Spatial and temporal variation in moss-associated dinitrogen fixation in coniferous- and deciduous-dominated Alaskan boreal forests

Dominant canopy tree species have strong effects on the composition and function of understory species, particularly bryophytes. In boreal forests, bryophytes and their associated microbes are a primary source of ecosystem nitrogen (N) inputs, and an important process regulating ecosystem productivity. We investigated how feather moss-associated N₂-fixation rates and contribution to N budgets vary in time and space among coniferous and broadleaf deciduous forests. We measured N₂-fixation rates using stable isotope (¹⁵N₂) labeling in two moss species (Pleurozium schreberi and Hylocomium splendens) in broadleaf deciduous (Alaska paper birch—Betula neoalaskana) and coniferous (black spruce—Picea mariana) stands near Fairbanks, interior Alaska, from 2013 to 2015. N₂-fixation rates showed substantial inter-annual variation among the 3 years. High N₂-fixation was more strongly associated with high precipitation than air temperature or light availability. Overall, contribution of N₂-fixation to N budgets was greater in spruce than in birch stands. Our results enhance the knowledge of the processes that drive N₂-fixation in boreal forests, which is important for predicting ecosystem consequences of changing forest composition.     

Absence of snow cover reduces understory plant cover and alters plant community composition in boreal forests

Snow regimes affect biogeochemistry of boreal ecosystems and are altered by climate change. The effects on plant communities, however, are largely unexplored despite their influence on relevant processes. Here, the impact of snow cover on understory community composition and below-ground production in a boreal Picea abies forest was investigated using a long-term (8-year) snow cover manipulation experiment consisting of the treatments: snow removal, increased insulation (styrofoam pellets), and control. The snow removal treatment caused longer (118 vs. 57 days) and deeper soil frost (mean minimum temperature −5.5 vs. −2.2°C) at 10 cm soil depth in comparison to control. Understory species composition was strongly altered by the snow cover manipulations; vegetation cover declined by more than 50% in the snow removal treatment. In particular, the dominant dwarf shrub Vaccinium myrtillus (−82%) and the most abundant mosses Pleurozium schreberi (−74%) and Dicranum scoparium (−60%) declined strongly. The C:N ratio in V. myrtillus leaves and plant available N in the soil indicated no altered nitrogen nutrition. Fine-root biomass in summer, however, was negatively affected by the reduced snow cover (−50%). Observed effects are attributed to direct frost damage of roots and/ or shoots. Besides the obvious relevance of winter processes on plant ecology and distribution, we propose that shifts in the vegetation caused by frost damage may be an important driver of the reported alterations in biogeochemistry in response to altered snow cover. Understory plant performance clearly needs to be considered in the biogeochemistry of boreal systems in the face of climate change.     

Statistical opportunities for analyzing spatial and temporal heterogeneity of field soils

Statistical techniques for analyzing data in the agricultural sciences have traditionally followed the pioneering efforts of R.A. Fisher who assumed that observations in the field were independent and identically distributed. Such techniques, proven useful in the past and still being used today for comparing the merits of different management practices or different treatments, are presently giving way to additional methods that are based upon observations being spatially or temporally correlated. It is physically more sensible to expect soil attributes to be correlated when they are measured at adjacent points in space or time. Spatially repetitious patterns of soil attributes for physical and biological processes occurring at distances of a few molecules to those of kilometers are also expected. Opportunities to use geostatistics, time series analyses, state-space models, spectral analyses of variance, lagged regression models and other alternative techniques for analyzing multidimensional random fields are available to enhance the understanding of agro-ecosystems. Approaches to modeling and fitting data using stochastic partial differential equations and scaling techniques also help reveal the underlying processes occurring in field soils. Inclusion of these alternatives in the development of an agro-ecological technology leads to improved sampling designs to better entire management units, rather than ascertaining the impact of particular, sometimes arbitrary treatments applied to a set of small plots using analysis of variance methods.     

