Bird community energetics in a boreal coniferous forest

The annual minimum energy consumption of the bird community was 2524 × 10³ kcal km⁻², of which 44% was consumed by wintering species and 73% by passerines. The daily energy consumption was in summer 14–16 × 10³ and in winter 1–2 × 10³ kcal km⁻². In spruce forests and in afforested swamps birds required approximately 0.12% of the net primary production. Their total annual energy consumption was covered by invertebrates (59%), vertebrates (2%) and vegetable matter (39%); the food derived from the ground (55%), from trees (44%) and from the air (1%). Arboreal insectivorous passerines, ground passerines and gallinaceous birds were the most important ecological guilds. Among passerines existence metabolism accounted for 73% of the annual energy consumption, extra activity for 17%, breeding activity for 1%, moult for 4% and nestlings for 4%.     

Changes in the occurrence and abundance of plant species in a Norwegian boreal coniferous forest, 1988–1993

Økland, R. H. 1995. Changes in the occurrence and abundance of Dlants sDecies in a Norwegian boreal coniferois forest, 1988–1993. — Nord. J. Bot. 15: 415–438. Copenhagen. ISSN 0107–055X.
Changes in the occurrence and abundance of vascular plants, bryophytes and lichens in a S Norwegian old-growth coniferous forest were studied, using 1 m² plots divided into 16 subplots. Fifty plots were analyzed for six consecutive years, 199 plots were analyzed at the beginning and the end of the study (in 1988 and 1993). Nonparametric tests were performed for combinations of species, data type (occurrence or abundance), forest type (spruce or pine), and time period (one-year or five-year). Results are interpreted by use of species persistence data, and by detailed comparisons with European long-term and experimental studies. Abundance changes in the pine forest are probably due to short-term (between-year) climatic fluctuations, notably drought. Some effects of the observers are also seen. True five-year trends were found in the spruce forest: (1) Deschampsia flexuosa (increase), (2) several vascular plant species with preference for richer sites, notably Oxalis acetosella (decrease), and (3) most bryophyte species (increase). True trends for vascular plants are discussed in relation to deposition of airborne nitrogen and soil acidification, while the increase of bryophytes probably is due to particularly favourable growth conditions in the five-year period.     

High diversity of arbuscular mycorrhizal fungi in a boreal herb-rich coniferous forest

* Here, the diversity of arbuscular mycorrhizal (AM) fungi was determined in a boreal herb-rich coniferous forest in relation to environmental variables. * Root samples of five plant species (Fragaria vesca, Galeobdolon luteum, Hepatica nobilis, Oxalis acetosella and Trifolium pratense) were analysed from stands differing in age and forest management intensity. * Thirty-four Glomeromycota taxa (small-subunit ribosomal RNA gene (SSU rDNA) sequence groups) were detected from 90 root samples (911 clones), including eight new taxa. Sequence groups related to Glomus intraradices were most common (MO-G3 and MO-G13). Samples of H. nobilis were colonized by more AM fungal taxa (3.68 +/- 0.31) than those of O. acetosella (2.69 +/- 0.34), but did not differ significantly in this respect from those of F. vesca (3.15 +/- 0.38). Effects of forest management, host plant species (except above) or season on the number or composition of fungal taxa in root samples were not detected, and neither were they explained by environmental variables (vegetation, soil and light conditions). * This is the most taxon-rich habitat described to date in terms of root-colonizing Glomeromycota. The data demonstrate the importance of temperate coniferous forests as habitats for AM fungi and plants. Lack of obvious fungal community patterns suggests more complex effects of biotic and abiotic factors, and possibly no adverse effect of common forest management practices on AM fungal diversity.     

