The development of planktonic biogeography in the Southern Ocean during the Cenozoic

Deep-sea drilling at high latitudes of the Southern Hemispheres has provided almost the only available data to evaluate the biogeographic development of the planktonic biota in the Southern Ocean during the Cenozoic (65 m.y. to Present Day). Paleontological investigations on Deep Sea Drilling Project (DSDP) materials have shown that the development of Cenozoic planktonic biogeography of the Southern Ocean is intimately linked with the evolution of the Southern Ocean water masses themselves. During the Cenozoic, this has included the development of the Circum-Antarctic Current system as obstructing land masses moved apart, the refrigeration and later extensive glaciation of the continent, and the development of the Antarctic Convergence (Polar Front) with related oceanic upwelling.Almost all evolution of calcareous planktonic microfossils has occurred outside of the Antarctic—Subantarctic region followed by limited migration into these water masses. Virtually no endemism occurs amongst calcareous microfossil groups at these latitudes. In contrast, conspicuous and widespread evolution has occurred within the siliceous microfossil groups especially during the Neogene. Low diversity and differences in stratigraphic ranges of Antarctic calcareous microfossils makes them only broadly useful for correlation. Relatively higher diversities within the Subantarctic provide a firmer basis for more detailed correlation, although the ranges of fossils are often different than at lower latitudes because of different paleoceanographic and paleoclimatic controls. Within the Antarctic water mass south of the Antarctic Convergence, siliceous microfossilsbiostratigraphy, oxygen isotopic stratigraphy and magnetostratigraphy, provide the only firm basis for correlation with low-latitude sequences.Eocene (55-38 Ma) sediments contain abundant calcareous microfossils even closely adjacent to the continent. Antarctic calcareous planktonic microfossils of this age exhibit relative high diversity, although this is lower than assemblages of equivalent age at middle and low latitudes. Within the Subantarctic region, Eocene planktonic foraminifera exhibit strong affinities with those in the temperate regions. Biogeographic differences exist between various sectors of the Southern Ocean related to biogeographic isolation preceding the development of the Circum-Antarctic Current. Subantarctic calcareous nannofossil assemblages of Paleocene and Eocene age exhibit higher diversity than Oligocene and Neogene assemblages. Siliceous microfossils are poorly represented or at best poorly known.One of the most dramatic changes in Southern Ocean planktonic biogeography occurred near the Eocene/Oligocene boundary (38 Ma). Since then, Antarctic planktonic foraminiferal assemblages have exhibited distinct polar characteristics, marked in particular by low diversity, and this event thus reflects the initiation of the Antarctic faunal and floral provinces. Profound paleoceanographic changes at this time, which triggered the biogeographic crisis, appear to be related to the initiation of widespread Antarctic sea-ice formation, and rapid cooling of deep and intermediate waters, in turn associated with increased Antarctic glaciation. During the Oligocene, planktonic microfossil diversity was low in all groups throughout the world's oceans. In Antarctic waters, the early Oligocene foraminiferal fauna is monospecific (Subbotina angiporoides), while in the later Oligocene two species (S. angiporoides and Catapsydrax dissimilis) were recorded. Calcareous nannofossil assemblages are of low diversity compared with the Eocene. Subantarctic foraminiferal faunas of Oligocene age display much higher diversity than those in the Antarctic, but early and middle Oligoceae faunas still exhibit the lowest diversities for the entire Cenozoic. Siliceous assemblages remain relatively inconspicuous in most regions of the Southern Ocean.The Paleogene-Neogene transition (22 Ma) is marked by a major change in the global planktonic biogeography, i.e. modern patterns developed in which permanent, steep faunal and floral diversity gradients existed between tropical and polar regions; a gradient which has persisted even during the most severe glacial episodes. Oligocene assemblages of low diversity and almost cosmopolitan distribution were replaced by distinctive belts of planktonic assemblages arranged latitudinally from the tropics to the poles. The establishment of the steep planktonic diversity gradients and latitudinal provinces near the beginning of the Neogene almost certainly were linked to the development of the Circum-Antarctic Current in the late Oligocene which effectively separated high- and low-latitude planktonic assemblages. These fundamental global circulation and biogeographic patterns have persisted through the Neogene.During the Neogene (22 Ma to Present Day), Antarctic calcareous microfossil assemblages exhibit persistent low diversity and high dominance, while Subantarctic assemblages are of much greater diversity. The beginning of the Neogene (= beginning of Miocene) heralded the development of the high-latitude siliceous microfossil assemblages towards their present-day dominant role. Siliceous biogenec productivity began to increase. These changes were linked to the initial development and later intensification of circulation associated with the Antarctic Convergence and Antarctic Divergence. The Antarctic Convergence sharply separates dominantly siliceous assemblages to the south from calcareous assemblages to the north. Radiolarian assemblages became more endemic. Relatively warm early and middle Miocene conditions are reflected by slightly higher diversity of planktonic foraminifera and by the presence, in the northern Subantarctic, of conspicuous discoasters in early Miocene sediments. In Antarctic waters, calcareous nannofossils become unimportant as biogenic elements after the middle Miocene.The latest Miocene ( 5 m.y. ago) was marked by northward movement of the Antarctic Convergence, corresponding expansion of the Antarctic water mass, and low diversity of calcareous assemblages. Pliocene planktonic foraminifera seem to be largely monospecific in Antarctic and southern Subantarctic sequences. During the Quaternary, Antarctic waters reached a maximum northward expansion and exhibit highest siliceous biogenic productivity for the Cenozoic. In the Subantarctic, Quaternary foraminiferal diversities are much higher than in Pliocene sequences. Although calcareous nannofossil diversity may be high, only a few species are abundant. Large northward shifts of Antarctic and Subantarctic water masses have occurred during the Quaternary although no southward penetrations have occurred much beyond that of the present day. Several radiolarian and foraminiferal species disappeared or appeared at or close to a number of paleomagnetic reversals during the last 4 m.y. These faunal events, which provide valuable datums, do not seem to be associated with major climatic changes.     

