Beavers alter stream macroinvertebrate communities in north-eastern Utah

1. Understanding changes in macroinvertebrate communities is important because they play a large role in stream ecosystem functioning, and they are an important food resource for fish. Beaver-induced changes to stream morphology could alter macroinvertebrate communities, which in turn could affect food webs and ecosystem function. However, studies investigating the effects of North American beaver activities on macroinvertebrates are rare in the inter-mountain west, an area with high potential for beaver-assisted restoration.
2. The aim of this study was to quantify differences in the macroinvertebrate community between unaltered segments of streams and within beaver ponds in north-eastern Utah, U.S.A. We assessed macroinvertebrate species richness, biomass, density, functional feeding group composition, mobility group composition, and macroinvertebrate habitat characteristics to test the hypothesis that macroinvertebrate communities will differ among habitat types (undammed stream segments and beaver ponds) in beaver-occupied streams.
3. Beaver pond communities significantly differed from lotic reach communities in many ways. Beaver ponds were less diverse with 25% fewer species. Although there was variability among streams, in general, beaver ponds had 75% fewer individuals and 90% lower total macroinvertebrate biomass compared to lotic reaches.
4. Regarding functional feeding groups, beaver ponds contained more engulfers, while lotic reaches contained more scrapers, filterers, and gatherers. For mobility groups, beaver ponds had more sprawlers, while lotic reaches had more clingers. Swimmers were also more prevalent in lotic reaches, although this is probably due to the abundance of Baetis within lotic reaches. More beaver pond taxa were classified as lentic-dwelling insects, while more lotic reach taxa were categorised as preferring lotic habitats.
5. The creation of ponds by beavers fundamentally altered the macroinvertebrate community in north-eastern Utah streams. Such changes to stream macroinvertebrate communities suggest that recolonisation of beavers across North America may be altering stream functioning and food webs. Our study highlights the need to further investigate the effects of beaver recolonisation on stream communities.

     

Thaw pond development and initial vegetation succession in experimental plots at a Siberian lowland tundra site

Background and aims

Permafrost degradation has the potential to change the Arctic tundra landscape. We observed rapid local thawing of ice-rich permafrost resulting in thaw pond formation, which was triggered by removal of the shrub cover in a field experiment. This study aimed to examine the rate of permafrost thaw and the initial vegetation succession after the permafrost collapse.

Methods

In the experiment, we measured changes in soil thaw depth, plant species cover and soil subsidence over nine years (2007–2015).

Results

After abrupt initial thaw, soil subsidence in the removal plots continued indicating further thawing of permafrost albeit at a much slower pace: 1 cm y^?1 over 2012–2015 vs. 5 cm y^?1 over 2007–2012. Grass cover strongly increased after the initial shrub removal, but later declined with ponding of water in the subsiding removal plots. Sedges established and expanded in the wetter removal plots. Thereby, the removal plots have become increasingly similar to nearby ‘natural’ thaw ponds.

Conclusions

The nine years of field observations in a unique shrub removal experiment at a Siberian tundra site document possible trajectories of small-scale permafrost collapse and the initial stage of vegetation recovery, which is essential knowledge for assessing future tundra landscape changes.

     

Ecosystem engineering by beavers affects mayfly life histories

1. The North American beaver has been studied as a model ecosystem engineer for many decades. Previous studies have documented physical, chemical and biological impacts attributed to beaver engineering in both aquatic and terrestrial environments. This study focused on the effects of ecosystem engineering by beavers on life histories of a common mayfly and on the potential consequences for mayfly populations. 2. We studied 18 montane beaver ponds of varying size and shape in western Colorado near the Rocky Mountain Biological Laboratory. Our goal was to test whether variation in beaver pond morphology (pond size and shape) explains downstream changes in stream temperature, mayfly size and timing of emergence. 3. Downstream water temperatures varied predictably with pond morphology, being colder downstream of high‐head dams and warmer downstream of low‐head dams. Pond morphology was also a significant predictor of variation in the size of mature female Baetis bicaudatus (the most abundant mayfly), with larger females emerging downstream of high‐head dams and smaller females downstream of low‐head dams. The size of male B. bicaudatus was not significantly related to pond morphology or stream temperature. There was no relationship between pond morphology and variation in the timing of emergence of Baetis (males or females) between upstream and downstream reaches. 4. Our results have implications for the effects of beaver ponds on Baetis individual fitness because large Baetis females are more fecund. Therefore, predictable female size variation associated with beaver pond morphology makes it possible to model the effects of beaver activity on local contributions of Baetis to the regional pool of reproductive adults at the catchment scale. Additionally, predictable changes in the size of emerging mayflies may have important consequences for the magnitude of aquatic to terrestrial resource subsidies in beaver‐modified systems.     

