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Articles by Timo Kuuluvainen

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
  • Jonsson, Mid Sweden University, Dept. of Natural Sciences, Sundsvall, Sweden ORCID ID:E-mail: bengt-gunnar.jonsson@miun.se (email)
  • Kouki, University of Eastern Finland, School of Forest Sciences, Joensuu, Finland ORCID ID:E-mail: jari.kouki@uef.fi
  • Kuuluvainen, University of Helsinki, Dept. of Forest Science, Helsinki, Finland ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi
article id 547, category Commentary
Timo Kuuluvainen. (2002). Introduction. Disturbance dynamics in boreal forests: defining the ecological basis of restoration and management of biodiversity. Silva Fennica vol. 36 no. 1 article id 547. https://doi.org/10.14214/sf.547
  • Kuuluvainen, Department of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi (email)

Category: Research article

article id 1279, category Research article
Andreas Kreutz, Tuomas Aakala, Russell Grenfell, Timo Kuuluvainen. (2015). Spatial tree community structure in three stands across a forest succession gradient in northern boreal Fennoscandia. Silva Fennica vol. 49 no. 2 article id 1279. https://doi.org/10.14214/sf.1279
Highlights: We studied the tree community spatial structure in three 1.2-ha plots representing naturally developed northern boreal forests of varying ages; Spatial structure showed little differences between the mid-successional, late-successional and old-growth stands; The occurrence of Picea abies relative to Betula spp. indicated a mosaic-like spatial assembly; Mosaics are likely maintained by species-specific replacement, not reciprocal replacement as thought earlier.
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.
  • Kreutz, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID ID:E-mail: andreas.kreutz@wald-rpl.de
  • Aakala, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID ID: http://orcid.org/0000-0003-0160-6410 E-mail: tuomas.aakala@helsinki.fi (email)
  • Grenfell, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID ID:E-mail: russell.grenfell@gmail.com
  • Kuuluvainen, Department of Forest Sciences, University of Helsinki, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi
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
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 ORCID ID:E-mail: russell.grenfell@helsinki.fi (email)
  • Aakala, University of Helsinki, Dept of Forest Sciences, Helsinki, Finland ORCID ID:E-mail:
  • Kuuluvainen, University of Helsinki, Dept of Forest Sciences, Helsinki, Finland ORCID ID:E-mail:
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
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 ORCID ID:E-mail: inari.yllasjarvi@ramk.fi (email)
  • Berglund, University of Helsinki, Department of Forest Sciences, Helsinki, Finland ORCID ID:E-mail:
  • Kuuluvainen, University of Helsinki, Department of Forest Sciences, Helsinki, Finland ORCID ID:E-mail:
article id 468, category Research article
Antti Lännenpää, Tuomas Aakala, Heikki Kauhanen, Timo Kuuluvainen. (2008). Tree mortality agents in pristine Norway spruce forests in northern Fennoscandia. Silva Fennica vol. 42 no. 2 article id 468. https://doi.org/10.14214/sf.468
We examined tree mortality agents in pristine old Norway spruce (Picea abies (L.) Karst.) forests in northern Finland and northwestern Russia. The data was collected on nine 40 m   400 m transects. The primary mortality agents of recently dead trees were recorded and their frequencies were calculated. The pattern of tree growth prior to death was studied based on increment core samples and compared with the growth of healthy dominant trees. Of all recently dead trees, 72% could be associated with a primary mortality agent. In both study areas the most common primary mortality agent was a Coniophora (Mérat) DC. -genus fungi, which was found on average in 33% of trees sampled. The fungi Phellinus chrysoloma (Fr.) Don and Onnia leporina (Fr.) H. Jahn as mortality agents were more common in the Finnish area compared to the Russian area. Analysis on the growth patterns indicated weak differences between different pathogens’ influence on prior-to-death growth of trees, so that fungi rotting the whole tree decreased tree growth more rapidly than fungi rotting only the heart wood. The results demonstrated that in old Norway spruce forests of northern Fennoscandia the most common primary tree mortality agents were wood rotting fungi, which weaken the mechanical stability of tree stems until they fall due to snow or wind, which should be considered only as secondary mortality agents. It is evident that tree death in pristine forest typically results from a long-lasting process involving both biotic and abiotic factors.
  • Lännenpää, Department of Forest Ecology, University of Helsinki, Finland ORCID ID:E-mail:
  • Aakala, Department of Forest Ecology, University of Helsinki, Finland ORCID ID:E-mail: tuomas.aakala@helsinki.fi (email)
  • Kauhanen, Kolari Research Unit, Finnish Forest Research Institute, Finland ORCID ID:E-mail:
  • Kuuluvainen, Department of Forest Ecology, University of Helsinki, Finland ORCID ID:E-mail:
article id 250, category Research article
