1. |
- Öberg, Lisa, 1958-
(författare)
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Treeline dynamics in short and long term perspectives : observational and historical evidence from the southern Swedish Scandes
- 2013
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Treelines in high-mountain regions are constrained by heat deficiency, although the working mechanisms are still not entirely understood. Observational and paleoecological studies on treeline performance may contribute to increased understanding of the treeline phenomenon in general. The present thesis addresses elevational shifts of alpine treelines in the Swedish Scandes. By various analytical tools, the studies embrace widely different temporal scales.The concept treeline refers to the elevation (m a.s.l.) at a specific site of the upper individual tree of a certain tree species, at least 2 m tall. All the principal tree species in the Scandes are concerned, i.e. mountain birch (Betula pubescens ssp. czerepanovii), Norway spruce (Picea abies) and Scots pine (Pinus sylvestris).Paper I deals with regional treeline dynamics at more than 100 sites over the past 100 years. Concurrent with temperature rise by c. 1.4 °C over the same period, maximum treeline advances of all species amount to about 200 m. Thus, under ideal conditions, treelines respond in close equilibrium with air temperature evolution. However, over most parts of the landscape, treeline upshifts have been much smaller than 200 m, which relates to the combined action of geomorphology, wind, snow distribution and soil depth. After 1975, the birch has lost its role as the most rapidly advancing tree species, being superseded by pine and spruce.Paper II is a short-term (2005/2007-2010/2011) study of mountain birch treeline performance along a regional maritimity-continentality gradient. Upshift by 3.0 yr-1 in the maritime part of the gradient contrasts to retreat by 0.4 m yr-1 in the continental part. In the latter area, earlier and more complete melting of late-lying snow patches has seemingly progressed to a state when soil drought sets back the vigour of existing birches and precludes sexual regeneration and upslope advance of the treeline. In the maritime area, extensive and deep snow packs still exist above the treeline and constrain its position, although some release is taking place in the current warm climate.Paper III explores treeline change by phenotypic transformation of old-established stunted and prostrate spruce individuals (krummholz) growing high above the treeline and is based on analyses of radiocarbon-dated megafossils, preserved in the soil underneath clonal groups of spruce. Living spruce clones, which in some cases may date back to the early Holocene (9500 cal. yr BP), suggests that spruce immigrated from “cryptic” ice age refugia much closer to Scandinavia than conventionally thought. As the krummholz form presupposes open and windy habitats, it is inferred that permanently open spots prevailed in the high-mountain landscape even during periods when treelines in general were much higher than today.Paper IV reports radiocarbon dates of wood samples, retrieved from newly exposed glacier forefields at three main sites, located high above the modern treelines and embracing the entire Swedish Scandes. It appears that pine colonized early emerging nunataks already during the Late Glacial. Around 9600-9500 cal. yr BP a first massive wave of tree establishment, birch and pine, took place in “empty” glacier cirques. Both species grew 400-600 m above their present day treeline position and accordingly, the summer temperatures may have been 3.5 °C warmer than present (uncorrected for land uplift). During the entire interval 9600 to 4400 cal. yr BP, birch prospered 100-150 m above the uppermost pines. In response to Neoglacial cooling, treelines of both birch and pine descended until their final disappearance from the record 4400 and 5900 cal. yr BP, respectively. Thereafter, these habitats experienced increased snow accumulation and glacier inception.
