a simple morphotype, which are considered like opportunistic taxa, crossed the boundary. This various responses of planktonic foraminifera and the fluctuation in diversity and in the number of species may be related to an increase in temperature and a deep outer shelf upper bathyal environment deposit.     

Palaeoceanographic and palaeoclimatic inferences from Late Cretaceous planktonic foraminiferal assemblages from the Exmouth Plateau (ODP Sites 762 and 763, eastern Indian Ocean)

The evolution of planktonic foraminifera during the Late Cretaceous is marked in the Santonian by the disappearance of complex morphotypes (the marginotruncanids), and the contemporary increasing importance and diversification of another group of complex taxa, the globotruncanids. Upper Turonian to lower Campanian planktonic foraminiferal assemblages from Holes 762C and 763B (Ocean Drilling Program, Leg 122, Exmouth Plateau, 47°S palaeolatitude) were studied in detail to evaluate the compositional variations at the genus and species level based on the assumption that, in the Cretaceous oceans as in the modern, any faunal change was associated with changes in the characteristics and the degree of stability of the oceanic surface waters. Three major groups were recognised based on gross morphology, and following the assumption that Cretaceous planktonic foraminifera, although extinct, had life-history strategies comparable to those of modern planktonics: 1 – r-selected opportunists; 2 – k-selected specialists; 3 – r/k intermediate morphotypes which include all genera that display a range of trophic strategies in-between opportunist and specialist taxa. Although planktonic foraminiferal assemblages are characterised by a progressive appearance of complex taxa, this trend is discontinuous. Variation in number of species and specimens within genera has allowed recognition of five discrete intervals each of them reflecting different oceanic conditions based on fluctuations in diversity and abundance of the major morphotypes. Planktonic forms show cyclical fluctuations in diversity and abundance of cold (r-strategists) and warm taxa (k-strategists), perhaps representing alternating phases of unstable conditions (suggesting a weakly stratified upper water column in a mesotrophic environment), and well-stratified surface and near-surface waters (indicating a more oligotrophic environment). Interval 1, middle Turonian to early Coniacian in age, is dominated by the r/k intermediate morphotypes which alternate with r-strategists. These cyclical alternations are used to identify three additional sub-intervals. Interval 2, aged middle to late Coniacian, is characterised by the increasing number of species and relative abundance of k-strategists. After this maximum diversification the k-strategists show a progressive decrease reaching a minimum value in Interval 3 (early to late Santonian), which corresponds to the extinction of the genus Marginotruncana. In the Interval 4, latest Santonian in age, the k-strategists, represented mainly by the genera Globotruncana, increase again in diversity and abundance. The last Interval 5 (early Campanian) is dominated by juvenile globotruncanids and r-strategists which fluctuate in opposite phase. The positive peak (Interval 2) related to the maximum diversification of warm taxa (k-strategists) in the Coniacian seems to correspond to a warmer episode. It is followed by a marked decrease in the relative abundance of warm taxa (k-strategists crisis) with a minimum in the late Santonian (Interval 3), reflecting a decrease in temperature. Detailed analysis of faunal variations allows the Santonian faunal turnover to be ascribed to a cooling event strong enough to cause the extinction of the marginotruncanids.     

Chloranthaceous floral structures from the Late Cretaceous of Sweden

New chloranthaceous floral structures from the Late Cretaceous (Late Santonian/Early Campanian) of Scania, southern Sweden, have provided important new information on theChloranthistemon plants. The material includes well preserved fragments of inflorescence axes with flowers in situ documenting thatChloranthistemon flowers were bisexual and closely resembled those of extantChloranthus (Chloranthaceae). An emended diagnosis is given for the type species of the genus,Chloranthistemon endressii, and a new species,C. alatus, is described. The flowers ofChloranthistemon are small, perianthless and strongly zygomorphic, consisting of a tripartite and broadened androecium borne in an abaxial to lateral position on the monocarpellate ovary, and arranged in the axils of decussate bracts. Stamens are either completely free (C. alatus), or free at the base and coherent at the apex (C. endressii). The apical connective is extensive in both species; elaborated into conspicuous wing-like structures inC. alatus, or into a massive and shield-like structure inC. endressii. Pollen grains ofC. endressii are spheroidal, and reticulate and spiraperturate, while those ofC. alatus are ellipsoidal, tectate and foveolate with a unique combination of a distal colpus and a proximal furrow (colpus?) perpendicular to each other. Ovaries observed in well preserved flowers of both species are small and undifferentiated. Larger, dispersed fruits of chloranthaceous affinity are abundant and distinct, and probably represent at least two or three species, but cannot be linked with certainty to any of theChloranthistemon species described here.     

