The Middle Miocene climatic transition in the Southern Ocean: Evidence of paleoclimatic and hydrographic changes at Kerguelen plateau from planktonic foraminifers and stable isotopes

Middle Miocene (14.8–11.9 Ma) deep-sea sediments from ODP Hole 747A (Kerguelen Plateau, southern Indian Ocean) contain abundant, well-preserved and diverse planktonic foraminiferal assemblages. A detailed study of the climatic and hydrographic changes that occurred in this region during the Middle Miocene Climatic Transition led to the identification of an intense cooling phase (the Middle Miocene Shift). Abundance fluctuations of planktonic foraminiferal species with different paleoclimatic affinities, and oxygen and carbon stable isotopes have been integrated in a multi-proxy approach. Reconstruction of changes in foraminiferal faunal composition and diversity through time were the basis for identification of three foraminiferal biofacies. The most prominent faunal change took place at 13.8 Ma, when a fauna with warm-water affinity (marked by high abundance of Globorotalia miozea group and Globoturborotalita woodi plexus) was replaced by an oligotypic, opportunistic fauna with typical polar characters and dominated by neogloboquadrinids. This faunal change is interpreted as the result of foraminiferal migration from adjacent bioprovinces, caused by modifications in climate and hydrography. A positive 2.0‰ shift in δ¹⁸O (interpreted as the Mi3 event) and a related positive 1.0‰ shift in δ¹³C (corresponding to the CM6 event) accompanied this faunal turnover. These are interpreted to reflect substantial reorganization of Southern Ocean waters, the northward migration of the Polar Front and a strong increase in primary productivity. The second faunal change took place at 12.9 Ma and was characterized by the gradual decrease in abundance of the neogloboquadrinids and the recovery of Globorotalia praescitula/scitula group and Globigerinita glutinata. A positive 1.5‰ shift in δ¹⁸O (interpreted as the Mi4 event) and a concurrent gradual negative shift in δ¹³C accompanied this faunal change, witnessing further modifications of the climate/ocean system. Variations in sea surface temperature, considered as the main factor causing changes of surface hydrography at the Kerguelen Plateau, seem to have been driven by obliquity and long-term eccentricity, thus suggesting a key role played by the astronomical forcing on the evolution of Southern Ocean dynamics during the Middle Miocene. Also an evident 1.2 Myr modulation of the δ¹³C record suggests a main control of the long-term obliquity cycles on the carbon cycle dynamics. Particularly, the Mi3/CM6 events exactly fit with a node of the 1.2 Myr modulation cycles. This confirms the key role played by orbital parameters on high-latitude temperatures and Antarctic ice volume, and indirectly on global carbon burial and/or productivity. This climatic transition was marked also by changes in surface hydrography. From 14.8 to 13.8 Ma an intermediate-strength thermocline controlled by seasonality developed just below the photic zone. Weaker seasonality characterized the interval from 13.8 to 12.9 Ma, when the thermocline became shallower and sharper and favored intermediate-water foraminifers. From 12.9 Ma, seasonality increased again and an intermediate-strength thermocline re-developed.     

Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified.Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in δ¹⁸O values coded as “Oi” and “Mi” events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific.Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.We also reviewed changes in other microfossil assemblages in the low latitudes during the late Oligocene through early Miocene. The microfossil assemblages of major groups show sequential changes near the Oligocene/Miocene (O/M) boundary (23.8 Ma). Many extinction events and some first occurrences of calcareous nannofossils and many occurrences of radiolarians are found from about 24.8 to 23.3 Ma, and first occurrences of planktic foraminifers and diatoms followed from 23.2 through 22 Ma. Hence, the O/M boundary is identified as a significant level for microfossil evolutions.     

Neogene planktonic foraminiferal biostratigraphy and evolution: Equatorial to subantarctic, South Pacific

