Impact of thermal stress on evolutionary trajectories of pathogen resistance in three-spined stickleback (Gasterosteus aculeatus), supplementary material

Background: Pathogens are a major regulatory force for host populations, especially under stressful conditions. Elevated temperatures may enhance the development of pathogens, increase the number of transmission stages, and can negatively influence host susceptibility depending on host thermal tolerance. As a net result, this can lead to a higher prevalence of epidemics during summer months. These conditions also apply to marine ecosystems, where possible ecological impacts and the population-specific potential for evolutionary responses to changing environments and increasing disease prevalence are, however, less known. Therefore, we investigated the influence of thermal stress on the evolutionary trajectories of disease resistance in three marine populations of three-spined sticklebacks Gasterosteus aculeatus by combining the effects of elevated temperature and infection with a bacterial strain of Vibrio sp. using a common garden experiment. Results: We found that thermal stress had an impact on fish weight and especially on survival after infection after only short periods of thermal acclimation. Environmental stress reduced genetic differentiation (QST) between populations by releasing cryptic within-population variation. While life history traits displayed positive genetic correlations across environments with relatively weak genotype by environment interactions (GxE), environmental stress led to negative genetic correlations across environments in pathogen resistance. This reversal of genetic effects governing resistance is probably attributable to changing environment-dependent virulence mechanisms of the pathogen interacting differently with host genotypes, i.e. GPathogenxGHostxE or (GPathogenxE)x(GHostxE) interactions, rather than to pure host genetic effects, i.e. GHostxE interactions. Conclusion: To cope with climatic changes and the associated increase in pathogen virulence, host species require wide thermal tolerances and pathogen-resistant genotypes. The higher resistance we found for some families at elevated temperatures showed that there is evolutionary potential for resistance to Vibrio sp. in both thermal environments. The negative genetic correlation of pathogen resistance between thermal environments, on the other hand, indicates that adaptation to current conditions can be a weak predictor for performance in changing environments. The observed feedback on selective gradients exerted on life history traits may exacerbate this effect, as it can also modify the response to selection for other vital components of fitness.

Age determination and median grain size of sediment core EN31-PC2

Hemipelagic muds deposited during the past 5.3 cal kyr in the northern Gulf of Mexico (Orca Basin) contain seven intervals punctuated by relatively coarse siliciclastic grain-size peaks, planktonic faunal turnovers, and negative d13C excursions. We believe these episodes represent megaflood deposits reflecting historically unprecedented outfall of North American floodwater and terrigenous mud plumes into the gulf, resulting in collapse of the open-ocean pelagic ecosystem. The deposits record multidecadal episodes of high continental precipitation and large Mississippi River floods at ~4.7, 3.5, 3.0, 2.5, 2.0, 1.2, and 0.3 cal ka (500-1200-year recurrence interval). Variations in tropical plankton frequencies define submillenial warming intervals that culminate in these fluvial episodes. Strengthened tropical currents in the gulf at these times appear to have increased sea surface temperatures and associated flow of moist gulf air to the midwest. Terrestrial paleohydrologic records support the marine evidence for millennial-scale changes in recurrence of large midwest flood episodes.

Isotopic ratios and geochemical composition of ODP Hole 176-735B minerals

The mineralogy and stable (O and C) and Sr isotopic compositions of low-temperature alteration phases were determined in Hole 735B gabbroic rocks in order to understand the processes of low-temperature alteration in this uplifted block of lower oceanic crust. Phyllosilicates include smectite (saponite, Mg montmorillonite, and nontronite), chlorite/smectite, chlorite, talc, and serpentine. Other phases include prehnite, albite, K-feldspar, analcite, natrolite, thompsonite, pyrite, and titanite. The low-grade mineral assemblages mainly represent zeolite facies and lower-temperature "seafloor weathering" processes. Phyllosilicates formed over a range of temperatures but may also reflect variable reaction progress. Alteration temperatures were probably somewhat greater below 1300 meters below seafloor. Mineralogy and isotopic data indicate that conditions were mostly reducing and that seawater solutions were rock dominated. Carbonates formed late from cold and generally oxidizing seawater solution, however, as seawater penetrated downward as the result of fracturing and faulting in the uppermost portion of the uplifted crustal block.

