Decomposition of mountain birch leaf litter at the forest-tundra ecotone in the Fennoscandian mountains in relation to climate and soil conditions

Litter decomposition is a key process in terrestrial ecosystems, releasing nutrients, returning CO₂ to the atmosphere, and contributing to the formation of humus. Litter decomposition is strongly controlled both by climate and by litter quality: global warming scenarios involving shifts in vegetation communities are therefore of particular interest in this context. The objective of the present study was to quantify the role of climatic environment and underlying substrate chemistry for the decomposition of standard mountain birch (Betula pubescens Ehrh. spp. czerepanovii) leaf litter at four sites, spanning the forest-tundra ecotone, in the Fennoscandian mountain range. Litter quality effects were thus held constant, but the study incorporated systematic changes in (i) latitude/altitude, (ii) `continentality', and (iii) vegetation community at each site, together with (iv) experimental manipulation of temperature using passive warming systems. The study was undertaken during a 3 year period, and forms part of a broader investigation of forest-tundra ecotone dynamics in the Fennoscandian mountains. Our results showed (1) higher decomposition rates in forest sites compared to tundra, (2) that the difference between the two vegetation communities was most pronounced at the more maritime sites, and (3) that chemistry of litter remaining after the three years experiment varied according to site and vegetation community (e.g. at the most southerly site, more lignin had decomposed at tundra communities compared with the forest). (4) Surface temperature explained 58% of the variation in mass loss at forest sites; at tundra sites, however, we hypothesise that litter moisture content was the more important factor. (5) Experimental warming lent weight to this hypothesis by reducing rates of mass loss: this reduction was likely the result of surface soil drying, an artefact of the warming treatment. We conclude that a replacement of tundra by forest would likely accelerate litter decomposition both via changes in surface and near-surface temperature and moisture regimes, although the strength of this response will vary between maritime and continental parts of the mountain range.     

Landscape patterns of litter decomposition in alpine tundra

A two-year study of the decomposition of alpine avens (Acomastylis rossii) foliage in alpine tundra of the Front Range of Colorado demonstrated a strong landscape-mediated effect on decay rates. Litter on sites with intermediate amounts of snowpack decayed more rapidly than litter on sites with larger or smaller amounts of snow. Annual decay constants (k-values) of this foliage ranged from-0.33 in dry tundra to-0.52 in moist tundra to-0.47 in the wettest habitat. No site differences in mass loss of litter were detected until late winter-early spring of the first year of decomposition, when significantly faster decomposition was observed for litter beneath the snowpack. In spite of obvious landscape-related patterns in rates of litter decomposition, total microarthropod densities in the top 5 cm of soil did not differ among habitats. However, the relative abundance of the oribatid and prostigmatid mites did vary significantly across the landscape in relation to the moisture gradient. Oribatid mites comprised a greater proportion of total mites in wetter areas. Microarthropod densities and composition, as well as patterns of decomposition, were compared with previous alpine, as well as arctic tundra, studies. The effects of soil invertebrates on decomposition rates in the alpine were evaluated with a mushroom litterbag decomposition experiment. Naphthalene was used to exclude fauna from a subset of litterbags placed in mesic and xeric habitats. Mushrooms without naphthalene additions decayed significantly faster in the mesic sites. Densities of invertebrates were also greater on mushrooms in these mesic sites. Mushrooms placed in xeric sites generally lacked fauna. Thus, both the activities and the composition of the detritus-based food web appear to change substantially across the moisture gradient found in alpine tundra.     

Initial Stages of Tundra Shrub Litter Decomposition May Be Accelerated by Deeper Winter Snow But Slowed Down by Spring Warming

