ALBERT

Description: The Arctic is exposed to even faster temperature changes than most other areas on Earth. Constantly increasing temperature will lead to thawing permafrost and changes in the methane (CH4) emissions from wetlands. One of the places exposed to those changes is the Abisko–Stordalen Mire in northern Sweden, where climate and vegetation studies have been conducted since the 1970s. In our study, we analyzed field-scale methane emissions measured by the eddy covariance method at Abisko–Stordalen Mire for 3 years (2014–2016). The site is a subarctic mire mosaic of palsas, thawing palsas, fully thawed fens, and open water bodies. A bimodal wind pattern prevalent at the site provides an ideal opportunity to measure mire patches with different permafrost status with one flux measurement system. The flux footprint for westerly winds was dominated by elevated palsa plateaus, while the footprint was almost equally distributed between palsas and thawing bog-like areas for easterly winds. As these patches are exposed to the same climatic and weather conditions, we analyzed the differences in the responses of their methane emission for environmental parameters. The methane fluxes followed a similar annual cycle over the 3 study years, with a gentle rise during spring and a decrease during autumn, without emission bursts at either end of the ice-free season. The peak emission during the ice-free season differed significantly for the two mire areas with different permafrost status: the palsa mire emitted 19 mg-C m−2 d−1 and the thawing wet sector 40 mg-C m−2 d−1. Factors controlling the methane emission were analyzed using generalized linear models. The main driver for methane fluxes was peat temperature for both wind sectors. Soil water content above the water table emerged as an explanatory variable for the 3 years for western sectors and the year 2016 in the eastern sector. The water table level showed a significant correlation with methane emission for the year 2016 as well. Gross primary production, however, did not show a significant correlation with methane emissions. Annual methane emissions were estimated based on four different gap-filing methods. The different methods generally resulted in very similar annual emissions. The mean annual emission based on all models was 3.1 ± 0.3 g-C m−2 a−1 for the western sector and 5.5 ± 0.5 g-C m−2 a−1 for the eastern sector. The average annual emissions, derived from these data and a footprint climatology, were 2.7 ± 0.5 and 8.2 ± 1.5 g-C m−2 a−1 for the palsa and thawing surfaces, respectively. Winter fluxes were relatively high, contributing 27 %–45 % to the annual emissions.

Description: The Arabian Sea (AS) hosts one of the most intense oxygen minimum zones (OMZs) in the world. Observations suggest a decline in O2 in the northern AS over the recent decades accompanied by an intensification of the suboxic conditions there. Over the same period, the local sea surface temperature has risen significantly, particularly over the Arabian Gulf (also known as Persian Gulf, hereafter the Gulf), while summer monsoon winds may have intensified. Here, we simulate the evolution of dissolved oxygen in the AS from 1982 through 2010 and explore its controlling factors, with a focus on changing atmospheric conditions. To this end, we use a set of eddy-resolving hindcast simulations forced with winds and heat and freshwater fluxes from an atmospheric reanalysis. We find a significant deoxygenation in the northern AS, with O2 inventories north of 20∘ N dropping by over 6 % per decade between 100 and 1000 m. These changes cause an expansion of the OMZ volume north of 20∘ N at a rate of 0.6 % per decade as well as an increase in the volume of suboxia and the rate of denitrification by 14 and 15 % per decade, respectively. We also show that strong interannual and decadal variability modulate dissolved oxygen in the northern AS, with most of the O2 decline taking place in the 1980s and 1990s. Using a set of sensitivity simulations we demonstrate that deoxygenation in the northern AS is essentially caused by reduced ventilation induced by the recent fast warming of the sea surface, including in the Gulf, with a contribution from concomitant summer monsoon wind intensification. This is because, on the one hand, surface warming enhances vertical stratification and increases Gulf water buoyancy, thus inhibiting vertical mixing and ventilation of the thermocline. On the other hand, summer monsoon wind intensification causes a rise in the thermocline depth in the northern AS that lowers O2 levels in the upper ocean. Our findings confirm that the AS OMZ is strongly sensitive to upper-ocean warming and concurrent changes in the Indian monsoon winds. Finally, our results also demonstrate that changes in the local climatic forcing play a key role in regional dissolved oxygen changes and hence need to be properly represented in global models to reduce uncertainties in future projections of deoxygenation.

