Natural disturbance impacts on ecosystem services and biodiversity in temperate and boreal forests

In many parts of the world forest disturbance regimes have intensified recently, and future climatic changes are expected to amplify this development further in the coming decades. These changes are increasingly challenging the main objectives of forest ecosystem management, which are to provide ecosystem services sustainably to society and maintain the biological diversity of forests. Yet a comprehensive understanding of how disturbances affect these primary goals of ecosystem management is still lacking. We conducted a global literature review on the impact of three of the most important disturbance agents (fire, wind, and bark beetles) on 13 different ecosystem services and three indicators of biodiversity in forests of the boreal, cool‐ and warm‐temperate biomes. Our objectives were to (i) synthesize the effect of natural disturbances on a wide range of possible objectives of forest management, and (ii) investigate standardized effect sizes of disturbance for selected indicators via a quantitative meta‐analysis. We screened a total of 1958 disturbance studies published between 1981 and 2013, and reviewed 478 in detail. We first investigated the overall effect of disturbances on individual ecosystem services and indicators of biodiversity by means of independence tests, and subsequently examined the effect size of disturbances on indicators of carbon storage and biodiversity by means of regression analysis. Additionally, we investigated the effect of commonly used approaches of disturbance management, i.e. salvage logging and prescribed burning. We found that disturbance impacts on ecosystem services are generally negative, an effect that was supported for all categories of ecosystem services, i.e. supporting, provisioning, regulating, and cultural services (P < 0.001). Indicators of biodiversity, i.e. species richness, habitat quality and diversity indices, on the other hand were found to be influenced positively by disturbance (P < 0.001). Our analyses thus reveal a ‘disturbance paradox’, documenting that disturbances can put ecosystem services at risk while simultaneously facilitating biodiversity. A detailed investigation of disturbance effect sizes on carbon storage and biodiversity further underlined these divergent effects of disturbance. While a disturbance event on average causes a decrease in total ecosystem carbon by 38.5% (standardized coefficient for stand‐replacing disturbance), it on average increases overall species richness by 35.6%. Disturbance‐management approaches such as salvage logging and prescribed burning were neither found significantly to mitigate negative effects on ecosystem services nor to enhance positive effects on biodiversity, and thus were not found to alleviate the disturbance paradox. Considering that climate change is expected to intensify natural disturbance regimes, our results indicate that biodiversity will generally benefit from such changes while a sustainable provisioning of ecosystem services might come increasingly under pressure. This underlines that disturbance risk and resilience require increased attention in ecosystem management in the future, and that new approaches to addressing the disturbance paradox in management are needed.     

Maintaining historic disturbance regimes increases species' resilience to catastrophic hurricanes

As habitat loss and fragmentation, urbanization, and global climate change accelerate, conservation of rare ecosystems increasingly relies on human intervention. However, any conservation strategy is vulnerable to unpredictable, catastrophic events. Whether active management increases or decreases a system's resilience to these events remains unknown. Following Hurricane Irma's landfall in our habitat restoration study sites, we found that rare ecosystems with active, human‐imposed management suffered less damage in a hurricane's path than unmanaged systems. At the center of Irma's landfall, we found Croton linearis' (a locally rare plant that is the sole host for two endangered butterfly species) survival and population growth rates in the year of the hurricane were higher in previously managed plots than in un‐managed controls. In the periphery of Irma's circulation, the effect of prior management was stronger than that of the hurricane. Maintaining the historical disturbance regime thus increased the resilience of the population to major hurricane disturbance. As climate change increases the probability and intensity of severe hurricanes, human management of disturbance‐adapted landscapes will become increasingly important for maintaining populations of threatened species in a storm's path. Doing nothing will accelerate extinction.     

