Moose (Alces alces L.) browsing was studied in young Scots pine (Pinus sylvestris L.) stands mixed with deciduous trees in high-density winter ranges. The proportional use of twig biomass decreased as the availability increased. The total as well as proportional biomass consumption were higher on the moist than on the dry type of forest. The per tree consumption of pine was higher on the moist type, where the availability of pine was lower. Deciduous trees were more consumed on the moist type, where their availability was relatively high. The consumption of pine saplings increased as the availability of birch increased. Pine stem breakages were most numerous when birch occurred as overgrowth above pine and at high birch densities. The availability of other deciduous tree species did not correlate with browsing intensity of Scots pine. Moose browsing had seriously inhibited the development of Scots pines in 6% of the stands, over 60% of available biomass having been removed. Rowan and aspen were commonly over-browsed and their height growth was inhibited, which occurred rarely by birch. There was no difference in the proportion of young stands in forest areas with high and low moose density. A high proportion of peatland forests was found to indicate relatively good feeding habitats in the high-density areas.
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The efficacy of pine tar as a moose (Alces alces L.) contact repellent was tested in young Scots pine (Pinus sylvestris L.) stands suffering from moose damage in Southern Finland during the winter 1981–82. Application of tar to shoots by spraying protected the trees satisfactorily throughout the winter.
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The main stem of young Scots pine (Pinus sylvestris L.) trees was cut off halfway along the current leading shoot and the two previous years’ leading shoots to simulate moose (Alces alces) damage. Trees of the same size were chosen as controls before treatments. The experiment was inspected ten years after artificial stem breakage. Removing the current leading shoot and the second shoot did not essentially affect the height and diameter growth of the trees. Removal down to the third shoot reduced the height as well as diameter growth. The average loss in growth was equivalent to less than one year’s growth. When the stem was cut off at the second or third shoot, stem crookedness and the presence of knots resulted in stem defects that will subsequently reduce the sawtimber quality. A high proportion of the stem defects will obviously still be visible at the first thinning cutting. Removing injured trees as pulpwood and pruning the remaining parts of cut stems evidently improves the quality of pine stand with moose damage.
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The light reflected from the crowns of Scots pine (Pinus sylvestris L.) clones was measured spectroradiometrically during and after growing season. Standard deviations of the spectra of pine clones showing differences in moose browsing intensity were compared. A new algorithm was developed for predicting the browsing intensity of moose (Alces alces).
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The effects of plantation characteristics on moose (Alces alces) browsing intensity was studied in 82 Scots pine (Pinus sylvestris L.) plantations in Southern Finland. Moose browsing occurred most commonly in plantations established on relatively fertile soil, and the degree of damage was at highest in plantations with openings. A high amount of brush, especially aspen, increased the risk of damage. Furthermore, damage was intensified in plantations situated on hills, slopes or at a long distance from main roads or settlements.
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The size, nutrient contents and terpene composition of needles of Scots pine (Pinus sylvestris L.) saplings untouched and repeatedly browsed by the moose (Alces alces L.) were compared. Material was collected from a 14-years old and 2.5 m high pine stand in Bromarv, Southern Finland. The average length and fresh and dry weight of the needles were measured, and nutrient content (N, P, K, Ca, Mg, B, Cu) was determined.
The needles of repeatedly browsed pines became long and robust. There was, however, no difference between the dry matter percentage between the needles. The average nitrogen content was higher in the rebrowsed trees. Nitrogen content is, however, not directly correlated with the palatability of pine needles. Even phosphorus and boron content were higher in the damaged trees. No difference was found in Ca, K, Mg and Cu contents of the browsed and control pine saplings.
