Spatial distribution of fine-roots in boreal forests in eastern Sweden

Investigations were carried out in six forest types in areas surrounding two Swedish nuclear power plants (Forsmark and Laxemar). The aim of the investigation was to determine the spatial distribution of fine-root biomass (live), necromass (dead) and standing crop (live + dead) and to test the use of the live/dead ratio as a vitality criterion. Soil cores were taken to depths with insignificant amounts of roots. The total amount of fine-root biomass (<1 mm in diameter) of tree species in the soil profile was 267, 317 and 235 g m^?2 for the Forsmark and 137, 371 and 50 g m^?2 for the Laxemar sites. The related necromass was 119, 226 and 184 g m^?2 and 87, 245 and 271 g m^?2. The biomass in the humus layer was 47, 7 and 48% for the Forsmark and 34, 26 and 7% for the Laxemar sites, as a percentage of the total live + dead fine roots in the soil profile. The related necromass in the humus layer was 13, 2 and 30% for the Forsmark and 13, 2 and 28% for the Laxemar sites. The live/dead ratio decreased with depth for both tree— and field-layer species and seems to be a most powerful vitality criterion of fine roots.     

Vegetation dynamics and disturbance history in three deciduous forests in boreal Sweden

Abstract. Remaining deciduous forests in the Fennoscandian boreal landscape have high ecological value, and are considered as key components of the forest landscape as well as remnants of a former natural forest type. To improve our understanding of the formation of deciduous forests, we studied past disturbance regimes and vegetation dynamics in three deciduous forests in boreal Sweden using dendro‐ecology, pollen analysis and charcoal analysis. We identified three stages in the development of the studied stands. Firstly, the coniferous period (pre 1800), a long‐lasting period characterized by frequent fires, livestock grazing and extensive agriculture during which Pinus sylvestris was dominant. Secondly, the transformation period (1800 ‐ 1900), when logging removed most pines from the sites while fire and grazing continued. At the time of the last fire, the sites lacked a local seed source of pines, resulting in a post‐fire succession dominated by deciduous species with the capacity to disperse over long distances. Thirdly, the deciduous period (1900 ‐ present), with little or no disturbance from fire, grazing or logging. Thus, the present deciduous stands have their origins in a complex interaction between changes in fire regime, extensive land use patterns and logging, contrary to earlier simplified explanations. We conclude that the complexity of historical patterns of land use, vegetation dynamics and disturbance should be acknowledged in the future when selecting areas for nature conservation and developing models for ecologically oriented forestry.     

High occurrence of red-listed bryophytes and lichens in mature managed forests in boreal Sweden