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2. |
- Öberg, Lisa, 1958-
(författare)
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Treeline dynamics in short and long term perspectives : observational and historical evidence from the southern Swedish Scandes
- 2010
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Licentiatavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Against the background of past, recent and future climate change, the present thesis addresses elevational shifts of alpine treelines in the Swedish Scandes. By definition, treeline refers to the elevation (m a.s.l.) at a specific site of the upper trees of a specific tree species, at least 2 m tall.Based on historical records, the first part of the thesis reports and analyzes the magnitude of treeline displacements for the main trees species (Betula pubescens ssp. czerepanovii, Picea abies and Pinus sylvestris) since the early 20th century. The study covered a large and heterogeneous region and more than 100 sites. Concurrent with temperature rise by c. 1.4 °C over the past century, maximum treeline advances of all species amount to about 200 m. That is virtually what should be predicted from the recorded temperature change over the same period of time. Thus, it appears that under ideal conditions, treelines respond in close equilibrium with air temperature evolution. However, over most parts of the landscape, conditions are not that ideal and treeline upshifts have therefore been much smaller. The main reason for that discrepancy was found to be topoclimatic constraints, i.e. the combined action of geomorphology, wind, snow distribution, soil depth, etc., which over large parts of the alpine landscape preclude treelines to reach their potential thermal limit.Recorded treeline advance by maximum 200 m or so over the past century emerges as a truly anomalous event in late Holocene vegetation history.The second part of the thesis is focused more on long-term changes of treelines and one specific and prevalent mechanism of treeline change. The first part of the thesis revealed that for Picea and Betula, treeline shift was accomplished largely by phenotypic transformation of old-established stunted and prostrate individuals (krummholz) growing high above the treeline. In obvious response to climate warming over the past century, such individuals have transformed into erect tree form, whereby the treeline (as defined here) has risen. As a means for deeper understanding of this mode of positional treeline change, extant clonal spruces, growing around the treeline, were radiocarbon dated from megafossil remains preserved in the soil underneath their canopies. It turned out that Picea abies in particular may attain almost eternal life due to its capability for vegetative reproduction and phenotypic plasticity. Some living clones were in fact inferred to have existed already 9500 years ago, and have thus persisted at the same spot throughout almost the entire Holocene. This contrasts with other tree species, which have left no living relicts from the early Holocene, when they actually grew equally high as the spruce. Thereafter they retracted by more than 300 m in elevation supporting that also on that temporal scale, treelines are highly responsive to climate change.The early appearance of Picea in the Scandes, suggests that Picea “hibernated” the last glacial phase much closer to Scandinavia than earlier thought. It has also immigrated to northern Sweden much earlier than the old-established wisdom.The experiences gained in this thesis should constitute essential components of any model striving to the project landscape ecological consequences of possible future climate shifts.
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3. |
- Öhmark, Sara, 1980-
(författare)
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Winter browsing by moose and hares in subarctic birch forest : Scale dependency and responses to food addition
- 2015
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Despite their difference in body size and morphology, the moose (Alces alces) andthe mountain hare (Lepus timidus) sustain themselves during winter on similar plantspecies and plant parts in in subarctic environments, namely apical twigs ofmountain birch (Betula pubescens ssp. czerepanovii). Herbivores must select areas anditems of food that provide sufficient intake rates and food nutritional quality whilebalancing this against their intake of dietary fiber and potentially detrimental plantsecondary metabolites. This selection takes place simultaneously at multiple spatialscales, from individual plants and plant parts to patches of food and parts of the wider landscape. While the herbivores must consider their need for food to sustaindaily activities, for body growth and reproduction it is also necessary to avoid predators and harsh environmental conditions. For managers, an understanding of key factors for animal foraging distributions is pivotal to reach intended goals ofmanagement and conservation plans. Knowledge in this area is also important formodels to make accurate predictions of foraging responses of herbivores to resource distributions. The mountain birch forest displays a naturally heterogeneous distribution of trees and shrubs which presents herbivores with a challenge to findgood feeding areas. In an investigation of the spatial distribution of moose browsing on birch and willows (Salix spp.) in two winter seasons separated in time by 14 years,it was found that moose browsing patterns in 1996 were correlated to those observed in 2010. It was also found that moose browsing was spatially clustered within the same distances (1000-2500 m) as densities