Chemical and isotope composition of carbonate nodules and host bottom sediments from Core DM9-666, Gulf of California

Results of mineralogical and isotopic analyzes of sulfur and carbon in carbonate nodules and host bottom sediments and results of 14C measurement in carbonate nodules are reported. It is proved that the carbonate nodules formed 11-22 thousand years ago in anaerobic diagenesis of bottom sediments rich in organic matter. Isotopic light metabolic carbon dioxide was a source of carbonate for nodules. It formed during microbial degradation of organic matter of bottom sediments.

Environment and habitat characteristics of wintering lemmings in the Arctic

Snow cover has dramatic effects on the structure and functioning of Arctic ecosystems in winter. In the tundra, the subnivean space is the primary habitat of wintering small mammals and may be critical for their survival and reproduction. We have investigated the effects of snow cover and habitat features on the distributions of collared lemming (Dicrostonyx groenlandicus) and brown lemming (Lemmus trimucronatus) winter nests, as well as on their probabilities of reproduction and predation by stoats (Mustela erminea) and arctic foxes (Vulpes lagopus). We sampled 193 lemming winter nests and measured habitat features at all of these nests and at random sites at two spatial scales. We also monitored overwinter ground temperature at a subsample of nest and random sites. Our results demonstrate that nests were primarily located in areas with high micro-topography heterogeneity, steep slopes, deep snow cover providing thermal protection (reduced daily temperature fluctuations) and a high abundance of mosses. The probability of reproduction increased in collared lemming nests at low elevation and in brown lemming nests with high availability of some graminoid species. The probability of predation by stoats was density dependent and was higher in nests used by collared lemmings. Snow cover did not affect the probability of predation of lemming nests by stoats, but deep snow cover limited predation attempts by arctic foxes. We conclude that snow cover plays a key role in the spatial structure of wintering lemming populations and potentially in their population dynamics in the Arctic.

Sediment investigations in the Kara Sea

AMS-14C dated sediment cores from the Ob and Yenisei estuaries and the adjacent inner Kara Sea were investigated to determine the siliclastic and organic carbon fluxes and their relationship to paleoenvironmental changes. The variability of sediment fluxes during Holocene times is related to the post-glacial sea-level rise and changes in river discharge and coastal erosion input. Whereas during the late/middle Holocene most of the terrigenous sediments were deposited in the estuaries and the areas directly off the estuaries, huge amounts of sediments accumulated on the Kara Sea shelf farther north during the early Holocene before about 9 Cal. kyrs. BP. The maximum accumulation at that time is related to the lowered sea level, increased coastal erosion, and increased river discharge due to the final stage of mountain deglaciation of the Putoran Massif. Increased supply of Yenisei-derived material indicated by peak magnetic susceptibility values probably occurred in climate-related pulses culminating near 11, 10, and 9 Cal. kyrs. BP. As sea level rose, the main Holocene depocenter migrated southward. Based on hydrogen index values and n-alkanes, the organic matter is predominantly of terrigenous origin. Maximum accumulation rates of 1.5 to more than 6 g/cm**2/y occurred in the early Holocene sediments, suggesting more humid climatic conditions with an increased vegetation cover in the source area at that time. In general, high organic carbon accumulation rates characterize the estuaries and the inner Kara Sea as important sink for terrigenous organic carbon. A high-resolution record of Holocene variability of magnetic susceptibility (MS) in an AMS14C-dated sediment core from the northern Yenisei estuary may indicate natural variability of Arctic climate change and river discharge on a centennial to millenial time scale. Short-term maxima in MS probably related to warmer climate, enhanced precipitation, intensified weathering/erosion and increased river discharge, display a frequency of about 300 to 700 years.

