Plant transport and methane production as controls on methane flux from arctic wet meadow tundra

The roles of plant transport and CH₄ production in controlling CH₄ flux from wet meadow tundra communities were investigated. Plant transport was the dominant pathway of CH₄ flux from this ecosystem. Most CH₄ production (measured within situ anaerobic incubations) occurred well below the water table, and C supply (estimated by anaerobic CO₂ production) was the best single predictor of CH₄ production rates. Plant transport of CH₄ was controlled both by CH₄ supply and the plant species.Eriophorum angustifolium transported substantially more CH₄ than didCarex aquatilis, due to differences in size and structure of the two species. The composition of the plant community was a greater control on CH₄ flux from the site than either water table height (which varied only slightly) or CH₄ production rates, indicating the importance of species-specific plant dynamics in controlling CH₄ flux from arctic wetlands.     

Carbon dioxide and methane exchange of a north-east Siberian tussock tundra

Carbon dioxide, energy flux measurements and methane chamber measurements were carried out in an arctic wet tussock grassland located on a flood plane of the Kolyma river in NE Siberia over a summer period of 155 days in 2002 and early 2003. Respiration was also measured in April 2004. The study region is characterized by late thaw of the top soil (mid of June) and periodic spring floods. A stagnant water table below the grass canopy is fed by thawing of the active layer of permafrost and by flood water. The climate is continental with average daily temperature in the warmest months of 13°C (maximum temperature at midday: 28°C by the end of July), dry air (maximum vapour pressure deficit at midday: 28 hPa) and low rainfall of 50 mm during summer (July–September). Summer evaporation (July–September: 103 mm) exceeded rainfall by a factor of 2. The daily average Bowen ratio (H/LE) was 0.62 during the growing season. Net ecosystem CO₂ uptake reached 10 μmol m⁻² s⁻¹ and was related to photon flux density (PFD) and vapour pressure deficit (VPD). The cumulative annual net carbon flux from the atmosphere to the terrestrial surface was estimated to be about −38 g C m⁻² yr⁻¹ (negative flux depicts net carbon sink). Winter respiration was extrapolated using the Lloyd and Taylor function. The net carbon balance is composed of a high rate of assimilation in a short summer and a fairly large but uncertain respiration mainly during autumn and spring. Methane flux (about 12 g C m⁻² measured over 60 days) was 25% of C uptake during the same period of time (end of July to end of September). Assuming that CH₄ was emitted only in summer, and taking the greenhouse gas warming potential of CH₄ vs. CO₂ into account (factor 23), the study site was a greenhouse gas source (at least 200 g C_equivalent m⁻² yr⁻¹). Comparing different studies in wetlands and tundra ecosystems as related to latitude, we expect that global warming would rather increase than decrease the CO₂-C sink.     

Local snow melt and temperature—but not regional sea ice—explain variation in spring phenology in coastal Arctic tundra

The Arctic is undergoing dramatic environmental change with rapidly rising surface temperatures, accelerating sea ice decline and changing snow regimes, all of which influence tundra plant phenology. Despite these changes, no globally consistent direction of trends in spring phenology has been reported across the Arctic. While spring has advanced at some sites, spring has delayed or not changed at other sites, highlighting substantial unexplained variation. Here, we test the relative importance of local temperatures, local snow melt date and regional spring drop in sea ice extent as controls of variation in spring phenology across different sites and species. Trends in long‐term time series of spring leaf‐out and flowering (average span: 18 years) were highly variable for the 14 tundra species monitored at our four study sites on the Arctic coasts of Alaska, Canada and Greenland, ranging from advances of 10.06 days per decade to delays of 1.67 days per decade. Spring temperatures and the day of spring drop in sea ice extent advanced at all sites (average 1°C per decade and 21 days per decade, respectively), but only those sites with advances in snow melt (average 5 days advance per decade) also had advancing phenology. Variation in spring plant phenology was best explained by snow melt date (mean effect: 0.45 days advance in phenology per day advance snow melt) and, to a lesser extent, by mean spring temperature (mean effect: 2.39 days advance in phenology per °C). In contrast to previous studies examining sea ice and phenology at different spatial scales, regional spring drop in sea ice extent did not predict spring phenology for any species or site in our analysis. Our findings highlight that tundra vegetation responses to global change are more complex than a direct response to warming and emphasize the importance of snow melt as a local driver of tundra spring phenology.     