Crustose lichens as indicators of forest continuity in boreal coniferous forests

The crustose lichen flora of fifteen areas in boreal coniferous forests from southern Sweden to Lapland was surveyed. Many of these areas are included in a national programme for monitoring long-term environmental change, the PMK programme. In all 190 species were recorded, 129 of them occurring on bark of conifers and lignum. Some species were found exclusively in forests with a long continuity. By using ordination and hierarchical classifications further species with similar habitat requirements were identified. An Indicator Species Index of Forest Continuity (ISIFC) was designed. Only two of the twenty index species occur on deciduous trees. The ISIFC was highly correlated with forest continuity and also with occurrence of species listed as threatened. Correlations with macroclimatic variables are considered sampling artifacts, and microclimatic factors are suggested to be of primary importance for species occurring in forests with a long continuity.     

Effects of simulated N deposition on understorey vegetation of a boreal coniferous forest

1. We report the short-term effects of simulated nitrogen (N) deposition on the understorey of a boreal forest in northern Sweden. Doses of ¹⁵N double-labelled NH₄NO₃ (0·5, 12·5, 25·0 and 50·0 kg N ha^–1) were applied to 1 m² plots in the summer of 1995 and plants were harvested the following autumn.
2. No significant treatment effects were found in either above- or below-ground biomass which was distributed as follows: the ericaceous shrub Vaccinium myrtillus contributed 76%, the grass Deschampsia flexuosa 4%, and the bryophytes Dicranum majus and Pleurozium schreberi 20%.
3. The recovery of applied N in these species was 24, 27, 27 and 32% of the 0·5, 12·5, 25·0 and 50·0 kg N ha^–1 applications, respectively, and thus the recovery increased with the N dose.
4. In the 0·5 kg N ha^–1 treated plots, the highest concentrations of fertilizer-derived ¹⁵N were found in the bryophytes, while in plots given the three higher N applications, leaves of D . flexuosa had the highest concentrations.
5. N application resulted in elevated concentrations of free amino acid N, which indicate increased storage of N in plant tissues. Furthermore, the N application resulted in increased damage to V . myrtillus by natural enemies. The severity of disease caused by two foliar parasitic fungi showed a clear numerical response to N treatment, as did the amount of herbivory as a result of Lepidoptera larvae.     

Sixty years of vegetation dynamics in a south boreal coniferous forest in southern Norway

Abstract. Vegetation data from permanent plots were collected in 1931, 1961 and 1991 in a south boreal forest 20 km north of Oslo in southern Norway. Major changes were found in the vegetation composition during those 60 years. The main changes were a reduction in the frequency of species and the frequency of joint occurrences of vascular species such as Andromeda polifolia, Calluna vulgaris, Cornus suecica, Eriophorum vaginatum, Maianthemum bifolium, Melampyrum pratense, Trientalis europaea, Vaccinium uliginosum and V. oxycoccus, and mosses, e.g. Dicranum fuscescens, Hylocomium splendens, Pleurozium schreberi, Ptilidium ciliare and Ptilium crista-castrensis. The observed changes were interpreted as being induced by internal processes e.g. notably a long-term change from paludified forest to mesic forest. In particular the growth of Picea abies seems to be a main driving force. The dominance of Picea abies and Vaccinium myrtillus appears to have rendered the conditions more unfavourable for other species. A doubling of the living stem biomass of P. abies during the last 67 yr shows that this old-growth forest has not yet reached a steady state. It was demonstrated that species such as Deschampsia flexuosa and Molinia caerulea did not increase in frequency in response to nitrogen deposition, as has occurred elsewhere in northern Europe. pH in the humus layer increased with 0.2 unit from 1961 to 1991. The results of this study indicate that protection from logging has initiated the reduction of species in the field layer and bottom layer. This study questions if monitoring of forest vegetation should be restricted to protected forests as is the practice in Scandinavia today. We recommend that also areas with some kind of selective cutting will be used for monitoring of forest vegetation.     

