Interactions among forestry regeneration treatments,plant vigour and browsing damage by deer.

In a large field experiment we studied the influence of regenerationmethods on the extent to which roe deer (Capreoluscapreolus L.) browse on spruce (Picea abies L.Karst) seedlings. Our objective was to evaluate if treatments that are good atstimulating regeneration may increase browsing damage by deer. Data werecollected between 1993 and 1995, and in the winters of 1993–94 and1994–95 the frequency of browsed seedlings was 13.2% and 13.4%,respectively. The frequency of browsing damage varied considerably betweensitesand clearcuts. Browsing damage was more frequent when the regeneration methodsincluded insecticide treatments. Among insecticide-free treatments, browsingwasheavier on scarified plots than on herbicide-treated or control plots (in whichseedlings were planted on untreated ground and given no post-plantingtreatment). Containerised seedlings were browsed more than bare-rooted ones.Neither the age of clearcut when planting, nor removal of slash, had any effecton browsing. It was demonstrated that regeneration methods that increased plantvigour (as assessed by leader growth and needle colour index) led to morebrowsing damage. Thus, our results support the plant vigour hypothesis, whichstates that many herbivores prefer to feed on vigorous plants because they aremore nutritious. However, the difference in browsing damage betweencontainerised and bare-rooted seedlings could not be explained solely by theplant vigour hypothesis.     

Frequencies and patterns of browsing by large herbivores on conifer seedlings

Browsing by large herbivores on planted and naturally regenerated conifer seedlings (Picea abies and Pinus sylvestris) was recorded in 104 clear‐cuts in east‐central Sweden during 1990 and in 47 clear‐cuts in 1991. The number of seedlings browsed and the browsing patterns were analysed in relation to seedling type. Browsing frequencies were also compared between forest stands with different site productivities and subjected to different management practices. The variation in the number of seedlings browsed in 1990 was explained mainly by seedling category. Among both planted and naturally regenerated seedlings, pine was browsed more than spruce. Two‐year‐old containerized seedlings of spruce was browsed more than 4‐yr‐old bare‐rooted spruce. In 1991, browsing was more equal among species and seedling types. Number of seedlings with their leader browsed and the amount of biomass left after browsing differed significantly between seedling types. Seedlings that had been browsed in 1990 experienced significantly higher browsing frequencies in 1991 when compared with unbrowsed seedlings. The effects of stand characteristics were not found to be significant.     

Height growth of young larch (Larix kaempferi) in relation to the frequency of deer browsing damage in Hokkaido, Japan

The effect of browsing frequency by Sika deer (Cervus nippon yezoensis) on the height growth of Japanese larch (Larix kaempferi) in Hokkaido, Japan, was analyzed. Tree heights at the time of the census and at the start of the current growing season were determined for 100 trees selected randomly at each site. The tallest shoot on each tree was identified and the number of browsing occurrences on the current-year shoot was determined. Young larches browsed by deer repeatedly develop new shoots during a growing season. Larch is a browsing-tolerant species, and the average height growth of the most heavily browsed trees was >15 cm. Although resprouting partly compensated for decreased height growth caused by browsing, the annual increase in height decreased with increased browsing frequency. Thus, browsing frequency during a growing season best reflects the intensity of browsing damage to larch trees.     

Mammalian browsing impact on regenerating <Emphasis Type="Italic">Eucalyptus</Emphasis> seedlings in a large commercially managed native forest estate

Excessive browsing by herbivorous mammals can adversely effect both forestry operations and ecosystem structure and function, and effective management of the problem across large forest estates requires the spatial extent of browsing impact to be determined. We conducted a browsing impact survey within 1.3 million ha of commercially managed Eucalyptus forest, and hypothesised that browsing impact would be related to variables reflecting habitat suitability for a major browsing species, the swamp wallaby (Wallabia bicolor Desmarest), at both large and small spatial scales. A simple spatial analysis showed that browsing severity varied across the landscape, but was high in a number of districts (30,000–100,000 ha land management units) grouped in a similar part of the surveyed area, identifying this region as a focus for management and research. Ordinal logistic regression showed that browsing impact was related to (a) the location of districts and (b) the number of times post-harvest regeneration had been attempted, with browsing impact generally worse in areas where regeneration had occurred twice. The latter result was consistent with our original hypothesis, as multiple regeneration attempts promote high quality habitat patches for wallabies. In most heavily browsed districts browsing severity varied substantially between harvested sites, suggesting that processes operating at the site scale were important. Our results describe the spatial extent of browsing within the surveyed area, and identify high risk localities and states, but additional data are required to understand the processes regulating browsing impact.     

