Patterns in fish radiation are compatible with Pleistocene desiccation of Lake Victoria and 14 600 year history for its cichlid species flock

Geophysical data are currently being interpreted as evidence for a late Pleistocene desiccation of Lake Victoria and its refilling 14 600 years ago. This implies that between 500 and 1000 endemic cichlid fish species must have evolved in 14 600 years, the fastest large-scale species radiation known. A recent review concludes that biological evidence clearly rejects the postulated Pleistocene desiccation of the lake: a 14 600 year history would imply exceptionally high speciation rates across a range of unrelated fish taxa. To test this suggestion, I calculated speciation rates for all 41 phylogenetic lineages of fish in the lake. Except for one cichlid lineage, accepting a 14 600 year history does not require any speciation rates that fall outside the range observed in fishes in other young lakes around the world. The exceptional taxon is a lineage of haplochromine cichlids that is also known for its rapid speciation elsewhere. Moreover, since it is unknown how many founding species it has, it is not certain that its speciation rates are really outside the range observed in fishes in other young lakes. Fish speciation rates are generally faster in younger than in older lakes, and those in Lake Victoria, by far the largest of the young lakes of the world, are no exception. From the speciation rates and from biogeographical observations that Lake Victoria endemics, which lack close relatives within the lake basin, have such relatives in adjacent drainage systems that may have had Holocene connections to Lake Victoria, I conclude that the composition of the fish assemblage does not provide biological evidence against Pleistocene desiccation. It supports a hypothesis of recent colonization from outside the lake basin rather than survival of a diverse assemblage within the basin.

Plio-Pleistocene diatom biostratigraphy from ODP Leg 177, Atlantic sector of the Southern Ocean

Seven sites were drilled during Ocean Drilling Program Leg 177 in the Atlantic sector of the Southern Ocean (SO) on a transect over the Antarctic Circumpolar Current from the Subantarctic to the Antarctic Zone. At four sites sediments were recovered with a Pliocene/Pleistocene sediment package of up to 580 m allowing the refinement of previous diatom zonation concepts. Samples were analyzed on stratigraphic distribution and abundance of diatom species. A refined diatom biozonation tied to the geomagnetic polarity record is proposed. For the middle and late Pleistocene two zonations applicable to the northern and southern area of the SO were constructed, considering different latitudinal distributions of biostratigraphic diatom marker species. The southern zonation for the Pleistocene relies on the occurrence of species of the genus Rouxia, R. leventerae and R. constricta n. sp. as well as on a revised last occurrence datum (LOD) of Actinocyclus ingens (0.38 Ma, late marine isotope stage (MIS) 11). The use of these new stratigraphic marker species refines the temporal resolution for biostratigraphic age assignment to up to 0.1 Myr. In particular the LOD of R. leventerae as an indicator for the MIS 6/5 boundary (Termination II) will improve future dating of carbonate-free Antarctic sediments. These new data were obtained from sediments of Sites 1093 and 1094 (Antarctic Zone). The northern zonation for the middle and late Pleistocene time interval is based on the Pleistocene abundance pattern of Hemidiscus karstenii which was already proposed by previous investigations (e.g. Gersonde and Barcena, 1998). One new species (R. constricta) and two new combinations (Fragilariopsis clementia, Fragilariopsis reinholdii) are proposed in this study.

Relative abundance and ranges of selected diatoms from Pliocene-Pleistocene sections of ODP Leg 177, Atlantic sector of the Southern Ocean

