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Silva Fennica vol. 45 no. 5 | 2011

Special issue: Northern Primeval Forests & Ecology, Conservation and Management

Category : Research article

article id 90, category Research article

Per Angelstam, Kjell Andersson, Robert Axelsson, Marine Elbakidze, Bengt Gunnar Jonsson, Jean-Michel Roberge. (2011). Protecting forest areas for biodiversity in Sweden 1991–2010: the policy implementation process and outcomes on the ground. Silva Fennica vol. 45 no. 5 article id 90. https://doi.org/10.14214/sf.90

Keywords: forest policy; forest protection; restoration ecology; connectivity; green infrastructure; umbrella species; forest disturbance regimes; participation and collaboration

Abstract | View details | Full text in PDF | Author Info

Swedish forest and environmental policies imply that forests should be managed so that all naturally occurring species are maintained in viable populations. This requires maintenance of functional networks of representative natural forest and cultural woodland habitats. We first review the policy implementation process regarding protected areas in Sweden 1991–2010, how ecological knowledge was used to formulate interim short-term and strategic long-term biodiversity conservation goals, and the development of a hierarchical spatial planning approach. Second, we present data about the amount of formally protected and voluntarily set aside forest stands, and evaluate how much remains in terms of additional forest protection, conservation management and habitat restoration to achieve forest and environmental policy objectives in the long-term. Third, a case study in central Sweden was made to estimate the functionality of old Scots pine, Norway spruce and deciduous forest habitats, as well as cultural woodland, in different forest regions. Finally, we assess operational biodiversity conservation planning processes. We conclude that Swedish policy pronouncements capture the contemporary knowledge about biodiversity and conservation planning well. However, the existing area of protected and set-aside forests is presently too small and with too poor connectivity. To bridge this gap, spatial planning, management and restoration of habitat, as well as collaboration among forest and conservation planners need to be improved.

Angelstam, School for Forest Management, Faculty of Forest Sciences, Swedish University of Agricultural Sciences, Skinnskatteberg, Sweden E-mail: per.angelstam@slu.se
Andersson, School for Forest Management, Faculty of Forest Sciences, Swedish University of Agricultural Sciences, Skinnskatteberg, Sweden E-mail: ka@nn.se
Axelsson, School for Forest Management, Faculty of Forest Sciences, Swedish University of Agricultural Sciences, Skinnskatteberg, Sweden E-mail: ra@nn.se
Elbakidze, School for Forest Management, Faculty of Forest Sciences, Swedish University of Agricultural Sciences, Skinnskatteberg, Sweden E-mail: me@nn.se
Jonsson, Dept of Natural Science, Engineering and Mathematics, Mid Sweden University, Sundsvall, Sweden E-mail: bgj@nn.se
Roberge, Dept of Wildlife, Fish and Environmental Studies, Faculty of Forest Sciences, Swedish University of Agricultural Sciences, Umeå, Sweden E-mail: jmr@nn.se

article id 89, category Research article

Russell Grenfell, Tuomas Aakala, Timo Kuuluvainen. (2011). Microsite occupancy and the spatial structure of understorey regeneration in three late-successional Norway spruce forests in northern Europe. Silva Fennica vol. 45 no. 5 article id 89. https://doi.org/10.14214/sf.89

Keywords: boreal forest; disturbance; microsite; spatial correlation; stand structure; natural forest

Abstract | View details | Full text in PDF | Author Info

We compared microsite occupancy and three spatial structure of regeneration in three primeval late-successional Norway spruce dominated forests. One area lay in the middle boreal zone in Russia (Dvina-Pinega) where larger-scale disturbance from bark beetles and drought had occurred; the other areas lay in the northern boreal zone, one in Finland (Pallas-Ylläs) had encountered only small-scale disturbance, and one in Russia (Kazkim) had been influenced by fire. We mapped all spruce (Picea abies) and birch (Betula pendula and Betula pubescens) trees with diameter at breast height (DBH) ≥ 10 cm on 40 m 400 m plots, and those with DBH < 10 cm on 2 m or 4 m 400 m subplots. On the subplots we also recorded microsite occupancy and estimated microsite availability. At all study areas small seedlings (h < 0.3 m) of both spruce and birch were found largely on disturbance-related microsites. Birch saplings (h ≥ 1.3 m, DBH < 10 cm) disproportionately occupied deadwood-related microsites at Dvina-Pinega. In contrast, spruce saplings at all study areas, and birch saplings at Kazkim and Pallas-Ylläs, showed less, or no, preference. Our results thus confirm the importance of disturbance-related microsites for regeneration establishment, but not necessarily for long-term survival. No spatial segregation between the overstorey (DBH ≥ 10 cm) and saplings (h ≥ 1.3 m, DBH < 10 cm) or seedlings (h < 1.3 m) was found at Pallas-Ylläs or Kazkim, and only three instances of very weak segregation were found at Dvina-Pinega. This suggests that the regeneration gap concept may not be useful for describing the regeneration dynamics of primeval boreal forests.

Grenfell, University of Helsinki, Dept of Forest Sciences, Helsinki, Finland E-mail: russell.grenfell@helsinki.fi
Aakala, University of Helsinki, Dept of Forest Sciences, Helsinki, Finland E-mail: ta@nn.fi
Kuuluvainen, University of Helsinki, Dept of Forest Sciences, Helsinki, Finland E-mail: tk@nn.fi

article id 88, category Research article

Seppo Rouvinen, Jari Kouki. (2011). Tree regeneration in artificial canopy gaps established for restoring natural structural variability in a Scots pine stand. Silva Fennica vol. 45 no. 5 article id 88. https://doi.org/10.14214/sf.88

