Diversity and species distribution of ectomycorrhizal fungi along productivity gradients of a southern boreal forest

Coniferous forests with diverse ectomycorrhizal fungus (EMF) communities are associated with nutrient-poor, acidic soils but there is some debate whether EMF can be equally adapted to more productive, nitrogen-rich sites. We compared EMF species distribution and diversity along a replicated productivity gradient in a southern boreal forest of British Columbia (Canada). Roots from subalpine fir (Abies lasiocarpa) saplings of the understory were sampled and EMF species were identified by morphotypes supplemented with ITS rDNA analysis. There were significant changes in the distribution and abundance of 74 EMF species along the productivity gradient, with as little as 24% community similarity among contrasting sites. Species richness per plot increased asymptotically with foliar nitrogen concentrations of subalpine fir, demonstrating that many EMF species were well suited to soils with high rates of nitrogen mineralization. EMF species abundance in relation to site productivity included parabolic, negative linear, and positive exponential curves. Both multi-site and more narrowly distributed EMF were documented, and a diverse mix of mantle exploration types was present across the entire productivity gradient. The results demonstrate strong associations of EMF fungal species with edaphic characteristics, especially nitrogen availability, and a specialization in EMF communities that may contribute to the successful exploitation of such contrasting extremes in soil fertility by a single tree host.     

Little evidence for niche partitioning among ectomycorrhizal fungi on spruce seedlings planted in decayed wood versus mineral soil microsites

Ectomycorrhizal fungal (EMF) communities vary among microhabitats, supporting a dominant role for deterministic processes in EMF community assemblage. EMF communities also differ between forest and clearcut environments, responding to this disturbance in a directional manner over time by returning to the species composition of the original forest. Accordingly, we examined EMF community composition on roots of spruce seedlings planted in three different microhabitats in forest and clearcut plots: decayed wood, mineral soil adjacent to downed wood, or control mineral soil, to determine the effect of retained downed wood on EMF communities over the medium and long term. If downed and decayed wood provide refuge habitat distinct from that of mineral soil, we would expect EMF communities on seedlings in woody habitats in clearcuts to be similar to those on seedlings planted in the adjacent forest. As expected, we found EMF species richness to be higher in forests than clearcuts (P ≤ 0.01), even though soil nutrient status did not differ greatly between the two plot types (P ≥ 0.05). Communities on forest seedlings were dominated by Tylospora spp., whereas those in clearcuts were dominated by Amphinema byssoides and Thelephora terrestris. Surprisingly, while substrate conditions varied among microsites (P ≤ 0.03), especially between decayed wood and mineral soil, EMF communities were not distinctly different among microhabitats. Our data suggest that niche partitioning by substrate does not occur among EMF species on very young seedlings in high elevation spruce-fir forests. Further, dispersal limitations shape EMF community assembly in clearcuts in these forests.     

Impact of moose population density on the production and composition of litter in boreal forests

Recent studies of ungulates have revealed that selective foraging seems to be an important mechanism by which they can affect the structure and species composition of the plant community, and thus quantity (dry mass) and quality (chemical composition) of litter available for decomposers. Such changes in litter production may be especially important in N-limited systems like boreal forests. We chose moose ( Alces alces ) as study species to investigate this mechanism. Moose browse mainly in the tree and shrub layers year round, and because of their wide distribution and often high population densities, they can have a significant effect on litter production of trees and shrubs in Swedish boreal forests. The effects of herbivores may also vary along productivity gradients. We therefore simulated browsing and urine and fecal deposition corresponding to 4 different moose densities in exclosures along a pre-existing forest productivity gradient. Both litter quantity (g dry mass per m² and year) and contributions of C and N (g dry mass per m² and year) decreased with increasing level of simulated moose density. High moose densities over extended time can therefore reduce N contributions to soil and therefore eventually reduce site productivity in Swedish boreal forests. This effect of moose was mainly a result of decreased litter quantity, because contradictory to studies from North America, litter quality (C:N ratio and N contribution per mass unit of litter) was not affected by level of simulated moose density.     

