Measurements of velocity-dependent and slip-dependent frictional strength in laboratory friction experiments on natural samples from IODP Expedition 316 and ODP Hole 190-1174B

Slowslip forms part of the spectrum of fault behaviour between stable creep and destructive earthquakes. Slow slip occurs near the boundaries of large earthquake rupture zones and may sometimes trigger fast earthquakes. It is thought to occur in faults comprised of rocks that strengthen under fast slip rates, preventing rupture as a normal earthquake, or on faults that have elevated pore-fluid pressures. However, the processes that control slow rupture and the relationship between slow and normal earthquakes are enigmatic. Here we use laboratory experiments to simulate faulting in natural rock samples taken from shallow parts of the Nankai subduction zone, Japan, where very low-frequency earthquakes - a form of slow slip - have been observed.We find that the fault rocks exhibit decreasing strength over millimetre-scale slip distances rather than weakening due to increasing velocity. However, the sizes of the slip nucleation patches in our laboratory simulations are similar to those expected for the very lowfrequency earthquakes observed in Nankai. We therefore suggest that this type of fault-weakening behaviour may generate slow earthquakes. Owing to the similarity between the expected behaviour of slow earthquakes based on our data, and that of normal earthquakes during nucleation, we suggest that some types of slow slip may represent prematurely arrested earthquakes.

Treeline dynamics in Siberia under changing climates as inferred from an individual-based model for Larix, supplementary material

Siberian boreal forests are expected to expand northwards in the course of global warming. However, processes of the treeline ecotone transition, as well astiming and related climate feedbacks are still not understood. Here, we present 'Larix Vegetation Simulator' LAVESI, an individual-based spatially-explicit model that can simulate Larix gmelinii (RUPR.) RUPR. stand dynamics in an attempt to improve our understanding about past and future treeline movements under changing climates. The relevant processes (growth, seed production and dispersal, establishment and mortality) are incorporated and adjusted to observation data mainly gained from the literature. Results of a local sensitivity analysis support the robustness of the model's parameterization by giving relatively small sensitivity values. We tested the model by simulating tree stands under modern climate across the whole Taymyr Peninsula, north-central Siberia (c. 64-80° N; 92-119° E). We find tree densities similar to observed forests in the northern to mid-treeline areas, but densities are overestimated in the southern parts of the simulated region. Finally, from a temperature-forcing experiment, we detect that the responses of tree stands lag the hypothetical warming by several decades, until the end of 21st century. With our simulation experiments we demonstrate that the newly-developed model captures the dynamics of the Siberian latitudinal treeline.

Solar radiation over and under sea ice and photographic quantification of Ice algal aggregates during the POLARSTERN cruise ARK-XXVII/3 (IceArc) in summer 2012

The ice cover of the Arctic Ocean has been changing dramatically in the last decades and the consequences for the sea-ice associated ecosystem remain difficult to assess. Algal aggregates underneath sea ice have been described sporadically but the frequency and distribution of their occurrence is not well quantified. We used upward looking images obtained by a remotely operated vehicle (ROV) to derive estimates of ice algal aggregate biomass and to investigate their spatial distribution. During the IceArc expedition (ARK-XXVII/3) of RV Polarstern in late summer 2012, different types of algal aggregates were observed floating underneath various ice types in the Central Arctic basins. Our results show that the floe scale distribution of algal aggregates in late summer is very patchy and determined by the topography of the ice underside, with aggregates collecting in dome shaped structures and at the edges of pressure ridges. The buoyancy of the aggregates was also evident from analysis of the aggregate size distribution. Different approaches used to estimate aggregate biomass yield a wide range of results. This highlights that special care must be taken when upscaling observations and comparing results from surveys conducted using different methods or on different spatial scales.