Description: It has long been assumed that the population dynamics of planktonic foraminifera is characterised by synchronous reproduction associated with ontogenetic vertical migration. However, due to contradictory observations, this concept became controversial, and subsequent studies provided evidence both in favour and against these phenomena. Here we present new observations from replicated vertically resolved profiles of abundance and shell size variation in four species of planktonic foraminifera from the tropical Atlantic to test for the presence, pattern, and extent of synchronised reproduction and ontogenetic vertical migration in this oceanic region. Specimens of Globigerinita glutinata, Globigerinoides ruber ruber, Globorotalia menardii and Orbulina universa were collected over the first 700 m resolved at nine depth intervals at nine stations over a period of 14 d. Dead specimens were systematically observed irrespective of the depth interval, sampling day and size. Conversely, specimens in the smaller size fractions dominated the sampled populations at all times and were recorded at all depths, indicating that reproduction might have occurred continuously and throughout the occupied part of the water column. However, a closer look at the vertical and temporal size distribution of specimens within each species revealed an overrepresentation of large specimens in depths at the beginning of the sampling (shortly after the full moon) and an overrepresentation of small individuals at the surface and subsurface by the end of the sampling (around new moon). These observations imply that a disproportionately large portion of the population followed for each species a canonical reproductive trajectory, which involved synchronised reproduction and ontogenetic vertical migration with the descent of progressively maturing individuals. This concept is consistent with the initial observations from the Red Sea, on which the reproductive dynamics of planktonic foraminifera has been modelled. Our data extend this model to non-spinose and microperforate symbiont-bearing species, but contrary to the extension of the initial observations on other species of foraminifera, we cannot provide evidence for ontogenetic vertical migration with ascent during maturation. We also show that more than half of the population does not follow the canonical trajectory, which helps to reconcile the existing contrasting observations. Our results imply that the flux of empty shells of planktonic foraminifera in the open ocean should be pulsed, with disproportionately large amounts of disproportionately large specimens being delivered in pulses caused by synchronised reproduction. The presence of a large population reproducing outside of the canonical trajectory implies that individual foraminifera in a fossil sample will record in the calcite of their shells a range of habitat trajectories, with the canonical trajectory emerging statistically from a substantial background range.

Description: Photosynthetically active foraminifera are prolific carbonate producers in warm, sunlit, surface waters of the oceans. Foraminifera have repeatedly developed mixotrophic strategies (i.e., the ability of an organism or holobiont to both feed and photosynthesize) by facultative or obligate endosymbiosis with microalgae or by sequestering plastids (kleptoplasts) of ingested algae. Mixotrophy provides access to essential nutrients (e.g., N, P) through feeding while providing carbohydrates and lipids produced through photosynthesis, resulting in substantial energetic advantage in warm, sunlit environments where food and dissolved nutrients are scarce. Our morphological as well as stable carbon isotope data provide, as of now, the earliest (Mid-Devonian) evidence for photosynthetic activity in the first advanced, multichambered, calcareous foraminifera, Semitextularia, from the tropical shelf of the Laurussia paleocontinent. This adaptation likely influenced the evolutionary radiation of calcareous Foraminifera in the Devonian (“Givetian revolution”), one of the most important evolutionary events in foraminiferal history, that coincided with the worldwide development of diverse calcifying marine communities inhabiting shelf environments linked with Devonian stromatoporoid coral reefs.

Description: The Arctic is warming rapidly, especially in winter, which is causing large-scale reductions in snow cover. Snow is one of the main controls on soil thermodynamics, and changes in its thickness and extent affect both permafrost thaw and soil biogeochemistry. Since soil respiration during the cold season potentially offsets carbon uptake during the growing season, it is essential to achieve a realistic simulation of the effect of snow cover on soil conditions to more accurately project the direction of arctic carbon–climate feedbacks under continued winter warming. The Lund–Potsdam–Jena General Ecosystem Simulator (LPJ-GUESS) dynamic vegetation model has used – up until now – a single layer snow scheme, which underestimated the insulation effect of snow, leading to a cold bias in soil temperature. To address this shortcoming, we developed and integrated a dynamic, multi-layer snow scheme in LPJ-GUESS. The new snow scheme performs well in simulating the insulation of snow at hundreds of locations across Russia compared to observations. We show that improving this single physical factor enhanced simulations of permafrost extent compared to an advanced permafrost product, where the overestimation of permafrost cover decreased from 10 % to 5 % using the new snow scheme. Besides soil thermodynamics, the new snow scheme resulted in a doubled winter respiration and an overall higher vegetation carbon content. This study highlights the importance of a correct representation of snow in ecosystem models to project biogeochemical processes that govern climate feedbacks. The new dynamic snow scheme is an essential improvement in the simulation of cold season processes, which reduces the uncertainty of model projections. These developments contribute to a more realistic simulation of arctic carbon–climate feedbacks.