Stable isotopic study of Oligocene-Miocene sediments from DSDP Site 354, Equatorial Atlantic

The oxygen- and carbon-isotope compositions of planktic and benthic foraminifera and calcareous nannofossils from Middle Oligocene-Early Miocene Equatorial Atlantic sediments (DSDP Site 354) indicate two important paleoceanographic changes, in the Late Oligocene (foraminiferal Zone P.21) and in the Early Miocene (foraminiferal Zone N.5). The first change, reflected by a δ¹⁸O increase of 1.45‰ inGlobigerina venezuelana, affected only intermediate pelagic and not surface, deep or bottom waters. The second change affected surface and intermediate waters, whereas deep and bottom waters showed only minor fluctuations. In the case of the former the isotope effect of the moderate ice accumulation on the Antarctic continent is amplified in the Equatorial Atlantic by changes in the circulation pattern. The latter paleoceanographic change, reflected by a significant increase in¹⁸O in both planktic and benthic forms (about 1.0‰ and 0.5‰, respectively), may have been caused by ice volume increase and temperature decrease. Both oxygen- and carbon-isotope compositions indicate a marked depth-habitat stratification for planktic foraminifera and calcareous nannofossils. Three different dwelling groups are recognized: shallowGlobigerinoides, Globoquadrina dehiscens, Globorotalia mayeri and nannofossils; intermediateGlobigerina venezuelana; and deepCatapsydrax dissimilis. The comparison of foraminifera and calcareous nannofossils suggests that the isotopic compositions of nannofossils are generally controlled by the same parameters which control the isotopic composition of shallow-dwelling foraminifera, but the former are more enriched in¹⁸O.     

Middle to late Miocene fluctuations in the incipient Benguela Upwelling System revealed by calcareous nannofossil assemblages (ODP Site 1085A)

Middle to late Miocene calcareous nannofossil data of ODP Site 1085 from the eastern South Atlantic off Namibia were analysed to document spatial and temporal changes in surface-ocean circulation, upwelling initiation, and associated productivity.

Our data show that calcareous nannofossils constitute a significant part of the carbonate fraction throughout the investigated interval from 12.5 to 7.7 million years (Ma). Highest numbers of calcareous nannofossils (up to 38,000 × 10⁶ nannofossils g^− 1 sediment) were observed during the intervals 9.9 to 9.7 and 8.7 to 8.0 Ma. These elevated numbers of calcareous nannofossils may generally be linked to the initiation of upwelling at about 10 Ma in the studied region. In contrast, diminished numbers of calcareous nannoplankton, as in the interval 9.6 to 9.0 Ma, probably characterise time intervals of weaker productivity resulting in a decrease of nannofossil carbonate contents in the sediments of Site 1085. This decrease in nannofossil production could be one possible explanation for the major CaCO₃ depression in between 9.6 and 9.0 Ma. Coccoliths of the genus Reticulofenestra are the most abundant taxa. Their occurrences are characterised by changes in the investigated time interval. In addition, Coccolithus pelagicus, Calcidiscus leptoporus and Umbilicosphaera spp. contribute a common part of the assemblage. Calcareous nannofossils account for more than half of the carbonate, with peak contribution up to 80% at 8.8 Ma.     

