Mari Jönsson

Demographics and disturbance history of a boreal old-growth Picea abies forest

Abstract Question: To what extent do tree growth, mortality, and long-term disturbance patterns affect stand structure and composition of an old-growth Picea abies forest? Location: Boreal Sweden. Methods: We linked data from three 50 m × 50 m permanent plots established in 1986 with dendrochronology data to evaluate tree growth and mortality over an 18-year period and to describe a several-hundred-year disturbance history for this forest type. Results: Averaged over all diameters, P. abies trees had an annual mortality rate of 0.60%; however, diameter had a striking effect on both growth and mortality, with trees of intermediate diameters (ca. 20–30 cm) showing faster growth and lower mortality. Their increased vigor gave rise to a diameter distribution resembling the ‘rotated sigmoid’ (not reverse-J) proposed for such conditions, and it led to a deficit of snags of intermediate diameters. Slow-growing trees had an increased likelihood of dying. Although recruitment occurred in most decades over the past 400 years, two prominent recruitment peaks occurred (mid 1700s and 1800s), neither of which appeared to cause a shift in tree species composition. The lack of fire evidence suggests that fire was not responsible for these recruitment peaks. Conclusions: Taken together, these results depict a rather impassive system, where canopy trees die slowly over decades. Field observations suggest that fungal infections, mediated by wind, account for much of the mortality during these periods of relative quiescence. However, these periods are at times punctuated by moderate-severity disturbances that foster abundant recruitment. Nomenclature: Mossberg & Stenberg (2003).

FUNGI AND WIND STRONGLY INFLUENCE THE TEMPORAL AVAILABILITY OF LOGS IN AN OLD-GROWTH SPRUCE FOREST

Coarse woody debris (CWD) is a key habitat for many species in forest ecosystems. To ensure the long-term survival of such species, forest management regimes must include measures that promote dead wood dynamics similar to those of natural forests. Thus, information on CWD dynamics under natural conditions is required, including data pertaining to the underlying agents of disturbance. This study examines modes of mortality, decay rates, and temporal patterns in the availability of Picea abies logs in a Swedish old-growth forest affected by internal, small-scale disturbance. All 684 logs in a 6.6-ha plot were mapped and classified into one of six decay classes. Logs in the early stages of decay were examined for the presence of heart-rot fungi. Six years later all logs were re-inventoried, including newly formed logs. Matrix models based on the transition rates between decay classes showed that it took about 60 years for 90% of the logs to decay beyond class 6 (a deformed trunk with soft wood). Large logs (> 26 cm) decayed 40% more slowly than small logs (< 25 cm). The initial volume of logs was 37.6 m3/ha but increased to 44.8 m3/ha after six years. In addition, there was a large shift in the decay-class distribution. The volume of logs in early and late decay classes increased by 71% and 45%, respectively, while the volume of logs in the intermediate decay classes decreased by 32%. The fluctuations appear to result from pulses in mortality, driven by a combination of strong winds and the heart-rot fungus, Phellinus chrysoloma, which was present in more than 30% of all logs at an early stage of decay. These results show that large temporal fluctuations in dead wood also occur in the absence of large-scale disturbance, and that heart-rot fungi are important factors driving the overall dynamics of dead wood. Since many wood-inhabiting species are naturally rare and have very specific substrate demands, such temporal variability in dead wood availability may have effects on biodiversity and should be taken into account when designing small, protected forest areas.

Lichen diversity and red-listed lichen species relationships with tree species and diameter in wooded meadows.

This study provides a unique large dataset of total epiphytic lichen diversity (fruticose, foliose and crustose species) and composition on 1,294 trees of 17 tree species in wooded meadows in Sweden and Estonia, the Baltic region. The inventory (25,380 observations and 246 lichen taxa) clearly illustrated that Ulmus minor, Quercus robur and Fraxinus excelsior contributed most significantly to epiphytic lichen richness and number of red-listed species. In Sweden, average single tree α richness was 22.2 on Ulmus (only in Sweden), 21.6 on Quercus (25.0 in Estonia) and 19.8 on Fraxinus (16.7 in Estonia), respectively. Ulmus hosted on average one red-listed species per tree, compared with 0.7 on Fraxinus (0.6 in Estonia), 0.4 on Quercus (0.7 in Estonia) and only 0.05 on Betula (same in Estonia). Lichen species composition and the average number of red-listed lichens were influenced by tree diameter on Fraxinus and Quercus, whilst no such pattern was evident on Ulmus. Randomized species accumulation curves of the dominating tree species illustrated that Fraxinus, Quercus and Ulmus supported α dominated lichen communities where individual trees hosted a substantial part of the total richness. Betula, on the other hand, supported β dominated communities where individual trees tended to be dissimilar and, therefore, more of the total richness existed as species turnover among host trees. Lichen species composition was influenced by tree species, and most notably, lichen species on Ulmus had a strong consistent clumping in ordination graphs, with many rare and red-listed lichens. The broadleaved deciduous trees within the wooded meadows clearly contribute greatly to the biodiversity of the Baltic region.

