Intensive land use in the Swedish mountains between AD 800 and 1200 led to deforestation and ecosystem transformation with long-lasting effects.

Anthropogenic deforestation has shaped ecosystems worldwide. In subarctic ecosystems, primarily inhabited by native peoples, deforestation is generally considered to be mainly associated with the industrial period. Here we examined mechanisms underlying deforestation a thousand years ago in a high-mountain valley with settlement artifacts located in subarctic Scandinavia. Using the Heureka Forestry Decision Support System, we modeled pre-settlement conditions and effects of tree cutting on forest cover. To examine lack of regeneration and present nutrient status, we analyzed soil nitrogen. We found that tree cutting could have deforested the valley within some hundred years. Overexploitation left the soil depleted beyond the capacity of re-establishment of trees. We suggest that pre-historical deforestation has occurred also in subarctic ecosystems and that ecosystem boundaries were especially vulnerable to this process. This study improves our understanding of mechanisms behind human-induced ecosystem transformations and tree-line changes, and of the concept of wilderness in the Scandinavian mountain range.

Diazotrophy in alluvial meadows of subarctic river systems.

There is currently limited understanding of the contribution of biological N2 fixation (diazotrophy) to the N budget of large river systems. This natural source of N in boreal river systems may partially explain the sustained productivity of river floodplains in Northern Europe where winter fodder was harvested for centuries without fertilizer amendments. In much of the world, anthropogenic pollution and river regulation have nearly eliminated opportunities to study natural processes that shaped early nutrient dynamics of large river systems; however, pristine conditions in northern Fennoscandia allow for the retrospective evaluation of key biochemical processes of historical significance. We investigated biological N2 fixation (diazotrophy) as a potential source of nitrogen fertility at 71 independent floodplain sites along 10 rivers and conducted seasonal and intensive analyses at a subset of these sites. Biological N2 fixation occurred in all floodplains, averaged 24.5 kg N ha(-1) yr(-1) and was down regulated from over 60 kg N ha(-1) yr(-1) to 0 kg N ha(-1) yr(-1) by river N pollution. A diversity of N2-fixing cyanobacteria was found to colonize surface detritus in the floodplains. The data provide evidence for N2 fixation to be a fundamental source of new N that may have sustained fertility at alluvial sites along subarctic rivers. Such data may have implications for the interpretation of ancient agricultural development and the design of contemporary low-input agroecosystems.

Boreal feather mosses secrete chemical signals to gain nitrogen.

The mechanistic basis of feather moss-cyanobacteria associations, a main driver of nitrogen (N) input into boreal forests, remains unknown. Here, we studied colonization by Nostoc sp. on two feather mosses that form these associations (Pleurozium schreberi and Hylocomium splendens) and two acrocarpous mosses that do not (Dicranum polysetum and Polytrichum commune). We also determined how N availability and moss reproductive stage affects colonization, and measured N transfer from cyanobacteria to mosses. The ability of mosses to induce differentiation of cyanobacterial hormogonia, and of hormogonia to then colonize mosses and re-establish a functional symbiosis was determined through microcosm experiments, microscopy and acetylene reduction assays. Nitrogen transfer between cyanobacteria and Pleurozium schreberi was monitored by secondary ion mass spectrometry (SIMS). All mosses induced hormogonia differentiation but only feather mosses were subsequently colonized. Colonization on Pleurozium schreberi was enhanced during the moss reproductive phase but impaired by elevated N. Transfer of N from cyanobacteria to their host moss was observed. Our results reveal that feather mosses likely secrete species-specific chemo-attractants when N-limited, which guide cyanobacteria towards them and from which they gain N. We conclude that this signalling is regulated by N demands of mosses, and serves as a control of N input into boreal forests.

Ecosystem feedbacks and nitrogen fixation in boreal forests.

Biological feedback mechanisms regulate fundamental ecosystem processes and potentially regulate ecosystem productivity. To date, no studies have documented the down-regulation of terrestrial nitrogen (N) fixation via an ecosystem-level feedback mechanism. Herein, we demonstrate such a feedback in boreal forests. Rapid cycling of N in early secondary succession forests yielded greater throughfall N deposition, which in turn decreased N fixation by cyanobacterial associates in feather moss carpets that reside on the forest floor. The forest canopy exerts a tight control on biotic N input at a period of high productivity.

