Forest disturbance by an ecosystem engineer : beaver in boreal forest landscapes

Natural disturbances are important for forest ecosystem dynamics and maintenance of biodiversity. In the boreal forest, large-scale disturbances such as wildfires and windstorms have been emphasized, while disturbance agents acting at smaller scales have received less attention. Especially in Europe beavers have long been neglected as forest disturbance agents because they were extirpated from most of their range centuries ago. However, now they are returning to many parts of their former distribution range. As a disturbance agent, beaver plays two roles: of an ecosystem engineer and of a herbivore. The engineering impact is realized through dam construction resulting in a transformation of an originally terrestrial ecosystem into an aquatic one. As herbivores, beavers affect stand structure and tree species composition by preferring deciduous trees over coniferous ones. After abandonment, a beaver pond gradually turns into a terrestrial habitat again. At well-drained sites, forest will return in due course, first dominated by deciduous trees. At poorly drained sites, moistness of beaver patches may result in fen development. We conclude that beaver has an important impact on ecosystem processes and biodiversity in boreal forest ecosystems because it creates and maintains a spatio-temporal mosaic of successional habitats and associated species communities that would otherwise not exist in the landscape.Peer reviewe

Representative boreal forest habitats in northern Europe, and a revised model for ecosystem management and biodiversity conservation

The natural range of variation of ecosystems provides reference conditions for sustainable management and biodiversity conservation. We review how the understanding of natural reference conditions of boreal forests in northern Europe has changed from earlier perceptions of even-aged dynamics driven by stand-replacing disturbances towards current understanding highlighting the role of non-stand-replacing disturbances and the resultant complex forest dynamics and structures. We show how earlier views and conceptual models of forest disturbance dynamics, including the influential ASIO model, provide estimates of reference conditions that are outside the natural range of variation. Based on a research synthesis, we present a revised forest reference model incorporating the observed complexity of ecosystem dynamics and the prevalence of old forests. Finally, we outline a management model and demonstrate its use in forest ecosystem management and show how regional conservation area needs can be estimated. We conclude that attaining favourable conservation status in northern Europe's boreal forests requires increasing emphasis on ecosystem management and conservation for old forest characteristics.Peer reviewe

Young and old forest in the boreal : critical stages of ecosystem dynamics and management under global change

Abstract The circumboreal forest encompasses diverse landscape structures, dynamics and forest age distributions determined by their physical setting, and historical and current disturbance regimes. However, due to intensifying forest utilisation, and in certain areas due to increasing natural disturbances, boreal forest age-class structures have changed rapidly, so that the proportion of old forest has substantially declined, while that of young post-harvest and post-natural-disturbance forest proportions have increased. In the future, with a warming climate in certain boreal regions, this trend may further be enhanced due to an increase in natural disturbances and large-scale use of forest biomass to replace fossil-based fuels and products. The major drivers of change of forest age class distributions and structures include the use of clearcut short-rotation harvesting, more frequent and severe natural disturbances due to climate warming in certain regions. The decline in old forest area, and increase in managed young forest lacking natural post-disturbance structural legacies, represent a major transformation in the ecological conditions of the boreal forest beyond historical limits of variability. This may introduce a threat to biodiversity, ecosystem resilience and long-term adaptive capacity of the forest ecosystem. To safeguard boreal forest biodiversity and ecosystem functioning, and to maintain the multiple services provided to societies by this forest biome, it is pivotal to maintain an adequate share and the ecological qualities of young post-disturbance stages, along with mature forest stages with old-growth characteristics. This requires management for natural post-disturbance legacy structures, and innovative use of diverse uneven-aged and continuous cover management approaches to maintain critical late-successional forest structures in landscapes

Ilmastonmuutos, monimuotoisuus ja metsien ekologinen kestävyys

Metsien ja niiden käytön roolista ilmastonmuutoksen hillinnässä on käyty vilkasta keskustelua. Tässä keskustelussa metsä on näyttäytynyt stabiilina ja ennustettavana systeeminä, jonka rakennetta ja hiilensidontaa voidaan säädellä ja muokata tarpeen mukaan metsien käsittelyn kautta (ks. esimerkiksi Suomen ilmastopaneelin raportit). Argumentointi on perustunut pääosin mallitarkasteluihin, joilla pyritään ennustamaan, miten metsät kasvavat ja sitovat hiiltä tulevaisuudessa suhteessa ympäristötekijöiden, lähinnä lämpötilan ja sadannan, ennustettuihin muutoksiin. Ilmastonmuutoksen vaikutuksia metsäekosysteemiin on kuitenkin tarkasteltu tällöin varsin yksipuolisesti

Conceptual models of forest dynamics in environmental education and management: keep it as simple as possible, but no simpler

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Tree diametric-species diversity is affected by human impact in old Scots pine dominated forest in boreal Fennoscandia

Background Forest structural and compositional variability is of fundamental importance for forest ecosystem functioning and species diversity. The purpose of this research was to examine how human impact has affected the compositional-structural diversity of mature pine-dominated boreal forest in boreal Fennoscandia. For this a new approach was used, based on the classification of tree sizes by the diameter at breast height (dbh) and tree species, resulting in a new variable, the diametric-species, the variation of which describes the compositional-structural diversity of the forest. This variable was used to compare the structural-compositional diversity among three forest classes with different degree of human influence, using rarefaction as the main tool of analysis, complemented by analyses based on common diversity indices. Results The results showed that the near-natural forest was the most diverse and the managed forest the least diverse. On the other hand, the diversity of near-natural and selectively logged forests were similar, suggesting that selectively logged forests are equal to the natural forest in their compositional-structural diversity. The analysis solely on tree species showed no significant differences among the forest classes of different human impact. The Shannon diversity index showed no significant difference between the forest classes for the diametric-species and tree species classifications only, but the Simpson index signaled a slight difference between the selectively logged and managed forest classes for the diametric-species classification. Furthermore, the Sorensen index detected a difference among forest classes in the diametric-species classification. Conclusions Forest utilization had an adverse impact on forest compositional-structural diversity of mature Scots pine forests. The analysis also shows that the novel approach based on diametric-species classification could be a useful tool for forest diversity analysis and comparison, especially in species-poor forests such as the boreal forest.Peer reviewe