Saara Lilja-Rothsten, Michelle de Chantal, Chris Peterson, Timo Kuuluvainen, Ilkka Vanha-Majamaa, Pasi Puttonen. (2008). Microsites before and after restoration in managed Picea abies stands in southern Finland: effects of fire and partial cutting with dead wood creation. Silva Fennica vol. 42 no. 2 article id 250. https://doi.org/10.14214/sf.250
Different types of microsites, e.g. CWD (coarse woody debris), mounds, and uprooting pits, are important for tree regeneration and biodiversity. However, microsite diversity is greatly reduced in managed stands. We studied how restoration treatments changed microsite distribution in mature managed Picea abies stands. Four cutting treatments were used: uncut, low-CWD (5 m3 ha–1 of down retention trees, DRT, and 50 m3 ha–1 of standing retention trees), intermediate-CWD (as previous but leaving 30 m3 ha–1 of DRT), and high-CWD (as previous but with 60 m3 ha–1 of DRT). Timber harvested from stands ranged from 108–168 m3 ha–1. Half of the stands were burned, and half remained unburned. Sampling was stratified into upland and paludified biotopes within each stand. The pre-treatment microsite distributions were dominated by level ground in both biotopes; mounds and microsites on or next to CWD or a stump were slightly more abundant in the paludified than in the upland biotopes. Microsites were more diverse after cutting, with and without fire. The cutting treatment increased the relative abundances of microsites on or next to CWD. Fire consumed small diameter dead wood and flattened mounds. Microsites were more diverse in paludified than in upland biotopes. The results demonstrate that microsite diversity can rapidly be restored to structurally impoverished managed Picea stands despite a large portion of wood volume being harvested.
  • Lilja-Rothsten, University of Helsinki, Dept. of Forest Ecology, Finland ORCID ID:E-mail: saara.lilja@helsinki.fi (email)
  • Chantal, University of Helsinki, Dept. of Forest Ecology, Finland ORCID ID:E-mail:
  • Peterson, Dept. of Plant Biology, University of Georgia, Athens, GA, USA ORCID ID:E-mail:
  • Kuuluvainen, University of Helsinki, Dept. of Forest Ecology, Finland ORCID ID:E-mail:
  • Vanha-Majamaa, The Finnish Forest Research Institute, Vantaa Unit, Finland ORCID ID:E-mail:
  • Puttonen, The Finnish Forest Research Institute, Vantaa Unit, Finland ORCID ID:E-mail:
article id 377, category Research article
Saara Lilja, Timo Kuuluvainen. (2005). Structure of old Pinus sylvestris dominated forest stands along a geographic and human impact gradient in mid-boreal Fennoscandia. Silva Fennica vol. 39 no. 3 article id 377. https://doi.org/10.14214/sf.377
Stand structural characteristics were examined in old Pinus sylvestris dominated sites in three regions along a broad geographic and human impact gradient in mid-boreal Fennoscandia. The study regions were: 1) Häme in south-western Finland, with a long history of forest utilization, 2) Kuhmo in north-eastern Finland, with a more recent history of intensive forest utilization, and 3) Vienansalo in Russian Karelia, still characterized by a large near-natural forest landscape. Within each region the sampled sites were divided into three human impact classes: 1) near-natural stands, 2) stands selectively logged in the past, and 3) managed stands treated with thinnings. The near-natural and selectively logged stands in Häme and Kuhmo had a significantly higher Picea proportion compared to stands in Vienansalo. In comparison, the proportions of deciduous tree volumes were higher in near-natural stands in Vienansalo compared to near-natural stands in Häme. The pooled tree diameter distributions, both in near-natural and selectively logged stands, were descending whereas managed stands had a bimodal diameter distribution. Structural diversity characteristics such as broken trunks were most common in near-natural stands and in stands selectively logged in the past. The results demonstrate the higher structural complexity of near-natural stands and stands selectively logged in the past compared to managed stands, and highlight that old near-natural stands and stands selectively logged in the past vary widely in their structures. This obviously reflects both their natural variability but also various combinations of pre-industrial land use and human impact on fire disturbance. These factors need to be acknowledged when using “natural” forest structures as a reference in developing strategies for forest management, restoration and nature conservation.
  • Lilja, University of Helsinki, Department of Forest Ecology, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID ID:E-mail: saara.lilja@helsinki.fi (email)
  • Kuuluvainen, University of Helsinki, Department of Forest Ecology, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID ID:E-mail:
article id 414, category Research article
Markku Larjavaara, Timo Kuuluvainen, Heidi Tanskanen, Ari Venäläinen. (2004). Variation in forest fire ignition probability in Finland. Silva Fennica vol. 38 no. 3 article id 414. https://doi.org/10.14214/sf.414