Successful biodiversity conservation needs to be based on reliable data regarding the distribution and abundance of species. Red Lists summarize the status of rare and declining species. Here we present the first detailed survey (10 m × 10 m resolution) of nationally red-listed bryophytes and lichens at stand level in boreal forests. 30 managed forest production stands in boreal Sweden, soon to be harvested, were surveyed in detail regarding red-listed bryophytes and lichens. On the 180 ha totally investigated, 33 red-listed species were found, representing 35% of all forest red-listed bryophytes and lichens so far recorded from this part of Sweden. The mean number of red-listed species per stand was 10 and varied between 5 and 16. Excluding three species which were so common that, given the time-limit, it was not possible to record all of their occurrences, there were on average 6 number of observations ha⁻¹. The presence and density exceed those earlier found in ‘key habitats’, defined as hot-spots for red-listed species. Almost all observations were made on trees, with 48% on living trees and 52% on dead. Altitude, area and stem volume explained 67% of the red-list species density. The results highlight the importance of unprotected areas for rare and declining species and point to a hitherto unacknowledged high biodiversity value of mature managed boreal forest stands, shaped by a less intense forestry than today's. It also raises questions regarding the use of red-listed species to identify areas of importance to biodiversity. Objective and systematic surveys of red-listed species in the forest landscape, including younger stands, are urgent.Der erfolgreiche Schutz der Biodiversität muss auf verlässlichen Daten über die Verbreitung und Abundanz der Arten basieren. Rote Listen fassen den Status seltener und abnehmender Arten zusammen. Hier präsentieren wir die erste detaillierte Erfassung (10 m × 10 m Auflösung) von Bryophyten und Flechten der nationalen Rote Liste auf Bestandsniveau in borealen Wäldern. Im borealen Schweden wurden in 30 bewirtschafteten Produktionswälder, die bald geerntet werden sollen, die Bryophyten und Flechten der Roten Liste im Detail erfasst. Auf den insgesamt untersuchten 180 ha wurden 33 Arten der Roten Liste gefunden, die 35% aller Wald-Bryophyten und-Flechten der Roten Liste repräsentieren, die bisher in diesem Teil Schwedens erfasst wurden. Die mittlere Anzahl derRote-Liste-Arten pro Bestand war 10 und variierte zwischen 5 und 16. Unter Ausschluss von drei Arten, die so gewöhnlich waren, dass es bei dem gegebenen Zeitlimit nicht möglich war alle ihre Vorkommen zu erfassen, gab es im Durchschnitt 6 Beobachtungen ha⁻¹. Vorkommen und Dichten übertrafen die früher in “Schlüsselhabitaten” gefundenen, die als ‘hot spots’ bezeichnet werden. Fast alle Beobachtungen wurden auf Bäumen gemacht, mit 48% auf lebenden und 52% auf toten Bäumen. Die Höhe, das Gebiet und das Stammvolumen erklärten 67% der Dichte der Rote-Liste-Arten. Die Ergebnisse heben die Wichtigkeit nicht-geschützter Gebiete für seltene und abnehmende Arten hervor und weisen auf einen bisher nicht wahrgenommenen hohen Biodiversitätswert von alten bewirtschafteten borealen Waldbeständen hin, die von einer weniger intensiven Forstwirtschaft als der heutigen geformt wurden. Es stellen sich ebenso Fragen, die die Verwendung der Rote-Liste-Arten betreffen, um für die Biodiversität wichtige Gebiete zu identifizieren. Objektive und systematische Erfassungen der Rote-Liste-Arten in Waldlandschaften inklusive der jüngeren Bestände sind dringend.     

The hunting habitats of Goshawks Accipiter gentilis in boreal forests of central Sweden

Radio-tracking was used to study the habitat utilization of hunting Goshawks Accipiter gentilis in central Swedish boreal forest. Data were collected during September-June, in 1977–1981. There was no major difference between autumn, winter or spring in any aspect of habitat preference, despite seasonal differences in prey composition. Despite the strong reversed sexual size-dimorphism, no habitat segregation between the sexes was found. It is argued that this is due to the lack of prey segregation between the sexes. Goshawks showed a strong preference for mature forest, which was chosen twice as often as would be expected from its prevalence in the environment, whereas the younger successional stages of the forest were under-used. Goshawks preferred large habitat patches, and showed no major preference with respect to tree species composition of the forest. Kills made by radio-tagged Goshawks showed the same habitat- and patch-size distribution as did Goshawk locations in general. It is argued that the profitability of different hunting habitats, and thus habitat preferences, is not only determined by prey density in the habitats, but also by other habitat features influencing the Goshawk's ability to hunt there. Winter home-ranges averaged 5700 ha, considerably larger than Goshawk ranges recorded in areas with richer prey supply.     

Summer movements,predation and habitat use of wolves in human modified boreal forests

Grey wolves (Canis lupus), formerly extirpated in Finland, have recolonized a boreal forest environment that has been significantly altered by humans, becoming a patchwork of managed forests and clearcuts crisscrossed by roads, power lines, and railways. Little is known about how the wolves utilize this impacted ecosystem, especially during the pup-rearing summer months. We tracked two wolves instrumented with GPS collars transmitting at 30-min intervals during two summers in eastern Finland, visiting all locations in the field, identifying prey items and classifying movement behaviors. We analyzed preference and avoidance of habitat types, linear elements and habitat edges, and tested the generality of our results against lower resolution summer movements of 23 other collared wolves. Wolves tended to show a strong preference for transitional woodlands (mostly harvested clearcuts) and mixed forests over coniferous forests and to use forest roads and low use linear elements to facilitate movement. The high density of primary roads in one wolf’s territory led to more constrained use of the home territory compared to the wolf with fewer roads, suggesting avoidance of humans; however, there did not appear to be large differences on the hunting success or the success of pup rearing for the two packs. In total, 90 kills were identified, almost entirely moose (Alces alces) and reindeer (Rangifer tarandus sspp.) calves of which a large proportion were killed in transitional woodlands. Generally, wolves displayed a high level of adaptability, successfully exploiting direct and indirect human-derived modifications to the boreal forest environment.     