A new species of Phoenicopsis (Leptostrobales) from the Maastrichtian–Danian of Chukotka,Russia

A new species, Phoenicopsis anadyrensis Nosova, is described based on the leaf morphology and cuticle features from the Maastrichtian–Danian Rarytkin Formation, Chukotka, North-East of Russia. It is characterized by hypostomatic leaves with stomatal bands on the abaxial side, undulate anticlinal epidermal cell walls and papillae on the epidermal cells and on the subsidiary cells of the stomata. The new species is the youngest reliable representative of the genus Phoenicopsis. This record extends the stratigraphic range of Phoenicopsis from its previously known latest records in the early Campanian up to Maastrichtian–Danian. The persistence of the genus Phoenicopsis up to the Cretaceous–Paleogene boundary is most likely related with vicinity to mountain floras associated with Late Cretaceous and Paleogene volcanic uplands.     

Revision of Cretaceous Radiolarians of the Family Hagiastridae from Lipman’s Collection

Based on the Lipman’s hagiastrid collection and original material, Late Cretaceous holotypes of the genera Histiastrum Ehrenberg, Hagiastrum Haeckel, and Tesserastrum Haeckel are revised. Histiastrum aster Lipman, H. cruciferum Lipman, H. membraniferum Lipman, H. latum Lipman, H. tetracanthum Lipman, H. tumeniense Lipman, Hagiastrum crux Lipman, and Tesserastrum quadratum Lipman are transferred to the genus Crucella Pessagno. A new species, Rhombastrum lipmanae sp. nov., is described. It is shown that Crucella lata (Lipman) and C. crux (Lipman) are Santonian–Campanian index species and C. crucifera (Lipman), C. tetracantha (Lipman), C. tumeniensis (Lipman), and Rhombastrum lipmanae sp. nov. are markers of the Campanian Boreal Realm.     

Organic-walled microfossils from Cambrian Stage IV in the Jiaobang section,eastern Guizhou,China

The lower Cambrian succession in the Jiaobang section, Jianhe County, eastern Guizhou, China, includes, in ascending order, the Bianmachong, Balang, and Tsinghsutung formations, with a total thickness of about 645 m. Twenty-six morphological genera (including one new genus) are identified from the Balang and the underlying Bianmachong formations, many of which are common and widely distributed. Six acritarch assemblages are discerned in the Balang Formation. They are, in ascending order, the Adara alea‒Skiagia ornata, the Acrum radiale‒Pterospermella velata, the Comasphaeridium molliculum‒Solisphaeridium baltoscandium, the Corrugasphaera perfecta n. sp.‒Pterospermella vinctusa n. sp., the Acrum novum‒Heliosphaeridium oligum, and the Acrum membranosum‒Adarve diafanum acritarch assemblages. An obvious change of organic-walled microfossil assemblages occurred in the interval between 84 m and 98 m from the bottom of the Balang Formation which roughly corresponds to the boundary between the Oryctacarella duyunensis trilobite Zone and the overlying Arthricocephalus chauveaui trilobite Zone. In addition, organic-walled microfossils are scarce in about 24 m thick from the bottom of the Balang Formation. One new genus and five new species including Plagasphaera balangensis n. gen. n. sp., Asteridium tubulus n. sp., Cymatiosphaera spina n. sp., Corrugasphaera perfecta n. sp., and Pterospermella vinctusa n. sp. are described.     

Calcareous and siliceous phytoplankton stratigraphy of Neogene marine sediments in central Crete (Greece)

Fine laminated diatomaceous beds in outcrops of the Finikia unit (Estavromenos and Athanatoi sections) in the northern part of the Heraklion district in Crete, have yielded a total of 55 marine diatom and 24 silicoflagellate taxa. The diatom assemblages, characterized by species of low latitudes, belong to the upper part of the Nitzschia jouseae biozone (early-late Pliocene). The silicoflagellate associations belong to the local Dictyocha fibula ausonia and Dictyocha hellenica subzones. These subzones correspond to the subzones CN11b (Discoaster asymmetricus) and CN12a (D. tamalis) of calcareous nannofossils respectively, as well as to the Globorotalia puncticulata/G. bononiensis biozones of planktic foraminifera. The sparse silicoflagellate association from fine sandy marls of the Panassos section in the central part of the Heraklion district (Aghia Varvara unit, upper Tortonian-Messinian) is assigned to the local biozone Distephanus speculum minutus. The latter corresponds approximately to the biozone C9 (Discoaster quinqueramus) of calcareous nannofossils and to the Globorotalia conomiozea zone of planktic foraminifera. No discoidal diatoms could be found in the samples from Panassos-section. The presence of silicoflagellates but simultaneous absence of diatoms is probably due to greater opal dissolution within the fine sandy marls of the Panassos section.     

Radiolaria and planktonic foraminifera from Sarama composite section of the Kannaviou Formation (Campanian,Upper Cretaceous,Cyprus)

Radiolarians and planktonic foraminifers were studied from a series of continuous outcrops of the Upper Cretaceous (upper Campanian) Kannaviou Formation near Sarama Village (southwestern Cyprus). The composite study section has a narrow stratigraphic interval that covers the upper Campanian, with a total thickness of more than 70 m. The Amphipyndax tylotus (radiolarian) Zone is found within the study section. According to foraminiferal data the studied interval corresponds to the middle-upper Campanian age (from Contusotruncana plummerae Zone to the lower part of Gansserina gansseri Zone).     