Detailed planktonic foraminiferal zonations have been established for the Neogene (Latest Oligocene through present) in six DSDP sites in the South Pacific ranging from equatorial to subantarctic latitudes (48°S). Two basic zonal schemes are readily recognized: tropical and temperate. The tropical zonation is best developed in DSDP Site 289 and the temperate zonation in Sites 206, 207A and 284. Tropical and temperate zonations can be linked by a warm subtropical scheme in Site 208, because this sequence includes a mixture of tropical and temperate elements. A site located close to the Subantarctic Convergence (Site 281) contains a zonation largely of temperate character, but the present of cooler elements and some differences in biostratigraphic ranges have required a slightly different biostratigraphic scheme.Although two broad schemes are recognized, none of the biostratigraphic sequences are identical between any of the sites. This reflects differences in biogeography, evolution and diachronous extinction at various latitudes during the entire Neogene. Diachronism in biostratigraphic ranges continue to create difficulties in correlation across such wide latitudes.Our detailed work has required the establishment of new biostratigraphic zonations in certain parts of the Neogene sequence and modifications in some other parts. Otherwise, previously established schemes are followed as closely as possible. In the temperate region, a new zonation has been established for the Early Miocene to early Middle Miocene. For the remainder of the Neogene the zonation of Kennett (1973) has been largely used. The tropical zonation of Blow (1969) is employed in the equatorial Site 289, but with further subdivisions for Zones N4 and N17. For areas intermediate between tropical and temperate latitudes (Site 208), a modified Early Miocene zonation is established based on changes in tropical and temperate elements.The zonal schemes are established on taxa that exhibit both diachronous and isochronous ranges across the latitudes. Zones that are at least partly diachronous include the Globigerinoides trilobus and Globorotalia miozea Zones of Early Miocene age; perhaps the Globorotalia mayeri Zone (its base) of the Middle Miocene; the Globorotalia conomiozea Zone of the Late Miocene; and the Globorotalia crassaformis Zone of the Early Pliocene.A large number of datum levels are recognized based on first evolutionary appearances or extinctions. The most widely applicable datums are as follows: latest Oligocene — Globigerinoides F.A.; Early Miocene — Globoquadrina dehiscens, F.A., Globorotalia kugleri L.A., Catapsydrax dissimilis L.A. and Praeorbulina glomerosa F.A.; Middle Miocene — Orbulina suturalis F.A., Globorotalia peripheroacuta F.A., Fohsella lineage L.A., Globorotalia mayeri L.A.; Late Miocene — “Neogloboquadrina” continuosa L.A., Globoquadrina dehiscens L.A., Globorotalia cibaoensis F.A.; Early Pliocene — Globorotalia puncticulata F.A., Globorotalia margaritae F.A.; Early Pleistocene — Globorotalia truncatulinoides F.A. A number of other datums are identified which assist with correlation over more restricted latitudinal ranges.The evolution of most Neogene planktonic foraminifera is now well established for a wide range of water masses. Evolutionary lineages are primarily centered in the temperate and tropical regions. Tropical lineages have recently been reviewed by Srinivasan and Kennett (1981) and are not discussed in detail here. However, Sphaeroidinellopsis seminulina is now considered to have evolved directly into S. paenedehiscens during the Late Miocene and S. subdehiscens Blow is considered to be junior synonym of S. seminulina.A new evolutionary lineage is recognized in the warm subtropics (Site 208) whereby Globigerina woodi woodi gave rise to Globigerinoides subquadratus via Globigerina brazieri. The discovery of this lineage clearly demonstrates that Globigerinoides is a polyphyletic “genus”. Another major phylogenetic lineage is recognized within the temperate globorotaliids of Early Miocene age as follows: “N.” continuosa → Globorotalia zealandica incognita → G. zealandica zelandica → G. praescitula → G. miozea. Although parts of this lineage have been recognized earlier, the entire phylogeny has previously been underscribed.A new Early to Middle Miocene lineage is recognized in the subantarctic to temperate areas which involve a transition from Globorotalia praescitula to G. challengeri n. sp. via intermediate forms.Two major Neogene globorotaliid lineages — the Menardella of the tropics and Middle Miocene to Recent forms of Globoconella of the temperate areas — are both considered to have evolved from Globorotalia praescitula beginning in the Early Miocene. This evolution initially was restricted to temperate areas but has since separated into distinctly tropical and temperate phylogenetic elements.     

First appearance of Globorotalia truncatulinoides: cladogenesis and immigration

The first appearance datum of the planktic foraminifer Globorotalia truncatulinoides is widely used to identify the base of the Quaternary Period. However, its appearance is not globally isochronous. We have previously shown that G. truncatulinoides evolved gradually from its ancestor Globorotalia crassaformis via the intermediate species Globorotalia tosaensis. This cladogenesis was documented in the southwest Pacific during the Late Pliocene in sympatric or parapatric populations. Based on qualitative observations, similar but younger, gradual transitions have been reported from other areas of the world's oceans. Therefore, this gradual evolutionary branching might have occurred in response to changing environments at different times in different ocean areas. To evaluate the hypothesis of a repeated, environmentally driven gradual, sympatric cladogenesis, we studied the morphological transitions of the three taxa, using image analytical techniques, in several deep-sea sections from various areas. Our study confirms that G. truncatulinoides evolved between 2.8 and 2.3 Ma sympatrically in large populations from its ancestor G. crassaformis in the southwest Pacific. Differentiated morphotypes of G. truncatulinoides subsequently immigrated into the Indian and Atlantic oceans between 2.3 and 1.9 Ma. Our morphometric data show these younger appearances outside the southwest Pacific to be punctuated.We hypothesize that the global cooling of surface waters, coinciding with the northern hemisphere glaciation, led to the formation of oceanographic barriers that could have retarded the expansion of G. truncatulinoides up to 2.3 Ma. At this time, a relative warming and subsequent transgression could have spurred the migration, possibly through the Indonesian passage. A direct link between the speciation and surface water changes linked to the northern hemisphere glaciation has not been proven so far and seems unlikely. In fact, stable isotope data in this lineage indicate that the three species' depth habitat preferences remained unchanged through the speciation and migration of G. truncatulinoides and that all three species were dominantly deep-dwellers, in agreement with their present environmental preferences.     