Keywords: ecosystems; diversity; restoration

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In Finland and elsewhere in Europe, many protected forest areas include also stands that were previously managed and that lack several naturally occurring stand characteristics. In these areas, ecosystem restoration can be used to facilitate and accelerate the formation of structural and habitat features resembling those of natural forests. For example, by creating small gaps it could be possible to diversify forest structure and tree species composition and to produce dead wood while still maintaining mostly continous canopy coverage. We examined experimentally the effects of artificial gap formation on post-disturbance tree regeneration in the gaps in a young protected, but formerly commercially managed pine (Pinus sylvestris L.) dominated forest. In the experimental sites, gap size and the portion of girdled trees out of all treated trees (girdled and felled trees combined) in the gaps varied. Natural and artificial (direct seeding of silver birch Betula pendula Roth) tree regeneration and development was monitored both on disturbed (scarified soil patches) and undisturbed forest floor during three growing seasons. Results show that gaps can be valuable in diversifying stand structure but to be successful and rapid, tree regeneration needs disturbed forest floor. Pine regenerated numerously, but birch had clearly lower regeneration, especially on small-sized gaps. In conclusion, increasing tree diversity in young pine-dominated forests seems to be difficult when only small artificial gaps are used. But even small gaps can be used to create and maintain diverse cohort structure of the dominant species and thus they can contribute to restoration goals.

Rouvinen, University of Eastern Finland, School of Forest Sciences, Joensuu, Finland E-mail: seppo.rouvinen@uef.fi
Kouki, University of Eastern Finland, School of Forest Sciences, Joensuu, Finland E-mail: jk@nn.fi

article id 87, category Research article

Radek Bace, Miroslav Svoboda, Pavel Janda. (2011). Density and height structure of seedlings in subalpine spruce forests of Central Europe: logs vs. stumps as a favourable substrate. Silva Fennica vol. 45 no. 5 article id 87. https://doi.org/10.14214/sf.87

Keywords: Norway spruce; Picea abies; coarse woody debris; microsite; natural regeneration

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Decaying logs and stumps provide an important seedling substrate in natural subalpine forests. However, only stumps present such a role in managed forests. The aim of this study was to assess the differences in the process of seedling colonization between logs and stumps. The study was carried out in the Czech Republic, in two old-growth subalpine spruce forests located in the Bohemian Forest and Ash Mts., dominated by Athyrium distentifolium Opiz and Vaccinium myrtillus L. undergrowth, respectively. Norway spruce (Picea abies (L.) Karst.) regeneration growing on logs, stumps and non-coarse woody debris (CWD) microsites was surveyed. Regeneration (height 0–2.0 m) densities exceeded 5000 individuals per ha on both sites. The average density of P. abies regeneration per square meter of substrate was 0.3-5.7-19.6 and 0.5-3.8-11.0 on non-CWD microsites, logs and stumps, located in A. distentifolium and V. myrtillus undergrowth, respectively. Stumps and non-CWD microsites dominated by V. myrtillus, supported a higher proportion of taller seedlings per plot compared to the small seedlings growing on logs and non-CWD dominated by A. distentifolium ground-cover. The disproportion in regeneration densities between the stumps and the original logs decreased with increasing stages of decay. The tallest regeneration growing on stumps (root-soil plates) was significantly older than that growing on the logs (stems). Based on these two latter findings, the stumps appeared to provide suitable seedling substrates several years earlier than the logs did. Therefore, we conclude that the stumps play a more important role (relative to their covered area, 21–28 m² ha^–1) in terms of suitable microsites for regeneration, than the logs do.

Bace, Czech University of Life Sciences, Faculty of Forestry and Wood Sciences, Prague, Czech Republic E-mail: bace@fld.czu.cz
Svoboda, Czech University of Life Sciences, Faculty of Forestry and Wood Sciences, Prague, Czech Republic E-mail: ms@nn.cz
Janda, Czech University of Life Sciences, Faculty of Forestry and Wood Sciences, Prague, Czech Republic E-mail: pv@nn.cz

article id 86, category Research article

Mats Jonsell, Jesper Hansson. (2011). Logs and stumps in clearcuts support similar saproxylic beetle diversity: implications for bioenergy harvest. Silva Fennica vol. 45 no. 5 article id 86. https://doi.org/10.14214/sf.86

Keywords: biodiversity; beetles; bioenergy; Coleoptera; logs; insects; stumps

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Stumps from clear cuts are increasingly used for bioenergy. Extracting this wood will reduce the habitat available for saproxylic (wood-living) organisms. As little is known about the species assemblages that will be affected, we investigated the diversity of saproxylic beetles in stumps on clear-felled sites and as a reference, we compared it with the diversity in downed logs. Stumps and logs of aspen (Populus tremula L.), birch (Betula pubescens Ehrh. and B. verrucosa Ehrh.[syn. B. pendula Roth]), spruce (Picea abies (L.) Karst.) and pine (Pinus sylvestris L.) were examined in clear cuts of two different ages: one summer old and 4–5 years old. The beetles were sampled by sieving bark (0.25 m²) peeled from the wood. The samples were taken in pairs of one log and one stump situated close together and of the same tree species, age since death and diameter. In total 3348 saproxylic beetles belonging to 124 species were found in 176 samples. The stumps had a similar number of species to the logs both as measured per sample and as an accumulated number. Exceptions were 4–5 years old wood of birch and pine where the number was significantly higher in the stumps. The number of red-listed species was also similar between stumps and logs. Species composition was more different between the stumps and logs of conifers than of deciduous trees. We conclude that clear-felled stumps have a diverse saproxylic insect fauna. This has to be taken into account if large scale extraction of logging stumps is implemented.

