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.

Appendix A. Whole-rock major, trace and rare earth element analyses of the Ulubey volcanics, Appendix B. Analytical standards and detection limits for analyses

Collisional and post-collisional volcanic rocks in the Ulubey (Ordu) area at the western edge of the Eastern Pontide Tertiary Volcanic Province (EPTVP) in NE Turkey are divided into four suites; Middle Eocene (49.4-44.6 Ma) aged Andesite-Trachyandesite (AT), Trachyandesite-Trachydacite-Rhyolite (TTR), Trachydacite-Dacite (TD) suites, and Middle Miocene (15.1 Ma) aged Trachybasalt (TB) suite. Local stratigraphy in the Ulubey area starts with shallow marine environment sediments of the Paleocene-Eocene time and then continues extensively with sub-aerial andesitic to rhyolitic and rare basaltic volcanism during Eocene and Miocene time, respectively. Petrographically, the volcanic rocks are composed primarily of andesites/trachyandesites, with minor trachydacites/rhyolites, basalts/trachybasalts and pyroclastics, and show porphyric, hyalo-microlitic porphyric and rarely glomeroporphyric, intersertal, intergranular, fluidal and sieve textures. The Ulubey (Ordu) volcanic rocks indicate magma evolution from tholeiitic-alkaline to calc-alkaline with medium-K contents. Primitive mantle normalized trace element and chondrite normalized rare earth element (REE) patterns show that the volcanic rocks have moderate light rare earth element (LREE)/heavy rare earth element (HREE) ratios relative to E-Type MORB and depletion in Nb, Ta and Ti. High Th/Yb ratios indicate parental magma(s) derived from an enriched source formed by mixing of slab and asthenospheric melts previously modified by fluids and sediments from a subduction zone. All of the volcanic rocks share similar incompatible element ratios (e.g., La/Sm, Zr/Nb, La/Nb) and chondrite-normalized REE patterns, indicating that the basic to acidic rocks originated from the same source. The volcanic rocks were produced by the slab dehydration-induced melting of an existing metasomatized mantle source, and the fluids from the slab dehydration introduced significant large ion lithophile element (LILE) and LREE to the source, masking its inherent HFSE-enriched characteristics. The initial 87Sr/86Sr (0.7044-0.7050) and eNd (-0.3 to +3.4) ratios of the volcanics suggest that they originated from an enriched lithospheric mantle source with low Sm/Nd ratios. Integration of the geochemical, petrological and isotopical with regional and local geological data suggest that the Tertiary volcanic rocks from the Ulubey (Ordu) area were derived from an enriched mantle, which had been previously metasomatized by fluids derived from subducted slab during Eocene to Miocene in collisional and post-collisional extension-related geodynamic setting following Late Mesozoic continental collision between the Eurasian plate and the Tauride-Anatolide platform.

Reconstructed glaciation limits in Eurasia (digitized from map Fig.1)

Based on field investigations in northern Russia and interpretation of offshore seismic data, we have made a preliminary reconstruction of the maximum ice-sheet extent in the Barents and Kara Sea region during the Early/Middle Weichselian and the Late Weichselian. Our investigations indicate that the Barents and Kara ice sheets attained their maximum Weichselian positions in northern Russia prior to 50 000 yr BP, whereas the northeastern flank of the Scandinavian Ice Sheet advanced to a maximum position shortly after 17000 calendar years ago. During the Late Weichselian (25 000-10000 yr BP), much of the Russian Arctic remained ice-free. According to our reconstruction, the extent of the ice sheets in the Barents and Kara Sea region during the Late Weichselian glacial maximum was less than half that of the maximum model which, up to now, has been widely used as a boundary condition for testing and refining General Circulation Models (GCMs). Preliminary numerical-modelling experiments predict Late Weichselian ice sheets which are larger than the ice extent implied for the Kara Sea region from dated geological evidence, suggesting very low precipitation.

Antagonistic interactions between the african weaver ant Oecophylla longinoda and the parasitoid Anagyrus pseudococci potentially limits suppression of the invasive mealybug Rastrococcus iceryoides

The ant Oecophylla longinoda Latreille forms a trophobiotic relationship with the invasive mealybug Rastrococus iceryoides Green and promotes the latter's infestations to unacceptable levels in the presence of their natural enemies. In this regard, the antagonistic interactions between the ant and the parasitoid Anagyrus pseudococci Girault were assessed under laboratory conditions. The percentage of parasitism of R. iceryoides by A. pseudococci was significantly higher on "ant-excluded" treatments (86.6% ± 1.27%) compared to "ant-tended" treatments (51.4% ± 4.13%). The low female-biased sex-ratio observed in the "ant-tended" treatment can be attributed to ants' interference during the oviposition phase, which disrupted parasitoids' ability to fertilize eggs. The mean foraging time, host handling time and number of successful oviposition in "ant-excluded" treatment were significantly higher compared to "ant-tended" treatments. When ant workers were allowed access to sterilized sand grains, mummified and unmummified R. iceryoides, they selectively removed the mummified mealybugs, indicating that they recognized the mummies as potential foods (1.2 ± 0.46 to 7.8 ± 1.17 mummies at 10 min intervals for 2 h). Percentage emergence from mummified R. iceryoides removed by the ants was significantly lower compared to emergence from mummies not exposed to ants. Although, host seeking parasitoids frequently evaded attacks, some were killed by the foraging ant workers (2.0 ± 0.38 to 6.0 ± 0.88 at 10 min intervals for 2 h). These results suggest for the first time that the presence of O. longinoda has a detrimental effect on the abundance, reproductive success and possibly oviposition strategy of female parasitoids, which might be a delimiting factor in field conditions if both natural enemies are to be recommended for use within the same agro-ecosystem.