Establishment of the western Pacific warm pool during the Pliocene: Evidence from planktic foraminifera, oxygen isotopes, and Mg/Ca ratios

This study presents new evidence of when and how the Western Pacific Warm Pool (WPWP) was established in its present form. We analyzed planktic foraminifera, oxygen isotopes, and Mg/Ca ratios in upper Miocene through Pleistocene sediments collected at Deep Sea Drilling Program (DSDP) Site 292. These data were then compared with those reported from Ocean Drilling Program (ODP) Site 806. Both drilling sites are located in the western Pacific Ocean. DSDP Site 292 is located in the northern margin of the modern WPWP and ODP Site 806 near the center of the WPWP. Three stages of development in surface-water conditions are identified in the region using planktic foraminferal data. During the initial stage, from 8.5 to 4.4 Ma, Site 806 was overlain by warm surface water but Site 292 was not, as indicated by the differences in faunal compositions and sea-surface temperature (SST) between the two sites. In addition, the vertical thermal gradient at Site 292 was weak during this period, as indicated by the small differences in the δ¹⁸O values between Globigerinoides sacculifer and Pulleniatina spp. During stage two, from 4.4 to 3.6 Ma, the SST at Site 292 rapidly increased to 27 °C, but the vertical thermal gradient had not yet be strengthened, as shown by Mg/Ca ratios and the presence of both mixed-layer dwellers and thermocline dwellers. Finally, a warm mixed layer with a high SST ca. 28 °C and a strong vertical thermal gradient were established at Site 292 by 3.6 Ma. This event is marked by the dominance of mixed-layer dwellers, a high and stable SST, and a larger differences in the δ¹⁸O values between G. sacculifer and Pulleniatina spp. Thus, evidence of surface-water evolution in the western Pacific suggests that Site 292 came under the influence of the WPWP at 3.6 Ma. The northward expansion of the WPWP from 4.4 to 3.6 Ma and the establishment of the modern WPWP by 3.6 Ma appear to be closely related to the closure of the Indonesian and Central American seaways.     

Late miocene magnetobiostratigraphy of Crete

Six Upper Miocene marine clay sections on Crete (Greece) have been subjected to a detailed magnetobiostratigraphic analysis. Six correlatable polarity zones are recognized and these demonstrate the regional synchrony of planktonic foraminiferal biohorizons.By way of correlation to the magnetic polarity time-scale of Lowrie and Alvarez (1981), the Cretan sequence is assigned to polarity chronozones 5 (anomaly 3A) and 6. The new chronology provides an age of 5.6 Ma for the first occurrence datum (FOD) of theGloborotalia conomiozea group in the Mediterranean, an age of 6.0 Ma for the FOD ofG. menardii form 5 and an age of 6.6 Ma for the last occurrence datum (LOD) ofG. menardii form 4.Correlating the polarity record of the New Zealand Blind River section (Kennett and Watkins, 1974), with the magnetic polarity time-scale provides an age of 6.0 Ma for the evolutionary appearance ofGloborotalia conomiozea, which is in complete agreement with the age of 6.1 ± 0.1Ma given by Loutit and Kennett (1979). The demonstrated diachrony of 0.4 Ma between the New Zealand FOD ofG. conomiozea and its Mediterranean counterpart is explicable in view of the different nature of the two events, the one in New Zealand being evolutionary and the one in the Mediterranean migrational.The FOD of theG. conomiozea group in the Tortonian/Messinian boundary stratotype section coincides with the level proposed by Colalongo et al. (1979) to mark the base of the Messinian. Since the FOD of theG. conomiozea group in Crete and in Sicily are most probably time-equivalent, the age of the Tortonian/Messinian boundary is fixed at 5.6 Ma.The youngest sediments incorporated in this study extend into the Gilbert chronozone and antedate the main evaporitic phase. Consequently, the Messinian evaporitic body is younger than the base of the Gilbert chronozone, the age of which is fixed at 5.3 Ma. Adopting an age of 5.0 Ma for the Miocene/Pliocene boundary would imply that evaporites and post-evaporitic Lago Mare sediments were deposited in some 300,000 years and suggests that in the central parts of the Mediterranean basins evaporites must have accumulated at rates of some 3 m per 1000 years.     