Sulfur speciation and distribution in soils and aboveground biomass of a boreal coniferous forest

Major sulfur pools are quantified in soils and aboveground biomass of a coniferous boreal forest. Total ecosystem S averages 1395 kg·ha⁻¹ of which 98% is found in the soil, with 89% being in the mineral horizons. Organic S dominates in soil, tree parts (trunks, branches + foliage, roots) and litterfall, ranging from 77 to 99% of total S concentration. Carbon-bonded S, ester sulfate and SO₄-S in soil profiles range respectively from 51–68%, 29–37% and 1–14% of total S concentrations and account respectively for 57, 33 and 10% of total S on an areal basis. Adsorbed SO₄ accounts for 82% of total SO₄, and can be predicted from Al bound to organic matter, amorphous Al and pH (r^{2 = 0.81}). There is a strong relationship between % carbon and carbon-bonded S in 4 of the 5 soil horizons studied which represent over 95% of the total soil organic matter, whereas ester sulfate is related to % carbon in 3 soil horizons representing only 37% of the soil organic matter. An analysis of atmospheric S loading and S data for 10 forested sites in Europe and North America suggests that the size of the organic S pool in forested systems is independent of atmospheric loading.     

Biodiversity of wood-inhabiting fungi in a boreal coniferous forest in Ser-Trendelag County, Central Norway

A survey of the patterns of lignicolous fungi to record occurrence of fruitbodies on fallen logs of Picea abies . has been performed. Ninetythree forest stands with different ages. ecology, and histories of management in Ser-Trendelag County, Central Norway were investigated. In each stand an investigation plot of 40 m diameter was established, in which five logs were randomly selected. Environmental variables of the logs and plots and geographical positions of the plots were recorded. The numerical methods used include detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA).
A total of 140 species of Aphyllophorales, Agaricales with eccentrically attached caps, heterobasiodomycetes, and prominent ascomycetes and deuteromycetes were registered. More than one third of the species were only found once. The main gradient in the data set was shown to be related to decay. The second gradient was related to size of the logs and some forest management variables (in addition to decay). Of the variation in composition of the species, 13.8 % can be ascribed to solely environmental variance, 6 % to solely spatial structure, and only 0.8 % to the combination of the environment and spatial component of variation. The results are discussed in the light of wood decay, forest management, and landscape fragmentation.     

Patterns of bryophyte associations at different scales in a Norwegian boreal spruce forest

Abstract. Patterns of associations between 36 bryophytes and their relationships with trends in α- and β-diversity were studied at five spatial scales (from 1 m² to 1/256 m²) in a Norwegian boreal spruce forest. The range and dispersion of α-diversity values in the data were significantly higher than in comparable model data sets, indicating varation from favourable to unfavourable conditions for bryophytes at all scales, particularly with increasing cover of the upper layers. The number of positive associations was significantly higher than predicted from a random distribution, for all sample plot sizes except the smallest. For the most frequent species, this number decreased with decreasing sample plot size. The excess of positive associations was due to the presence of (1) α-diversity trends, as demonstrated by a stochastic simulation, and (2) β-diversity (variation along environmental gradients). A microscale coenocline from dominance of large forest floor mosses to small liverworts, is added to two broad-scale coenoclines demonstrated earlier. Facilitation - positive density-dependence caused by more favourable moisture conditions within dense stands - is discussed as a possible third cause of excess positive associations. The number of significant negative associations was generally low, and deviated neither from theoretical values assuming random distribution of species, nor from predicted values accounting for α-diversity trends. The low proportion of negative associations, even in the presence of β-diversity trends, suggests that interspecific competition is not important in this vegetation. Several alternatives to competition are discussed; (1) static mechanisms for avoidance of competition, (2) mobility, and (3) the importance of density-independent mortality, in particular due to fine-scale disturbance. Simulation studies for assessing the effects of α-diversity trends on species associations are emphasized.     