Description: The Cretaceous evolution of angiosperm leaves towards higher vein densities enables unprecedented leaf stomatal conductance. Still, simulating and quantifying the impact of such change on plant productivity and transpiration in the specific environmental conditions of the Cretaceous remain challenging. Here, we address this issue by combining a paleo proxy-based model with a full atmosphere–vegetation model that couples stomatal conductance to carbon assimilation. Based on the fossil record, we build and evaluate three consistent proto-angiosperm vegetation parameterizations under two end-member scenarios of pCO2 (280 and 1120 ppm) for the mid-Cretaceous: a reduction of leaf hydraulic or photosynthetic capacity and a combination of both, supported by a likely coevolution of stomatal conductance and photosynthetic biochemistry. Our results suggest that decreasing leaf hydraulic and/or photosynthetic capacities always generates a reduction of transpiration that is predominantly the result of plant productivity variations modulated by light, water availability in the soil, atmospheric evaporative demand and pCO2. The high pCO2 acts as a fertilizer on plant productivity that strengthens plant transpiration and water-use efficiency. However, we show that proto-angiosperm physiology does not allow vegetation to grow under low pCO2 because of a positive feedback between leaf stomatal conductance and leaf area index. Our modeling approach stresses the need to better represent paleovegetation physiological traits. It also confirms the hypothesis of a likely evolution of angiosperms from a state of low leaf hydraulic and photosynthetic capacities at high pCO2 to a state of high leaf hydraulic and photosynthetic capacities linked to leaves with more and more veins together with a more efficient biochemistry at low pCO2.

Description: Within the last years pollen grains have gained increasing attention due to their cloud-forming potential. Especially the discovery that ice nucleating macromolecules (INMs) or subpollen particles (SPPs) obtained from pollen grains are able to initiate freezing has stirred up interest in pollen. INMs and SPPs are much smaller and potentially more numerous than pollen grains and could significantly affect cloud formation in the atmosphere. However, INMs and SPPs are not clearly distinguished. This has motivated the present study, which focuses on birch pollen and investigates the relationship between pollen grains, INMs, and SPPs. According to the usage of the term SPP in the medical fields, we define SPPs as the starch granules contained in pollen grains. We show that these insoluble SPPs are only obtained when fresh pollen grains are used to generate aqueous extracts from pollen. Due to the limited seasonal availability of fresh pollen grains, almost all studies have been conducted with commercial pollen grains. To enable the investigation of the SPPs we develop an alternative extraction method to generate large quantities of SPPs from commercial pollen grains. We show that INMs are not bonded to SPPs (i.e. can be washed off with water). Further, we find that purified SPPs are not ice nucleation active: after several times of washing SPPs with ultrapure water the ice nucleation activity completely disappears. To our knowledge, this is the first study to investigate the ice nucleation activity of isolated SPPs. To study the chemical nature of the INMs, we use fluorescence spectroscopy. Fluorescence excitation–emission maps indicate a strong signal in the protein range (maximum around λex = 280 nm and λem = 330 nm) with all ice nucleation active samples. In contrast, with purified SPPs the protein signal is lost. We also quantify the protein concentration with the Bradford assay. The protein concentration ranges from 77.4 µg mL−1 (highly concentrated INMs) to below 2.5 µg mL−1 (purified SPPs). Moreover, we investigate the connection between proteins and ice nucleation activity by treating the ice nucleation active samples with subtilisin A and urea to unfold and digest the proteins. After this treatment the ice nucleation activity clearly diminished. The results indicate a linkage between ice nucleation activity and protein concentration. The missing piece of the puzzle could be a glycoprotein which exhibits carboxylate functionalities, can bind water in tertiary structures, and displays degeneration and unfolding of its secondary structure due to heat treatment or reaction with enzymes. Even though purified SPPs are not ice nucleation active they could act as carriers of INMs and distribute those in the atmosphere.