Miocene biochronology and paleoceanography of the North Pacific

Biostratigraphic correlation based on microfossil datum levels, directly or indirectly tied to the paleomagnetic time scale, provides a high resolution time control for the Miocene in the equatorial and middle latitude North Pacific. Faunal changes and abundance fluctuations of planktic foraminiferal species combined with the oxygen Pacific. Faunal changes and abundance fluctuations of planktic foraminiferal species combined with the oxygen isotope record of foraminifers, reveal the paleoclimatic and paleoceanographic history. The planktic foraminiferal assemblage change in the early Miocene, extinction of Oligocene fauna and rise of a highly diverse Neogene fauna, appears to be related to increased water mass stratification in the world oceans presumably resulting from the establishment of circum-Antarctic circulation. An increase in the siliceous productivity in the eastern equatorial Pacific region between 20 and 18 Ma suggests that the vertical and horizontal circulation was intensified at that time. Climates cooled rapidly during the middle Miocene between 14 and 13 Ma suggesting the growth of a major east Antarctic ice sheet. Paleoclimatic conditions remained generally cool, although oscillating, during the late Miocene. In the late early to middle Miocene faunal provincialism developed between low and middle latitudes, and by late Miocene time a distinct provincialism similar to the present was established.     

Fertility tracers and monsoon forcing at an equatorial site of the Somali Basin (Northwest Indian Ocean)

Temporal changes in biological and isotopic tracers have successfully been used to monitor monsoon variability in the Arabian Sea. Convincing evidence that fertility indicators, such as the relative abundance of the planktic foraminifer Globigerina bulloides, and SiO₂ and Ba fluxes, can monitor monsoon changes, has also been published. Time series of the ¹³C content of the thermocline foraminifer Neogloboquadrina dutertrei, and of an Upwelling Radiolarian Index (URI), have been used to reconstruct upwelling changes in the Somali Basin at 5 °N for the last 160 kyr. In order to establish a reference site for comparison with other upwelling and/or fertility records from the same area, a time series of 8 climatic and fertility proxies (¹³C content of N. dutertrei and Globorotalia menardii, percentages of G. bulloides and of a thermocline foraminiferal group, minimal δ¹⁸ values of Globigerinoides sacculifer, N. dutertrei and G. menardii) are investigated in the spectral domain for the last 360 kyr, at a site located at the equator and outside the Somali upwelling centers (Core MD 85668).Chronostratigraphies for these records are developed by correlation of oxygen isotope record of G. sacculifer to the Martinson standard stacked benthic oxygen isotope record. The temporal resolution of the data, is between 2 and 5 kyr.Cross spectral comparisons with ETP (Eccentricity + Tilt + Precession-composite signal) were used to estimate coherences in conjunction with phase relationships and to quantify relationships between fertility and climatic indicators. The timing of fertility indicator cycles at the equator shows little (obliquity band) or no (other orbital bands) coincidence with monsoon proxies of the Arabian Sea. At the equator, fertility responses are dissociated, and present important leads or lags with ETP signal at the three orbital bands (with the exception of the thermocline foraminifera, in phase with maximum eccentricity). Due to significant leads with ETP in the timing of ¹³C minima and URI cycles, no relationships with the Northern Hemisphere radiation can be deduced. At the obliquity and precessional frequency bands, thermocline foraminifera cycles display significant lag with ETP and ice volume minima, which suggests that radiation is not the sole forcing mechanism. Global climatic and ice volume changes may be an important forcing mechanism for these fertility proxies. The G. bulloides record is more complicated with cycles in phase with Arabian Sea monsoon proxies at the obliquity band, and ¹⁸O minima at the precession frequency.It appears that, at the equator, the southwest monsoon winds are not the main force driving radiolarian and foraminiferal productivity.     

Globorotalia puncticulata: Population divergence, dispersal and extinction related to Pliocene–Quaternary water masses

