Articles by Ari Nikula

The European Pine Sawfly (Neodiprion sertifer Geoffroy) is one of the most serious defoliators of Scots pine (Pinus sylvestris L.) in northern Europe. We studied the pattern in the regional occurrence of the outbreaks of N. sertifer in Finland in years 1961-90, and made predictions about the outbreak pattern to the year 2050 after predicted winter warming. We tested whether minimum winter temperatures and forest type and soil properties could explain the observed outbreak pattern. We analysed outbreak patterns at two different spatial levels: forest board- and municipal-level.

The proportion of coniferous forests on damage-susceptible soils (dry and infertile sites) explained a significant part of the variation in outbreak frequency at small spatial scale (municipalities) but not at large spatial scale (forest boards). At the forest board level, the incidence of minimum temperatures below -36 °C (= the critical value for egg mortality) explains 33% of the variation in the outbreak pattern, and at the municipal level the incidence of cold winters was also the most significant explaining variable in northern Finland. Egg mortality due to cold winters seems to be the most parsimonious factor explaining why there have been so few N. sertifer outbreaks in northern and north-eastern Finland. We predict that climate change (increased winter temperatures) may increase the frequency of outbreaks in eastern and northern Finland in the future.

• Virtanen, E-mail: tv@mm.unknown
• Neuvonen, E-mail: sn@mm.unknown
• Niemelä, E-mail: pn@mm.unknown
• Nikula, E-mail: an@mm.unknown
• Varama, E-mail: mv@mm.unknown

The moose (Alces alces L.), a common large herbivore in the boreal region, impairs forest regeneration by browsing on tree seedlings and saplings. Moose prefer deciduous species, but during winter more coniferous seedlings are used. We used meta-analyses, separately for deciduous and coniferous seedlings, for evaluating whether excluding moose browsing affected seedling density and height. In addition, we compared (1) deciduous seedling proportion, (2) stand density, (3) elapsed time from fencing and (4) estimated moose density with moose exclusion effect sizes. Fencing had a positive effect on coniferous seedling height. With more deciduous trees in a seedling stand, the fencing effect for both seedling height and density of coniferous seedlings decreased. On the other hand, the fencing effects increased with denser stands. At some point effect sizes turned to negative, and conifer species varied in their response to browsing. This implies that deciduous seedlings can protect conifers from browsing by moose up to some mixing ratio, but when deciduous seedling densities are too high, their negative effect increases, presumably through increased competition. Our results suggest that a moderate deciduous admixture in conifer-dominated mixed seedling stands can decrease moose damage but also underline the significance of timely silvicultural measures to minimize the negative effects of excessive deciduous seedlings and too dense stands. Due to differences in coniferous and deciduous species, as well as their compositions and amounts in studied experiments, more studies adjusted to local conditions are still needed to give exact measures for silvicultural recommendations.

• Domisch, Natural Resources Institute Finland (Luke), Natural Resources, Yliopistokatu 6B, FI-80100 Joensuu, Finland https://orcid.org/0000-0001-7026-1087 E-mail: timo.domisch@luke.fi
• Huuskonen, Natural Resources Institute Finland (Luke), Natural Resources, Latokartanonkaari 9, FI-00790 Helsinki, Finland E-mail: saija.huuskonen@luke.fi
• Matala, Natural Resources Institute Finland (Luke), Natural Resources, Yliopistokatu 6B, FI-80100 Joensuu, Finland https://orcid.org/0000-0002-5867-5057 E-mail: juho.matala@luke.fi
• Nikula, Natural Resources Institute Finland (Luke), Natural Resources, Ounasjoentie 6, FI-96200 Rovaniemi, Finland https://orcid.org/0000-0001-8372-8440 E-mail: ari.nikula@luke.fi

An important modifier of forests and forestry practices is browsing by cervids. As high populations of moose (Alces alces L.) cause extensive forest damage in the Fennoscandian boreal forests, models should be able to predict the susceptibility of projected forest structures to browse damage. We augmented the European Forestry Dynamics Model (EFDM) for the area of seedling stands damaged by moose. The augmented model was tested in projecting both forest resources and moose damage for 18 million hectares of forest land in Finland, based on input data from the National Forest Inventory (NFI). Modeling the area of seedling stands damaged as a function of moose population density, forest characteristics, and region-specific interactions of these variables was found to work realistically for 30 years, predicting that the area of seedling stands damaged by moose would increase by up to a third from the last NFI observation. Our work lays the groundwork for modeling consequential, large-scale ecological and socio-economic effects of moose browsing.

