Natural variability of forests as a reference for restoring and managing biological diversity in boreal Fennoscandia

In Fennoscandia, use of the natural forest as a reference for restoration and management of forest biodiversity has been widely accepted. However, limited understanding of the structure and dynamics of the natural forest has hampered the applications of the natural variability approach. This is especially the case in areas, where the natural forests have almost totally vanished. This review was motivated by the idea that despite these diffi culties the essential features of the natural forest can be reconstructed based on biological archives, historical documents, research done in adjacent natural areas, and modeling. First, a conceptual framework for analyzing the relationship between forest structure, dynamics and biodiversity is presented. Second, the current understanding of the structure and dynamics of natural forests at different spatiotemporal scales in boreal Fennoscandia is reviewed. Third, the implications of this knowledge, and gaps in knowledge, on research and on practical restoration and management methods aimed at forest biodiversity conservation are discussed. In conclusion, naturally dynamic forest landscapes are complex, multiscaled hierarchical systems. Current forest management methods create disturbance and successional dynamics that are strongly scale-limited when compared with the natural forest. To restore some of the essential characteristics of the natural forest’s multiscale heterogeneity, diversifi cation of silvicultural and harvesting treatments, as guided by natural disturbance dynamics, is needed to produce more variation in disturbance severity, quality, extent, and repeatability.

[1]  T. Kuuluvainen,et al.  Seedling establishment in relation to microhabitat variation in a windthrow gap in a boreal Pinus sylvestris forest , 1998 .

[2]  J. Niemelä,et al.  Forestry and the boreal fauna: matching management with natural forest dynamics , 1994 .

[3]  A. Gromtsev Natural disturbance dynamics in the boreal forests of European Russia : a review , 2002 .

[4]  T. Hokkanen,et al.  Properties of top soil and the relationship between soil and trees in a boreal Scots pine stand , 1995 .

[5]  J. Franklin Preserving Biodiversity: Species, Ecosystems, or Landscapes? , 1993, Ecological applications : a publication of the Ecological Society of America.

[6]  M. Ohlson,et al.  Stand dynamics, regeneration patterns and long-term continuity in boreal old-growth Picea abies swamp-forests , 1995 .

[7]  T. Pukkala,et al.  Factors related to seedling growth in a boreal Scots pine stand: a spatial analysis of a vegetation-soil system , 1993 .

[8]  L. Östlund,et al.  Retrospective gap analysis in a Swedish boreal forest landscape using historical data , 2001 .

[9]  T. Allen,et al.  Will similar forests develop on similar sites , 1985 .

[10]  M. Kuusinen Epiphyte Flora and Diversity on Basal Trunks of Six Old-Growth Forest Tree Species in Southern and Middle Boreal Finland , 1996, The Lichenologist.

[11]  T. Kuuluvainen Gap disturbance, ground microtopography, and the regeneration dynamics of boreal coniferous forests in Filand: a review , 1994 .

[12]  P. White,et al.  The Ecology of Natural Disturbance and Patch Dynamics , 1986 .

[13]  H. Morin,et al.  Key issues in disturbance dynamics in boreal forests: Introduction , 1998 .

[14]  Jerry F. Franklin,et al.  The diversity and maintenance of old-growth forests. , 1996 .

[15]  L. Ericson,et al.  Mosaic boreal landscapes with open and forested wetlands , 1997 .

[16]  Knut Rydgren,et al.  Single-tree influence on understorey vegetation in a Norwegian boreal spruce forest , 1999 .

[17]  E. Johnson,et al.  Wildfires in the western Canadian boreal forest: Landscape patterns and ecosystem management , 1998 .

[18]  L. Walker,et al.  Ecosystems of Disturbed Ground , 1999 .

[19]  Malcolm L. Hunter,et al.  Paleoecology and the Coarse-Filter Approach to Maintaining Biological Diversity , 1988 .

[20]  Per‐Anders Esseen,et al.  Epiphytic Lichen Biomass in Managed and Old-Growth Boreal Forests: Effect of Branch Quality , 1996 .

[21]  Lars Östlund,et al.  The history and transformation of a Scandinavian boreal forest landscape since the 19th century , 1997 .

[22]  B. Økland,et al.  What factors influence the diversity of saproxylic beetles? A multiscaled study from a spruce forest in southern Norway , 2004, Biodiversity & Conservation.