Nonlinear responses of ecosystem carbon fluxes to nitrogen deposition in an old-growth boreal forest

Nitrogen (N) deposition is known to increase carbon (C) sequestration in N-limited boreal forests. However, the long-term effects of N deposition on ecosystem carbon fluxes have been rarely investigated in old-growth boreal forests. Here we show that decade-long experimental N additions significantly stimulated net primary production (NPP) but the effect decreased with increasing N loads. The effect on soil heterotrophic respiration (Rh) shifted from a stimulation at low-level N additions to an inhibition at higher levels of N additions. Consequently, low-level N additions resulted in a neutral effect on net ecosystem productivity (NEP), due to a comparable stimulating effect on NPP and Rh, while NEP was increased by high-level N additions. Moreover, we found nonlinear temporal responses of NPP, Rh and NEP to low-level N additions. Our findings imply that actual N deposition in boreal forests likely exerts a minor contribution to their soil C storage.     

Ectomycorrhizal community composition and function in a spruce forest transitioning between nitrogen and phosphorus limitation

Nitrogen is the main limiting nutrient in boreal ecosystems, but studies in southwest Sweden suggest that certain forests approach phosphorus (P) limitation driven by nitrogen (N) deposition. We added N, P or N + P to a Norway spruce forest in this region, to push the system to N or P limitation. Tree growth and needle nutrient concentrations indicated that the trees are P limited. EMF biomass was reduced only by N + P additions. Soil EMF communities responded more strongly to P than to N. Addition of apatite to ingrowth meshbags altered EMF community composition and enhanced the abundance of Imleria badia in the control and N plots, but not when P was added. The ecological significance of this species is discussed. Effects on tree growth, needle chemistry, and EMF communities indicate a dynamic interaction between EMF fungi and the nutrient status of trees and soils.     

The essential role of biodiversity in the key axes of ecosystem function

Biodiversity is essential for maintaining the terrestrial ecosystem multifunctionality (EMF). Recent studies have revealed that the variations in terrestrial ecosystem functions are captured by three key axes: the maximum productivity, water use efficiency, and carbon use efficiency of the ecosystem. However, the role of biodiversity in supporting these three key axes has not yet been explored. In this study, we combined the (i) data collected from more than 840 vegetation plots across a large climatic gradient in China using standard protocols, (ii) data on plant traits and phylogenetic information for more than 2,500 plant species, and (iii) soil nutrient data measured in each plot. These data were used to systematically assess the contribution of environmental factors, species richness, functional and phylogenetic diversity, and community-weighted mean (CWM) and ecosystem traits (i.e., traits intensity normalized per unit land area) to EMF via hierarchical partitioning and Bayesian structural equation modeling. Multiple biodiversity attributes accounted for 70% of the influence of all the variables on EMF, and ecosystems with high functional diversity had high resource use efficiency. Our study is the first to systematically explore the role of different biodiversity attributes, including species richness, phylogenetic and functional diversity, and CWM and ecosystem traits, in the key axes of ecosystem functions. Our findings underscore that biodiversity conservation is critical for sustaining EMF and ultimately ensuring human well-being.     

Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island,Magellan Strait

Glacier foreland moraines provide an ideal model to examine the patterns of ecosystem development and the evolution of nitrogen and phosphorous limitation over successional time. In this paper, we focus on a 400‐year soil chronosequence in the glacier forelands of Santa Inés Island in the Magellan Strait, southern Chile by examining forest development on phosphorus (P)‐poor substrates in a uniquely unpolluted region of the world. Results show a steady increase in tree basal area and a humped trend in tree species richness over four centuries of stand development. The increase in basal area suggests that the late successional tree species were more efficient nutrient users than earlier successional ones. Total contents of carbon (C) and nitrogen (N) in soils increased during the chronosequence, reaching an asymptote in late succession. The net increases in soil C : N, C : P and N : P ratios observed over successional time suggest that nutrient limitation is maximal in 400‐year‐old substrates. Foliar C : N and C : P ratios also increased over time to reach an asymptote in old‐growth stages, following soil stoichiometric relationships; however the foliar N‐to‐P ratio remained constant throughout the chronosequence. Biological N fixation was greater in early postglacial succession, associated with the presence of the symbiotic N‐fixer Gunnera magellanica. Declining trends of δ¹⁵N in surface soils through the 400‐year chronosequence are evidence of decreasing N losses in old‐growth forests. In synthesis, glacier foreland chronosequences at this high South American latitude provide evidence for increasing efficiency of N and P use in the ecosystem, with the replacement of shade‐intolerant pioneers by more efficient, shade‐tolerant tree species. This pattern of ecosystem development produces a constant foliar N : P ratio, regardless of variation in soil N‐to‐P ratio over four centuries.     