Acute survivorship of the deep-sea coral Lophelia pertusa from the Gulf of Mexico under acidification,warming,and deoxygenation

Changing global climate due to anthropogenic emissions of CO2 are driving rapid changes in the physical and chemical environment of the oceans via warming, deoxygenation, and acidification. These changes may threaten the persistence of species and populations across a range of latitudes and depths, including species that support diverse biological communities that in turn provide ecological stability and support commercial interests. Worldwide, but particularly in the North Atlantic and deep Gulf of Mexico, Lophelia pertusa forms expansive reefs that support biological communities whose diversity rivals that of tropical coral reefs. In this study, L. pertusa colonies were collected from the Viosca Knoll region in the Gulf of Mexico (390 to 450 m depth), genotyped using microsatellite markers, and exposed to a series of treatments testing survivorship responses to acidification, warming, and deoxygenation. All coral nubbins survived the acidification scenarios tested, between pH of 7.67 and 7.90 and aragonite saturation states of 0.92 and 1.47. However, calcification generally declined with respect to pH, though a disparate response was evident where select individuals net calcified and others exhibited net dissolution near a saturation state of 1. Warming and deoxygenation both had negative effects on survivorship, with up to 100% mortality observed at temperatures above 14ºC and oxygen concentrations of approximately 1.5 ml·l-1. These results suggest that, over the short-term, climate change and OA may negatively impact L. pertusa in the Gulf of Mexico, though the potential for acclimation and the effects of genetic background should be considered in future research.

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.

PEECE II mesocosm experiment: Dynamics of extracellular enzyme activities in seawater under changed atmospheric pCO2, 2011

As part of the PeECE II mesocosm project, we investigated the effects of pCO2 levels on the initial step of heterotrophic carbon cycling in the surface ocean. The activities of microbial extracellular enzymes hydrolyzing 4 polysaccharides were measured during the development of a natural phytoplankton bloom under pCO2 conditions representing glacial (190 µatm) and future (750 µatm) atmospheric pCO2. We observed that (1) chondroitin hydrolysis was variable throughout the pre-, early- and late-bloom phases, (2) fucoidanase activity was measurable only in the glacial mesocosm as the bloom developed, (3) laminarinase activity was low and constant, and (4) xylanase activity declined as the bloom progressed. Concurrent measurements of microbial community composition, using denaturing-gradient gel electrophoresis (DGGE), showed that the 2 mesocosms diverged temporally, and from one another, especially in the late-bloom phase. Enzyme activities correlated with bloom phase and pCO2, suggesting functional as well as compositional changes in microbial communities in the different pCO2 environments. These changes, however, may be a response to temporal changes in the development of phytoplankton communities that differed with the pCO2 environment. We hypothesize that the phytoplankton communities produced dissolved organic carbon (DOC) differing in composition, a hypothesis supported by changing amino acid composition of the DOC, and that enzyme activities responded to changes in substrates. Enzyme activities observed under different pCO2 conditions likely reflect both genetic and population-level responses to changes occurring among multiple components of the microbial loop.

Grazing under experimental hypercapnia and elevated temperature does not affect the radula of a chiton (Mollusca, Polyplacophora, Lepidopleurida)

Chitons (class Polyplacophora) are benthic grazing molluscs with an eight-part aragonitic shell armature. The radula, a serial tooth ribbon that extends internally more than half the length of the body, is mineralised on the active feeding teeth with iron magnetite apparently as an adaptation to constant grazing on rocky substrates. As the anterior feeding teeth are eroded they are shed and replaced with a new row. The efficient mineralisation and function of the radula could hypothetically be affected by changing oceans in two ways: changes in seawater chemistry (pH and pCO2) may impact the biomineralisation pathway, potentially leading to a weaker or altered density of the feeding teeth; rising temperatures could increase activity levels in these ectothermic animals, and higher feeding rates could increase wear on the feeding teeth beyond the animals' ability to synthesise, mineralise, and replace radular rows. We therefore examined the effects of pH and temperature on growth and integrity in the radula of the chiton Leptochiton asellus. Our experiment implemented three temperature (10, 15, 20 °C) and two pCO2 treatments (400 µatm, pH 8.0; 2000 µatm, pH 7.5) for six treatment groups. Animals (n = 50) were acclimated to the treatment conditions for a period of 4 weeks. This is sufficient time for growth of ca. 7-9 new tooth rows or 20% turnover of the mineralised portion. There was no significant difference in the number of new (non-mineralised) teeth or total tooth row count in any treatment. Examination of the radulae via SEM revealed no differences in microwear or breakage on the feeding cusps correlating to treatment groups. The shell valves also showed no signs of dissolution. As a lineage, chitons have survived repeated shifts in Earth's climate through geological time, and at least their radulae may be robust to future perturbations.

