Effects of roe deer browsing and site preparation on performance of planted broadleaved and conifer seedlings when using temporary fences

Abstract

The effects of fencing and site preparation on performance of birch (Betula pendula Roth), oak (Quercus robur L.), pine (Pinus sylvestris L.) and spruce [Picea abies (L.) Karst.] seedlings were studied over four growing seasons in southern Sweden. The experiment was composed of four browsing treatments, from which roe deer (Capreolus capreolus L.) were excluded for 0, 12, 24 or 42 months, and four soil treatments. Natural browsing was combined with artificial browsing (clipping). It was demonstrated that browsing by roe deer could be isolated from browsing by moose (Alces alces L.) by selective fencing. Except for birch, browsing had little effect on seedling survival. Browsing reduced height growth by more than 100% for oak and pine, and more than 60% of pine seedling developed multiple stems. Except for oak, site preparation increased seedling survival. Inverting site preparation, in combination with or without fertilization, produced positive growth responses in seedlings, whereas patch scarification did not. There was no interaction between browsing and soil treatments on seedling performance. It was concluded that long-term protection against browsing by roe deer is needed for oak and pine, whereas short-term fences might be used for birch, and that spruce is relatively unaffected by browsing.     

Incidence of Cronartium ribicola in Finnish botanical gardens and five-needle pine plantations in northern Finland

Abstract

Stump removal procedure increases the extent of exposed mineral soil in the clear felled areas. In this study, our aim was to find out whether the early growth and mycorrhization of Norway spruce (Picea abies (L.) Karst.) seedlings differ between stump removal and mounding sites. Stumps were harvested from five one-hectare study sites and other five sites were mounded after clear felling. Twenty seedlings were planted on mounds at each study site. Although the height of spruce seedlings did not differ between the treatments after three growing periods, their mean growth was ca. 10% higher at the stump removal sites. The community of ectomycorrhizal (ECM) in the seedling roots did not differ between the treatments. The high mycorrhization rate in the nursery still remained after two and three growing seasons in the field and diversity of ECM community slightly increased during the first years. Our results indicate some positive although rather small and transient effects of stump removal on the performance of planted spruce seedlings. Longer-term studies are needed to fully understand the effects of stump removal on the later seedling development and their symbiotic interactions with mycorrhizal fungi.     

Susceptibility of different provenances of Pinus sylvestris,Pinus Contorta and Picea abies to Gremmeniella abietina

In a randomized block factorial experiment, 1200 seedlings from four provenances, each of Pinus sylvestris, Pinus contorta and Picea abies were inoculated with conidia (2 × 10⁴ and 1 × 10⁶ conidia/seedling) of Gremmeniella abietina (Brunchorstia pinea), isolated from P. contorta plantations in northern Sweden. A further 600 seedlings were left as controls. The occurrence of symptoms and the extension of dead tissues on the annual shoots were recorded 13 months after inoculation. Only the higher spore dose resulted in significant infection. P. sylvestris and P. contorta seedlings were equally susceptible (53% infected), and significantly more infected than P. abies seedlings (39%). The annual shoots of P. sylvestris and P. abies were affected to 43% and 37% of their length, which was significantly more than 15% of the length of P. Contorta shoots. The frequency of affected seedlings differed between the most southern and northern provenances of P. sylvestris and P. abies, and between the northwestern and the south-eastern provenance of P. contorta Twenty-six months after inoculation, a higher proportion of P. contorta seedlings than P. sylvestris and P. abies seedlings had recovered, and a lower proportion of P. contorta than P. sylvestris and P. abies seedlings had died. This paper discusses why P. sylvestris was found to be more susceptible to G. abietina than P. contorta in this experiment, while the reverse is found in plantations in northern Sweden.     

