Aquatic biomass is a major source to particulate organic matter export in large Arctic rivers.

Arctic rivers provide an integrated signature of the changing landscape and transmit signals of change to the ocean. Here, we use a decade of particulate organic matter (POM) compositional data to deconvolute multiple allochthonous and autochthonous pan-Arctic and watershed-specific sources. Constraints from carbon-to-nitrogen ratios (C:N), δ13C, and Δ14C signatures reveal a large, hitherto overlooked contribution from aquatic biomass. Separation in Δ14C age is enhanced by splitting soil sources into shallow and deep pools (mean ± SD: -228 ± 211 vs. -492 ± 173‰) rather than traditional active layer and permafrost pools (-300 ± 236 vs. -441 ± 215‰) that do not represent permafrost-free Arctic regions. We estimate that 39 to 60% (5 to 95% credible interval) of the annual pan-Arctic POM flux (averaging 4,391 Gg/y particulate organic carbon from 2012 to 2019) comes from aquatic biomass. The remainder is sourced from yedoma, deep soils, shallow soils, petrogenic inputs, and fresh terrestrial production. Climate change-induced warming and increasing CO2 concentrations may enhance both soil destabilization and Arctic river aquatic biomass production, increasing fluxes of POM to the ocean. Younger, autochthonous, and older soil-derived POM likely have different destinies (preferential microbial uptake and processing vs. significant sediment burial, respectively). A small (~7%) increase in aquatic biomass POM flux with warming would be equivalent to a ~30% increase in deep soil POM flux. There is a clear need to better quantify how the balance of endmember fluxes may shift with different ramifications for different endmembers and how this will impact the Arctic system.

Mercury Export from Arctic Great Rivers.

Land-ocean linkages are strong across the circumpolar north, where the Arctic Ocean accounts for 1% of the global ocean volume and receives more than 10% of the global river discharge. Yet estimates of Arctic riverine mercury (Hg) export constrained from direct Hg measurements remain sparse. Here, we report results from a coordinated, year-round sampling program that focused on the six major Arctic rivers to establish a contemporary (2012-2017) benchmark of riverine Hg export. We determine that the six major Arctic rivers exported an average of 20 000 kg y-1 of total Hg (THg, all forms of Hg). Upscaled to the pan-Arctic, we estimate THg flux of 37 000 kg y-1. More than 90% of THg flux occurred during peak river discharge in spring and summer. Normalizing fluxes to watershed area (yield) reveals higher THg yields in regions where greater denudation likely enhances Hg mobilization. River discharge, suspended sediment, and dissolved organic carbon predicted THg concentration with moderate fidelity, while suspended sediment and water yields predicted THg yield with high fidelity. These findings establish a benchmark in the face of rapid Arctic warming and an intensifying hydrologic cycle, which will likely accelerate Hg cycling in tandem with changing inputs from thawing permafrost and industrial activity.

Negative feedback processes following drainage slow down permafrost degradation.

The sustainability of the vast Arctic permafrost carbon pool under climate change is of paramount importance for global climate trajectories. Accurate climate change forecasts, therefore, depend on a reliable representation of mechanisms governing Arctic carbon cycle processes, but this task is complicated by the complex interaction of multiple controls on Arctic ecosystem changes, linked through both positive and negative feedbacks. As a primary example, predicted Arctic warming can be substantially influenced by shifts in hydrologic regimes, linked to, for example, altered precipitation patterns or changes in topography following permafrost degradation. This study presents observational evidence how severe drainage, a scenario that may affect large Arctic areas with ice-rich permafrost soils under future climate change, affects biogeochemical and biogeophysical processes within an Arctic floodplain. Our in situ data demonstrate reduced carbon losses and transfer of sensible heat to the atmosphere, and effects linked to drainage-induced long-term shifts in vegetation communities and soil thermal regimes largely counterbalanced the immediate drainage impact. Moreover, higher surface albedo in combination with low thermal conductivity cooled the permafrost soils. Accordingly, long-term drainage effects linked to warming-induced permafrost degradation hold the potential to alleviate positive feedbacks between permafrost carbon and Arctic warming, and to slow down permafrost degradation. Self-stabilizing effects associated with ecosystem disturbance such as these drainage impacts are a key factor for predicting future feedbacks between Arctic permafrost and climate change, and, thus, neglect of these mechanisms will exaggerate the impacts of Arctic change on future global climate projections.