Mg/Ca-temperature proxy in benthic foraminifera from the Florida Strait

Over the past decade, the ratio of Mg to Ca in foraminiferal tests has emerged as a valuable paleotemperature proxy. However, large uncertainties remain in the relationships between benthic foraminiferal Mg/Ca and temperature. Mg/Ca was measured in benthic foraminifera from 31 high-quality multicore tops collected in the Florida Straits, spanning a temperature range of 5.8° to 18.6°C. New calibrations are presented for Uvigerina peregrina, Planulina ariminensis, Planulina foveolata, and Hoeglundina elegans. The Mg/Ca values and temperature sensitivities vary among species, but all species exhibit a positive correlation that decreases in slope at higher temperatures. The decrease in the sensitivity of Mg/Ca to temperature may potentially be explained by Mg/Ca suppression at high carbonate ion concentrations. It is suggested that a carbonate ion influence on Mg/Ca may be adjusted for by dividing Mg/Ca by Li/Ca. The Mg/Li ratio displays stronger correlations to temperature, with up to 90% of variance explained, than Mg/Ca alone. These new calibrations are tested on several Last Glacial Maximum (LGM) samples from the Florida Straits. LGM temperatures reconstructed from Mg/Ca and Mg/Li are generally more scattered than core top measurements and may be contaminated by high-Mg overgrowths. The potential for Mg/Ca and Mg/Li as temperature proxies warrants further testing.

Carbon/Nitrogen ratio of plant tissues and biomass of belowground consumers after 11 years of experimental treatment, Toolik lake

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The Exploitation Ecosystem Hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.

Geochemistry and morphometry on planktonic foraminifera

About one third of the anthropogenic carbon dioxide (CO2) released into the atmosphere in the past two centuries has been taken up by the ocean. As CO2 invades the surface ocean, carbonate ion concentrations and pH are lowered. Laboratory studies indicate that this reduces the calcification rates of marine calcifying organisms, including planktic foraminifera. Such a reduction in calcification resulting from anthropogenic CO2 emissions has not been observed, or quantified in the field yet. Here we present the findings of a study in the Western Arabian Sea that uses shells of the surface water dwelling planktic foraminifer Globigerinoides ruber in order to test the hypothesis that anthropogenically induced acidification has reduced shell calcification of this species. We found that light, thin-walled shells from the surface sediment are younger (based on 14C and d13C measurements) than the heavier, thicker-walled shells. Shells in the upper, bioturbated, sediment layer were significantly lighter compared to shells found below this layer. These observations are consistent with a scenario where anthropogenically induced ocean acidification reduced the rate at which foraminifera calcify, resulting in lighter shells. On the other hand, we show that seasonal upwelling in the area also influences their calcification and the stable isotope (d13C and d18O) signatures recorded by the foraminifera shells. Plankton tow and sediment trap data show that lighter shells were produced during upwelling and heavier ones during non-upwelling periods. Seasonality alone, however, cannot explain the 14C results, or the increase in shell weight below the bioturbated sediment layer. We therefore must conclude that probably both the processes of acidification and seasonal upwelling are responsible for the presence of light shells in the top of the sediment and the age difference between thick and thin specimens.

Sea surafe temperture calculation for the Arabian Sea

Sea surface temperature (SST) and seawater d18O (d18Ow) were reconstructed in a suite of sediment cores from throughout the Arabian Sea for four distinct time intervals (0 ka, 8 ka, 15 ka, and 20 ka) with the aim of understanding the history of the Indian Monsoon and the climate of the Arabian Sea region. This was accomplished through the use of paired Mg/Ca and d18O measurements of the planktonic foraminifer Globigerinoides ruber. By analyzing basin-wide changes and changes in cross-basinal gradients, we assess both monsoonal and regional-scale climate changes. SST was colder than present for the majority of sites within all three paleotime slices. Furthermore, both the Indian Monsoon and the regional Arabian Sea mean climate have varied substantially over the past 20 kyr. The 20 ka and 15 ka time slices exhibit average negative temperature anomalies of 2.5°-3.5°C attributable, in part, to the influences of glacial atmospheric CO2 concentrations and large continental ice sheets. The elimination of the cross-basinal SST gradient during these two time slices likely reflects a decrease in summer monsoon and an increase in winter monsoon strength. Changes in d18Ow that are smaller than the d18O signal due to global ice volume reflect decreased evaporation and increased winter monsoon mixing. SSTs throughout the Arabian Sea were still cooler than present by an average of 1.4°C in the 8 ka time slice. These cool SSTs, along with lower d18Ow throughout the basin, are attributed to stronger than modern summer and winter monsoons and increased runoff and precipitation. The results of this study underscore the importance of taking a spatial approach to the reconstruction of processes such as monsoon upwelling.