Exploring potential biodiversity indicators in boreal forests

The present study evaluates indicators in Swedish spruce forests. We ask whether different species groups co-vary in their occurrence and to what extent species richness and composition is predictable from habitat structures. We studied 10 boreal spruce forest stands constituting a gradient in degree of selective logging. Occurrences of vascular plants, bryophytes, epiphytic lichens and wood-inhabiting fungi as well as habitat structures was inventoried. In addition, in five of the stands, beetles were sampled with windows traps. Total species richness was correlated with several habitat factors, mainly particular substrates and degree of forestry impact. However, the richness of a set of species regularly used as indicators did not correlate with habitat factors. Correlation in species richness among different organism groups were few and scale dependent. Only lichens and vascular plants formed nested subset patterns (i.e. species composition at poorer sites is subsets of the species present at richer sites) among the study sites. The study shows that in this forest type one cannot a priori assume that richness in one group of species correlated with richness in other, and measures of single habitat features may be relevant only to particular groups of species. Instead, monitoring and inventories should be based on a set of factors reflecting important aspects for different groups of organisms and if indicator species are to be used these should be chosen from several species groups.     

The history and future of fungi as biodiversity surrogates in forests

Biodiversity surrogates are commonly used in conservation biology. Here we review how fungi have been used as such in forest conservation, emphasizing proposed surrogate roles and practical applications. We show that many fungal surrogates have been suggested based on field experience and loose concepts, rather than on rigorously collected scientific data. Yet, they have played an important role, not only in forest conservation, but also in inspiring research in fungal ecology and forest history. We argue that, even in times of ecosystem oriented conservation planning and molecular tools to analyze fungal communities, fruit bodies of macrofungi have potential as convenient conservation shortcuts and easy tools to communicate complex biodiversity for a broader audience. To improve the reliability of future fungal surrogates we propose a three step protocol for developing evidence based schemes for practical application in forest conservation.     

Vegetation dynamics and disturbance history in three deciduous forests in boreal Sweden

Abstract. Remaining deciduous forests in the Fennoscandian boreal landscape have high ecological value, and are considered as key components of the forest landscape as well as remnants of a former natural forest type. To improve our understanding of the formation of deciduous forests, we studied past disturbance regimes and vegetation dynamics in three deciduous forests in boreal Sweden using dendro‐ecology, pollen analysis and charcoal analysis. We identified three stages in the development of the studied stands. Firstly, the coniferous period (pre 1800), a long‐lasting period characterized by frequent fires, livestock grazing and extensive agriculture during which Pinus sylvestris was dominant. Secondly, the transformation period (1800 ‐ 1900), when logging removed most pines from the sites while fire and grazing continued. At the time of the last fire, the sites lacked a local seed source of pines, resulting in a post‐fire succession dominated by deciduous species with the capacity to disperse over long distances. Thirdly, the deciduous period (1900 ‐ present), with little or no disturbance from fire, grazing or logging. Thus, the present deciduous stands have their origins in a complex interaction between changes in fire regime, extensive land use patterns and logging, contrary to earlier simplified explanations. We conclude that the complexity of historical patterns of land use, vegetation dynamics and disturbance should be acknowledged in the future when selecting areas for nature conservation and developing models for ecologically oriented forestry.     

Significant increase in natural disturbance impacts on European forests since 1950

Over the last decades, the natural disturbance is increasingly putting pressure on European forests. Shifts in disturbance regimes may compromise forest functioning and the continuous provisioning of ecosystem services to society, including their climate change mitigation potential. Although forests are central to many European policies, we lack the long-term empirical data needed for thoroughly understanding disturbance dynamics, modeling them, and developing adaptive management strategies. Here, we present a unique database of >170,000 records of ground-based natural disturbance observations in European forests from 1950 to 2019. Reported data confirm a significant increase in forest disturbance in 34 European countries, causing on an average of 43.8 million m³ of disturbed timber volume per year over the 70-year study period. This value is likely a conservative estimate due to under-reporting, especially of small-scale disturbances. We used machine learning techniques for assessing the magnitude of unreported disturbances, which are estimated to be between 8.6 and 18.3 million m³/year. In the last 20 years, disturbances on average accounted for 16% of the mean annual harvest in Europe. Wind was the most important disturbance agent over the study period (46% of total damage), followed by fire (24%) and bark beetles (17%). Bark beetle disturbance doubled its share of the total damage in the last 20 years. Forest disturbances can profoundly impact ecosystem services (e.g., climate change mitigation), affect regional forest resource provisioning and consequently disrupt long-term management planning objectives and timber markets. We conclude that adaptation to changing disturbance regimes must be placed at the core of the European forest management and policy debate. Furthermore, a coherent and homogeneous monitoring system of natural disturbances is urgently needed in Europe, to better observe and respond to the ongoing changes in forest disturbance regimes.     