Incidence of Gremmeniella abietina in planted seedlings of Picea abies and Pinus sylvestris in northern Finland

The ascomycete Gremmeniella abietina causes a disease (scleroderris canker) on conifers. The pathogen kills terminal shoots and branches of Pinus sylvestris and Picea abies, sporulates on host shoots and causes leader changes (i.e. dieback). Damage caused by G. abietina was investigated in young P. abies and P. sylvestris plantations in northern Finland. Side branches from seedlings of both species and the main stems of P. sylvestris were collected from three sites in two locations (Poksa 1 and 2, Kivalo) in northern Finland. The number of cankers and leader changes was counted in branches of each age (i.e. year) in a total of ca. 6300 shoots of P. abies and 1200 shoots of P. sylvestris. Cankers were common on both P. abies and P. sylvestris in Poksa 1 (2002–2008) and on P. abies in Poksa 2 (1998–2001). In Kivalo, cankers occurred only sporadically on P. sylvestris. Leader changes were most frequent in 1999–2009 in Poksa 1, in 2001–2009 in Poksa 2 and in 2003 in Kivalo. Cankers and other symptoms of infection were more obscure on P. abies than on P. sylvestris. According to both conidial morphology and molecular analysis, the strain of G. abietina infecting both P. abies and P. sylvestris was small‐tree type (STT). This is the first report of G. abietina, STT or B type, injuring P. abies in plantations.     

Predicting regeneration establishment in Norway spruce plantations using a multivariate multilevel model

This study predicts the regeneration establishment on 3-year-old Norway spruce (Picea abies (L.) Karst.) plantations in southern Finland using regeneration survey data. Regeneration establishment was described by seven response variables: number of planted spruces, natural Scots pines (Pinus sylvestris L.), natural spruces, natural seed-origin birches (Betula pubescens Ehrh. and B. pendula Roth.) and other broadleaves (i.e. sprout-origin birches and other broadleaves than birch), as well as height of crop-tree spruce and dominant height of broadleaves. Due to the multivariate (several responses for each plot) and multilevel (plot, stand, municipality, forest centre) structure, regeneration establishment was modelled by fitting a multivariate multilevel model with explanatory variables such as temperature sum, site fertility, soil quality and method of site preparation. In the model, the numbers of tree seedlings were modelled using over-dispersed Poisson distributed equations, and the tree heights were modelled using normally distributed linear equations. The estimated fixed and random parameters of the equations were logical, and there was no serious bias in predicting the regeneration establishment in the independent test data set. This modelling approach can be used to predict the regeneration establishment stochastically by taking into account the large unexplained variation in regeneration models.     

Effect of uninucleate Rhizoctonia on Scots pine and Norway spruce seedlings

One‐year‐old container‐grown seedlings were planted in spring on clear cut areas: the Norway spruce (Picea abies) on a moist upland site (Myrtillus‐type) and Scots pine (Pinus sylvestris) on a dryish upland site (Vaccinium‐type). While still in the nursery, half of the seedlings of each species had been inoculated during the previous summer, with a uninucleate Rhizoctonia sp., a root dieback fungus. At outplanting all the seedlings appeared healthy and had a normal apical bud, although the height of the inoculated seedlings was less than that of the uninoculated control seedlings. At the end of the first growing season after planting, the mortality of inoculated Scots pine and Norway spruce seedlings was 25 and 69%, respectively. After two growing seasons the mortality of inoculated seedlings had increased to 38% for Scots pine and 93% for Norway spruce. The mortality of control seedlings after two growing seasons in the forest was 2% for Scots pine and 13% for Norway spruce. After outplanting the annual growth of inoculated seedlings was poor compared with the growth of control seedlings. These results show that, although Rhizoctonia‐affected seedlings are alive and green in the nursery, the disease subsequently affects both their survival and growth in the forest.     

Reduction of frost heaving of Norway spruce and Scots pine seedlings by planting in mounds or in humus

This study was carried out in northern Sweden to determine the effects of frost heaving on the establishment of Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings in relation to planting methods. For this purpose, one year old containerised seedlings were planted on two sites and on two dates: during the spring (early planting) and during the fall (late planting). In each case, two planting depths (normal and deep planting) and four planting sites (in mineral soil in the depressions, in the scalp/trench area, on the top of the mound and in the untreated humus layer) were used. On each site, 50 seedlings were planted for each treatment. Frost heaving was observed and measured during two years. The amount of heaving was highest in the hole and almost insignificant on the top of the mound and in the humus layer. Planting depth influenced the degree of heaving only for Scots pine planted in the hole and was not related to the planting time.     