of willow and birch, but at other spatial scales, browsing was mostly randomly distributed. It was concluded that foragedensity is a cue for moose but only at certain spatial scales. Similarly, a comparison of foraging distribution by hare and moose showed that high birch density was a key factor for both species. In spite of this, hares and moose used different parts ofthe same environment because they respond to food resource distribution at different spatial scales. Hares fed from smaller plants, and focused their foraging activity on smaller spatial scales than moose. These results emphasize the importance of taking into account the distribution of food resources at spatial scales relevant for each species in plans for conservation and management. In an experimental study it was found that intensified browsing on natural forage by mountain hares can be induced locally through placement of food. The induced browsing varied with the amount and quality of the added food, but also with thedensity of natural food plants and natural foraging distribution by hares. Finally, ina last experiment habitat preference of mountain hares across edges between open and forested areas was studied. The results were not consistent; hares utilized baitto a greater extent within forested areas than bait placed on a nearby lake ice, butbait on mires and heaths was either preferred over bait in nearby forest, or utilizedto a similar extent. A possible explanation is that hares have knowledge of their environment such that both forested areas and subarctic mires and heaths are partof its natural home range, whilst the extreme environment on the lake ice is not. During recent decades arctic areas have had an increase in vegetation density andwill be affected by future climate warming and therefore, factors that determineforaging ecology of key herbivores need to be identified. This thesis sheds some light on these factors in relation to spatial scale and forage distribution for two high profile herbivores in the subarctic.
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4. |
- Campbell, Charles, 1982-
(författare)
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Sphagnum limits : Physiology, morphology and climate
- 2019
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Sphagnum is the most important plant genus in terms of terrestrial carbon cycling. It and the habitats it creates store an equivalent of ~68% of the CO2 in the atmosphere. The genus has little dispersal limitation and the mire habitats are functionally similar at global scales. Sphagnum species are limited by water deficit at local and biogeographic scales, but this alone is not sufficient to explain local and global scale species patterns. As Sphagnum shoots are long-lived they may be limited by stochastic periods of cold temperature. Within Europe, species are associated with climate gradients along north-south (cold-warm) and oceanic-continental (wet-dry) clines. Within mires, species are sorted along a moisture (hummock-hollow) gradient.In this thesis I examined species responses to and recovery from freezing (I). I compared species with different water level niches in traits related to water management of individual shoots and colonies (II). Using distribution modelling of GBIF data, I estimated how different aspects of climate contributed to Sphagnum species distributions in Europe (III). Combining the approaches in papers II and III, I modelled the climatic distributions of the parapatric species S. cuspidatum and S. lindbergii and assessed how traits of water economy varied across the distribution boundary (IV).Species responses to winter stress were largely allied to both their hydrological niche and geographic range. Generally, hollow species managed better than hummock species, but species from intermediate positions were less consistent in their response. Species associated with boreal regions were generally less affected than those from temperate regions. Hardening against low temperature was triggered by shorter days and cold nights. Cold temperatures during late autumn may be more important for Sphagnum limits than the minimum temperature during winter.Water-related traits split the species into two groups; hollows species with large capitula and hummock species with small capitula. However, inter- and intra-specific trait variation and trait trends along the hydrological gradient were not necessarily the same at the shoot and canopy scale. Some trait correlations were common to all species. Canopy traits, which were emergent traits of colonies of shoots, had the strongest trait associations with the species position along the hummock-hollow gradient.At the continental scale the distribution of most Sphagnum species could be successfully modelled by a combination of annual degree days and water balance and the degree of seasonality in these two variables. Individual species distributions were shaped more by the seasonality in degree days than in water balance.Across the distributional border of S. cuspidatum and S. lindbergii divergence in the measured traits was mostly seen in the capitula indicating that limits to Sphagnum species are strongly linked to the functioning of the capitulum. Capitulum mass of both species was lower in sympatry than in allopatry, even though the measured values were similar. Canopy traits most strongly separated the species though did not change across the species boundaries.In summary, Sphagnum species in general are limited by the availability of water. Low temperature, particularly during late autumn are probably decisive for the biogeographic limits and for the distribution of species along the hydrological gradient.