FOSSILS AND A LARGE MOLECULAR PHYLOGENY SHOW THAT THE EVOLUTION OF SPECIES RICHNESS,GENERIC DIVERSITY,AND TURNOVER RATES ARE DISCONNECTED

The magnitude and extent of global change during the Cenozoic is remarkable, yet the impacts of these global changes on the biodiversity and evolutionary dynamics of species diversification remain poorly understood. To investigate this question, we combine paleontological and neontological data for the angiosperm order Fagales, an ecologically important clade of about 1370 species of trees with an exceptional fossil record. We show differences in patterns of accumulation of generic diversity, species richness, and turnover rates for Fagales. Generic diversity evolved rapidly since the Late Cretaceous and peaked during the Eocene or Oligocene. Turnover rates were high during periods of extreme global climate change, but relatively low when the climate remained stable. Species richness accumulated gradually throughout the Cenozoic, possibly at an accelerated pace after the Middle Miocene. Species diversification occurred in new environments: Quercoids radiating in Oligocene subtropical seasonally arid habitats, Casuarinaceae in Australian pyrophytic biomes, and Betula in Late Neogene holarctic habitats. These radiations were counterbalanced by regional extinctions in Late Neogene mesic warm‐temperate forests. Thus, the overall diversification at species level is linked to regional radiations of clades with appropriate ecologies exploiting newly available habitats.     

长江三角洲DY03孔颗石藻及沉积环境浅析

本文研究了上海奉贤DY03孔的颗石藻化石。钻孔60.8-2m深度产颗石藻化石,涵盖上更新统的川沙组、南汇组,全新统的娄塘组、上海组和青浦组。颗石藻化石赋存于灰色泥、泥质粉砂或粉细砂中,但化石种类贫乏。群落面貌以Gephyrocapsa oceanica占绝对数量,Braarudosphaera bigelowii,Helicosphaera hyalina和Heli-cosphaera wallichii等偶现,属典型的单种类组合。研究依据颗石藻定量统计,划分出两个海相层:第一海相层,孔深61-40m,属上更新统川沙组和南汇组,颗石藻丰度低,属种单一,代表了海侵过程中的潮上带环境;第二海相层,孔深24.8-2.0m,属全新统娄塘组、上海组和青浦组,颗石藻化石丰度较高,并有数个属种共生,反映当时的滨岸潮上带-河口滨海-浅海近岸海湾的环境演化过程。综合前人的研究成果,认为长江三角洲全新世海侵之强度和范围都超过晚更新世发生的海侵。此外,本文仔细讨论了Gephyrocapsa oceanica的生态特征,显示其为一个盐度适应能力较强、分布广泛的暖水种。     

Quaternary calcareous nannofossils from Periantarctic basins: Paleoecological and paleoclimatic implications

Currently, coccolithophores have a widespread oceanic distribution and are reported from most latitudes, but not those higher than 65°S. Fifteen piston cores were sampled with the aim of investigating the distribution and abundance variation of Quaternary calcareous nannofossils of the Antarctic region, south of the Antarctic Divergence (> 65°S), particularly from Maud Rise, Bausan Bank, and from Weddell, Ross and Bellingshausen Seas. A calcareous nannofossil cold-taxa association is present in most of the cores examined and their discontinuous occurrence is thought to indicate key environmental relationships. The presence of calcareous nannofossils is correlated with interglacial intervals with warmer SSTs and may indicate high productivity and an open-ocean environment. Our results confirm that, during short periods of the late Quaternary, coccolithophorids occurred at southern high latitudes, in the western Antarctic basins, while in the eastern Antarctic basins they are nearly absent, suggesting more variable SSTs near West Antarctic Ice Sheet.     

