The biological controls of soil carbon accumulation following wildfire and harvest in boreal forests: A review.

Boreal forests are frequently subjected to disturbances, including wildfire and clear-cutting. While these disturbances can cause soil carbon (C) losses, the long-term accumulation dynamics of soil C stocks during subsequent stand development is controlled by biological processes related to the balance of net primary production (NPP) and outputs via heterotrophic respiration and leaching, many of which remain poorly understood. We review the biological processes suggested to influence soil C accumulation in boreal forests. Our review indicates that median C accumulation rates following wildfire and clear-cutting are similar (0.15 and 0.20 Mg ha-1 year-1, respectively), however, variation between studies is extremely high. Further, while many individual studies show linear increases in soil C stocks through time after disturbance, there are indications that C stock recovery is fastest early to mid-succession (e.g. 15-80 years) and then slows as forests mature (e.g. >100 years). We indicate that the rapid build-up of soil C in younger stands appears not only driven by higher plant production, but also by a high rate of mycorrhizal hyphal production, and mycorrhizal suppression of saprotrophs. As stands mature, the balance between reductions in plant and mycorrhizal production, increasing plant litter recalcitrance, and ectomycorrhizal decomposers and saprotrophs have been highlighted as key controls on soil C accumulation rates. While some of these controls appear well understood (e.g. temporal patterns in NPP, changes in aboveground litter quality), many others remain research frontiers. Notably, very little data exists describing and comparing successional patterns of root production, mycorrhizal functional traits, mycorrhizal-saprotroph interactions, or C outputs via heterotrophic respiration and dissolved organic C following different disturbances. We argue that these less frequently described controls require attention, as they will be key not only for understanding ecosystem C balances, but also for representing these dynamics more accurately in soil organic C and Earth system models.

Diversity of forest structures important for biodiversity is determined by the combined effects of productivity, stand age, and management.

In forests, the amount and diversity of structural features with high value for biodiversity, such as large trees and dead wood, are affected by productivity, stand age, and forest management. For efficient conservation of forest biodiversity, it is essential to understand the combined effects of these drivers. We used data from the Swedish National Forest Inventory to study the combined effects of productivity, stand age, and management for wood production on structures with high value for biodiversity: tree species richness, large living trees, dead wood volume, and specific dead wood types. Forest management changed the relationship between productivity and amount or diversity of some of the structures. Most structures increased with productivity and stand age, but decreased due to management. The negative effect of management was greatest for structures occurring mainly in high-productivity forests, such as deciduous dead wood. Thus, biodiversity conservation should target high-productivity forests to preserve these structures.

Prescribed fire is an effective restoration measure for increasing boreal fungal diversity.

Intensive forestry practices have had a negative impact on boreal forest biodiversity; as a consequence, the need for restoration is pressing. Polypores (wood-inhabiting fungi) are key decomposers of dead wood, but, due to a lack of coarse woody debris (CWD) in forest ecosystems, many species are under threat. Here, we study the long-term effects on polypore diversity of two restoration treatments: creating CWD by felling whole trees and prescribed burning. This large-scale experiment is located in spruce-dominated boreal forests in southern Finland. The experiment has a factorial design (n = 3) including three levels of created CWD (5, 30, and 60 m3 ha-1 ) crossed with burning or no burning. In 2018, 16 years after launching the experiment, we inventoried polypores on 10 experimentally cut logs and 10 naturally fallen logs per stand. We found that overall polypore community composition differed between burned and unburned stands. However, only red-listed species abundances and richness were positively affected by prescribed burning. We found no effects of CWD levels created mechanically by felling of trees. We show, for the first time, that prescribed burning is an effective measure for restoring polypore diversity in a late-successional Norway spruce forest. Burning creates CWD with certain characteristics that differ from what is created by CWD restoration by felling trees. Prescribed burning promotes primarily red-listed species, demonstrating its effectiveness as a restoration measure to promote diversity of threatened polypore species in boreal forests. However, because the CWD that the burning creates will decrease over time, to be functional, prescribed burns need to be applied regularly on the landscape scale. Large-scale and long-term experimental studies, such as this one, are invaluable for establishing evidence-based restoration strategies.

Fire severity as a key determinant of aboveground and belowground biological community recovery in managed even-aged boreal forests.