Age determination and pollen profile of sediment core Trentelmoor, Peine, northern Germany

The filling up of the lake which existed in the basin of the Trentelmoor (40 km E of Hannover, Germany) - in Preboreal times was finished 2000 years ago. Since then fen vegetation has covered the former lake's surface. The postglacial development of the vegetation follows the pattern which is typical of Central Europe. However, due to the poorness of the soils around the Trentelmoor, the frequencies of some tree species differ. Beech for example never reached - for the benefit of oak - that importance which this tree species usually gains on better soils. Human impact becomes recognisable in the upper Neolithic for the first time. The area has been settled continuously, but with changing intensities, throughout the last 3000 years. When the manuscript of this paper went to press the results of two radiocarbon age determinations only were completed. An additional three determinations were completed somewhat later. See the accompanying table for results.

Major element concentrations in spinel, olivine and pyroxene from peridotites of ODP Hole 210-1277 of the Newfoundland margin

Serpentinized spinel peridotites of the Newfoundland margin drilled during ODP Leg 210 at Site 1277 have preserved, relic mineral compositions similar to the most depleted abyssal peridotites worldwide and different from those of the conjugate Iberian margin. The samples are derived from mass flows containing clasts of peridotite and gabbro and from in-situ basement, and are mostly mylonitic cpx-poor spinel harzburgites with Cr-rich spinels (Cr#0.35-0.66). Melting of the Newfoundland mantle occurred in the spinel peridotite field and probably exceeded the cpx-out phase boundary for some samples. Using proposed spinel peridotite melting models and experimentally derived phase diagrams, the Newfoundland harzburgites can be modeled as a residue after extraction of 14 to 20-25% melting. Basalts that are interleaved with mass flow deposits on top of the peridotite basement resemble normal to transitional mid-ocean ridge basalt. This, together with the unusually high Cr# of some spinel harzburgites suggest that the formation of basalts and partial melting of the underlying peridotite are not cogenetic. Among mantle samples some of the Newfoundland harzburgites approach mineral compositions of the Bay of island ophiolite and ophiolites from Japan that represent peridotites formed in an arc-setting. Thus, the peridotites drilled at Site 1277 may represent inherited (Caledonian or older) subarc mantle that was exhumed close to the ocean floor during the rifting evolution of the Atlantic. Compared to the spinel harzburgites from Newfoundland, the peridotites from the conjugate Iberian margin are, on average, less depleted and provide evidence for local equilibration in the plagioclase stability field. This can either be explained by an inherited, primary, Ca-richer composition of the Iberia peridotite, or, alternatively, by local melt impregnation and stagnation during continental rifting, and thus refertilizing previously depleted (arc-related) peridotite.

Separating ITCZ- and ENSO-related rainfall changes in the Galápagos over the last 3 kyr using D/H ratios of multiple lipid biomarkers

We present a 3000-yr rainfall reconstruction from the Galápagos Islands that is based on paired biomarker records from the sediment of El Junco Lake. Located in the eastern equatorial Pacific, the climate of the Galápagos Islands is governed by movements of the Intertropical Convergence Zone (ITCZ) and the El Niño-Southern Oscillation (ENSO). We use a novel method for reconstructing past ENSO- and ITCZ-related rainfall changes through analysis of molecular and isotopic biomarker records representing several types of plants and algae that grow under differing climatic conditions. We propose that ?D values of dinosterol, a sterol produced by dinoflagellates, record changes in mean rainfall in El Junco Lake, while dD values of C34 botryococcene, a hydrocarbon unique to the green alga Botryococcus braunii, record changes in rainfall associated with moderate-to-strong El Niño events. We use these proxies to infer changes in mean rainfall and El Niño-related rainfall over the past 3000 yr. During periods in which the inferred change in El Niño-related rainfall opposed the change in mean rainfall, we infer changes in the amount of ITCZ-related rainfall. Simulations with an idealized isotope hydrology model of El Junco Lake help illustrate the interpretation of these proxy reconstructions. Opposing changes in El Niño- and ITCZ-related rainfall appear to account for several of the largest inferred hydrologic changes in El Junco Lake. We propose that these reconstructions can be used to infer changes in frequency and/or intensity of El Niño events and changes in the position of the ITCZ in the eastern equatorial Pacific over the past 3000 yr. Comparison with El Junco Lake sediment grain size records indicates general agreement of inferred rainfall changes over the late Holocene.