Isotopic insights into methane production,oxidation, and emissions in Arctic polygon tundra

Arctic wetlands are currently net sources of atmospheric CH₄. Due to their complex biogeochemical controls and high spatial and temporal variability, current net CH₄ emissions and gross CH₄ processes have been difficult to quantify, and their predicted responses to climate change remain uncertain. We investigated CH₄ production, oxidation, and surface emissions in Arctic polygon tundra, across a wet‐to‐dry permafrost degradation gradient from low‐centered (intact) to flat‐ and high‐centered (degraded) polygons. From 3 microtopographic positions (polygon centers, rims, and troughs) along the permafrost degradation gradient, we measured surface CH₄ and CO₂ fluxes, concentrations and stable isotope compositions of CH₄ and DIC at three depths in the soil, and soil moisture and temperature. More degraded sites had lower CH₄ emissions, a different primary methanogenic pathway, and greater CH₄ oxidation than did intact permafrost sites, to a greater degree than soil moisture or temperature could explain. Surface CH₄ flux decreased from 64 nmol m^?2 s^?1 in intact polygons to 7 nmol m^?2 s^?1 in degraded polygons, and stable isotope signatures of CH₄ and DIC showed that acetate cleavage dominated CH₄ production in low‐centered polygons, while CO₂ reduction was the primary pathway in degraded polygons. We see evidence that differences in water flow and vegetation between intact and degraded polygons contributed to these observations. In contrast to many previous studies, these findings document a mechanism whereby permafrost degradation can lead to local decreases in tundra CH₄ emissions.     

北极苔原土壤中可培养细菌的分离及其抗菌活性测定

【目的】北极地区具有高纬度、低温、高辐射等独特的环境条件。北冰洋及周围大面积的陆地区域鲜有人类踪迹,其中微生物数量不可低估。本研究旨在了解北极土壤中的可培养微生物的多样性及其抗菌活性。【方法】对来源于北极黄河站附近的7份不同植物根下苔原土壤进行直接涂布和富集培养后涂布。【结果】共获得细菌菌株721株,对其中608株进行细菌16S rRNA基因序列测定,归属于86个属,229个种,主要分布于变形菌门(Proteobacteria,54.3%)、放线菌门(Actinobacteria,21.2%)、拟杆菌门(Bacteroidetes,12.8%)、厚壁菌门(Firmicutes,10.0%)和奇异球菌门(Deinococcus-Thermus,1.6%)。其中从16S rRNA基因序列同源性推测有22株细菌菌株为潜在新种/属。从分离菌株中筛选出16株可抑制金黄色葡萄球菌(Staphylococcusaureus)或鲍氏不动杆菌(Acinetobacterbaumannii)生长的拮抗菌。【结论】获得了北极土壤地区特有的微生物菌株资源,为进一步筛选拮抗菌的活性物质提供了菌株基础。     

Inter-annual variability of NDVI in response to long-term warming and fertilization in wet sedge and tussock tundra

This study explores the relationship between the normalized difference vegetation index (NDVI) and aboveground plant biomass for tussock tundra vegetation and compares it to a previously established NDVI–biomass relationship for wet sedge tundra vegetation. In addition, we explore inter-annual variation in NDVI in both these contrasting vegetation communities. All measurements were taken across long-term experimental treatments in wet sedge and tussock tundra communities at the Toolik Lake Long Term Ecological Research (LTER) site, in northern Alaska. Over 15 years (for wet sedge tundra) and 14 years (for tussock tundra), N and P were applied in factorial experiments (N, P and N+P), air temperature was increased using greenhouses with and without N+P fertilizer, and light intensity was reduced by 50% using shade cloth. during the peak growing seasons of 2001, 2002, and 2003, NDVI measurements were made in both the wet sedge and tussock tundra experimental treatment plots, creating a 3-year time series of inter-annual variation in NDVI. We found that: (1) across all tussock experimental tundra treatments, NDVI is correlated with aboveground plant biomass (r ²=0.59); (2) NDVI–biomass relationships for tussock and wet sedge tundra communities are community specific, and; (3) NDVI values for tussock tundra communities are typically, but not always, greater than for wet sedge tundra communities across all experimental treatments. We suggest that differences between the response of wet sedge and tussock tundra communities in the same experimental treatments result from the contrasting degree of heterogeneity in species and functional types that characterize each of these Arctic tundra vegetation communities.     