Changes in fire regime of boreal forests in response to climate warming are expected to impact postfire recovery. However, quantitative data on how managed forests sustain and recover from recent fire disturbance are limited.Two years after a large wildfire in managed even-aged boreal forests in Sweden, we investigated how recovery of aboveground and belowground communities, that is, understory vegetation and soil microbial and faunal communities, responded to variation in the severity of soil (i.e., consumption of soil organic matter) and canopy fires (i.e., tree mortality).While fire overall enhanced diversity of understory vegetation through colonization of fire adapted plant species, it reduced the abundance and diversity of soil biota. We observed contrasting effects of tree- and soil-related fire severity on survival and recovery of understory vegetation and soil biological communities. Severe fires that killed overstory Pinus sylvestris promoted a successional stage dominated by the mosses Ceratodon purpureus and Polytrichum juniperinum, but reduced regeneration of tree seedlings and disfavored the ericaceous dwarf-shrub Vaccinium vitis-idaea and the grass Deschampsia flexuosa. Moreover, high tree mortality from fire reduced fungal biomass and changed fungal community composition, in particular that of ectomycorrhizal fungi, and reduced the fungivorous soil Oribatida. In contrast, soil-related fire severity had little impact on vegetation composition, fungal communities, and soil animals. Bacterial communities responded to both tree- and soil-related fire severity. Synthesis: Our results 2 years postfire suggest that a change in fire regime from a historically low-severity ground fire regime, with fires that mainly burns into the soil organic layer, to a stand-replacing fire regime with a high degree of tree mortality, as may be expected with climate change, is likely to impact the short-term recovery of stand structure and above- and belowground species composition of even-aged P. sylvestris boreal forests.

Increase in dead wood, large living trees and tree diversity, yet decrease in understory vegetation cover: The effect of three decades of biodiversity-oriented forest policy in Swedish forests.

In Sweden, the majority of forest area has been altered by industrial forestry over the decades. Almost 30 years ago, a shift towards biodiversity-oriented forest management practices occurred. Here we took advantage of long-term data collected by the Swedish National Forest Inventory to track developmental changes in forest structural components over this time. We assessed changes in structural components that play an important role in biodiversity (dead wood, large living trees, tree species composition, and understory vegetation) in four forest types with descending tiers of biodiversity protection: protected areas, woodland key habitats, low-productivity forests and production forests. Overall, we found a positive trend in the volumes of dead wood and large living trees, as well as in tree species diversity, while there was a general decline in understory vegetation coverage. Most observed changes were consistent with the intended outcomes of the current forest policy, adapted in the early 1990s. The implementation of retention forestry is likely driving some of the observed changes in forest structural components in the south. In contrast, we observed no changes in any of the focal structural components in the north, which could be attributed to the ongoing clear-cutting of forests previously managed less intensively. Dead wood and large living trees increased not only in managed, but also in unmanaged forests, likely reflecting historical management. The increased tree species diversity can be explained through current forest management practices that encourages maintenance of additional tree species. Decreasing understory vegetation coverage in both dense managed and unmanaged forests suggests that factors other than forestry contribute to the ongoing changes in understory vegetation in Swedish forests. Overall, the observed increase in structural components has not yet been reflected in documented improvements for red-listed forest species, which may be due to delays in species responses to small improvements, as well as a lack of detailed monitoring. Similarly, the increased availability of forest structural components might still be insufficient to meet the specific habitat requirements of red-listed species.

Do different growth rates of trees cause distinct habitat qualities for saproxylic assemblages?

In production forests, a common silvicultural objective is enhancing tree growth rates. The growth rate influences both mechanical and biochemical properties of wood, which may have an impact on dead wood inhabiting (i.e. saproxylic) species. In this study, we tested for the first time whether tree growth rates affect dead-wood associated assemblages in general and the occurrence of red-listed species in particular. We sampled saproxylic beetles (eclector traps) and fungi (DNA metabarcoding of wood samples) in dead trunks of Norway spruce (Picea abies), which had different growth rates within the same hemiboreal forests in Sweden. A high proportion of fungi showed a positive association to increasing tree growth. This resulted in higher fungal richness in fast-grown trees both at the trunk scale and across multiple studied trunks. Such patterns were not observed for saproxylic beetles. However, a set of species (both beetles and fungi) preferred slow-grown wood. Moreover, the total number of red-listed species was highest in slow-grown trunks. We conclude that dead wood from slow-grown trees hosts relatively fewer saproxylic species, but a part of these may be vulnerable to production forestry. It implies that slow-grown trees should be a target in nature conservation. However, where slow-grown trees are absent, for instance in forests managed for a high biomass production, increasing the volumes of dead wood from fast-grown trees may support many species.