A new duck from the Middle Miocene of Mongolia, with comments on Miocene evolution of ducks

A new relatively large duck, Chenoanas deserta gen. et sp. nov. from the Middle Miocene of the Sharga locality, which is morphologically intermediate between the extant genera Chenonetta and Tachyeres, is described. The diversity of Early and Middle Miocene ducks is discussed. It is noted that some Middle Miocene duck remains are incorrectly referred to the genus Mionetta. The distribution of morphological characters of the humerus in Neogene and extant ducks shows that the present day diversity of ducks apparently results from extinction of some taxa which were formed in the Oligocene-Early Miocene. The distribution of morphological characters in the evolution of diving ducks is evidence that not only the formation of different morphotypes but also so-called ??evolutionary maturation?? of taxa also explains the modern diversity.     

New fossil leaves and fruits of Lauraceae from the Middle Miocene of Fujian,southeastern China differentiated using a cluster analysis

The fossil record of Lauraceae can be traced back to the Early Cretaceous of eastern Asia based on fossil flowers. Here, we refer a number of new occurrences of leaf and fruit fossils of Lauraceae from the Middle Miocene of Zhangpu, Fujian, China, to seven species. These data provide evidence supporting the fact that a diverse subtropical, or tropical, Lauraceae-dominated evergreen forest surrounded this region 15 million years ago (Mya). The Lauraceae fossils presented in this paper provide evidence for the evolution of this group as well as new materials that enable the study of the Fujian Province Neogene flora. The fossils described in this paper fill in the gaps in studies about Lauraceae pollen in the Middle Miocene from Fotan, Fujian, China. In addition, these fossils also enrich the Middle Miocene fossil records of Lauraceae in eastern Asia, especially improving the study of the macrostructures and reproductive organs of fossil Lauraceae from southern China. The similarity between fossil and modern fruits shows that during the Middle Miocene the fruit morphological of Lauraceae have changed very little. We also identify families where the fossils we report belong to their closest relatives and can be used to reconstruct the paleoenvironment of Fujian in the Middle Miocene.     

Diversity and biogeography of the Antarctic flora

Aim To establish how well the terrestrial flora of the Antarctic has been sampled, how well the flora is known, and to determine the major patterns in diversity and biogeography. Location Antarctica south of 60° S, together with the South Sandwich Islands, but excluding South Georgia, Bouvetøya and the periantarctic islands. Methods Plant occurrence data were collated from herbarium specimens and literature records, and assembled into the Antarctic Plant Database. Distributional patterns were analysed using a geographic information system. Biogeographical patterns were determined with a variety of multivariate statistics. Results Plants have been recorded from throughout the Antarctic, including all latitudes between 60° S and 86° S. Species richness declines with latitude along the Antarctic Peninsula, but there was no evidence for a similar cline in Victoria Land and the Transantarctic mountains. Multi‐dimensional scaling ordinations showed that the species compositions of the South Orkney, South Shetland Islands and the north‐western Antarctic Peninsula are very similar to each other, as are the floras of different regions in continental Antarctica. They also suggest, however, that the eastern Antarctic Peninsula flora is more similar to the flora of the southern Antarctic Peninsula than to the continental flora (with which it has traditionally been linked). The South Sandwich Islands have a flora that is very dissimilar to that in all Antarctic regions, probably because of their isolation and volcanic nature. Main conclusions The Antarctic flora has been reasonably well sampled, but certain areas require further floristic surveys. Available data do, however, allow for a number of robust conclusions. A diversity gradient exists along the Antarctic Peninsula, with fewer species (but not fewer higher taxa) at higher latitudes. Multi‐dimensional scaling ordination suggests three major floral provinces within Antarctica: northern maritime, southern maritime, and continental. Patterns of endemism suggest that a proportion of the lichen flora may have an ancient vicariant distribution, while most bryophytes are more recent colonists.     

Dabbling ducks (Aves: Anatidae) from the Middle Miocene of Mongolia

A new genus and two new species of ducks (Aves: Anatidae) from the Middle Miocene Sharga locality are described. Mioquerquedula minutissima gen. et sp. nov. is a very small duck. Anas velox Milne-Edwards, 1868 from the Middle Miocene of France is transferred to the genus Mioquerquedula. Aix praeclara sp. nov. described here is the oldest record of the modern genus Aix. A revision of the previously described small duck Anas soporata Kurochkin, 1976 shows that only the specimens from the Sharga locality should be referred to this species. The status of other small ducks from the Neogene of Europe and North America is discussed. The diversity of herbivorous and diving ducks in the Sharga locality indicates that Miocene Shargyn Govi Lake was rich in food resources.     

On the adaptive pattern of "Ramapithecus".