The Arctic climate is projected to change during the coming century, with expected higher air temperatures and increased winter snowfall. These climatic changes might alter litter decomposition rates, which in turn could affect carbon (C) and nitrogen (N) cycling rates in tundra ecosystems. However, little is known of seasonal climate change effects on plant litter decomposition rates and N dynamics, hampering predictions of future arctic vegetation composition and the tundra C balance. We tested the effects of snow addition (snow fences), warming (open top chambers), and shrub removal (clipping), using a full-factorial experiment, on mass loss and N dynamics of two shrub tissue types with contrasting quality: deciduous shrub leaf litter (Salix glauca) and evergreen shrub shoots (Cassiope tetragona). We performed a 10.5-month decomposition experiment in a low-arctic shrub tundra heath in West-Greenland. Field incubations started in late fall, with harvests made after 249, 273, and 319 days of field incubation during early spring, summer and fall of the next year, respectively. We observed a positive effect of deeper snow on winter mass loss which is considered a result of observed higher soil winter temperatures and corresponding increased winter microbial litter decomposition in deep-snow plots. In contrast, warming reduced litter mass loss during spring, possibly because the dry spring conditions might have dried out the litter layer and thereby limited microbial litter decomposition. Shrub removal had a small positive effect on litter mass loss for C. tetragona during summer, but not for S. glauca. Nitrogen dynamics in decomposing leaves and shoots were not affected by the treatments but did show differences in temporal patterns between tissue types: there was a net immobilization of N by C. tetragona shoots after the winter incubation, while S. glauca leaf N-pools were unaltered over time. Our results support the widely hypothesized positive linkage between winter snow depth and litter decomposition rates in tundra ecosystems, but our results do not reveal changes in N dynamics during initial decomposition stages. Our study also shows contrasting impacts of spring warming and snow addition on shrub decomposition rates that might have important consequences for plant community composition and vegetation-climate feedbacks in rapidly changing tundra ecosystems.     

Winter regulation of tundra litter carbon and nitrogen dynamics

Mass and nitrogen (N) dynamics of leaf litter measured in Alaskan tussock tundra differed greatly from measurements of these processes made in temperate ecosystems. Nearly all litter mass and N loss occurred during the winter when soils were mostly frozen. Litter lost mass during the first summer, but during the subsequent two summers when biological activity was presumably higher than it is during winter, litter mass remained constant and litter immobilized N. By contrast, litter lost significant mass and N over both winters of measurement. Mass loss and N dynamics were unaffected by microsite variation in soil temperature and moisture. Whether wintertime mass and N loss resulted from biological activity during winter or from physical processes (e.g., fragmentation or leaching) associated with freeze-thaw is unknown, but has implications for how future climate warming will alter carbon (C) and N cycling in tundra. We hypothesize that spring runoff over permafrost as soils melt results in significant losses of C and N from litter, consistent with the observed influx of terrestrial organic matter to tundra lakes and streams after snow melt and the strong N limitation of terrestrial primary production.     

Vegetation responses in Alaskan arctic tundra after 8 years of a summer warming and winter snow manipulation experiment

We used snow fences and small (1 m²) open‐topped fiberglass chambers (OTCs) to study the effects of changes in winter snow cover and summer air temperatures on arctic tundra. In 1994, two 60 m long, 2.8 m high snow fences, one in moist and the other in dry tundra, were erected at Toolik Lake, Alaska. OTCs paired with unwarmed plots, were placed along each experimental snow gradient and in control areas adjacent to the snowdrifts. After 8 years, the vegetation of the two sites, including that in control plots, had changed significantly. At both sites, the cover of shrubs, live vegetation, and litter, together with canopy height, had all increased, while lichen cover and diversity had decreased. At the moist site, bryophytes decreased in cover, while an increase in graminoids was almost entirely because of the response of the sedge Eriophorum vaginatum. These community changes were consistent with results found in studies of responses to warming and increased nutrient availability in the Arctic. However, during the time period of the experiment, summer temperature did not increase, but summer precipitation increased by 28%. The snow addition treatment affected species abundance, canopy height, and diversity, whereas the summer warming treatment had few measurable effects on vegetation. The interannual temperature fluctuation was considerably larger than the temperature increases within OTCs (<2°C), however. Snow addition also had a greater effect on microclimate by insulating vegetation from winter wind and temperature extremes, modifying winter soil temperatures, and increasing spring run‐off. Most increases in shrub cover and canopy height occurred in the medium snow‐depth zone (0.5–2 m) of the moist site, and the medium to deep snow‐depth zone (2–3 m) of the dry site. At the moist tundra site, deciduous shrubs, particularly Betula nana, increased in cover, while evergreen shrubs decreased. These differential responses were likely because of the larger production to biomass ratio in deciduous shrubs, combined with their more flexible growth response under changing environmental conditions. At the dry site, where deciduous shrubs were a minor part of the vegetation, evergreen shrubs increased in both cover and canopy height. These changes in abundance of functional groups are expected to affect most ecological processes, particularly the rate of litter decomposition, nutrient cycling, and both soil carbon and nitrogen pools. Also, changes in canopy structure, associated with increases in shrub abundance, are expected to alter the summer energy balance by increasing net radiation and evapotranspiration, thus altering soil moisture regimes.     