During Ocean Drilling Program (ODP) Leg 177, seven sites were drilled aligned on a transect across the Antarctic Circumpolar Current in the Atlantic sector of the Southern Ocean. The primary scientific objective of Leg 177 was the study of the Cenozoic paleoceanographic and paleoclimatic history of the southern high latitudes and its relationship with the Antarctic cryosphere development. Of special emphasis was the recovery of Pliocene-Pleistocene sections, allowing paleoceanographic studies at millennial or higher time resolution, and the establishment of refined biostratigraphic zonations tied to the geomagnetic polarity record and stable isotope records. At most sites, multiple holes were drilled to ensure complete recovery of the section. A description of the recovered sections and the construction of a multihole splice for the establishment of a continuous composite is presented in the Leg 177 Initial Reports volume for each of the sites (Gersonde, Hodell, Blum, et al., 1999). Here we present the relative abundance pattern and the stratigraphic ranges of diatom taxa encountered from shore-based light microscope studies completed on the Pliocene-Pleistocene sequences from six of the drilled sites (Sites 1089-1094). No shore-based diatom studies have been conducted on the Pliocene-Pleistocene sediments obtained at Site 1088, located on the northern crest of the Agulhas Ridge, because of the scattered occurrence and poor preservation of diatoms in these sections (Shipboard Scientific Party, 1999b). The data included in our report present the baseline of a diatom biostratigraphic study of Zielinski and Gersonde (2002), which (1) includes a refinement of the southern high-latitude Pliocene-Pleistocene diatom zonation, in particular for the middle and late Pleistocene, and (2) presents a biostratigraphic framework for the establishment of age models of the recovered sediment sections. Zielinski and Gersonde (2002) correlated the diatom ranges with the geomagnetic polarity record established shipboard (Sites 1090 and 1092) (Shipboard Scientific Party, 1999c, 1999d) and on shore (Sites 1089, 1091, 1093, and 1094) by Channell and Stoner (2002). The Pliocene-Pleistocene diatom zonation proposed by Zielinski and Gersonde (2002) relies on a diatom zonation from Gersonde and Bárcena (1998) for the northern belt of the Southern Ocean. Because of latitudinal differentiation of sea-surface temperature, nutrients, and salinity between Antarctic and Subantarctic/subtropical water masses, the Pliocene-Pleistocene stratigraphic marker diatoms are not uniformly distributed in the Southern Ocean (Fenner, 1991; Gersonde and Bárcena, 1998). As a consequence, Zielinski and Gersonde (2002) propose two diatom zonations for application in the Antarctic Zone south of the Polar Front (Southern Zonation, Sites 1094 and 1093) and the area encompassing the Polar Front Zone (PFZ) and the Subantarctic Zone (Northern Zonation, Sites 1089-1092). This accounts especially for the Pleistocene zonation where Hemidiscus karstenii, whose first abundant occurrence datum and last occurrence datum defines the subzonation of the northern Thalassiosira lentiginosa Zone, occurs only sporadically in the cold-water realm south of the PFZ and thus is not applicable in sections from this area. However, newly established marker species assigned to the genus Rouxia (Rouxia leventerae and Rouxia constricta) are more related to cold-water environments and allow a refinement of the Pleistocene stratigraphic zonation for the southern cold areas. A study relying on quantitative counts of both Rouxia species confirms the utility of these stratigraphic markers for the identification of sequences attributed to marine isotope Stages 6 and 8 in the southern Southern Ocean (Zielinski et al., 2002).

Plio-Pleistocene age-depth points and occurrence of hiatuses in ODP sites of the Southern Ocean

Changes in Atlantic deep water circulation were reconstructed by comparing the benthic foraminiferal delta13C record at ODP Site 1090 in the South Atlantic with similar records from the North Atlantic (Sites 982, 607, 925, 929) and deep Pacific (Site 849) oceans. Important deep water circulation changes occurred in the early Pleistocene at 1.55 Myr and during the Mid-Pleistocene Transition at 0.9 Myr. At 1.55 Myr, glacial delta13C values in the Southern Ocean became significantly lower than those in the deep Pacific, establishing a pattern that persisted throughout the late Pleistocene. We propose that the lowering of delta13C values of Southern Component Water (SCW) at this time resulted from expansion of sea ice and reduced ventilation of deep water during glacial periods after marine isotope stage 52. Accompanying this change in Southern Ocean deep water circulation was enhanced interhemispheric coupling between the North and South Atlantic after 1.55 Myr. At ~0.9 Myr, the magnitude of glacial-to-interglacial variabilityin delta13C increased and shifted to a longer frequency (100 kyr) along with oceanic delta18O (ice volume). Calculation of percent Northern Component Water (NCW) using Site 1090 as the SCW end member yielded 20-30% less reduction of NCW during glacial periods of the late Pleistocene. Also, a trend toward reduced glacial suppression of NCW during the past 400 kyr is not evident. The apparent decoupling of ice volume and deep water circulation reported previously maybe an artifact of using a Pacific, rather than a Southern Ocean, carbon isotopic record to calculate past mixing ratios of NCW and SCW.