Volatile emissions from thawing permafrost soils are influenced by meltwater drainage conditions

Vast amounts of carbon are bound in both active layer and permafrost soils in the Arctic. As a consequence of climate warming, the depth of the active layer is increasing in size and permafrost soils are thawing. We hypothesize that pulses of biogenic volatile organic compounds are released from the near‐surface active layer during spring, and during late summer season from thawing permafrost, while the subsequent biogeochemical processes occurring in thawed soils also lead to emissions. Biogenic volatile organic compounds are reactive gases that have both negative and positive climate forcing impacts when introduced to the Arctic atmosphere, and the knowledge of their emission magnitude and pattern is necessary to construct reliable climate models. However, it is unclear how different ecosystems and environmental factors such as drainage conditions upon permafrost thaw affect the emission and compound composition. Here we show that incubations of frozen B horizon of the active layer and permafrost soils collected from a High Arctic heath and fen release a range of biogenic volatile organic compounds upon thaw and during subsequent incubation experiments at temperatures of 10°C and 20°C. Meltwater drainage in the fen soils increased emission rates nine times, while having no effect in the drier heath soils. Emissions generally increased with temperature, and emission profiles for the fen soils were dominated by benzenoids and alkanes, while benzenoids, ketones, and alcohols dominated in heath soils. Our results emphasize that future changes affecting the drainage conditions of the Arctic tundra will have a large influence on volatile emissions from thawing permafrost soils – particularly in wetland/fen areas.     

Ecosystem engineering by invasive exotic beavers reduces in-stream diversity and enhances ecosystem function in Cape Horn,Chile

Species invasions are of global significance, but predicting their impacts can be difficult. Introduced ecosystem engineers, however, provide an opportunity to test the underlying mechanisms that may be common to all invasive engineers and link relationships between changes in diversity and ecosystem function, thereby providing explanatory power for observed ecological patterns. Here we test specific predictions for an invasive ecosystem engineer by quantifying the impacts of habitat and resource modifications caused by North American beavers (Castor canadensis) on aquatic macroinvertebrate community structure and stream ecosystem function in the Cape Horn Biosphere Reserve, Chile. We compared responses to beavers in three habitat types: (1) forested (unimpacted) stream reaches, (2) beaver ponds, and (3) sites immediately downstream of beaver dams in four streams. We found that beaver engineering in ponds created taxonomically simplified, but more productive, benthic macroinvertebrate assemblages. Specifically, macroinvertebrate richness, diversity and number of functional feeding groups were reduced by half, while abundance, biomass and secondary production increased three- to fivefold in beaver ponds compared to forested sites. Reaches downstream of beaver ponds were very similar to natural forested sections. Beaver invasion effects on both community and ecosystem parameters occurred predominantly via increased retention of fine particulate organic matter, which was associated with reduced macroinvertebrate richness and diversity (via homogenization of benthic microhabitat) and increased macroinvertebrate biomass and production (via greater food availability). Beaver modifications to macroinvertebrate community structure were largely confined to ponds, but increased benthic production in beaver-modified habitats adds to energy retention and flow for the entire stream ecosystem. Furthermore, the effects of beavers on taxa richness (negative) and measures of macroinvertebrate biomass (positive) were inversely related. Thus, while a generally positive relationship between diversity and ecosystem function has been found in a variety of systems, this work shows how they can be decoupled by responding to alterative mechanisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Climate‐associated tundra thaw pond formation and range expansion of boreal zooplankton predators

