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.

Lake Baikal amphipods under climate change: Thermal constraints and ecological consequences, links to supplementary material

Lake Baikal, the world's most voluminous freshwater lake, has experienced unprecedented warming during the last decades. A uniquely diverse amphipod fauna inhabits the littoral zone and can serve as a model system to identify the role of thermal tolerance under climate change. This study aimed to identify sublethal thermal constraints in two of the most abundant endemic Baikal amphipods, Eulimnogammarus verrucosus and Eulimnogammarus cyaneus, and Gammarus lacustris, a ubiquitous gammarid of the Holarctic. As the latter is only found in some shallow isolated bays of the lake, we further addressed the question whether rising temperatures could promote the widespread invasion of this non-endemic species into the littoral zone. Animals were exposed to gradual temperature increases (4 week, 0.8 °C/d; 24 h, 1 °C/h) starting from the reported annual mean temperature of the Baikal littoral (6 °C). Within the framework of oxygen- and capacity-limited thermal tolerance (OCLTT), we used a nonlinear regression approach to determine the points at which the changing temperature-dependence of relevant physiological processes indicates the onset of limitation. Limitations in ventilation representing the first limits of thermal tolerance (pejus (= "getting worse") temperatures (Tp)) were recorded at 10.6 (95% confidence interval; 9.5, 11.7), 19.1 (17.9, 20.2), and 21.1 (19.8, 22.4) °C in E. verrucosus, E. cyaneus, and G. lacustris, respectively. Field observations revealed that E. verrucosus retreated from the upper littoral to deeper and cooler waters once its Tp was surpassed, identifying Tp as the ecological thermal boundary. Constraints in oxygen consumption at higher than critical temperatures (Tc) led to an exponential increase in mortality in all species. Exposure to short-term warming resulted in higher threshold values, consistent with a time dependence of thermal tolerance. In conclusion, species-specific limits to oxygen supply capacity are likely key in the onset of constraining (beyond pejus) and then life-threatening (beyond critical) conditions. Ecological consequences of these limits are mediated through behavioral plasticity in E. verrucosus. However, similar upper thermal limits in E. cyaneus (endemic, Baikal) and G. lacustris (ubiquitous, Holarctic) indicate that the potential invader G. lacustris would not necessarily benefit from rising temperatures. Secondary effects of increasing temperatures remain to be investigated.

Alkenones and SST of surface sediments (Appendix)

We have analysed alkenones in 149 surface sediments from the eastern South Atlantic in order to establish a sediment-based calibration of the U37K' paleotemperature index. Our study covers the major tropical to subpolar production systems and sea-surface temperatures (SST's) between 0° and 27°C. In order to define the most suitable calibration for this region, the U37K' values were correlated to seasonal, annual, and production-weighted annual mean atlas temperatures and compared to previously published culture and core-top calibrations. The best linear correlation between U37K' and SST was obtained using annual mean SST from 0 to 10 m water depth (U37K' = 0.033 T + 0.069, r**2 = 0.981). Data scattering increased significantly using temperatures of waters deeper than 20 m, suggesting that U37K' reflects mixed-layer SST and that alkenone production at thermocline depths was not high enough to significantly bias the mixed-layer signal. Regressions based on both production-weighted and on actual annual mean atlas SST were virtually identical, indicating that regional variations in the seasonality of primary production have no discernible effect on the U37K' vs. SST relationship. Comparison with published core-top calibrations from other oceanic regions revealed a high degree of accordance. We, therefore, established a global core-top calibration using U37K' data from 370 sites between 60°S and 60°N in the Atlantic, Indian, and Pacific Oceans and annual mean atlas SST (0-29°C) from 0 m water depth. The resulting relationship (U37K' = 0.033 T + 0.044, r**2 = 958) is identical within error limits to the widely used E. huxleyi calibrations of and attesting their general applicability. The observation that core-top calibrations extending over various biogeographical coccolithophorid zones are strongly linear and in better accordance than culture calibrations suggests that U37K' is less species-dependent than is indicated by culture experiments. The results also suggest that variations in growth rate of algae and nutrient availability do not significantly affect the sedimentary record of U37K' in open ocean environments.

Alkenone data of sediment traps off Cape Blanc, NW Africa

We analysed long-chain alkenones in sinking particles and surface sediments from the filamentous upwelling region off Cape Blanc, NW Africa, to evaluate the transfer of surface water signals into the geological record. Our study is based on time-series sediment trap records from 730 m (1990-1991) to 2195-3562 m depth (1988-1991). Alkenone fluxes showed considerable interannual variations and no consistent seasonality. The average flux of C37 and C38 alkenones to the deep traps was 1.9 µg/m**2/d from March 1988 to October 1990 and sevenfold higher in the subsequent year. Alkenone fluxes to the shallower traps were on average twice as high and showed similar temporal variations. The alkenone unsaturation indices UK'37, UK38Me and UK38Et closely mirrored the seasonal variations in sea-surface temperature (weekly Reynolds SST). Time lags of 10-48 days between the SST and unsaturation maxima suggest particle sinking rates of about 80 and 280 m/d for the periods of low and high alkenone fluxes, respectively. The average flux-weighted UK'37 temperature for the 4-year time series of the deeper traps was 22.1°C, in perfect agreement with the mean weekly SST for the same period. This and the comparison with seasonal temperature variations in the upper 100 m of the water column suggests that UK'37 records principally the yearly average of the mixed-layer temperature in this region. A comparison between the average annual alkenone fluxes to the lower traps (2400 µg/m**2/yr) and into the underlying sediments (4 µg/m**2/yr) suggests that only about 0.2% of the alkenones reaching the deep ocean became preserved in the sediments. The flux-weighted alkenone concentrations also decreased considerably, from 2466 µg/gC in the water column to 62 µg/gC in the surface sediments. Such a low degree of alkenone preservation is typical for slowly accumulating oxygenated sediments. Despite these dramatic diagenetic alkenone losses, the UK'37 ratio was not affected. The average UK'37 value of the sediments (0.796±0.010 or 22.3±0.3°C) was identical within error limits to the 4-year average of the lower traps. The unsaturation indices for C38 alkenones and the ratio between C37 and C38 alkenones also revealed a high degree of stability. Our results do not support the hypothesis that UK'37 is biased towards higher values during oxic diagenesis.

Accumulation rates of bulk sediment and opal in surface sediments of the South Atlantic

A set of 114 samples from the sediment surface of the Atlantic, eastern Pacific and western Indian sectors of the Southern Ocean has been analyzed for 230Th and biogenic silica. Maps of opal content, Th-normalized mass flux, and Th-normalized biogenic opal flux into the sediment have been derived. Significant differences in sedimentation patterns between the regions can be detected. The mean bulk vertical fluxes integrated into the sediment in the open Southern Ocean are found in a narrow range from 2.9 g/m**2 yr (Eastern Weddell Gyre) to 15.8 g/m**2 yr (Indian sector), setting upper and lower limits to the vertically received fraction of open ocean sediments. The silica flux to sediments of the Atlantic sector of the Southern Ocean is found to be 4.2 ± 1.4 * 10**11 mol/yr, just one half of the last estimate. This adjustment represents 6% of the output term in the global marine silica budget.