The isolating effect of water mass partitioning of populations on the morphology, stratigraphic distribution and extinction of planktonic foraminifera is assessed from the Pliocene–Quaternary record of Globorotalia puncticulata. Southern Hemisphere, Mediterranean and North Atlantic data on these aspects of its history are examined and appear consistent with a limited dispersal biogeographic model wherein populations are largely confined by hydrographic barriers.Earliest populations appeared during the latest Miocene in Southern Hemisphere middle latitude water masses. However, morphometric analysis shows that significant differentiation in the axial shape of shells had developed by 4 Ma between Southwest Pacific populations from ODP Site 1123 (temperate water) and ODP Site 1119 (subantarctic water). These sites are in close proximity but separated by the Subtropical Front. At Site 1123 inflation of late-formed chambers and reduction in the number included in the outer whorl created shell profiles that anticipated the globose form of Globorotalia inflata. The latter's gradual evolution from G. puncticulata s.s. took place in this temperate water mass, with the earliest morphotypes with three chambers in the outer whorl present by 4.1 Ma. In contrast, subantarctic populations at Site 1119 retained four chambers but their axial shape was modified. The development of a large, highly arched aperture and increase in the number of chambers in the outer whorl in Mediterranean–North Atlantic Globorotalia puncticulata bononiensis is an example of population differentiation later in the Pliocene.Chronostratigraphy shows that the northward expansion of central temperate water populations commenced with their occupation of Southwest Pacific subtropical water about 4.8 Ma. The rather abrupt entry of substantial populations into Mediterranean and North Atlantic water at 4.5 Ma marked a major biogeographic expansion and established G. puncticulata as a bipolar species. It was widely distributed about 3 Ma, with major populations in several water masses during a period of middle Pliocene warmth.After this acme North Atlantic and Mediterranean G. puncticulata bononiensis populations collapsed as late Pliocene Northern Hemisphere glacials intensified. They were extinguished in MIS 96 (2.4 Ma). Concurrently, G. puncticulata s.s became extinct in the warm subtropical Southwest Pacific. Subantarctic populations persisted but in turn were decimated in severe glacials during the Middle Pleistocene Transition. Most had disappeared by MIS 16 (0.66 Ma). However, at Sites 594 and 1119 there was a small Lazarus-like revival in MIS 11 (0.41 Ma). The highest known occurrence is in MIS 9 (0.33 Ma) at Site 1119. Confinement of the species to subantarctic water in the Pleistocene may have raised its vulnerability to extinction.While stable isotope data indicate that the lineage's evolution is related to depth habitat selection about the thermocline, its biogeography suggests that hydrographic barriers significantly isolated populations and probably facilitated speciation. Eddies such as the North Brazil Current rings provide conduits for inter-water mass transfer of populations but the history of G. puncticulata suggests that such mechanisms seldom operated successfully.The morphology of the lectotype of G. puncticulata s.s., from beach sand at Rimini, Italy, is consistent with a lower Pliocene source. Reports of living occurrences are poorly documented and the species is considered to be extinct.     

Small mammal (rodents and lagomorphs) European biogeography from the Late Oligocene to the mid Pliocene

Aim To analyse the fossil species assemblages of rodents and lagomorphs from the European Neogene in order to assess what factors control small mammal biogeography at a deep‐time evolutionary time‐scale. Location Western Europe: 626 fossil‐bearing localities located within 31 regions and distributed among 18 successive biochronological units ranging from c. 27 Ma (million years ago; Late Oligocene) to c. 3 Ma (mid Pliocene). Methods Taxonomically homogenized pooled regional assemblages are compared using the Raup and Crick index of faunal similarity; then, the inferred similarity matrices are visualized as neighbour‐joining trees and by projecting the statistically significant interregional similarities and dissimilarities onto palaeogeographical maps. The inferred biogeographical patterns are analysed and discussed in the light of known palaeogeographical and palaeoclimatic events. Results Successive time intervals with distinct biogeographical contexts are identified. Prior to c. 18 Ma (Late Oligocene and Early Miocene), a relative faunal homogeneity (high interregional connectivity) is observed all over Europe, a time when major geographical barriers and a weak climatic gradient are known. Then, from the beginning of the Middle Miocene onwards, the biogeography is marked by a significant decrease in interregional faunal affinities which matches a drastic global climatic degradation and leads, in the Late Miocene (c. 11 Ma), to a marked latitudinal pattern of small mammal distribution. In spite of a short rehomogenization around the Miocene/Pliocene boundary (6–4 Ma), the biogeography of small mammals in the mid Pliocene (c. 3 Ma) finally closely reflects the extant situation. Main conclusions The resulting biogeographical evolutionary scheme indicates that the extant endemic situation has deep historical roots corresponding to global tectonic and climatic events acting as primary drivers of long‐term changes. The correlation of biogeographical events with climatic changes emphasizes the prevalent role of the climate over geography in generating heterogeneous biogeographical patterns at the continental scale.     

Stepwise mass extinctions and impact events: Late Eocene to early Oligocene

Species ranges and relative abundances of dominant planktonic foraminifers of eight late Eocene to early Oligocene deep-sea sections are discussed to determine the nature and magnitude of extinctions and to investigate a possible cause-effect relationship between impact events and mass extinctions.Late Eocene extinctions are neither catastrophic nor mass extinctions, but occur stepwise over a period of about 1–2 million years. Four stepwise extinctions are identified at the middle/late Eocene boundary, the upperGlobigerapsis semiinvoluta zone, theG. semiinvoluta/Globorotalia cerroazulensis zone boundary and at the Eocene/Oligocene boundary. Each stepwise extinction event represents a time of accelerated faunal turnover characterized by generally less than 15% species extinct and in itself is not a significant extinction event. Relative species abundance changes at each stepwise extinction event, however, indicate a turnover involving > 60% of the population implying major environmental changes.There microtektite horizons are present in late Eocene sediments; one in the upperG. semiinvoluta zone (38.2 Ma) and two closely spaced layers only a few thousand years apart in the lower part of theGloborotalia cerroazulensis zone (37.2 Ma). Each of the three impact events appears to have had some effect on microplankton communities. However, the overriding factor that led to the stepwise mass extinctions may have been the result of multiple causes as there is no evidence of impacts associated with the step preceding, or the step following the deposition of the presently known microtektite horizons.     