Most of the freshwater component of the Earth's surface is composed of shallow tundra ponds. These high latitude ecosystems have been exposed to a variety of abiotic disturbances associated with recent environmental change. However, the biological significance of these changes remains poorly understood. Here, we characterize the abiotic disturbance to the shallow tundra ponds of northwest Alaska. We used historical aerial imagery to determine that up to 53% of the sampled ponds have formed during the recent warmer decades (since the 1970s). We discovered that two top predator species (phantom midges of the genus Chaoborus) of the freshwater zooplankton have recently undergone range expansion, forming widespread (a scale of hundreds of km) stable tundra populations. We assessed the population persistence of these boreal predators by longitudinal sampling over 14 yr. Recent thaw ponds had significantly dissimilar zooplankton communities to communities of ponds that formed before 1950. Both predator and herbivore species differed by age of pond. Younger pond ages and warmer surface temperatures were the significant predictors of the presence of temperate Chaoborus americanus in tundra ponds. Ponds containing tundra populations of C. americanus and C. cf. flavicans were associated with recent formation (83–90%). Recent ponds in river valleys appeared more important than recent ponds near roads as colonization corridors for C. americanus. Only 24% of the tundra keystone predator, Heterocope septentrionalis, populations were from recent ponds. Our results suggest that climate‐associated disturbance can lead to a widespread stable range expansion of boreal species despite the propinquity of older ponds with top‐down control exerted by an endemic keystone predator.     

Does the morphology of beaver ponds alter downstream ecosystems?

Differences among lake morphologies often explain variation in characteristics of lentic ecosystems. Although beaver ponds also vary in morphology, previous studies have not examined the effects of such variation on downstream ecosystems. This study evaluated downstream effects of multiple beaver ponds in the Colorado Rocky Mountains during one low and one high-flow year. Beaver pond morphology was described as the natural log transformed ratio of beaver dam height (which determines hydraulic head) to pond surface area and related to pond spillover phytoplankton and characteristics of the ecosystem downstream (nutrient concentrations, limiting nutrients, periphyton, benthic organic matter (BOM), and benthic invertebrate consumers). Nitrate concentration increased systematically downstream of beaver ponds, but only in the low flow year when groundwater influences predominated. Effects of beaver ponds on soluble reactive phosphorus concentration depended on pond morphology, increasing downstream of small ponds with high dams, but only during the low-flow year. In situ experiments showed that neither beaver activity nor pond morphology predicted periphyton-limiting nutrients downstream. Both periphyton biomass and BOM decreased downstream of small ponds with high dams but pond morphology did not predict abundance of invertebrate grazers or detritus-feeding consumers. While suspension feeding invertebrates increased downstream from small ponds with high dams, variation in chlorophyll a from water spilling over beaver dams did not follow a similar pattern. We conclude that the effects of beaver ponds on downstream nutrients, resources and consumers are rarely systematic, but instead depend on variation in pond morphology and on annual hydrologic variation.     

Fish recruitment, dispersal, and trophic interactions in a heterogeneous lotic environment

I examine how dispersal of juvenile creek chubs (Semotilusatromaculatus) from beaver ponds into adjacent stream environments interacts with temporal abiotic variability to influence fish foraging, growth, and long-term persistence in the lotic ecosystem. Minnow trapping in upstream and downstream beaver ponds, along with weir traps used to monitor directional movement, indicated that most chubs colonized the stream from the downstream beaver pond. Large annual fluctuations in density of age 0 creek chubs occurred in the stream over a 10-year sampling period. Multiple regression analysis indicated that stream temperature, precipitation, and the density of reproductive creek chubs were not correlated with summer density of age 0 chubs in the stream. The factor most strongly associated with increased density of age 0 creek chubs was creation of the downstream beaver pond during the 6th–7th years of the study, suggesting dispersal from the pond was the primary factor determining age 0 fish density in the stream. Most individuals in the strong year classes neither persisted in the stream through their first winter nor resulted in an increased abundance of older age classes in later years. Comparison of age 0 fish density in summer to the proportion of fish surviving to age 1 in spring suggested that overwinter mortality was density dependent. Furthermore, a comparison of the size structure for age 0 individuals in summer to age 1 individuals the following spring indicated that winter mortality was size dependent. Experiments in an artificial stream adjacent to the natural channel revealed that fish growth was strongly density dependent, decreasing as fish density increased across both spring and summer, and elevated and low discharge. The decline in invertebrate prey captured by the fish and the subsequent decline in fish growth appeared to be particularly pronounced under low discharge in summer. Changes in juvenile creek chub density had no significant effect on benthic insect or crustacean abundance, suggesting that exploitative competition for limited invertebrate drift resources was a more important cause of density- dependent growth than depressed local benthic invertebrate abundance. These results suggest that lotic regions adjacent to beaver ponds act as potential reproductive “sinks” for dispersing juveniles confronting seasonal and flow-mediated restrictions on resource acquisition and growth, and the occurrence of seasonal bottlenecks to their survival, especially harsh winter conditions. Received: 9 September 1996 / Accepted: 8 August 1997     