Field screening for genetic-based susceptibility to mammalian browsing

Browsing by mammalian herbivores is a major problem in plantation forestry. Seedlings are most vulnerable to browsing during establishment, making protection crucial during this period. Aside from reducing herbivore numbers, browsing can be controlled through the application of tree guards or chemical repellents. These methods are generally short-term options. A promising, potentially longer-term method of reducing browsing damage on plantations is to plant individuals with enhanced natural browse resistance. This requires the development of a rapid, cost-effective means of identifying germplasm with enhanced resistance. Here we present such a screening methodology.We planted Eucalyptus globulus seedlings from 22 different seedlots in randomised blocks along edge rows of six operational plantations. Seedlings were monitored for mammal browsing damage and growth for 2 years. Natural variation between E. globulus seedlots resulted in significant variation in the extent to which they were browsed. Differential browsing was evident just 10 weeks after planting, and caused a significant effect on tree growth after 2 years. Differential browsing was thought to be due to variation in levels of plant secondary metabolites (PSMs). The identification and selective planting of high PSM seedlots in high risk areas could be an effective management tool to reduce browsing in plantation forestry.     

Growth limitation of mountain birch caused by sheep browsing at the altitudinal treeline

Treelines are advancing on a global scale and encroaching upon alpine ecosystems. Browsing by vertebrate herbivores could affect treeline dynamics and forest expansion by limiting growth of trees. However, this has not been experimentally investigated, and due to a combination of herbivore behaviour and plant responses to herbivory, is unlikely to be a simple relationship. A long term (8 years of manipulation), large scale (0.3 km²) sheep enclosure experiment, in the Scandes of Norway, was used to investigate the effect of three densities of sheep (no sheep, low and high densities with respectively 0, 25 and 80 sheep km⁻²) on tree growth within a mountain birch (Betula pubescens tortuosa) treeline ecotone. Birch trees were most likely to be browsed at intermediate heights of around 100-130 cm and the likelihood of an individual being browsed did not differ between low and high sheep densities. However, when browsed, the intensity of browsing increased with sheep density. High densities of sheep reduced the net annual shoot growth of young mountain birch in comparison to growth in the absence of sheep. The long-term height growth of smaller mountain birch (stem diameter < 15 mm) was also reduced at high densities of sheep, but the height growth of larger birch was resistant to browsing, even though these trees were on average still within browsing height. This study shows that susceptibility to browsing in birch decreases with size, but by restraining the height growth of trees high densities of sheep can keep trees at heights at which they remain susceptible to browsing. Reductions in livestock density over a period as short as 8 years can thus facilitate altitudinal forest expansion.     

Effects of browsing on recruitment and mortality of European aspen (Populus tremula L.)

Abstract

High browsing pressure may reduce or halt regeneration of browsing-prone tree species. In this study, exclosures were used to quantify the effects of browsing on vital demographic parameters in European aspen (Populus tremula L.) stands. The experiment focused on stands located in managed forest landscape with densities of 0.85 moose (Alces alces L.)km^?2. Other herbivores such as mountain hare (Lepus timidus L.) were present at densities common for this region and roe deer (Capreolus capreolus L.) at low densities. Recruitment and mortality of aspen ramets were recorded before and after the growing season, and over four consecutive years. Seven to 19% of the ramets within browsing reach were browsed annually. No differences were found in recruitment or mortality attributable to browsing per se. Instead, there were large among-stand variability in vital population parameters unrelated to browsing incidence. In general, aspen stands with high ramet density showed higher recruitment rates than stands with low ramet density. There was substantial variability in self-replacement capability unrelated to ecological factors. The results suggest that more attention should be given to finding and managing stands with high capacity for suckering, as these stands may have a disproportionate impact on aspen recruitment at the landscape scale.     