(Table 1) Gas tie points used to correlate NorthGRIP and Vostol records

North Atlantic sediment records (MD95-2042), Greenland (Greenland Ice Core Project (GRIP)) and Antarctica (Byrd and Vostok) ice core climate records have been synchronized over marine isotopic stage 3 (MIS 3) (64 to 24 kyr B.P.) (Shackleton et al., 2000). The resulting common timescale suggested that MD95-2042 d18Obenthic fluctuations were synchronous with temperature changes in Antarctica (dDice or d18Oice records). In order to assess the persistency of this result we have used here the recent Greenland NorthGRIP ice core covering the last glacial inception. We transfer the Antarctic Vostok GT4 timescale to NorthGRIP d18Oice and MD95-2042 d18Oplanktonic records and precisely quantify all the relative timing uncertainties. During the rapid warming of Dansgaard-Oeschger 24, MD95-2042 d18Obenthic decrease is in phase with d18Oplanktonic decrease and therefore with NorthGRIP temperature increase, but it takes place 1700 ± 1100 years after the Antarctic warming. Thus the present study reveals that the results obtained previously for MIS 3 cannot be generalized and demonstrates the need to improve common chronologies for marine and polar archives.

Middle Miocene to Quaternary diatom biostratigraphy of DSDP Site 90-594

A diatom biostratigraphy is presented for middle Miocene through Quaternary sediments recovered from the Chatham Rise east of New Zealand's South Island. The upper 590 m of the 639.5-m composite-section Site 594 represents approximately 16 m.y. and is characterized by moderately to very poorly preserved diatoms of antarctic to temperate affinity. Pliocene through Quaternary assemblages are poorly preserved and dominated by antarctic-subantarctic species which provide detailed biostratigraphic control. Recognized are 11 of 14 zones of the middle upper Miocene to Quaternary Neogene Southern Ocean diatom zonation (NSD 7-NSD 20) of Ciesielski (1983; this chapter). Four Neogene Southern Ocean diatom zones (NSD 3-NSD 6) are recognized in the lower middle Miocene to middle upper Miocene of Site 594. Assemblages of this interval have a mixed high-latitude and temperate affinity; however, poor preservation limits correlation to high- and temperate-latitude zonal schemes. Neogene North Pacific diatom zones and subzones of NNPD 3 through NNPD 5 (Barron, in press, b) are correlated to Neogene Southern Ocean diatom zones NSD 3 through NSD 7: the upper portions of the Actinocyclus ingens Zone (NNPD 3) is correlative to the upper Nitzschia maleinterpretaria Zone (NSD 3); the Denticulopsis lauta Zone (NNPD 4) and Subzones a and b are correlative to the lower Coscinodiscus lewisianus Zone (NSD 4); and the D. hustedtü-D. lauta Zone (NNPD 5) and its Subzones a through d encompass the upper C. lewisianus Zone (NSD 4), N. grossepunctata Zone (NSD 5), N. denticuloides Zone (NSD 6), and the lower D. hustedtii-D. lauta Zone (NSD 7). A major disconformity spans the late Gilbert to early Gauss Chron (3.9-2.8 Ma). A second disconformity brackets the Miocene/Pliocene boundary; the section missing covers late Chron 5 and the early Gilbert chron (5.5-4.6 Ma). The remainder of the siliceous-fossil-bearing Miocene sediments at Site 594 appear to be correlative to lower paleomagnetic Chronozone 5 through upper Chronozone 16. Uppermost lower Miocene or lowermost middle Miocene sediments in the basal 50 m of Hole 594A are barren of diatoms.