Effects of microclimate,litter type,and mesh size on leaf litter decomposition along an elevation gradient in the Wuyi Mountains,China

Litter decomposition is an important ecosystem process regulated by both biotic factors (e.g., decomposers and litter types) and abiotic factors (e.g., temperature and moisture). This study examined the regulatory effects of soil fauna and microclimate on decomposition of two substrates (Castanopsis carlesii and Pinus taiwanensis) along an elevation gradient in four ecosystems of zonal vegetation types in southeastern China: evergreen broadleaf forest (EVB), coniferous forest (COF), dwarf forest (DWF), and alpine meadow (ALM). Our objective was to identify the mechanisms by which microclimate, substrate, and fauna control litter decomposition, especially where variations in ecosystem structure and environment are markedly shown across an elevation gradient. The hypotheses were as follows: (1) litter decomposition within the same litter type would decrease across the elevation gradient, (2) litter decomposition would be lower in poorer nutrient quality substrate across the four sites, and (3) litter dynamics, influenced by strong interactions among ecosystem type, litter type, and decomposers, would vary by elevation gradient due to microclimate effects (i.e., temperature and moisture). The decomposition rates of C. carlesii were significantly higher than those of P. taiwanensis at EVB, COF, and DWF sites; however, they were not significantly different at the ALM site. Low elevation forests possessed a microclimate (warm and humid) that favors decomposer activities and also appeared to possess a decomposer community adapted to consuming large amounts of leaf litter, as indicated by the rapid leaf litter loss. Litter decomposition in micro-mesh bags proceeded more slowly compared to litter in meso-mesh and macro-mesh litterbags across the elevation gradient, indicating that restricting some detritivore access to litter reduced litter mass loss. We suggest that microclimate and faunal contributions to plant litter decomposition differ markedly across the ecosystems in the Wuyi Mountains.     

湿地枯落物分解及其对全球变化的响应

综述了当前湿地枯落物分解及其对全球变化响应的研究动态。湿地枯落物分解研究已随研究方法的改进而不断深化;当前湿地枯落物分解过程研究主要集中在有机质组分和元素含量变化特征的探讨上;湿地枯落物分解同时受生物因素（即枯落物性质以及参与分解的异养微生物和土壤动物的种类、数量和活性等）和非生物因素（即枯落物分解过程的外部环境条件,包括气候条件、水分条件、酸碱度与盐分条件以及湿地沉积的行为与特征等）的制约;模型已成为湿地枯落物分解研究的重要手段,对其研究也在不断深化。还讨论了湿地枯落物分解对于全球变化的响应,指出全球变暖、大气CO2浓度上升、干湿沉降及其化学组成改变可能对枯落物分解产生的直接、间接和综合影响。最后,指出了当前该领域研究尚存在的问题以及今后亟需加强的几个研究方面。     