Short-term browsing by roe deer has little effect on survival and growth of sessile oak seedlings

We studied the effect of roe deer (Capreolus capreolus) browsing on the growth and mortality of sessile oak (Quercus petraea) seedlings. First, we measured the height and basal diameter, and counted the number of shoots of the 849 seedlings of an experimental plantation, 60 of which were individually protected by fencing. We then introduced two tame roe deer females every day for three weeks from late May to mid-June. Measurements and counts were repeated after roe deer removal, then once again the following winter. We found no significant effect of browsing on mortality, total height, or basal diameter of the seedlings. However, the browsed seedlings exhibited a lower number of shoots despite regrowth observed after roe deer removal. We propose to use the tolerance of oak to deer browsing to promote natural and inexpensive methods, to protect oak during regeneration, such as the spatial arrangement of plants or the use of slash or surrounding vegetation in favourable environments.     

Can Eucalyptus invade native forest fragments close to commercial stands?

Some Eucalyptus species are widely used as a plantation crop in tropical and subtropical regions. One reason for this is the diversity of end uses, but the main reason is the high level of wood production obtained from commercial plantings. With the advancement of biotechnology it will be possible to expand the geographical area in which eucalypts can be used as commercial plantation crops, especially in regions with current climatic restrictions. Despite the popularity of eucalypts and their increasing range, questions still exist, in both traditional planting areas and in the new regions: Can eucalypts invade areas of native vegetation, causing damage to natural ecosystems biodiversity?The objective of this study it was to assess whether eucalypts can invade native vegetation fragments in proximity to commercial stands, and what factors promote this invasive growth. Thus, three experiments were established in forest fragments located in three different regions of Brazil. Each experiment was composed of 40 plots (1 m² each one), 20 plots located at the border between the forest fragment and eucalypts plantation, and 20 plots in the interior of the forest fragments. In each experimental site, the plots were paired by two soil exposure conditions, 10 plots in natural conditions and 10 plots with soil exposure (no plant and no litter). During the rainy season, 2 g of eucalypts seeds were sown in each plot, including Eucalyptus grandis or a hybrid of E. urophylla × E. grandis, the most common commercial eucalypt species planted in the three region. At 15, 30, 45, 90, 180, 270 and 360 days after sowing, we assessed the number of seedlings of eucalypts and the number of seedlings of native species resulting from natural regeneration. Fifteen days after sowing, the greatest number of eucalypts seedlings (37 m⁻²) was observed in the plots with lower luminosity and exposed soil. Also, for native species, it was observed that exposed soil improved natural germination reaching the highest number of 163 seedlings per square meter. Site and soil exposure were the factors that have the greatest influence on seed germination of both eucalypt and native species. However, 270 days after sowing, eucalypt seedlings were not observed at any of the three experimental sites. The result shows the inability of eucalypts to adapt to condition outside of their natural range. However, native species demonstrated their strong capacity for natural regeneration in forest fragments under the same conditions where eucalypts were seeded.     

Influences of seedling size,container type and mammal browsing on the establishment of <Emphasis Type="Italic">Eucalyptus globulus</Emphasis> in plantation forestry

Rapid early growth of tree seedlings is critical to the success of plantation establishment. We investigated the effects of seedling size (small and large) and container types (small [Lannen 121], medium [Lannen 81] and large [Forestry Tube]) in the nursery and the effects of mammal browsing after planting on growth of Eucalyptus globulus in Tasmania’s Southern Forests. After planting, seedlings were either exposed to browsing or protected from browsing by wire-mesh cages until age 6 months. Low browsing pressure resulted in around 20% and 5–10% of foliage being browsed in the large and small size categories, respectively, between 1 and 3 months after planting the uncaged treatment. 6 months after planting, height growth increment was lower, and 4 years after planting, mortality was higher in uncaged large than caged large seedling treatments. Six and twelve months after planting, seedlings raised in Forestry Tube containers had significantly greater height increment and root collar diameter relative to other treatments. By 4 years after planting, trees of the small seedling treatment had significantly greater diameter than those of the large seedling treatment, but there was no effect of container type treatment. Four years after planting there was no effect on diameter growth by browsing of <30% of foliage up to 3 months after planting, although there was greater incidence of double leaders in trees that had been browsed as seedlings. Small seedlings produced more growth 4 years after planting than large seedlings of E. globulus.     