We examined climate-caused spatio-temporal variation of forest fire ignition probability in Finland based on empirical ignition experiments and 37 years of meteorological data from 26 meteorological stations scattered across Finland. First, meteorological data was used in order to estimate the variation in forest fuel moisture content with the model of the Finnish forest fire risk index. Second, based on data from empirical ignition experiments, fuel moisture content was linked with forest fire ignition probability. In southern Finland average forest fire ignition probability typically peaks in late May and early June, whereas in the northern part of the country the peak occurs at the end of June. There was a three-fold difference in the average annual ignition probability between the north-eastern part (3%) and south-western part of the country (9%). The observed differences in fire ignition probability suggest that the characteristics of the natural fire regime also vary considerably in the southern versus the northern part of the country.
  • Larjavaara, University of Helsinki, Dept of Forest Ecology, FI-00014 University of Helsinki, Finland ORCID ID:E-mail: markku.larjavaara@helsinki.fi (email)
  • Kuuluvainen, University of Helsinki, Dept of Forest Ecology, FI-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Tanskanen, University of Helsinki, Dept of Forest Ecology, FI-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Venäläinen, University of Helsinki, Dept of Forest Ecology, FI-00014 University of Helsinki, Finland ORCID ID:E-mail:
article id 557, category Research article
Tuomo Wallenius, Timo Kuuluvainen, Raimo Heikkilä, Tapio Lindholm. (2002). Spatial tree age structure and fire history in two old-growth forests in eastern Fennoscandia. Silva Fennica vol. 36 no. 1 article id 557. https://doi.org/10.14214/sf.557
Two near natural old-growth forests, one dominated by Picea abies and the other by Pinus sylvestris, were studied for their fire history, and spatial patterns of trees and tree ages. The spatial tree age structure and the disturbance history of the forests were examined by drawing age class maps based on mapped and aged trees and by dating fires based on fire scars, and by using spatial analyses at tree scale. The tree age structures of the Picea and Pinus dominated forests were different, mainly due to differences in fire history and sensitivity of the dominant tree species to fire. Fire histories and tree age structures of both sites have probably been affected by human in the ancient past. However, in the Picea dominated site, the fires had been severe, killing most of the trees, whereas in the Pinus dominated site the severity of fires had been more variable, leaving some Pinus and even Picea trees alive. In the Pinus dominated site, the tree age distribution was multimodal, consisting of two Pinus cohorts, which were established after fires and a later Picea regeneration. The Picea dominated site was composed of four patches of different disturbance history. In the oldest patch, the tree age distribution was unimodal, with no distinct cohorts, while a single cohort that regenerated after severe fire disturbances dominated the three other patches. In both sites the overall spatial patterns of living and dead trees were random and the proportion of spatially autocorrelated variance of tree age was low. This means that trees of different age grew more or less mixed in the forest without forming spatially distinct regeneration patches, even in the oldest patch of Picea dominated Liimatanvaara, well over 200 years after a fire. The results show that detail knowledge of disturbance history is essential for understanding the development of tree age structures and their spatial patterns.
  • Wallenius, Department of Ecology and Systematics, University of Helsinki, P.O. Box 65, FIN-00014, Helsinki, Finland ORCID ID:E-mail: tuomo.wallenius@helsinki.fi (email)
  • Kuuluvainen, Department of Forest Ecology, University of Helsinki, P.O. Box 27, FIN-00014, Helsinki, Finland ORCID ID:E-mail:
  • Heikkilä, Research Centre of Friendship Park, Tönölä, FIN-88900 Kuhmo, Finland ORCID ID:E-mail:
  • Lindholm, Finnish Environment Institute, Nature and Land Use Division, P.O. Box 140, FIN-00251 Helsinki, Finland ORCID ID:E-mail:
article id 556, category Research article
Timo Kuuluvainen, Juha Mäki, Leena Karjalainen, Hannu Lehtonen. (2002). Tree age distributions in old-growth forest sites in Vienansalo wilderness, eastern Fennoscandia. Silva Fennica vol. 36 no. 1 article id 556. https://doi.org/10.14214/sf.556
The age and size of trees was sampled and measured on eight sample plots (0.2 ha each) within a Pinus sylvestris -dominated boreal forest landscape in Vienansalo wilderness, Russian Karelia. The fire history of these plots was obtained from a previous dendrochronological study. All the studied sample plots showed a wide and uneven distribution of tree ages, but the shape of the age distributions of trees as well as tree species composition varied substantially. Trees over 250 years of age occurred in every studied plot, despite its small size. This suggests that old Pinus were common and rather evenly distributed in the landscape matrix. The oldest Pinus tree was 525 years of age. The correlations between tree age and size were often weak or even nil. In Pinus the correlation between age and diameter was stronger than that between age and height. In the dominant tree species Pinus and Picea, the largest trees were not the oldest trees. The tree age distributions together with the fire history data indicated that the past fires have not been stand replacing, as many of the older Pinus had survived even several fires. Tree age classes that had regenerated after the last fire were most abundant and dominated by Picea and/or deciduous trees, while the trees established before the last fire were almost exclusively Pinus. The results suggest that periodic occurrence of fire is important for the maintenance of the Pinus-dominated landscape. This is because fire kills most Picea and deciduous trees and at the same time enhances conditions for Pinus regeneration, facilitated by available seed from the continuous presence of old fire-tolerant Pinus trees.