The biostratigraphy and paleobiology of Oligocene planktonic foraminifera from the equatorial Pacific Ocean (ODP Site 1218)

Planktonic foraminifera from a continuous Oligocene succession with clear magnetochronology and sediment cycles at Ocean Drilling Program Site 1218 (equatorial Pacific Ocean) were studied in the interval from 27 to 30 Ma. Paragloborotalia taxa are common and we examined their size, relative abundance, and stable isotopes. Multispecies stable isotope data indicate the depth habitats of Oligocene planktonic foraminifera and suggest that “Globoquadrina” venezuelana and Dentoglobigerina globularis were probably mixed-layer dwellers, with paragloborotaliids recording heavier δ¹⁸O signatures consistent with a thermocline habitat. Cyclic variations in the abundance of Paragloborotalia match eccentricity (100 kyr) variations in percent carbonate and δ¹³C, suggesting orbitally forced upwelling in the equatorial Pacific Ocean and that Paragloborotalia were responding directly to changes in surface water productivity. The high-resolution biostratigraphy calibrated to the magnetochronology constrains the extinction of Paragloborotalia opima which marks the top of Planktonic Foraminifera Biozone O5 (P21b) at 27.456 Ma. The highest occurrence of P. opima is associated with a 50% size decrease in Paragloborotalia pseudocontinuosa taxa within Chron 9n. In addition, we find the extinction of Chiloguembelina cubensis is consistent with other deep-sea sections within Chron 10n at 28.426 Ma marking the O4/O5 (P21a/P21b) boundary.     

Surface-water cooling and salinity decrease during the Middle Miocene climate transition at Southern Ocean ODP Site 747 (Kerguelen Plateau)

Paleoceanographic variability at southern high latitude Ocean Drilling Program (ODP) Site 747 was investigated in this study through the interval which spans the Middle Miocene Climate Transition (MMCT). Between 15.0 and 12.2 million years ago (Ma), foraminiferal δ¹⁸O records derived from both benthic (Cibicidoides spp.) and planktonic taxa (Globorotalia praescitula and Globigerina bulloides) reveal a history of changes in water column thermal and salinity structure and a strong imprint of seasonality. Prior to the MMCT, in the interval between 14.35 and 13.9 Ma, G. bulloides displays relatively high δ¹⁸O values similar to those of G. praescitula, interpreted to indicate weakening of the thermocline and/or increased seasonality with cooler early-spring and/or late-fall temperatures. Following this interval, G. bulloides δ¹⁸O values diverge significantly from benthic and G. praescitula values, with G. bulloides values remaining relatively low for at least 600 kyr following the benthic foraminiferal δ¹⁸O shift during the MMCT at ~ 13.9 Ma. This divergence in δ¹⁸O records occurs in direct association with the Mi3 cooling and glaciation event and may suggest: (1) a strengthening of the vertical temperature gradient, with greater cooling of deep waters than surface waters, (2) changes in the depth habitat of G. bulloides, (3) changes in the dominant season of G. bulloides calcification, (4) modification of surface-water δ¹⁸O values in association with enhanced sea-ice formation, (5) increased surface-water carbonate ion concentration, and/or (6) a significant decrease in surface-water salinity across the MMCT. The first of these possible scenarios is not likely, particularly in light of recent Mg/Ca evidence for significant surface-water cooling in the Southern Ocean associated with the MMCT. Of the remaining possibilities, we favor a change in surface salinity to explain the observed trends in δ¹⁸O values and hypothesize that surface salinity may have decreased by up to 2 salinity units at ~ 13.9 Ma. In this scenario, the development of a lower-salinity Antarctic surface layer coincided with regional cooling of both surface and deep waters of the Southern Ocean during the Mi3 glaciation of East Antarctica, and contributed into the dominance of Neogloboquadrina spp. between 13.8 and 13.2 Ma. Additionally, the distinct patterns observed in planktonic foraminiferal δ¹⁸O records spanning the MMCT correspond with changes in the vertical δ¹³C gradient between planktonic and benthic foraminiferal records and major changes in planktonic foraminiferal assemblages at Site 747, providing further evidence of the environmental significance of this climatic transition.     

The replacement of the“Globorotalia menardii” group by theGloborotalia miotumida group: An aid to recognizing the Tortonian-Messinian boundary in the Mediterranean and adjacent Atlantic

Different authors working in the Mediterranean area have reported a distinctive change in the assemblages of keeled globorotaliids coinciding with the FAD ofGloborotalia conomiozea Kennett. This change which has now been observed in the Guadalquivir basin consists of the abrupt disappearance of the “Globorotalia menardii” group, which is then replaced by theGloborotalia miotumida group.This event has been found in a number of areas of the Atlantic Ocean and it is proposed that it can be used to recognize the Tortonian-Messinian boundary in this realm whereG. conomiozea occurs only sporadically.     