Responses of High Arctic wet sedge tundra to climate warming since 1980

The global climate is changing rapidly and Arctic regions are showing responses to recent warming. Responses of tundra ecosystems to climate change have been examined primarily through short‐term experimental manipulations, with few studies of long‐term ambient change. We investigated changes in above‐ and belowground biomass of wet sedge tundra to the warming climate of the Canadian High Arctic over the past 25 years. Aboveground standing crop was harvested from five sedge meadow sites and belowground biomass was sampled from one of the sites in the early 1980s and in 2005 using the same methods. Aboveground biomass was on average 158% greater in 2005 than in the early 1980s. The belowground biomass was also much greater in 2005: root biomass increased by 67% and rhizome biomass by 139% since the early 1980s. Dominant species from each functional group (graminoids, shrubs and forbs) showed significant increases in aboveground biomass. Responsive species included the dominant sedge species Carex aquatilis stans, C. membranacea, and Eriophorum angustifolium, as well as the dwarf shrub Salix arctica and the forb Polygonum viviparum. However, diversity measures were not different between the sample years. The greater biomass correlated strongly with increased annual and summer temperatures over the same time period, and was significantly greater than the annual variation in biomass measured in 1980–1983. Increased decomposition and mineralization rates, stimulated by warmer soils, were likely a major cause of the elevated productivity, as no differences in the mass of litter were found between sample periods. Our results are corroborated by published short‐term experimental studies, conducted in other wet sedge tundra communities which link warming and fertilization with elevated decomposition, mineralization and tundra productivity. We believe that this is the first study to show responses in High Arctic wet sedge tundra to recent climate change.     

雪被斑块对高山森林两种灌木凋落叶质量损失的影响

高山/亚高山森林灌木层植物凋落物的分解对于系统物质循环等过程具有重要意义, 并可能受到冬季不同厚度雪被斑块下冻融格局的影响。该文采用凋落物分解袋法, 研究了高山森林典型灌层植物华西箭竹(Fargesia nitida)和康定柳(Salix paraplesia)凋落物在沿林窗-林下形成的冬季雪被厚度梯度(厚型雪被斑块、较厚型雪被斑块、中型雪被斑块、薄型雪被斑块、无雪被斑块)上在第一年不同关键时期(冻结初期、冻结期、融化期、生长季节初期和生长季节后期)的质量损失特征。在整个冻融季节, 华西箭竹和康定柳凋落叶的平均质量损失分别占全年的(48.78 ± 2.35)%和(46.60 ± 5.02)%。冻融季节雪被覆盖斑块下凋落叶的失重率表现出厚型雪被斑块大于薄型雪被斑块的趋势,而生长季节无雪被斑块的失重率明显较高。尽管如此, 华西箭竹凋落物第一年分解表现出随冬季雪被厚度增加而增加的趋势, 但康定柳凋落物第一年失重率以薄型雪被斑块最高, 而无雪被斑块最低。同时, 相关分析表明冻融季节凋落叶的失重率与平均温度和负积温呈极显著正相关, 生长季节凋落叶的失重率与所调查的温度因子并无显著相关关系, 但全年凋落物失重率与平均温度和正/负积温均显著相关。这些结果清晰地表明, 未来冬季变暖情境下高山森林冬季雪被格局的改变将显著影响灌层植物凋落物分解, 影响趋势随着物种的差异具有明显差异。     

Experimental icing affects growth,mortality, and flowering in a high Arctic dwarf shrub

Effects of climate change are predicted to be greatest at high latitudes, with more pronounced warming in winter than summer. Extreme mid‐winter warm spells and heavy rain‐on‐snow events are already increasing in frequency in the Arctic, with implications for snow‐pack and ground‐ice formation. These may in turn affect key components of Arctic ecosystems. However, the fitness consequences of extreme winter weather events for tundra plants are not well understood, especially in the high Arctic. We simulated an extreme mid‐winter rain‐on‐snow event at a field site in high Arctic Svalbard (78°N) by experimentally encasing tundra vegetation in ice. After the subsequent growing season, we measured the effects of icing on growth and fitness indices in the common tundra plant, Arctic bell‐heather (Cassiope tetragona). The suitability of this species for retrospective growth analysis enabled us to compare shoot growth in pre and postmanipulation years in icing treatment and control plants, as well as shoot survival and flowering. Plants from icing treatment plots had higher shoot mortality and lower flowering success than controls. At the individual sample level, heavily flowering plants invested less in shoot growth than nonflowering plants, while shoot growth was positively related to the degree of shoot mortality. Therefore, contrary to expectation, undamaged shoots showed enhanced growth in ice treatment plants. This suggests that following damage, aboveground resources were allocated to the few remaining undamaged meristems. The enhanced shoot growth measured in our icing treatment plants has implications for climate studies based on retrospective analyses of Cassiope. As shoot growth in this species responds positively to summer warming, it also highlights a potentially complex interaction between summer and winter conditions. By documenting strong effects of icing on growth and reproduction of a widespread tundra plant, our study contributes to an understanding of Arctic plant responses to projected changes in winter climatic conditions.     