  • Kuuluvainen, Department of Forest Ecology, University of Helsinki, P.O. Box 24, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi (email)
  • Mäki, Department of Forest Ecology, University of Helsinki, P.O. Box 24, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Karjalainen, Department of Forest Ecology, University of Helsinki, P.O. Box 24, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Lehtonen, Faculty of Forestry, University of Joensuu, P.O. Box 24, FIN-80101 Joensuu, Finland ORCID ID:E-mail:
article id 555, category Research article
Leena Karjalainen, Timo Kuuluvainen. (2002). Amount and diversity of coarse woody debris within a boreal forest landscape dominated by Pinus sylvestris in Vienansalo wilderness, eastern Fennoscandia. Silva Fennica vol. 36 no. 1 article id 555. https://doi.org/10.14214/sf.555
The amount, variability, quality and spatial pattern of coarse woody debris (CWD) on mineral soil sites was studied within a natural Pinus sylvestris L. dominated boreal forest landscape in Russian Viena Karelia. Data on the total CWD was collected on 27 sample plots (20 m x 100 m) and data on large CWD was surveyed along four transects (40 m wide and up to 1000 m long). The mean volume of CWD (standing and down combined) was 69.5 m3 ha–1, ranging from 22.2 m3 ha–1 to 158.7 m3 ha–1 from plot to plot. On average, 26.9 m3 ha–1 (39%) of CWD was standing dead wood and 42.7 m3 ha–1 (61%) down dead wood. The CWD displayed a wide range of variation in tree species, tree size, stage of decay, dead tree type and structural characteristics, creating a high diversity of CWD habitats for saproxylic organisms. Large CWD was almost continuously present throughout the landscape and its overall spatial distribution was close to random, although a weak autocorrelation pattern was found at distances less than about 50 m. On small spatial scales total CWD showed wide variation up to a sample area of about 0.1 ha, beyond which the variation stabilized. The fire history variables of the sample plots were not related to the amount of CWD. This and the spatial pattern of CWD suggest that the CWD dynamics in this landscape was not driven by fire, but by more or less random mortality of trees due to autogenic causes of death.
  • Karjalainen, Department of Forest Ecology, P.O. Box 24, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Kuuluvainen, Department of Forest Ecology, P.O. Box 24, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi (email)
article id 554, category Research article
Seppo Rouvinen, Timo Kuuluvainen, Juha Siitonen. (2002). Tree mortality in a Pinus sylvestris dominated boreal forest landscape in Vienansalo wilderness, eastern Fennoscandia. Silva Fennica vol. 36 no. 1 article id 554. https://doi.org/10.14214/sf.554
Tree mortality and its causes and spatial pattern were examined along four transects (width 40 m, length 2550–3960 m), with a total length of 12 190 m and area of 48.8 ha, in a Pinus sylvestris L. dominated, boreal forest landscape. Tree mortality was determined within a time window of 3 years by identifying those trees (dbh ≥ 10 cm) along the transects that fitted into one of the three categories: 1) current mortality: trees that had died during the year of survey (1998), 2) recent mortality: trees that had died during the year (1997) before the survey year, and 3) predicted mortality: trees that were expected to die during the year (1999) following the survey year. Long-term tree mortality was studied on 10 plots (20 m x 100 m) by dating 87 dead trees using dendrochronological methods. The mean current mortality was 1.4 m3 ha–1 (3.7 trees ha–1). Both the recent and predicted mortalities were also 1.4 m3 ha–1. Mortality was, on the average, higher on peatlands than on mineral soils. The highest mortality was found within an area recently flooded by beavers. Over half of the examined trees (52%) were judged to die without any visible signs of an external abiotic cause. At the landscape scale, tree mortality was continuous although somewhat aggregated in space. Of the 66 dated standing dead Pinus trees, 23 (35%) had died during the 19th century and two during the 18th century, demonstrating that dead Pinus can remain standing for long periods of time before falling. Our results show that autogenic mortality of individual trees or small groups of trees was the predominant mode of disturbance in this Pinus dominated landscape.
  • Rouvinen, University of Joensuu, Faculty of Forestry, P.O. Box 111, FIN-80101 Joensuu, Finland ORCID ID:E-mail: seppo.rouvinen@forest.joensuu.fi (email)
  • Kuuluvainen, University of Helsinki, Department of Forest Ecology, P.O. Box 27, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail:
  • Siitonen, Finnish Forest Research Institute, P.O. Box 18, FIN-01301 Vantaa, Finland ORCID ID:E-mail: juha.siitonen@metla.fi