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An inventory of Scots pine (Pinus sylvestris L.) graft collection in Kolari (67°15’ N, 23°45’ S) showed that severe damage by arctic hare (Lepus timidus L.), root and bank vole (Microtus oeconomus Pallas and M. agrestis L.) and moose (Alces alces L.) was done to grafts in size and in rather poor condition. Furthermore, the damage by arctic hare was dependent on the dry matter content of the needles. Another inventory in a fertilization experiment in a pine pole-stage forest showed that nitrogen fertilization increased the damage by arctic hare. On the basis of the present results, an assumption was made that the formation of repellent substances against herbivorous mammals is connected with wintering process of northern pines.
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Monoterpene hydrocarbon contents of needles in Scots pine (Pinus sylvestris L.) plants both damaged and untouched by the moose (Alces alces L.) were compared in the study. The material was collected in an 8-year-old plantation in Central Finland. Needle samples were taken from the topmost shoot whorl of the plants in the middle of April, 1976. Only minor differences were found between the plant groups. Thus, terpenes in pine presumably play no important role in the browsing preference by moose.
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The winter food of moose (Alces alces L.) was examined in 1967-68 in the Saariselkä fell area in the communes of Inari and Sodankylä, in the northern parts of the communes of Salla and Savukoski, and in the central part of the commune of Salla in eastern Lapland in Finland.
In northern parts of Salla and Savukoski 25 moose were followed during 3.-13.4.1968. This area is typical wintering terrain of moose in north-east Lapland. According to the estimate, 45% food taken by the moose was Scots pine shoots and needles, 28% birch, 17% juniper sprigs and needles, 9% willow, and 1.5% bear moss. According to observations of the researchers in 26.1.-16.5.1968, moose seemed to avoid birch, even if it was available in the area, and eat Scots pine shoots and needles and juniper.
Moose seemed to prefer willow in as a winter feed in the southern part of the area studied, where it accounts according to the present and earlier studies 50-90% of the winter food. In the northern wintering areas of moose, where willow is not as common, willow seemed to account for less than 10% of the winter food. There Scots pine is the most important winter food for moose.
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The article reviews the occurrence of damage causes by elk (Alces alces L.) in young Scots pine (Pinus sylvestris L.) stands established by direct seeding in the Ostrobothia region in Finland. The data was collected by random sampling, and consists of 110 sample plots in pine stands, established in 1930-1944.
Signs of elk damages could be observed in 20% of the stands. In more than half of the damaged stands pine seedlings were damaged by elk, on the rest of the stands the damage was targeted to hardwood saplings only. With the present density of elk population, the damage has an insignificant bearing upon the development of pine seedling stands in Ostrobothnia. The weaknesses of silvicultural state of the stands have been caused by other factors than elk.
Silviculturally weak stands were more liable to elk damage than strong ones. The occurrence of elk damage was more usual in stands with hardwood mixture than in pure pine stands. Especially goat willow, mountain ash and aspen, but also to some decree birch, seem to attract elk. Those factors that promote hardwood growth: fertility of the site, swampiness and the presence of seeding hardwoods in the area, increase the stand’s liability to elk damage.
The article includes an abstract in English and a summary in Swedish.
Increase in the elk (Alces alces L.) population and the problems of its grazing has called for detailed research. The present study concentrated on three observation areas representing northern, western and eastern parts of Finland. There were 28 field observers watching 68 elks.
Earlier investigations in Finland indicate that aspen (Populus tremula L.) is the staple diet of elk. This study reached different conclusions, probably largely because of aspen is gradually becoming an increasingly rare tree species in Finland. According to this study, the principal food of elk in the winter is willow (Salix sp.). In the whole country, willow accounts for about 70% of elk’s nutrition. In the Far North the percentage is approx. 90. Of the other tree species, the order of preference is: aspen, Scots pine, mountain ash, juniper and birch. In addition, in Western Finland where snow is less deep, lingonberry and blueberry shrubs are on the menu. Beard moss on the spruce was frequently eaten locally. Elk seems to have eaten mainly the last annual shoot of trees and bushes. In few cases it has gnawed the bark of Scots pine, aspen and willow. Elk consumes in average 340 twigs or terminal shoots per day in the winter. This corresponds to about 1.8 kg of food.
