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.

Organic-geochemical proxies (UK'37 and TEXH86) records in surface sediments of the tropical Eastern Indian Ocean

In this study we reconstruct sea surface temperatures (SSTs) using two lipid-based biomarker proxies (alkenone unsaturation index UK'37 and TEX86 index based on glycerol dibiphytanyl glycerol tetraethers) in 36 surface sediment samples from the Indonesian continental margin off west Sumatra and south of Java and the Lesser Sunda Islands. Comparison of measured temperatures (World Ocean Atlas 09) to reconstructed temperatures suggests that SST-UK'37 reflects the SE monsoon SST in the upwelling area south of Java and the Lesser Sunda Islands, whereas Temp-TEXH86 estimates are up to 2°C lower than SST-UK'37. This offset is possibly related to either one or a combination of two factors: i) the depth habitats of the source organisms; ii) different seasonal production and/ or seasonality of export associated with phytoplankton blooming triggered by primary productivity. In the non-upwelling area off west Sumatra, the alkenone-based SSTs are cooler than measured temperatures during the entire year, likely due to the reduced sensitivity of the UK'37 proxy beyond 28°C. However, reconstructed temperatures based on TEXH86 are consistent with mean annual SST, implying that the Temp-TEXH86 reflects the mean annual SST in the non-upwelling area of the tropical Eastern Indian Ocean.

Pollen from 50 lake surface samples in Brandenburg, Germany, in 2009

Past changes in plant and landscape diversity can be evaluated through pollen analysis, however, pollen based diversity indexes are potentially biased by differential pollen production and deposition. Studies examining the relationship between pollen and landscape diversity are therefore needed. The aim of this study is to evaluate how different pollen based indexes capture aspects of landscape diversity. Pollen counts were obtained from surface samples of 50 small to medium sized lakes in Brandenburg (Northeast Germany) and compiled into two sets, with one containing all pollen counts from terrestrial plants and the second restricted to wind-pollinated taxa. Both sets were adjusted for the pollen production/dispersal bias using the REVEALS model. A high resolution biotope map was used to extract the density of total biotopes and different biotopes per area as parameters describing landscape diversity. In addition tree species diversity was obtained from forest inventory data. The Shannon index and the number of taxa in a sample of 10 pollen grains are highly correlated and provide a useful measure of pollen type diversity which corresponds best to landscape diversity within one km of the lake and the proportion of non-forested area within seven km. Adjustments of the pollen production/dispersal bias only slightly improve the relationships between pollen diversity and landscape diversity for the restricted dataset as well as for the forest inventory data and corresponding pollen types. Using rarefaction analysis, we propose the following convention: pollen type diversity is represented by the number of types in a small sample (low count e.g. 10), pollen type richness is the number of types in a large sample (high count e.g. 500) and pollen sample evenness is characterized by the ratio of the two. Synthesis. Pollen type diversity is a robust index that captures vegetation structure and landscape diversity. It is ideally suited for between site comparisons as it does not require high pollen counts. In concert with pollen type richness and evenness, it helps evaluating the effect of climate change and human land use on vegetation structure on long timescales.

Characteristics of introduced Aphididae species, an endemic fly (Anatalanta aptera) and an introduced ground beetle (Merizodus soledadinus) on the Kerguelen Islands

The suite of environments and anthropogenic modifications of sub-Antarctic islands provide key opportunities to improve our understanding of the potential consequences of climate change and biological species invasions on terrestrial ecosystems. The profound impact of human introduced invasive species on indigenous biota, and the facilitation of establishment as a result of changing thermal conditions, has been well documented on the French sub-Antarctic Kerguelen Islands (South Indian Ocean). The present study provides an overview of the vulnerability of sub-Antarctic terrestrial communities with respect to two interacting factors, namely climate change and alien insects. We present datasets assimilated by our teams on the Kerguelen Islands since 1974, coupled with a review of the literature, to evaluate the mechanism and impact of biological invasions in this region. First, we consider recent climatic trends of the Antarctic region, and its potential influence on the establishment, distribution and abundance of alien insects, using as examples one fly and one beetle species. Second, we consider to what extent limited gene pools may restrict alien species' colonisations. Finally, we consider the vulnerability of native communities to aliens using the examples of one beetle, one fly, and five aphid species taking into consideration their additional impact as plant virus vectors. We conclude that the evidence assimilated from the sub-Antarctic islands can be applied to more complex temperate continental systems as well as further developing international guidelines to minimise the impact of alien species.