• Vauhkonen, University of Eastern Finland, School of Forest Sciences, Yliopistokatu 7, FI-80101 Joensuu, Finland; University of Helsinki, Department of Forest Sciences, Latokartanonkaari 7, FI-00014 Helsingin yliopisto, Finland E-mail: jari.vauhkonen@uef.fi
• Matala, Natural Resources Institute Finland (Luke), Natural resources, Yliopistokatu 6 B, FI-80100 Joensuu, Finland E-mail: juho.matala@luke.fi
• Nikula, Natural Resources Institute Finland (Luke), Natural resources, Ounasjoentie 6, FI-96200 Rovaniemi, Finland https://orcid.org/0000-0001-8372-8440 E-mail: ari.nikula@luke.fi

We modelled the effect of habitat composition and roads on the number and occurrence of moose (Alces alces L.) damage in Ostrobothnia and Lapland using a zero-inflated count model. Models were developed for 1 km², 25 km² and 100 km² landscapes consisting of equilateral rectangular grid cells. Count models predict the number of damage, i.e. the number of plantations and zero models the probability of a landscape being without damage for a given habitat composition. The number of moose damage in neighboring grid cells was a significant predictor in all models. The proportion of mature forest was the most frequent significant variable, and an increasing admixture of mature forests among plantations increased the number and occurrence of damage. The amount of all types of plantations was the second most common significant variable predicting increasing damage along with increasing amount of plantations. An increase in thinning forests as an admixture also increased damage in 1 km² landscapes in both areas, whereas an increase in pine-dominated thinning forests in Lapland reduced the number of damage in 25 km² landscapes. An increasing amount of inhabited areas in Ostrobothnia and the length of connecting roads in Lapland reduced the number of damage in 1 and 25 km² landscapes. Differences in model variables between areas suggest that models of moose damage risk should be adjusted according to characteristics that are specific to the study area.

• Nikula, Natural Resources Institute Finland (Luke), Bioeconomy and Environment, Ounasjoentie 6, FI-96200 Rovaniemi, Finland E-mail: ari.nikula@luke.fi
• Nivala, Natural Resources Institute Finland (Luke), Bioeconomy and Environment, Ounasjoentie 6, FI-96200 Rovaniemi, Finland E-mail: vesa.nivala@luke.fi
• Matala, Natural Resources Institute Finland (Luke), Natural resources, Yliopistokatu 6, FI-80100 Joensuu, Finland E-mail: juho.matala@luke.fi
• Heliövaara, University of Helsinki, Department of Forest Sciences, P.O. Box 27, FI-00014 University of Helsinki, Finland E-mail: kari.heliovaara@helsinki.fi

There is evidence that moose are attracted to fertile growth habitats apparently due to better quality and larger quantities of food. The nutrients in mineral soils originate from the weathering of bedrock and the composition of parental bedrock affects the fertility of produced mineral soil, thus affecting also the import of nutrients into the whole food web. We surveyed the connection between moose damage in forest plantations and the composition of bedrock and surficial deposits in Finnish Lapland. We used a database of compensated moose damage in private forests in years 1997−2010. Undamaged stands in National Forest Inventories (NFI) from years 1986–2008 served as a control data and moose-damaged NFI-stands as a reference data. Bedrock and surficial depositions and the location of studied stands in relation to ancient shorelines were explored by using the digital databases of the Geological Survey of Finland. Moose-damaged stands were concentrated in southwestern and east Lapland in the areas of the Peräpohja Schist Belt and Lapland’s Greenstone Belt that are both composed of nutrient-rich rocks. The bedrock of damaged stands contained a higher proportion of mafic and alkaline rocks than did the control stands. Moose-damaged stands were pine-dominated and grew in more fertile forest sites than did control stands. Part of pine stands probably located in soils formerly occupied by spruce, which may increase the stands’ vulnerability to biotic threats. Especially, there were relatively more moose damage in pine plantations regenerated on fine-grained mineral soils derived from nutrient rich rocks than in less fertile soils.