Bacterial communities and greenhouse gas emissions of shallow ponds in the High Arctic

Permafrost thawing in lowland Arctic tundra results in a polygonal patterned landscape and the formation of numerous small ponds. These ponds emit biologically mediated carbon dioxide (CO₂) and methane (CH₄). Their greenhouse gas (GHG) emissions are variable, for reasons that are not well understood. Emissions are related to a balance between GHG producers and consumers, as well as to physical properties of the water column controlling gas exchange rates with the atmosphere. Here, we investigated the bacterial diversity of polygonal and runnel ponds, two geomorphologically distinct pond types commonly found in continuous permafrost regions. Using a combination of 16S rRNA Sanger sequencing and high-throughput amplicon sequencing, we found that bacterial communities in overlying waters were clearly dominated by carbon degraders and were similar in both pond types, despite their variable physical and chemical properties. However, surface sediment communities in the two pond types were significantly different. Polygonal pond sediment was colonized by carbon degraders (46–29 %), cyanobacteria (20–27 %) that take up CO₂ and produce oxygen, and methanotrophs (11–20 %) that consume CH₄ and require oxygen. In contrast, cyanobacteria were effectively absent from the surface sediment of runnel ponds, which in addition to carbon degraders (65–81 %), were colonized by purple non-sulfur bacteria (5–21 %), and by fewer methanotrophs (1–5 %). The link between the methanotrophic community and the type of ponds could potentially be used to improve upscale estimates of GHG emissions based on landscape morphology in such remote regions.     

Persistence at distributional edges: Columbia spotted frog habitat in the arid Great Basin,USA

A common challenge in the conservation of broadly distributed, yet imperiled species is understanding which factors facilitate persistence at distributional edges, locations where populations are often vulnerable to extirpation due to changes in climate, land use, or distributions of other species. For Columbia spotted frogs (Rana luteiventris) in the Great Basin (USA), a genetically distinct population segment of conservation concern, we approached this problem by examining (1) landscape‐scale habitat availability and distribution, (2) water body‐scale habitat associations, and (3) resource management‐identified threats to persistence. We found that areas with perennial aquatic habitat and suitable climate are extremely limited in the southern portion of the species’ range. Within these suitable areas, native and non‐native predators (trout and American bullfrogs [Lithobates catesbeianus]) are widespread and may further limit habitat availability in upper‐ and lower‐elevation areas, respectively. At the water body scale, spotted frog occupancy was associated with deeper sites containing abundant emergent vegetation and nontrout fish species. Streams with American beaver (Castor canadensis) frequently had these structural characteristics and were significantly more likely to be occupied than ponds, lakes, streams without beaver, or streams with inactive beaver ponds, highlighting the importance of active manipulation of stream environments by beaver. Native and non‐native trout reduced the likelihood of spotted frog occupancy, especially where emergent vegetation cover was sparse. Intensive livestock grazing, low aquatic connectivity, and ephemeral hydroperiods were also negatively associated with spotted frog occupancy. We conclude that persistence of this species at the arid end of its range has been largely facilitated by habitat stability (i.e., permanent hydroperiod), connectivity, predator‐free refugia, and a commensalistic interaction with an ecosystem engineer. Beaver‐induced changes to habitat quality, stability, and connectivity may increase spotted frog population resistance and resilience to seasonal drought, grazing, non‐native predators, and climate change, factors which threaten local or regional persistence.     