Interannual variability of plant phenology in tussock tundra: modelling interactions of plant productivity, plant phenology, snowmelt and soil thaw

We present a linked model of plant productivity, plant phenology, snowmelt and soil thaw in order to estimate interannual variability of arctic plant phenology and its effects on plant productivity. The model is tested using 8 years of soil temperature data, and three years of bud break data of Betula nana. Because the factors that trigger the end of the growing season of arctic vegetation are less well known than those of the start of the growing season, three hypotheses were formulated and tested for their effects on productivity and its sensitivity to climate change; the hypothesised factors determining the end of the growing season were frost, photoperiod and periodic constraints. The performance of the soil thermal model was good; both the onset of soil thaw in spring and the initiation of freezing in autumn were predicted correctly in most cases. The phenology model predicted the bud break date of Betula nana closely for the three different years. The soil thaw model predicted similar growing season start dates under current climate as the models based on sum of temperatures, but it made significantly different predictions under climate change scenarios, probably because of the non‐linear interactions between snowmelt and soil thaw. The uncertainty about the driving factors for the end of the growing season, in turn, resulted in uncertainty in the interannual variability of the simulated annual gross primary productivity (GPP). The interannual variability ranged from ? 25 to + 26% of the mean annual GPP for the frost hypothesis, from ? 20 to + 20% for the photoperiod hypothesis and only from ? 7 to + 7% for the periodic hypothesis. The different hypotheses also resulted in different sensitivity to climate change, with the frost hypothesis resulting in 30% higher annual GPP values than the periodic hypothesis when air temperatures were increased by 3 °C.     

Seasonal variation in enzyme activities and temperature sensitivities in Arctic tundra soils

Arctic soils contain large amounts of organic matter due to very slow rates of detritus decomposition. The first step in decomposition results from the activity of extracellular enzymes produced by soil microbes. We hypothesized that potential enzyme activities are low relative to the large stocks of organic matter in Arctic tundra soils, and that enzyme activity is low at in situ temperatures. We measured the potential activity of six hydrolytic enzymes at 4 and 20 °C on four sampling dates in tussock, intertussock, shrub organic, and shrub mineral soils at Toolik Lake, Alaska. Potential activities of N‐acetyl glucosaminidase, β‐glucosidase, and peptidase tended to be greatest at the end of winter, suggesting that microbes produced enzymes while soils were frozen. In general, enzyme activities did not increase during the Arctic summer, suggesting that enzyme production is N‐limited during the period when temperatures would otherwise drive higher enzyme activity in situ. We also detected seasonal variations in the temperature sensitivity (Q₁₀) of soil enzymes. In general, soil enzyme pools were more sensitive to temperature at the end of the winter than during the summer. We modeled potential in situβ‐glucosidase activities for tussock and shrub organic soils based on measured enzyme activities, temperature sensitivities, and daily soil temperature data. Modeled in situ enzyme activity in tussock soils increased briefly during the spring, then declined through the summer. In shrub soils, modeled enzyme activities increased through the spring thaw into early August, and then declined through the late summer and into winter. Overall, temperature is the strongest factor driving low in situ enzyme activities in the Arctic. However, enzyme activity was low during the summer, possibly due to N‐limitation of enzyme production, which would constrain enzyme activity during the brief period when temperatures would otherwise drive higher rates of decomposition.     

Spatio-temporal variability and environmental controls of methane fluxes at the forest–tundra ecotone in the Fennoscandian mountains

We report on temporal and spatial variability in net methane (CH₄) fluxes measured during the thaw period of 1999 and 2000 at three study sites along a c. 8° latitudinal gradient in the Fennoscandian mountain range and across the mountain birch‐tundra ecotone. All of the sites studied here were underlain by well‐drained mesic soils. In addition, we conducted warming experiments in the field to simulate future climate change. Our results show significant CH₄ uptake at mesic sites spanning the forest‐tundra ecotone: on average 0.031 and 0.0065 mg CH₄ m^?2 h^?1 during the 1999 and 2000 thaw periods, respectively, in Abisko (Sweden), and 0.019 and 0.032 mg CH₄ m^?2 h^?1 during 2000 in Dovrefjell and Joatka (Norway), respectively. These values were both temporally and spatially highly variable, and multiple regression analysis of data from Abisko showed no consistent relationship with soil‐moisture status and temperature. Also, there was no consistent difference in CH₄ fluxes between forest and tundra plots; our data, therefore, provide no support for the hypothesis that conversion of tundra to mountain birch forest, or vice versa, would result in a systematic change in the magnitude or direction of net CH₄ fluxes in this region. Experimental warming treatments were associated with a 2.4 °C increase in soil temperatures (5 cm depth) in 1999 in Abisko, but no consistent soil warming was noted at any of the three field locations during 2000. In spite of this, there were significant treatment effects, principally early during the thaw period, with increased CH₄ uptake compared with control (ambient) plots. These results suggest that direct effects of air warming on vegetation processes (e.g. transpiration, root exudation and nutrient assimilation) can influence CH₄ fluxes even in predominantly methanotrophic environments. We conclude that net CH₄ oxidation is significant in these cold, mesic soils and could be strengthened in an environmental change scenario involving a combination of (i) an increase in the length of the thaw period and (ii) increased mean temperatures during this period in combination with decreased soil‐moisture content.     