The effect of girdling on the moisture content of small-sized trees

In this study, the effect of girdling on the moisture content of small-sized trees for heat energy production was clarified. The moisture content was measured for Scots pine (Pinus sylvestris), Norway spruce (Picea abies), and Downy birch (Betula pubescens) during two growing seasons after girdling. The trees were girdled at breast height for around 30 cm by removing the bark, phloem, and cambium from around the stem. At the beginning of the growing season the mean moisture content of the living Scots pine (P. sylvestris) and Norway spruce (P. abies) was 60%, and for Downy birch (B. pubescens) it was 50%. During the first growing season the effect of girdling on the moisture content was low, but during the second growing season the moisture content decreased significantly. The moisture content of the Norway spruce (P. abies) (23%) and Downy birch (B. pubescens) (33%) was at its lowest point at 14 months after girdling. There were no significant changes in the moisture content of the Scots pine (P. sylvestris) in this study. The results of this study can be used in basic research and in the development of energy wood production.     

Outdoor winter storage of container stock on raised pallets—effects on root zone temperatures and seedling growth

Containerized seedlings of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) were overwintered on the ground and above ground on pallets. Soil temperatures in elevated containers were lower and showed greater fluctuation than containers on the ground. The lowest temperatures (‐15 to ‐16°C) were observed in containers stored on pallets with little or no snow cover during the winter. Temperatures in the edge rows of containers were lower than interior bed soil temperatures. Lower temperatures were also observed in the top than in the bottom of the container. The storage on pallets resulted in reduced shoot and root growth. Although insulation preventing air movements beneath the container units improved soil temperature conditions and subsequent seedling growth, the best result was obtained when seedlings were stored directly on the ground surface.     

Regeneration of Picea abies forests on highly productive peatlands—clearcutting or selective cutting?

The study was focused on severe forest regeneration problems that are typical for highly productive peatlands. The aim of the study was to give recommendations for practical forestry on how to renew the forests. Experiments with different forest regeneration methods on highly productive peatlands were set up in nine mature Norway spruce (Picea abies (L.) Karst.) forests in northern, central and southern Sweden. The treatments in the study were natural regeneration in shelterwoods (at densities of 140 and 200 stems ha^‐1), planting of bare‐root spruce seedlings after site preparation (mounding) in the shelterwoods and on clearcuts, and planting without site preparation on clearcuts.

Judging from the extent of windthrow in the denser shelterwoods and the stocking of natural regeneration under remaining shelter trees 4–5 years after final cut, shelterwood regeneration is a promising method. On clearcuts, planting without site preparation resulted in poor seedling survival, large extent of damage to the seedlings and small height increment. Planting in mounds on the clearcuts was much more successful. The best survival of planted seedlings was observed in the shelterwoods. Both the denser and the sparse shelterwoods seemed to give satisfactory protection to the seedlings.

Final recommendations on the most suitable forest renewal method(s) will be made in about five years after the removal of the shelter trees.     

Root growth capacity and field performance of Pinus sylvestris and Picea abies seedlings

Correlations between root growth capacity (RGC), at the time of planting, and field performance were studied for Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings. Before planting a gradient in seedling viability was generated through exposure to low root temperatures and different winter storage regimes. The hypothesis that high RGC values would improve field performance was to some extent verified for pine seedlings while no correlations could be registered for spruce. Reasons for these results are discussed.     