The problem of elks damaging Scots pine seedlings has been observed in Western Finland, were the elk population is higher. The article suggests that suitable feeding places would be left for elk in places that are unsuitable for agriculture or forestry. Leaving, for instance, birch seedlings in Scots pine stands has been noticed to attract elks and to increase the damage to pine.
The article includes an abstract in English and a summary in Swedish.
The study was carried out at Padasjoki, Southern Finland, where moose (Alces alces L.) density on the winter range had been over 1.5 individuals/km2. Moose browsing intensity, expressed in terms of number of twigs eaten and biomass used, increased with stand density (biomass). Total biomass consumption (dry weight) per sample plot and per sapling. The number of bites increased, but the percentage biomass removed did not differ when stand density increased. A relatively large bite size was observed on the plots of low stand density. The quantity of food, which on average was of relatively low quality, was obviously important due to the benefit gained through reducing the search time.
The nutritive value of the browse, expressed in terms of chemical compounds indicating low food digestibility, was lower in the dense than in the sparse Scots pine stand. However, the amount of crude protein and arginine were relatively high in the dense stand. We concluded that shading affected the nutritional status of saplings on high density plots.
Although the biomass removed by moose per sapling was high for low density plots, the remaining biomass was larger than that on the high-density plots owing to the relatively large twig biomass of saplings. The number of saplings per hectare without main stem breakage increased significantly as stand density increased.
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The utilization of available food resources by the moose (Alces alces L.) was studied in a Scots pine (Pinus sylvestris L.) plantation containing an admixture of deciduous species. Rowan (Sorbus aucuparia L.) and aspen (Populus tremula L.) were highly utilized compared to pine and both silver birch (Betula pendula Roth) and downy birch (B. pubescens Ehrh.). However, they were not capable of withstanding continuous browsing by moose owing to their diminished biomass. In total, the browsing intensity (number of browsed twigs/tree) on pine and birch was about double of that on rowan and aspen.
The number of browsed twigs per tree increased as the amount of available main branches increased. The number of bites per available branch, as well as the maximum diameter of the bites, decreased as the density of the plantation increased. Silver birch was more used by moose than pubescent birch as well as planted silver birch compared with naturally regenerated trees.
Main stem breakage was especially common in winter 1988, the average height of the pine and birch trees being over two meters. The tops of broken stems were commonly utilized as food. The increase in moose density and the relatively deep snow cover evidently promoted the incidence of serious damage. The number of undamaged trees/ha was greater in dense than in sparse parts of the stand.
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The establishment of moose (Alces alces L.) winter feeding sites, their utilization and their effect on damage to young Scots pine (Pinus sylvestris L.) plantations was studied in Ruokolahti-Imatra area in Eastern Finland in 1987–89. During the period, the density in the area was about 3–5 moose/ 1,000 ha.
Six feeding sites were established by fertilization, offering mineral lics and the tops of aspen and Scot pine and by salting the tops of pine. The moose preferred the feeding site to control areas during both summer and winter. In winter browsing was very heavy, especially in those areas located in or close to traditional wintering areas. In winter no moose were seen in the summer habitats.
The extent of, and fluctuations in moose damage were studied in 47 Scots pine plantations in the immediate surroundings of the feeding sites (29 plantations), control areas (18 plantations) and also 68 randomly selected pine plantations. Before the experiment began in 1987 four plantations had been seriously damaged. During the study period only one plantation was seriously damaged. However, it could not be conclusively proved that damage to the pine plantations had been reduced as a result of the feeding sites. The results of the study can be put into practice elsewhere to create better living conditions for moose in their winter habitats. However, the food offered at the feeding site should be in the right proportion to the number of animals wintering in the area, so that the risk of damage to nearby plantations would be kept as small as possible.
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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.