Paleocene-Pleistocene benthic foraminiferal evidence of major paleoceanographic events in the eastern South Atlantic (DSDP Site 525, Walvis Ridge)

Benthic foraminifers in the size-fraction greater than 0.073 mm were studied in 88 Paleocene to Pleistocene samples from Deep Sea Drilling Project Site 525 (Hole 525A, Walvis Ridge, eastern south Atlantic). Clustering of the samples on the basis of the 86 most abundant foraminifers (in total, 331 taxa were identified) allowed separating two major assemblage zones: the Paleocene to Eocene interval, and the Oligocene to Pleistocene interval. Each of these, in turn, were subdivided into three minor subzones as follows: lower upper Paleocene (approx. 62.4 to 57.8 Ma); upper upper Paleocene (56.6 to 56.2 Ma); lower and middle Eocene (55.3 to 46.8 Ma); upper Oligocene to middle Miocene (25.3 to 16 Ma); middle Miocene to Pliocene (15.7 to 4.2 Ma); and lower Pleistocene (0.4 to 0.02 Ma), with only minor differences with the previous zone. Some very abundant taxa span most of the column studies (Bolivina huneri, Cassidulina subglobosa, Eponides bradyi, E. weddellensis, Gavelinella micra, Oridorsalis umbonatus, etc.). Several of the faunal breaks recorded coincide with conspicuous minima in the specific diversity curve, thus suggesting that the corresponding turnovers signal the final stages of periods of faunal impoverishment. At least one major bottom-water temperature drop (as derived from δ¹⁸O data) is synchronous with a decrease in the foraminiferal specific diversity. On the other hand, a specific diversity maximum in the middle Miocene might be associated with a δ¹³C increase at approx. 16 to 12 Ma. Highest foraminiferal abundances (up to 600–800 individuals per gram of dry sediment) occurred in the late Paleocene and in the early Pleistocene, in coincidence with the lowest diversity figures calculated. The magnitude of the most important faunal turnover recorded, between the middle Eocene and the late Oligocene, is magnified in our data set by the large hiatus which separates the middle Eocene from the upper Oligocene sediments. Considerably smaller overturns occurred within the late Paleocene (in coincidence with changes in the specific diversity, absolute abundance of foraminiferal tests, and δ¹³C), and in the middle Miocene (in coincidence with a specific diversity maximum and a δ¹³C excursion). New information on the morphology and the stratigraphic ranges of several species is furnished. For all the taxa recorded the number of occurrences, total number of individuals identified and first and last appearances are listed.     

Middle Eocene–Early Pliocene Subantarctic planktic foraminiferal biostratigraphy of Site 1090, Agulhas Ridge

The analysis of planktic foraminiferal assemblages from Site 1090 (ODP Leg 177), located in the central part of the Subantarctic Zone south of South Africa, provided a geochronology of a 330-m-thick sequence spanning the Middle Eocene to Early Pliocene. A sequence of discrete bioevents enables the calibration of the Antarctic Paleogene (AP) Zonation with lower latitude biozonal schemes for the Middle–Late Eocene interval. In spite of the poor recovery of planktic foraminiferal assemblages, a correlation with the lower latitude standard planktic foraminiferal zonations has been attempted for the whole surveyed interval. Identified bioevents have been tentatively calibrated to the geomagnetic polarity time scale following the biochronology of Berggren et al. (1995). Besides planktic foraminiferal bioevents, the disappearance of the benthic foraminifera Nuttallides truempyi has been used to approximate the Middle/Late Eocene boundary. A hiatus of at least 11.7 Myr occurs between 78 and 71 m composite depth extending from the Early Miocene to the latest Miocene–Early Pliocene. Middle Eocene assemblages exhibit a temperate affinity, while the loss of several planktic foraminiferal species by late Middle to early Late Eocene time reflects cooling. During the Late Eocene–Oligocene intense dissolution caused impoverishment of planktic foraminiferal assemblages possibly following the emplacement of cold, corrosive bottom waters. Two warming peaks are, however, observed: the late Middle Eocene is marked by the invasion of the warmer water Acarinina spinuloinflata and Hantkenina alabamensis at 40.5 Ma, while the middle Late Eocene experienced the immigration of some globigerinathekids including Globigerinatheka luterbacheri and Globigerinatheka cf. semiinvoluta at 34.3 Ma. A more continuous record is observed for the Early Miocene and the Late Miocene–Early Pliocene where planktic foraminiferal assemblages show a distinct affinity with southern mid- to high-latitude faunas.     