Natural regeneration on planted clear‐cuts in boreal Sweden

Natural regeneration was registered on 97 clear‐felled and scarified sites in northeastern Sweden ten growing seasons after planting. On each site, three fix plots were laid out at planting and the planted seedlings were mapped. Ten years later all healthy non‐planted seedlings were registered in an anulus of size 15.7 m². Non‐planted seedlings were also observed in four 4 m² squares on each fix‐plot to estimate the zero‐square frequency. Species were classified as Pinus sylvestris, L., Picea abies (L) Karst. and deciduous species (Betula pendulajpubescens Roth./Ehrh., Populus tremula L., Alnus incana (L) Moench.). On average, 7 133 non‐planted seedlings ha^‐1 were found, of which 29% were conifers. Among the conifers, 71% were P. abies. More than 2 500 conifer non‐planted seedlings ha^‐1 were found on 27% of the plots. The number of non‐planted seedlings was correlated to the presence of neighbouring seed‐producing stands within 100 m and decreased with rising altitude. Pinus sylvestris seedlings were more frequent on dry soils while the deciduous seedlings were more frequent on wet sites. In the 4 m² squares, the zero‐square frequency was 29% if both conifers were and deciduous seedlings were considered and 63% if only conifers were considered. Mean height of the tallest seedling per species and square was 62 cm for P. sylvestris, 42 cm for P. abies and 107 cm for the deciduous seedlings. Key words: Alnus incana, Betula, broad‐leaves, deciduous, Norway spruce, Picea abies, Pinus sylvestris, Populus tremula, reforestation, Scots pine.     

Effects of nickel and copper on growth and mycorrhiza of Scots pine seedlings inoculated with Gremmeniella abietina

Scots pine (Pinus sylvestris) seedlings were planted in soil originating from two localities with different background levels of nickel and copper. In addition, some of the seedlings were exposed to additional nickel (20.5 mg Ni/l of soil) or copper (63.5 mg Cu/l of soil), or a combination of both Ni and Cu, via soil without direct shoot exposure during their second growing period. The seedlings were either irrigated with spring water (pH 6) or got only natural rain during the whole field experiment. All seedlings were inoculated with conidia of a shoot‐pathogen Gremmeniella abietina during their third growing season, and harvested the following spring. Lengths of shoots of different year‐classes were used as growth estimates. In roots, the proportion of fungal (assumedly mycorrhizal) biomass was estimated by measuring ergosterol concentration. Guajacol peroxidase activity was measured. Short roots were classified into two groups according to their condition and the composition of the mycorrhizal community was expressed as a proportion of morphotypes in the roots. The seedlings exposed to additional Ni had higher shoot growth than the seedlings in the other treatments. The mean Ni concentration in the roots of seedlings exposed to additional Ni was 79 p.p.m. and in other seedlings 16 p.p.m. Additional Ni also decreased the frequency of clearly senescent short roots and the proportion of the mycorrhizal morphotype with the thinnest mantle. These results indicate that the Ni exposure levels used in this experiment had some positive effects on the seedlings. The relative fungal biomass was about 6% lower (p = 0.0981) in the fine roots of seedlings treated with additional Cu. The mean Cu concentration in the roots of seedlings exposed to additional Cu was 256 p.p.m. and in other seedlings 29 p.p.m. Peroxidase activity, which was used as a general stress indicator in this study, was not affected by any of the treatments. The shoot growth and the relative biomass of fungi in the fine roots were positively correlated in all seedlings, and this correlation was stronger in seedlings exposed to additional Ni that were not irrigated compared with seedlings not exposed to additional Ni that were irrigated. The frequency of asymptomatic infections of G. abietina was positively correlated with the proportion of senescent short roots in the irrigated seedlings but not in not‐irrigated seedlings. The general condition of seedlings may be an important factor for infection by G. abietina when moisture is high enough for the fungi to infect seedlings by conidia.     