Restocking a keystone species in a biodiversity hotspot: Recovering the European rabbit on a landscape scale

Rabbit populations in Iberia remain at low densities in several areas in which their endangered predators still coexist, and the recovery of these populations is therefore urgent if the integrity of Iberian Mediterranean ecosystems is to be maintained. The enhancement of wild rabbit populations has been attempted through the use of in situ extensive rabbit captive breeding enclosures (restocking plots), which reduce mortality caused by terrestrial predators and dispersal movements and permit the breeding of young individuals which can then naturally disperse to settle in the surrounding areas. However, their effectiveness, the role of its size, the optimal habitat management that should be promoted around them and the habitat features remains uncertain. Here, we show results from a four year study of an ambitious rabbit restocking plan on a landscape scale. We measured rabbit abundance in a vast area in which thirty-two restocking plots were built to create an initial rabbit population for further dispersion, in addition to an intensive habitat management program. We also compared rabbit abundance between managed and unmanaged UTM cells of 2.5 km × 2.5 km. Our results showed that rabbit abundance was three times higher in managed cells, but four years after restocking, rabbit abundances had only reached the threshold needed to support stable Iberian lynx presence (at least 10 latrines per km^?1) in 9 of the 23 managed cells. Rabbit abundance was strongly affected by habitat treatment and scrub coverage. The increase of shelter was useful in low cover areas but ineligible in places with high scrub cover, where the increase of refuge plus scrub clearing to create pastures improve rabbit abundance more effectively. In the light of our results, restocking plots should be built only in places with suitable habitat, whereas pastures should be created in dense scrublands and refuge in low cover areas.     

Trade‐offs between carbon stocks and biodiversity in European temperate forests

Policies to mitigate climate change and biodiversity loss often assume that protecting carbon‐rich forests provides co‐benefits in terms of biodiversity, due to the spatial congruence of carbon stocks and biodiversity at biogeographic scales. However, it remains unclear whether this holds at the scales relevant for management, and particularly large knowledge gaps exist for temperate forests and for taxa other than trees. We built a comprehensive dataset of Central European temperate forest structure and multi‐taxonomic diversity (beetles, birds, bryophytes, fungi, lichens, and plants) across 352 plots. We used Boosted Regression Trees (BRTs) to assess the relationship between above‐ground live carbon stocks and (a) taxon‐specific richness, (b) a unified multidiversity index. We used Threshold Indicator Taxa ANalysis to explore individual species’ responses to changing above‐ground carbon stocks and to detect change‐points in species composition along the carbon‐stock gradient. Our results reveal an overall weak and highly variable relationship between richness and carbon stock at the stand scale, both for individual taxonomic groups and for multidiversity. Similarly, the proportion of win‐win and trade‐off species (i.e., species favored or disadvantaged by increasing carbon stock, respectively) varied substantially across taxa. Win‐win species gradually replaced trade‐off species with increasing carbon, without clear thresholds along the above‐ground carbon gradient, suggesting that community‐level surrogates (e.g., richness) might fail to detect critical changes in biodiversity. Collectively, our analyses highlight that leveraging co‐benefits between carbon and biodiversity in temperate forest may require stand‐scale management that prioritizes either biodiversity or carbon in order to maximize co‐benefits at broader scales. Importantly, this contrasts with tropical forests, where climate and biodiversity objectives can be integrated at the stand scale, thus highlighting the need for context‐specificity when managing for multiple objectives. Accounting for critical change‐points of target taxa can help to deal with this specificity, by defining a safe operating space to manipulate carbon while avoiding biodiversity losses.     

Fire disturbance and forest structure in old‐growth mixed conifer forests in the northern Sierra Nevada,California