Late Neogene carbonate sedimentation and stable isotope record of ODP Site 121-758

The evolution of oceanic and climatic conditions the northeast Indian Ocean during the last 7 m.y. is revealed in the sediments from Site 758. We present detailed and continuous records of d18O and d13C from planktonic foraminifers, weight percent calcium carbonate, weight percent coarse fraction, magnetic susceptibility, and geomagnetic reversals. Sample spacing of the records ranges from 3 to 10 cm and is equivalent to an average time interval of 2000 to 6000 yr. Despite the fact that core recovery ranged between 100% and 105%, recovery gaps as large as 2.7 m occurred at nearly every break between advanced hydraulic piston cores. Approximately 12% of the late Neogene sequence was not recovered in each of the two holes drilled at Site 758. To circumvent the discontinuity introduced by the gaps, a composite depth section was constructed from multiple cores taken from offset holes at Site 758. The resulting composite depth section extends continuously from 0 to 116 mbsf, from the Holocene to the upper Miocene. A detailed chronostratigraphy is based on geomagnetic reversals which extend from the Brunhes Chron to Chron 6, and on d18O stages 1 through 105, which span from 0 to 2.5 Ma. The d18O record is dominated by a ~40-k.y. cycle in the late Pliocene and early Pleistocene, and is followed by a change to a ~100-k.y. cycle in the late Pleistocene. The mid-Pleistocene transition between these two modes of variability occurs between d18O stages 25 and 22 (between 860 and 800 Ka). Thirteen major volcanic ash horizons from the Indonesian arc are observed throughout the sedimentary section and are dated by their relative position within the geomagnetic reversals and the d18O chronostratigraphy. Since 5 Ma, there has been a long-term decline in weight percent CaCO3 and CaCO3 mass accumulation rates, and an associated rise in non-CaCO3 mass accumulation rates. We attribute these changes to a decrease in CaCO3 productivity and an increase in terrigenous sedimentation through enhanced riverine input. Such input may be linked to rapid tectonic uplift of the Himalayas and the Tibetan Plateau via mechanisms such as the intensification of the monsoonal rains, increased fluvial erosion, and regional glaciation. The long-term increase in percent coarse fraction since 5 Ma suggests a gradual increase in CaCO3 preservation. Higher frequency fluctuations in CaCO3 preservation are superimposed on the long-term trend and are related to climate fluctuations. The abrupt drop (-50%) in CaCO3 accumulation at 3.4 Ma signals a dramatic decrease in CaCO3 production that occurred over much of the Indian Ocean.

Mid-Pleistocene planktonic foraminiferal composition and IRD content in ODP Sites 162-980 and 162-984

We investigated surface and deep ocean variability in the subpolar North Atlantic from 1000 to 500 thousand years ago (ka) based on two Ocean Drilling Program (ODP) sites, Feni drift site 980 (55°29'N, 14°42'W) and Bjorn drift site 984 (61°25'N, 24°04'W). Benthic foraminiferal stable isotope data, planktic foraminiferal faunas, ice-rafted debris data, and faunally based sea-surface temperature estimates help test the hypothesis that oceanographic changes in the North Atlantic region were associated with the onset of the 100-kyr world during the mid-Pleistocene revolution. Based on percentage of Neogloboquadrina pachyderma (s) records from both sites, surface waters during interglacials and glacials were cooler in the mid-Pleistocene than during marine isotope stages (MIS) 5 and 6. In particular, interglaciations at Bjorn drift site 984 were significantly cooler. Faunal evidence suggests that the interglacial Arctic front shifted from a position between the two sites to a position northwest of Bjorn drift site 984 after ca. 610 ka. As during the late Pleistocene, we find faunal evidence for lagging surface warmth at most of the glacial initiations during the mid-Pleistocene. Each initiation is associated with high benthic d13C values that are maintained into the succeeding glaciation, which we term "lagging NADW production." These findings indicate that lagging warmth and lagging NADW production are robust features of the regional climate system that persist in the middle to late Pleistocene.

