Planktonic foraminifera in the Maastrichtian of Hor-Hahar

The lengthy warm, stable climate of the Cretaceous terminated in the Campanian with a cooling trend, interrupted in the early and latest Maastrichtian by two events of global warming, at ~70-68 Ma and at 65.78-65.57 Ma. These climatic oscillations had a profound effect on pelagic ecosystems, especially on planktic foraminiferal populations. Here we compare biotic responses in the tropical-subtropical (Tethyan) open ocean and mesotrophic (Zin Valley, Israel) and oligotrophic (Tunisia) slopes, which correlate directly with global warming and cooling. The two warming events coincide with blooms of Guembelitria, an extreme opportunist genus best known as the main survivor of the Cretaceous-Paleogene (K-Pg) catastrophe. In the Maastrichtian, Guembelitria bloomed in the uppermost surface water above shelf and slope environments but failed to reach the open ocean as it did at K-Pg. The coldest interval of the late Maastrichtian (~68-65.78 Ma) is marked by an acme of the otherwise rare species Gansserina gansseri, a deep-dwelling keeled globotruncanid. The G. gansseri acme event can be traced from the deep ocean even onto the Tethyan slope, marking copious production and circulation of cold intermediate water. This acme is abruptly terminated by extinction of the species, a dramatic reversal attributed to a short-term global warming episode. This extinction corresponds precisely with the second bloom of Guembelitria that began ~300 kyr prior to the K-Pg event. The antithetical relationship between blooming of Guembelitria and the G. gansseri acme reflects planktic foraminiferal sensitivity to warm-cool-warm-cool climatic oscillations marking the end of the Cretaceous.

Stable carbon and oxygen isotope record of Cretaceous planktonic foraminifera from DSDP (Table 1)

The Cretaceous Heterohelix moremani (Cushman) was the only biserial planktonic foraminiferal species from its first appearance in the late Albian up to the Cenomanian/Turonian boundary. Within that time, it increased gradually in abundance relative to other planktonic foraminifera in five Circum-North Atlantic sections. It is generally rare in upper Albian sediments, common in most of the Cenomanian and very abundant in sediments representing the latest Cenomanian Oceanic Anoxic Event. Short-term variations on the overall abundance trend correlate with positive excursions in the bulk carbonate delta13C record. Maximum rain rates of H. moremani during OAE2 show that this species was an opportunist that did well in extreme conditions, but its overall distribution indicates that it is not necessarily a marker for very high palaeoproductivity environments. Stable oxygen and carbon isotope measurements on foraminiferal species indicate that H. moremani was a surface water dweller at least in part of its geographic range, but incorporated 13C out of equilibrium with ambient seawater. It is depleted in delta13C relative to other planktonic foraminifera, which is attributed to vital effects related to its opportunistic character.

Maastrichtian calcareous nannofossils of the Indian Ocean and Shatsky Rise

The biotic effects of volcanism have long been the unknown factors in creating biotic stress, and the contribution of the Deccan volcanism to the K-T mass extinction remains largely unknown. Detailed studies of the volcanic-rich sediments of Indian Ocean Ninetyeast Ridge Sites 216 and 217 and Wharton Basin Site 212 reveal that the biotic effects of late Maastrichtian volcanism on planktic foraminifera and calcareous nannofossils are locally as severe as those of the K-T mass extinction. The biotic expressions of these high stress environments are characterized by the Lilliput effect, which includes reduced diversity by eliminating most K-strategy species, and reduction in specimen size (dwarfing), frequently to less than half their normal adult size of both r-strategy and surviving K-strategy species. In planktic foraminifera, the most extreme biotic stress results are nearly monospecific assemblages dominated by the disaster opportunist Guembelitria, similar to the aftermath of the K-T mass extinction. The first stage of improving environmental conditions results in dominance of dwarfed low oxygen tolerant Heterohelix species and the presence of a few small r-strategy species (Hedbergella, Globigerinelloides). Calcareous nannofossil assemblages show similar biotic stress signals with the dominance of Micula decussata, the disaster opportunist, and size reduction in the mean length of subordinate r-strategy species particularly in Arkhangelskiella cymbiformis and Watznaueria barnesiae. These impoverished and dwarfed late Maastrichtian assemblages appear to be the direct consequences of mantle plume volcanism and associated environmental changes, including high nutrient influx leading to eutrophic and mesotrophic waters, low oxygen in the water column and decreased watermass stratification.

Benthic foraminiferal accumulation rates of the late Pliocene-Pleistocene in the northern Indian Ocean

During the late Pliocene-middle Pleistocene, 63 species of elongate, bathyal-upper abyssal benthic foraminifera (Extinction Group = Stilostomellidae, Pleurostomellidae, some Nodosariidae) declined in abundance and finally disappeared in the northern Indian Ocean (ODP Sites 722, 758), as part of the global extinction of at least 88 related species at this time. The detailed record of withdrawal of these species differs by depth and geography in the Indian Ocean. In northwest Indian Ocean Site 722 (2045 m), the Extinction Group of 54 species comprised 2-15% of the benthic foraminiferal fauna in the earliest Pleistocene, but declined dramatically during the onset of the mid-Pleistocene Transition (MPT) at 1.2-1.1 Ma, with all but three species disappearing by the end of the MPT (~0.6 Ma). In northeast Indian Ocean Site 758 (2925 m), the Extinction Group of 44 species comprised 1-5% of the benthic foraminiferal fauna at ~3.3-2.6 Ma, but declined in abundance and diversity in three steps, at ~2.5, 1.7, and 1.2 Ma, with all but one species disappearing by the end of the MPT. At both sites there are strong positive correlations between the accumulation rate of the Extinction Group and proxies indicating low-oxygen conditions with a high organic carbon input. In both sites, there was a pulsed decline in Extinction Group abundance and species richness, especially in glacial periods, with some partial recoveries in interglacials. We infer that the glacial declines at the deeper Site 758 were a result of increased production of colder, well-ventilated Antarctic Bottom Water (AABW), particularly in the late Pliocene and during the MPT. The Extinction Group at shallower water depths (Site 722) were not impacted by the deeper water mass changes until the onset of the MPT, when cold, well-ventilated Glacial North Atlantic Intermediate Water (GNAIW) production increased and may have spread into the Indian Ocean. Increased chemical ventilation at various water depths since late Pliocene, particularly in glacial periods, possibly in association with decreased or more fluctuating organic carbon flux, might be responsible for the pulsed global decline and extinction of this rather specialised group of benthic foraminifera.

Late Cretaceous planktonic foraminiferal assemblages from the Exmouth Plateau, ODP Sites 122-762 and 122-763, 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 subintervals. 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.