Jonsell, Swedish University of Agrarian Sciences, Dept of Ecology, Uppsala, Sweden E-mail: mats.jonsell@ekol.slu.se
Hansson, Swedish University of Agrarian Sciences, Dept of Ecology, Uppsala, Sweden E-mail: jh@nn.se

article id 85, category Research article

Ville A.O. Selonen, Maija Mussaari, Tero Toivanen, Janne S. Kotiaho. (2011). The conservation potential of brook-side key habitats in managed boreal forests. Silva Fennica vol. 45 no. 5 article id 85. https://doi.org/10.14214/sf.85

Keywords: forest management; saproxylic species; dead wood; vegetation

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Today, maintaining biodiversity is included in the targets of boreal forest management. A widespread approach in northern Europe is to identify and preserve woodland key habitats within managed forests. Woodland key habitats are expected to be patches that host populations of threatened and declining species, and the preservation of these patches is assumed to enable the persistence of the focal species in the landscape. In Finland, the criteria for selecting woodland key habitats are defined in the Finnish Forest Act, and the selection has been done by forest practitioners. Our objective was to determine whether the surroundings of boreal brooks and rivulets qualified as key habitats are truly different from brook-side habitats not granted the key habitat status, and whether the brook-side habitats of the two types differ from the forest matrix managed for timber production. We found that the two brook-side habitats were in most aspects rather alike but there was a difference in the composition of ground vegetation assemblages. In contrast, the control forests were distinct from the brook-sides in terms of dead wood, species richness and assemblages of polypores, species richness of epiphytic mosses, and the composition of beetle assemblages. We conclude that brook-sides in general provide an important habitat clearly diverging from the surrounding matrix but that the conservation value of the brook-sides granted the key habitat status may not be substantially larger than that of the brook-sides without the status.

Selonen, University of Jyväskylä, Department of Biological and Environmental Sciences, Lahti, Finland E-mail: ville.selonen@juy.fi
Mussaari, University of Jyväskylä, Department of Biological and Environmental Sciences, Lahti, Finland E-mail: mm@nn.fi
Toivanen, University of Jyväskylä, Department of Biological and Environmental Sciences, Lahti, Finland E-mail: tt@nn.fi
Kotiaho, University of Jyväskylä, Department of Biological and Environmental Sciences, Lahti, Finland E-mail: jsk@nn.fi

article id 84, category Research article

Asko Lõhmus, Piret Lõhmus. (2011). Old-forest species: the importance of specific substrata vs. stand continuity in the case of calicioid fungi. Silva Fennica vol. 45 no. 5 article id 84. https://doi.org/10.14214/sf.84

Keywords: forest management; conservation; continuity; habitat; lichen; limiting factor; structural diversity

Abstract | View details | Full text in PDF | Author Info

Appropriate conservation management of old-forest species depends on the causes of their old-forest affinity, which, however, are insufficiently known. Calicioid fungi are often considered old-forest dependent because of their special requirements for microhabitat, microclimate, and stand continuity for at least two tree generations. We demonstrate that, for several methodological or interpretational problems, published studies do not provide unequivocal evidence for such mechanisms and even for old-forest dependency of calicioids in general. We then analyse a large Estonian dataset (ca. 2300 records of 32 species) representing various management types and site types to answer whether old forests have more calicioid species, and any specific species, than could be expected for the substratum availability observed. Although old growth had more species and records than mature managed stands or cutover sites, those substratum types that occurred at roughly similar abundances also hosted comparable numbers of species in different management types. The characteristic substrata adding extra species to old growth were snags and root-plates of treefall mounds; wood surfaces in general comprised more than half of all calicioid records. Although substratum abundance did not fully explain the species-richness contrast between old growth and mature stands, additional evidence suggested that the unexplained variance is rather due to small-scale habitat characteristics than stand-scale continuity or microclimate. Finally, we review the evidence for old-forest affinity of calicioid species and distinguish a set of threatened species. We conclude that the availability of specific substrata is the main limiting factor for calicioid fungi in forests, and its quantitative and stochastic nature explains the large random and region-specific variation in the published lists of ‘old-forest species’.

Lõhmus, Department of Zoology, Institute of Ecology and Earth Sciences, University of Tartu, Vanemuise st. 46, EE-51014, Tartu, Estonia E-mail: asko.lohmus@ut.ee
Lõhmus, Department of Botany, Institute of Ecology and Earth Sciences, University of Tartu, Tartu, Estonia E-mail: pl@nn.ee

article id 83, category Research article

Inari Ylläsjärvi, Håkan Berglund, Timo Kuuluvainen. (2011). Relationships between wood-inhabiting fungal species richness and habitat variables in old-growth forest stands in the Pallas-Yllästunturi National Park, northern boreal Finland. Silva Fennica vol. 45 no. 5 article id 83. https://doi.org/10.14214/sf.83

Keywords: species-area relationship; species richness; old-growth forest; indicators; northern Fennoscandia; polyporaceae; polypores

Abstract | View details | Full text in PDF | Author Info

Indicators for biodiversity are needed for efficient prioritization of forests selected for conservation. We analyzed the relationships between 86 wood-inhabiting fungal (polypore) species richness and 35 habitat variables in 81 northern boreal old-growth forest stands in Finland. Species richness and the number of red-listed species were analyzed separately using generalized linear models. Most species were infrequent in the studied landscape and no species was encountered in all stands. The species richness increased with 1) the volume of coarse woody debris (CWD), 2) the mean DBH of CWD and 3) the basal area of living trees. The number of red-listed species increased along the same gradients, but the effect of basal area was not significant. Polypore species richness was significantly lower on western slopes than on flat topography. On average, species richness was higher on northern and eastern slopes than on western and southern slopes. The results suggest that a combination of habitat variables used as indicators may be useful in selecting forest stands to be set aside for polypore species conservation.

Ylläsjärvi, Rovaniemi University of Applied Sciences, School of Forestry and Rural Industries, Jokiväylä 11 c, FI-96300 Rovaniemi, Finland E-mail: inari.yllasjarvi@ramk.fi
Berglund, University of Helsinki, Department of Forest Sciences, Helsinki, Finland E-mail: hb@nn.fi
Kuuluvainen, University of Helsinki, Department of Forest Sciences, Helsinki, Finland E-mail: tk@nn.fi

article id 82, category Research article

Nicole J. Fenton, Yves Bergeron. (2011). Dynamic old-growth forests? A case study of boreal black spruce forest bryophytes. Silva Fennica vol. 45 no. 5 article id 82. https://doi.org/10.14214/sf.82

Keywords: disturbances; old-growth forest; succession; boreal; bryophytes; climax; gap dynamics