Fire-derived charcoal causes loss of forest humus.

Fire is a global driver of carbon storage and converts a substantial proportion of plant biomass to black carbon (for example, charcoal), which remains in the soil for thousands of years. Black carbon is therefore often proposed as an important long-term sink of soil carbon. We ran a 10-year experiment in each of three boreal forest stands to show that fire-derived charcoal promotes loss of forest humus and that this is associated with enhancement of microbial activity by charcoal. This result shows that charcoal-induced losses of belowground carbon in forests can partially offset the benefits of charcoal as a long-term carbon sink.

Long-term fire frequency not linked to prehistoric occupations in northern Swedish boreal forest.

Knowledge of past fire regimes is crucial for understanding the changes in fire frequency that are likely to occur during the coming decades as a result of global warming and land-use change. This is a key issue for the sustainable management of forest biodiversity because fire regimes may be controlled by vegetation, human activities, and/or climate. The present paper aims to reconstruct the pattern of fire frequency over the Holocene at three sites located in the same region in the northern Swedish boreal forest. The fire regime is reconstructed from sedimentary charcoal analysis of small lakes or ponds. This method allows fire events to be characterized, after detrending the charcoal influx series, and allows estimation of the time elapsed between fires. The long-term fire regime, in terms of fire-free intervals, can thus be elucidated. At the three sites, the mean fire-free intervals through the Holocene were long and of similar magnitude (approximately 320 years). This similarity suggests that the ecological processes controlling fire ignition and spread were the same. At the three sites, the intervals were shorter before 8600 cal yr BP (calibrated years before present), between 7500 and 4500 cal yr BP, and after 2500 cal yr BP. Geomorphological and vegetation factors cannot explain the observed change, because the three sites are located in the same large ecological region characterized by Pinus sylvestris-Ericaceae mesic forests, established on morainic deposits at the same elevation. Archaeological chronologies also do not match the fire chronologies. A climatic interpretation is therefore the most likely explanation of the long-term regional pattern of fire. Although recent human activities between the 18th and the 20th centuries have clearly affected the fire regime, the dominant factor controlling it for 10000 years in northern Sweden has probably been climatic.

Ecosystem controls on nitrogen fixation in boreal feather moss communities.

N fixation in feather moss carpets is maximized in late secondary successional boreal forests; however, there is limited understanding of the ecosystem factors that drive cyanobacterial N fixation in feather mosses with successional stage. We conducted a reciprocal transplant experiment to assess factors in both early and late succession that control N fixation in feather moss carpets dominated by Pleurozium schreberi. In 2003, intact microplots of moss carpets (30 cm x 30 cm x 10-20 cm deep) were excavated from three early secondary successional (41-101 years since last fire) forest sites and either replanted within the same stand or transplanted into one of three late successional (241-356 years since last fire) forest sites and the transverse was done for late successional layers of moss. Moss plots were monitored for changes in N-fixation rates by acetylene reduction (June 2003-September 2005) and changes in the presence of cyanobacteria on moss shoots by microscopy (2004). Forest nutrient status was measured using ionic resin capsules buried in the humus layer. Late successional forests exhibit high rates of N fixation and consistently high numbers of cyanobacteria on moss shoots, but low levels of available N. Conversely, early successional forests have higher N availability and have low rates of N fixation and limited presence of cyanobacteria on moss shoots. Transplantation of moss carpets resulted in a significant shift in presence and activity of cyanobacteria 1 year after initiation of the experiment responding to N fertility differences in early versus late successional forests.

Physiological and molecular diversity of feather moss associative N2-fixing cyanobacteria.