Disturbance interval modulates the starting point for vegetation succession.

Increased frequency and new types of disturbances caused by global change calls for deepened insights into possible alterations of successional pathways. Despite current interest in disturbance interactions there is a striking lack of studies focusing on the implication of decreasing times between disturbances. We surveyed forest-floor vegetation (vascular plants and bryophytes) in a Pinus sylvestris-dominated, even-aged production forest landscape, unique because of the presence of stands under a precisely dated disturbance interval gradient, ranging from 0 to 123 yr between clearcutting and a subsequent megafire. Despite a dominance of early-successional species in all burned stands 5 yr after fire, progression of succession was linked to time since the preceding clearcutting disturbance. This was most clearly seen in increased frequency with time since clearcutting of the dominant, late-successional dwarf shrub Vaccinium myrtillus, with surviving rhizomes as an important mechanism for postfire recovery. Our results demonstrate the role of legacy species as significant drivers of succession. We conclude that the starting point for succession is modulated by disturbance interval, so that shortened intervals risk reducing development towards late-successional stages. We suggest that a decrease in long successional sequences caused by more frequent disturbances may represent a general pattern, relevant also for other forest types and ecosystems.

Increasing the amount of dead wood by creation of high stumps has limited value for lichen diversity.

Artificial creation of dead wood in managed forests can be used to mitigate the negative effects of forestry on biodiversity. For this to be successful, it is essential to understand the conservation value that the created dead wood has in comparison to naturally occurring dead wood, and, furthermore, where in the landscape addition of dead wood is most beneficial, i.e. how landscape composition influences species occurrence on dead wood. We examined these questions by surveying epixylic lichens on artificially created high stumps of Scots pine (Pinus sylvestris) in 3-17 years old clear-cuts. We compared lichen assemblages on high stumps to those on other types of pine dead wood in mature forests, and examined how stump age, the amount of dead wood at the clear-cuts, and landscape composition at 500 m - 2.5 km scale influenced the assemblages. In comparison to other dead wood types, high stumps hosted lower lichen richness and less variable assemblages containing mainly common generalist species. Species richness increased with stump age, whereas dead wood amount and landscape composition were not important; only the total amount of forests in the landscape had a minor positive effect. We conclude that at the studied timescale high stumps of Scots pine are not particularly valuable for epixylic lichens and provide a poor substitute for naturally occurring dead wood in mature forests, although their value may increase with age. Furthermore, directing dead wood creation to specific stands or landscapes does not appear beneficial for lichen biodiversity, given the minor effect of landscape composition found at scales below 2.5 km.

Partitioning the colonization and extinction components of beta diversity across disturbance gradients.

Changes in species diversity often result from species losses and gains. The dynamic nature of beta diversity (spatial variation in species composition) that derives from such temporal species turnover, however, has received relatively little attention. Here, we disentangled extinction and colonization components of beta diversity by using the sets of species that went locally extinct and that newly colonized the study sites. We applied this concept of extinction and colonization beta diversity to ground vegetation communities that have been repeatedly surveyed in forests where fire and harvesting were experimentally applied. We first found that fire and harvesting caused no effect on beta diversity 2 yr after the treatments. From this result, we might conclude that they did not alter the ways in which species assemble across space. However, when we analyzed the extinction and colonization beta diversity between pre-treatment and 2 yr after the treatments, both measures were found to be significantly lower in burned sites compared to unburned sites (i.e., the groups of excluded and newly colonized species both showed low beta diversity in the burned sites). These results indicate that the fire excluded similar subsets of species across space, making communities become more heterogeneous, but at the same time induced spatially uniform colonization of new species, causing communities to homogenize. Consequently, the effects of these two processes canceled each other out. The relative importance of extinction and colonization components per se also changed temporally after the treatments. Fire and harvesting showed synergetic negative impacts on extinction beta diversity between pre-treatment and 10 yr after the treatments. Overall, analyses using extinction and colonization beta diversity allowed us to detect nonrandom disassembly and reassembly dynamics in ground vegetation communities. Our results suggest that common practices of analyzing beta diversity at one point in time can mask significant variation driven by disturbance. Acknowledging the extinction-colonization dynamics behind beta diversity is essential for understanding the spatiotemporal organization of biodiversity.