Exploring potential biodiversity indicators in boreal forests

The present study evaluates indicators in Swedish spruce forests. We ask whether different species groups co-vary in their occurrence and to what extent species richness and composition is predictable from habitat structures. We studied 10 boreal spruce forest stands constituting a gradient in degree of selective logging. Occurrences of vascular plants, bryophytes, epiphytic lichens and wood-inhabiting fungi as well as habitat structures was inventoried. In addition, in five of the stands, beetles were sampled with windows traps. Total species richness was correlated with several habitat factors, mainly particular substrates and degree of forestry impact. However, the richness of a set of species regularly used as indicators did not correlate with habitat factors. Correlation in species richness among different organism groups were few and scale dependent. Only lichens and vascular plants formed nested subset patterns (i.e. species composition at poorer sites is subsets of the species present at richer sites) among the study sites. The study shows that in this forest type one cannot a priori assume that richness in one group of species correlated with richness in other, and measures of single habitat features may be relevant only to particular groups of species. Instead, monitoring and inventories should be based on a set of factors reflecting important aspects for different groups of organisms and if indicator species are to be used these should be chosen from several species groups.     

Competitive interactions among raptors in boreal forests

We examined inter-specific interactions among goshawks (Accipiter gentilis), common buzzards (Buteo buteo) and honey buzzards (Pernis apivorus) in western Finland in 1983–1996. Because goshawks are among the largest birds of prey species in boreal forests they may take over the nest of smaller and less-competitive forest-dwelling raptors when searching for suitable places for breeding. Accordingly, more than half of newly established goshawk territories were found on the territories previously occupied by the common buzzard and the honey buzzard. Otherwise, territory sharing between these species was rare. Fledgling production of honey buzzards was not associated with the presence of goshawks, probably owing to the almost 2 months later onset of breeding. This probably decreases competitive interactions between these two species. An intensive interference competition, instead, seemed to be evident between common buzzards and goshawks, because the fledgling production of common buzzards was decreased by 20% as a result of failures during incubation and nestling period in the vicinity (<1 km) of occupied goshawk nests. Similarly, territory occupancy of common buzzards till the next breeding season was significantly reduced in the presence of goshawks. Relatively high proportions of occupied buzzard territories (17%) in the study area were shared by breeding goshawks on the same territory. This suggests that although their diets are dissimilar they inhabit similar habitats and might compete for the available prime nesting habitats within forest landscapes. In addition, goshawks benefit from taking over the complete nests of other raptors, imposing upon the original owners of the nest, because building a large stick nest is probably energetically costly. As a large raptor, the goshawk apparently has a competitive advantage over smaller ones, and may have an ever-increasing impact on smaller birds of prey, if there is a lack of sheltered forests inducing competition for the available nest sites.     

High temporal but low spatial heterogeneity of bacterioplankton in the Chesapeake Bay

Compared to freshwater and the open ocean, less is known about bacterioplankton community structure and spatiotemporal dynamics in estuaries, particularly those with long residence times. The Chesapeake Bay is the largest estuary in the United States, but despite its ecological and economic significance, little is known about its microbial community composition. A rapid screening approach, ITS (internal transcribed spacer)-LH (length heterogeneity)-PCR, was used to screen six rRNA operon (16S rRNA-ITS-23S rRNA) clone libraries constructed from bacterioplankton collected in three distinct regions of the Chesapeake Bay over two seasons. The natural length variation of the 16S-23S rRNA gene ITS region, as well as the presence and location of tRNA-alanine coding regions within the ITS, was determined for 576 clones. Clones representing unique ITS-LH-PCR sizes were sequenced and identified. Dramatic shifts in bacterial composition (changes within subgroups or clades) were observed for the Alphaproteobacteria (Roseobacter clade, SAR11), Cyanobacteria (Synechococcus), and Actinobacteria, suggesting strong seasonal variation within these taxonomic groups. Despite large gradients in salinity and phytoplankton parameters, a remarkably homogeneous bacterioplankton community was observed in the bay in each season. Stronger seasonal, rather than spatial, variation of the bacterioplankton population was also supported by denaturing gradient gel electrophoresis and LH-PCR analyses, indicating that environmental parameters with stronger seasonal, rather than regional, dynamics, such as temperature, might determine bacterioplankton community composition in the Chesapeake Bay.     