Ice-rafted debris of Late Cenozoic sediments from ODP Leg 104 holes

The abundance and composition of the upper Cenozoic terrigenous coarse-sand fraction (250 µm-2 mm) at ODP Sites 642, 643, and 644 were investigated to date the onset of significant ice-rafting in the Norwegian Sea, establish the regional chronology of ice-rafting, and determine the relative importance of global vs. regional controls on ice-rafting in this area. The first input of ice-rafted debris (IRD) occurs at approximately 2.9 Ma, with significant ice-rafting beginning at about 2.5 Ma. IRD abundances increase significantly in sediments younger than 0.9 Ma at all three holes, indicating climatic deterioration in the late Pleistocene. Differences in the timing of this IRD increase between holes result from regional patterns of IRD supply and surface circulation. Variations in IRD sources and dispersal patterns may also explain the slightly higher background level of IRD abundance at Hole 642B, a seaward site. Major peaks in the generalized IRD records from the Norwegian Sea are tentatively correlated to glacial stages or glacial-to-interglacial transitions in the globally defined oxygen isotope record. This correlation indicates the effect of global conditions on the regional climate of the Norwegian Sea, although the detailed IRD records at these sites are also affected by local/regional processes (e.g., circulation patterns and source area differences).

Pleistocene sedimentation in the Benguela Current, southeastern Atlantic

We infer variations in paleoproductivity and eolian input at ODP Site 1082 in the Walvis Basin from stable oxygen isotope compositions of the planktonic foraminifera Globorotalia inflata, total organic carbon mass accumulation rates (TOC MAR), and X-ray fluorescence analyses of Fe content. The most pronounced paleoclimatic changes correspond to the time at about 0.9 Ma, when glacial conditions in the northern hemisphere (NH) led to the onset of pronounced 100-kyr glacial-interglacial cycles. We used Fe intensity as a proxy for eolian terrigenous input, and TOC MAR as a paleoproductivity indicator. Paleoproductivity and eolian input show generally higher-amplitude variations of glacial-interglacial cyclicity from 1.5 to 0.58 Ma, indicating pronounced variations in upwellingfavorable winds in this area. At 0.58 Ma, paleoproductivity and eolian input shifted abruptly to lower-amplitude variations with a periodicity of 100 kyr while delta18O values show a trend toward more negative isotope values for the past 0.65 Myr. Especially during glacial periods, oxygen isotope values indicate increasingly warmer sea-surface temperatures toward the end of the Pleistocene. To evaluate the relative influences of NH glaciation and southern hemisphere (SH) insolation as potential forcing mechanisms for variations of eolian input and productivity in the northern Benguela system, we filtered our proxy records at orbital frequencies. The filtered records of Fe intensity and TOC MAR indicate a strong influence of the 100-kyr and 41-kyr frequency bands, supporting our assumption that strong ice buildup in the NH is the dominant trigger for climate changes on the continent and probably in trade-wind intensity. SH insolation and low-latitude precession-related insolation changes were important for paleoproductivity variations in the northern Benguela system, modifying the nutrient supply by southern ocean intermediate waters and the zonal direction of upwelling-inducing trades by the African monsoon system, respectively.