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5. |
- Lindmark, Matilda
(författare)
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Increased resilience in managed boreal forests : Controlling damage by spruce bark beetle and moose in Sweden
- 2022
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Most of the forests growing in Europe today have been planted, or heavily affected, by humans. This is because the forest industry provides crucial raw materials for diverse products, including timber, pulp and paper, cellulose-based substances and renewable energy sources. However, forest management has turned heterogenic natural forests into homogenous forest stands that are highly susceptible to various kinds of damage. This thesis focuses on moose (Alces alces L.) browsing and attacks by spruce bark beetle (Ips typographus L.), both of which cause large economic losses. Empirically, it is based on field experiments conducted in managed forests in Sweden. Moose browsing on vulnerable top shoots of young pines during wintertime is a substantial problem in northern Europe. My colleagues and I (hereafter we) found that extracts of spruce trees (which moose prefer less than pine) can act as efficient natural browsing repellents.Spruce bark beetles attack spruce (Picea abies) trees throughout Europe. Thus, we also evaluated the field behavior of the beetles exposed to non-host- and fungal volatiles in efforts to identify more efficient ways to protect susceptible spruce stands. In addition, we evaluated the predator:prey ratio between the beetles and Thanasimus predators in representative forest stands, to assist efforts to enhance the biological control of spruce bark beetles in managed forests. Finally, we investigated changes in bacterial communities in wood after attacks by spruce bark beetles to improve knowledge of their ecology.Changes in forest management regime are needed to increase structural and tree species diversity, and thus raise forests’ resilience and ability to cope with ongoing climate change. Results presented in this thesis indicate that changing forest management to increase the variation in tree species variation will enhance the resilience of managed forests, thereby reducing damaging by both moose and spruce bark beetle.
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6. |
- Lönnell, Niklas, 1969-
(författare)
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Dispersal of bryophytes across landscapes
- 2014
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- Dispersal, especially long-distance dispersal, is an important component in many disciplines within biology. Many species are passively dispersed by wind, not least spore-dispersed organisms.In this thesis I investigated the dispersal capacity of bryophytes by studying the colonization patterns from local scales (100 m) to landscape scales (20 km). The dispersal distances were measured from a known source (up to 600 m away) or inferred from a connectivity measure (1–20 km). I introduced acidic clay to measure the colonization rates over one season of a pioneer moss, Discelium nudum (I–III). I also investigated which vascular plants and bryophytes that had colonized limed mires approximately 20–30 years after the first disturbance (IV).Discelium effectively colonized new disturbed substrates over one season. Most spores were deposited up to 50 meters from a source but the relationship between local colonization rates and connectivity increased with distance up to 20 km (I–III). Also calcicolous wetland bryophyte species were good colonizers over similar distances, while vascular plants in the same environment colonized less frequently. Common bryophytes that produce spores frequently were more effective colonizers, while no effect of spore size was detected (IV). A mechanistic model that take into account meteorological parameters to simulate the trajectories for spores of Discelium nudum fitted rather well to the observed colonization pattern, especially if spore release thresholds in wind variation and humidity were accounted for (III).This thesis conclude that bryophytes in open habitats can disperse effectively across landscapes given that the regional spore source is large enough (i.e. are common in the region and produce spores abundantly). For spore-dispersed organisms in open landscapes I suggest that it is often the colonization phase and not the transport that is the main bottle-neck for maintaining populations across landscapes.