Cyanobacteria colonizing the feather moss Pleurozium schreberi were isolated from moss samples collected in northern Sweden and subjected to physiological and molecular characterization. Morphological studies of isolated and moss-associated cyanobacteria were carried out by light microscopy. Molecular tools were used for cyanobacteria identification, and a reconstitution experiment of the association between non-associative mosses and cyanobacteria was conducted. The influence of temperature on N2 fixation in the different cyanobacterial isolates and the influence of light and temperature on N2-fixation rates in the moss were studied using the acetylene reduction assay. Two different cyanobacteria were effectively isolated from P. schreberi: Nostoc sp. and Calothrix sp. A third genus, Stigonema sp. was identified by microscopy, but could not be isolated. The Nostoc sp. was found to fix N2 at lower temperatures than Calothrix sp. Nostoc sp. and Stigonema sp. were the predominant cyanobacteria colonizing the moss. The attempt to reconstitute the association between the moss and cyanobacteria was successful. The two isolated genera of cyanobacteria in feather moss samples collected in northern Sweden differ in their temperature optima, which may have important ecological implications.

Effects of species and functional group loss on island ecosystem properties.

Considerable recent attention has focused on predicting how the losses of species and functional groups influence ecosystem properties, but the extent to which these effects vary among ecosystems remains poorly understood. Island systems have considerable scope for studying how biotic and abiotic factors influence processes in different ecosystems, because they enable the simultaneous study of large numbers of independent replicate systems at ecologically meaningful spatial scales. We studied a group of 30 islands in northern Sweden, for which island size determined disturbance history, and therefore vegetation successional stage and biotic and abiotic ecosystem properties. On each island we conducted a seven-year study that involved experimental removals of combinations of both plant functional groups and plant species. We show that although losses of functional groups and species often impaired key ecosystem processes, these effects were highly context-dependent and strongly influenced by island size. Our study provides evidence that the consequences of biotic loss for ecosystem functioning vary greatly among ecosystems and depend on the specific abiotic and biotic attributes of the system.

Potential toxic effect on aquatic fauna by the dwarf shrub Empetrum hermaphroditum.

The common evergreen dwarf shrub Empetrum hermaphroditum has influence on the functioning of boreal terrestrial ecosystems in northern Sweden. The negative effects of E. hermaphroditum are partly attributed to the production of the dihydrostilbene, batatasin-III, which is released from leaves and litter by rain and snowmelt. In this study, we investigated whether batatasin-III is carried by runoff into streams and lakes during the snowmelt period and whether it is also potentially hazardous to aquatic fauna. Sampling of water from streams and a lake for which the surrounding terrestrial vegetation is dominated by E. hermaphroditum was done during the snowmelt period in May 1993 and in 1998, and analyzed for batatasin-III. Using 24- and 48-hr standard toxicity tests, we analyzed toxicity to brown trout (Salmo trutta) alevins and juvenile water fleas (Daphnia magna). Toxicity (proportion of dead individuals) to trout was tested at pH 6.5 and compared with that of a phenol within a range of concentrations. In the toxicity (proportion of immobilized individuals) test on D. magna, the interactive effect of pH (pH 5.5-7.0) was included. Concentration of batatasin-III was generally higher in 1998 than in 1993 and showed peak levels during snowmelt. Concentration in ephemeral runnels > the lake > streams running through clear-cuts dominated by E. hermaphroditum > control streams lacking adjacent E. hermaphroditum vegetation. The maximum concentration of batatasin-III found was 1.06 mg l(-1). The proportion of dead yolk sac alevins increased significantly (P < 0.001) with increasing concentrations of batatasin-III and time of exposure. After 24 hr, EC50 was 10 mg l(-1). It was 2 mg l(-1) after 48 hr. The effect of phenol was negligible, indicating a specific phytotoxic effect of the bibenzyl structure of batatasin-III. The proportion of mobile D. magna became significantly smaller (P < 0.001) with increasing concentrations of batatasin-III, with decreasing pH, and with increasing exposure time. EC50 varied between 7 and 17 mg l(-1) at pH 5.5 and 7.0, respectively. After 24 hr EC50 decreased and was 2.5 at pH 5.5 and 12 mg l(-1) at pH 7.0. The levels of batatasin-III found in the field samples were below the lowest EC50 in acute toxicity tests. However, in view of the interactive effect of pH and exposure time, this study suggests that this stable plant metabolite may impose a lethal effect on the aquatic fauna in small streams.

The use of plants as regular food in ancient subarctic economies: a case study based on Sami use of Scots pine innerbark.