A reconstruction of the adaptive pattern of the Miocene hominoid "Ramapithecus," supported by comparative data and morphological comparisons, is presented. It is based upon the completely known dentition, inferior portion of the face, and on the paleoecology and geographic distribution of "Ramapithecus" bearing deposits. The known adaptive and morphological patterns of the "Ramapithecus" species are almost indistinguishable from those of the contemporary species of Sivapithecus, which has taxonomic priority. The "Ramapithecus" species are therefore referred to Sivapithecus, a dryopithecine genus which apparently radiated throughout Eurasia and Africa during the Middle Miocene.     

Biochronologie et corrélation des bassins néogènes du Couloir sud-rifain (Maroc) fondées sur les évènements de foraminifères planctoniques et de nannofossiles calcaires

Qualitative and quantitative analyses of planktonic foraminifera (completed by calcareous nannofossils in some localities) from 8 sections and boreholes located in 7 Neogene basins of the South Rifian Corridor (Morocco) enable us to identify 16 bioevents calibrated with the geomagnetic polarity time scale. These events are useful for (1) assigning the sediments to the Tortonian (bioevents 1 to 7), the Messinian (bioevents 9 to 14, the event 8 indicates the Tortonian/Messinian boundary), the Early Pliocene (bioevent 15) and the Middle Pliocene (bioevent 16), (2) establishing high resolution correlations between the sections and boreholes studied herein, (3) locating the South Rifian Corridor sections and boreholes within the framework of biostratigraphic events recognized during the latest ten years in time-equivalent Atlantic and Mediterranean sequences, and (4) estimating the variation of sedimentation rates in the studied basins. With respect to Morocco, previous detailed studies concerned only the Rabat area (Bou Regreg valley) and Guercif basin. Our results extend correlations to the other basins of the South Rifian Corridor, a key area to understand connections between the Atlantic Ocean and the Mediterranean Sea at the end of the Miocene.     

Triton's trident: cryptic Neogene divergences in a marine clam (Lasaea australis) correspond to Australia's three temperate biogeographic provinces

The southern coast of Australia is composed of three distinct biogeographic provinces distinguished primarily by intertidal community composition. Several ecological mechanisms have been proposed to explain their formation and persistence, but no consensus has been reached. The marine clam Lasaea australis is arguably the most common bivalve on southern Australian rocky shores and occurs in all three provinces. Here, we tested if this species exhibits cryptic genetic structuring corresponding to the provinces and if so, what mechanisms potentially drove its divergence. Variation in two mitochondrial genes (16S and COIII) and one nuclear gene (ITS2) was assayed to test for genetic structuring and to reconstruct the clam's phylogenetic history. Our results showed that L. australis is comprised of three cryptic mitochondrial clades, each corresponding almost perfectly to one of the three biogeographic provinces. Divergence time estimates place their cladogenesis in the Neogene. The trident‐like topology and Neogene time frame of L. australis cladogenesis are incongruent with Quaternary vicariance predictions: a two‐clade topology produced by Pleistocene Bass Strait land bridge formation. We hypothesize that the interaction of the Middle Miocene Climate Transition with the specific geography of the southern coastline of Australia was the primary cladogenic driver in this clam lineage. Additional in‐depth studies of the endemic southern Australian marine biota across all three provinces are needed to establish the generality of this proposed older framework for regional cladogenesis.     

Ostracodes et stratigraphie du néogène et du quaternaire méditerranéens

A stratigraphical chart of marine ostracoda from Lower Miocene to Recent is established. Selected species (approximatively 220) are those morphologicaly well characterized and known from different parts of the Mediterranean area. It appears that: • lower Miocene ostracodes are still poorly known; • specific diversity is high during the Tortonian and the Lower Messinian before the complete disappearance of marine Mediterranean species during the Upper Messinian évaporitic episodes; • during the early Pliocene, about half of the Upper Miocene marine species are reintroduced with the Atlantic waters; other species migrate for the first time in the Mediterranean Sea by the same way; • at the end of the Pliocene or at the beginning of the Pleistocene several species known in Mediterranean since the Middle Miocene or before, such as Cytherella sp. gr. transversa and Ruggieria tetraptera, as well some “nordic guests” such as Hemicythere villosa and Cythere lutea, appear. This work is an opportunity to confirme a Late Miocene age for the Neogene of Skyros (Aegean Sea), to assign the “Upper Pliocene” of Terquem to the Lower Pleistocene and to refute the existence of a pliocene psychrosphere.     