• Ruuhola, Natural Resources Institute Finland (Luke), Natural Resources and Bioproduction, Yliopistokatu 6, FI-80101 Joensuu, Finland; University of Eastern Finland, Faculty of Science and Forestry, P.O. Box 111, FI-80101 Joensuu, Finland E-mail: teija.ruuhola@uef.fi
• Nikula, Natural Resources Institute Finland (Luke), Economics and Society, Eteläranta 55, FI-96300 Rovaniemi, Finland E-mail: ari.nikula@luke.fi
• Vesa, Natural Resources Institute Finland (Luke), Economics and Society, Eteläranta 55, FI-96300 Rovaniemi, Finland E-mail: vesa.nivala@luke.fi
• Nevalainen, Natural Resources Institute Finland (Luke), Natural Resources and Bioproduction, Yliopistokatu 6, FI-80101 Joensuu, Finland E-mail: seppo.nevalainen@luke.fi
• Matala, Natural Resources Institute Finland (Luke), Natural Resources and Bioproduction, Yliopistokatu 6, FI-80101 Joensuu, Finland E-mail: juho.matala@luke.fi

The occurrence of moose damage was studied using data from three National Forest Inventories (NFIs) accomplished between 1986 and 2008 in Finland. The combined data included a total of 97 390 young stands. The proportion of moose damage increased from 3.6% to 8.6% between the 8th NFI (1986–1994) and the 10th NFI (2004–2008). The majority (75%) of the damage occurred in Scots pine-dominated stands. The proportion of damage was higher in aspen-dominated stands than in stands dominated by any other tree species. The tree species mixture also had a clear effect on the occurrence of damage. Pure Scots pine stands had less damage than mixed Scots pine stands, and moose damage decreased linearly with the increasing proportion of Scots pine. Stands on mineral soil had more frequent moose damage than stands on peatlands. The fertility class of the site had no straightforward effect on the damage frequency. Artificially regenerated stands had more damage than naturally regenerated stands. Accomplished soil preparation measures and the need for thinning or clearing operations increased moose damage. High proportions of moose damage in young stands were found around the country. In the 10th NFI, the largest concentration of damage was found in southwestern Finland. Our study shows the temporal and spatial changes in the occurrence of moose damage and pinpoints some important silvicultural factors affecting the relative risk of young stands over a large geographical area.

• Nevalainen, Natural Resources Institute Finland (Luke), Management and Production of Renewable Resources, P.O. Box 68, FI-80101 Joensuu, Finland E-mail: seppo.nevalainen@luke.fi
• Matala, Natural Resources Institute Finland (Luke), Management and Production of Renewable Resources, P.O. Box 68, FI-80101 Joensuu, Finland E-mail: juho.matala@luke.fi
• Korhonen, Natural Resources Institute Finland (Luke), Economics and society, P.O. Box 68, FI-80101 Joensuu, Finland E-mail: kari.t.korhonen@luke.fi
• Ihalainen, Natural Resources Institute Finland (Luke), Economics and society, P.O. Box 18, FI-01301 Vantaa, Finland E-mail: antti.ihalainen@luke.fi
• Nikula, Natural Resources Institute Finland (Luke), Economics and society, P.O. Box 16, FI-96301 Rovaniemi, Finland E-mail: ari.nikula@luke.fi

• Miettinen, Finnish Game and Fisheries Research Institute, Finland E-mail: janne.miettinen@rktl.fi
• Helle, Finnish Game and Fisheries Research Institute, Finland E-mail: ph@nn.fi
• Nikula, Finnish Forest Research Institute, Finland E-mail: an@nn.fi
• Niemelä, University of Turku, Dept of Biology E-mail: pn@nn.fi

• Miettinen, Kankurinhaka 14, FI-90450 Kempele, Finland E-mail: janne.miettinen@rktl.fi
• Helle, Finnish Game and Fisheries Research Institute, Tutkijantie 2 E, FI-90570 Oulu, Finland E-mail: ph@nn.fi
• Nikula, Finnish Forest Research Institute, Rovaniemi Research Unit, Eteläranta 55, FI-99600 Rovaniemi, Finland E-mail: an@nn.fi
• Niemelä, University of Turku, Department of Biology, FI-20014 University of Turku, Finland E-mail: pn@nn.fi

• Nikula, Finnish Forest Research Institute, Rovaniemi Research Unit, Eteläranta 55, FI-96300 Rovaniemi, Finland E-mail: ari.nikula@metla.fi
• Hallikainen, Finnish Forest Research Institute, Rovaniemi Research Unit, Eteläranta 55, FI-96300 Rovaniemi, Finland E-mail: vh@nn.fi
• Jalkanen, Finnish Forest Research Institute, Rovaniemi Research Unit, Eteläranta 55, FI-96300 Rovaniemi, Finland E-mail: rj@nn.fi
• Hyppönen, Finnish Forest Research Institute, Rovaniemi Research Unit, Eteläranta 55, FI-96300 Rovaniemi, Finland E-mail: mh@nn.fi
• Mäkitalo, Finnish Forest Research Institute, Rovaniemi Research Unit, Eteläranta 55, FI-96300 Rovaniemi, Finland E-mail: km@nn.fi