Contrasting patterns of soil N-cycling in model ecosystems of Fennoscandian boreal forests

The low plant productivity of boreal forests in general has been attributed to low soil N supply and low temperatures. Exceptionally high productivity occurs in toe-slope positions, and has been ascribed to influx of N from surrounding areas and higher rates of soil N turnover in situ. Despite large apparent natural variations in forest productivity, rates of gross soil N mineralization and gross nitrification have never been compared in Fennoscandian boreal forests of contrasting productivity. We report contrasting patterns of soil N turnover in three model ecosystems, representing the range in soil C-to-N ratios (19–41) in Fennoscandian boreal forests and differences in forest productivity by a factor close to 3. Gross N mineralization was seven times higher when soil, microbial, and plant C-to-N ratios were the lowest compared to the highest. This process, nitrification and potential denitrification correlated with inorganic, total and microbial biomass N, but not microbial C. There was a constant ratio between soil and microbial C-to-N ratio of 3.7±0.2, across wide ratios of soil C-to-N and fungi-to-bacteria. Soil N-cycling should be controlled by the supplies of C and N to the microbes. In accordance with plant allocation theory, we discuss the possibility that the high fungal biomass at high soil C-to-N ratio reflects a particularly high supply of plant photosynthates, substrates of high-quality C, to mycorrhizal fungi. Methods to study soil N turnover and N retention should be developed to take into account the impact of mycorrhizal fungi on soil N-cycling.     

Diversity patterns of fungivorous insects: comparison between glaciated vs. refugial boreal forests

Aim The aim of this study was to compare diversity patterns of insect communities associated with the wood‐decaying Fomitopsis fungi in north‐east (NE) Asia and Fennoscandia. We hypothesized that the diversity of the fungal–insect communities is greater in NE Asia, because the region was one of the largest refugia of boreal species during the last Pleistocene glaciation. Location This study was conducted in boreal forests in NE People's Republic of China, and in south‐central Finland. Methods Fruiting bodies of three Fomitopsis species were collected from the field in order to rear the insects inhabiting the fruiting bodies. Taxonomic uniqueness, diversity and food web structure of the insect assemblages were analysed using hierarchical cluster analysis, diversity indices and analyses of food web compartmentalization, respectively. Results Contrary to the expectations, the richness of families, genera and species was greater in Finland than in NE China. This applied for the overall Fomitopsis‐associated insect assemblage, as well as for the fungal species separately. The taxonomic composition at the level of families and genera was similar between the two regions. The level of monophagy was higher in Finland and the food web was divided into compartments corresponding to the two Fomitopsis species. Main conclusion The often‐suggested higher diversity in refugial areas does not apply for all taxa in boreal forests, such as fungivorous insects associated with Fomitopsis fungi.     

Tree cover at fine and coarse spatial grains interacts with shade tolerance to shape plant species distributions across the Alps

The role of competition for light among plants has long been recognised at local scales, but its importance for plant species distributions at larger spatial scales has generally been ignored. Tree cover modifies the local abiotic conditions below the canopy, notably by reducing light availability, and thus, also the performance of species that are not adapted to low‐light conditions. However, this local effect may propagate to coarser spatial grains, by affecting colonisation probabilities and local extinction risks of herbs and shrubs. To assess the effect of tree cover at both the plot‐ and landscape‐grain sizes (approximately 10‐m and 1‐km), we fit generalised linear models (GLMs) for the plot‐level distributions of 960 species of herbs and shrubs using 6935 vegetation plots across the European Alps. We ran four models with different combinations of variables (climate, soil and tree cover) at both spatial grains for each species. We used partial regressions to evaluate the independent effects of plot‐ and landscape‐grain tree cover on plot‐level plant communities. Finally, the effects on species‐specific elevational range limits were assessed by simulating a removal experiment comparing the species distributions under high and low tree cover. Accounting for tree cover improved the model performance, with the probability of the presence of shade‐tolerant species increasing with increasing tree cover, whereas shade‐intolerant species showed the opposite pattern. The tree cover effect occurred consistently at both the plot and landscape spatial grains, albeit most strongly at the former. Importantly, tree cover at the two grain sizes had partially independent effects on plot‐level plant communities. With high tree cover, shade‐intolerant species exhibited narrower elevational ranges than with low tree cover whereas shade‐tolerant species showed wider elevational ranges. These findings suggest that forecasts of climate‐related range shifts for herb and shrub species may be modified by tree cover dynamics.     