Late Miocene to Holocene calcareous nannofossils from ODP sites of Leg 125

During Leg 125 of the Ocean Drilling Program, nine sites were drilled in the Mariana and Izu-Bonin areas. The sediments recovered range in age from early Pliocene to late Pleistocene in the Mariana Region and from middle Eocene to late Pleistocene in the Izu-Bonin region. This contribution concerns the biostratigraphic study of the latest Miocene (CN9b Subzone) to late Pleistocene interval. Aquantitative analysis of all calcareous nannofossil associations was conducted for the interval encompassing late Miocene to the top of the early Pliocene. Moreover, the genera Discoaster, Amaurolithus, and Ceratolithus were quantitatively investigated from the late Miocene to late Pliocene interval. Some bioevents were identified, and variations in the composition of assemblages were linked to climatic changes.

The occurrence and significance of Pleistocene and Upper Pliocene sapropels

Numerous sapropels and sapropelic strata from Upper Pliocene and Pleistocene hemipelagic sediments of the Tyrrhenian Sea show that intermittent anoxia, possibly related to strongly increased biological productivity, was not restricted to the eastern Mediterranean basins and may be a basin-wide result of Late Pliocene-Pleistocene climatic variability. Even though the sapropel assemblage of the Tyrrhenian Sea clearly originates from multiple processes such as deposition under anoxic conditions or during spikes in surface water productivity and lateral transport of organic-rich suspensates, many "pelagic sapropels" have been recognized. Stratigraphic ages calculated for the organic-rich strata recovered during ODP Leg 107 indicate that the frequency of sapropel formation increased from the lowermost Pleistocene to the base of the Jaramillo magnetic event, coinciding with a period when stable isotope records of planktonic foraminifera indicate the onset of climatic cooling in the Mediterranean. A second, very pronounced peak in sapropel formation occurred in the Middle to Late Pleistocene (0.73-0.26 Ma). Formainifers studied in three high-resolution sample sets suggest that changes in surface-water temperature may have been responsible for establishing anoxic conditions, while salinity differences were not noted in the faunal assemblage. However, comparison of sapropel occurrence at Site 653 with the oxygen isotopic record of planktonic foraminifers established by Thunell et al. (1990, doi:10.2973/odp.proc.sr.107.155.1990) indicates that sapropel occurrences coincide with negative d18O excursions in planktonic foraminifers in thirteen of eighteen sapropels recognized in Hole 653A. A variant of the meltwater hypothesis accepted for sapropel formation in the Late Pleistocene eastern Mediterranean may thus be the cause of several "anoxic events" in the Tyrrhenian as well. Model calculations indicate that the amount of oxygen advection from Western Mediterranean Deep Water exerts the dominant control on the oxygen content in deep water of the Tyrrhenian Sea. Inhibition of deep-water formation in the northern Adriatic and the Balearic Basin by increased meltwater discharge and changing storm patterns during climatic amelioration may thus be responsible for sapropel formation in the Tyrrhenian Sea.

