The ecology of the mountain hare (Lepus timidus L.) in managed boreal forests: habitat associations at different forest scales

Aims of this thesis This study is part of a larger hare project in Finland, which provides answers to basic ecological questions regarding the mountain hare. This study of the ecology of the mountain hare focuses in particular on different levels of managed boreal forest. The feeding habits and intensity of mountain hares in winter are explored, and the connections between mountain hares versus the forest structure are also studied (e.g. habitat use and the importance of different forest layers for hares). The use of the environment by hares at the landscape level was examined (forest patch structures), and the home ranges of mountain hares were studied. Finally, the productivity and survival rate of mountain hare populations were also studied (discussion e.g. predator effects on hare populations). Conclusions Feeding intensity seemed to be highest in the spring-winter, when home ranges were also largest. Favourable food species are covered by snow in winter and the mobility of hares is highest during late winter. A shortage of suitable food species may be problematic for hares, especially during the winter period. In this study mountain hares preferred a dense shrub layer at local level and deciduous and mixed tree forest over coniferous forest at the landscape level. Food and shelter are vital for hares and the preference for particular habitats may also affect the population dynamics of the mountain hare. It would be possible to improve the quality of food and shelter or at least prevent the most negative habitat changes through forest management. At a local level it is also possible to add supplementary food for hares through the winter period. The intensive clearing of young sapling stands and especially the removal of deciduous shrubs and trees reduces the quality of habitats for the mountain hare. Mountain hares primarily live in forest habitat and it is possible that changes in the forest structure play a crucial role in mountain hare habitat preference. Ecological knowledge of the mountain hare is vital to create habitat structure more suitable for the species. More deciduous trees should be saved in managing forests and the mechanical clearing of the shrub layer should be done carefully.

Disturbance in boreal spruce forest - immediate dynamics from stand to understorey level

The immediate effects of two human-related vegetation disturbances, (1) green tree retention (GTR) patch felling and scarification by harrowing and (2) experimental understorey vegetation layer removal, were examined in boreal forest stands in Finland. Effects of GTR patch felling and scarification on tree uprootings, on coarse woody debris (CWD) and on epixylic plant community were followed in upland and in paludified forest types. Uprootings increased considerably during 2-3 years after the fellings and were more frequent (47%) in the paludified than in the upland forest (13%). Scarification reduced 68% of the CWD in the felling area. Cover and especially species richness of epixylics declined in the both areas during 1-2 years after the felling. The increasing size of GTR patch correlated positively with the species richness. Regeneration of understorey vegetation community and Vaccinium myrtillus and Vaccinium vitis-idaea after different removals of vegetation layers in an old-growth forest took four years. The regeneration occurred mainly by vegetative means and it was faster in the terms of species richness than in the cover. In the most severe treatment, recovery occurred merely by sexual reproduction. V. myrtillus recovered mainly by producing new shoots. V. vitis-idaea recovered faster than V. myrtillus, mainly by increasing length growth. For ecological reasons, use of larger GTR patches on paludified biotope would be recommendable. In felling areas, scarification by harrowing could be replaced with some other spot-wise method. After moderate intensity level disturbance, recovery occurs rapidly by vegetative regrowth of the dominating species. High level of intensity may prevent the recovery of vegetation community for years, while enabling also the genetic regeneration of the initial species. Local anthropogenic-related disturbances are currently increasing and they can interact during temporally short times, which should be taken in to account in the future forest management plans.

Temperature sensitivity of soil organic matter decomposition in boreal soils

The temperature sensitivity of decomposition of different soil organic matter (SOM) fractions was studied with laboratory incubations using 13C and 14C isotopes to differentiate between SOM of different age. The quality of SOM and the functionality and composition of microbial communities in soils formed under different climatic conditions were also studied. Transferring of organic layers from a colder to a warmer climate was used to assess how changing climate, litter input and soil biology will affect soil respiration and its temperature sensitivity. Together, these studies gave a consistent picture on how warming climate will affect the decomposition of different SOM fractions in Finnish forest soils: the most labile C was least temperature sensitive, indicating that it is utilized irrespective of temperature. The decomposition of intermediate C, with mean residence times from some years to decades, was found to be highly temperature sensitive. Even older, centennially cycling C was again less temperature sensitive, indicating that different stabilizing mechanisms were limiting its decomposition even at higher temperatures. Because the highly temperature sensitive, decadally cycling C, forms a major part of SOM stock in the organic layers of the studied forest soils, these results mean that these soils could lose more carbon during the coming years and decades than estimated earlier. SOM decomposition in boreal forest soils is likely to increase more in response to climate warming, compared to temperate or tropical soils, also because the Q10 is temperature dependent. In the northern soils the warming will occur at a lower temperature range, where Q10 is higher, and a similar increase in temperature causes a higher relative increase in respiration rates. The Q10 at low temperatures was found to be inversely related to SOM quality. At higher temperatures respiration was increasingly limited by low substrate availability.