Abstract | View details | Full text in PDF | Author Info

Old-growth forests have sparked significant interest over the last twenty years and definitions have evolved from structure based to process based, acknowledging the diversity of forests that could be considered old growth. However studies frequently group all forests over a certain age into a single type, negating the dynamic processes that create old growth. In this study we examine a 2350-year chronosequence in boreal black spruce forests in northwestern Quebec to determine whether continued community change can be observed in the bryophyte layer. Bryophytes dominate the understory of boreal forests and influence ecosystem functioning, particularly in paludified forests where production exceeds decomposition in the organic layer. Community composition and richness changed throughout the chronosequence with no evidence of a steady state associated with an old-growth phase. In contrast the bryophyte community continued to evolve with multiple phases being evident. These results suggest that old-growth forests on the Clay Belt of northwestern Quebec and northeastern Ontario, Canada, should be regarded as part of the continuous gradient in forest development rather than a single state. This complicates conservation of these forests as multiple phases should be considered when planning forest reserves.

Fenton, Université du Québec en Abitibi-Témiscamingue, 445 Boulevard de l’Université, Rouyn-Noranda, Québec, Canada J9X 4E5 E-mail: nicole.fenton@uqat.ca
Bergeron, Université du Québec en Abitibi-Témiscamingue, 445 Boulevard de l’Université, Rouyn-Noranda, Québec, Canada J9X 4E5 E-mail: yb@nn.ca

article id 81, category Research article

Tuomas Aakala. (2011). Temporal variability of deadwood volume and quality in boreal old-growth forests. Silva Fennica vol. 45 no. 5 article id 81. https://doi.org/10.14214/sf.81

Keywords: forest dynamics; Picea abies; coarse woody debris; range of variability

Abstract | View details | Full text in PDF | Author Info

Reference deadwood volumes from natural forests for forest management and restoration are often derived from one-time measurements or from repeated measurements over short time-scales. Such an approach often assumes an equilibrium state between tree mortality and decomposition, which is questionable in many boreal forest ecosystems due to the occurrence of allogenic disturbances. Using a simulation model based on empirical estimates of tree mortality, disturbance chronologies and models of wood decay class dynamics, this study aimed at characterizing variability in the volume and quality of deadwood for the past 200 years. The variability of deadwood volumes in old-growth forests, arising from differences in disturbance regimes and differing decay rates, was exemplified in two areas of Picea abies-dominated forests in northern Europe. The results imply that with stable deadwood input and slow decay rates the deadwood volume may be in an equilibrium state. On the contrary, if moderate-severity disturbances occur such a state seems improbable. Both study areas displayed continuity in deadwood availability, although temporary paucity in the early decay classes with shortest residence times was also observed. The results stress the importance of understanding the dynamic nature of deadwood in old-growth forests, instead of the traditional view of deadwood as a static ecosystem component.

Aakala, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: tuomas.aakala@helsinki.fi

article id 80, category Research article

Mari T. Jönsson, Shawn Fraver, Bengt Gunnar Jonsson. (2011). Spatio-temporal variation of coarse woody debris input in woodland key habitats in central Sweden. Silva Fennica vol. 45 no. 5 article id 80. https://doi.org/10.14214/sf.80

Keywords: boreal forest; Picea abies; dendrochronology; disturbance dynamics; tree mortality; woodland key habitats; windstorms

Abstract | View details | Full text in PDF | Author Info

The persistence of many saproxylic (wood-living) species depends on a readily available supply of coarse woody debris (CWD). Most studies of CWD inputs address stand-level patterns, despite the fact that many saproxylic species depend on landscape-level supplies of CWD. In the present study we used dated CWD inputs (tree mortality events) at each of 14 Norway spruce (Picea abies) dominated woodland key habitat sites to analyze the spatial and temporal patterns of CWD additions between 1950 and 2002 within a small landscape in central Sweden. We found that inputs were episodic within sites, where local windstorms created pulses in CWD input. Pulses occurred simultaneously in many sites, yielding landscape-level synchrony of CWD input. These synchronous pulses, and importantly, the breaks between pulses, may have negative implications for saproxylic species that are dependent on large volume inputs of freshly killed Norway spruce. In addition, the inherent small size and relative isolation of these sites may further increase extinction risks due to stochastic events. However, background CWD input rates occurring between pulses varied substantially among sites, presumably the result of the sites’ varied histories and structural characteristics. This finding suggests that the different sites have varied abilities to provide habitat for saproxylic species during periods with low landscape-level input of CWD.

Jönsson, Department of Ecology, SLU, P.O. Box 7044, SE-750 07 Uppsala, Sweden (current); Department of Natural Sciences, Engineering and Mathematics, Mid Sweden University, Sundsvall, Sweden E-mail: mari.jonsson@slu.se
Fraver, U.S. Forest Service, Northern Research Station, Grand Rapids, Minnesota, USA (current); Department of Natural Sciences, Engineering and Mathematics, Mid Sweden University, Sundsvall, Sweden E-mail: sf@nn.us
Jonsson, Department of Natural Sciences, Engineering and Mathematics, Mid Sweden University, Sundsvall, Sweden E-mail: bgj@nn.se

article id 79, category Research article

Jean-Philippe Légaré, Christian Hébert, Jean-Claude Ruel. (2011). Alternative silvicultural practices in irregular boreal forests: response of beetle assemblages. Silva Fennica vol. 45 no. 5 article id 79. https://doi.org/10.14214/sf.79

Keywords: biodiversity; old-growth forest; irregular stands; selection cutting; coarse woody debris; sustainable management

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In the process of implementing sustainable management in the eastern Canadian boreal forest, we tested two selection cutting methods and compared them with two widely used practices in the boreal forest: clearcutting with protection of the advanced growth and soils and irregular shelterwood cutting leaving small merchantable stems. We used old-growth irregular stands as references in comparing the impact of these silvicultural treatments on the diversity and abundance of beetles. Three groups were targeted: saproxylic flying beetles, epigaeic saproxylic beetles and epigaeic non-saproxylic beetles. A sampling design including 320 pitfall traps and 80 multidirectional flight-interception traps was deployed in 2007. A total of 26 906 beetles was captured including 407 taxa distributed among 52 families. We found that clearcutting with protection of the advanced growth and soils and irregular shelterwood cutting leaving small merchantable stems had a greater impact on beetle communities than both selection cuttings. Canopy opening as well as the presence of snags and downed woody debris appear as important attributes for several saproxylic and non-saproxylic species. Beetle communities in selection cuttings remained more similar to those found in controls; these silvicultural treatments are new tools to implement ecosystemic and sustainable management in irregular boreal forests.