Ecological impact of Eurasian beaver (<Emphasis Type="Italic">Castor fiber</Emphasis>) activity on macroinvertebrate communities in Lithuanian trout streams

Our study found that beaver activity affects macroinvertebrate assemblages of both beaver ponds and downstream sites. The percentage composition of the invertebrate faunae of beaver ponds was strikingly different from the invertebrate faunae of upstream forested and downstream sites. The number of EPT (ephemeropteran, plecopteran, trichopteran) taxa in the upstream forested sites in all streams was higher than in beaver pond and downstream sites. Statistically significant differences were found in absolute and relative abundances of EPT and Chironomidae between different streams sites. The absolute and relative abundance of pollution-sensitive EPT was significantly higher in forested sites than in beaver pond and downstream sites in all measured streams. Beaver ponds had a significantly higher absolute and relative abundance of Chironomidae compared with upstream forested and downstream sites. We found that Plecoptera and Coleoptera were absent from beaver pond sites. The absolute abundance of Plecoptera was significantly higher in upstream forested sites than in downstream sites in all three streams. Gatherers were the dominant functional feeding group in relative abundance in all three habitat types. The percentage of gatherers was higher in beaver ponds than in forested and downstream sites.     

Cumulative geoecological effects of 62 years of infrastructure and climate change in ice‐rich permafrost landscapes,Prudhoe Bay Oilfield,Alaska

Many areas of the Arctic are simultaneously affected by rapid climate change and rapid industrial development. These areas are likely to increase in number and size as sea ice melts and abundant Arctic natural resources become more accessible. Documenting the changes that have already occurred is essential to inform management approaches to minimize the impacts of future activities. Here, we determine the cumulative geoecological effects of 62 years (1949–2011) of infrastructure‐ and climate‐related changes in the Prudhoe Bay Oilfield, the oldest and most extensive industrial complex in the Arctic, and an area with extensive ice‐rich permafrost that is extraordinarily sensitive to climate change. We demonstrate that thermokarst has recently affected broad areas of the entire region, and that a sudden increase in the area affected began shortly after 1990 corresponding to a rapid rise in regional summer air temperatures and related permafrost temperatures. We also present a conceptual model that describes how infrastructure‐related factors, including road dust and roadside flooding are contributing to more extensive thermokarst in areas adjacent to roads and gravel pads. We mapped the historical infrastructure changes for the Alaska North Slope oilfields for 10 dates from the initial oil discovery in 1968–2011. By 2010, over 34% of the intensively mapped area was affected by oil development. In addition, between 1990 and 2001, coincident with strong atmospheric warming during the 1990s, 19% of the remaining natural landscapes (excluding areas covered by infrastructure, lakes and river floodplains) exhibited expansion of thermokarst features resulting in more abundant small ponds, greater microrelief, more active lakeshore erosion and increased landscape and habitat heterogeneity. This transition to a new geoecological regime will have impacts to wildlife habitat, local residents and industry.     

A spatially explicit analysis to extrapolate carbon fluxes in upland tundra where permafrost is thawing

One of the most important changes in high‐latitude ecosystems in response to climatic warming may be the thawing of permafrost soil. In upland tundra, the thawing of ice‐rich permafrost can create localized surface subsidence called thermokarst, which may change the soil environment and influence ecosystem carbon release and uptake. We established an intermediate scale (a scale in between point chamber measurements and eddy covariance footprint) ecosystem carbon flux study in Alaskan tundra where permafrost thaw and thermokarst development had been occurring for several decades. The main goal of our study was to examine how dynamic ecosystem carbon fluxes [gross primary production (GPP), ecosystem respiration (R_eco), and net ecosystem exchange (NEE)] relate to ecosystem variables that incorporate the structural and edaphic changes that co‐occur with permafrost thaw and thermokarst development. We then examined how these measured ecosystem carbon fluxes responded to upscaling. For both spatially extensive measurements made intermittently during the peak growing season and intensive measurements made over the entire growing season, ecosystem variables including degree of surface subsidence, thaw depth, and aboveground biomass were selected in a mixed model selection procedure as the ‘best’ predictors of GPP, R_eco, and NEE. Variables left out of the model (often as a result of autocorrelation) included soil temperature, moisture, and normalized difference vegetation index. These results suggest that the structural changes (surface subsidence, thaw depth, aboveground biomass) that integrate multiple effects of permafrost thaw can be useful components of models used to estimate ecosystem carbon exchange across thermokarst affected landscapes.     