Below-ground herbivory mitigates biomass loss from above-ground herbivory of nitrogen fertilized plants.

Herbivorous insects can influence grassland ecosystem functions in several ways, notably by altering primary production and nutrient turnover. Interactions between above- and belowground herbivory could affect these functions; an effect that might be modified by nitrogen (N) addition, an important global change driver. To explore this, we added above- (grasshoppers) and belowground (wireworms) insect herbivores and N into enclosed, equally composed, grassland plant communities in a fully factorial field experiment. N addition substantially altered the impact of above- and belowground herbivory on ecosystem functioning. Herbivory and N interacted such that biomass was reduced under above ground herbivory and high N input, while plant biomass remained stable under simultaneous above- and belowground herbivory. Aboveground herbivory lowered nutrient turnover rate in the soil, while belowground herbivory mitigated the effect of aboveground herbivory. Soil decomposition potential and N mineralization rate were faster under belowground herbivory at ambient N, but at elevated N this effect was only observed when aboveground herbivores were also present. We found that N addition does not only influence productivity directly (repeatedly shown by others), but also appears to influence productivity by herbivory mediated effects on nutrient dynamics, which highlights the importance of a better understanding of complex biotic interactions.

Half a century of multiple anthropogenic stressors has altered northern forest understory plant communities.

Boreal forests form the largest and least disturbed forest biome in the northern hemisphere. However, anthropogenic pressure from intensified forest management, eutrophication, and climate change may alter the ecosystem functions of understory vegetation and services boreal forests provide. Swedish forests span long gradients of climate, nitrogen deposition, and management intensity. This makes them ideal to study how the species composition and functions of other, more pristine, boreal forests might change under increased anthropogenic pressure. Moreover, the National Forest Inventory (NFI) has collected systematic data on Swedish forest vegetation since the mid-20th century. We use this data to quantify changes in vegetation types between two periods, 1953-1962 and 2003-2012. The results show changes in forest understory vegetation since the 1950s at scales not previously documented in the boreal biome. The spatial extent of most vegetation types changed significantly. Shade-adapted and nutrient-demanding species (those with high specific leaf area) have become more common at the expense of light-demanding and nutrient-conservative (low specific leaf area) species. The cover of ericaceous dwarf shrubs decreased dramatically. These effects were strongest where anthropogenic impacts were greatest, suggesting links to drivers such as nitrogen deposition and land-use change. These changes may impact ecosystem functions and services via effects on higher trophic levels and faster plant litter decomposition in the expanding vegetation types. This, in turn, may influence nutrient dynamics, and consequently ecosystem productivity and carbon sequestration.

Conservation value of low-productivity forests measured as the amount and diversity of dead wood and saproxylic beetles.

In many managed landscapes, low-productivity land comprises most of the remaining relatively untouched areas, and is often over-represented within protected areas. The relationship between the productivity and conservational value of a site is poorly known; however, it has been hypothesized that biodiversity increases with productivity due to higher resource abundance or heterogeneity, and that the species communities of low-productivity land are a nested subset of communities from more productive land. We tested these hypotheses for dead-wood-dependent beetles by comparing their species richness and composition, as well as the amount and diversity of dead wood, between low-productivity (potential forest growth <1 m3 ·ha-1 ·yr-1 ) and high-productivity Scots pine-dominated stands in Sweden. We included four stand types: stands situated on (1) thin soils and (2) mires (both low-productivity), (3) managed stands, and (4) unmanaged stands set aside for conservation purposes (both high-productivity). Beetle species richness and number of red-listed species were highest in the high-productivity set-asides. Species richness was positively correlated with the volume and diversity of dead wood, but volume appeared to be a better predictor than diversity for the higher species richness in set-asides. Beetle species composition was similar among stand types, and the assemblages in low-productivity stands were largely subsets of those in high-productivity set-asides. However, 11% of all species and 40% of red-listed species only occurred in high-productivity stands, while no species were unique to low-productivity stands. We conclude that low-productivity forests are less valuable for conservation than high-productivity forest land. Given the generally similar species composition among stand types, a comparable conservational effect could be obtained by setting aside a larger area of low-productivity forest in comparison to the high-productivity. In terms of dead wood volumes, 1.8-3.6 ha of low-productivity forest has the same value as 1 ha of unmanaged high-productivity forest. This figure can be used to estimate the conservation value of low-productivity forests; however, as high-productivity forests harbored some unique species, they are not completely exchangeable.