植物多样性与生态系统土壤保持功能关系及其生态学意义

测定物种丰富度呈梯度变化的半湿润常绿阔叶林不同次生演替阶段小区地表径流、土壤侵蚀和总磷流失及影响这些过程的植物群落郁闭度、个体密度、胸高断面积、植物叶吸附水,分析物种多样性与生态系统土壤保持功能、稳定性及直接影响土壤保持功能的群落结构、树冠截留间的关系。结果表明,在降雨、坡度、坡向、坡位、土壤类型等水土保持影响因子相同条件下,随着各小区物种多样性的增加,地表产流次数不断下降;在3个降雨季节,物种多样性最低的小区产生地表径流77次,而物种多样性最高小区产生地表径流才9次;系列小区地表径流、土壤侵蚀和总磷流失随着物种多样性增加呈幂指数下降;物种多样性最低的小区地表径流、土壤侵蚀和总磷流失分别为960.20 m³·hm^-2·a^-1,11.4 t·hm^-2·a^-1,127.69 kg·hm^-2·a^-1,而物种多样性最高的小区为75.55 m³·hm^-2·a^-1、0.28 t·hm^-2·a^-1、4.71 kg·hm^-2·a^-1,分别相差12、50和25倍;地表径流、土壤侵蚀和总磷流失变异系数也呈幂指数下降,物种多样性最高的小区地表径流、土壤侵蚀和总磷流失的变异系数分别为57.93、187.94和 59.2,而物种多样性最低的小区变异系数高达287.6、534.21、315.47,分别相差4、3和5倍。物种多样性与影响土壤保持功能的群落郁闭度、密度和胸高断面积呈正相关关系。不同演替阶段植物叶吸附水量差异显著,吸附水量最高的演替阶段是次生半湿润常绿阔叶林,为12.28 t·hm^-2·a^-1, 最低是云南松(Pinus yunnanensis)林, 为4.15 t·hm^-2·a^-1。“植物多样性-土壤保持功能相关群落结构因子及树冠截留效应-生态系统土壤保持功能”的耦合关系表明了植物多样性通过植物群落结构削弱了降雨动能,减少了地表径流,减轻了土壤及营养元素的流失,以间接方式调控生态系统土壤保持功能,维持系统营养的持续性,在不同尺度上实现生态系统生产力。物种多样性的提高,促进了生态系统土壤保持功能的稳定性。植物多样性-生态系统土壤保持过程的研究可能是生态系统稳定性研究的好方法。用植物叶吸附水测定可评价群落树冠截留效应。由于植物多样性与生态系统土壤保持功能间存在相关关系,基于植物多样性对生态系统土壤保持功能作用模式,可增进对生命系统和地球系统界面间相互作用关系的了解。     

The relationship between species richness and biomass changes from boreal to subtropical forests in China