Pollen availability for predaceous mites on apple: spatial and temporal heterogeneity

It has been suggested that an abundance of alternate food early in the spring may be critical to the ability of generalist predaceous mites to suppress spider mite pests. One alternate food that is typically very abundant in spring is wind-dispersed pollen. Here we investigate, at several spatial scales, the heterogeneity in the availability of pollen to predaceous mites on apple. We found pollen to be abundant on apple leaves very soon after they opened (>100 grains/cm 2 ), and that the dominant pollen types at this time were wind-dispersed tree pollens (Betulaceae and Pinaceae). We found that most of the spatial variation in pollen abundance occurred at either small spatial scales (within trees) or very large spatial scales (among orchard blocks). Variability among orchards was clearly influenced by the surrounding vegetation, and probably also by the management regime (frequency of mowing). Spatial heterogeneity in pollen availability may affect the build-up of predatory mite populations in the spring, as we found early season abundances of Typhlodromus pyri (Phytoseiidae) and Zetzellia mali (Stigmaeidae) to be better correlated with early season pollen density than with abundance of mite prey (Aculus schlechtendali).     

Carbon and nitrogen cycling in North American boreal forests

A model of boreal forest dynamics was adapted to examine the factors controlling carbon and nitrogen cycling in the boreal forests of interior Alaska. Empirical relationships were used to simulate decomposition and nitrogen availability as a function of either substrate quality, the soil thermal regime, or their interactive effects. Test comparisons included black spruce forests growing on permafrost soils and black spruce, birch, and white spruce forests growing on permafrost-free soils. For each forest, simulated above-ground tree biomass, basal area, density, litterfall, moss biomass, and forest floor mass, turnover, thickness, and nitrogen concentration were compared to observed data. No one decay equation simulated forests entirely consistent with observed data, but over the range of upland forest types in interior Alaska, the equation that combined the effects of litter quality and the soil thermal regime simulated forests that were most consistent with observed data. For black spruce growing on permafrost soils, long-term simulated forest dynamics in the absence of fire resulted in unproductive forests with a thick forest floor and low nitrogen mineralization. Fires were an important means to interrupt this sequence and to restart forest succession.     

The importance of small-scale heterogeneity in boreal forests: variation in diversity in forest-floor invertebrates across the succession gradient

Heterogeneity in species assemblages of forest-floor arthropods — carabid beetles, ants and spiders — within and between different forest age classes was studied in the southern Finnish taiga. The importance of processes operating on the local scale (within the movement radius of the species) vs on the regional scale (among forest stands) in determining the observed variation was assessed. Four data sets with different spatial resolutions in mesic forests in the same general study area were used. The material consists of 18 283 carabids of 51 species, 48 769 spiders of 212 species, and 126 718 worker ants of 23 species. Analyses of abundance variation and species complementarity among successional stages revealed that in all the three taxa species occurring in the mature forest were prevalent in the younger successional stages as well, constituting more than half of the catch in any age class. A great majority of carabid and spider species were widely distributed across the forest age classes, whereas ants include a higher proportion of species with a narrower amplitude across the succession gradient. Comparisons of similarity between samples at increasing distance from one another on the local scale within forest stands (a few tens of meters to a few hundreds of meters) showed a quite consistent pattern in carabids and spiders: there was more variation between sampling sites in young successional forests than in the mature forest. Furthermore, only in the mature forest a slight, albeit statistically not significant, negative relationship between similarity of samples and distance between sampling sites was detected. In carabids and spiders, comparisons between samples located at a distance of 10–15 m from each other showed considerable heterogeneity, the mean percentage similarity being c. 0.6 (in ants c. 0.8). On the regional scale, systematic variation between young and mature forest stands is a major element increasing the total diversity (species turnover c. 50% in carabids and spiders; compositional similarity c. 0.3–0.4 in carabids, 0.2–0.3 in spiders), but variation within forest stands on a spatial scale of 10–15 meters is another important component in the total heterogeneity. The results suggest that regional abundance variation is a primary factor influencing the composition of local assemblages; a set of hypotheses elaborating this conclusion is formulated. The result implies that maintenance of habitat heterogeneity on a small scale (10–15 m) is needed to preserve biodiversity in managed forests.     