Coordination through databases can improve prescribed burning as a conservation tool to promote forest biodiversity.

Prescribed fires are a common nature conservation practice. They are executed by several parties with limited coordination among them, and little consideration for wildfire occurrences and habitat requirements of fire-dependent species. Here, we gathered data on prescribed fires and wildfires in Sweden during 2011-2015 to (i) evaluate the importance and spatial extent of prescribed fires compared to wildfires and (ii) illustrate how a database can be used as a management tool for prescribed fires. We found that on average only 0.006% (prescribed 65%, wildfires 35%) of the Swedish forest burns per year, with 58% of the prescribed fires occurring on clearcuts. Also, both wildfires and prescribed fires seem to be important for the survival of fire-dependent species. A national fire database would simplify coordination and make planning and evaluation of prescribed fires more efficient. We propose an adaptive management strategy to improve the outcome of prescribed fires.

Above- and belowground insect herbivory modifies the response of a grassland plant community to nitrogen eutrophication.

Understanding the role that species interactions play in determining the rate and direction of ecosystem change due to nitrogen (N) eutrophication is important for predicting the consequences of global change. Insects might play a major role in this context. They consume substantial amounts of plant biomass and can alter competitive interactions among plants, indirectly shaping plant community composition. Nitrogen eutrophication affects plant communities globally, but there is limited experimental evidence of how insect herbivory modifies plant community response to raised N levels. Even less is known about the roles of above- and belowground herbivory in shaping plant communities, and how the interaction between the two might modify a plant community's response to N eutrophication. We conducted a 3-yr field experiment where grassland plant communities were subjected to above- and belowground insect herbivory with and without N addition, in a full-factorial design. We found that herbivory modified plant community responses to N addition. Aboveground herbivory decreased aboveground plant community biomass by 21%, but only at elevated N. When combined, above- and belowground herbivory had a stronger negative effect on plant community biomass at ambient N (11% decrease) than at elevated N (4% decrease). In addition, herbivory shifted the functional composition of the plant community, and the magnitude of the shifts depended on the N level. The N and herbivory treatments synergistically conferred a competitive advantage to forbs, which benefited when both herbivory types were present at elevated N. Evenness among the plant species groups increased when aboveground herbivory was present, but N addition attenuated this increase. Our results demonstrate that a deeper understanding of how plant-herbivore interactions above and below ground shape the composition of a plant community is crucial for making reliable predictions about the ecological consequences of global change.

A cross-continental comparison of plant and beetle responses to retention of forest patches during timber harvest.

Timber harvest can adversely affect forest biota. Recent research and application suggest that retention of mature forest elements (retention forestry), including unharvested patches (or aggregates) within larger harvested units, can benefit biodiversity compared to clearcutting. However, it is unclear whether these benefits can be generalized among the diverse taxa and biomes in which retention forestry is practiced. Lack of comparability in methods for sampling and analyzing responses to timber harvest and edge creation presents a challenge to synthesis. We used a consistent methodology (similarly spaced plots or traps along transects) to investigate responses of vascular plants and ground-active beetles to aggregated retention at replicate sites in each of four temperate and boreal forest types on three continents: Douglas-fir forests in Washington, USA; aspen forests in Minnesota, USA; spruce forests in Sweden; and wet eucalypt forests in Tasmania, Australia. We assessed (1) differences in local (plot-scale) species richness and composition between mature (intact) and regenerating (previously harvested) forest; (2) the lifeboating function of aggregates (capacity to retain species of unharvested forest); and whether intact forests and aggregates (3) are susceptible to edge effects and (4) influence the adjacent regenerating forest. Intact and harvested forests differed in composition but not richness of plants and beetles. The magnitude of this difference was generally similar among regions, but there was considerable heterogeneity of composition within and among replicate sites. Aggregates within harvest units were effective at lifeboating for both plant and beetle communities. Edge effects were uncommon even within the aggregates. In contrast, effects of forest influence on adjacent harvested areas were common and as strong for aggregates as for larger blocks of intact forest. Our results provide strong support for the widespread application of aggregated retention in boreal and temperate forests. The consistency of pattern in four very different regions of the world suggests that, for forest plants and beetles, responses to aggregated retention are likely to apply more widely. Our results suggest that through strategic placement of aggregates, it is possible to maintain the natural heterogeneity and biodiversity of mature forests managed for multiple objectives.