Carnivoran resource and habitat use in the context of a Late Miocene faunal turnover episode

We investigate resource and habitat use by apex predators through stable isotope analysis at two Spanish Late Miocene localities: Los Valles de Fuentidueña (~9.6 Ma, LVF) and Cerro de los Batallones (~9.1 Ma, BAT). The temporal window represented by LVF and BAT was crucial in the shaping of the current Iberian mammalian structure because it corresponds to the initial stages of a faunal turnover episode and regional environmental change at ~9.5–8.5 Ma (Vallesian–Turolian transition), associated with an increase in the seasonality of precipitation. Herbivore and carnivore δ¹³C and δ¹⁸O values do not point to significant changes in either the vegetation cover (a woodland to mesic C₃ grassland) or the hydrological regime during the time lapse represented between LVF and BAT. This suggests that the environmental shift recorded around the Vallesian–Turolian boundary may have occurred later in time, since LVF and BAT ages are synchronic with the onset of the turnover event. From the standpoint of predator–prey evaluation by means of stable isotope analysis, statistical post hoc tests, mixing model output, and the assessment of niche occupation by LVF and BAT carnivores point to high levels of interspecific competition among large active carnivores, albeit some genera, such as the amphicyonid Magericyon and specially the hyaenid Lycyaena, seemed to avoid competition by taking prey from a more open habitat. Despite the drop in diversity and change in faunal components observed between the LVF and BAT assemblages, a high degree of resource and habitat competition is evident from stable isotope data.     

Progenesis in Late Eocene populations ofSubbotina linaperta (Foraminifera) from the western Atlantic

It has been argued that the successive appearance of a distinct set of test morphologies by planktic foraminifera represents evidence for the multiple evolution of mechanical optima (Steineck and Fleisher, 1978). However, morphometric, isotopic and biostratigraphic analyses of the globigerine speciesSubbotina linaperta from Middle-Late Eocene Atlantic Ocean deep sea cores suggests that changes in developmental pathways may also play an important role in planktic foraminiferal evolution. During a period of prolonged and global change in the Middle-Late Eocene marine environment, at least one western Atlantic population ofS. linaperta was characterized by a marked decrease in mean test size that persisted throughout the remainder of this species' teilzone. In addition, this Late Eocene population exhibited anomalous relative abundances and a pronounced change in depth habitat when compared to conspecific populations in the Gulf of Mexico and South Atlantic. Quantitative morphometric analyses indicate the observed size reduction to be a secondary result of selection for the ontogenetically accelerated attainment of sexual maturity; a mode of developmentally mediated evolution termed progenesis. The temporal persistence of this progenetically dwarfed population throughout the remainder of the Late Eocene at this locality serves to illustrate the potential of this evolutionary mechanism to produce sustained morphological and ecological changes within populations of marine plankton as well as implying that this population was at least partially isolated from the general circulation of the Gulf Stream during this time interval. In addition, similarities between the nature of phenotypic change in this population and the common anecdotal observation of test size reduction in a number of planktic foraminiferal lineages at the Cretaceous-Tertiary boundary and during the Paleogene-Neogene faunal transition suggest that selection for different developmental patterns may provide an alternative explanation for the reappearance of simple globigerine morphotypes after major planktic foraminiferal extinction events.     

Impact of climate changes from Middle Miocene onwards on evolutionary diversification in Eurasia: Insights from the mesobuthid scorpions

The aridification from Middle Miocene onwards has transformed the Asian interior into an arid environment, and the Pleistocene glacial–interglacial oscillations exerted further ecological impact. Therefore, both aridification and glaciation would have considerably influenced the evolution of many mid‐latitude species in temperate Asia. Here, we tested this perspective by a phylogeographic study of the mesobuthid scorpions across temperate Asia using one mitochondrial and three nuclear genes. Concordant mitochondrial and nuclear gene trees were obtained, which are consistent with species tree inferred using a Bayesian approach. The age of the most recent common ancestor (MRCA) of all the studied scorpions was estimated to be 12.49 Ma (late Middle Miocene); Mesobuthus eupeus diverged from the clade composing Mesobuthus caucasicus and Mesobuthus martensii in early Late Miocene (10.21 Ma); M. martensii diverged from M. caucasicus at 5.53 Ma in Late Miocene. The estimated MRCA ages of M. martensii and the Chinese lineage of M. eupeus were 2.37 and 0.68 Ma, respectively. Central Asia was identified as the ancestral area for the lineage leading to M. martensii and M. caucasicus and the Chinese lineage of M. eupeus. The ancestral habitat of the genus Mesobuthus is likely to have been characterized by an arid environment; a shift towards more humid habitat occurred in the MRCA of M. martensii and a lineage of M. caucasicus, finally leading to the adaptation of M. martensii to humid environment. Our data strongly support the idea that the stepwise intensified aridifications from Mid‐Miocene onwards drove the diversification of mesobuthid scorpions, and suggest that M. martensii and M. eupeus observed today in China originated from an ancestral lineage distributed in Central Asia. Both the colonization and the ensuing evolution of these species in East Asia appear to have been further moulded by Quaternary glaciations.     

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.