Decomposition of Senesced Leaf Litter is Faster in Tall Compared to Low Birch Shrub Tundra

Many Low Arctic tundra regions are currently undergoing a vegetation shift towards increasing growth and groundcover of tall deciduous shrubs due to recent climate warming. Vegetation change directly affects ecosystem carbon balance, but it can also affect soil biogeochemical cycling through physical and biological feedback mechanisms. Recent studies indicate that enhanced snow accumulation around relatively tall shrubs has negligible physical effect on litter decomposition rates. However, these investigations were no more than 3 years, and therefore may be insufficient to detect differences in inherently slow biogeochemical processes. Here, we report a 5-year study near Daring Lake, Canada, comparing Betula neoalaskana foliar litter decay rates within unmanipulated and snowfenced low-stature birch (height: ~?0.3 m) plots to test the physical effect of experimentally deepened snow, and within tall birch (height: ~?0.8 m) plots to test the combined physical and biological effects, that is, deepened snow plus strong birch dominance. Having corrected for carbon gain by the colonizing decomposers, actual litter carbon loss increased by approximately 25% in the tall birch relative to both low birch sites. Decay of lignin-like acid unhydrolizable litter residues also accelerated in the tall birch site, and a similar but lower magnitude response in the snowfenced low birch site indicated that physical effects of deepened snow were at least partially responsible. In contrast, deepened snow alone did not affect litter carbon loss. Our findings suggest that a combination of greater litter inputs, altered soil microbial community, enhanced soil nutrient pools, and warmer winter soils together promote relatively fast decomposition of recalcitrant litter carbon in tall birch shrub environments.     

Long‐term experimental warming alters community composition of ascomycetes in Alaskan moist and dry arctic tundra

Arctic tundra regions have been responding to global warming with visible changes in plant community composition, including expansion of shrubs and declines in lichens and bryophytes. Even though it is well known that the majority of arctic plants are associated with their symbiotic fungi, how fungal community composition will be different with climate warming remains largely unknown. In this study, we addressed the effects of long‐term (18 years) experimental warming on the community composition and taxonomic richness of soil ascomycetes in dry and moist tundra types. Using deep Ion Torrent sequencing, we quantified how OTU assemblage and richness of different orders of Ascomycota changed in response to summer warming. Experimental warming significantly altered ascomycete communities with stronger responses observed in the moist tundra compared with dry tundra. The proportion of several lichenized and moss‐associated fungi decreased with warming, while the proportion of several plant and insect pathogens and saprotrophic species was higher in the warming treatment. The observed alterations in both taxonomic and ecological groups of ascomycetes are discussed in relation to previously reported warming‐induced shifts in arctic plant communities, including decline in lichens and bryophytes and increase in coverage and biomass of shrubs.     

Biotic and Abiotic Constraints on the Decomposition of Fagus sylvatica Leaf Litter Along an Altitudinal Gradient in Contrasting Land-Use Types

Climate change can affect the process of carbon cycling and leaf litter decomposition in multiple ways, both directly and indirectly, though the strength and direction of this relationship is often context dependent. In this experiment, we followed decomposition of a standard litter type—senescent leaves of Fagus sylvatica collected from a single location—along a 1000 m altitudinal gradient of four sites over 2.5 years. To control the edaphic conditions, we transplanted intact turf mesocosms from three different land-use types [densely wooded, sparsely wooded, and unwooded (UW) pastures] from the highest altitude site into UW pastures along the altitudinal gradient from the moist, cool high-elevation site to the dry, warm low-elevation site, using shade cloth to mimic the light conditions in the original habitats. Decomposition in the drier UW pasture mesocosms increased with altitude, likely because of higher moisture at the highest sites. Decomposition in the more mesic mesocosms from sparsely and densely wooded sites was insensitive to altitude, suggesting an overriding moisture, rather than temperature, constraint on decomposition across these sites. The functional composition of decomposer microbial communities (fungal/bacterial ratio) was similarly insensitive to altitude. Our findings bring substantial evidence for the controlling role of soil moisture on litter decomposition, as well as for the indirect effects of climate through changes in the decomposer community.     