Effects of retrogressive permafrost thaw slumping on sediment chemistry and submerged macrophytes in Arctic tundra lakes

1. Global warming is predicted to cause changes in permafrost cover and stability in the Arctic. Zones of high ion concentration in regions of ice‐rich permafrost are a reservoir of chemicals that can potentially be transferred to fresh waters during thawing. Consequently, input of enriched runoff from the thaw and sediment and vegetation from the landscape could alter lakes by affecting their geochemistry and biological production. 2. Three undisturbed lakes and five lakes disturbed by retrogressive permafrost thaw slumps were sampled during late summer of 2006 to assess the potential effects of thermokarst shoreline slumping on water and sediment chemistry, the underwater light regime, and benthic macrophyte biomass and community structure. 3. Undisturbed lakes had sediments rich in organic material and selected micronutrients, while disturbed lakes had sediments richer in calcium, magnesium and strontium, greater transparency of the water column, and a well‐developed submerged macrophyte community. 4. It is postulated that enriched runoff chemistry may alter nutrient availability at the sediment–water interface and also the degradation of organic material, thus affecting lake transparency and submerged macrophytes. The results suggest that retrogressive permafrost slumping can significantly affect food webs in arctic tundra lakes through an increase in macrophyte biomass and development of a more complex benthic habitat.     

Community structure – Ecosystem function relationships in the Congo Basin methane cycle depend on the physiological scale of function

Belowground ecosystem processes can be highly variable and difficult to predict using microbial community data. Here, we argue that this stems from at least three issues: (a) complex covariance structure of samples (with environmental conditions or spatial proximity) can make distinguishing biotic drivers a challenge; (b) communities can control ecosystem processes through multiple mechanisms, making the identification of these controls a challenge; and (c) ecosystem function assessments can be broad in physiological scale, encapsulating multiple processes with unique microbially mediated controls. We test these assertions using methane (CH₄)‐cycling processes in soil samples collected along a wetland‐to‐upland habitat gradient in the Congo Basin. We perform our measurements of function under controlled laboratory conditions and statistically control for environmental covariates to aid in identifying biotic drivers. We divide measurements of microbial communities into four attributes (abundance, activity, composition, and diversity) that represent different forms of community control. Lastly, our process measurements differ in physiological scale, including broader processes (gross methanogenesis and methanotrophy) that involve more mediating groups, to finer processes (hydrogenotrophic methanogenesis and high‐affinity CH₄ oxidation) with fewer mediating groups. We observed that finer scale processes can be more readily predicted from microbial community structure than broader scale processes. In addition, the nature of those relationships differed, with broad processes limited by abundance while fine‐scale processes were associated with diversity and composition. These findings demonstrate the importance of carefully defining the physiological scale of ecosystem function and performing community measurements that represent the range of possible controls on ecosystem processes.     

Hydrologic regulation of gross methyl chloride and methyl bromide uptake from Alaskan Arctic tundra