Drainage enhances modern soil carbon contribution but reduces old soil carbon contribution to ecosystem respiration in tundra ecosystems.

Warming temperatures are likely to accelerate permafrost thaw in the Arctic, potentially leading to the release of old carbon previously stored in deep frozen soil layers. Deeper thaw depths in combination with geomorphological changes due to the loss of ice structures in permafrost, may modify soil water distribution, creating wetter or drier soil conditions. Previous studies revealed higher ecosystem respiration rates under drier conditions, and this study investigated the cause of the increased ecosystem respiration rates using radiocarbon signatures of respired CO2 from two drying manipulation experiments: one in moist and the other in wet tundra. We demonstrate that higher contributions of CO2 from shallow soil layers (0-15 cm; modern soil carbon) drive the increased ecosystem respiration rates, while contributions from deeper soil (below 15 cm from surface and down to the permafrost table; old soil carbon) decreased. These changes can be attributed to more aerobic conditions in shallow soil layers, but also the soil temperature increases in shallow layers but decreases in deep layers, due to the altered thermal properties of organic soils. Decreased abundance of aerenchymatous plant species following drainage in wet tundra reduced old carbon release but increased aboveground plant biomass elevated contributions of autotrophic respiration to ecosystem respiration. The results of this study suggest that drier soils following drainage may accelerate decomposition of modern soil carbon in shallow layers but slow down decomposition of old soil carbon in deep layers, which may offset some of the old soil carbon loss from thawing permafrost.

Plants, microorganisms, and soil temperatures contribute to a decrease in methane fluxes on a drained Arctic floodplain.

As surface temperatures are expected to rise in the future, ice-rich permafrost may thaw, altering soil topography and hydrology and creating a mosaic of wet and dry soil surfaces in the Arctic. Arctic wetlands are large sources of CH4 , and investigating effects of soil hydrology on CH4 fluxes is of great importance for predicting ecosystem feedback in response to climate change. In this study, we investigate how a decade-long drying manipulation on an Arctic floodplain influences CH4 -associated microorganisms, soil thermal regimes, and plant communities. Moreover, we examine how these drainage-induced changes may then modify CH4 fluxes in the growing and nongrowing seasons. This study shows that drainage substantially lowered the abundance of methanogens along with methanotrophic bacteria, which may have reduced CH4 cycling. Soil temperatures of the drained areas were lower in deep, anoxic soil layers (below 30 cm), but higher in oxic topsoil layers (0-15 cm) compared to the control wet areas. This pattern of soil temperatures may have reduced the rates of methanogenesis while elevating those of CH4 oxidation, thereby decreasing net CH4 fluxes. The abundance of Eriophorum angustifolium, an aerenchymatous plant species, diminished significantly in the drained areas. Due to this decrease, a higher fraction of CH4 was alternatively emitted to the atmosphere by diffusion, possibly increasing the potential for CH4 oxidation and leading to a decrease in net CH4 fluxes compared to a control site. Drainage lowered CH4 fluxes by a factor of 20 during the growing season, with postdrainage changes in microbial communities, soil temperatures, and plant communities also contributing to this reduction. In contrast, we observed CH4 emissions increased by 10% in the drained areas during the nongrowing season, although this difference was insignificant given the small magnitudes of fluxes. This study showed that long-term drainage considerably reduced CH4 fluxes through modified ecosystem properties.