Estimating Coextinction Risks from Epidemic Tree Death: Affiliate Lichen Communities among Diseased Host Tree Populations of Fraxinus excelsior

At least 10% of the world’s tree species are threatened with extinction and pathogens are increasingly implicated in tree threats. Coextinction and threats to affiliates as a consequence of the loss or decline of their host trees is a poorly understood phenomenon. Ash dieback is an emerging infectious disease causing severe dieback of common ash Fraxinus excelsior throughout Europe. We utilized available empirical data on affiliate epiphytic lichen diversity (174 species and 17,800 observations) among 20 ash dieback infected host tree populations of F. excelsior on the island Gotland in the Baltic Sea, Sweden. From this, we used structured scenario projections scaled with empirical data of ash dieback disease to generate probabilistic models for estimating local and regional lichen coextinction risks. Average coextinction probabilities (Ā) were 0.38 (95% CI ±0.09) for lichens occurring on F. excelsior and 0.14 (95% CI ±0.03) when considering lichen persistence on all tree species. Ā was strongly linked to local disease incidence levels and generally increasing with lichen host specificity to F. excelsior and decreasing population size. Coextinctions reduced affiliate community viability, with significant local reductions in species richness and shifts in lichen species composition. Affiliates were projected to become locally extirpated before their hosts, illuminating the need to also consider host tree declines. Traditionally managed open wooded meadows had the highest incidence of ash dieback disease and significantly higher proportions of affiliate species projected to go extinct, compared with unmanaged closed forests and semi-open grazed sites. Most cothreatened species were not previously red-listed, which suggest that tree epidemics cause many unforeseen threats to species. Our analysis shows that epidemic tree deaths represent an insidious, mostly overlooked, threat to sessile affiliate communities in forested environments. Current conservation and management strategies must account for secondary extinctions associated with epidemic tree death.

Diverse ecological roles within fungal communities in decomposing logs of Picea abies

Fungal communities in Norway spruce (Picea abies) logs in two forests in Sweden were investigated by 454-sequence analyses and by examining the ecological roles of the detected taxa. We also investigated the relationship between fruit bodies and mycelia in wood and whether community assembly was affected by how the dead wood was formed. Fungal communities were highly variable in terms of phylogenetic composition and ecological roles: 1910 fungal operational taxonomic units (OTUs) were detected; 21% were identified to species level. In total, 58% of the OTUs were ascomycetes and 31% basidiomycetes. Of the 231 337 reads, 38% were ascomycetes and 60% basidiomycetes. Ecological roles were assigned to 35% of the OTUs, accounting for 62% of the reads. Wood-decaying fungi were the most common group; however, other saprotrophic, mycorrhizal, lichenized, parasitic and endophytic fungi were also common. Fungal communities in logs formed by stem breakage were different to those in logs originating from butt breakage or uprooting. DNA of specific species was detected in logs many years after the last recorded fungal fruiting. Combining taxonomic identification with knowledge of ecological roles may provide valuable insights into properties of fungal communities; however, precise ecological information about many fungal species is still lacking.

Biological legacies buffer local species extinction after logging.

Summary Clearcutting has been identified as a main threat to forest biodiversity. In the last few decades, alternatives to clearcutting have gained much interest. Living and dead trees are often retained after harvest to serve as structural legacies to mitigate negative effects of forestry. However, this practice is widely employed without information from systematic before–after control‐impact studies to assess the processes involved in species responses after clearcutting with retention. We performed a large‐scale survey of the occurrence of logging‐sensitive and red‐listed bryophytes and lichens before and after clearcutting with the retention approach. A methodology was adopted that, for the first time in studies on retention approaches, enabled monitoring of location‐specific substrates. We used uncut stands as controls to assess the variables affecting the survival of species after a major disturbance. In total, 12 bryophyte species and 27 lichen species were analysed. All were classified as sensitive to logging, and most species are also currently red‐listed. We found that living and dead trees retained after final harvest acted as refugia in which logging‐sensitive species were able to survive for 3 to 7 years after logging. Depending on type of retention and organism group, between 35% and 92% of the species occurrences persisted on retained structures. Most species observed outside retention trees or patches disappeared. Larger pre‐harvest population sizes of bryophytes on dead wood increased the survival probability of the species and hence buffered the negative effects of logging. Synthesis and applications. Careful spatial planning of retention structures is required to fully embrace the habitats of logging‐sensitive species. Bryophytes and lichens persisted to a higher degree in retention patches compared to solitary trees or in the clearcut area. Retaining groups of trees in logged areas will help to sustain populations of species over the clearcut phase. When possible, old logs should be moved into retention patches to provide a more beneficial environment for dead wood‐dependent species. Our study also highlights the need for more before–after control‐impact studies of retention forestry to explore factors influencing the survival of species after logging.