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
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; ORCID ID:E-mail: guntis.brumelis@lu.lv (email)
  • Jonsson, Department of Natural Sciences, Engineering and Mathematics, Mid Sweden University, Sundsvall, Sweden ORCID ID:E-mail:
  • Kouki, School of Forest Sciences, University of Eastern Finland, Joensuu, Joensuu ORCID ID:E-mail:
  • Kuuluvainen, Department of Forest Sciences, University of Helsinki, Finland ORCID ID:E-mail:
  • Shorohova, Saint-Petersburg State Forest Academy, Saint-Petersburg, Russia & Finnish Forest Research Institute, Vantaa Research Unit, Vantaa, Finland ORCID ID:E-mail:
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
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 ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi (email)
  • Aakala, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland ORCID ID:E-mail:
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
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) ORCID ID:E-mail: shorohova@ES13334.spb.edu (email)
  • Kneeshaw, Université du Québec à Montréal, Centre d’étude de la forêt, Montreal, Canada ORCID ID:E-mail:
  • Kuuluvainen, University of Helsinki, Department of Forest Sciences, Helsinki, Finland ORCID ID:E-mail:
  • Gauthier, Natural Resources Canada, Canadian Forest Service, Laurentian Forestry Centre, Québec, Canada ORCID ID:E-mail:
article id 552, category Review article
Timo Kuuluvainen. (2002). Natural variability of forests as a reference for restoring and managing biological diversity in boreal Fennoscandia. Silva Fennica vol. 36 no. 1 article id 552. https://doi.org/10.14214/sf.552
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 difficulties 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, diversification of silvicultural and harvesting treatments, as guided by natural disturbance dynamics, is needed to produce more variation in disturbance severity, quality, extent, and repeatability.
  • Kuuluvainen, Department of Forest Ecology, P.O. Box 24, FIN-00014 University of Helsinki, Finland ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi (email)