Dead standing pine trees in a boreal forest landscape in the Kalevala National park, northern Fennoscandia : amount, population characteristics and spatial pattern

Background: After their death, Scots pine trees can remain standing for decades and sometimes up to 200 years, forming long-lasting and ecologically important structures in boreal forest landscapes. Standing dead pines decay very slowly and with time develop into 'kelo' trees, which are characterized by hard wood with silvery-colored appearance. These kelo trees represent an ecologically important, long lasting and visually striking element of the structure of natural pine-dominated forests in boreal Fennoscandia that is nowadays virtually absent from managed forest landscapes. Methods: We examined and mapped the amount, structural features, site characteristics and spatial distribution of dead standing pine trees over a ten hectare area in an unmanaged boreal forest landscape in the Kalevala National Park in Russian Viena Karelia. Results: The mean basal area of dead standing pine trees in the forested part of the landscape was 1.7 m(2).ha(-1) and the estimated volume 12.7 m(3).ha(-1). From the total number of standing dead pine trees 65% were kelo trees, with a basal area of 1.1 m(2).ha(-1) and volume of 8.0 m(3).ha(-1), the remainder consisting of standing dead pines along the continuum between a recently dead tree and a kelo tree. Overall, standing dead pines were distributed throughout the study area, but there was a tendency towards spatial clustering up to <100 m distances. Standing dead pines were most commonly situated on flat ground or in the mid slope in the local topography. In addition, standing dead pines contributed to substrate diversity also by commonly having charred wood and broken tops. Based on the presence of dead pine snags in different stage of transition from a recently dead pine to a kelo with silvery surface, it seems evident that the process of kelo recruitment was continuously in action in the studied landscape. Conclusions: Kelo trees are an omnipresent feature in natural pine-dominated forest landscapes with important contribution to forest structural and substrate diversity. Because of their longevity and extremely slow turnover dynamics and importance for biodiversity, protection of vulnerable kelo tree populations, and ensuring their continuous recruitment, should be of high priority in forest restoration and sustainable management.Peer reviewe

Preface to the special issue : The impact of ungulates and other mammalian herbivores on forest ecosystems

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North Fennoscandian mountain forests : History, composition, disturbance dynamics and the unpredictable future

North Fennoscandian mountain forests are distributed along the Scandes Mountains between Sweden and Norway, and the low-mountain regions of northern Norway, Sweden and Finland, and the adjacent northwestern Russia. Regionally, these forests are differentiated into spruce, pine or birch dominance due to climatic differences. Variation in tree species dominance within these regions is generally caused by a combination of historical and prevailing disturbance regimes, including both chronic and episodic disturbances, their magnitude and frequency, as well as differences in edaphic conditions and topography. Because of their remoteness, slow growth and restrictions of use, these mountain forests are generally less affected by human utilization than more productive and easily utilizable forests at lower elevations and/or latitudes. As a consequence, these northern forests of Europe are often referred to as "Europe's last wilderness", even if human influence of varying intensity has been ubiquitous through historical time. Because of their naturalness, the North Fennoscandian mountain forests are of paramount importance for biodiversity conservation, monitoring of ecosystem change and for their sociocultural values. As such, they also provide unique reference areas for basic and applied research, and for developing methods of forest conservation, restoration and ecosystem-based management for the entire Fennoscandia. However, the current rapid change in climate is predicted to profoundly affect the ecology and dynamics of these forests in the future. (C) 2016 Elsevier B.V. All rights reserved.Peer reviewe

Spatial tree community structure in three stands across a forest succession gradient in northern boreal Fennoscandia

Development of species composition during succession is well studied in natural boreal forests, but empirical assessments of how within-stand spatial structure develops in late-successional stages are few. Here, we quantified spatial patterns in three unmanaged stands consisting of Picea abies (L.) Karst. and Betula pubescens Ehrh. and Betula pendula Roth (hereafter Betula spp.) in northern boreal Fennoscandia. We conducted a comprehensive analysis of small-scale spatial point patterns in three fully mapped 1.2-ha sample plots, representing different forest developmental stages: mid-successional, late-successional and old-growth forest. We used several variants of Ripley’s K-function to analyze the spatial point patterns along the successional gradient. Univariate analyses showed that mature trees of both species were either randomly distributed or clumped. P. abies saplings were clumped, and Betula spp. saplings occurred in a random or clumped manner. In the bivariate analyses, saplings were more likely to be found in the surroundings of mature trees of the same species, but occurred independent of the individuals of other tree species. Mature trees showed interspecific repulsion. Only modest differences occurred in the univariate patterns between the three successional stages, but in the bivariate analyses the most evident patterns, i.e. intraspecific attraction and interspecific repulsion, were stronger in the older successional stages. Overall, the studied stands appear structured as species-specific mosaics. These mosaics, along with mixed species composition, seem to be maintained by species self-replacement, which contrasts with findings from earlier studies.Peer reviewe