Regeneration in windthrow areas in hemiboreal forests: the influence of microsite on the height growths of different tree species

Natural regeneration of windthrow areas is an important issue when planning forestry measures after forest disturbances. Seedling recruitment was investigated in storm-damaged hemiboreal mixed forests in eastern Estonia. The establishment and growth of seedlings from natural regeneration was registered for tree species in soil pits and in mounds of uprooted trees in stands that were either heavily or moderately damaged. Seedling growth is expected to be better in large but shallow soil pits created by uprooted Norway spruce [Picea abies (L.) Karst.] and poorer in small but deep pits created by the hardwoods in the area, silver birch (Betula pendula Roth.) and European aspen (Populus tremula L.). The most abundant regenerating species was birch. Pits hosted larger seedling numbers than mounds, due to soil instability in mounds. Rowan (Sorbus aucuparia L.) showed significantly faster growth than the other seedling species. Norway spruce pits were preferred to pits of other species by both birch and spruce seedlings. Black alder [Alnus glutinosa (L.) J. Gaertn.] did not show a preference for pits of a certain species of uprooted tree. Both spruce and rowan preferred hardwood mounds over spruce mounds. Storm severity also affected species composition: birch predominantly occurred on pits and mounds in heavily disturbed areas, while spruce was more abundant in the moderately damaged areas. The effects of advance regeneration and surrounding stands on seedling microsite preferences should be considered in future research and subsequent management recommendations.     

Influence of soil scarification on reindeer foraging and damage to planted Pinus sylvestris seedlings

Abstract

The effects of soil scarification on reindeer lichen cover and re-establishment, reindeer foraging and damage possibly caused by reindeer to planted Scots pine seedlings were studied during six growing periods in a field experiment established in a reindeer lichen site that was grazed by a herd of 500–600 reindeer each winter. Seedlings (at a density equivalent to 2000 ha^?1) were planted in mounds with mineral soil on top, in tracks with exposed mineral soil, in tracks with mixed organic material and mineral soil and in intact lichen mat (control) with no soil disturbance. The disturbed area varied from 0 to 28%. After six growing periods, the reindeer lichen cover and volume were 10–20% lower in the scarified plots than in the control plots. There was no clear evidence that the reindeer avoided foraging even in plots with the highest levels of soil disturbance. However, the behaviour of the reindeer during winter grazing seemed to be affected by both coverage of reindeer lichen (positively) and the proportion of exposed mineral soil (negatively). Damage possibly caused by reindeer trampling affected 8.2% of the living seedlings each year and approximately 50% of these seedlings were subsequently infected by fungal diseases (compared with 20% of viable undamaged seedlings). Scarification treatments that exposed mineral soil mildly resulted in higher survival and growth rates than the other treatments, especially the control.     

Evaluating the effect of plant water availability on inner alpine coniferous trees based on sap flow measurements

Climate simulations anticipate an increase in mean summer temperature with synchronous decrease in summer precipitation during the course of the current century in Central Europe. As a consequence, transpiration of forest trees and stands may be altered along with soil water availability. In this study, the effect of reduced plant water availability to conifers was investigated into an open Pinus sylvestris forest (Erico-Pinetum typicum; P. sylvestris 60 %, Picea abies 20 %; and Larix decidua 20 %) within the inner alpine dry valley of the Inn River in Tyrol, Austria. For reducing plant water availability, we installed a transparent roof construction above the forest floor to prevent precipitation to reach the soil. The roofed area covered 240 m² and included 10 trees. A respective number of 11 trees served as controls in the absence of any manipulation. Roofing significantly reduced plant water availability as indicated in lower predawn needle water potentials. Sap flow density (Q _s) was 63, 47, and 24 % lower in roofed P. sylvestris, P. abies, and L. decidua trees, respectively, as compared to control trees. Our findings suggest that P. sylvestris and P. abies behaves “isohydric” as they close their stomata relatively early under conditions of reduced plant water availability and thus stabilize their water relations, whereas L. decidua behaves “anisohydric” and maintains high transpiration rates.     

Beeting in Picea abies (L) karst. Growth and damage in a field experiment

A field experiment was established in two Picea abies (L.) Karst. plantations in southern Sweden to study yield and effects of competition on growth in beeted stands. Beeting was carried out in gaps where originally planted seedlings had been removed. Tree species used for beeting were P. abies, Pinus sylvestris L., Pinus contorta Dougl. and Larix decidua Mill × L. leptolepis Gord. The plantations were beeted when mean heights were 95 cm (Ullasjo) and 40 cm (Knäred). Effects of competition on diameter growth of P. abies were found when originally planted stands were 2.5 and 3.5 m in height. On both sites, diameter growth in stump height was significantly lower for seedlings beeted in small gaps (approx. 20 m²) than for seedlings beeted in the middle of large gaps (approx. 155 m²). This difference was larger at Ullasjo than at Knared. About 50 % of the P. sylvestris 70 % of the P. contorta and 80% of the L. decidua x L. leptolepis were dead or severely damaged six years after beeting. The main causes of damage were fraying and browsing by moose and roe‐deer.     