Diversity‐manipulation experiments suggest a positive effect of biodiversity on ecosystem properties (EPs), but variable relationships between species richness and EPs have been reported in natural communities. An explanation for this discrepancy is that observed richness–EPs relationships in natural communities change with environment and species functional identities. But how the relationships change along broad‐scale climatic gradients has rarely been examined. In this paper, we sampled 848 plots of 20 × 30 m² from boreal to tropical forests across China. We examined plot biomass with respect to environmental factors, tree species richness and functional group identity (FGI, i.e. evergreen vs deciduous, and coniferous vs broadleaf). Variation partitioning was used to evaluate the relative effects of the three classes of factors. We found that, most of the ‘effects’ (percentage of variation explained) of richness and FGI on forest biomass were shared with environmental factors, but species richness and FGI still revealed significant effects in addition to environment for plots across China. Richness and FGI explained biomass mainly through their shared effects instead of independent effects, suggesting that the positive biodiversity effect is closely associated with a sampling effect. The relative effects of richness, FGI and environment varied latitudinally: the independent effects of environment and richness decreased from boreal to subtropical forests, whereas the total effect of FGI increased. We also found that the slope of richness–biomass relationship decreased monotonically from boreal to subtropical forests, possibly because of decreasing complementarity and increasing competition with increasing productivity. Our results suggest that while species richness does have significant effects on forest biomass it is less important than environmental gradients and other biotic factors in shaping large‐scale biomass patterns. We suggest that understanding how and why the diversity–EPs relationships change along climatic gradient would be helpful for a better understanding of real biodiversity effects in natural communities.     

Ectomycorrhizal-Dominated Boreal and Tropical Forests Have Distinct Fungal Communities,but Analogous Spatial Patterns across Soil Horizons

Fungi regulate key nutrient cycling processes in many forest ecosystems, but their diversity and distribution within and across ecosystems are poorly understood. Here, we examine the spatial distribution of fungi across a boreal and tropical ecosystem, focusing on ectomycorrhizal fungi. We analyzed fungal community composition across litter (organic horizons) and underlying soil horizons (0–20 cm) using 454 pyrosequencing and clone library sequencing. In both forests, we found significant clustering of fungal communities by site and soil horizons with analogous patterns detected by both sequencing technologies. Free-living saprotrophic fungi dominated the recently-shed leaf litter and ectomycorrhizal fungi dominated the underlying soil horizons. This vertical pattern of fungal segregation has also been found in temperate and European boreal forests, suggesting that these results apply broadly to ectomycorrhizal-dominated systems, including tropical rain forests. Since ectomycorrhizal and free-living saprotrophic fungi have different influences on soil carbon and nitrogen dynamics, information on the spatial distribution of these functional groups will improve our understanding of forest nutrient cycling.     

Naturally Saline Boreal Communities as Models for Reclamation of Saline Oil Sand Tailings

Reclaimed landscapes after oil sands mining have saline soils; yet, they are required to have similar biodiversity and productivity as the predisturbance nonsaline landscape. Given that many species in the boreal forest are not tolerant of salinity, we studied the effects of soil salinity on plant communities in natural saline landscapes to understand potential plant responses during the reclamation process. Vegetation–soil relationships were measured along transects from flooded wetlands to upland forest vegetation in strongly saline, slightly saline, nonsaline, and reclaimed boreal landscapes. In strongly saline landscapes, surface soil salinity was high (>10 dS/m) in flooded, wet‐meadow, and dry‐meadow vegetation zones as compared to slightly saline (<5 dS/m) and nonsaline (<2 dS/m) landscapes. Plant communities in these vegetation zones were quite different from nonsaline boreal landscapes and were dominated by halophytes common to saline habitats of the Great Plains. In the shrub and forest vegetation zones, surface soil salinity was similar between saline and nonsaline landscapes, resulting in similar plant communities. In strongly saline landscapes, soils remained saline at depth through the shrub and forest vegetation zones (>10 dS/m), suggesting that forest vegetation can establish over saline soils as long as the salts are below the rooting zone. The reclaimed landscape was intermediate between slightly saline and nonsaline landscapes in terms of soil salinity but more similar to nonsaline habitats with respect to species composition. Results from this study suggest it may be unrealistic to expect that plant communities similar to those found on the predisturbance landscape can be established on all reclaimed landscapes after oil sands mining.     