The overall increase of ungulate populations in modern Europe has contributed to conflicts in national economies, particularly between game management and the forestry sector. This study assessed damage risks to young pine (Pinus sylvestris L.), spruce (Picea abies (L.) H. Karst.) and aspen (Populus tremula L.) stands at two spatial scales. One level assessed the interaction between sex-age structure of cervid populations, measured by pellet group density, and forest damages, measured as the percentage of heavily browsed trees in 2040 stand surveys distributed proportionally throughout the country. The second level compared pellet counts and trail-camera-based records of moose (Alces alces L.) and red deer (Cervus elaphus L.) presence in a pilot study area. We examined whether 1) there is a correlation between damage amount and ungulate population structure and 2) are the data from trail cameras suitable for wider use in monitoring ungulate population structure. The study confirmed significantly higher moose pellet group densities in pine than in spruce and aspen stands. Pine damages were greater in stands with higher moose pellet group density, especially with female moose prevailing over male moose density index. The red deer pellet group densities were significantly higher in heavily damaged pine and spruce stands, regardless of sex-age structure. In most cases, there were no statistically significant differences between the two survey methods of ungulate population structure by using pellet count transects and trail camera fixations. Both methods provide comparable data on sex-age structure in moose and red deer populations if seasonal and habitat-predicted biases are considered. However, trail cameras are more widely applicable and easier to use by hunters than pellet counts.
Moose (Alces alces L.) browsing causes severe damage in Scots pine (Pinus sylvestris L.) seedling stands. The effects of this damage on the quality of sawlogs were studied in a long-term controlled experiment. This article reports the stem size and external quality characteristics of Scots pine stems 34 years after artificial moose browsing damage. Damaging the trees by clipping the main stem at the seedling stage reduced the diameter, height, and tree volume of the trees at the end of the experiment. The tree growth reduction was dependent on the severity of clipping. The differences between the damaged and the control trees were more obvious in diameter than in height at the time of final felling. Stem form defects and vertical branches were the most typical externally detectable defects caused by clipping. Defects in the butt logs were detected in 71–89% of the damaged trees, depending on the clipping treatment severity. The stronger the clipping treatment, the more likely the stem form was defected and the more commonly were vertical branches and crooks detected in the stems. The results indicate that both tree dimensions and stem quality suffer from moose browsing. The findings of this controlled experiment more likely underestimate than overestimate the damage in comparison to real moose browsing. Further analyses are required to assess the effects of browsing damage on the internal quality of sawlogs and subsequent economic outcomes.
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.
This study aimed at establishing calibrations to predict nutrient concentrations of solid moose (Alces alces L.) rumen content using near infrared spectroscopy (NIRS), as an alternative to expensive chemical analyses. NIR reflectance spectra of 148 dry pulverized samples were recorded. The scanned samples were then analyzed for crude protein, available protein, microbial nitrogen (N), ash, acid-detergent fiber (ADF), neutral detergent fiber (NDF) and lignin contents following standard chemical analysis procedures. The calibration models were derived by Orthogonal Projection to Latent Structure (OPLS) and validated using external prediction sets. The calibration models accurately predicted crude protein, available protein and ash contents (R2 = 0.99, 0.96, and 0.92, prediction error = 0.39, 0.72 and 0.53% dry matter, respectively) while NDF (R2 = 0.92; prediction error = 2.23% dry matter) and ADF (R2 = 0.89; prediction error = 1.94% dry matter) were predicted with sufficient accuracy and that of microbial-N (R2 = 0.81; prediction error = 1.25 mg yeast-RNA g–1 dry matter) and lignin (R2 = 0.84; prediction error = 1.05% dry matter) were acceptable. The ratio of performance to deviation values were > 3.0 for crude protein and available protein, between 3.0 and 2.5 for ADF, NDF and lignin, and 2.32 for microbial-N; attesting the robustness of the calibration models. It can be concluded that NIR spectroscopy offers a quick and inexpensive procedure for prediction of nutrient concentrations of solid rumen contents in wild herbivores.
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.
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.