凋落物多样性及组成对凋落物分解和土壤微生物群落的影响

凋落物分解是生态系统营养物质循环的核心过程,而土壤微生物群落在凋落物分解过程中扮演着极其重要且不可替代的角色。随着生物多样性的丧失日益严峻,探讨凋落物多样性及组成对凋落物分解和土壤微生物群落的影响,不仅有助于了解凋落物分解的内在机制,而且可为退化草原生态系统的恢复提供参考。以内蒙古呼伦贝尔草原退化恢复群落中的草本植物为研究对象,依据植物多度、盖度、频度和物种的重要值及其在群落中的恢复程度筛选出排序前4的羊草（Leymus chinensis）、茵陈蒿（Artemisia capillaris）、麻花头（Serratula centauroides）、二裂委陵菜（Potentilla bifurca）的凋落物为实验材料,通过设置3种凋落物多样性水平（1,2,4）,包括11种凋落物组合（单物种凋落物共4种,两物种凋落物混合共6种,四物种凋落物混合共1种）,利用磷脂脂肪酸（PLFA）方法来研究分解60 d后凋落物多样性及组成对凋落物分解和土壤微生物群落的影响。结果表明：（1）凋落物物种多样性仅对C残余率具有显著影响,表现在两物种混合凋落物C残余率显著低于单物种凋落物,而凋落物组成对所观测的4个凋落物分解参数（质量、C、N残余率以及C/N）均具有显著影响;（2）凋落物物种多样性对细菌（B）含量具有显著影响,而凋落物组成对真菌（F）含量具有显著影响,两者对F/B以及微生物总量均无显著影响;（3）冗余分析结果表明凋落物组成与凋落物分解相关指标（凋落物质量、C、N残余率及C/N）和土壤微生物（真菌、细菌含量）的相关关系高于凋落物多样性。（4）进一步建立结构方程模型（Structural Equation Model,SEM）发现,凋落物初始C含量对凋落物质量、C、N残余率及C/N有显著正的直接影响;凋落物木质素含量对凋落物质量、C、N残余率有显著正的直接影响;凋落物初始N含量对N残余率有显著正的直接影响,而对C残余率及C/N有显著负的直接影响;凋落物初始C/N对凋落物质量、N残余率有显著正的直接影响,而对C/N有显著负的直接影响。此外,凋落物初始C、N、木质素含量及C/N均对真菌含量具有显著正影响,并可通过真菌对凋落物质量分解产生显著负的间接影响。以上结果表明该退化恢复区域优势种凋落物分解以初始C、木质素为主导,主要通过土壤真菌影响凋落物的分解进程,这将减缓凋落物的分解速率进而减慢草原生态系统的进程。这些结果为凋落物多样性及组成对自身分解和土壤微生物群落的影响提供了实验依据,也为进一步分析凋落物分解内在机制以及草原生态系统的恢复提供了数据参考。     

Acute seasonal drought does not permanently alter mass loss and nitrogen dynamics during decomposition of red maple (Acer rubrum L.) litter

Litter decomposition is a major component of nutrient cycling in forest ecosystems and is known to be sensitive to both temperature and moisture. A drought during periods of the year when moisture is normally abundant may impact decomposition more than moisture limitation during seasons that are routinely dry. We excluded precipitation for a 3‐month period during each of four seasons and measured mass loss and nitrogen (N) dynamics in Acer rubrum leaf litter by using a litterbag approach. We hypothesized that rainfall exclusion would retard mass loss during the exclusion period, with the maximum effect occurring during the spring and summer months when moisture was normally adequate and temperatures optimum for decomposition, and that exclusion during these two periods would have the greatest impact on subsequent timing of decomposition events, particularly N mineralization. Mass loss in litterbags was initially reduced by precipitation exclusion in the spring treatment only. For each season, a set of bags exposed to drying in the same way and allowed to remain in the field under ambient conditions for an additional 9 months no longer showed a difference in mass loss relative to control bags. Nitrogen accumulation was immediately and significantly reduced in the exclusion treatments in all seasons; however, like mass loss, there was no difference 9 months later. We conclude that short‐term effects of drought do occur, especially with regard to N accumulation, and that these effects do not persist and are unlikely to have any substantial effect on nutrient cycling.     