Légaré, Université Laval, Faculté de foresterie, de géographie et de géomatique, Pavillon Abitibi-Price, Québec, Canada E-mail: jpl@nn.ca
Hébert, Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, 1055 du P.E.P.S., P.O. Box 10380, Stn. Sainte-Foy, Québec (Québec), G1V 4C7, Canada E-mail: christian.hebert@rncan.gc.ca
Ruel, Université Laval, Faculté de foresterie, de géographie et de géomatique, Pavillon Abitibi-Price, Québec, Canada E-mail: jcr@nn.ca

article id 78, category Research article

Kris Vandekerkhove, Luc De Keersmaeker, Ruben Walleyn, Frank Köhler, Luc Crevecoeur, Leen Govaere, Arno Thomaes, Kris Verheyen. (2011). Reappearance of old-growth elements in lowland woodlands in northern Belgium: Do the associated species follow? Silva Fennica vol. 45 no. 5 article id 78. https://doi.org/10.14214/sf.78

Keywords: colonisation; saproxylic species; dead wood; very large trees

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The forest cover of the western European lowland plain has been very low for centuries. Remaining forests were intensively managed, and old-growth elements like veteran trees and coarse woody debris became virtually absent. Only over the last decades have these old-growth elements progressively redeveloped in parks, lanes and forests, and have now reached their highest level over the last 500–1000 years. Biodiversity associated with these old-growth elements makes up an important part of overall forest biodiversity. The ability of species to recolonise the newly available habitat is strongly determined by limitations in their dispersal and establishment. We analyse the current status and development of old-growth elements in Flanders (northern Belgium) and the process of recolonisation by means of specific cases, focussing on saproxylic fungi and saproxylic beetles. Our results show that ‘hotspots’ of secondary old growth, even isolated small patches, may have more potential for specialised biodiversity than expected, and may provide important new strongholds for recovery and recolonisation of an important share of old-growth related species.

Vandekerkhove, INBO, Gaverstraat 4, B-9500 Geraardsbergen, Belgium E-mail: kris.vandekerkhove@inbo.be
De Keersmaeker, INBO, Gaverstraat 4, B-9500 Geraardsbergen, Belgium E-mail: ldk@nn.be
Walleyn, INBO, Gaverstraat 4, B-9500 Geraardsbergen, Belgium E-mail: rw@nn.be
Köhler, Koleopterologisches Forschungsbüro, Bornheim, Germany E-mail: fk@nn.de
Crevecoeur, Genk, Belgium E-mail: lc@nn.be
Govaere, Agency of Nature and Forests, Brussels, Belgium E-mail: lg@nn.be
Thomaes, INBO, Gaverstraat 4, B-9500 Geraardsbergen, Belgium E-mail: at@nn.be
Verheyen, Ghent University, Laboratory of Forestry, Gontrode, Belgium E-mail: kv@nn.be

article id 77, category Research article

Annie Claude Bélisle, Sylvie Gauthier, Dominic Cyr, Yves Bergeron, Hubert Morin. (2011). Fire regime and old-growth boreal forests in central Quebec, Canada: an ecosystem management perspective. Silva Fennica vol. 45 no. 5 article id 77. https://doi.org/10.14214/sf.77

Keywords: black spruce-feather moss; fire history; ecosystem management; dendrochronology

Abstract | View details | Full text in PDF | Author Info

Boreal forest management in Eastern Canada has caused depletion and fragmentation of old-growth ecosystems, with growing impacts on the associated biodiversity. To mitigate impacts of management while maintaining timber supplies, ecosystem management aims to narrow the gap between natural and managed landscapes. Our study describes the fire history and associated natural old-growth forest proportions and distribution of a 5000 km² area located in the black spruce-feather moss forest of central Quebec. We reconstructed a stand-origin map using archival data, aerial photos and dendrochronology. According to survival analysis (Cox hazard model), the mean fire cycle length was 247 years for the 1734–2009 period. Age-class distribution modelling showed that old-growth forests were present on an average of 55% of the landscape over the last 275 years. The mean fire size was 10 113 ha, while most of the burned area was attributable to fires larger than 10 000 ha, leading to old-growth agglomerations of hundreds of square kilometres. In regards to our findings, we propose ecosystem management targets and strategies to preserve forest diversity and resilience.

Bélisle, Centre for Forest Research, Université du Québec à Montréal, Montréal, Québec, Canada E-mail: annieclaude_b@hotmail.com
Gauthier, Laurentian Forestry Centre, Canadian Forest Service, Natural Resources Canada, Sainte-Foy, Québec, Canada E-mail: sg@nn.ca
Cyr, Institut Québécois d’Aménagement de la Fort Feuillue, Université du Québec en Outaouais, Ripon, Québec, Canada E-mail: dc@nn.ca
Bergeron, Centre for Forest Research, Université du Québec à Montréal, Montréal, Québec, Canada & NSERC-UQAT-UQAM Industrial Chair in Sustainable Forest Management, Université du Québec en Abitibi-Témiscamingue, Rouyn-Noranda, Québec, Canada E-mail: yb@nn.ca
Morin, Département des Sciences Fondamentales, Université du Québec à Chicoutimi, Chicoutimi, Québec, Canada E-mail: hm@nn.can

article id 76, category Research article

Alessandra Bottero, Matteo Garbarino, Vojislav Dukic, Zoran Govedar, Emanuele Lingua, Thomas A. Nagel, Renzo Motta. (2011). Gap-phase dynamics in the old-growth forest of Lom, Bosnia and Herzegovina. Silva Fennica vol. 45 no. 5 article id 76. https://doi.org/10.14214/sf.76