The Arctic tundra has been shown to be a potentially significant regional sink for methyl chloride (CH₃Cl) and methyl bromide (CH₃Br), although prior field studies were spatially and temporally limited, and did not include gross flux measurements. Here we compare net and gross CH₃Cl and CH₃Br fluxes in the northern coastal plain and continental interior. As expected, both regions were net sinks for CH₃Cl and CH₃Br. Gross uptake rates (−793 nmol CH₃Cl m⁻² day⁻¹ and −20.3 nmol CH₃Br m⁻² day⁻¹) were 20–240% greater than net fluxes, suggesting that the Arctic is an even greater sink than previously believed. Hydrology was the principal regulator of methyl halide flux, with an overall trend towards increasing methyl halide uptake with decreasing soil moisture. Water table depth was one of the best predictors of net and gross uptake, with uptake increasing proportionately with water table depth. In drier areas, gross uptake was very high, averaging −1201 nmol CH₃Cl m⁻² day⁻¹ and −34.9 nmol CH₃Br m⁻² day⁻¹; in flooded areas, gross uptake was significantly lower, averaging −61 nmol CH₃Cl m⁻² day⁻¹ and −2.3 nmol CH₃Br m⁻² day⁻¹. Net and gross uptake was greater in the continental interior than in the northern coastal plain, presumably due to drier inland conditions. Within certain microtopographic features (low‐ and high‐centered polygons), uptake rates were positively correlated with soil temperature, indicating that temperature played a secondary role in methyl halide uptake. Incubations suggested that the inverse relationship between water content and methyl halide uptake was the result of mass transfer limitation in saturated soils, rather than because of reduced microbial activity under anaerobic conditions. These findings have potential regional significance, as the Arctic is expected to become warmer and drier due to anthropogenic climate forcing, potentially enhancing the Arctic sink for CH₃Cl and CH₃Br.     

The processes of methane production and oxidation in the soils of the Russian Arctic tundra

Methane emission from the following types of tundra soils was studied: coarse humic gleyey loamy cryo soil, peaty gleyey soil, and peaty gleyey midloamy cryo soil of the arctic tundra. All the soils studied were found to be potential sources of atmospheric methane. The highest values of methane emission were recorded in August at a soil temperature of 8–10°C. Flooded parcels were the sources of atmospheric methane throughout the observation period. The rates of methane production and oxidation in tundra soils of various types were studied by the radioisotope method at 5 and 15°C. Methane oxidation was found to occur in bog water, in the green part of peat moss, and in all the soil horizons studied. Methane production was recorded in the horizons of peat, in clay with plant roots, and in peaty moss dust of the bogey parcels. At both temperatures, the methane oxidation rate exceeded the rate of methane production in all the horizons of the mossy-lichen tundra and of the hillock tundra with flat-bottom depressions. Methanogenesis prevailed only in a sedge-peat moss bog at 15°C. Bacterial enrichment cultures oxidizing methane at 5 and 15°C were obtained. Different types of methanotrophic bacteria were shown to be responsible for methane oxidation under these conditions. A representative of type I methylotrophs oxidized methane at 5°C, and Methylocella tundrae, a psychroactive representative of an acidophilic methanotrophic genus Methylocella, at 15°C.__________Translated from Mikrobiologiya, Vol. 74, No. 2, 2005, pp. 261–270.Original Russian Text Copyright © 2005 by Berestovskaya, Rusanov, Vasileva, Pimenov.     

Growth form and leaf habit drive contrasting effects of Arctic amplification in long-lived woody species

Current global change is inducing heterogeneous warming trends worldwide, with faster rates at higher latitudes in the Northern Hemisphere. Consequently, tundra vegetation is experiencing an increase in growth rate and uneven but expanding distribution. Yet, the drivers of this heterogeneity in woody species responses are still unclear. Here, applying a retrospective approach and focusing on long-term responses, we aim to get insight into growth trends and climate sensitivity of long-lived woody species belonging to different functional types with contrasting growth forms and leaf habits (shrub vs. tree and deciduous vs. evergreen). A total of 530 samples from 7 species (common juniper, dwarf birch, woolly willow, Norway spruce, lodgepole pine, rowan, and downy birch) were collected in 10 sites across Iceland. We modelled growth trends and contrasted yearly ring-width measurements, filtering in high- and low-frequency components, with precipitation, land- and sea-surface temperature records (1967–2018). Shrubs and trees showed divergent growth trends, with shrubs closely tracking the recent warming, whereas trees, especially broadleaved, showed strong fluctuations but no long-term growth trends. Secondary growth, particularly the high-frequency component, was positively correlated with summer temperatures for most of the species. On the contrary, growth responses to sea surface temperature, especially in the low frequency, were highly diverging between growth forms, with a strong positive association for shrubs and a negative for trees. Within comparable vegetation assemblage, long-lived woody species could show contrasting responses to similar climatic conditions. Given the predominant role of oceanic masses in shaping climate patterns in the Arctic and Low Arctic, further investigations are needed to deepen the knowledge on the complex interplay between coastal tundra ecosystems and land-sea surface temperature dynamics.     