Phosphorus retention in forest soils and the functioning of buffer zones used in forestry

Phosphorus (P) retention properties of soils typical for boreal forest, i.e. podzolic soil and peat soils, vary significantly, but the range of this variation has not been sufficiently documented. To assess the usefulness of buffer zones used in forestry in removing P from the discharge by chemical sorption in soil, and to estimate the risk of P leaching after forestry operations, more data is needed on soil P retention properties. P retention properties of soils were studied at clear-cut areas, unharvested buffer zones adjoining the clear-cut and at peatland buffer zone areas. Desorption-sorption isotherms were determined for the humus layer, the mineral soil horizons E, B and C of the Podzol profile and for the surface layer peat (0-15 cm) and the subsurface layer peat (15-30 cm). The efficiency of buffer zones in retaining P was studied at six peatland buffer zone areas by adding P-containing solute in the inflow. A tracer study was conducted at one of the buffer zone areas to determine the allocation of the added P in soil and vegetation. Measured sorption or desorption rather than parameter values of fitted sorption equations described P desorption and sorption behaviour in soil. The highest P retention efficiency was in the B horizon and consequently, if contact occurred or was established between the soluble P in the water and the soil B horizon, the risk of P leaching was low. Humus layer was completely incapable of retaining P after clear-cutting. In the buffer zones, the decrease in P retention properties in the humus layer and the low amount of P sorbed by it indicated that the importance of the layer in the functioning of buffer zones is low. The peatland buffer zone areas were efficient in retaining soluble P from inflow. P sorption properties of the peat soil at the buffer zone areas varied largely but the contribution of P sorption in the peat was particularly important during high flow in spring, when the vegetation was not fully developed. Factors contributing to efficient P retention were large buffer size and low hydrological load whereas high hydrological load combined with the formation of preferential flow paths, especially during early spring or late autumn was disadvantageous. However, small buffer zone areas, too, may be efficient in reducing P load.

Spatial variation of climate and the impact of disturbances on local climate and forest recovery in northern Finland

The structure and function of northern ecosystems are strongly influenced by climate change and variability and by human-induced disturbances. The projected global change is likely to have a pronounced effect on the distribution and productivity of different species, generating large changes in the equilibrium at the tree-line. In turn, movement of the tree-line and the redistribution of species produce feedback to both the local and the regional climate. This research was initiated with the objective of examining the influence of natural conditions on the small-scale spatial variation of climate in Finnish Lapland, and to study the interaction and feedback mechanisms in the climate-disturbances-vegetation system near the climatological border of boreal forest. The high (1 km) resolution spatial variation of climate parameters over northern Finland was determined by applying the Kriging interpolation method that takes into account the effect of external forcing variables, i.e., geographical coordinates, elevation, sea and lake coverage. Of all the natural factors shaping the climate, the geographical position, local topography and altitude proved to be the determining ones. Spatial analyses of temperature- and precipitation-derived parameters based on a 30-year dataset (1971-2000) provide a detailed description of the local climate. Maps of the mean, maximum and minimum temperatures, the frost-free period and the growing season indicate that the most favourable thermal conditions exist in the south-western part of Lapland, around large water bodies and in the Kemijoki basin, while the coldest regions are in highland and fell Lapland. The distribution of precipitation is predominantly longitudinally dependent but with the definite influence of local features. The impact of human-induced disturbances, i.e., forest fires, on local climate and its implication for forest recovery near the northern timberline was evaluated in the Tuntsa area of eastern Lapland, damaged by a widespread forest fire in 1960 and suffering repeatedly-failed vegetation recovery since that. Direct measurements of the local climate and simulated heat and water fluxes indicated the development of a more severe climate and physical conditions on the fire-disturbed site. Removal of the original, predominantly Norway spruce and downy birch vegetation and its substitution by tundra vegetation has generated increased wind velocity and reduced snow accumulation, associated with a large variation in soil temperature and moisture and deep soil frost. The changed structural parameters of the canopy have determined changes in energy fluxes by reducing the latter over the tundra vegetation. The altered surface and soil conditions, as well as the evolved severe local climate, have negatively affected seedling growth and survival, leading to more unfavourable conditions for the reproduction of boreal vegetation and thereby causing deviations in the regional position of the timberline. However it should be noted that other factors, such as an inadequate seed source or seedbed, the poor quality of the soil and the intensive logging of damaged trees could also exacerbate the poor tree regeneration. In spite of the failed forest recovery at Tunsta, the position and composition of the timberline and tree-line in Finnish Lapland may also benefit from present and future changes in climate. The already-observed and the projected increase in temperature, the prolonged growing season, as well as changes in the precipitation regime foster tree growth and new regeneration, resulting in an advance of the timberline and tree-line northward and upward. This shift in the distribution of vegetation might be decelerated or even halted by local topoclimatic conditions and by the expected increase in the frequency of disturbances.