Effects of fire severity on plant nutrient uptake reinforce alternate pathways of succession in boreal forests

Fire activity in the North American boreal region is projected to increase under a warming climate and trigger changes in vegetation composition. In black spruce forests of interior Alaska, fire severity impacts residual organic layer depth which is strongly linked to the relative dominance of deciduous versus coniferous trees in early succession. These alternate successional pathways may be reinforced by biogeochemical processes that affect the relative ability of deciduous versus coniferous trees to acquire limiting nutrients. To test this hypothesis, we examined changes in soil inorganic nitrogen (N) supply and in situ ¹⁵N root uptake by aspen (Populus tremuloides) and black spruce (Picea mariana) saplings regenerating in lightly and severely burned sites, 16 years following fire. Fire severity did not impact the composition or magnitude of N supply, and nitrate represented nearly 40 % of total N supply. Both aspen and spruce took up more N in severely burned than in lightly burned sites. Spruce exhibited only a moderately lower rate of nitrate uptake, and a higher ammonium uptake rate than aspen in severely burned sites. At the stand level, differences in species nutrient uptake were magnified, with aspen taking up nearly an order-of-magnitude more N per m² in severely burned than in lightly burned sites. We suggest that differences in nutrient sinks (biomass) established early in succession and effects of post-fire organic layer depth on nutrient uptake, are key mechanisms reinforcing the opposing stand dominance patterns that have developed in response to variations in organic layer depth.     

Plant acquisition of organic nitrogen in boreal forests

Research on plant nitrogen (N) uptake and metabolism has more or less exclusively concerned inorganic N, particularly nitrate. Nevertheless, recent as well as older studies indicate that plants may have access to organic N sources. Laboratory studies have shown that ectomycorrhizal and ericoid mycorrhizal plants can degrade polymeric N and absorb the resulting products. Recent studies have also shown that some non‐mycorrhizal plants are able to absorb amino acids. Moreover, amino acid transporters have been shown to be present in both plant roots and in mycorrhizal hyphae. Although both mycorrhizal and non‐mycorrhizal plants appear to have a capacity for absorbing a range of organic N compounds, is this capacity realized in the field? Several lines of evidence show that plants are outcompeted by microorganisms for organic N sources. Such studies, however, have not addressed the issue of spatial and temporal separation between plants and microorganisms. Moreover, competition studies have not been able to separate uptake by symbiotic and non‐symbiotic microorganisms. Qualitative assessment of organic N uptake by plants has been performed with dual‐labelled glycine in several studies. These studies arrive at different conclusions: some indicate that plants do not absorb this organic N source when competing with other organisms in soil, while others conclude that significant fractions of amino acid N are absorbed as intact amino acid. These variable results may reflect species differences in the ability to absorb glycine as well as differences in experimental conditions and analytical techniques. Although theoretical calculations indicate that organic N might add significant amounts of N to plant N uptake, direct quantitative assessment of the fraction of plant N derived from uptake by organic N sources is a challenge for future research.     