Keywords: disturbance regime; Lom; old-growth forest; canopy gaps; beech-spruce-fir; small-scale

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We investigated forest canopy gaps in the mixed beech (Fagus sylvatica L.), silver fir (Abies alba Miller), and Norway spruce (Picea abies (L.) Karst.) old-growth forest of Lom in the Dinaric Mountains of Bosnia and Herzegovina. Gap size, age, gap fraction, gapmaker characteristics and the structure and composition of gapfillers were documented to investigate gap dynamics. The percentages of forest area in canopy and expanded gaps were 19% and 41%, respectively. The median canopy gap size was 77 m², and ranged from 11 to 708 m². Although there were many single tree-fall gaps, the majority had multiple gapmakers that were often in different stages of decay, suggesting gap expansion is important at the study site. Of the gapmakers recorded, 14% were uprooted stems, 60% snapped stems, and 26% were standing dead trees. Dendroecological analysis suggests that gap formation varied in time. The density of gapfillers was not correlated to gap size, and the species composition of gapfillers varied between seedling, sapling, and tree life stages. The results suggest that gaps are mainly formed by endogenous senescence of single canopy trees. Exogenous disturbance agents, most likely related to wind and snow, act mainly as secondary agents in breaking weakened trees and in expanding previously established gaps. Although the findings are partially consistent with other studies of gap disturbance processes in similar old-growth forests in central Europe, the observed gap dynamic places the Lom core area at the end of a gradient that ranges from forests controlled by very small-scale processes to those where large, stand replacing disturbances predominate.

Bottero, University of Turin, Department Agroselviter, Via Leonardo da Vinci 44, 10095 Grugliasco (TO), Italy E-mail: alessandra.bottero@unito.it
Garbarino, University of Turin, Department Agroselviter, Via Leonardo da Vinci 44, 10095 Grugliasco (TO), Italy E-mail: mg@nn.it
Dukic, University of Banja Luka, Faculty of Forestry, Banja Luka, Bosnia and Herzegovina E-mail: vd@nn.ba
Govedar, University of Banja Luka, Faculty of Forestry, Banja Luka, Bosnia and Herzegovina E-mail: zg@nn.ba
Lingua, University of Padua, Department of TeSAF, Legnaro (PD), Italy E-mail: el@nn.it
Nagel, University of Ljubljana, Department of Forestry and Renewable Forest Resources, Ljubljana, Slovenia E-mail: tan@nn.si
Motta, University of Turin, Department Agroselviter, Via Leonardo da Vinci 44, 10095 Grugliasco (TO), Italy E-mail: rm@nn.it

article id 75, category Research article

Silvia Lamedica, Emanuele Lingua, Ionel Popa, Renzo Motta, Marco Carrer. (2011). Spatial structure in four Norway spruce stands with different management history in the Alps and Carpathians. Silva Fennica vol. 45 no. 5 article id 75. https://doi.org/10.14214/sf.75

Keywords: Picea abies; LISA; old-growth forests; point pattern analyses; random mortality; spatial structure

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In Europe most Norway spruce (Picea abies (L.) Karst.) mountain forests have been altered by human activities, leading to a lack of reference condition concerning their original structure. Nonetheless, remnants of Norway spruce primeval forests still exist in the Carpathians. Our objective was to assess the differences in structure between managed and unmanaged stands, concerning diameter distributions, amount of standing deadwood, spatial distributions and spatial structure of trees. We established four permanent plots: one in a virgin forest in the Eastern Carpathians and three in a previously managed forest in the Alps. In each plot, species, DBH, and position of the live and dead standing trees were collected. Spatial distribution and structure of all the trees was analysed through several indices. In the Carpathians forest there are clear signs of natural density-dependent mortality processes whereas in the Alpine plots such dynamics are less evident. In these latter plots, the lower snags volume and the random trees spatial distribution can be considered the legacies of past management. Nonetheless, despite the different history of the four stands, they all seem to converge towards a similar spatial structure with the presence of groups (30–40 m) of trees of similar size.

Lamedica, Forest Ecology Research Unit, Dept TeSAF, University of Padova, Legnaro (PD), Italy E-mail: sl@nn.it
Lingua, Forest Ecology Research Unit, Dept TeSAF, University of Padova, Legnaro (PD), Italy E-mail: el@nn.it
Popa, Forest Research and Management Institute, Research Station for Norway Spruce Silviculture, Câmpulung Moldovenesc, Romania E-mail: ip@nn.ro
Motta, Department AGROSELVITER, University of Torino, Grugliasco (TO), Italy E-mail: rm@nn.it
Carrer, Forest Ecology Research Unit, Dept TeSAF, University of Padova, Legnaro (PD), Italy E-mail: marco.carrer@unipd.it

Category : Review article

article id 446, category Review article

Guntis Brumelis, Bengt Gunnar Jonsson, Jari Kouki, Timo Kuuluvainen, Ekaterina Shorohova. (2011). Forest naturalness in northern Europe: perspectives on processes, structures and species diversity. Silva Fennica vol. 45 no. 5 article id 446. https://doi.org/10.14214/sf.446

Keywords: disturbances; dead wood; natural forest; biological diversity

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Saving the remaining natural forests in northern Europe has been one of the main goals to halt the ongoing decline of forest biodiversity. To facilitate the recognition, mapping and efficient conservation of natural forests, there is an urgent need for a general formulation, based on ecological patterns and processes, of the concept of “forest naturalness”. However, complexity, structural idiosyncracy and dynamical features of unmanaged forest ecosystems at various spatio-temporal scales pose major challenges for such a formulation. The definitions hitherto used for the concept of forest naturalness can be fruitfully grouped into three dimensions: 1) structure-based concepts of natural forest, 2) species-based concepts of natural forest and 3) process-based concepts of natural forest. We propose that explicit and simultaneous consideration of all these three dimensions of naturalness can better cope with the natural variability of forest states and also aid in developing strategies for forest conservation and management in different situations. To become operational, criteria and indicators of forest naturalness need to integrate the three dimensions by combining species (e.g. red-listed-, indicator- and umbrella species) with stand and landscape level structural features that are indicative of disturbance and succession processes.