Differing responses of three Southern Ocean Emiliania huxleyi ecotypes to changing seawater carbonate chemistry

The invasion of anthropogenic carbon dioxide into the surface ocean is altering seawater carbonate speciation, a process commonly called ocean acidification. The high latitude waters of the Southern Ocean are one of the primary and most severely affected regions. Coccolithophores are an important phytoplankton group, responsible for the majority of pelagic calcium carbonate production in the world's oceans, with a distribution that ranges from tropical to polar waters. Emiliania huxleyi is numerically the most abundant coccolithophore species and appears in several different ecotypes. We tested the effects of ocean acidification on 3 carefully selected E. huxleyi ecotypes isolated from the Southern Ocean. Their responses were measured in terms of growth, photosynthesis, calcification, cellular geometry, and stoichiometry. The 3 ecotypes exhibited differing sensitivities in regards to seawater carbonate chemistry when cultured at the same temperature (14°C) and continuous light (110 µmol photons/m2/s). Under future ocean acidification scenarios, particulate inorganic to organic carbon ratios (PIC:POC) decreased by 38-44, 47-51 and 71-98% in morphotype A 'over-calcified' (A o/c), A and B/C, respectively. All ecotypes reduced their rate of calcification, but the cold-water adapted ecotype (morphotype B/C) was by far the most sensitive, and almost ceased calcification at partial pressure of carbon dioxide ( pCO2) levels above 1000 µatm. We recommend that future surveys for E. huxleyi cells in the Southern Ocean should include the capability of recognising 'naked cells' by molecular and microscopic tools. The distinct differences in the physiological responses of these 3 dominant Southern Ocean coccolithophore ecotypes are likely to have consequences for future coccolithophore community structures and thereby the Southern Ocean carbon cycle.

European sea bass, Dicentrarchus labrax, in a changing ocean

Ocean acidification, caused by rising concentrations of carbon dioxide (CO2), is widely considered to be a major global threat to marine ecosystems. To investigate the potential effects of ocean acidification on the early life stages of a commercially important fish species, European sea bass (Dicentrarchus labrax), 12 000 larvae were incubated from hatch through metamorphosis under a matrix of two temperatures (17 and 19 °C) and two seawater pCO2 levels (ambient and 1,000 µatm) and sampled regularly for 42 days. Calculated daily mortality was significantly affected by both temperature and pCO2, with both increased temperature and elevated pCO2 associated with lower daily mortality and a significant interaction between these two factors. There was no significant pCO2 effect noted on larval morphology during this period but larvae raised at 19 °C possessed significantly larger eyes and lower carbon:nitrogen ratios at the end of the study compared to those raised under 17 °C. Similarly, when the incubation was continued to post-metamorphic (juvenile) animals (day 67-69), fish raised under a combination of 19 °C and 1000 µatm pCO2 were significantly heavier. However, juvenile D. labrax raised under this combination of 19 °C and 1000 µatm pCO2 also exhibited lower aerobic scopes than those incubated at 19 °C and ambient pCO2. Most studies investigating the effects of near-future oceanic conditions on the early life stages of marine fish have used incubations of relatively short durations and suggested that these animals are resilient to ocean acidification. Whilst the increased survival and growth observed in this study supports this view, we conclude that more work is required to investigate whether the differences in juvenile physiology observed in this study manifest as negative impacts in adult fish.