Effects of beaver impoundments on dissolved organic matter quality and biodegradability in boreal riverine systems

Beaver impoundments modify the structure of river reaches and lead to changes in ecosystem function and biogeochemical processes. Here, we assessed the changes in dissolved organic matter (DOM) quality and the biodegradation patterns in a set of beaver systems across Sweden. As the effect of beaver impoundments might be transient and local, we compared DOM quality and biodegradability of both pond and upstream sections of differentially aged beaver systems. Newly established dams shifted the sources and DOM biodegradability patterns. In particular, humic-like DOM, most likely leached from surrounding soils, characterized upstream sections of new beaver impoundments. In contrast, autochthonous and processed compounds, with both higher biodegradation rates and a broader spectrum of reactivities, differentiated DOM in ponds. DOM in recently established ponds seemed to be more humic and less processed compared to older ponds, but system idiosyncrasies determined by catchment particularities influenced this ageing effect.     

Microbial biogeography of permafrost thaw ponds across the changing northern landscape

Microbial diversity varies at multiple spatial scales, but little is known about how climate change may influence this variation. Here we assessed the free‐living bacterioplankton composition of thaw ponds over a north‐south gradient of permafrost degradation in the eastern Canadian subarctic. Three nested spatial scales were compared: 1) among ponds within individual valleys 2) between two valleys within each landscape type, and 3) between landscape types (southern sporadic versus northern discontinuous permafrost). As a reference point, we sampled rock‐basin lakes whose formation was not related to permafrost thawing. β‐diversity was low at the smallest scale despite marked differences in limnological properties among neighboring ponds. β‐diversity was high among valleys, associated with greater environmental heterogeneity. The largest differences were between landscape types and appeared to reflect the concomitant effects of environmental filtering and dispersal limitation. Raup–Crick β‐diversity indicated that community assembly was driven by both stochastic (random extinction, dispersal, ecological drift) and deterministic (environmental filtering) processes. Communities sampled in the most degraded valley appeared primarily assembled through stochastic processes, while environmental filtering played a greater role at the other valleys. These results imply that climate warming and ongoing permafrost degradation will influence microbial community assembly, which in turn is likely to affect the functioning of thaw pond ecosystems.     

Seasonal and Long-Term Changes to Active-Layer Temperatures after Tall Shrubland Expansion and Succession in Arctic Tundra

Vegetation, active-layer soils, and snow cover regulate energy exchange between the atmosphere and permafrost. Therefore, interactions between changes to tundra vegetation and soil thermal regime will fundamentally affect permafrost in a warmer world. We recorded soil temperatures for approximately 1 year in a Siberian Low Arctic landscape with a known history of alder (Alnus) shrub expansion on disturbed microsites in patterned ground. We recorded near-surface soil temperatures and measured physical properties of soils and vegetation on sorted-circle microsites in four stages of shrubland development: (1) tundra lacking tall shrubs; (2) shrub colonization zones; (3) mature shrublands; and (4) paludified, long-established shrublands with thick soil organic layers. Summer soil temperatures declined with increasing shrub cover and soil organic thickness; shrub colonization suppressed cryoturbation, facilitating the development of continuous vegetation and a surface organic mat on circles. Compared to open tundra, mature shrubs cooled soils by up to 9 °C during summer, but warmed soils by greater than 10 °C in winter presumably because they developed highly insulative snowpacks. Paludified shrublands had the coldest summer active layers, but winter soil temperatures were much colder than mature shrublands and were similar to earlier stages. Our results indicate that although tall shrub establishment dramatically warms winter soils within decades, much of this warming is transient at paludification-prone sites because the buildup of wet peat favors cooling in winter and the stature and snow-trapping capacity of shrubs diminish over time. In the ecosystem we studied, shrub expansion has contrasting effects on active-layer temperatures both seasonally and over longer timescales due to successional processes.     