Limited evidence that mixing leaf litter accelerates decomposition or increases diversity of decomposers in streams of eastern Canada

Most studies of terrestrial litter decomposition in streams and rivers have used leaves from a single tree species, but leaf packs in streams in eastern North America are usually mixtures of two or more species. Litter mixtures may decay more quickly than either of the component species. If so, estimates of stream energy and nutrient budgets may be inaccurate. In northern Nova Scotia, Canada, we measured mass loss from binary mixtures (1:1 mass ratio) of leaf litter in mesh bags, using freshly fallen or air-dried litter from five species of canopy trees. We repeated the experiment eight times, in summer and fall, in two streams and a small river, over 3 years. In some trials we enumerated benthic invertebrate and fungal colonization of decaying litter. Although there were marked differences in mass loss rates among litter types, decomposition was accelerated in mixtures relative to the mean of the component species in only three of eight trials, and only in mixtures containing N-rich speckled alder leaves. Mixing yellow birch and red maple leaves inhibited decomposition. Diversity (Shannon–Weaver Index), species richness, and abundance of aquatic hyphomycete fungi, as indexed by conidial production, were never greater (and sometimes less) on litter mixtures than on the component species. Total numbers, taxonomic richness and diversity of benthic invertebrates generally, and litter-feeding species in particular, were not augmented by mixing litter types. Litter mixtures appear to dilute a preferred substrate with patches of a less preferred substrate. Our results provide only weak support for the contention that combining two litter types leads to acceleration of decomposition rates. Handling editor: K. Martens     

Seasonal climate manipulations have only minor effects on litter decomposition rates and N dynamics but strong effects on litter P dynamics of sub-arctic bog species

Litter decomposition and nutrient mineralization in high-latitude peatlands are constrained by low temperatures. So far, little is known about the effects of seasonal components of climate change (higher spring and summer temperatures, more snow which leads to higher winter soil temperatures) on these processes. In a 4-year field experiment, we manipulated these seasonal components in a sub-arctic bog and studied the effects on the decomposition and N and P dynamics of leaf litter of Calamagrostis lapponica, Betula nana, and Rubus chamaemorus, incubated both in a common ambient environment and in the treatment plots. Mass loss in the controls increased in the order Calamagrostis < Betula < Rubus. After 4 years, overall mass loss in the climate-treatment plots was 10 % higher compared to the ambient incubation environment. Litter chemistry showed within each incubation environment only a few and species-specific responses. Compared to the interspecific differences, they resulted in only moderate climate treatment effects on mass loss and these differed among seasons and species. Neither N nor P mineralization in the litter were affected by the incubation environment. Remarkably, for all species, no net N mineralization had occurred in any of the treatments during 4 years. Species differed in P-release patterns, and summer warming strongly stimulated P release for all species. Thus, moderate changes in summer temperatures and/or winter snow addition have limited effects on litter decomposition rates and N dynamics, but summer warming does stimulate litter P release. As a result, N-limitation of plant growth in this sub-arctic bog may be sustained or even further promoted.     