General responses of Thoracosphaera heimii grown under a range of pCO2

Ocean acidification is considered a major threat to marine ecosystems and may particularly affect calcifying organisms such as corals, foraminifera and coccolithophores. Here we investigate the impact of elevated pCO2 and lowered pH on growth and calcification in the common calcareous dinoflagellate Thoracosphaera heimii. We observe a substantial reduction in growth rate, calcification and cyst stability of T. heimii under elevated pCO2. Furthermore, transcriptomic analyses reveal CO2 sensitive regulation of many genes, particularly those being associated to inorganic carbon acquisition and calcification. Stable carbon isotope fractionation for organic carbon production increased with increasing pCO2 whereas it decreased for calcification, which suggests interdependence between both processes. We also found a strong effect of pCO2 on the stable oxygen isotopic composition of calcite, in line with earlier observations concerning another T. heimii strain. The observed changes in stable oxygen and carbon isotope composition of T. heimii cysts may provide an ideal tool for reconstructing past seawater carbonate chemistry, and ultimately past pCO2. Although the function of calcification in T. heimii remains unresolved, this trait likely plays an important role in the ecological and evolutionary success of this species. Acting on calcification as well as growth, ocean acidification may therefore impose a great threat for T. heimii.

Being young in a changing world: how temperature and salinity changes interactively modify the performance of larval stages of the barnacle Amphibalanus improvisus

The fate of key species, such as the barnacle Amphibalanus improvisus, in the course of global change is of particular interest since any change in their abundance and/or performance may entail community-wide effects. In the fluctuating Western Baltic, species typically experience a broad range of environmental conditions, which may preselect them to better cope with climate change. In this study, we examined the sensitivity of two crucial ontogenetic phases (naupliar, cypris) of the barnacle toward a range of temperature (12, 20, and 28°C) and salinity (5, 15, and 30 psu) combinations. Under all salinity treatments, nauplii developed faster at intermediate and high temperatures. Cyprid metamorphosis success, in contrast, was interactively impacted by temperature and salinity. Survival of nauplii decreased with increasing salinity under all temperature treatments. Highest settlement rates occurred at the intermediate temperature and salinity combination, i.e., 20°C and 15 psu. Settlement success of "naive" cyprids, i.e., when nauplii were raised in the absence of stress (20°C/15 psu), was less impacted by stressful temperature/salinity combinations than that of cyprids with a stress history. Here, settlement success was highest at 30 psu particularly at low and high temperatures. Surprisingly, larval survival was not highest under the conditions typical for the Kiel Fjord at the season of peak settlement (20°C/15 psu). The proportion of nauplii that ultimately transformed to attached juveniles was, however, highest under these "home" conditions. Overall, only particularly stressful combinations of temperature and salinity substantially reduced larval performance and development. Given more time for adaptation, the relatively smooth climate shifts predicted will probably not dramatically affect this species.

Solar radiation over and under sea ice during the POLARSTERN cruise ARK-XXVI/3 (TransArc) in summer 2011

Arctic sea ice has declined and become thinner and younger (more seasonal) during the last decade. One consequence of this is that the surface energy budget of the Arctic Ocean is changing. While the role of surface albedo has been studied intensively, it is still widely unknown how much light penetrates through sea ice into the upper ocean, affecting sea-ice mass balance, ecosystems, and geochemical processes. Here we present the first large-scale under-ice light measurements, operating spectral radiometers on a remotely operated vehicle (ROV) under Arctic sea ice in summer. This data set is used to produce an Arctic-wide map of light distribution under summer sea ice. Our results show that transmittance through first-year ice (FYI, 0.11) was almost three times larger than through multi-year ice (MYI, 0.04), and that this is mostly caused by the larger melt-pond coverage of FYI (42 vs. 23%). Also energy absorption was 50% larger in FYI than in MYI. Thus, a continuation of the observed sea-ice changes will increase the amount of light penetrating into the Arctic Ocean, enhancing sea-ice melt and affecting sea-ice and upper-ocean ecosystems.