Stress tolerance of soil fungal communities from native Atlantic forests,reforestations, and a sand mining degraded area

Microorganisms are essential to the functionality of the soil, particularly in organic matter decomposition and nutrient cycling, which regulate plant productivity and shape the soil structure. However, biotic and abiotic stresses greatly disrupt soil fungal communities and, thereby, disturb the ecosystem. This study quantified seasonal tolerances to UV-B radiation and heat of fungal communities, which could be cultured, found in soil from two native Atlantic forest fragments called F1 and F2, five reforested areas (RA) planted in 1994, 1997, 2004, 2007, and 2009 with native species of the Atlantic forest, and one sand mining degraded soil (SMDS). The cold activity of the soil fungal communities (FC) from the eight different areas was also studied. Higher tolerance to UV-B radiation and heat was found in the FC from the SMDS and the 2009RA, where the incidence of heat and UV radiation from sun was more intense, which caused selection for fungal taxa that were more UV-B and heat tolerant in those areas. Conversely, the FC from the native forests and older reforested sites were very susceptible to heat and UV-B radiation. The cold activity of the soil FC from different areas of the study showed an erratic pattern of responses among the sampling sites. Little difference in tolerance to UV-B radiation and heat was found among the FC of soil samples collected in different seasons; in general soil FC collected in winter were less tolerant to UV-B radiation, but not for heat. In conclusion, FC from SMDS soil that receive intense heat and UV radiation, as well as with low nutrient availability, were more tolerant to both UV-B radiation and heat.     

Carbon and Nitrogen Dynamics of Boreal Jack Pine Stands With and Without a Green Alder Understory

We compared the species composition, structure and selected components of the carbon (C) and nitrogen (N) budgets of similar-aged, mature boreal jack pine (Pinus banksiana Lamb.) forests with and without green alder [Alnus crispa (Ait.) Pursh.] in two different boreal environments. The C and N content of the overstory biomass components (for example, stem, branch, and foliage), total vegetation, forest floor, and mineral soil were greater (P= 0.05 to P= 0.10) for jack pine with alder (JPA) stands than for jack pine without alder (JP) stands at both study areas. Jack pine foliage N isotopic discrimination (δ¹⁵N) and annual litterfall N content were significantly greater (P < 0.05) in the JPA than the JP stands at both study areas, suggesting that alder was fixing N and that N availability was greater in the JPA than the JP stands. The greater leaf area index (LAI) and overstory C accumulation in the JPA than the JP stands (P < 0.05) is likely because of the greater N availability in the JPA stands, but the effect of soil texture discontinuity on water availability in the JPA stands can not be dismissed. Percent ground cover by feathermoss varied among the jack pine communities and was positively correlated with overstory LAI (r ²= 0.83, P< 0.05). One index of N-use efficiency (NUE), defined as aboveground net primary productivity (ANPP) per litterfall N, was significantly greater (P < 0.05) for the JP than the JPA stands, but a second index of NUE, ANPP/N uptake, did not differ between the two jack pine communities. Jack pine trees growing without alder produced more organic matter per unit of N, but percent N retranslocation from senescing foliage and N mean residence time in the overstory did not differ between the JPA and the JP stands. A conceptual model is presented that illustrates the potential influence of alder on the species composition, structure, and function of boreal jack pine forests. Received 6 January 1998; accepted 15 April 1998.     

Negative impacts of high temperatures on growth of black spruce forests intensify with the anticipated climate warming

An increasing number of studies conclude that water limitations and heat stress may hinder the capacity of black spruce (Picea mariana (Mill.) B.S.P.) trees, a dominant species of Canada's boreal forests, to grow and assimilate atmospheric carbon. However, there is currently no scientific consensus on the future of these forests over the next century in the context of widespread climate warming. The large spatial extent of black spruce forests across the Canadian boreal forest and associated variability in climate, demography, and site conditions pose challenges for projecting future climate change responses. Here we provide an evaluation of the impacts of climate warming and drying, as well as increasing [CO₂], on the aboveground productivity of black spruce forests across Canada south of 60°N for the period 1971 to 2100. We use a new extensive network of tree‐ring data obtained from Canada's National Forest Inventory, spatially explicit simulations of net primary productivity (NPP) and its drivers, and multivariate statistical modeling. We found that soil water availability is a significant driver of black spruce interannual variability in productivity across broad areas of the western to eastern Canadian boreal forest. Interannual variability in productivity was also found to be driven by autotrophic respiration in the warmest regions. In most regions, the impacts of soil water availability and respiration on interannual variability in productivity occurred during the phase of carbohydrate accumulation the year preceding tree‐ring formation. Results from projections suggest an increase in the importance of soil water availability and respiration as limiting factors on NPP over the next century due to warming, but this response may vary to the extent that other factors such as carbon dioxide fertilization, and respiration acclimation to high temperature, contribute to dampening these limitations.     