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7. |
- Olofsson, Madelen, 1981-
(författare)
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On the investigation of chemical parameters reflecting microbial activity linked to nutrient availability in forest soil
- 2015
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- As agrarian society developed, the most fertile soils able to sustain the nutritional requirements needed for high crop yield were assigned to farming, while the more penurious soils were left to uphold the forest ecosystems. Some temperate forests are developed on acidic soils considered to be nutrient poor, as much of the inorganic nutrients are entrapped in poorly weatherable soil minerals and not easily accessed by plant roots. In an undisturbed ecosystem, the largest contribution of available nutrients comes from the recycling of organically bound nutrients via the decomposition of dead plant material. If biomass is removed, for instance with a more intensified exploitation of the forest ecosystems including whole tree harvesting, this source of nutrients is consequently decreased. The importance of soil mineral weathering as a source of nutrients, and especially that promoted by soil biota, is thereby emphasized.This thesis addresses biotic parameters associated with mineral weathering. Different aspects of soil solution sampling strategies and analysis of different organic ligands as well as biomarkers for the estimation of fungal biomass were investigated. These chemical parameters were also evaluated as indicators of microbial activity in relation to mineral nutrient availability in soil.With the assumption that the current nutrient status of a soil will affect the microbial interest of certain minerals as sources of inorganic nutrients, a mineral amendment trial was performed in a Swedish boreal forest soil. Overall, the amended soil presented good nutrient status, but with a possible shortage of iron. Due to this, it was hypothesized that the amended mineral with the highest iron content i.e. biotite would cause an elevation of microbial activity in its vicinity when compared to the bulk soil.The level of microbial activity in the vicinity of the amended minerals was evaluated via quantification of organic acids and siderophores, as well as estimation of fungal biomass and enzymatic activity.The highest microbial activity was measured for the O horizon of the investigated podzol, although nothing indicated an elevated association with the amended minerals. In the E horizon, however, elevation in microbial activity was observed in the vicinity of the biotite mineral when compared with bulk soil, although only a few of the investigated parameters differed significantly when evaluated separately. To enable this study, a highly sensitive analytical method employing liquid chromatography and mass spectrometry was developed to quantify a number of hydroxamate siderophores. On-line pre-concentration enabled detection of these organic ligands in the pico-molar range – a necessity when analyzing natural samples.Furthermore, an analytical method was developed for the estimation of fungal biomass via quantification of chitin-derived glucosamine, which also employed liquid chromatography and tandem mass spectrometry. Unlike currently available methods, the one presented in this thesis did not involve analyte derivatization, which resulted in high sample throughput while simultaneously avoiding complications involved with the additional derivatization procedure.The distribution of a group of organic ligands known as aromatic low molecular mass organic acids was also studied in a boreal forest podzol soil. Different sampling and samples preparation techniques, namely tension-lysimeters, soil centrifugation and liquid-soil extraction, were compared when analyzing soil solution components. Significant differences in analyte amount and species type were found between these sampling techniques. Some of the differences could be accounted for by variation in soil composition at different depths of the investigated podzol, but others could be attributed to structural differences within the studied analyte group. This clearly illustrated the intricacy of sampling and analysis when working with a sample matrix as complex and diverse as soil.As previously, liquid chromatography and mass spectrometry was used to quantify the analytes of interest. A highly sensitive analytical method was developed that was able to detect eleven aromatic low molecular mass organic acids in the nano-molar range. High selectivity was ensured by applying multiple reaction monitoring enabled by collision induced fragmentation of the analytes.