Planimetric ice-flow convergence and flow-orthonormal strain rate across the Antarctic Ice Sheet, with links to model result files

Fast-flowing ice streams discharge most of the ice from the interior of the Antarctic Ice Sheet coastward. Understanding how their tributary organisation is governed and evolves is essential for developing reliable models of the ice sheet's response to climate change. Despite much research on ice-stream mechanics, this problem is unsolved, because the complexity of flow within and across the tributary networks has hardly been interrogated. Here I present the first map of planimetric flow convergence across the ice sheet, calculated from satellite measurements of ice surface velocity, and use it to explore this complexity. The convergence map of Antarctica elucidates how ice-stream tributaries draw ice from the interior. It also reveals curvilinear zones of convergence along lateral shear margins of streaming, and abundant convergence ripples associated with nonlinear ice rheology and changes in bed topography and friction. Flow convergence on ice-stream tributaries and their feeding zones is markedly uneven, and interspersed with divergence at distances of the order of kilometres. For individual drainage basins as well as the ice sheet as a whole, the range of convergence and divergence decreases systematically with flow speed, implying that fast flow cannot converge or diverge as much as slow flow. I therefore deduce that flow in ice-stream networks is subject to mechanical regulation that limits flow-orthonormal strain rates. These properties and the gridded data of convergence and flow-orthonormal strain rate in this archive provide targets for ice- sheet simulations and motivate more research into the origin and dynamics of tributarization.

Atterberg limits of sediments from DSDP Leg 65 Holes

Atterberg limits have been determined for 32 unconsolidated sediment samples, ranging in composition from silty clay to sandy silt and recovered from four sites drilled at the mouth of the Gulf of California during DSDP Leg 65. The liquid limit of the samples ranged from 41.5% to 157.5%, and the plastic limit from 32.8% to 65.1%. The plasticity index ranged from 5.9% to 102.0%. In some samples, the water content was less than the liquid limit. The liquidity index averaged 76% while the flow and toughness indices averaged 35% and 2.18%, respectively. On the basis of these limits, the sediments analyzed can be classified as inorganic clays of high plasticity, organic clays of moderate to high plasticity, and diatomaceous sands, silts, and silty clays of low plasticity.

Temperature tolerance of different larval stages of the spider crab Hyas araneus exposed to elevated seawater PCO2

Exposure to elevated seawater PCO2 limits the thermal tolerance of crustaceans but the underlying mechanisms have not been comprehensively explored. Larval stages of crustaceans are even more sensitive to environmental hypercapnia and possess narrower thermal windows than adults. In a mechanistic approach, we analysed the impact of high seawater CO2 on parameters at different levels of biological organization, from the molecular to the whole animal level. At the whole animal level we measured oxygen consumption, heart rate and activity during acute warming in zoea and megalopa larvae of the spider crab Hyas araneus exposed to different levels of seawater PCO2. Furthermore, the expression of genes responsible for acid-base regulation and mitochondrial energy metabolism, and cellular responses to thermal stress (e.g. the heat shock response) was analysed before and after larvae were heat shocked byrapidly raising the seawater temperature from 10°C rearing temperature to 20°C. Zoea larvae showed a high heat tolerance, which decreased at elevated seawater PCO2, while the already low heat tolerance of megalopa larvae was not limited further by hypercapnic exposure. There was a combined effect of elevated seawater CO2 and heat shock in zoea larvae causing elevated transcript levels of heat shock proteins. In all three larval stages, hypercapnic exposure elicited an up-regulation of genes involved in oxidative phosphorylation, which was, however, not accompanied by increased energetic demands. The combined effect of seawater CO2 and heat shock on the gene expression of heat shock proteins reflects the downward shift in thermal limits seen on the whole animal level and indicates an associated capacity to elicit passive thermal tolerance. The up-regulation of genes involved in oxidative phosphorylation might compensate for enzyme activities being lowered through bicarbonate inhibition and maintain larval standard metabolic rates at high seawater CO2 levels. The present study underlines the necessity to align transcriptomic data with physiological responses when addressing mechanisms affected by an interaction of elevated seawater PCO2 and temperature extremes.

Metabolic responses to temperature stress under elevated pCO2 in Crepidula fornicata

In the current context of environmental change, ocean acidification is predicted to affect the cellular processes, physiology and behaviour of all marine organisms, impacting survival, growth and reproduction. In relation to thermal tolerance limits, the effects of elevated pCO2 could be expected to be more pronounced at the upper limits of the thermal tolerance window. Our study focused on Crepidula fornicata, an invasive gastropod which colonized shallow waters around European coasts during the 20th century. We investigated the effects of 10 weeks' exposure to current (380 µatm) and elevated (550, 750, 1,000 µatm) pCO2 on this engineer species using an acute temperature increase (1 °C/12 h) as the test. Respiration rates were measured on both males (small individuals) and females (large individuals). Mortality increased suddenly from 34 °C, particularly in females. Respiration rate in C. fornicata increased linearly with temperature between 18 and 34 °C, but no differences were detected between the different pCO2 conditions either in the regressions between respiration rate and temperature or in Q10 values. In the same way, condition indices were similar in all the pCO2 treatments at the end of the experiment, but decreased from the beginning of the experiment. This species was highly resistant to acute exposure to high temperature regardless of pCO2 levels, even though food was limited during the experiment. Crepidula fornicata appears to have either developed resistance mechanisms or a strong phenotypic plasticity to deal with fluctuations of physicochemical parameters in its habitat. This suggests that invasive species may be more resistant to future environmental changes than its native competitors.