Plant Community Composition as a Predictor of Regional Soil Carbon Storage in Alaskan Boreal Black Spruce Ecosystems

The boreal forest is the largest terrestrial biome in North America and holds a large portion of the world’s reactive soil carbon. Therefore, understanding soil carbon accumulation on a landscape or regional scale across the boreal forest is useful for predicting future soil carbon storage. Here, we examined the relationship between floristic composition and ecosystem parameters, such as soil carbon pools, the carbon-to-nitrogen (C/N) ratio of live black spruce needles, and normalized basal area increment (NBAI) of trees in black spruce communities, the most widespread forest type in the boreal forest of Alaska. Variability in ecosystem properties among black spruce stands was as large as that which had previously been documented among all forest types in the central interior of Alaska; we found an eightfold range in NBAI and fivefold range in mineral soil carbon and nitrogen pools. Acidic black spruce communities had significantly more carbon in the organic soil horizon than did nonacidic black spruce communities, but did not differ in any other measured ecosystem parameter. We explained 48% of the variation in total soil carbon with a combination of plant community indices and abiotic and biotic factors. Plant community composition was at least as effective as any single environmental factor or stand characteristic in predicting soil C pools in Alaskan black spruce ecosystems. We conclude that among the community properties analyzed, the presence of key groups of species, overall species composition, and diversity of certain functional types, especially Sphagnum moss species, are important predictors of soil carbon sequestration in the black spruce forest type.     

Effects of Light Gaps and Litter Removal on the Seedling Performance of Six African Timber Species1

Valuable timber tree species frequently show poor regeneration after selective logging in tropical forests. Small size of logging gaps, lack of soil disturbance, and limited seed availability have each been blamed for observed regeneration failures. We investigated seed germination and seedling performance using a split‐plot factorial design involving light availability and litter removal for six Central African timber tree species, hypothesizing that canopy gaps and litter removal would improve seedling establishment, and that less shade‐tolerant species would show stronger responses to both factors. Contrary to our expectations, significantly more germinants established on intact litter than on exposed mineral soil 3 mo after seeding. After 18 mo, seedling survival, height and diameter growth, leaf area, and rooting depth were all much higher in gap plots than in the understory for all species, with the exception of Gilbertiodendron dewevrei, a highly shade‐tolerant species whose survival was higher in the understory. Leaf production was negatively influenced by litter removal in the least shade‐tolerant species, Nauclea diderrichii, with weak or positive effects in other species. G. dewevrei, while displaying a low‐light threshold for growth, exhibited a surprisingly high growth response to increasing light comparable to more shade‐intolerant species, a response that may help explain its local competitive dominance in the region. Due to the rapid closure of small gaps, we suggest that shade‐intolerant species such as N. diderrichii, Khaya anthotheca, and Entandrophragma utile might benefit from more intensive silvicultural practices that create larger canopy gaps.     

Organic mound-building ants: their impact on soil properties in temperate and boreal forests

Ants are important components of most soil invertebrate communities, and can affect the flow of energy, nutrients and water through many terrestrial ecosystems. The vast majority of ant species build nests in the mineral soil, but a small group of ants in temperate and boreal forests of Eurasia and North America build large parts of their nests above‐ground using organic materials collected from the surrounding soil. Many studies have shown that ants nesting in mineral soil can affect water infiltration rates, soil organic matter (OM) content, and nutrient cycling, but much less is known on how mound‐building ants influence soil physical and chemical properties. In this paper we summarize what is known on the soil impacts of organic mound‐building ants in temperate and boreal forests, and how these ants could be affected by ecosystem disturbance and future climate change. Much of this information comes from studies on Formica rufa group ants in Europe, which showed that CO₂ emissions and concentrations of C, N, and P are usually higher in ant mounds than in the surrounding forest soil. However, ant mounds are a minor component of total soil C and nutrient pools, but they do increase spatial heterogeneity of soil water and available nutrients. Mound‐building ants can also impact tree growth, which could change the quantity and quality of OM added to soil. Forest management, fire, and projected climate change, especially in boreal forests, could affect mound‐building ant population dynamics, and indirectly, soil properties.