Aboveground insect herbivory increases plant competitive asymmetry, while belowground herbivory mitigates the effect.

Insect herbivores can shift the composition of a plant community, but the mechanism underlying such shifts remains largely unexplored. A possibility is that insects alter the competitive symmetry between plant species. The effect of herbivory on competition likely depends on whether the plants are subjected to aboveground or belowground herbivory or both, and also depends on soil nitrogen levels. It is unclear how these biotic and abiotic factors interactively affect competition. In a greenhouse experiment, we measured competition between two coexisting grass species that respond differently to nitrogen deposition: Dactylis glomerata L., which is competitively favoured by nitrogen addition, and Festuca rubra L., which is competitively favoured on nitrogen-poor soils. We predicted: (1) that aboveground herbivory would reduce competitive asymmetry at high soil nitrogen by reducing the competitive advantage of D. glomerata; and (2), that belowground herbivory would relax competition at low soil nitrogen, by reducing the competitive advantage of F. rubra. Aboveground herbivory caused a 46% decrease in the competitive ability of F. rubra, and a 23% increase in that of D. glomerata, thus increasing competitive asymmetry, independently of soil nitrogen level. Belowground herbivory did not affect competitive symmetry, but the combined influence of above- and belowground herbivory was weaker than predicted from their individual effects. Belowground herbivory thus mitigated the increased competitive asymmetry caused by aboveground herbivory. D. glomerata remained competitively dominant after the cessation of aboveground herbivory, showing that the influence of herbivory continued beyond the feeding period. We showed that insect herbivory can strongly influence plant competitive interactions. In our experimental plant community, aboveground insect herbivory increased the risk of competitive exclusion of F. rubra. Belowground herbivory appeared to mitigate the influence of aboveground herbivory, and this mechanism may play a role for plant species coexistence.

A Pine Is a Pine and a Spruce Is a Spruce--The Effect of Tree Species and Stand Age on Epiphytic Lichen Communities.

With an increasing demand for forest-based products, there is a growing interest in introducing fast-growing non-native tree species in forest management. Such introductions often have unknown consequences for native forest biodiversity. In this study, we examine epiphytic lichen species richness and species composition on the trunks of non-native Pinus contorta and compare these to the native Pinus sylvestris and Picea abies in managed boreal forests in northern Sweden across a chronosequence of age classes. Overall, we recorded a total of 66,209 lichen occurrences belonging to 57 species in the 96 studied forest stands. We found no difference in species richness of lichens between stands of P. contorta and P. sylvestris, but stands of P. abies had higher total species richness. However, species richness of lichens in stands of P. abies decreased with increasing stand age, while no such age effect was detected for P. contorta and P. sylvestris. Lichen species composition progressively diverged with increasing stand age, and in 30-year-old stands all three tree species showed species-specific assemblages. Epiphytic lichen assemblages in stands of 30-year-old P. contorta were influenced by greater basal area, canopy closure, and average diameter at breast height, P. abies stands by higher branch density and canopy closure, and stands of P. sylvestris by greater bark crevice depth. Differences in lichen species richness and composition were mainly explained by canopy closure and habitat availability, and the greater canopy closure in mature P. abies stands promoted the colonization and growth of calicioid lichen species. Our results indicate that the non-native P. contorta have similar species richness as the native P. sylvestris. The main difference in lichen species richness and composition is between P. abies and Pinus spp. in managed forests of boreal Sweden.

Underdispersion and overdispersion of traits in terrestrial snail communities on islands.