Biodiversity and historical biogeography of stalked crinoids (Echinodermata) in the deep sea

About 95 species of stalked crinoids are now described from 60m to hadal depths, but our knowledge remains far from complete. Depending on which species concept is used, estimates of species richness can be dramatically different. It is necessary to have a homogeneous concept for taxonomic units. The abundance of the crinoid fossil record allows a discussion of the ancestry of deep sea crinoid fauna. Stalked crinoids have a horizontal diversity pattern with three regional centres of high diversity (i.e. western tropical Pacific, western tropical Atlantic and north-eastern Atlantic). Vertical patterns show two faunal strata which vary in importance among provinces. The epibathyal stratum has apparently remained relatively similar in intertropical areas since the Mesozoic. Despite environmental changes related to glaciation since the Middle Miocene, the deepest crinoid fauna (i.e. the deep sea fauna sensu stricto at depths more than 1000 ± 200 m) have a very ancient origin with a dispersion closely related to plate tectonics. The bathyal fauna on hard substrates includes a few living fossils and has a high historical interest.     

苏北盆地阜宁组钙质超微化石由来初探

对苏北盆地4个钻孔300余块心样品进行了分析,在阜宁组的8块样品中发现了钙质超微化石,计3属1种及3个比较种,化石属种单调,保存较 ,受到严重的溶蚀或增生,结合这些种类的分布时代和产出层位,很难排除苏北盆地阜宁组钙质超微化石再沉积的属性。     

New data on the distribution of calcareous nannofossils in the Upper Jurassic Loino section (Kirov region,Russia)

The middle Volgian clayey deposits of the panderi Zone in the Loino section of the Kirov region have been found to contain an assemblage of calcareous nannofossils never studied in such high latitudes of the Boreal Realm. The species constituting this assemblage are characterized by wide geographic and stratigraphic distributions.     

中国陆相新第三系的初步划分与对比

本文着重介绍了欧洲地中海区新第三纪研究的进展、分期和对比,并提出我国新第三纪哺乳动物群的分期及与世界对比的初步设想。文中着重阐明将蓬蒂期(Pontian(s. s. )相当于我国保德三趾马红土)置于中新世的根据,并建议把新第三纪/第四纪的界限放在1.8百万年(相当奥杜威事件)。对中新世,作者建议划分为五个不同的期:早中新世,谢家期;中中新世早期,山旺期;中中新世晚期,通古尔期;晚中新世,坝河期和最晚中新世,保德期。对上新世的划分目前尚有困难,建议暂时保留静乐期,代表上新世早期,大体相当欧洲的路西南期(Ruscinian, MN14—MN15)。借用游河期代表上新世晚期,约相当于欧洲的维兰尼期(Villanyian,MN16—MN17)。     

A new Oligocene-Miocene calcareous nannofossil species: Pontosphaera wallacei

A high-resolution study of sediments from DSDP Leg 72 Site 516 provided the opportunity to reconstruct a detailed biostratigraphy for the late Oligocene - early Miocene time interval. The detailed taxonomic investigation of calcareous nannofossils, performed by a quantitative analysis, highlighted the presence of a species of Pontosphaera that does not show the morphologic features of any recognized species of this genus. This species, which was previously classified as Pontosphaera sp., is here described as Pontosphaera wallacei nov. sp.     

Plankton stratigraphy of the Early and Middle Miocene Kareem and Rudeis Formations in the central part of the Gulf of Suez,Egypt

The planktonic foraminifera and nannofossils of three wells in the Gulf of Suez penetrating the Early to Middle Miocene Upper Rudeis and Kareem Formations are attributed (from top to base) to the Middle Miocene Globorotalia peripheroronda Partial Range Zone (M6), the earliest Middle Miocene Praeorbulina sicana–Orbulina suturalis Interval Zone (M5), subdivided into the Praeorbulina glomerosa s. strict.–O. suturalis Interval Subzone (M5b) and the P. sicana–P. glomerosa s. str. Interval Subzone (M5a) and the Early Miocene Globigerinoides bisphericus Partial Range Subzone (M4b). The appearance of O. suturalis at the base of Subzone M5b represents the final stage of evolution of the Globigerinoides trilobus–Praeorbulina–Orbulina Lineage. In addition, the calcareous nannoplankton assemblages indicate the Sphenolithus heteromorphus Zone (NN5) and the Helicosphaera ampliaperta Zone (NN4). These biozones are well correlatable with those established by El-Heiny and Martini (1981, Geol Mediterr. Tome, VIII(2): 101–108) from the southwestern flank of the Gulf of Suez.