Colonization of recent coniferous versus deciduous forest stands by vascular plants at the local scale

Questions: 1. Are there differences among species in their preference for coniferous vs. deciduous forest? 2. Are tree and shrub species better colonizers of recent forest stands than herbaceous species? 3. Do colonization patterns of plant species groups depend on tree species composition? Location: Three deciduous and one coniferous recent forest areas in Brandenburg, NE Germany. Methods: In 34 and 21 transects in coniferous and deciduous stands, respectively, we studied the occurrence and percentage cover of vascular plants in a total of 150 plots in ancient stands, 315 in recent stands and 55 at the ecotone. Habitat preference, diaspore weight, generative dispersal potential and clonal extension were used to explain mechanisms of local migration. Regression analysis was conducted to test whether migration distance was related to species’ life‐history traits. Results: 25 species were significantly associated with ancient stands and ten species were significantly more frequent in recent stands. Tree and shrub species were good colonizers of recent coniferous and deciduous stands. In the coniferous stands, all herbaceous species showed a strong dispersal limitation during colonization, whereas in the deciduous stands generalist species may have survived in the grasslands which were present prior to afforestation. Conclusions: The fast colonization of recent stands by trees and shrubs can be explained by their effective dispersal via wind and animals. This, and the comparably efficient migration of herbaceous forest specialists into recent coniferous stands, implies that the conversion of coniferous into deciduous stands adjacent to ancient deciduous forests is promising even without planting of trees.     

Small scale pH buffering in organic horizons of two boreal coniferous forest stands

Factors behind the small-scale variaton of pH were examined in O horizons (humus layers) developed under two stands of Picea abies (L.) Karst. (Site F and K) by combining data on the composition of the cation exchange complex with data from titrations of corresponding H⁺-saturated samples. Cations extractable in 0.5 M CuCl₂ (S=cmol_c kg^–1 [2Ca+2Mg+2Mn+K+Na]), aluminium extractable in 1.0 M KCl (Al_e=cmol_c kg^–1 [3Al]) and in 0.5 M CuCl₂ (Al_org=cmol_c kg^–1 [3Al]), as well as pH measured in 0.01 M CaCl₂ (pH_Ca) were analysed in one-cm-layers of 13 O horizon cores at each site. Composite samples representing each of the one-cm-layers at each site, as well as samples with two different levels of Al saturation at Site K, were H⁺ saturated and titrated with NaOH to chosen end points of pH_Ca=4.0 and 5.5 in a 0.01 M CaCl₂ ionic medium. The Acid Neutralisation Capacity (ANC) was estimated as the amount of base needed to increase pH_Ca of the composite H⁺-saturated samples to the mean pH_Ca of the corresponding natural samples. The ANC was found to be similar in magnitude or higher than the amount of sites binding S+Al_e, which suggests that 1.0 M KCl exchangeable Al ions are nonacidic in acid O horizons. The relative contribution from i) the capacity of acidic functional groups, ii) their acid strength and iii) their degree of neutralisation to differences in pH_Ca between sites, among cm-layers and between samples with different levels of Al saturation were estimated from titration curves adjusted to hold two out of three factors (i, ii and iii) to be constant. The degree of neutralisation explained most of the differences in pH_Ca between the two sites, as well as between samples with different levels of Al saturation at Site K. The pH_Ca decrease by depth at site F was, however, partly explained by an increasing acid strength. The study emphasizes the importance of examining not only changes in the degree of neutralisation, but also changes in the acid strength and the capacity of buffering functional groups before conclusions about causes behind acidification processes can be made. Difficulties of accurately estimating the degree of neutralisation (base saturation) of acidic functional groups from the composition of adsorbed cations, owing to the unknown acidity of adsorbed Al, was also demonstrated.     

林火干扰对北方针叶林林下植被的影响

林下植被在北方针叶林植被群落中的物种多样性最高, 且具有较高的生物量周转率和地上部分净初级生产力, 对北方针叶林生态系统功能起着重要作用。火干扰是决定北方针叶林林下植被结构与功能的一个重要景观过程。该文综述了火干扰是如何通过与地形、火前林冠组成的交互作用而影响环境资源和林下植被的。最近的研究表明: 林下植被能够影响火后树木更新苗的定植、重建速率及森林演替轨迹; 林下植被还会通过影响元素的生物地球化学过程(凋落物降解和养分循环)影响林下环境资源的数量与异质性。因此, 研究火后初期北方针叶林林下植被的动态变化, 对于物种多样性保护和森林管理具有重要意义。     