Canopy processes, fluxes and microclimate in a pine forest

Interaction between forests and the atmosphere occurs by radiative and turbulent transport. The fluxes of energy and mass between surface and the atmosphere directly influence the properties of the lower atmosphere and in longer time scales the global climate. Boreal forest ecosystems are central in the global climate system, and its responses to human activities, because they are significant sources and sinks of greenhouse gases and of aerosol particles. The aim of the present work was to improve our understanding on the existing interplay between biologically active canopy, microenvironment and turbulent flow and quantify. In specific, the aim was to quantify the contribution of different canopy layers to whole forest fluxes. For this purpose, long-term micrometeorological and ecological measurements made in a Scots pine (Pinus sylvestris) forest at SMEAR II research station in Southern Finland were used. The properties of turbulent flow are strongly modified by the interaction between the canopy elements: momentum is efficiently absorbed in the upper layers of the canopy, mean wind speed and turbulence intensities decrease rapidly towards the forest floor and power spectra is modulated by spectral short-cut . In the relative open forest, diabatic stability above the canopy explained much of the changes in velocity statistics within the canopy except in strongly stable stratification. Large eddies, ranging from tens to hundred meters in size, were responsible for the major fraction of turbulent transport between a forest and the atmosphere. Because of this, the eddy-covariance (EC) method proved to be successful for measuring energy and mass exchange inside a forest canopy with exception of strongly stable conditions. Vertical variations of within canopy microclimate, light attenuation in particular, affect strongly the assimilation and transpiration rates. According to model simulations, assimilation rate decreases with height more rapidly than stomatal conductance (gs) and transpiration and, consequently, the vertical source-sink distributions for carbon dioxide (CO2) and water vapor (H2O) diverge. Upscaling from a shoot scale to canopy scale was found to be sensitive to chosen stomatal control description. The upscaled canopy level CO2 fluxes can vary as much as 15 % and H2O fluxes 30 % even if the gs models are calibrated against same leaf-level dataset. A pine forest has distinct overstory and understory layers, which both contribute significantly to canopy scale fluxes. The forest floor vegetation and soil accounted between 18 and 25 % of evapotranspiration and between 10 and 20 % of sensible heat exchange. Forest floor was also an important deposition surface for aerosol particles; between 10 and 35 % of dry deposition of particles within size range 10 30 nm occurred there. Because of the northern latitudes, seasonal cycle of climatic factors strongly influence the surface fluxes. Besides the seasonal constraints, partitioning of available energy to sensible and latent heat depends, through stomatal control, on the physiological state of the vegetation. In spring, available energy is consumed mainly as sensible heat and latent heat flux peaked about two months later, in July August. On the other hand, annual evapotranspiration remains rather stable over range of environmental conditions and thus any increase of accumulated radiation affects primarily the sensible heat exchange. Finally, autumn temperature had strong effect on ecosystem respiration but its influence on photosynthetic CO2 uptake was restricted by low radiation levels. Therefore, the projected autumn warming in the coming decades will presumably reduce the positive effects of earlier spring recovery in terms of carbon uptake potential of boreal forests.