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8. |
- Sandström, Jennie
(författare)
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Old-Growth Forests in the High Coast Region in Sweden and Active Management in Forest Set-Asides
- 2018
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- In today´s intensively managed landscape, very few forests with old-growth characteristics and little human impact exist. One of the rare exceptions is pine forests on rocky soils, a forest type which has probably escaped extensive human use because of its low productivity. Our objective was to investigate the structure, dynamics, and history as well as the abundance and richness of wood-inhabiting fungi in these types of forest. We chose rocky pine forests situated in the High Coast Region to exemplify this forest type since the regional County Administration had already made surveys of the conservation value in 26 rocky pine forest stands in this region. We investigated the forests by recording tree species and measuring tree size and age in eight of the stands that were ranked with the highest conservation value. We also sampled dead wood to examine time since death and we sampled living and dead trees with fire scars to date fires. In addition, we made an inventory of wood-inhabiting fruiting bodies and took woodchip samples from logs to learn (by DNA analysis) whether five rare wood inhabiting fungi species were present as mycelia in logs.We found that rocky pine forests in the High Coast Region have a multi-sizedand multi-aged structure and old pine trees (approximately 13 ha-1 older than 300 years) are present. Fire has been common (an average of 42 years betweenfires) but they were likely to have been low-intense and small. Although the amount of dead wood is relatively low (4.4 m3 ha-1 on average) compared to many other boreal forests with old-growth characteristics, the share of deadwood of the total tree basal area (18%) was in line with other pine forests with low levels of human impact. The low dead wood volume is therefore likely to be an effect of the low productivity rather than dead wood extraction by humans. We also discovered that dead wood can be present for a really longtime without totally decomposing; we found logs and snags that had been dead for 500 years. This continuity of dead wood might be important for organisms dependent on dead wood as a substrate and even though we found that the species richness of wood-inhabiting fungi was somewhat low, we did find some rare species. Cinereomyces lenis and Hyphodontia halonata were present as fruiting bodies and we also found Antrodia albobrunnea, Antrodiainfirma, Crustoderma corneum and Anomoporia kamtschatica present as myceliain logs.The second part of this thesis reports two systematic reviews studying the effects of active management on the biodiversity in boreal and temperate forests. A systematic review follows certain guidelines and aims to compile the evidence base in well-defined topics, so that managers, researchers and policymakers can gain access to a high-quality compilation of current research. In our systematic map, we found almost 800 relevant papers but the set of papers turned out to be too heterogenic (many intervention types, e.g. thinning, burning, grazing and many types of outcomes) to allow any quantitative analysis. However, this map identified knowledge gaps and several detailed research questions that had sufficient data to provide aquantitative statistical analysis.One of these questions was: What is the impact of dead wood creation or addition on dead wood-dependent species? We focused on three types of interventions: creation of dead wood, addition of dead wood from elsewhere and prescribed burning. The selected outcomes were: saproxylic insects (rareand pest species), saproxylic fungi (rare species), ground-living insects and cavity-nesting birds. There was no significant negative effect on any of the investigated species groups but a positive effect on the abundance and richness of saproxylic insects and fungi. We also found that, although the amount of dead wood created was much less (50%) with prescribed burning, the abundance and richness of saproxylic insects showed similar positive effects to those of other intervention methods. A likely explanation for this is that burning results in a diversity of dead wood of various levels of quality (e.g. dense and/or charred wood), which creates a heterogeneity of dead woodtypes having a positive effect on the diversity of species dependent on deadwood. In summary, active management generally has a positive effect on biodiversity but the choice of management type should always be made carefully, and in consideration of the effect you want to achieve. In addition, there is a need for more long-term primary studies and more species groups in more geographical areas need to be incorporated so that the systematic reviews in this field will be even more informative in the future.