Chemical composition of zoobenthos and bottom sediments from the gas-hydrate seep area near the Paramushir Island, Kuril Island arc

Bioaccumulation of metals by zoobenthos was investigated during cruise 11A of R/V Akademik Mstislav Keldysh in the vicinity of a gas-hydrate seep off Paramushir Island in the Sea of Okhotsk. Atomic absorption studies of concentrations of Al, Fe, Mn, Ni, Cu and Zn in zoobenthos (polychaetes, bivalves, ophiurans and echinoderms) collected from depths of 700-800 m indicated that their concentrations in individuals near the seep were not significantly different from those in individuals from other communities. Obtained results indicate that sea urchins and holothurians (non-sorting bottom-feeders) can separate mineral fraction of ingested bottom material.

Elemental contents, non-carbonate, fatty acids and alcohols analysis from different Holes of IODP Expedition 310

During IODP Expedition 310 (Tahiti Sea Level), drowned Pleistocene-Holocene barrier-reef terraces were drilled on the slope of the volcanic island. The deglacial reef succession typically consists of a coral framework encrusted by coralline algae and later by microbialites; the latter make up < 80% of the rock volume. Lipid biomarkers were analyzed in order to identify organisms involved in reef-microbialite formation at Tahiti, as the genesis of deglacial microbialites and the conditions favoring their formation are not fully understood. Sterols plus saturated and monounsaturated short-chain fatty acids predominantly derived from both marine primary producers (algae) and bacteria comprise 44 wt% of all lipids on average, whereas long-chain fatty acids and long-chain alcohols derived from higher land plants represent an average of only 24 wt%. Bacterially derived mono-O-alkyl glycerol ethers (MAGEs) and branched fatty acids (10-Me-C16:0; iso- and anteiso-C15:0 and -C17:0) are exceptionally abundant in the microbial carbonates (average, 19 wt%) and represent biomarkers of intermediate-to-high specificity for sulfate-reducing bacteria. Both are relatively enriched in 13C compared to eukaryotic lipids. No lipid biomarkers indicative of cyanobacteria were preserved in the microbialites. The abundances of Al, Si, Fe, Mn, Ba, pyroxene, plagioclase, and magnetite reflect strong terrigenous influx with Tahitian basalt as the major source. Chemical weathering of the basalt most likely elevated nutrient levels in the reefs and this fertilization led to an increase in primary production and organic matter formation, boosting heterotrophic sulfate reduction. Based on the observed biomarker patterns, sulfate-reducing bacteria were apparently involved in the formation of microbialites in the coral reefs off Tahiti during the last deglaciation.

Two intertidal, non-calcifying macroalgae (Palmaria palmata and Saccharina latissima) show complex and variable responses to short-term CO2 acidification

Ocean acidification, the result of increased dissolution of carbon dioxide (CO2) in seawater, is a leading subject of current research. The effects of acidification on non-calcifying macroalgae are, however, still unclear. The current study reports two 1-month studies using two different macroalgae, the red alga Palmaria palmata (Rhodophyta) and the kelp Saccharina latissima (Phaeophyta), exposed to control (pHNBS = 8.04) and increased (pHNBS = 7.82) levels of CO2-induced seawater acidification. The impacts of both increased acidification and time of exposure on net primary production (NPP), respiration (R), dimethylsulphoniopropionate (DMSP) concentrations, and algal growth have been assessed. In P. palmata, although NPP significantly increased during the testing period, it significantly decreased with acidification, whereas R showed a significant decrease with acidification only. S. latissima significantly increased NPP with acidification but not with time, and significantly increased R with both acidification and time, suggesting a concomitant increase in gross primary production. The DMSP concentrations of both species remained unchanged by either acidification or through time during the experimental period. In contrast, algal growth differed markedly between the two experiments, in that P. palmata showed very little growth throughout the experiment, while S. latissima showed substantial growth during the course of the study, with the latter showing a significant difference between the acidified and control treatments. These two experiments suggest that the study species used here were resistant to a short-term exposure to ocean acidification, with some of the differences seen between species possibly linked to different nutrient concentrations between the experiments.