Category: Discussion article

article id 572, category Discussion article
Timo Kuuluvainen, Kaisu Aapala, Petri Ahlroth, Mikko Kuusinen, Tapio Lindholm, Tapani Sallantaus, Juha Siitonen, Harri Tukia. (2002). Principles of ecological restoration of boreal forested ecosystems: Finland as an example. Silva Fennica vol. 36 no. 1 article id 572. https://doi.org/10.14214/sf.572
  • Kuuluvainen, Department of Forest Ecology, University of Helsinki, P.O.Box 27 FIN-00014, Finland ORCID ID:E-mail: timo.kuuluvainen@helsinki.fi (email)
  • Aapala, Finnish Environment Institute, Expert Services Department, Nature Division, P.O. Box 140, FIN-00251 Helsinki ORCID ID:E-mail:
  • Ahlroth, University Museum, Section of Natural History, P.O. Box 35, FIN-40351, Jyväskylä, Finland ORCID ID:E-mail:
  • Kuusinen, Ministry of the Environment, Land Use Department, P.O.Box 380, FIN-00131 Helsinki, Finland ORCID ID:E-mail:
  • Lindholm, Finnish Environment Institute, Expert Services Department, Nature Division, P.O. Box 140, FIN-00251 Helsinki ORCID ID:E-mail:
  • Sallantaus, Pirkanmaa Regional Environment Centre, P.O. Box 297, FIN-33101 Tampere, Finland ORCID ID:E-mail:
  • Siitonen, Finnish Forest Research Institute, Vantaa Research Centre, P.O. Box 18, FIN-01301 Vantaa, Finland ORCID ID:E-mail: juha.siitonen@metla.fi
  • Tukia, Finnish Environment Institute, Expert Services Department, Nature Division, P.O. Box 140, FIN-00251 Helsinki ORCID ID:E-mail:

Category: Article

article id 5598, category Article
Timo Kuuluvainen, Kari Leinonen, Markku Nygren, Antti Penttinen. (1996). Statistical opportunities for comparing stand structural heterogeneity in managed and primeval forests: an example from boreal spruce forest in southern Finland. Silva Fennica vol. 30 no. 2–3 article id 5598. https://doi.org/10.14214/sf.a9243

The horizontal and vertical stand structure of living trees was examined in a managed and in a primeval Norway spruce-dominated forest in Southern Finland. Tree size distributions (DBHs, tree height) were compared using frequency histograms. The vertical distribution of tree heights was illustrated as tree height plots and quantified as the tree height diversity (THD) using the Shannon-Weaver formula. The horizontal spatial pattern of trees was described with stem maps and quantified with Ripley's K-function. The spatial autocorrelation of tree sizes was examined with semivariogram analysis. In the managed forest the DBH and height distributions of trees were bimodal, indicating a two-layered vertical structure with a single dominant tree layer and abundant regeneration in the understory. The primeval forest had a much higher total number of trees which were rather evenly distributed in different diameter and tree height classes. The K-function summaries for trees taller than 15 m indicated that the primeval stand was close to complete random pattern. The managed stand was regular at small distances (up to 4 m). The semivariograms of tree sizes (DBH tree height) showed that the managed forest had a clear spatial dependence in tree sizes up to inter-tree distances of about 12 meters. In contrast, the primeval spruce forest had a variance peak at very short inter-tree distances (< 1 m) and only weak spatial autocorrelation at short inter-tree distances (1–5 m). Excluding the understory trees (h < 15 m) from the analysis drastically changed the spatial structure of the forest as revealed by semivariograms. ln general, the structure of the primeval forest was both horizontally and vertically more variable and heterogeneous compared to the managed forest. The applicability of the used methods in describing fine-scale forest structure i discussed.

  • Kuuluvainen, ORCID ID:E-mail:
  • Leinonen, ORCID ID:E-mail:
  • Nygren, ORCID ID:E-mail:
  • Penttinen, ORCID ID:E-mail:
article id 5556, category Article
Timo J. Hokkanen, Erkki Järvinen, Timo Kuuluvainen. (1995). Properties of top soil and the relationship between soil and trees in a boreal Scots pine stand. Silva Fennica vol. 29 no. 3 article id 5556. https://doi.org/10.14214/sf.a9207

One-hectare plot in a Scots pine (Pinus sylvestris L.) forest was systemically sampled for surface soil characteristics: humus layer thickness, soil carbon and nitrogen content, pH, electrical conductivity and respiration were determined from 106 samples. The effects of large trees on the plot were mapped and their joint influences at the locations of soil sampling were described as the influence potential, derived from the ecological field theory, and were calculated based on the locations and dimensions of trees.

The range of variation of soil characteristics was from three to sevenfold; no spatial autocorrelation was detected. The calculated influence potential of trees, as determined by their size and spatial distribution, was related to the spatial variation of top soil properties. Top soil properties were also related to thickness of the humus layer but they were poorly correlated with underlying mineral soil characteristics. Humus layer thickness, with the calculated influence potential of trees, may provide a means to predict top soil characteristics in specific microenvironments in the forest floor.