Understanding and disentangling different processes underlying the assembly and diversity of communities remains a key challenge in ecology. Species can assemble into communities either randomly or due to deterministic processes. Deterministic assembly leads to species being more similar (underdispersed) or more different (overdispersed) in certain traits than would be expected by chance. However, the relative importance of those processes is not well understood for many organisms, including terrestrial invertebrates. Based on knowledge of a broad range of species traits, we tested for the presence of trait underdispersion (indicating dispersal or environmental filtering) and trait overdispersion (indicating niche partitioning) and their relative importance in explaining land snail community composition on lake islands. The analysis of community assembly was performed using a functional diversity index (Rao's quadratic entropy) in combination with a null model approach. Regression analysis with the effect sizes of the assembly tests and environmental variables gave information on the strength of under- and overdispersion along environmental gradients. Additionally, we examined the link between community weighted mean trait values and environmental variables using a CWM-RDA. We found both trait underdispersion and trait overdispersion, but underdispersion (eight traits) was more frequently detected than overdispersion (two traits). Underdispersion was related to four environmental variables (tree cover, habitat diversity, productivity of ground vegetation, and location on an esker ridge). Our results show clear evidence for underdispersion in traits driven by environmental filtering, but no clear evidence for dispersal filtering. We did not find evidence for overdispersion of traits due to diet or body size, but overdispersion in shell shape may indicate niche differentiation between snail species driven by small-scale habitat heterogeneity. The use of species traits enabled us to identify key traits involved in snail community assembly and to detect the simultaneous occurrence of trait underdispersion and overdispersion.

Rapid ecosystem shifts in peatlands: linking plant physiology and succession.

Stratigraphic records from peatlands suggest that the shift from a rich fen (calcareous fen) to an ombrotrophic bog can occur rapidly. This shift constitutes a switch from a species-rich ecosystem to a species-poor one with greater carbon storage. In this process, the invasion and expansion of acidifying bog species of Sphagnum (peat mosses) play a key role. To test under what conditions an acidifying bog species could invade a rich fen, we conducted three experiments, contrasting the bog species S. fucsum with the rich-fen species S. warnstorfii and S. teres. We first tested the effect of calcareous water by growing the three species at different constant height above the water table (HWT; 2, 7, and 14 cm) in a rich-fen pool and measured maximum photosynthetic rate and production and difference in length growth as an indicator of competition. In none of the species was the photosynthetic capacity negatively affected when placed at low HWT, but S. fuscum was a weaker competitor at low HWT. In our second experiment we transplanted the three species into microhabitats with different and naturally varying HWT in a rich fen. Here, S. fuscum nearly ceased to photosynthesize when transplanted to low HWT (brown moss carpet), while it performed similarly to the two rich-fen species at the intermediate level (S. warnstorfii hummock level). In contrast to S. fuscum, the rich-fen sphagna performed equally well in both habitats. The brown moss carpet was seasonally flooded, and in our third experiment we found that S. fuscum, but not S. teres, was severely damaged when submerged in rich-fen water. Our results suggest two thresholds in HWT affecting the ecosystem switch: one level that reduces the risk of submergence and a higher one that makes bog sphagna competitive against the rich-fen species.

Physiological responses to nitrogen and sulphur addition and raised temperature in Sphagnum balticum.

Sphagnum, the main genus which forms boreal peat, is strongly affected by N and S deposition and raised temperature, but the physiological mechanisms behind the responses are largely unknown. We measured maximum photosynthetic rate (NP(max)), maximum efficiency of photosystem II [variable fluorescence (F (v))/maximum fluorescence yield (F (m))] and concentrations of N, C, chlorophyll and carotenoids as responses to N and S addition and increased temperature in Sphagnum balticum (a widespread species in the northern peatlands) in a 12-year factorial experiment. NP(max) did not differ between control (0.2 g N m(-2) year(-1)) and high N (3.0 g N m(-2) year(-1)), but was higher in the mid N treatment (1.5 g N m(-2) year(-1)). N, C, carotenoids and chlorophyll concentration increased in shoot apices after N addition. F (v)/F (m) did not differ between N treatments. Increased temperature (+3.6 degrees C) had a small negative effect on N concentration, but had no significant effect on NP(max) or F (v)/F (m). Addition of 2 g S m(-2) year(-1) showed a weak negative effect on NP(max) and F (v)/F (m). Our results suggest a unimodal response of NP(max) to N addition and tissue N concentration in S. balticum, with an optimum N concentration for photosynthetic rate of ~13 mg N g(-1). In conclusion, high S deposition may reduce photosynthetic capacity in Sphagnum, but the negative effects may be relaxed under high N availability. We suggest that previously reported negative effects on Sphagnum productivity under high N deposition are not related to negative effects on the photosynthetic apparatus, but differences in optimum N concentration among Sphagnum species may affect their competitive ability under different N deposition regimes.