(Table 1) Basin area, average late-summer speedup, number of sinks and runoff of West Greenland Ice Sheet catchments

We use interferometric synthetic aperture radar observations recorded in a land-terminating sector of western Greenland to characterise the ice sheet surface hydrology and to quantify spatial variations in the seasonality of ice sheet flow. Our data reveal a non-uniform pattern of late-summer ice speedup that, in places, extends over 100 km inland. We show that the degree of late-summer speedup is positively correlated with modelled runoff within the 10 glacier catchments of our survey, and that the pattern of late-summer speedup follows that of water routed at the ice sheet surface. In late-summer, ice within the largest catchment flows on average 48% faster than during winter, whereas changes in smaller catchments are less pronounced. Our observations show that the routing of seasonal runoff at the ice sheet surface plays an important role in shaping the magnitude and extent of seasonal ice sheet speedup.

(Supplement Table 1) Individual measurements of prosome length, dry weight and lipid sacs (length, width, area) in arctic calanoid copepods

We present an accurate, fast, simple and non-destructive photographic method to estimate wax ester and lipid content in single individuals of the calanoid copepod genus Calanus and test this method against gas-chromatographic lipid measurements.

n-Alkanes and lignin in bottom sediments of the West European Basin (area of the battleship Bismark wreck site)

Three bottom sediment cores were collected from the top, slope, and foot of a small topographic high located near the West European continental rise within the Porcupine abyssal plain at the battleship Bismark wreck site. Using high-efficient gas chromatography technique we determined content and examined molecular composition of n-alkane fraction of hydrocarbons and phenol compounds of lignin. n-Alkane and phenol concentrations in bottom sediments of all three cores were low both in values per unit mass of sediments and in organic matter composition that is typical for pelagic deposits of the World Ocean. They vary from 0.07 to 2.01 µg/g of dry sediment and from 0.0001 to 0.01% of TOC; phenol ranges are from 1.43 to 11.1 µg/g and from 0.03 to 0.6%. Non-uniform supply of terrigenous matter to the bottom under conditions of changes in sedimentation environment in different geological epochs is the principal reason for significant variations in n-alkane and lignin concentrations with depth in the cores. Lignin and its derivatives make the main contribution to formation of organic matter composition of the region in study. With respect to n-alkane and lignin concentrations organic matter of deposits of the West European Basin is composed of remains of higher plants and of autochtonous organic matter of marine flora; they have mixed terrigenous-autochtonous (terrigenous-planktonogenic) origin.

Pollen and non-pollen palynomorph records of the Holocene part of the composite sediment core TMD from lake Tso Moriri (NW India) and of modern surface samples from the Tso Moriri region

The high-altitude lake Tso Moriri (32°55'46'' N, 78°19'24'' E; 4522 m a.s.l.) is situated at the margin of the ISM and westerly influences in the Trans-Himalayan region of Ladakh. Human settlements are rare and domestic and wild animals are concentrating at the alpine meadows. A set of modern surface samples and fossil pollen from deep-water TMD core was evaluated with a focus on indicator types revealing human impact, grazing activities and lake system development during the last ca. 12 cal ka BP. Furthermore, the non-pollen palynomorph (NPP) record, comprising remains of limnic algae and invertebrates as well as fungal spores and charred plant tissue fragments, were examined in order to attest palaeolimnic phases and human impact, respectively. Changes in the early and middle Holocene limnic environment are mainly influenced by regional climatic conditions and glacier-fed meltwater flow in the catchment area. The NPP record indicates low lake productivity with high influx of freshwater between ca. 11.5 and 4.5 cal ka BP which is in agreement with the regional monsoon dynamics and published climate reconstructions. Geomorphologic observations suggest that during this period of enhanced precipitation the lake had a regular outflow and contributed large amounts of water to the Sutlej River, the lower reaches of which were integral part of the Indus Civilization area. The inferred minimum fresh water input and maximum lake productivity between ca. 4.5-1.8 cal ka BP coincides with the reconstruction of greatest aridity and glaciation in the Korzong valley resulting in significantly reduced or even ceased outflow. We suggest that lowered lake levels and river discharge on a larger regional scale may have caused irrigation problems and harvest losses in the Indus valley and lowlands occupied by sedentary agricultural communities. This scenario, in turn, supports the theory that, Mature Harappan urbanism (ca. 4.5-3.9 cal ka BP) emerged in order to facilitate storage, protection, administration, and redistribution of crop yields and secondly, the eventual collapse of the Harappan Culture (ca. 3.5-3 cal ka BP) was promoted by prolonged aridity. There is no clear evidence for human impact around Tso Moriri prior to ca. 3.7 cal ka BP, with a more distinct record since ca. 2.7 cal ka BP. This suggests that the sedimentary record from Tso Moriri primarily archives the regional climate history.