Response of photosynthetic carbon gain to ecosystem retrogression of vascular plants and mosses in the boreal forest

In the long-term absence of rejuvenating disturbances, forest succession frequently proceeds from a maximal biomass phase to a retrogressive phase characterized by reduced nutrient availability [notably nitrogen (N) and phosphorus (P)] and net primary productivity. Few studies have considered how retrogression induces changes in ecophysiological responses associated with photosynthetic carbon (C) gain, and only for trees. We tested the hypothesis that retrogression would negatively impact photosynthetic C gain of four contrasting species, and that this impact would be greater for vascular plants (i.e., trees and shrubs) than for non-vascular plants (i.e., mosses). We used a 5,000-year-old chronosequence of forested islands in Sweden, where retrogression occurs in the long-term absence of lightning-ignited wildfires. Despite fundamental differences in plant form and ecological niche among species, vascular plants and mosses showed similar ecophysiological responses to retrogression. The most common effects of retrogression were reductions in photosynthesis and respiration per unit foliar N, increases in foliar N, δ(13)C and δ(15)N, and decreases in specific leaf areas. In contrast, photosynthesis per unit mass or area generally did not change along the chronosequence, but did vary many-fold between vascular plants and mosses. The consistent increases in foliar N without corresponding increases in mass- or area-based photosynthesis suggest that other factor(s), such as P co-limitation, light conditions or water availability, may co-regulate C gain in retrogressive boreal forests. Against our predictions, traits of mosses associated with C and N were generally highly responsive to retrogression, which has implications for how mosses influence ecosystem processes in boreal forests.     

Amino acid uptake: a widespread ability among boreal forest plants

Amino acids constitute a potentially important source of nitrogen for plants in boreal forest ecosystems. Accordingly, it may be suggested that distinct plant species differing abilities to take up amino acids constitutes an important factor in determining plant ecosystem composition. Using GC-MS and isotopically labelled amino acids, we measured the simultaneous uptake of 15 different amino acids by 31 common boreal forest plant species. The results from this study show that all plant species tested, representing a wide variety of plant types, have the ability to take up amino acids from an incubation solution. Furthermore, uptake rates were unrelated to mycorrhizal associations as well as habitat soil amino acid concentrations and plant nitrogen availability dependence as measured by Ellenberg nitrogen indicator values. These results suggest that mycorrhiza is of minor importance for discrete plant amino acid uptake rates and further points out the potential importance of amino acids to plant nitrogen nutrition in boreal forest ecosystems.     

Exotic vascular plant invasiveness and forest invasibility in urban boreal forest types

The riverine forests of the northern city of Edmonton, Alberta, Canada display strong resilience to disturbance and are similar in species composition to southern boreal mixedwood forest types. This study addressed questions such as, how easily do exotic species become established in urban boreal forests (species invasiveness) and do urban boreal forest structural characteristics such as, native species richness, abundance, and vertical vegetation layers, confer resistance to exotic species establishment and spread (community invasibility)? Eighty-four forest stands were sampled and species composition and mean percent cover analyzed using ordination methods. Results showed that exotic tree/shrub types were of the most concern for invasion to urban boreal forests and that exotic species type, native habitat and propagule supply may be good indicators of invasive potential. Native forest structure appeared to confer a level of resistance to exotic species and medium to high disturbance intensity was associated with exotic species growth and spread without a corresponding loss in native species richness. Results provided large-scale evidence that diverse communities are less vulnerable to exotic species invasion, and that intermediate disturbance intensity supports species coexistence. From a management perspective, the retention of native species and native forest structure in urban forests is favored to minimize the impact of exotic species introductions, protect natural succession patterns, and minimize the spread of exotic species.     