Evidence for key enzymatic controls on metabolism of Arctic river organic matter.

Permafrost thaw in the Arctic driven by climate change is mobilizing ancient terrigenous organic carbon (OC) into fluvial networks. Understanding the controls on metabolism of this OC is imperative for assessing its role with respect to climate feedbacks. In this study, we examined the effect of inorganic nutrient supply and dissolved organic matter (DOM) composition on aquatic extracellular enzyme activities (EEAs) in waters draining the Kolyma River Basin (Siberia), including permafrost-derived OC. Reducing the phenolic content of the DOM pool resulted in dramatic increases in hydrolase EEAs (e.g., phosphatase activity increased >28-fold) supporting the idea that high concentrations of polyphenolic compounds in DOM (e.g., plant structural tissues) inhibit enzyme synthesis or activity, limiting OC degradation. EEAs were significantly more responsive to inorganic nutrient additions only after phenolic inhibition was experimentally removed. In controlled mixtures of modern OC and thawed permafrost endmember OC sources, respiration rates per unit dissolved OC were 1.3-1.6 times higher in waters containing ancient carbon, suggesting that permafrost-derived OC was more available for microbial mineralization. In addition, waters containing ancient permafrost-derived OC supported elevated phosphatase and glucosidase activities. Based on these combined results, we propose that both composition and nutrient availability regulate DOM metabolism in Arctic aquatic ecosystems. Our empirical findings are incorporated into a mechanistic conceptual model highlighting two key enzymatic processes in the mineralization of riverine OM: (i) the role of phenol oxidase activity in reducing inhibitory phenolic compounds and (ii) the role of phosphatase in mobilizing organic P. Permafrost-derived DOM degradation was less constrained by this initial 'phenolic-OM' inhibition; thus, informing reports of high biological availability of ancient, permafrost-derived DOM with clear ramifications for its metabolism in fluvial networks and feedbacks to climate.

Assessing contemporary Arctic habitat availability for a woolly mammoth proxy.

Interest continues to grow in Arctic megafaunal ecological engineering, but, since the mass extinction of megafauna ~ 12-15 ka, key physiographic variables and available forage continue to change. Here we sought to assess the extent to which contemporary Arctic ecosystems are conducive to the rewilding of megaherbivores, using a woolly mammoth (M. primigenius) proxy as a model species. We first perform a literature review on woolly mammoth dietary habits. We then leverage Oak Ridge National Laboratories Distributive Active Archive Center Global Aboveground and Belowground Biomass Carbon Density Maps to generate aboveground biomass carbon density estimates in plant functional types consumed by the woolly mammoth at 300 m resolution on Alaska's North Slope. We supplement these analyses with a NASA Arctic Boreal Vulnerability Experiment dataset to downgrade overall biomass estimates to digestible levels. We further downgrade available forage by using a conversion factor representing the relationship between total biomass and net primary productivity (NPP) for arctic vegetation types. Integrating these estimates with the forage needs of woolly mammoths, we conservatively estimate Alaska's North Slope could support densities of 0.0-0.38 woolly mammoth km-2 (mean 0.13) across a variety of habitats. These results may inform innovative rewilding strategies.

Protection of Permafrost Soils from Thawing by Increasing Herbivore Density.

Climate change will cause a substantial future greenhouse gas release from warming and thawing permafrost-affected soils to the atmosphere enabling a positive feedback mechanism. Increasing the population density of big herbivores in northern high-latitude ecosystems will increase snow density and hence decrease the insulation strength of snow during winter. As a consequence, theoretically 80% of current permafrost-affected soils (<10 m) is projected to remain until 2100 even when assuming a strong warming using the Representative Concentration Pathway 8.5. Importantly, permafrost temperature is estimated to remain below -4 °C on average after increasing herbivore population density. Such ecosystem management practices would be therefore theoretically an important additional climate change mitigation strategy. Our results also highlight the importance of new field experiments and observations, and the integration of fauna dynamics into complex Earth System models, in order to reliably project future ecosystem functions and climate.