(Table 1) Percentage of non-opaque heavy minerals, heavy residue, magnetite, and opaque heavy minerals, DSDP Sites 58-445 and 58-446

Heavy-mineral analyses were made for 39 samples, 27 from DSDP Site 445 and 12 from Site 446. About one-fourth of the samples were so loose that they were easily disaggregated in water. The amount of heavy residue and the magnetite content of the heavy fraction were very high, 0.2 to 44 per cent and (on the average) more than 20 per cent, respectively. Among the non-opaque heavy minerals, common hornblende (0 to 80%) and augite (0 to 98%) are most abundant. Pale-green and bluish-green amphiboles (around 10%) and the epidote group (a few to 48%) are next in abundance. Euhedral apatite and biotite and irregularly shaped chromite are not abundant, but are present throughout the sequence. Hacksaw structure is developed in pale-green amphibole and augite. At Site 445, a fair amount of chlorite and a few glauconite(?) grains are present from Core 445-81 downward. The content of common hornblende and opaque minerals also changes from Core 445-81 downward. A geological boundary may exist between Cores 445-77 and 445-81. Source rocks of the sediments at both sites were basaltic volcanic rocks (possibly alkali suite), schists, and ultramafic rocks. The degree of lithification and amount of heavy residue, and the content of magnetite, non-opaque heavy minerals (excluding mafic minerals), and mafic minerals in the cores were compared with Eocene, Oligocene, and Miocene sandstones of southwest Japan. In many respects, the sediments at Sites 445 and 446 are quite different from those of southwest Japan. From the early Eocene to the early Miocene, the area of these sites belonged to a different geologic province than southwest Japan.

Pedogeomorphology, geochronology and paleobotany of soil profiles obtained in the Nagqu area, central Tibet

Vast areas on the Tibetan Plateau are covered by alpine sedge mats consisting of different species of the genus Kobresia. These mats have topsoil horizons rich in rhizogenic organic matter which creates turfs. As the turfs have recently been affected by a complex destruction process, knowledge concerning their soil properties, age and pedogenesis are needed. In the core area of Kobresia pygmaea mats around Nagqu (central Tibetan Plateau, ca. 4500 m a.s.l.), four profiles were subjected to pedological, paleobotanical and geochronological analyses concentrating on soil properties, phytogenic composition and dating of the turf. The turf of both dry K. pygmaea sites and wet Kobresia schoenoides sites is characterised by an enrichment of living (dominant portion) and dead root biomass. In terms of humus forms, K. pygmaea turfs can be classified as Rhizomulls mainly developed from Cambisols. Wet-site K. schoenoides turfs, however, can be classified as Rhizo-Hydromors developed from Histic Gleysols. At the dry sites studied, the turnover of soil organic matter is controlled by a non-permafrost cold thermal regime. Below-ground remains from sedges are the most frequent macroremains in the turf. Only a few pollen types of vascular plants occur, predominantly originating from sedges and grasses. Large amounts of microscopic charcoal (indeterminate) are present. Macroremains and pollen extracted from the turfs predominantly have negative AMS 14C ages, giving evidence of a modern turf genesis. Bulk-soil datings from the lowermost part of the turfs have a Late Holocene age comprising the last ca. 2000 years. The development of K. pygmaea turfs was most probably caused by an anthropo(zoo)-genetically initiated growth of sedge mats replacing former grass-dominated vegetation ('steppe'). Thus the turfs result from the transformation of pre-existing topsoils comprising a secondary penetration and accumulation of roots. K. schoenoides turfs, however, are characterised by a combined process of peat formation and penetration/accumulation of roots probably representing a (quasi) natural wetland vegetation.