Dynamics of understory vegetation in an old-growth boreal coniferous forest, 1988–1993

Abstract. Understorey vegetation changes in a South Norwegian old-growth coniferous forest were studied between 1988 and 1993 in 200 1-m² vegetation plots. Our aims were to quantify the amount of between-year compositional change, and to elaborate the environmental basis for long-term vegetation change, including the previously identified gradient structure with a major gradient related to topography (and soil nutrient status and soil depth) and a minor gradient reflecting paludification and canopy coverage. Species richness (yearly mean and cumulative species number) and change in species richness differed between vascular plants and cryptogams, and between forest types. The number of vascular plant species decreased in pine forest in dry years; bryophyte species number increased in spruce forest. Statistically significant vegetation change, as tested by constrained ordination (CCA) with time as the constraining variable, is demonstrated for most one-year periods and for the five-year period in most forest types. Vegetation change along identified gradients, measured as plot displacement along DCA ordination axes, also occurred. The magnitude of year-to-year vegetation change was related neither to forest type nor to one-year period; different responses to climatic and environmental change were observed in each forest type. The largest average displacement observed, from medium-rich spruce forest towards poor spruce forest, was interpreted as a long-term trend. Humus-layer pH decreased by ca. 0.25 units from 1988 to 1993, most strongly in medium-rich spruce forest where exchangeable Ca decreased and Al and Mn increased strongly. Our study supports the hypothesis that vascular plants show a long-term and broad-scale response to soil acidification. Change in bryophyte composition is linked to some very long growing-seasons. Detailed analysis of short-term vegetation dynamics enhances the interpretation of long-term changes and stresses the complementarity of univariate and multivariate methods in the analysis of vegetation change.     

Gross nitrogen transformations in soils from uncut and cut boreal upland and peatland coniferous forest stands

Gross and net nitrogen (N) ammonification and nitrification were measured in soils from an uncut and recently cut upland and peatland conifer stand in northwestern Ontario, Canada. Rates of gross total inorganic N immobilization were similar to gross mineralization, resulting in low net mineralization rates in soils from all four upland and peatland conifer stands. Gross ammonification rates were variable but similar in soils from uncut and cut peatland hollows (18–19mgNkg^–1day^–1) and upland forest floor soils (14–19mgNkg^–1day^–1). Gross ammonium ( ) immobilization rates were also variable but similar to ammonification rates. Median gross nitrification rates were within 0–2mgNkg^–1day^–1 in soils from all four upland and peatland cut and uncut stands, although rates were consistently higher for the soils from the cut stands. Large variability in gross nitrification rates were observed in peatland soils, however the highest gross nitrification rates were measured in saturated peatland soils. Net rates remained low in the soils from all four stands due to high nitrate ( ) immobilization and very fast turnover (<0.2 day). Our results suggest that potential losses may be negated by high immobilization in uncut and cut boreal forest stands. This study reveals the potential for the interaction of N production and consumption processes in regulating N retention in upland and peatland conifer forests, and the resilience of the boreal forest to disturbance.     

Element concentrations in boreal,coniferous forest humus layers in relation to moss chemistry and soil factors

Concentrations of 15 elements (Al, Ca, Cd, Cr, Cu, Fe, K, Mg, Mn, Na, Ni, P, Pb, S and Zn) in the current annual growth increments of the ectohydric feather moss, Hylocomium splendens, were compared with those in the humus layer (Of+Oh) of coniferous forests at 353 sites throughout Finland. Moss concentrations in the moss increments were considered measures of atmospheric deposition of these elements. Concentrations in both moss and humus layer samples were negatively correlated with latitude, except Mn (moss) and Cr and Ni (humus layer), indicating a direct effect of deposition. Spearman correlations between concentrations in moss and humus layer samples were significant (P<0.05) for all elements; that for Pb concentrations was the highest (r_s=0.60) while those for the other elements were ≤0.48. Using thickness and organic matter content of the humus layer, clay content of the mineral soil (5–20 cm layer) and the concentration in the moss explained between 8% (Cd) and 44% (Cr, Mg) of the variation in the concentrations in the humus layer samples. It was concluded that, for most elements, concentration in the humus layer are a poor indicator of current atmospheric deposition levels, at least in relatively unpolluted regions such as Finland.