  • Hokkanen, ORCID ID:E-mail:
  • Järvinen, ORCID ID:E-mail:
  • Kuuluvainen, ORCID ID:E-mail:
article id 5385, category Article
Timo Kuuluvainen. (1989). Branching dynamics in young Scots pine. Silva Fennica vol. 23 no. 3 article id 5385. https://doi.org/10.14214/sf.a15542

The development of shoot number and shoot properties was examined in successive shoot cohorts of young widely-spaced Scots pine trees (Pinus sylvestris L.) growing in a progeny trial. This was accomplished by reconstructing the branching process of the trees over a period of five years, from tree age 4 to 8. During this time the number of shoots in successive shoot cohorts increased rapidly, while at the same time the mean shoot length decreased. The decrease in shoot lengths from older to younger shoots was accompanied by a decline in the bifuraction frequency of the shoots. In general, rapid changes occurred in the branching characteristics during the yearly development of the trees. The variation in the branching characteristics was reflected in the development of the architecture and biomass production of the trees.

The PDF includes an abstract in Finnish.

  • Kuuluvainen, ORCID ID:E-mail:
article id 5379, category Article
Timo Kuuluvainen, Timo Pukkala. (1989). Effect of Scots pine seed trees on the density of ground vegetation and tree seedlings. Silva Fennica vol. 23 no. 2 article id 5379. https://doi.org/10.14214/sf.a15536

The study uses the methodology of ecological field theory to model the effect of Scots pine (Pinus sylvestris L.) seed trees on the density of tree seedlings and other plants in the field layer. The seed trees had a clear effect on the expected value of the amount and distribution of the ground vegetation. The vicinity of seed trees had an adverse effect on the growth of grasses, herbs and seedlings, while mosses were most abundant near the trees. Models based on the ecological field approach were derived to describe the effect of seed trees on the ground vegetation.

The PDF includes an abstract in Finnish.

  • Kuuluvainen, ORCID ID:E-mail:
  • Pukkala, ORCID ID:E-mail:
article id 5347, category Article
Timo Kuuluvainen, Markku Kanninen, Juha-Pekka Salmi. (1988). Tree architecture in young Scots pine: properties, spatial distribution and relationships of components of tree architecture. Silva Fennica vol. 22 no. 2 article id 5347. https://doi.org/10.14214/sf.a15504

The architecture of Scots pine (Pinus sylvestris L.) was studied in an eight-year-old progeny test. The measurements included characteristics of crown structure, spatial distribution of shoots and yield components. The spatial distribution of shoots showed striking between-tree differences, and two extreme distribution patterns were detected. One represented a non-layered structure with a vertically relative even shoot distribution, and the other a layered structure with a vertically highly uneven shoot distribution.

Close correlations existed between several components of tree architecture and it is suggested that changes in the phenotypic architecture in Scots pine follow an epigenetic pattern, which enables the prediction of adaptational changes in structural components. The structural characteristics related to high above-ground biomass were a long crown, high total shoot length, high number of branches per whorl and big shoots of low needle density occupying a big share of the crown volume.

The PDF includes a summary in Finnish.

  • Kuuluvainen, ORCID ID:E-mail:
  • Kanninen, ORCID ID:E-mail:
  • Salmi, ORCID ID:E-mail:
article id 5316, category Article
Timo Pukkala, Timo Kuuluvainen. (1987). Effect of canopy structure on the diurnal interception of direct solar radiation and photosynthesis in a tree stand. Silva Fennica vol. 21 no. 3 article id 5316. https://doi.org/10.14214/sf.a15473

The utilization of direct radiation was studied in five model stands of Poisson-type tree distribution and cone-shaped crowns. The radiation extinction depended on the self-shading of the crown and the shading caused by other trees. The results indicate that at low sun elevation a stand populated by very narrow-crowned trees is most effective in light interception and photosynthesis. At high sun elevation a broad-crowned canopy is best illuminated and most favourable for photosynthesis. A stand with a two-storey canopy is effective in all latitudes when the crowns are moderately narrow. In two-storey canopies the foliage of the lower storey can be better illuminated than in the lower parts of the upper storey, because of the smaller self-shading in the small crowns of the lower storey. A canopy where the crown volume is concentrated on few big crowns is less effective than a canopy consisting of many small crowns.

The PDF includes an abstract in Finnish.

  • Pukkala, ORCID ID:E-mail:
  • Kuuluvainen, ORCID ID:E-mail:

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