Carbonate composition of Pliocene-Pleistocene sediments of the Maledives

Ocean Drilling Program (ODP) Leg 115 post-cruise research was focused on two Maldives sites, more precisely on the top 108 m of Hole 716B (water depth, 540 m), equivalent to the past 3.5 m.y., and the top 19.5 m of Hole 714A (water depth, 2195 m), equivalent to the past 0.55 m.y. These sediments consist of mostly unaltered and undisturbed, turbidite-free, periplatform ooze. Results of our research are compared with existing data on Hole 633A (water depth, 1681 m), drilled in the Bahamas during ODP Leg 101, using age/depth models built on the basis of oxygen isotope, nannofossil, and magnetic stratigraphies. Climate-induced, long-term (roughly 0.5 m.y.) aragonite cycles, superposed on short-term (roughly 0.04 and 0.1 m.y.) aragonite cycles, have been established at least during the past 2.0 m.y., in the Maldives and the Bahamas. Our most interesting result is the clear correlation among the aragonite long-term cycles in the Maldives and the Bahamas and the carbonate-preservation, long-term cycles from the open Pacific, Indian, and North Atlantic oceans. The mid-Brunhes dissolution interval, corresponding to the youngest preservation minima of the carbonate-preservation, longterm cycles, is clearly defined by fine aragonite minimum values in the deep periplatform sites, and by maximum fragmentation of pteropod tests in the shallow sites. Aragonite and planktonic d18O records, usually in phase during the late Pleistocene, display, further back in time, discreet intervals where the two records do not match with one another. Major mismatches between both records occur synchronously in the Maldives and Bahamas periplatform sites and seem to correspond to extreme events of either carbonate-preservation or dissolution in the deep pelagic carbonate sites of the equatorial Pacific Ocean. Based on our findings, short- and long-term aragonite cycles can no longer be explained only by variations of aragonite input from the nearby shallow carbonate banks, in response to their alternate flooding and exposure through cyclic sea-level fluctuations. The aragonite long-term cycles in the periplatform environments are interpreted as carbonatepreservation cycles at intermediate-water depths. Their occurrence shows, therefore, that the carbonate chemistry of the entire water column has been influenced by long-term (0.5 m.y.) cyclic variations during the past 2.0 m.y. These major changes of the water-column carbonate chemistry are linked to the climate-induced carbon cycling among the different atmospheric, oceanic, and sedimentary carbon reservoirs.

Distribution of late Quaternary calcareous nannofossils in DSDP Leg 96 sites

Sediments from holes drilled at 11 sites in the northern Gulf of Mexico during Deep Sea Drilling Project Leg 96 were analyzed for calcareous nannofossil content. All sediments recovered are Holocene and late Pleistocene in age and are within the Emiliania huxleyi Zone. The datum level represented by the lowest stratigraphic occurrence of dominant E. huxleyi occurs at two sites (Sites 615 and 619) and can be dated at approximately 84,000 yr. ago at Site 619. Reworked Cretaceous nannofossils are generally common or abundant and dominate the floral assemblages of the late Wisconsin glacial sediments. When present, indigenous late Quaternary species are rare or few in abundance. Slight increases in the contemporaneous Quaternary component of the floral assemblages can be documented by the use of a calculated in s/fu/reworked ratio. This ratio, based on the relative abundances of the indigenous Quaternary taxa and reworked taxa, shows potential both for local correlations between drill sites and for correlation with glacio-eustatic fluctuations during the late Pleistocene.

Late Pliocene age models of Agulhas Plateau sediment cores

An ensemble of new, high-resolution records of surface ocean hydrography from the Indian-Atlantic oceanic gateway, south of Africa, demonstrates recurrent and high-amplitude salinity oscillations in the Agulhas Leakage area during the penultimate glacial-interglacial cycle. A series of millennial-scale salinification events, indicating strengthened salt leakage into the South Atlantic, appear to correlate with abrupt changes in the North Atlantic climate and Atlantic Meridional Overturning Circulation (AMOC). This interhemispheric coupling, which plausibly involved changes in the Hadley Cell and midlatitude westerlies that impacted the interocean transport at the tip of Africa, suggests that the Agulhas Leakage acted as a source of negative buoyancy for the perturbed AMOC, possibly aiding its return to full strength. Our finding points to the Indian-to-Atlantic salt transport as a potentially important modulator of the AMOC during the abrupt climate changes of the Late Pleistocene.

A new extinct primate among the Pleistocene megafauna of Bahia, Brazil.

A nearly complete skeleton of a robust-bodied New World monkey that resembles living spider monkeys was recovered from undisturbed Pleistocene deposits in the Brazilian state of Bahia. The skeleton displays the highly specialized postcranial pattern typical of spider and woolly spider monkeys and shares cranial similarities to the spider monkey exclusively. It is generically distinct on the basis of its robustness (>20 kg) and on the shape of its braincase. This new genus indicates that New World monkeys nearly twice the size of those living today were part of the mammalian biomass of southern Amazonia in the late Pleistocene. The discovery of this specimen expands the known adaptive diversity of New World monkeys and demonstrates that they underwent body size expansion in the terminal Pleistocene, as did many other types of mammals.