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9. |
- Skytt, Torbjörn, 1964-
(författare)
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Fossil Free or Not - That is the Question
- 2021
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- This is an evaluation of Jämtland County’s ability to achieve its climate targets to reduce greenhouse gas (GHG) emissions with 10% annually, and to become fossil fuel free 2030. There is a risk that the target to become fossil fuel-free, relying primarily on outtake of forest biomass, contradicts with the target in the UN Paris Agreement to keep the global temperature increase below 1.5°C. Neither the GHG emissions reduction target, nor the target to become fossil fuel free, takes forest carbon content into account. To reach the target to become fossil fuel free, forest harvest volumes may need to be increased. This can result in lower short-term net climate benefits than if fossil fuels are substituted to a less extent, and harvest volumes are kept at lower levels. Lower harvest volumes contribute to higher net climate benefits, and in the short term this effect is greater the lower the harvest level. In any case, forest production should be optimized towards products that give high substitution effects from the replacement of other materials. To be able to reach the short-term reductions of GHG emissions, efforts should be made to decrease regional energy demand and general consumption. The transition to new technologies is slow, and is unlikely to contribute to short-term reductions. By the end of 2023, according to the county strategy, the GHG emissions should be max 73% of the emissions by the end of 2019. This is a historically huge reduction. The potential to increase uptake of carbon dioxide in forests should be considered as a complement to the other climate targets. Assessments of social/welfare risks have to be weighed against the risks of increased temperatures, which is challenging, but needs to be done. The complexity of differences in short-term and long-term carbon balance effects makes risk assessments of climate mitigation strategies difficult. Compromises will be required between risks with different time horizons. To rely on techno-optimistic visions increase the risks that the emission reduction target will not be reached. From some perspectives, it is positive to spread techno-optimistic future visions, but there is a risk that we will keep on increasing consumption if we believe that problems with increased atmospheric concentrations of GHG can be solved by continuous consumption of ‘sustainable’ products. A simple 2% rule of thumb is that 2 kg of CO2eq is emitted for each 100 SEK consumed.
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10. |
- Berglund, Håkan, 1967-
(författare)
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Biodiversity in fragmented boreal forests : assessing the past, the present and the future
- 2004
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Doktorsavhandling (övrigt vetenskapligt/konstnärligt)abstract
- The aims of this thesis are to (1) analyze the predictability (indicators) of plant and fungal species diversity in old-growth forests, and (2) assess the history and biodiversity of woodland key habitats (WKHs) and their potential to maintain species diversity in fragmented boreal forest landscapes. Predictability was explored in Granlandet nature reserve, an unexploited landscape composed of discrete old-growth Picea forest patches of varying size isolated by wetland, reflecting conditions of insular biota at stochastic equilibrium. Data from 46 patches (0.2-12 ha) showed that most species were rare. However, species richness and composition patterns exhibited a high degree of predictability, which strengthen the possibility to apply biodiversity indicators in old-growth forest stands. Area was a key factor. The increase in species richness starts to level out at 2-3 ha. Large patches host more Red-list species in their interiors than do small ones, i.e. stand size is an important qualitative aspect of old-growth habitat. Nestedness emerged in relation to area but also in equal-sized plots. Structural complexity and habitat quality were important for species richness and compositional patterns, and small habitats of high quality could harbor many rare species. Monitoring of wood-fungi on downed logs showed that species diversity on downed logs changed over periods of 5-10 years and that the occurrences of annual species were unpredictable. It is suggested that monitoring of species with durable fruit bodies (mainly polypores) is likely to be a feasible approach to obtain comparable data over time.Assessments of biodiversity of WKHs were performed in two areas with contrasting histories of forest exploitation, namely in south boreal and north boreal Sweden. Analyses of the history of 15 south boreal WKHs showed that fire-suppression, selective logging until mid-20th century and abandonment by modern forestry has shaped their forest structure. These WKHs are not untouched forests, they lack key structural components and harbor few Red-list species. Artificial interventions to restore natural processes and patterns are needed to further increase their suitability for threatned species. Modeling analyses of species richness in 32 WKHs in north boreal Sweden, some of which have not been isolated by modern forestry until recently, indicated an excess of crustose lichen species, i.e. WKHs may face delayed species extinctions. By contrast, the results indicate that wood-fungi have tracked the environmental changes. Differences in substrate dynamics between epiphytes on living trees and species growing on decaying logs may explain the diffeence between species groups. The results indicate that population densities of Red-list species were low, which may result in further depletion of species diversity.Continuing species declines and extinctions are likely if not conservation of WKHs are combined with other considerations in th managed forest landscape. Both WKHs and their surroundings must be managed and designed to maintain biodiversity over time. For a successful future conservation of boreal forest biodiversity monitoring of WKHs must be combined with monitoring of refeence areas.
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