Methanogenic activity and biomass in Holocene permafrost deposits of the Lena Delta, Siberian Arctic and its implication for the global methane budget

Permafrost environments within the Siberian Arctic are natural sources of the climate relevant trace gas methane. In order to improve our understanding of the present and future carbon dynamics in high latitudes, we studied the methane concentration, the quantity and quality of organic matter, and the activity and biomass of the methanogenic community in permafrost deposits. For these investigations a permafrost core of Holocene age was drilled in the Lena Delta (72°22′N, 126°28′E). The organic carbon of the permafrost sediments varied between 0.6% and 4.9% and was characterized by an increasing humification index with permafrost depth. A high CH₄ concentration was found in the upper 4 m of the deposits, which correlates well with the methanogenic activity and archaeal biomass (expressed as PLEL concentration). Even the incubation of core material at −3 and −6°C with and without substrates showed a significant CH₄ production (range: 0.04–0.78 nmol CH₄ h⁻¹ g⁻¹). The results indicated that the methane in Holocene permafrost deposits of the Lena Delta originated from modern methanogenesis by cold‐adapted methanogenic archaea. Microbial generated methane in permafrost sediments is so far an underestimated factor for the future climate development.     

The palatability of Arctic willow for greater snow geese: the role of nutrients and deterring factors

Temporary feeding on willow buds and leaves by nesting greater snow geese provided us with an opportunity to test the relative importance of nutrients and deterrents in affecting the palatability for geese of a food plant with a high phenol content. Protein, total phenol and fiber (neutral and acid detergent fiber, and lignin) were analyzed in closed and open buds and in rolled and open leaves. Geese feed on willows at the open-buds and rolled-leaf stages but not at the closed-bud and open-leaf stages. Protein content was higher in open buds and rolled leaves (25–27%) than in closed buds and open leaves (19–21%). Phenol content increased during leaf emergence but was already high (14%) in rolled leaves. All plant fibers were very high in closed buds but declined rapidly during leaf emergence. The increase in phenol: protein ratio appeared to be more important than phenol concentration alone in explaining the cessation of feeding by geese on willow leaves whereas the high fiber content of closed buds may explain why they were not eaten. Our results illustrate the value of a multifactorial approach in the study of the food selection process in herbivores.     

Contrasting effects of warming and increased snowfall on Arctic tundra plant phenology over the past two decades

Recent changes in climate have led to significant shifts in phenology, with many studies demonstrating advanced phenology in response to warming temperatures. The rate of temperature change is especially high in the Arctic, but this is also where we have relatively little data on phenological changes and the processes driving these changes. In order to understand how Arctic plant species are likely to respond to future changes in climate, we monitored flowering phenology in response to both experimental and ambient warming for four widespread species in two habitat types over 21 years. We additionally used long‐term environmental records to disentangle the effects of temperature increase and changes in snowmelt date on phenological patterns. While flowering occurred earlier in response to experimental warming, plants in unmanipulated plots showed no change or a delay in flowering over the 21‐year period, despite more than 1 °C of ambient warming during that time. This counterintuitive result was likely due to significantly delayed snowmelt over the study period (0.05–0.2 days/yr) due to increased winter snowfall. The timing of snowmelt was a strong driver of flowering phenology for all species – especially for early‐flowering species – while spring temperature was significantly related to flowering time only for later‐flowering species. Despite significantly delayed flowering phenology, the timing of seed maturation showed no significant change over time, suggesting that warmer temperatures may promote more rapid seed development. The results of this study highlight the importance of understanding the specific environmental cues that drive species’ phenological responses as well as the complex interactions between temperature and precipitation when forecasting phenology over the coming decades. As demonstrated here, the effects of altered snowmelt patterns can counter the effects of warmer temperatures, even to the point of generating phenological responses opposite to those predicted by warming alone.     

Bryoflora of the Alpine Tundra of Changbai Mountain and Its Floristic Relationship to the Bryofloras of Arctic Tundras

The alpine tundra of Changbai Mountain is situa,ed in 41º53′–42º04′N, 127º57′–128º11′E and at elevations ranging from 1950(2000) –2749.2 m. There are 67 genera and 135 species of bryophytes in the alpine tundra, belonging to following five distribution types (DT): Cosmo-politan DT (50 genera, 29 species), North Temperate DT (14, 79), Eastern Asian-North American Discontinuous DT (1, 1), Eurasian Temperate DT (1, 10), Eastern Asian DT (1, 16). The alpine tundra of Changbai Mountain shares 55 genera and 103 species of bryophytes with arctic tundras.     

Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.