Question: This study evaluates how fire regimes influence stand structure and dynamics in old‐growth mixed conifer forests across a range of environmental settings. Location: A 2000‐ha area of mixed conifer forest on the west shore of Lake Tahoe in the northern Sierra Nevada, California. Methods: We quantified the age, size, and spatial structure of trees in 12 mixed conifer stands distributed across major topographic gradients. Fire history was reconstructed in each stand using fire scar dendrochronology. The influence of fire on stand structure was assessed by comparing the fire history with the age, size, and spatial structure of trees in a stand. Results: There was significant variation in species composition among stands, but not in the size, age and spatial patterning of trees. Stands had multiple size and age classes with clusters of similar aged trees occurring at scales of 113 ‐ 254 m². The frequency and severity of fires was also similar, and stands burned with low to moderate severity in the dormant season on average every 9–17 years. Most fires were not synchronized among stands except in very dry years. No fires have burned since ca. 1880. Conclusions: Fire and forest structure interact to perpetuate similar stand characteristics across a range of environmental settings. Fire occurrence is controlled primarily by spatial variation in fuel mosaics (e.g. patterns of abundance, fuel moisture, forest structure), but regional drought synchronizes fire in some years. Fire exclusion over the last 120 years has caused compositional and structural shifts in these mixed conifer forests.     

Tree species dynamics and disturbance in three Swedish boreal forest stands during the last two thousand years

Abstract. Data from three forest stands for the past 2000 yr show how the shade-intolerant species Pinus sylvestris and Betula pubescens maintain significant populations in the Swedish boreal landscape. Age structure data from a northern stand close to the range limits of Picea abies and Pinus complement a local pollen diagram, and reveal cyclic population fluctuations which can be related to periods of climatic stress and fire. Pollen data from two southern stands show that high fire frequencies in the past prevented the expansion of Picea populations. Reduction of the fire frequency during the last 200 yr has favoured Picea. A long time perspective reveals the population dynamics of long-lived species and indicates the controlling factors. Such knowledge permits assessment of the current status and likely future of forest stands.     

The use of a functional approach as surrogate of Collembola species richness in European perennial crops and forests

Collembolans are known indicators of soil disturbance, used in several soil biodiversity monitoring programmes. As for other groups of soil microarthropods, taxonomic determination of Collembola species requires a huge effort and expert knowledge. In this study, we evaluated whether identification of Collembola species to the morphotype level, using an eco-morphological index which classifies individuals based on their adaptation to the soil, can be used as a surrogate of species richness in extensive monitoring schemes. The same evaluation was performed for higher taxa surrogates, using taxonomic categories at the genus and family levels. Additionally, sampling effort for perennial crops and forest systems was determined. Species data were collected from 35 sites sampled within different projects with the same number of samples (16 per site). Results showed that, on average, 8–12 samples are enough to have a good estimate of species richness for this type of systems, averaging 80–90%, but varying considerably with site habitat heterogeneity and local Collembola species pool. GLM models (using species richness as response variable) fitted for family, genus or morphotype levels (explanatory variables) were all significant (p < 0.05) but only the latter two had a pseudo R² higher than 0.75. This indicates that when a rapid, cost-effective assessment of Collembola richness in different sites is required, this eco-morphological trait approach, as well as the determination to the genus level, could be used.     

Understorey plant and soil responses to disturbance and increased nitrogen in boreal forests

Question: How do N fertilization and disturbance affect the understorey vegetation, microbial properties and soil nutrient concentration in boreal forests? Location: Kuusamo (66°22′N; 29°18′E) and Oulu (65°02′N; 25°47′E) in northern Finland. Methods: We conducted a fully factorial experiment with three factors: site (two levels), N fertilization (four levels) and disturbance (two levels). We measured treatment effects on understorey biomass, vegetation structure, and plant, soil and microbial N and C concentrations. Results: The understorey biomass was not affected by fertilization either in the control or in the disturbance treatment. Fertilization reduced the biomass of deciduous Vaccinium myrtillus. Disturbance had a negative effect on the biomass of V. myrtillus and evergreen Vaccinium vitis‐idaea and decreased the relative proportion of evergreen species. Fertilization and disturbance increased the biomass of grass Deschampsia flexuosa and the relative proportion of graminoids. The amount of NH₄⁺ increased in soil after fertilization, and microbial C decreased after disturbance. Conclusions: Our results suggest that the growth of slow‐growing Vaccinium species and soil microbes in boreal forests are not limited by N availability. However, significant changes in the proportion of dwarf shrubs to graminoids and a decrease in the biomass of V. myrtillus demonstrate the susceptibility of understorey vegetation to N enrichment. N enrichment and disturbance seem to have similar effects on understorey vegetation. Consequently, increasing N does not affect the rate or the direction of recovery after disturbance. Moreover, our study demonstrates the importance of understorey vegetation as a C source for soil microbes in boreal forests.     