Effects of 13-year fencing on browsing by sika deer on seedlings on Mt. Ohdaigahara, central Japan

We examined the effects of fencing on deer browsing on seedlings 13 years after the building the experimental fence on Mt. Ohdaigahara, central Japan, where the sika deer (Cervus nippon) population is high. There was no difference in the species number of seedlings between 1991 and 2004. The density of seedlings in the fenced plot in 2004 was three times higher than in 1991. Although the unfenced plots had no seedlings higher than 20 cm, there were seedlings up to 120 cm in the fenced plot. These results suggest that 13-year fencing promoted regeneration of seedlings in this area.     

Winter Browsing by Large Herbivores on Planted Deciduous Seedlings in Southern Sweden

Deciduous seedlings are valuable for the forest industry and, at the same time, often palatable for large herbivores. Therefore, browsing on seedlings of seven deciduous tree species and on Picea abies(L.) Karst., as a reference species, was tested in a field experiment with the species presented at feeding stations. Twenty stations, each with one seedling per species, were established in each of 15 clear-cuts in southern Sweden. The planting took place in October. Depending on species, between 16 and 85% of the seedlings were browsed after one winter. The descending order, based on number of browsed seedlings, was Quercus robur L . > Alnus glutinosa (L.) Gaertn.> Fagus silvatica L.> Tilia cordata Mill.> Prunus avium L.> Betula pendula Roth> Picea abies (L.) Karst. > Fraxinus excelsior L. On 83% (range 45-100%) of the browsed seedlings the leader was damaged. On average, browsing reduced seedling hight by 32% (range 12-47%). Within species, initially taller seedlings were more often selected than shorter ones.     

Biotic injuries on conifer seedlings planted in forest understory environments

We investigated how partial overstory retention, understory vegetationmanagement, and protective Vexar^® tubing affected the frequency andseverityof biotic injuries in a two-storied stand underplanted with western redcedar(Thuja plicata Donn ex D. Don), Douglas-fir(Pseudotsuga menziesii (Mirb.) Franco, grand fir(Abies grandis (Dougl.) Lindl), and western hemlock(Tsuga heterophylla (Raf.) Sarg.). The most prevalentsource of damage was browsing by black-tailed deer (Odocoileushemionis columbiana); deer browsed over 74% of Douglas-fir and over36% of western redcedar seedlings one or more times over the four years of thisstudy. Neither the spatial pattern of thinning (even or uneven) nor the densityof residual overstory affected browsing frequency. Spraying subplots may haveslightly increased browsing frequency, but the resulting reduction of theadjacent understory vegetation increased the volume of all seedlings by 13%,whether or not they were browsed. Vexar^® tubing did not substantiallyaffectseedling survival, browsing damage frequency, or fourth-year volume. Greaterlevels of overstory retention reduced frequency of second flushing. Chafing bydeer and girdling by rodents and other small mammals began once seedlingssurpassed 1 m in height. Essentially all grand fir seedlingsexhibited a foliar fungus infection.     

Young conifer stands form a deer browsing refuge for an oak admixture: silvicultural implications for forest regeneration under herbivore pressure

There is little knowledge how ungulate pressure on forest regeneration may be mitigated by silvicultural methods. The knowledge is especially needed for artificially regenerated, deciduous tree species. We studied factors affecting browsing incidence by deer in the Pisz Forest District in Poland, an area where 10,000 ha of forest was damaged by a 2002 hurricane. In 2006, we established three experimental plots (in total, 22.6 ha), in which the main species was Scots pine (Pinus sylvestris) admixed with pedunculate oak (Quercus robur). The data on browsing were collected in 2008–2015. In general, oak browsing incidence was unrelated to oak planting density. On a plantation scale, it was significantly affected by the pine age. Although in each variant all the oaks were browsed for four consecutive years (2009–2012), in 2013 browsing incidence began to decrease. When the pines grew higher and formed a physical barrier, it was harder for deer—roe deer (Capreolus capreolus), red deer (Cervus elaphus) and moose (Alces alces)—to move through and locate the oaks. Moreover, within plantations, oak browsing incidence was higher in the patches with shorter pines. Browsing of individual saplings or small groups of saplings was also negatively affected by the height of neighbouring pine saplings. Oak density influenced deer selectivity depending on the tree height. In a low oak tree density, browsing incidence was unrelated to oak height, while in higher tree density, deer selected oaks of the height between 40 and 100 cm. We postulate that deciduous admixture in a coniferous (unattractive) stand can be planted with a few year delay. Older coniferous trees should impede locating of attractive tree species by deer and the browsing incidence.     