Root system traits of Norway spruce, Scots pine, and silver birch in mixed boreal forests: an analysis of root architecture, morphology, and anatomy

The aim of this thesis was to unravel the functional-structural characteristics of root systems of Betula pendula Roth., Picea abies (L.) Karst., and Pinus sylvestris L. in mixed boreal forest stands differing in their developmental stage and site fertility. The root systems of these species had similar structural regularities: horizontally-oriented shallow roots defined the horizontal area of influence, and within this area, each species placed fine roots in the uppermost soil layers, while sinker roots defined the maximum rooting depth. Large radial spread and high ramification of coarse roots, and the high specific root length (SRL) and root length density (RLD) of fine roots indicated the high belowground competitiveness and root plasticity of B. pendula. Smaller radial root spread and sparser branching of coarse roots, and low SRL and RLD of fine roots of the conifers could indicate their more conservative resource use and high association with and dependence on ectomycorrhiza-forming fungi. The vertical fine root distributions of the species were mostly overlapping, implying the possibility for intense belowground competition for nutrients. In each species, conduits tapered and their frequency increased from distal roots to the stem, from the stem to the branches, and to leaf petioles in B. pendula. Conduit tapering was organ-specific in each species violating the assumptions of the general vascular scaling model (WBE). This reflects the hierarchical organization of a tree and differences between organs in the relative importance of transport, safety, and mechanical demands. The applied root model was capable of depicting the mass, length and spread of coarse roots of B. pendula and P. abies, and to the lesser extent in P. sylvestris. The roots did not follow self-similar fractal branching, because the parameter values varied within the root systems. Model parameters indicate differences in rooting behavior, and therefore different ecophysiological adaptations between species.

Habitat selection of three rodents in a frequently burned boreal environment : productivity and habitat diversity

How does fire affect the plant and animal community of the boreal forest? This study attempted to examine the changes in plant composition and productivity, and small mammal demography brought about by fire in the northern boreal environment at Chick Lake, N.W.T. (65053fN, 128°14,W). Two 5*6 ha plots measuring 375m x 150m were selected for study during the summers of 1973 and 197^. One had been unburned for 120 years, the other was part of a fire which burned in the spring of 1969. Grids of 15m x 15m were established in each plot and meter square quadrats taken at each of the 250 grid intersections in order to determine plant composition and density. Aerial primary production was assessed by clipping and drying 80 samples of terminal new production for each species under investigation. Small mammal populations were sampled by placing a Sherman live trap at each grid intersection for ten days in every month. The two plots were similar in plant species composition which suggested that most regrowth in the burned area was from rootstocks which survived the fire. The plant data were submitted to a cluster analysis that revealed nine separate species associations, six of which occured in the burned area and eight of which occured in the control. These were subsequently treated as habitats for purposes of comparison with small mammal distributions. The burned area showed a greater productivity in flowers and fruits although total productivity in the control area was higher due to a large contribution from the non-vascular component. Maximum aerial productivity as dry wieght was measured at 157.1 g/m and 207.8 g/m for the burn and control respectively. Microtus pennsylvanicus and Clethrionomys rutilus were the two most common small mammals encountered; Microtus xanthognathus, Synaptomys borealis, and Phenacomys intermedius also occured in the area. Populations of M. pennsylvanicus and C. rutilus were high during the summer of 1973; however, M. pennsylvanicus was rare on the control but abundant on the burn, while C. rutilus was rare on the burn but abundant in the control. During the summer of 197^ populations declined, with the result that few voles of any species were caught in the burn while equal numbers of the two species were caught in the control. During the summer of 1973 M. pennsylvanicus showed a positive association to the most productive habitat type in the burn which was avoided by C. rutilus. In the control £• rutilus showed a similar positive association to the most productive habitat type which was avoided by M. pennsylvanicus. In all cases for the high population year of 1973# the two species never overlapped in habitat preference. When populations declined in 197^f "both species showed a strong association for the most productive habitat in the control. This would suggest that during a high population year, an abundant species can exclude competitors from a chosen habitat, but that this dominance decreases as population levels decrease. It is possible that M. pennsylvanicus is a more efficient competitor in a recently burned environment, while C. rutilus assumes this role once non-vascular regrowth becomes extensive.