[23]  P. Gemmel,et al.  Development of nature-based silvicultural models in southern Sweden: the scientific background , 1994 .

[24]  T. Kuuluvainen,et al.  Structure and asymmetry of tree crowns in relation to local competition in a natural mature Scots pine forest , 1997 .

[25]  B. Palik,et al.  USING LANDSCAPE HIERARCHIES TO GUIDE RESTORATION OF DISTURBED ECOSYSTEMS , 2000 .

[26]  T. Kuuluvainen,et al.  Tree age distributions in old-growth forest sites in Vienansalo wilderness, eastern Fennoscandia , 2002 .

[27]  John H. Lawton,et al.  What Do Species Do in Ecosystems , 1994 .

[28]  Pauline Stenberg,et al.  Crown structure, light interception and productivity of pinetrees and stands , 1994 .

[29]  K. Woods Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests , 1984, Vegetatio.

[30]  A. Cajander,et al.  Theory of forest types , 1926 .

[31]  Robert K. Wayne,et al.  The preservation of process: the missing element of conservation programs , 1993 .

[32]  Julia A. Jones,et al.  Windthrow disturbance, forest composition, and structure in the Bull Run basin, Oregon , 2000 .

[33]  Herman H. Shugart,et al.  A Systems Analysis of the Global Boreal Forest: Contents , 1992 .

[34]  Antti Penttinen,et al.  Statistical opportunities for comparing stand structural heterogeneity in managed and primeval forests: An example from boreal spruce forest in southern Finland , 1996 .

[35]  M. Gilpin,et al.  Metapopulation Biology: Ecology, Genetics, and Evolution , 1997 .

[36]  Monica G. Turner,et al.  Maintaining Biodiversity in Forest Ecosystems: Dynamic forest mosaics , 1999 .

[37]  K. Rogers Operationalizing Ecology under a New Paradigm: An African Perspective , 1997 .

[38]  Gene E. Likens,et al.  The Ecological Basis of Conservation: Heterogeneity, Ecosystems, and Biodiversity , 1997 .

[39]  O. Zackrisson,et al.  Influence of forest fires on the North Swedish boreal forest , 1977 .

[40]  L. Musacchio Spatial Modeling of Forest Landscape Change: Approaches and Applications , 2000 .

[41]  James H. Brown,et al.  The Report of the Ecological Society of America Committee on the Scientific Basis for Ecosystem Management , 1996 .

[42]  M. Abrams,et al.  Multiple successional pathways on recently disturbed jack pine sites in Michigan , 1985 .

[43]  R. Heikkilä,et al.  Spatial tree age structure and fire history in two old-growth forests in eastern Fennoscandia , 2002 .

[44]  T. Kuuluvainen,et al.  Tree mortality in a Pinus sylvestris dominated boreal forest landscape in Vienansalo wilderness, eastern Fennoscandia , 2002 .

[45]  Malcolm L. Hunter,et al.  Maintaining Biodiversity in Forest Ecosystems: Contents , 1999 .

[46]  E. Hansen,et al.  Effects of a native forest pathogen, Phellinus weirii, on Douglas-fir forest composition in western Oregon , 1993 .

[47]  O. Laiho,et al.  Diversity-oriented silviculture in the Boreal Zone of Europe , 1999 .

[48]  Wiktor L. Adamowicz,et al.  Timber supply implications of natural disturbance management , 1999 .

[49]  Timo Kuuluvainen,et al.  Post-fire understorey regeneration in boreal Pinus sylvestris forest sites with different fire histories , 2000 .

[50]  Tapio Linkosalo,et al.  Estimation of a spatial tree-influence model using iterative optimization , 1998 .

[51]  A. Hofgaard Structure and regeneration patterns in a virgin Picea abies forest in northern Sweden , 1993 .

[52]  A. Axelsson Forest landscape change in boreal Sweden 1850-2000 , 2001 .

[53]  W. Baker Restoration of Landscape Structure Altered by Fire Suppression , 1994 .

[54]  T. Kuuluvainen,et al.  Structure of a pristine Picea abies forest in northeastern Europe , 1998 .

[55]  Mattias Carlsson,et al.  A review of conceptual landscape planning models for multiobjective forestry in Sweden , 1998 .

[56]  Martti E. Pesonen,et al.  University of Joensuu, Finland , 2001 .