A dwarf bamboo (<Emphasis Type="Italic">Pleioblastus chino</Emphasis>) and winter browsing by Japanese hare (<Emphasis Type="Italic">Lepus brachyurus</Emphasis>) combine to limit establishment of transplanted native tree seedlings in an abandoned agricultural field

Natural forest recovery on abandoned farmland is hindered by a variety of factors and active restoration plays an important role when quick afforestation is desired. We investigated seedling survival of four transplanted native tree species (Quercus myrsinifolia, Quercus serrata, Aphananthe aspera, and Rhus sylvestris) by experimentally manipulating the vegetation cover, which was mainly dominated by dwarf bamboo (Pleioblastus chino), and herbivore access to the planting sites on farmland that had been abandoned for 15 years at the start of the study. Few transplanted seedlings of any species survived under intact vegetation cover, irrespective of herbivore presence. In gaps in the vegetation cover, winter browsing by Japanese hare (Lepus brachyurus) damaged all species. However, lower browsing frequency and higher resprouting ability after grazing of the seedlings enabled both Quercus species to survive better than the other species. These results indicate that dwarf bamboo and the hare jointly limit the establishment of native trees in old fields. If active afforestation by transplanting seedlings at sites dominated by dwarf bamboo is planned, a combination of vegetation removal, selection of suitable species, and temporary seedling protection will be most effective.     

Long-term browsing impact around diversionary feeding stations for moose in Southern Norway

Diversionary winter feeding of browsing ungulates is an increasingly common management practice although evidence for its efficacy to reduce excessive browsing remains ambiguous. Moreover, comparative estimates on changes in browsing pressure (proportion of available shoots browsed) following long-term winter feeding are currently lacking. We quantified spatiotemporal changes in browsing pressure of moose (Alces alces L.) on commercial and non-commercial tree species around 30 feeding stations after 15–20 years of winter feeding. The results were compared with browsing pressure indices recorded at the same feeding stations 10 years previously. We expected leader stem and lateral twig browsing to have increased over time at a fine spatial scale (≤200 m from feeding station) and to have increased in spatial extent away from feeding stations. Furthermore, we tested whether moose browsing patterns conformed to central-place foraging theory. Despite 2–3-fold higher faecal pellet group numbers in the vicinity of feeding stations, leader stem browsing increased only on the commercially valuable Norway spruce (Picea abies (L.) Karst), a species normally avoided by moose. Lateral twig browsing largely decreased within 200 m of feeding stations while at a broader scale (≤1 km from feeding stations), leader stem browsing was high on most tree species (ca. 60% of available stems browsed) as expected, Peak browsing on lateral twigs of Scots pine (Pinus sylvestris L.) occurred further from feeding stations after 15–20 years of winter feeding than 10 years earlier. Browsing pressure on active feeding stations (n = 18) was comparable to that on inactive feeding stations (n = 12), suggesting the occurrence of rebrowsing. The ability of central-place foraging theory to explain fine-scale browsing patterns around feeding stations decreased as diversionary feeding continued over time. Long-term diversionary winter feeding of moose in unproductive boreal forests has serious implications for the intensity of fine-scale browsing pressure, which may lead to resource depletion close to feeding stations followed by high browsing pressure at distances further away from feeding stations. Our study emphasizes the importance of considering the longevity of a winter feeding programme at its inception.     