Effects of Spruce Budworm (Choristoneura fumiferana (Clem.)) Outbreaks on Boreal Mixed-Wood Bird Communities

This study examined the influence of a spruce budworm (Choristoneura fumiferana (Clem.)) outbreak on a boreal mixed-wood bird community in forest stands ranging in age from 0 to 223 yr. We asked if (1) patterns of species response were consistent with the existence of spruce budworm specialists, i.e., species that respond in a stronger quantitative or qualitative way than other species; (2) the superabundance of food made it possible for species to expand their habitat use in age classes that were normally less used; and (3) the response to budworm was limited to specialists or was it more widespread. Results here indicated that three species, specifically the Bay-breasted Warbler (Dendroica castanea), Tennessee Warbler (Vermivora peregrina), and Cape May Warbler (Dendroica tigrina), had a larger numerical response to the budworm outbreak. They responded with increases in density of up to tenfold over 4 or 5 yr. No other species responded with more than a twofold increase in the same time period. These species also showed a functional response by breeding more frequently in young stands aged 1–21 yr and intermediate stands aged 22–36 yr as budworm numbers increased. Our data also suggested that many species profited to a lesser extent from budworm outbreaks, but that this effect may be too subtle to detect in most studies. We found evidence of a positive numerical effect in at least 18 additional species in one or two stand-age categories but never in all three for any one species. Given the numerical response in many species and the potential influence of budworm on bird populations because of the vast extent of outbreaks, we believe that the population cycle of spruce budworm should be considered in any evaluation of population trends in eastern boreal birds.

Ovenbird (Seiurus aurocapilla) Territory Placement Near Seismic Lines is Influenced by Forest Regeneration and Conspecific Density

The boreal forest of western Canada is being dissected by seismic lines used for oil and gas exploration. The vast amount of edge being created is leading to concerns that core habitat will be reduced for forest interior species for extended periods of time. The Ovenbird (Seiurus aurocapilla) is a boreal songbird known to be sensitive to newly created seismic lines because it does not include newly cut lines within its territory. We examined multiple hypotheses to explain potential mechanisms causing this behavior by mapping Ovenbird territories near lines with varying states of vegetation regeneration. The best model to explain line exclusion behavior included the number of neighboring conspecifics, the amount of bare ground, leaf-litter depth, and canopy closure. Ovenbirds exclude recently cut seismic lines from their territories because of lack of protective cover (lower tree and shrub cover) and because of reduced food resources due to large areas of bare ground. Food reduction and perceived predation risk effects seem to be mitigated once leaf litter (depth and extent of cover) and woody vegetation cover are restored to forest interior levels. However, as conspecific density increases, lines are more likely to be used as landmarks to demarcate territorial boundaries, even when woody vegetation cover and leaf litter are restored. This behavior can reduce territory density near seismic lines by changing the spatial distribution of territories. Landmark effects are longer lasting than the effects from reduced food or perceived predation risk because canopy height and tree density take >40 years to recover to forest interior levels. Mitigation of seismic line impacts on Ovenbirds should focus on restoring forest cover as quickly as possible after line cutting.

How well do regional or national Breeding Bird Survey data predict songbird population trends at an intact boreal site?

A study to monitor boreal songbird trends was initiated in 1998 in a relatively undisturbed and remote part of the boreal forest in the Northwest Territories, Canada. Eight years of point count data were collected over the 14 years of the study, 1998-2011. Trends were estimated for 50 bird species using generalized linear mixed-effects models, with random effects to account for temporal (repeat sampling within years) and spatial (stations within stands) autocorrelation and variability associated with multiple observers. We tested whether regional and national Breeding Bird Survey (BBS) trends could, on average, predict trends in our study area. Significant increases in our study area outnumbered decreases by 12 species to 6, an opposite pattern compared to Alberta (6 versus 15, respectively) and Canada (9 versus 20). Twenty-two species with relatively precise trend estimates (precision to detect > 30% decline in 10 years; observed SE ≤ 3.7%/year) showed nonsignificant trends, similar to Alberta (24) and Canada (20). Precision-weighted trends for a sample of 19 species with both reliable trends at our site and small portions of their range covered by BBS in Canada were, on average, more negative for Alberta (1.34% per year lower) and for Canada (1.15% per year lower) relative to Fort Liard, though 95% credible intervals still contained zero. We suggest that part of the differences could be attributable to local resource pulses (insect outbreak). However, we also suggest that the tendency for BBS route coverage to disproportionately sample more southerly, developed areas in the boreal forest could result in BBS trends that are not representative of range-wide trends for species whose range is centred farther north.