Description: The surface mixed layer (ML) in the Mediterranean Sea is a well-stratified domain characterized by low macronutrients and low chlorophyll content for almost 6 months of the year. In this study we characterize the biogeochemical cycling of nitrogen (N) in the ML by analyzing simultaneous in situ measurements of atmospheric deposition, nutrients in seawater, hydrological conditions, primary production, heterotrophic prokaryotic production, N2 fixation and leucine aminopeptidase activity. Dry deposition was continuously measured across the central and western open Mediterranean Sea, and two wet deposition events were sampled, one in the Ionian Sea and one in the Algerian Basin. Along the transect, N budgets were computed to compare the sources and sinks of N in the mixed layer. In situ leucine aminopeptidase activity made up 14 % to 66 % of the heterotrophic prokaryotic N demand, and the N2 fixation rate represented 1 % to 4.5 % of the phytoplankton N demand. Dry atmospheric deposition of inorganic nitrogen, estimated from dry deposition of nitrate and ammonium in aerosols, was higher than the N2 fixation rates in the ML (on average 4.8-fold). The dry atmospheric input of inorganic N represented a highly variable proportion of biological N demand in the ML among the stations, 10 %–82 % for heterotrophic prokaryotes and 1 %–30 % for phytoplankton. As some sites were visited on several days, the evolution of biogeochemical properties in the ML and within the nutrient-depleted layers could be followed. At the Algerian Basin site, the biogeochemical consequences of a wet dust deposition event were monitored through high-frequency sampling. Notably, just after the rain, nitrate was higher in the ML than in the nutrient-depleted layer below. Estimates of nutrient transfer from the ML into the nutrient-depleted layer could explain up to a third of the nitrate loss from the ML. Phytoplankton did not benefit directly from the atmospheric inputs into the ML, probably due to high competition with heterotrophic prokaryotes, also limited by N and phosphorus (P) availability at the time of this study. Primary producers decreased their production after the rain but recovered their initial state of activity after a 2 d lag in the vicinity of the deep chlorophyll maximum layer.

Description: Recent observations have shown that phytoplankton biomass increases in the North Atlantic during winter, even when the mixed layer is deepening and light is limited. Current theories suggest that this is due to a release from grazing pressure. Here we demonstrate that the often-used grazing models that are linear at low phytoplankton concentration do not allow for a wintertime increase in phytoplankton biomass. However, mathematical formulations of grazing as a function of phytoplankton concentration that are quadratic at low concentrations (or more generally decrease faster than linearly as phytoplankton concentration decreases) can reproduce the fall to spring transition in phytoplankton, including wintertime biomass accumulation. We illustrate this point with a minimal model for the annual cycle of North Atlantic phytoplankton designed to simulate phytoplankton concentration as observed by BioGeoChemical-Argo (BGC-Argo) floats in the North Atlantic. This analysis provides a mathematical framework for assessing hypotheses of phytoplankton bloom formation.

Description: Although the burrowing activity of some species (e.g., gophers) is well studied, a comprehensive inventory of burrowing animals in adjacent biomes is not yet known, despite the potential importance of burrowing activity on the physical and chemical evolution of Earth's surface. In this study, we review the available information with a focus on the following: (a) an inventory of burrowing vertebrates and invertebrates along the climate and ecological gradient in Chile; (b) the dimensions and characteristics of burrows; and (c) calculation of excavation rates by local species compositions. Methods used include a literature compilation (〉 1000 studies) of Chilean burrowing animal species integrated with global, species-specific excavation rates. A field study augments literature findings with quantification of the zoogeomorphic effects on hillslope mass transport at the animal community level and along the arid to humid–temperate climate gradient within the Chilean Coastal Cordillera (27–38∘ S latitude). The literature review indicates a minimum of 45 vertebrate and 345 invertebrate burrowing species distributed across Chile in different biomes. Burrowing depths for Chilean mammals range between 3 m (e.g., for skunks, Conepatus) and 0.25 m (for rock rats, Aconaemys). For invertebrates, burrowing depths in Chile range between 1 m for scorpions to 0.3 m for spiders. In comparison, globally documented maximum burrow depths reach up to more than 6 m for vertebrates (gopher tortoises and aardvarks) and 4 m for invertebrates (ants). Minimum excavation rates of local animal communities observed from field sites in Chile are 0.34 m3 ha−1 yr−1 for the arid site, 0.56 m3 ha−1 yr−1 for the semiarid site, 0.93 m3 ha−1 yr−1 for the mediterranean site and 0.09 m3 ha−1 yr−1 for the humid–temperate site, with the latter likely an underestimation. The calculated minimum Chilean excavation rates are within the large range of globally observed single species rates ranging between 0.01 and 56.20 m3 ha−1 yr−1 for vertebrates and from 0.01 to 37.31 m3 ha−1 yr−1 for invertebrates. Taken together, results not only highlight the diverse and latitudinally varying number of burrowing vertebrates and invertebrates present in different biomes, but also foster the understanding of how burrowing activity changes over a gradient and is influenced by mean annual temperature, mean annual precipitation, slope aspect and latitudinal-related incoming solar energy.