Effects of a nursery CO2 enriched atmosphere on the germination and seedling morphology of two Mediterranean oaks with contrasting leaf habit

The use of an enriched CO₂ atmosphere in tree nurseries has been envisaged as a promising technique to increase productivity and to obtain seedlings with a higher root/shoot ratio, an essential trait to respond to water stress in Mediterranean-type ecosystems. In that framework, we have analyzed the effects of three levels of atmospheric CO₂ concentration (350, 500 and 700ppm) on the germination rate, growth and morphology of seedlings of two Mediterranean oaks used in reforestation programs: the evergreen Quercus ilex L. and the deciduous Quercus cerrioides Wilk. et Costa. CO₂ enrichment increased the germination rate of Q. cerrioides (from 70±7 to 81±3%) while it decreased that of Q. ilex (from 71±10 to 41±12%). Seedlings of both species increased approximately 60% their total biomass in response to CO₂ enrichment but at two different CO₂ concentrations: 500ppm for Q. cerrioides and 700ppm for Q. ilex. This increase in seedlings biomass was entirely due to an augmentation of root biomass. Considering germination and biomass partitioning, an enriched CO₂ atmosphere might not be appropriate for growing Mediterranean evergreen oaks, such as Q. ilex, since it reduces acorn germination and the only gains in root biomass occur at a high concentration (700ppm). On the other hand, a moderate CO₂ enrichment (500ppm) appears as a promising nursery technique to stimulate the germination, growth and root/shoot ratio of deciduous oaks, such as Q. cerrioides.     

In vitro effects of four ectomycorrhizal fungi, Boletus edulis, Rhizopogon roseolus, Laccaria laccata and Lactarius deliciosus on Fusarium damping off in Pinus nigra seedlings

Species of Fusarium cause damping off of forest seedlings resulting in considerable losses and oftentimes fungicides are used to control the disease. Such chemicals may be detrimental to the environment. Protection of forest seedlings against this disease must be focused on integrated management for in which biological control is one of the most important tools. Mycorrhizal fungi may provide protection against pathogen invasion of seedlings. In this study, the interactions between the mycorrhizal fungi Boletus edulis, Rhizopogon roseolus, Laccaria laccata and Lactarius deliciosus and damping off pathogens (Fusarium oxysporum and F. moniliforme) were investigated in vitro and mechanisms involved in the protection of damping off in P. nigra seedlings are also discussed. The effect of filtrates from mycorrhizal species on spore germination of F. oxysporum varied throughout the tests. At the end of the assay, the inhibitory effect only could be observed in the L.␣deliciosus treatment. None of the filtrates reduced spore germination in F. moniliforme. Finally, three of the four mycorrhizal species grown for 2 months in the substrate, L. laccata, L.␣deliciosus and B. edulis, increased the survival of Fusarium inoculated P.␣nigra seedlings.     

Can spines deter deer browsing?: a field experiment using a shrub <Emphasis Type="Italic">Damnacanthus indicus</Emphasis>

We examined the defensive function of spines of Damnacanthus indicus against deer herbivory by experimentally removing spines in the field where deer density is high. Individuals of D. indicus whose spines had been removed had a higher probability of being browsed by deer than control individuals. In addition, plant height was significantly lower for individuals that were browsed than those that were not. These results support the notion that spines of D. indicus have a function to deter deer browsing. This study is the first to demonstrate experimentally the defensive function of spines against large mammals in temperate areas.     

Prevalence and incidence of the root-inhabiting fungi, Fusarium, Cylindrocarpon and Pythium, on container-grown Douglas-fir and spruce seedlings in British Columbia

Surveys were made at the end of the 1990 and 1991 growing seasons for root-inhabiting fungi in the genera Fusarium, Cylindrocarpon and Pythium from the roots of one year-old container-grown Douglas-fir and spruce seedlings grown under greenhouse conditions. In the 1990 survey of four nurseries, it was found that 61–97% of both Douglas-fir and spruce roots were colonized with Fusarium, Cylindrocarpon or Pythium. There were significantly (p0.05) more Douglas-fir roots than spruce roots colonized by Fusarium at all nurseries, however, there were significantly (p0.05) more spruce roots than Douglas-fir roots colonized by Cylindrocarpon and Pythium. Root colonization of Douglas-fir and spruce by the three fungal genera during 1991 varied from 0–82% at three nurseries, however, only at a south coastal nursery was there significantly (p0.05) more spruce than Douglas-fir roots colonized by Cylindrocarpon. Significantly more seedlings were infected in 1990 than in 1991. In 1991, there were few significant differences between Douglas-fir and spruce, in the percentage of seedlings with colonized roots and in the percentage of growth medium colonized by the fungi. However, there were significant differences between nurseries.