Biodiversity and resilience of arthropod communities after fire disturbance in temperate forests

Changes in ecosystem functions following disturbances are of central concern in ecology and a challenge for ecologists is to understand the factors that affect the resilience of community structures and ecosystem functions. In many forest ecosystems, one such important natural disturbance is fire. The aim of this study was to understand the variation of resilience in six functional groups of invertebrates in response to different fire frequencies in southern Switzerland. We measured resilience by analysing arthropod species composition, abundance and diversity in plots where the elapsed time after single or repeated fires, as determined by dendrochronology, varied. We compared data from these plots with data from plots that had not burned recently and defined high resilience as the rapid recovery of the species composition to that prior to fire. Pooling all functional groups showed that they were more resilient to single fires than to repeated events, recovering 6–14 years after a single fire, but only 17–24 years after the last of several fires. Flying zoophagous and phytophagous arthropods were the most resilient groups. Pollinophagous and epigaeic zoophagous species showed intermediate resilience, while ground-litter saprophagous and saproxylophagous arthropods clearly displayed the lowest resilience to fire. Their species composition 17–24 years post-burn still differed markedly from that of the unburned control plots. Depending on the fire history of a forest plot, we found significant differences in the dominance hierarchy among invertebrate species. Any attempt to imitate natural disturbances, such as fire, through forest management must take into account the recovery times of biodiversity, including functional group composition, to ensure the conservation of multiple taxa and ecosystem functions in a sustainable manner.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Significance of woodland key habitats for polypore diversity and red-listed species in boreal forests

Preservation of woodland key habitats has become an integral part of biodiversity-oriented forest management in northern Europe. In Finland, brook-side spruce forests constitute the most important key-habitat type in terms of total area and timber volume. Our aim was to compare polypore diversity and the occurrence of red-listed species between brook-side key habitats and comparable sites of managed forest as controls. Furthermore, we assessed the importance of stand structural features to polypore communities, and contrasted regions with different land-use history. Altogether 69 key habitats and 70 controls were inventoried in four regions across southern Finland. A total of 28,023 surveyed dead-wood units had 3,307 occurrences of 114 species of which 25 species were red-listed. Key habitats hosted on average 28% (about two species) more polypore species (mainly deciduous-specialists) than controls because of a larger average volume of dead wood and a higher proportion of dead deciduous wood. However, the average number of red-listed species was low and did not differ between the two forest categories. We detected a landscape effect connected with the intensity and length of forest management history, with the highest numbers of all and red-listed species per site in eastern Finland. Volume and diversity of dead wood were the most important variables explaining variation in species richness. Presently, key habitats in managed forests appear to contribute only little to the conservation of red-listed polypore species.     

Sampling efficiency of the pale grass blue butterfly Zizeeria maha (Lepidoptera: Lycaenidae): A versatile indicator species for environmental risk assessment in Japan

The pale grass blue butterfly, Zizeeria maha (Lepidoptera, Lycaenidae), has successfully been employed as an indicator species to evaluate acute biological changes in polluted human-living environments after the Fukushima nuclear accident. Here, we quantitatively examined the field sampling efficiency of the butterfly and its associated sexual sampling bias under different conditions, which may provide important information for environmental risk assessment. Sampling data were obtained in 2014 and 2015 from 87 localities across Japan. Across the localities, the mean capture rate was 17 individuals per hour per person, and males represented approximately 80% of the collected butterflies. No significant difference was detected in the capture rate under different weather conditions. Among the four habitats, the riverside showed a significantly higher capture rate than the city park. A shorter sampling time tended to yield a higher capture rate. Comparisons among prefectures and districts of collection localities revealed that the Kanto district had significantly higher percentages of collected males than 4 other districts. Fukushima prefecture also had high percentage, although not significant. A generalized linear mixed model indicated that the capture rate was significantly negatively affected by the agricultural village, the city park, the rainy weather, and latitude and positively by longitude. Together, the present study showed high sampling efficiency and versatility of this butterfly as an indicator for environmental risk assessment but simultaneously suggested preferable conditions for a field sampling design. This study also suggested a possibility that the Fukushima nuclear accident or other factors might have affected behavioral or population dynamics of the butterfly.