Bouleiceras (Hildoceratidae,Ammonitina) from the lower Toarcian (Jurassic) of the Iberian Range (Spain): Taxonomy,stratigraphic distribution and insights on its dispersal

Bouleiceras is a very rare genus among the rich assemblages of ammonoids from the lower Toarcian of the Iberian Range. So far, only two dozen specimens have been recorded in numerous field campaigns carried out since 1965 by different authors. The interest of this taxon lies in its peculiar paleogeographical distribution in comparison with most other ammonoids of the same age. A review of these specimens is carried out, including those obtained in previous works and others recently collected in selected localities. Based mainly on the differences in the shape of the ventral section and the suture line, seven species have been identified; two of which are new: Bouleiceras ibericum nov. sp. and Bouleiceras? betetensis nov. sp. All the reviewed specimens are recorded in the Semicelatum Subzone of the Tenuicostatum Zone and the Elegantulum Subzone of the Serpentinum Zone from the Central Sector and the Levantine Sector of the Iberian Range. The global distribution of the genus is summarized from the known data, and its possible dispersal routes are analyzed, as well as the factors that could have conditioned them.     

Upper Cretaceous sulcate pollen from the Timerdyakh Formation,Vilui Basin (Siberia)

One inaperturate and 16 monosulcate pollen types are described from the latest Campanian to earliest Maastrichtian sediments of the Vilui basin, Siberia, using both light and scanning electron microscopy, and assigned systematically when possible to modern families or subfamilies. Despite their scant occurrence, the pollen grains show considerable diversity. Two new genera have been erected. Lasioideaecidites with two species, Lasioideaecidites hessei and Lasioideaecidites bogneri, represents the earliest record of the subfamily Lasioideae (Araceae). Aristolochiacidites with one species, Aristolochiacidites viluiensis, is assigned to the subfamily Aristolochioideae (Aristolochiaceae) and represents the first fossil pollen record of the family. A new species of Liliacidites, Liliacidites goldblattii, is closely similar to pollen of Isophysioideae (Iridaceae) and/or Doryanthaceae and is the first fossil evidence of the clade that includes these closely related families. A further nine new fossil species are described (Clavatipollenites timerdyakhensis, Liliacidites palaeofritillaria, Retimonocolpites longosucatus, R. microreticulatus, R. microrugulatus, R. lysichitonoides, R. chapmaniae, Monosulcites parvus and Arecipites tyungensis) and assigned to Chloranthaceae, Araceae (Orontioideae, ?Pothoideae), Liliaceae, ?Hypoxidaceae and Arecaceae. The Hydatellaceae (Nymphaeales) may also be present in the palynoflora of the Vilui Basin, represented by pollen similar to M. rivularis Braman from the Santonian to Campanian (Upper Cretaceous) Milk River Formation, Canada.     

<Emphasis Type="Italic">Weitschataster intermedius</Emphasis> gen. et sp. nov., a goniasterid starfish (Echinodermata: Asteroidea) from the Upper Cretaceous of Germany

Weitschataster intermedius gen. et sp. nov., is described from the Upper Campanian (Upper Cretaceous) of Hannover (Lower Saxony, N Germany). The new species is characterized by its conspicuous rimmed pedicellariae scars covering almost the complete actinal surface and by small body size. W. intermedius represents a transitional state between W. decipiens comb. nov. from the Lower Campanian and the well-known W. undulatus comb. nov. from the Lower Maastrichtian of NW Europe. Among Spencer’s types of W. undulatus, one paratype is recognized as belonging to the genus Parametopaster.     

Late Cretaceous to early tertiary Dinoflagellate cysts from Narasapur Well-1, Godavari-Krishna basin,South India

Dinoflagellate cyst assemblages are described fromsome of the richest horizons in the Narasapur Well-1, Godavari-Krishna basin, Andhra-Pradesh, India. Twenty-five taxa belonging to nineteen genera are recorded. The majority exhibit close morphological similarity with species described earlier from Europe, North America and Australia. One new genus, Godavariella, and three new species, Godavariella venkatachalae, Fibrocysta variabilis and Cyclonephelium indicum, are proposed; several more new forms are described but, since meagrely represented, are not named. The dinoflagellate assemblages, along with spore and pollen studies carried out earlier by Venkatachala & Sharma (1984), suggest a Late Cretaceous (Maastrichtian) to Early Tertiary (Palaeocene) age for these sediments.     

A new enantiornithine bird from Upper Cretaceous non-marine deposits at Villespassans (Hérault,southern France)

A coracoid of an enantiornithine bird from Upper Cretaceous (probably late Campanian) fluvial sediments at Castigno (Villespassans, Hérault, southern France) is described. It differs from all hitherto reported enantiornithine coracoids and is referred a new genus and species, Castignovolucris sebei. This bone is large and robust, indicating a bird that was among the largest known enantiornithines, possibly the size of a Canada Goose (Branta canadensis). The new taxon is an addition to the short list of Late Cretaceous birds from France and confirms that enantiornithines were an important component of European avifaunas until late in the Cretaceous.