Geochemical analysis of nine sections from the Nama Group, Namibia

The first appearance of skeletal metazoans in the late Ediacaran (~550 million years ago; Ma) has been linked to the widespread development of oxygenated oceanic conditions, but a precise spatial and temporal reconstruction of their evolution has not been resolved. Here we consider the evolution of ocean chemistry from ~550 to ~541 Ma across shelf-to-basin transects in the Zaris and Witputs Sub-Basins of the Nama Group, Namibia. New carbon isotope data capture the final stages of the Shuram/Wonoka deep negative C-isotope excursion, and these are complemented with a reconstruction of water column redox dynamics utilising Fe-S-C systematics and the distribution of skeletal and soft-bodied metazoans. Combined, these inter-basinal datasets provide insight into the potential role of ocean redox chemistry during this pivotal interval of major biological innovation. The strongly negative d13C values in the lower parts of the sections reflect both a secular, global change in the C-isotopic composition of Ediacaran seawater, as well as the influence of 'local' basinal effects as shown by the most negative d13C values occurring in the transition from distal to proximal ramp settings. Critical, though, is that the transition to positive d13C values postdates the appearance of calcified metazoans, indicating that the onset of biomineralization did not occur under post-excursion conditions. Significantly, we find that anoxic and ferruginous deeper water column conditions were prevalent during and after the transition to positive d13C that marks the end of the Shuram/Wonoka excursion. Thus, if the C isotope trend reflects the transition to global-scale oxygenation in the aftermath of the oxidation of a large-scale, isotopically light organic carbon pool, it was not sufficient to fully oxygenate the deep ocean. Both sub-basins reveal highly dynamic redox structures, where shallow, inner ramp settings experienced transient oxygenation. Anoxic conditions were caused either by episodic upwelling of deeper anoxic waters or higher rates of productivity. These settings supported short-lived and monospecific skeletal metazoan communities. By contrast, microbial (thrombolite) reefs, found in deeper inner- and mid-ramp settings, supported more biodiverse communities with complex ecologies and large skeletal metazoans. These long-lived reef communities, as well as Ediacaran soft-bodied biotas, are found particularly within transgressive systems, where oxygenation was persistent. We suggest that a mid-ramp position enabled physical ventilation mechanisms for shallow water column oxygenation to operate during flooding and transgressive sea-level rise. Our data support a prominent role for oxygen, and for stable oxygenated conditions in particular, in controlling both the distribution and ecology of Ediacaran skeletal metazoan communities.

Stable isotopc record of ODP Site 114-704

We report a near-continuous, stable isotopic record for the Pliocene-Pleistocene (4.8 to 0.8 Ma) from Ocean Drilling Program Site 704 in the sub-Antarctic South Atlantic (47°S, 7°E). During the early to middle Pliocene (4.8 to 3.2 Ma), variation in delta18O was less than ~0.5 per mil, and absolute values were generally less than those of the Holocene. These results indicate some warming and minor deglaciation of Antarctica during intervals of the Pliocene but are inconsistent with scenarios calling for major warming and deglaciation of the Antarctic ice sheet. The climate System operated within relatively narrow limits prior to ~3.2 Ma, and the Antarctic cryosphere probably did not fluctuate on a large scale until the late Pliocene. Benthic oxygen isotopic values exceeded 3 per mil for the first time at 3.16 Ma. The amplitude and mean of the delta18O signal increased at 2.7 Ma, suggesting a shift in climate mode during the latest Gauss. The greatest delta18O values of the Gaus anti Gilbert chrons occurred at ~2.6 Ma, just below a hiatus that removed the interval from ~2.6 to 2.3 Ma in Site 704. These results agree with those from Subantarctic Site 514, which suggest that the latest Gauss (2.68 to 2.47 Ma) was the time of greatest change in Neogene climate in the northern Antarctic and Subanthtic regions. During this period, surface water cooled as the Polar Front Zone (PFZ) migrated north and perennial sea ice Cover expanded into the Subantarctic region. Antarctic ice volume increased and the ventilation rate of Southern Ocean deep water decreased during glacial events after 2.7 Ma. We suggest that these changes in the Southern Ocean were related to a gradual lowering of sea level and a reduction in the flux of North Atlantic Deep Water (NADW) with the Initiation of ice growth in the northern hemisphere. The early Matuyama Chron (~ 2.3 to 1.7 Ma) was marked by relatively warm climates in the Southern Ocean except for strong glacial events associated with isotopic stages 82 (2.027 Ma), 78 (1.941 Ma), and 70 (1.782 Ma). At 1.67 Ma (stage 65/64 transition), surface waters cooled as the PFZ migrated equatorward and oscillated about a far northerly position for a prolonged interval between 1.67 and 1.5 Ma (stages 65 to 57). Beginning at ~1.42 Ma (stage 52), all parameters (delta18O, delta13C, %opal, %CaCO3) in Hole 704 become highly correlated with each other and display a very strong 41-kyr cyclicity. This increase in the importance of the 41-kyr cycle is attributed to an increase in the amplitude of the Earth's obliquity cycle that was likely reinforced by increased glacial suppression of NADW, which may explain the tightly coupled response that developed between the Southern Ocean and the North Atlantic beginning at ~1.42 Ma (stage 52).