The biogeochemical variability of Arctic thermokarst ponds is reflected by stochastic and niche-driven microbial community assembly processes

Shallow thermokarst ponds are a conspicuous landscape element of the Arctic Siberian tundra with high biogeochemical variability. Little is known about how microbes from the regional species pool assemble into local pond communities and how the resulting patterns affect functional properties such as dissolved organic carbon (DOC) remineralization and greenhouse gas (GHG) turnover. We analysed the pelagic microbiomes of 20 ponds in north-eastern Siberia in the context of their physico-chemical properties. Ponds were categorized as polygonal or trough according to their geomorphological origin. The diversity of bacteria and eukaryotic microbes was assessed by ribosomal gene tag sequencing. Null model analysis revealed an important role of stochastic assembly processes within ponds of identical origin, in particular for genotypes only occurring in few systems. Nevertheless, the two pond types clearly represented distinct niches for both the bacterial and eukaryotic microbial communities. Carbon dioxide concentration, indicative of heterotrophic microbial processes, varied greatly, especially in the trough ponds. Methane concentrations were lower in polygonal ponds and were correlated with the estimated abundance of methanotrophs. Thus, the overall functional variability of Arctic ponds reflects the stochastic assembly of their microbial communities. Distinct functional subcommunities can, nevertheless, be related to GHG concentrations.     

The Cooling Capacity of Mosses: Controls on Water and Energy Fluxes in a Siberian Tundra Site

Arctic tundra vegetation composition is expected to undergo rapid changes during the coming decades because of changes in climate. Higher air temperatures generally favor growth of deciduous shrubs, often at the cost of moss growth. Mosses are considered to be very important to critical tundra ecosystem processes involved in water and energy exchange, but very little empirical data are available. Here, we studied the effect of experimental moss removal on both understory evapotranspiration and ground heat flux in plots with either a thin or a dense low shrub canopy in a tundra site with continuous permafrost in Northeast Siberia. Understory evapotranspiration increased with removal of the green moss layer, suggesting that most of the understory evapotranspiration originated from the organic soil layer underlying the green moss layer. Ground heat flux partitioning also increased with green moss removal indicating the strong insulating effect of moss. No significant effect of shrub canopy density on understory evapotranspiration was measured, but ground heat flux partitioning was reduced by a denser shrub canopy. In summary, our results show that mosses may exert strong controls on understory water and heat fluxes. Changes in moss or shrub cover may have important consequences for summer permafrost thaw and concomitant soil carbon release in Arctic tundra ecosystems.     

Plant carbon allocation drives turnover of old soil organic matter in permafrost tundra soils

Carbon cycle feedbacks from permafrost ecosystems are expected to accelerate global climate change. Shifts in vegetation productivity and composition in permafrost regions could influence soil organic carbon (SOC) turnover rates via rhizosphere (root zone) priming effects (RPEs), but these processes are not currently accounted for in model predictions. We use a radiocarbon (bomb‐¹⁴C) approach to test for RPEs in two Arctic tall shrubs, alder (Alnus viridis (Chaix) DC.) and birch (Betula glandulosa Michx.), and in ericaceous heath tundra vegetation. We compare surface CO₂ efflux rates and ¹⁴C content between intact vegetation and plots in which below‐ground allocation of recent photosynthate was prevented by trenching and removal of above‐ground biomass. We show, for the first time, that recent photosynthate drives mineralization of older (>50 years old) SOC under birch shrubs and ericaceous heath tundra. By contrast, we find no evidence of RPEs in soils under alder. This is the first direct evidence from permafrost systems that vegetation influences SOC turnover through below‐ground C allocation. The vulnerability of SOC to decomposition in permafrost systems may therefore be directly linked to vegetation change, such that expansion of birch shrubs across the Arctic could increase decomposition of older SOC. Our results suggest that carbon cycle models that do not include RPEs risk underestimating the carbon cycle feedbacks associated with changing conditions in tundra regions.