Pleistocene Arctic megafaunal ecological engineering as a natural climate solution?

Natural climate solutions (NCS) in the Arctic hold the potential to be implemented at a scale able to substantially affect the global climate. The strong feedbacks between carbon-rich permafrost, climate and herbivory suggest an NCS consisting of reverting the current wet/moist moss and shrub-dominated tundra and the sparse forest-tundra ecotone to grassland through a guild of large herbivores. Grassland-dominated systems might delay permafrost thaw and reduce carbon emissions-especially in Yedoma regions, while increasing carbon capture through increased productivity and grass and forb deep root systems. Here we review the environmental context of megafaunal ecological engineering in the Arctic; explore the mechanisms through which it can help mitigate climate change; and estimate its potential-based on bison and horse, with the aim of evaluating the feasibility of generating an ecosystem shift that is economically viable in terms of carbon benefits and of sufficient scale to play a significant role in global climate change mitigation. Assuming a megafaunal-driven ecosystem shift we find support for a megafauna-based arctic NCS yielding substantial income in carbon markets. However, scaling up such projects to have a significant effect on the global climate is challenging given the large number of animals required over a short period of time. A first-cut business plan is presented based on practical information-costs and infrastructure-from Pleistocene Park (northeastern Yakutia, Russia). A 10 yr experimental phase incorporating three separate introductions of herds of approximately 1000 individuals each is costed at US$114 million, with potential returns of approximately 0.3-0.4% yr-1 towards the end of the period, and greater than 1% yr-1 after it. Institutional friction and the potential role of new technologies in the reintroductions are discussed. This article is part of the theme issue 'Climate change and ecosystems: threats, opportunities and solutions'.

Particulate Organic Matter Dynamics in a Permafrost Headwater Stream and the Kolyma River Mainstem.

Ongoing rapid arctic warming leads to extensive permafrost thaw, which in turn increases the hydrologic connectivity of the landscape by opening up subsurface flow paths. Suspended particulate organic matter (POM) has proven useful to trace permafrost thaw signals in arctic rivers, which may experience higher organic matter loads in the future due to expansion and increasing intensity of thaw processes such as thermokarst and river bank erosion. Here we focus on the Kolyma River watershed in Northeast Siberia, the world's largest watershed entirely underlain by continuous permafrost. To evaluate and characterize the present-day fluvial release of POM from permafrost thaw, we collected water samples every 4-7 days during the 4-month open water season in 2013 and 2015 from the lower Kolyma River mainstem and from a small nearby headwater stream (Y3) draining an area completely underlain by Yedoma permafrost (Pleistocene ice- and organic-rich deposits). Concentrations of particulate organic carbon generally followed the hydrograph with the highest concentrations during the spring flood in late May/early June. For the Kolyma River, concentrations of dissolved organic carbon showed a similar behavior, in contrast to the headwater stream, where dissolved organic carbon values were generally higher and particulate organic carbon concentrations lower than for Kolyma. Carbon isotope analysis (δ13C, Δ14C) suggested Kolyma-POM to stem from both contemporary and older permafrost sources, while Y3-POM was more strongly influenced by in-stream production and recent vegetation. Lipid biomarker concentrations (high-molecular-weight n-alkanoic acids and n-alkanes) did not display clear seasonal patterns, yet implied Y3-POM to be more degraded than Kolyma-POM.

Understory vegetation mediates permafrost active layer dynamics and carbon dioxide fluxes in open-canopy larch forests of northeastern Siberia.