Estimating net primary production of boreal forests in Finland and Sweden from field data and remote sensing

We calculated annual mean stem volume increment (AMSVI) and total litter fall to produce forest net primary production (NPP) maps at 1‐km² and half‐degree resolutions in Finland and Sweden. We used a multi‐scale methodology to link field inventory data reported at plot and forestry district levels through a remotely sensed total plant biomass map derived from 1‐km² AVHRR image. Total litter fall was estimated as function of elevation and latitude. Leaf litter fall, a surrogate for fine root production, was estimated from total litter fall by forest type. The gridded NPP estimates agreed well with previously reported NPP values, based on point measurements. Regional NPP increases from northeast to southwest. It is positively related to annual mean temperature and annual mean total precipitation (strongly correlated with temperature) and is negatively related to elevation at broad scale. Total NPP (TNPP) values for representative cells selected based on three criteria were highly correlated with simulated values from a process‐based model (CEVSA) at 0.5°× 0.5° resolution. At 1‐km² resolution, mean above‐ground NPP in the region was 408 g/m²/yr ranging from 172 to 1091 (standard deviation (SD) = 134). Mean TNPP was 563 (252 to 1426, SD = 176). Ranges and SD were reduced while the mean values of the estimated NPP stayed almost constant as cell size increased from 1‐km² to 0.5°× 0.5°, as expected. Nordic boreal forests seem to have lower productivity among the world boreal forests.     

Landscape ecology of boreal forests

It is becoming increasingly clear that densities and dynamics of various organisms often cannot be understood from the processes occurring within separate habitat patches. Effects from surrounding areas also have to be considered; a landscape perspective has to be employed. Habitat mosaics affect diversity and dynamics both in pristine and managed boreal landscapes. There is increasing interest in the landscape ecology of boreal forests and many important processes have been identified as being driven or affected by environmental heterogeneity. Comparisons between ecological processes in original and managed boreal landscapes are urgently needed.     

Red-listed lichens in Sweden: habitats, threats, protection, and indicator value in boreal coniferous forests

The three major threats to lichens in Sweden; forestry, air pollution, and structural changes in the agricultural landscape, are outlined. Forestry is the most serious. The majority of the red-listed lichens in forests are found in productive forests with a continuity in tree growth, but nearly 70% of the protected area in Sweden consists of alpine landscapes. Examples of important habitats with red-listed lichens are described, e.g. Alnus glutinosa marshes, alvar-vegetation, broad-leaved deciduous forests and different Picea abies forests. The use of selected indicator species of lichens and macrofungi (mainly red-listed species) in boreal coniferous forests provides information on continuity in tree growth of an area, and on earlier influences of forestry. Examples of indicator species for different types of forest are presented and discussed. In spite of the Swedish right of common accesso, collecting of lichens is forbidden without permission from the land-owner. Collecting in protected areas is allowed only with a permit obtained from the authorities.     

The boreal forests of north-eastern Eurasia

The distribution, diversity and structure of boreal forests of monsoon Asia are reviewed. The boreal zone is circumscribed by the isotherms of 15°C and 45°C of Kira's Warmth Index (WI) and includes the territory of the Amur basin, the seashores of the Sea of Okhotsk, Kamchatka, Sakhalin and the Kuril islands. Boreal forests also occur on the mountains of Hokkaido, Honshu, Korea and NE China. The oceanic sectors of the boreal zone are composed mainly of birch forests, the marine sectors of spruce-fir forests and the continental sectors of larch and pine forests. The upper limit of the boreal forests ascends from the lowest level in the North to ca. 1500 m a.s.l. in the South of the zone, and reaches above 2500 m in the mountains of the temperate zone. In the boreal zone the secondary forests are common. They form after fires and cutting. Volcanism is an essential factor in the forest dynamics in the oceanic part of the region. It is suggested that as a result of global warming one may expect a strong alteration in vegetation pattern, especially in the northern and mountain areas.Paper presented at the International Symposium on Global Change Impacts on Terrestrial Ecosystems in Monsoon Asia (TEMA) Waseda University, Tokyo, 4–6 September 1993.     

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.     

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.     

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.     

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.     

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.     

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.