Climate sensitivity across marine domains of life: limits to evolutionary adaptation shape species interactions, supplementary material

Organisms in all domains, Archaea, Bacteria, and Eukarya will respond to climate change with differential vulnerabilities resulting in shifts in species distribution, coexistence, and interactions. The identification of unifying principles of organism functioning across all domains would facilitate a cause and effect understanding of such changes and their implications for ecosystem shifts. For example, the functional specialization of all organisms in limited temperature ranges leads us to ask for unifying functional reasons. Organisms also specialize in either anoxic or various oxygen ranges, with animals and plants depending on high oxygen levels. Here, we identify thermal ranges, heat limits of growth, and critically low (hypoxic) oxygen concentrations as proxies of tolerance in a meta-analysis of data available for marine organisms, with special reference to domain-specific limits. For an explanation of the patterns and differences observed, we define and quantify a proxy for organismic complexity across species from all domains. Rising complexity causes heat (and hypoxia) tolerances to decrease from Archaea to Bacteria to uni- and then multicellular Eukarya. Within and across domains, taxon-specific tolerance limits likely reflect ultimate evolutionary limits of its species to acclimatization and adaptation. We hypothesize that rising taxon-specific complexities in structure and function constrain organisms to narrower environmental ranges. Low complexity as in Archaea and some Bacteria provide life options in extreme environments. In the warmest oceans, temperature maxima reach and will surpass the permanent limits to the existence of multicellular animals, plants and unicellular phytoplankter. Smaller, less complex unicellular Eukarya, Bacteria, and Archaea will thus benefit and predominate even more in a future, warmer, and hypoxic ocean.

Experimental data on photophysiology and growth rates of Pararotalia calcariformata

The eastern Mediterranean is a hotspot of biological invasions. Numerous species of Indo-pacific origin have colonized the Mediterranean in recent times, including tropical symbiont-bearing foraminifera. Among these is the species Pararotalia calcariformata. Unlike other invasive foraminifera, this species has been discovered only two decades ago and is restricted to the eastern Mediterranean coast. Combining ecological, genetic and physiological observations, we attempt to explain the recent invasion of this species in the Mediterranean Sea. Using morphological and genetic data, we confirm the species attribution to P. calcariformata McCulloch 1977 and identify its symbionts as a consortium of diatom species dominated by Minutocellus polymorphus. We document photosynthetic activity of its endosymbionts using Pulse Amplitude Modulated Fluorometry and test the effects of elevated temperatures on growth rates of asexual offspring. The culturing of asexual offspring for 120 days shows a 30-day period of rapid growth followed by a period of slower growth. A subsequent 48-day temperature sensitivity experiment indicates a similar developmental pathway and high growth rate at 28°C, whereas an almost complete inhibition of growth was observed at 20°C and 35°C. This indicates that the offspring of this species may have lower tolerance to cold temperatures than what would be expected for species native to the Mediterranean. We expand this hypothesis by applying a Species Distribution Model (SDM) based on modern occurrences in the Mediterranean using three environmental variables: irradiance, turbidity and yearly minimum temperature. The model reproduces the observed restricted distribution and indicates that the range of the species will drastically expand westwards under future global change scenarios. We conclude that P. calcariformata established a population in the Levant because of the recent warming in the region. In line with observations from other groups of organisms, our results indicate that continued warming of the eastern Mediterranean will facilitate the invasion of more tropical marine taxa into the Mediterranean, disturbing local biodiversity and ecosystem structure.

Diatom abundances in sediment core CESAR_83-006

The modern Arctic Ocean is regarded as a barometer of global change and amplifier of global warming (Graversen et al., 2008, doi:10.1038/nature06502) and therefore records of past Arctic change are critical for palaeoclimate reconstruction. Little is known of the state of the Arctic Ocean in the greenhouse period of the Late Cretaceous epoch (65-99 million years ago), yet records from such times may yield important clues to Arctic Ocean behaviour in near-future warmer climates. Here we present a seasonally resolved Cretaceous sedimentary record from the Alpha ridge of the Arctic Ocean. This palaeo-sediment trap provides new insight into the workings of the Cretaceous marine biological carbon pump. Seasonal primary production was dominated by diatom algae but was not related to upwelling as was previously hypothesized (Kitchell and Clark, 1982, doi:10.1016/0031-0182(82)90087-6). Rather, production occurred within a stratified water column, involving specially adapted species in blooms resembling those of the modern North Pacific subtropical gyre (Dore et al., 2008, doi:10.1016/j.pocean.2007.10.002), or those indicated for the Mediterranean sapropels (Kemp et al., 1999, doi:10.1038/18001). With increased CO2 levels and warming currently driving increased stratification in the global ocean (Sarmiento et al., 1998, doi:10.1038/30455), this style of production that is adapted to stratification may become more widespread. Our evidence for seasonal diatom production and flux testify to an ice-free summer, but thin accumulations of terrigenous sediment within the diatom ooze are consistent with the presence of intermittent sea ice in the winter, supporting a wide body of evidence for low temperatures in the Late Cretaceous Arctic Ocean (Falcon-Lang et al., 2004, doi:10.1016/j.palaeo.2004.05.016; Amiot et al., 2004, doi:10.1016/j.epsl.2004.07.015; Otto-Bliesner et al., 2002, doi:10.1029/2001JD000821), rather than recent suggestions of a 15 °C mean annual temperature at this time (Jenkyns et al., 2004, doi:10.1038/nature03143).