Arctic ecosystems are characterized by a broad range of plant functional types that are highly heterogeneous at small (~1-2 m) spatial scales. Climatic changes can impact vegetation distribution directly, and also indirectly via impacts on disturbance regimes. Consequent changes in vegetation structure and function have implications for surface energy dynamics that may alter permafrost thermal dynamics, and are therefore of interest in the context of permafrost related climate feedbacks. In this study we examine small-scale heterogeneity in soil thermal properties and ecosystem carbon and water fluxes associated with varying understory vegetation in open-canopy larch forests in northeastern Siberia. We found that lichen mats comprise 16% of understory vegetation cover on average in open canopy larch forests, and lichen abundance was inversely related to canopy cover. Relative to adjacent areas dominated by shrubs and moss, lichen mats had 2-3 times deeper permafrost thaw depths and surface soils warmer by 1-2°C in summer and less than 1°C in autumn. Despite deeper thaw depths, ecosystem respiration did not differ across vegetation types, indicating that autotrophic respiration likely dominates areas with shrubs and moss. Summertime net ecosystem exchange of CO2 was negative (i.e. net uptake) in areas with high shrub cover, while positive (i.e. net loss) in lichen mats and areas with less shrub cover. Our results highlight relationships between vegetation and soil thermal dynamics in permafrost ecosystems, and underscore the necessity of considering both vegetation and permafrost dynamics in shaping carbon cycling in permafrost ecosystems.

Utilization of ancient permafrost carbon in headwaters of Arctic fluvial networks.

Northern high-latitude rivers are major conduits of carbon from land to coastal seas and the Arctic Ocean. Arctic warming is promoting terrestrial permafrost thaw and shifting hydrologic flowpaths, leading to fluvial mobilization of ancient carbon stores. Here we describe (14)C and (13)C characteristics of dissolved organic carbon from fluvial networks across the Kolyma River Basin (Siberia), and isotopic changes during bioincubation experiments. Microbial communities utilized ancient carbon (11,300 to >50,000 (14)C years) in permafrost thaw waters and millennial-aged carbon (up to 10,000 (14)C years) across headwater streams. Microbial demand was supported by progressively younger ((14)C-enriched) carbon downstream through the network, with predominantly modern carbon pools subsidizing microorganisms in large rivers and main-stem waters. Permafrost acts as a significant and preferentially degradable source of bioavailable carbon in Arctic freshwaters, which is likely to increase as permafrost thaw intensifies causing positive climate feedbacks in response to on-going climate change.

Role of megafauna and frozen soil in the atmospheric CH4 dynamics.

Modern wetlands are the world's strongest methane source. But what was the role of this source in the past? An analysis of global 14C data for basal peat combined with modelling of wetland succession allowed us to reconstruct the dynamics of global wetland methane emission through time. These data show that the rise of atmospheric methane concentrations during the Pleistocene-Holocene transition was not connected with wetland expansion, but rather started substantially later, only 9 thousand years ago. Additionally, wetland expansion took place against the background of a decline in atmospheric methane concentration. The isotopic composition of methane varies according to source. Owing to ice sheet drilling programs past dynamics of atmospheric methane isotopic composition is now known. For example over the course of Pleistocene-Holocene transition atmospheric methane became depleted in the deuterium isotope, which indicated that the rise in methane concentrations was not connected with activation of the deuterium-rich gas clathrates. Modelling of the budget of the atmospheric methane and its isotopic composition allowed us to reconstruct the dynamics of all main methane sources. For the late Pleistocene, the largest methane source was megaherbivores, whose total biomass is estimated to have exceeded that of present-day humans and domestic animals. This corresponds with our independent estimates of herbivore density on the pastures of the late Pleistocene based on herbivore skeleton density in the permafrost. During deglaciation, the largest methane emissions originated from degrading frozen soils of the mammoth steppe biome. Methane from this source is unique, as it is depleted of all isotopes. We estimated that over the entire course of deglaciation (15,000 to 6,000 year before present), soils of the mammoth steppe released 300-550 Pg (10(15) g) of methane. From current study we conclude that the Late Quaternary Extinction significantly affected the global methane cycle.