[57]  O. Johansson,et al.  Silvicultural models to maintain and restore natural stand structures in Swedish boreal forests , 1997 .

[58]  T. Wallenius Forest age distribution and traces of past fires in a natural landscape dominated by Picea abies , 2002 .

[59]  P. Angelstam Maintaining and restoring biodiversity in European boreal forests by developing natural disturbance regimes , 1998 .

[60]  Alain Leduc,et al.  Forest management guidelines based on natural disturbance dynamics: Stand- and forest-level considerations , 1999 .

[61]  E. David Ford,et al.  A Model of Competition Incorporating Plasticity through Modular Foliage and Crown Development , 1993 .

[62]  P. Renvall Community structure and dynamics of wood-rotting Basidiomycetes on decomposing conifer trunks in northern Finland , 1995 .

[63]  Timo Kuuluvainen,et al.  Amount and spatial distribution of standing and downed dead trees in two areas of different fire history in a boreal Scots pine forest , 2001 .

[64]  J. Lawton,et al.  POSITIVE AND NEGATIVE EFFECTS OF ORGANISMS AS PHYSICAL ECOSYSTEM ENGINEERS , 1997 .

[65]  J. Pennanen Forest age distribution under mixed-severity fire regimes : a simulation-based analysis for middle boreal Fennoscandia , 2002 .

[66]  Pierre Dutilleul,et al.  Spatial Heterogeneity against Heteroscedasticity: An Ecological Paradigm versus a Statistical Concept , 1993 .

[67]  V. T. Parker,et al.  Ecological scale : theory and applications , 1999 .

[68]  R. Macarthur,et al.  The Theory of Island Biogeography , 1969 .

[69]  Carl J. Walters,et al.  Adaptive Management of Renewable Resources , 1986 .

[70]  D. DeAngelis,et al.  Individual-Based Models and Approaches in Ecology , 1992 .

[71]  Steven K. Friedman,et al.  Neighbourhood effects in forests: implications for within‐stand patch structure , 1998 .

[72]  Frederick J. Swanson,et al.  CONSERVING BIODIVERSITY IN MANAGED FORESTS : LESSONS FROM NATURAL FORESTS , 1991 .

[73]  T. Kuuluvainen,et al.  Amount and diversity of coarse woody debris within a boreal forest landscape dominated by Pinus sylvestris in Vienansalo wilderness, eastern Fennoscandia , 2002 .

[74]  Per‐Anders Esseen,et al.  Invertebrate communities in boreal forest canopies as influenced by forestry and lichens with implications for passerine birds , 1995 .

[75]  S. Levin THE PROBLEM OF PATTERN AND SCALE IN ECOLOGY , 1992 .

[76]  O. Engelmark Boreal forest disturbances , 1999 .

[77]  C. E. Van Wagner,et al.  Age-class distribution and the forest fire cycle , 1978 .

[78]  T. Niemelä,et al.  Interactions of fungi at late stages of wood decomposition. , 1995 .

[79]  Peter B. Reich,et al.  Minireviews: Neighborhood Effects, Disturbance Severity, and Community Stability in Forests , 1999, Ecosystems.

[80]  Ulf Dieckmann,et al.  A DYNAMICAL SYSTEM FOR NEIGHBORHOODS INPLANT COMMUNITIES , 2000 .

[81]  Mats Niklasson,et al.  NUMBERS AND SIZES OF FIRES: LONG-TERM SPATIALLY EXPLICIT FIRE HISTORY IN A SWEDISH BOREAL LANDSCAPE , 2000 .

[82]  B. Jonsson Availability of coarse woody debris in a boreal old‐growth Picea abies forest , 2000 .

[83]  E. Tuittila,et al.  Seedling establishment after prescribed burning of a clear-cut and a partially cut mesic boreal forest in southern Finland. , 1996 .

[84]  Alain Leduc,et al.  Natural fire regime : a guide for sustainable management of the Canadian boreal forest , 2002 .

[85]  J. Wiens The Emerging Role of Patchiness in Conservation Biology , 1997 .

[86]  David J. Mladenoff,et al.  A Systems Analysis of the Global Boreal Forest: The southern boreal–northern hardwood forest border , 1992 .

[87]  S. Kellomäki,et al.  Tree survival in wildfires. , 1993 .