陆地生态系统凋落物分解对全球气候变暖的响应

陆地生态系统凋落物分解是全球碳收支的一个重要组成部分, 主要受气候、凋落物质量和土壤生物群落的综合控制。科学家们普遍认为全球气候变化将对陆地生态系统凋落物分解产生复杂而深远的影响。该文结合凋落物分解试验的常用方法——缩微试验、原位模拟实验和自然环境梯度实验, 归纳现有研究结果, 意在揭示全球气候变化对陆地生态系统凋落物分解的直接影响(温度对凋落物分解速率的影响)和间接影响(温度对凋落物质量、土壤微生物群落及植被型的影响)的普遍规律。各种研究方法都表明: 在水分条件理想的情况下, 温度升高往往能加快凋落物的分解速率; 原位模拟实验中, 凋落物分解速率因物种、增温方法和地理方位而异; 全球气候变化能改变凋落物质量, 但可能不会在短期内影响凋落物的分解速率; 凋落物质量和可分解性的种间差异远大于增温所引发的表型响应差异, 那么, 气候变化所引发的植物群落结构和物种组成的变化将对陆地生态系统凋落物分解产生更强烈的影响; 土壤生物群落如何响应全球气候变化, 进而怎样影响凋落物分解过程, 这些都还存在着极大的不确定性。     

高山林线交错带高山杜鹃的凋落物分解

凋落物分解是维持生态系统生产力、养分循环、土壤有机质形成的关键生态过程。高山林线交错带是陆地生态系统中对气候变化响应的敏感区域。季节变化和海拔梯度上的植被类型差异可能会影响该区域凋落物的分解,进而对高山生态系统的碳氮循环产生重要影响。采用凋落物分解袋的方法,研究了川西高山林线交错带优势种高山杜鹃(Rhododendron lapponicum)凋落叶在雪被期和生长季的分解特征。结果显示:(1)季节变化和植被类型对高山杜鹃凋落物的分解均具有显著影响(P0.05),凋落叶的质量损失主要发生在生长季且在高山林线最大,暗针叶林中雪被期的质量损失略高于生长季,但差异不显著;(2)林线交错带上高山杜鹃凋落叶分解缓慢,一年干物质失重率为9.62%,拟合分解系数k为0.145;(3)高山杜鹃凋落叶的质量变化主要体现在纤维素降解显著且集中在雪被期,木质素无明显降解,在高山林线上C/N、C/P、木质素/N变化幅度较小且C、N、P的释放表现得稳定而持续。结果表明,季节性雪被对林线交错带内高山杜鹃分解的影响不仅局限在雪被期内,雪被融化期间频繁的冻融作用和雪融水淋洗作用可能会促进高山杜鹃凋落物在生长季初期的分解。总的来看,在气候变暖的情景下,雪被的缩减、生长季的延长和高山杜鹃群落的扩张可能加速高山林线交错带高山杜鹃凋落物的分解。     

Contrasting above‐ and belowground organic matter decomposition and carbon and nitrogen dynamics in response to warming in High Arctic tundra

Tundra regions are projected to warm rapidly during the coming decades. The tundra biome holds the largest terrestrial carbon pool, largely contained in frozen permafrost soils. With warming, these permafrost soils may thaw and become available for microbial decomposition, potentially providing a positive feedback to global warming. Warming may directly stimulate microbial metabolism but may also indirectly stimulate organic matter turnover through increased plant productivity by soil priming from root exudates and accelerated litter turnover rates. Here, we assess the impacts of experimental warming on turnover rates of leaf litter, active layer soil and thawed permafrost sediment in two high‐arctic tundra heath sites in NE‐Greenland, either dominated by evergreen or deciduous shrubs. We incubated shrub leaf litter on the surface of control and warmed plots for 1 and 2 years. Active layer soil was collected from the plots to assess the effects of 8 years of field warming on soil carbon stocks. Finally, we incubated open cores filled with newly thawed permafrost soil for 2 years in the active layer of the same plots. After field incubation, we measured basal respiration rates of recovered thawed permafrost cores in the lab. Warming significantly reduced litter mass loss by 26% after 1 year incubation, but differences in litter mass loss among treatments disappeared after 2 years incubation. Warming also reduced litter nitrogen mineralization and decreased the litter carbon to nitrogen ratio. Active layer soil carbon stocks were reduced 15% by warming, while soil dissolved nitrogen was reduced by half in warmed plots. Warming had a positive legacy effect on carbon turnover rates in thawed permafrost cores, with 10% higher respiration rates measured in cores from warmed plots. These results demonstrate that warming may have contrasting effects on above‐ and belowground tundra carbon turnover, possibly governed by microbial resource availability.