Biased representation of disturbance rates in the roadside sampling frame in boreal forests: implications for monitoring design

The North American Breeding Bird Survey (BBS) is the principal source of data to inform researchers about the status of and trend for boreal forest birds. Unfortunately, little BBS coverage is available in the boreal forest, where increasing concern over the status of species breeding there has increased interest in northward expansion of the BBS. However, high disturbance rates in the boreal forest may complicate roadside monitoring. If the roadside sampling frame does not capture variation in disturbance rates because of either road placement or the use of roads for resource extraction, biased trend estimates might result. In this study, we examined roadside bias in the proportional representation of habitat disturbance via spatial data on forest “loss,” forest fires, and anthropogenic disturbance. In each of 455 BBS routes, the area disturbed within multiple buffers away from the road was calculated and compared against the area disturbed in degree blocks and BBS strata. We found a nonlinear relationship between bias and distance from the road, suggesting forest loss and forest fires were underrepresented below 75 and 100 m, respectively. In contrast, anthropogenic disturbance was overrepresented at distances below 500 m and underrepresented thereafter. After accounting for distance from road, BBS routes were reasonably representative of the degree blocks they were within, with only a few strata showing biased representation. In general, anthropogenic disturbance is overrepresented in southern strata, and forest fires are underrepresented in almost all strata. Similar biases exist when comparing the entire road network and the subset sampled by BBS routes against the amount of disturbance within BBS strata; however, the magnitude of biases differed. Based on our results, we recommend that spatial stratification and rotating panel designs be used to spread limited BBS and off-road sampling effort in an unbiased fashion and that new BBS routes be established where sufficient road coverage exists.

Modelling the effects of nitrogen deposition and carbon dioxide enrichment on forest carbon balance

Atmospheric CO2 concentration ([CO2]) has increased over the last 250 years, mainly due to human activities. Of total anthropogenic emissions, almost 31% has been sequestered by the terrestrial biosphere. A considerable contribution to this sink comes from temperate and boreal forest ecosystems of the northern hemisphere, which contain a large amount of carbon (C) stored as biomass and soil organic matter. Several potential drivers for this forest C sequestration have been proposed, including increasing atmospheric [CO2], temperature, nitrogen (N) deposition and changes in management practices. However, it is not known which of these drivers are most important. The overall aim of this thesis project was to develop a simple ecosystem model which explicitly incorporates our best understanding of the mechanisms by which these drivers affect forest C storage, and to use this model to investigate the sensitivity of the forest ecosystem to these drivers. I firstly developed a version of the Generic Decomposition and Yield (G’DAY) model to explicitly investigate the mechanisms leading to forest C sequestration following N deposition. Specifically, I modified the G’DAY model to include advances in understanding of C allocation, canopy N uptake, and leaf trait relationships. I also incorporated a simple forest management practice subroutine. Secondly, I investigated the effect of CO2 fertilization on forest productivity with relation to the soil N availability feedback. I modified the model to allow it to simulate short-term responses of deciduous forests to environmental drivers, and applied it to data from a large-scale forest Free-Air CO2 Enrichment (FACE) experiment. Finally, I used the model to investigate the combined effects of recent observed changes in atmospheric [CO2], N deposition, and climate on a European forest stand. The model developed in my thesis project was an effective tool for analysis of effects of environmental drivers on forest ecosystem C storage. Key results from model simulations include: (i) N availability has a major role in forest ecosystem C sequestration; (ii) atmospheric N deposition is an important driver of N availability on short and long time-scales; (iii) rising temperature increases C storage by enhancing soil N availability and (iv) increasing [CO2] significantly affects forest growth and C storage only when N availability is not limiting.