[88]  R. J. Pool,et al.  Plant Succession. An Analysis of the Development of Vegetation , 1917 .

[89]  Risto Kalliola,et al.  Landscape structure and forest dynamics in subcontinental Russian European taiga , 1994 .

[90]  Lena Gustafsson,et al.  Dying and dead trees. A review of their importance for biodiversity , 1994 .

[91]  Douglas H. Deutschman,et al.  Details That Matter: The Spatial Distribution of Individual Trees Maintains Forest Ecosystem Function , 1995 .

[92]  P. Attiwill The disturbance of forest ecosystems: the ecological basis for conservative management , 1994 .

[93]  Lynne Boddy,et al.  Fungal decomposition of wood. Its biology and ecology. , 1988 .

[94]  R. Ostfeld,et al.  The Ecological Basis of Conservation: Heterogeneity, Ecosystems, and Biodiversity , 1999 .

[95]  Norman L. Christensen,et al.  Managing for Heterogeneity and Complexity on Dynamic Landscapes , 1997 .

[96]  P. White,et al.  The Ecology of Natural Disturbance and Patch Dynamics , 1986 .

[97]  W. J. Mattson,et al.  Dynamics of forest herbivory: quest for pattern and principle. , 1996 .

[98]  B. S. Lindgren,et al.  A conceptual model of biotic disturbance ecology in the central interior of B.C.: How forest management can turn Dr. Jekyll into Mr. Hyde , 2000 .

[99]  Robert K. Peet,et al.  Plant succession : theory and prediction , 1993 .

[100]  A. Granström Spatial and temporal variation in lightning ignitions in Sweden , 1993 .

[101]  Per Angelstam,et al.  Woodpecker assemblages in natural and managed boreal and hemiboreal forest: a review , 1994 .

[102]  Carl J. Walters,et al.  Large‐Scale Management Experiments and Learning by Doing , 1990 .

[103]  P. Zinke,et al.  The Pattern of Influence of Individual Forest Trees on Soil Properties , 1962 .

[104]  Gene E. Likens,et al.  The Ecological Basis of Conservation , 1997, Springer US.

[105]  Frederick J. Swanson,et al.  OVERVIEW OF THE USE OF NATURAL VARIABILITY CONCEPTS IN MANAGING ECOLOGICAL SYSTEMS , 1999 .

[106]  M. Nilsson,et al.  Regeneration Pulses and Climate-Vegetation Interactions in Nonpyrogenic Boreal Scots Pine Stands , 1995 .

[107]  Timo Saksa,et al.  Structure and yield of all-sized and even-sized Scots pine-dominated stands , 1994 .

[108]  Juha Siitonen,et al.  Forest management, coarse woody debris and saproxylic organisms: Fennoscandian boreal forests as an example , 2001 .

[109]  W. D. Hoskins,et al.  Local competition in a naturally established jack pine stand , 1989 .

[110]  H. Lehtonen,et al.  Forest Fire History in Viena Karelia, Russia , 2000 .

[111]  G. Sirén The development of Spruce forest on raw humus sites in northern Finland and its ecology. , 1955 .

[112]  Per‐Anders Esseen,et al.  Treefall disturbance maintains high bryophyte diversity in a boreal spruce forest. , 1990 .

[113]  J. Siitnen Decaying wood and saproxylic Coleoptera in two old spruce forests: a comparison based on two sampling methods , 1994 .

[114]  T. Kuuluvainen,et al.  Principles of ecological restoration of boreal forested ecosystems: Finland as an example , 2002 .

[115]  V. Radeloff,et al.  EFFECTS OF INTERACTING DISTURBANCES ON LANDSCAPE PATTERNS: BUDWORM DEFOLIATION AND SALVAGE LOGGING , 2000 .

[116]  Timo Kuuluvainen,et al.  Structural heterogeneity and spatial autocorrelation in a natural mature Pinus sylvestris dominated forest , 1998 .

[117]  Per Linder,et al.  Structural changes in three mid-boreal Swedish forest landscapes, 1885–1996 , 1998 .

[118]  D. Mladenoff,et al.  Spatial Modeling of Forest Landscape Change: Approaches and Applications , 1999 .

[119]  L. Söderström The occurrence of epixylic bryophyte and lichen species in an old natural and a managed forest stand in Northeast Sweden , 1988 .