The influence of different forest organic matter on the growth of one-year old planted Norway spruce seedlings in a greenhouse experiment

A greenhouse trial was carried out to improve the knowledge of how forest organic matter could be utilized in site preparation and the choice of planting spot for Picea abies (L.) Karst. One-year old Norway spruce seedlings were grown for 26 weeks in pots containing pure mineral soil, forest organic matter in three different states of decomposition and combined treatments where the organic matter was mixed with or placed on the surface of the mineral soil (volume proportions of organic:mineral of 1:2). Watering was adapted to the water retention characteristics of each growing medium in order to keep the soil water potential between -4 and -5 kPa in all pots.In its pure form as well as combined with mineral soil, the moderately decomposed mor organic matter gave a higher seedling dry weight increment than the corresponding growing media containing the more decomposed humus. Adding mor or humus to mineral soil increased the seedling dry weight increment by 30–140% and 10–40%, respectively. The largest seedlings grew in the pure mor and humus. Mixing these forms of organic matter with mineral soil appeared to reduce seedling growth as compared to placing them on the mineral soil surface. The increment increase compared to plain mineral soil mostly consisted of proleptic growth and was probably due to the improved nutrient availability of the growing media. Fresh organic matter (chopped needles and twigs) seemed to impair seedling root function and reduced the dry weight increment by 30–50% as compared with pure mineral soil.The results suggest that as long as the water and temperature requirements are fulfilled planting Norway spruce seedlings without scarification and/or gathering extra mor and humus to the planting position should improve seedling growth as compared to the growth following scarification. Unmixed fresh needles and twigs or fresh needles and twigs combined with plain mineral soil should be avoided when planting Norway spruce seedlings.     

Pathogenitätstests mit Bursaphelenchus mucronatus an Kiefern und Fichtensämlingen in Deutschland

Pathogenicity tests with Bursaphelenchus mucronatus on pine and spruce seedlings in Germany Inoculation tests on Pinus sylvestris seedlings with a German, a Siberian and a French isolate of Bursaphelenchus mucronatus and on Picea abies seedlings using only the German isolate have clearly shown, under German field conditions, that the nematode can cause wilt symptoms on the apex of about 60% of the inoculated P. sylvestris plants. The nematodes in all these tests mostly remained near the inoculation site. However, further spread of the nematodes and wilt symptoms occured when a climate chamber, at 25°C, was used or when the French isolate was used outdoors. At 25°C, 10% of the inoculated plants with at least 1000 nematodes per seedling died. Seedlings' apices wilted if more than 50 nematodes per trunk were present, whereas smaller numbers of nematodes could cause partial wilting. Nevertheless, a few pine seedlings did tolerate high nematode-population densities, up to 966 per plant. No further progress of wilt of plants with symptoms was observed the following year. Inoculations of P. abies with a German isolate resulted in population development near the site of inoculation. Spruces largely tolerated nematode infestations without any development of wilt symptoms.     

First report of Calonectria montana causing damping‐off disease on pine and spruce seedlings in Europe

In 2015–2016, damping‐off symptoms were observed on Pinus sylvestris and Picea abies seedlings in four forest nurseries in Poland. Some Calonectria isolates were obtained from symptomatic seedlings by culturing pieces of necrotic tissue on malt extract agar.Morphological and molecular analyses revealed that isolates represent Calonectria montana, a recently described species from natural forests in China. The isolates represent the first detection of this species outside of Asia. Pathogenicity tests demonstrated that C. montana is a severe pathogen of P. sylvestris seedlings.