This study combines ethnological, historical, and dendroecological data from areas north of the Arctic Circle to analyze cultural aspects of Sami use of Scots pine (Pinus sylvestris L.) inner bark as regular food. Bark was peeled in June when trees were at the peak of sapping, leaving a strip of undamaged cambium so the tree survived. As a result, it is possible to date bark-peeling episodes using dendrochronology. The paper argues that the use of Scots pine inner bark reflects Sami religious beliefs, ethical concerns, and concepts of time, all expressed in the process of peeling the bark. A well-developed terminology and a set of specially designed tools reveal the technology involved in bark peeling. Consistent patterns with respect to the direction and size of peeling scars found across the region demonstrate common values and standards. Peeling direction patterns and ceremonial meals relating to bark probably reflect ritual practices connected to the sun deity, Biejvve.

Long-term effects of wildfire on ecosystem properties across an island area gradient.

Boreal forest soils play an important role in the global carbon cycle by functioning as a large terrestrial carbon sink or source, and the alteration of fire regime through global change phenomena may influence this role. We studied a system of forested lake islands in the boreal zone of Sweden for which fire frequency increases with increasing island size. Large islands supported higher plant productivity and litter decomposition rates than did smaller ones, and, with increasing time since fire, litter decomposition rates were suppressed sooner than was ecosystem productivity. This contributes to greater carbon storage with increasing time since fire; for every century without a major fire, an additional 0.5 kilograms per square meter of carbon becomes stored in the humus.

Quantifying nitrogen-fixation in feather moss carpets of boreal forests.

Biological nitrogen (N) fixation is the primary source of N within natural ecosystems, yet the origin of boreal forest N has remained elusive. The boreal forests of Eurasia and North America lack any significant, widespread symbiotic N-fixing plants. With the exception of scattered stands of alder in early primary successional forests, N-fixation in boreal forests is considered to be extremely limited. Nitrogen-fixation in northern European boreal forests has been estimated at only 0.5 kg N ha(-1) yr(-1); however, organic N is accumulated in these ecosystems at a rate of 3 kg N ha(-1) yr(-1) (ref. 8). Our limited understanding of the origin of boreal N is unacceptable given the extent of the boreal forest region, but predictable given our imperfect knowledge of N-fixation. Herein we report on a N-fixing symbiosis between a cyanobacterium (Nostoc sp.) and the ubiquitous feather moss, Pleurozium schreberi (Bird) Mitt. that alone fixes between 1.5 and 2.0 kg N ha(-1) yr(-1) in mid- to late-successional forests of northern Scandinavia and Finland. Previous efforts have probably underestimated N-fixation potential in boreal forests.

Role of warm stratification in promoting germination of seeds of Empetrum hermaphroditum (Empetraceae), a circumboreal species with a stony endocarp.

The broad objective of this research was to define the role of warm (≥15°C) stratification in breaking dormancy in seeds with stony endocarps that require warm-plus-cold (∼0°-10°C) stratification for germination. This question was addressed using seeds (true seed + endocarp, hereafter called seeds) of Empetrum hermaphroditum. Only 2-5% of freshly matured seeds collected in September and October at five sites in Sweden germinated in light at daily alternating temperature regimes of 15°/6°, 20°/10°, and 25°/15°C. Dormancy was not due to impermeability of the stony endocarp surrounding each seed, and embryos did not grow prior to radicle emergence. Thus, seeds did not have physical, morphological, or morphophysiological dormancy. Long periods of either cold stratification (20 or 32 wk) or warm stratification (16 wk) resulted in a maximum of 22-38 and 10% germination, respectively, in light at 25°/15°C. After 12 wk warm stratification plus 20 wk cold stratification, 83-93% of the seeds germinated in light at the three temperature regimes. For a cold stratification period of 20 wk, germination increased with increase in length of the preceding warm stratification treatment. Gibberellic acid (GA(3)) promoted germination of 77-87% of the seeds. Based on dormancy-breaking requirements and response to GA(3), 62-78% of the seeds had intermediate physiological dormancy; the others had nondeep physiological dormancy. Contrary to suggestions of several other investigators that warm stratification is required to make the endocarp permeable to water via its breakdown by microorganisms, our results with E. hermaphroditum show that this is not the case. In this species, warm stratification is part of the dormancy-breaking requirement of embryos in seeds with intermediate physiological dormancy.