Bryophytes attenuate anthropogenic nitrogen inputs in boreal forests

Productivity in boreal ecosystems is primarily limited by available soil nitrogen (N), and there is substantial interest in understanding whether deposition of anthropogenically derived reactive nitrogen (N_r) results in greater N availability to woody vegetation, which could result in greater carbon (C) sequestration. One factor that may limit the acquisition of N_r by woody plants is the presence of bryophytes, which are a significant C and N pool, and a location where associative cyanobacterial N‐fixation occurs. Using a replicated stand‐scale N‐addition experiment (five levels: 0, 3, 6, 12, and 50 kg N ha^?1 yr^?1; n=6) in the boreal zone of northern Sweden, we tested the hypothesis that sequestration of N_r into bryophyte tissues, and downregulation of N‐fixation would attenuate N_r inputs, and thereby limit anthropogenic N_r acquisition by woody plants. Our data showed that N‐fixation per unit moss mass and per unit area sharply decreased with increasing N addition. Additionally, the tissue N concentrations of Pleuorzium schreberi increased and its biomass decreased with increasing N addition. This response to increasing N addition caused the P. schreberi N pool to be stable at all but the highest N addition rate, where it significantly decreased. The combined effects of changed N‐fixation and P. schreberi biomass N accounted for 56.7% of cumulative N_r additions at the lowest N_r addition rate, but only a minor fraction for all other treatments. This ‘bryophyte effect’ can in part explain why soil inorganic N availability and acquisition by woody plants (indicated by their δ¹⁵N signatures) remained unchanged up to N addition rates of 12 kg ha^?1 yr^?1 or greater. Finally, we demonstrate that approximately 71.8% of the boreal forest experiences N_r deposition rates at or below 3 kg ha^?1 yr^?1, suggesting that bryophytes likely limit woody plant acquisition of ambient anthropogenic N_r inputs throughout a majority of the boreal forest.     

Classification of ultracontinental boreal forests in central Yakutia

Using the Braun-Blanquet approach, five associations of boreal forests were distinguished in central Yakutia, the most continental part of eastern Siberia. Ecological features of the syntaxa were explained with the use of the DCA ordination of 50 relevés. All available data from eastern Siberia were involved in the study for syntaxonomic analysis. Central Yakutian boreal forests were classified into two classes:Rhytidio-Laricetea sibiricae Korotkov etErmakov 1999 — ultracontinental light coniferous hemiboreal forests, andVaccinio-Piceetea Br.-Bl. inBr.-Bl. et al. 1939 — typical coniferous taiga forests of northern Eurasia. A new concept of higher syntaxonomic units of the classVaccinio-Piceetea in eastern Siberia has been developed. Three orders represent the diversity of taiga forests: (1)Cladonio-Vaccinietalia Kielland-Lund 1967 (with alliancesHieracio umbellati-Pinion sylvestris Anenkhonov etChytry 1998 andSaxifrago bronchialis-Pinion sylvestris all. nov.) — light coniferous boreal forests occurring in dry and moderately dry oligotrophic sites in various climatic sectors of Northern Eurasia; (2)Lathyro humilis-Laricetalia cajanderi ord. nov. (with alliancesAulacomnio acuminati-Laricion cajanderi all. nov. andRhododendro daurici-Laricion gmelinii all. nov.) — zonal boreal forests with xeric elements, which are typical of regions of northern Asia with cold, dry ultracontinental climate; (3)Ledo-Laricetalia cajanderi ord. prov. (with allianceLedo-Laricion cajanderi prov.) — North Eurasian boreal forests occurring in cold sites with excessive soil moisture, sometimes water-logged. Phytogeography and ecology of these orders are discussed in comparison with other regions of northern Asia.     

A new source of methane in boreal forests

Methane was found among the gases evolved during natural wood decay caused by bracket fungi in boreal forests. Methane was detected both in decaying wood and fungal fruiting bodies. A scheme of symbiotic association of wood-degrading fungi and anaerobic microorganisms providing the methanogenesis in the wood was proposed. The scale of mycogenic methane emission has to be consistent with the huge volume of decaying wood in boreal forest ecosystems.     

A new source of methane in boreal forests

Methane was found among the gases evolved during natural wood decay caused by bracket fungi in boreal forests. Methane was detected both in decaying wood and fungal fruiting bodies. A scheme of symbiotic association of wood-degrading fungi and anaerobic microorganisms providing the methanogenesis in the wood was proposed. The scale of mycogenic methane emission has to be consistent with the huge volume of decaying wood in boreal forest ecosystems.