Brumelis, Faculty of Biology, University of Latvia, Kronvalda bulv. 4, Riga, LV-1586, Latvia; E-mail: guntis.brumelis@lu.lv
Jonsson, Department of Natural Sciences, Engineering and Mathematics, Mid Sweden University, Sundsvall, Sweden E-mail: bgj@nn.se
Kouki, School of Forest Sciences, University of Eastern Finland, Joensuu, Joensuu E-mail: jk@nn.fi
Kuuluvainen, Department of Forest Sciences, University of Helsinki, Finland E-mail: tk@nn.fi
Shorohova, Saint-Petersburg State Forest Academy, Saint-Petersburg, Russia & Finnish Forest Research Institute, Vantaa Research Unit, Vantaa, Finland E-mail: es@nn.ru

article id 74, category Review article

Philip J. Burton, S. Ellen Macdonald. (2011). The restorative imperative: challenges, objectives and approaches to restoring naturalness in forests. Silva Fennica vol. 45 no. 5 article id 74. https://doi.org/10.14214/sf.74

Keywords: afforestation; disturbance regime; ecological restoration; forest rehabilitation; native species; reclamation

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Many of the world’s forests are not primeval; forest restoration aims to reverse alterations caused by human use. Forest restoration (including reforestation and forest rehabilitation) is widely researched and practiced around the globe. A review of recent literature reveals some common themes concerning forest restoration motivations and methods. In some parts of the world, forest restoration aims mainly to re-establish trees required for timber or fuelwood; such work emphasizes the propagation, establishment and growth of trees, and equates with the traditional discipline of silviculture. Elsewhere, a recent focus on biocentric values adopts the goal of supporting full complements of indigenous trees and other species. Such ecosystem-based restoration approaches consider natural templates and a wide array of attributes and processes, but there remains an emphasis on trees and plant species composition. Efforts to restore natural processes such as nutrient cycling, succession, and natural disturbances seem limited, except for the use of fire, which has seen widespread adoption in some regions. The inherent challenges in restoring “naturalness” include high temporal and spatial heterogeneity in forest conditions and natural disturbances, the long history of human influence on forests in many regions of the world, and uncertainty about future climate and disturbance regimes. Although fixed templates may be inappropriate, we still have a reasonably clear idea of the incremental steps required to make forests more natural. Because most locations can support many alternative configurations of natural vegetation, the restoration of forest naturalness necessarily involves the setting of priorities and strategic directions in the context of human values and objectives, as informed by our best understanding of ecosystem structure and function now and in the future.

Burton, Canadian Forest Service, Natural Resources Canada, 3333 University Way, Prince George, British Columbia, Canada V2N 4Z9 E-mail: Phil.Burton@NRCan-RNCan.gc.ca
Macdonald, Department of Renewable Resources, University of Alberta, Edmonton, Alberta, Canada E-mail: sem@nn.ca

article id 73, category Review article

Timo Kuuluvainen, Tuomas Aakala. (2011). Natural forest dynamics in boreal Fennoscandia: a review and classification. Silva Fennica vol. 45 no. 5 article id 73. https://doi.org/10.14214/sf.73

Keywords: disturbance; age structure; stand replacement; cohort; gap; patch; natural disturbance emulation

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The aim here was to review and summarize the findings of scientific studies concerning the types of forest dynamics which occur in natural forests (i.e. forests with negligible human impact) of boreal Fennoscandia. We conducted a systematic search for relevant studies from selected reference databases, using search terms describing the location, structure and processes, and degree of naturalness of the forest. The studies resulting from these searches were supplemented with other known works that were not indexed in the databases. This procedure yielded a total of 43 studies. The studies were grouped into four types of forest dynamics according to the information presented on the characteristics of the native disturbance-succession cycle: 1) even-aged stand dynamics driven by stand-replacing disturbances, 2) cohort dynamics driven by partial disturbances, 3) patch dynamics driven by tree mortality at intermediate scales (> 200 m²) and 4) gap dynamics driven by tree mortality at fine scales (< 200 m²). All four dynamic types were reported from both spruce and pine dominated forests, but their commonness differed. Gap dynamics was most commonly reported in spruce forests, and cohort dynamics in pine forests. The studies reviewed provide the best obtainable overall picture of scientific findings concerning the characteristics and variability of the unmanaged boreal forest dynamics in Fennoscandia. The results demonstrate that the unmanaged Fennoscandian forests are characterized by more diverse and complex dynamics than has traditionally been acknowledged.

Kuuluvainen, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: timo.kuuluvainen@helsinki.fi
Aakala, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: ta@nn.fi

article id 72, category Review article

Ekaterina Shorohova, Daniel Kneeshaw, Timo Kuuluvainen, Sylvie Gauthier. (2011). Variability and dynamics of old-growth forests in the circumboreal zone: implications for conservation, restoration and management. Silva Fennica vol. 45 no. 5 article id 72. https://doi.org/10.14214/sf.72

Keywords: late-successional forest; disturbance regimes; stand dynamic types; cohort dynamics

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Due to the unprecedented loss of old-growth forests to harvesting throughout circumboreal regions an understanding of similarities and differences in old-growth dynamics is needed to design effective restoration, management and conservation efforts. This paper reviews concepts, prevalence and variability of old-growth forests across landscapes, and evaluates different stand scale dynamics at the old-growth stage across the circumboreal zone. Old-growth historically dominated many boreal forest landscapes in both Eurasia and North America. Throughout much of North America, and to some extent in western Siberia, the natural prevalence and development of old-growth forests are regulated by the occurrence of stand-replacing fires. In eastern North America and Siberia, insect outbreaks may, however, be more important. Insect outbreaks as well as recurrent non-stand replacing surface fires and windthrows, when occurring at the old-growth stage, often form stands characterized by several tree age-class cohorts. This multi age-class forest development type is common in Europe and eastern Siberia but its prevalence and importance in boreal North-America is not well documented. Similarities in successional dynamics across the circumboreal region are found in the development of mono-dominant even-aged stands, the replacement of shade intolerant tree species by shade tolerant species, as well as in all-aged stands driven by small-scale gap dynamics. The message to land managers is that the focus should not only be on setting aside remaining old-growth forests or in restoring static old-growth attributes, but also in emulating natural disturbances and successional dynamics at landscape and regional scales to maintain natural variability in old-growth attributes through time.