Evaluating citizen science data for forecasting species responses to national forest management

Abstract The extensive spatial and temporal coverage of many citizen science datasets (CSD) makes them appealing for use in species distribution modeling and forecasting. However, a frequent limitation is the inability to validate results. Here, we aim to assess the reliability of CSD for forecasting species occurrence in response to national forest management projections (representing 160,366 km2) by comparison against forecasts from a model based on systematically collected colonization–extinction data. We fitted species distribution models using citizen science observations of an old‐forest indicator fungus Phellinus ferrugineofuscus. We applied five modeling approaches (generalized linear model, Poisson process model, Bayesian occupancy model, and two MaxEnt models). Models were used to forecast changes in occurrence in response to national forest management for 2020‐2110. Forecasts of species occurrence from models based on CSD were congruent with forecasts made using the colonization–extinction model based on systematically collected data, although different modeling methods indicated different levels of change. All models projected increased occurrence in set‐aside forest from 2020 to 2110: the projected increase varied between 125% and 195% among models based on CSD, in comparison with an increase of 129% according to the colonization–extinction model. All but one model based on CSD projected a decline in production forest, which varied between 11% and 49%, compared to a decline of 41% using the colonization–extinction model. All models thus highlighted the importance of protected old forest for P. ferrugineofuscus persistence. We conclude that models based on CSD can reproduce forecasts from models based on systematically collected colonization–extinction data and so lead to the same forest management conclusions. Our results show that the use of a suite of models allows CSD to be reliably applied to land management and conservation decision making, demonstrating that widely available CSD can be a valuable forecasting resource.

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.

A decision support model for individual tree stump harvesting options based on criteria for economic return and environmental protection

ABSTRACT Based on principles of multi-criteria analysis techniques, a model (MAPStump-E) for decision support on stump harvesting at stand level was developed. The model applies the concept that each stump can be attributed production values (economic) and environmental values (here soil protection and water quality). Individual tree stump information was incorporated directly from the production reports of harvesters and combined with high-resolution Geographical Information System data on topography and soil type to create a production submodel and a soil and water vulnerability submodel (SWM). To test the model, it was applied to a 45-ha study forest in south-central Sweden and the outcome of nine scenarios with varying bioenergy prices and environmental protection levels was examined. Combined analysis of the effects of production and environmental criteria on total dry mass of harvestable stumps at the study site showed that biomass prices had a stronger influence than environmental criteria. Conflict stumps were defined as stumps suitable for harvest based on production criteria, but unsuitable based on soil and water protection criteria. In a “medium” price scenario, the proportion of conflict stumps at the study site ranged from 6% to 18%, depending on protection level set.

Transient trade-off between climate benefit and biodiversity loss of harvesting stumps for bioenergy

To replace fossil fuel and thereby mitigate climate change, harvesting of wood such as stumps for bioenergy will likely increase. Coarse deadwood is an important resource for biodiversity and stumps comprise the main part of the coarse deadwood in managed forests. We provide the first integrated analysis of the long‐term climate and biodiversity impacts of a whole landscape. We simultaneously project climate and biodiversity impacts of harvesting stumps to substitute for fossil coal, assuming scenarios with different proportions of the landscape with stump harvest (10, 50, 80%) the coming 50 years. A life cycle approach was used to calculate future global temperature changes and future metapopulation changes in six epixylic lichens. Metapopulation dynamics were projected using colonization and extinction models based on times series data. Harvesting stumps from ≥50% of the clear‐cut forest land benefits climate with a net global temperature reduction >0.5·10−9 K ha−1 after 50 years if assuming substitution of fossil coal. For all scenarios, using stump bioenergy leads to immediate (within 1 year) reductions in temperature of 50% compared to using fossil coal, increasing to 70% reduction after 50 years. However, large‐scale stump harvest inflicted substantial metapopulation declines for five of six lichens. High stump harvest levels (≥50%) put common lichens at risk of becoming red‐listed following the IUCN criteria. The net temperature reduction (cooling effect) from substituting fossil coal with stumps harvested for bioenergy increased over time, while lichen metapopulations stabilized at lower equilibria after two to three decades. This indicates that trade‐offs between climate and metapopulations of commons species are transient, where climate benefits become more prevalent in the long term. As both objectives are important for meeting (inter‐)national climate and biodiversity targets, integrated analyses such as this should be encouraged and urged to guide policymaking about large‐scale implementation of stump harvest.