CPI, d13C, Corg and diploptene from ODP Hole 169-1034B varved sediments

Saanich Inlet has been a highly productive fjord since the last glaciation. During ODP Leg 169S, nearly 70 m of Holocene sediments were recovered from Hole 1034 at the center of the inlet. The younger sediments are laminated, anaerobic, and rich in organic material (1-2.5 wt.% Corg), whereas the older sediments below 70 mbsf are non-laminated, aerobic, with glacio-marine characteristics and have a significantly lower organic matter content. This difference is also reflected in the changes of interstitial fluids, and in biomarker compositions and their carbon isotope signals. The bacterially-derived hopanoid 17alpha(H),21beta(H)-hop-22(29)-ene (diploptene) occurs in Saanich Inlet sediments throughout the Holocene but is not present in Pleistocene glacio-marine sediments. Its concentration increases after ~6000 years BP up to present time to about 70 µg/g Corg, whereas terrigenous biomarkers such as the n-alkane C31 are low throughout the Holocene (<51 µg/g Corg) and even slightly decrease to 36 µg/g Corg at the most recent time. The increasing concentrations of diploptene in sediments younger than ~6000 years BP separate a recent period of higher primary productivity, stronger anoxic bottom waters, and higher bacterial activity from an older period with lesser activity, heretofore undifferentiated. Carbon isotopic compositions of diploptene in the Holocene are between ~31.5 and ~39.6 per mil PDB after ~6000 years BP. These differences in the carbon isotopic record of diploptene probably reflect changes in microbial community structure of bacteria living at the oxic-anoxic interface of the overlying water column. The heavier isotope values are consistent with the activity of nitrifying bacteria and the lighter isotope values with that of aerobic methanotrophic bacteria. Therefore, intermediate delta13C values probably represent mixtures between the populations. In contrast, carbon isotopic compositions of n-C31 are roughly constant at ~31.4 ± 1.1 per mil PDB throughout the Holocene, indicating a uniform input from cuticular waxes of higher plants. Prior to ~6000 years BP, diploptene enriched in 13C of up to -26.3 per mil PDB is indicative of cyanobacteria living in the photic zone and suggests a period of lower primary productivity, more oxygenated bottom waters, and hence lower bacterial activity during the earliest Holocene.

Representative analyses of pre-tectonic metabasic rocks and metapelitic gneiss from Sverdrupfjella, East Antarctica

Extensive high-grade polydeformed metamorphic provinces surrounding Archaean cratonic nuclei in the East Antarctic Shield record two tectono-thermal episodes in late Mesoproterozoic and late Neoproterozoic-Cambrian times. In Western Dronning Maud Land, the high-grade Mesoproterozoic Maud Belt is juxtaposed against the Archaean Grunehogna Province and has traditionally been interpreted as a Grenvillian mobile belt that was thermally overprinted during the Early Palaeozoic. Integration of new U-Pb sensitive high-resolution ion microprobe and conventional single zircon and monazite age data, and Ar-Ar data on hornblende and biotite, with thermobarometric calculations on rocks from the H.U. Sverdrupfjella, northern Maud Belt, resulted in a more complex P-T-t evolution than previously assumed. A c. 540?Ma monazite, hosted by an upper ampibolite-facies mineral assemblage defining a regionally dominant top-to-NW shear fabric, provides strong evidence for the penetrative deformation in the area being of Pan-African age and not of Grenvillian age as previously reported. Relics of an eclogite-facies garnet-omphacite assemblage within strain-protected mafic boudins indicate that the peak metamorphic conditions recorded by most rocks in the area (T = 687-758°C, P = 9·4-11·3?kbar) were attained subsequent to decompression from P > 12·9?kbar. By analogy with limited U-Pb single zircon age data and on circumstantial textural grounds, this earlier eclogite-facies metamorphism is ascribed to subduction and accretion around 565?Ma. Post-peak metamorphic K-metasomatism under amphibolite-facies conditions is ascribed to the intrusion of post-orogenic granite at c. 480?Ma. The recognition of extensive Pan-African tectonism in the Maud Belt casts doubts on previous Rodinia reconstructions, in which this belt takes a pivotal position between East Antarctica, the Kalahari Craton and Laurentia. Evidence of late Mesoproterozoic high-grade metamorphism during the formation of the Maud Belt exists in the form of c. 1035?Ma zircon overgrowths that are probably related to relics of granulite-facies metamorphism recorded from other parts of the Maud Belt. The polymetamorphic rocks are largely derived from a c. 1140?Ma volcanic arc and 1072 ± 10?Ma granite.