Shorohova, Saint-Petersburg State Forest University, Saint-Petersburg, Russia & Finnish Forest Research Institute, Vantaa Research Unit, Vantaa, Finland (ekaterina.shorohova@metla.fi) E-mail: shorohova@ES13334.spb.edu
Kneeshaw, Université du Québec à Montréal, Centre d’étude de la forêt, Montreal, Canada E-mail: dk@nn.ca
Kuuluvainen, University of Helsinki, Department of Forest Sciences, Helsinki, Finland E-mail: tk@nn.fi
Gauthier, Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, Québec, Canada E-mail: sg@nn.fi

Category : Research note

article id 92, category Research note

Adam Felton, Erik Andersson, David Ventorp, Matts Lindbladh. (2011). A comparison of avian diversity in spruce monocultures and spruce-birch polycultures in southern Sweden. Silva Fennica vol. 45 no. 5 article id 92. https://doi.org/10.14214/sf.92

Keywords: biodiversity; conservation; sustainable forestry

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The replacement of some spruce monocultures with stands composed of planted Norway spruce (Picea abies) and naturally regenerated birch (Betula spp.) has a range of potential benefits, but the implications for biodiversity are generally unknown. Here we conduct a paired replicated study in southern Sweden of the avian biodiversity found within Norway spruce monocultures, and within Norway spruce stands possessing approximately 20% birch. Our research leads us to three findings. First, avian diversity was significantly higher in the spruce–birch polycultures. Second, spruce–birch polycultures exclusively attracted broadleaf-associated bird species and retained the majority of conifer-associated bird species found in the spruce monocultures. Third, avian biodiversity within the spruce–birch polycultures did not incorporate threatened taxa. We suggest that in addition to the apparent benefits for stand level diversity, widespread use of spruce–birch polycultures could provide a means of softening the matrix for broadleaved-associated species, while concurrently providing an increased broadleaf base from which future conservation actions could be implemented. Our results are relevant to multi-use forestry, and recent policy initiatives by forest certification agencies which aim to increase broadleaf-associated biodiversity within conifer-dominated production forest landscapes.

Felton, Swedish University of Agricultural Sciences, Southern Swedish Forest Research Centre, Alnarp, Sweden E-mail: adam.felton@ess.slu.se
Andersson, Swedish University of Agricultural Sciences, Southern Swedish Forest Research Centre, Alnarp, Sweden E-mail: ea@nn.se
Ventorp, Swedish University of Agricultural Sciences, Southern Swedish Forest Research Centre, Alnarp, Sweden E-mail: dv@nn.se
Lindbladh, Swedish University of Agricultural Sciences, Southern Swedish Forest Research Centre, Alnarp, Sweden E-mail: ml@nn.se

article id 91, category Research note

Raisa Mäkipää, Tapio Linkosalo. (2011). A non-destructive field method for measuring wood density of decaying logs. Silva Fennica vol. 45 no. 5 article id 91. https://doi.org/10.14214/sf.91

Keywords: coarse woody debris; carbon stock; decaying wood; wood decomposition; penetrometer; pilodyn

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Decaying dead wood density measurements are a useful indicator for multiple purposes, such as for estimating the amount of carbon in dead wood and making predictions of potential diversity of dead wood inhabiting fungi and insects. Currently, qualitative decay phases are used as wood density estimates in many applications, since measuring the density is laborious. A quantitative measure of density would, however, be preferred over the qualitative one. Penetrometers, which are commonly used for measuring the density of standing trees, might also be applicable to dead wood density measurements. We tested the device for making quick, quantitative measurements of decaying logs. The penetrometer measures the depth into which a pre-loaded spring forces a pin in the wood. We tested pins of 5 and 10 mm diameter together with an original 2.5 mm pin and compared the results with gravimetric density measurements of the sample logs. Our results suggest that the standard pin works for less decayed wood, but for more decomposed wood, the thicker 5 mm pin gave more reliable estimates when the penetration measures were converted to densities with a linear regression function (R² = 0.62, F = 82.9, p = 0.000). The range of wood densities successfully measured with the 5 mm pin was from 180 to 510 kg m^–3. With the 10 mm pin, the measuring resolution of denser wood was compromised, while the improvement at the other end of density scale was not large. As a conclusion, the penetrometer seems to be a promising tool for quick density testing of decaying logs in field, but it needs to be modified to use a thicker measuring pin than the standard 2.5 mm pin.

Mäkipää, The Finnish Forest Research Institute, Vantaa, Finland E-mail: raisa.makipaa@metla.fi
Linkosalo, The Finnish Forest Research Institute, Vantaa, Finland E-mail: ts@nn.fi

Category : Commentary

article id 445, category Commentary

Bengt Gunnar Jonsson, Jari Kouki, Timo Kuuluvainen. (2011). Northern Primeval Forests – Ecology, Conservation and Management. Silva Fennica vol. 45 no. 5 article id 445. https://doi.org/10.14214/sf.445

View details | Full text in PDF | Author Info

Jonsson, Mid Sweden University, Dept. of Natural Sciences, Sundsvall, Sweden E-mail: bengt-gunnar.jonsson@miun.se
Kouki, University of Eastern Finland, School of Forest Sciences, Joensuu, Finland E-mail: jari.kouki@uef.fi
Kuuluvainen, University of Helsinki, Dept. of Forest Science, Helsinki, Finland E-mail: timo.kuuluvainen@helsinki.fi

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