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.     

Habitat use of large ungulates in northeastern Germany in relation to forest management

Estimating large herbivore density has been a major area of research in recent decades. Previous studies monitoring ungulate density, however, focused mostly on determining animal abundance, and did not interpret animal distribution in relation to habitat parameters. We surveyed large ungulates in the Biodiversity Exploratory Schorfheide-Chorin using faecal pellet group counts. This allowed us to explore the link between relative ungulate abundance, habitat use, and browsing damage on trees in a region with several types of forest, including unharvested and age-class beech forests, as well as age-class pine forests. Our results demonstrate that roe deer and fallow deer relative abundance is negatively correlated with large tree cover, and positively correlated with the cover of small shrubs (Rubus spec., Vaccinium spec.), and winter food supply. Habitat use of roe deer and fallow deer, as estimated by counting faecal pellet groups, revealed a preference for mature pine forests, and avoidance of deciduous forests. This differential habitat use is explained by different distributions of high quality food resources during winter. The response of deer to understory cover differed between roe deer and fallow deer at high cover percentages. The amount of browsing damage we observed on coniferous trees was not consistent with the relative deer abundance. Browsing damage was consistently higher on most deciduous trees, except for beech saplings which sustained less damage when roe deer density was low. Because roe deer is a highly selective feeder, it was reported to affect tree diversity by feeding only on trees with high nutritional value. Consequently, we propose that managing the number of all deer species by hunting is necessary to allow successful forest regeneration. Such an adjustment to deer numbers would need to account for both current tree diversity and alternative food resources. Our findings may be applicable to other forest landscapes in northeastern Germany including mature pine stands and differently harvested deciduous forests.     

Variation of stem density and vegetation carbon pool in subtropical forests of Northwestern Himalaya

The present study was conducted in five forest types of subtropical zone in the Northwestern Himalaya, India. Three forest stands of 0.1 ha were laid down in each forest type to study the variation in vegetation carbon pool, stem density, and ecosystem carbon density. The stem density in the present study ranged from (483 to 417 trees ha^?1) and stem biomass from (262.40 to 39.97 tha^?1). Highest carbon storage (209.95 t ha^?1) was recorded in dry Shiwalik sal forest followed by Himalayan chir forest > chir pine plantation > lower Shiwalik pine forest > northern mixed dry deciduous forest. Maximum tree above ground biomass is observed in dry Shiwalik sal forests (301.78 t ha^?1), followed by upper Himalayan chir pine forests (194 t ha^?1) and lower in Shiwalik pine forests (138.73 t ha^?1). The relationship with stem volume showed the maximum adjusted r² (0.873), followed by total density (0.55) and average DBH (0.528). The regression equation of different parameters with shrub biomass showed highest r² (0.812) and relationship between ecosystem carbon with other parameters of different forest types, where cubic function with stem volume showed highest r² value of 0.873 through cubic functions. Our results suggest that biomass and carbon stocks in these subtropical forests vary greatly with forest type and species density. This variation among forests can be used as a tool for carbon credit claims under ongoing international conventions and protocols.     

Soil aggregate characteristics and stability of soil carbon stocks in a <Emphasis Type="Italic">Pinus tabulaeformis</Emphasis> plantation

Pinus tabulaeformis has been widely planted in order to conserve soil and water and improve the ecological environment in China. This study aimed to unravel how soil aggregates and soil carbon (C) stock stability of a P. tabulaeformis plantation change after 60 years of natural development and was performed in Vitex negundo var. heterophylla and Ziziphus jujuba var. spinosa shrub (shrub), a P. tabulaeformis forest (pine), and a coniferous broadleaf P. tabulaeformis mixed forest (pine-oak). Afforestation increased the stability of soil aggregates in the 0–10 cm soil layer but resulted in a decrease in the 10–20 cm soil layer. However, the presence of deciduous broadleaf species in the pine plantation improved the stability of soil aggregates. The total soil C stock was increased by afforestation, especially in the pine-oak forest, where it reached a significant level. The mineral soil C stock in the shrub stand was higher than that in pine and lower than that in pine-oak forests, but the C fractions had a different change. Afforestation increased the C fraction of macroaggregates in the 0–10 cm soil layer but decreased it in the 10–20 cm soil layer. This result suggested that afforestation could improve soil C stabilization in deeper soil. However, the pine-oak forest had a higher C fraction of macroaggregates than the pine forest in the 10–20 cm soil layer, indicating that soil C stabilization of the P. tabulaeformis plantation decreased when deciduous broadleaf species were present and thus formed the coniferous broadleaf mixed forest.     

Effects of silvicultural cleaning in mixed pine‐deciduous stands on moose damage to scots pine (Pinus sylvestris)

Silvicultural cleaning is commonly carried out in young stands to control competition between conifers and deciduous tree species. However, it has been questioned whether this practice affects moose (Alces alces L.) browsing on Scots pine (Pinus sylvestris L.) with respect to damage at the stand level. In this study, moose winter browsing was examined in relation to food availability in 19 Scots pine stands divided into silviculturally cleaned and non‐cleaned blocks. Cleaning had a negative effect on moose foraging in terms of reduced density, biomass availability, and height of the deciduous tree species. The total biomass consumed by moose was significantly lower on the cleaned blocks than on the non‐cleaned ones. There were no differences between the treatments in the amounts of pine consumed and in moose damage to pines. The density or biomass available of different deciduous tree species did not explain the variation in moose browsing on pine. In conclusion, timing of silvicultural cleaning with reference to moose browsing should be taken into account. If pines are not overtopped by deciduous trees, as in the present study, silvicultural cleaning is not necessary to prevent moose damage.     

Browsing of sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) in the context of life history strategies: a literature review

Sallow (Salix caprea L.) and rowan (Sorbus aucuparia L.) constitute small proportions of the deciduous tree volume in Scandinavia, but are highly preferred winter forage for moose and red deer, which occur at historically high densities. Thus, a possible decline of these tree species has been indicated. Against this background, we have reviewed the life histories of relevance for browsing, as well as the basic biology and genetics of sallow and rowan. The species show similarities with respect to short lifespan, small size and sympodial growth pattern, which are risk factors in a browsing context. They also have high juvenile growth rate, important for growing quickly out of reach of browsers. Sallow depends strongly on disturbance for establishment and is more demanding with respect to soil and light conditions than rowan, possibly important for the substantially lower abundance of sallow on the Norwegian Forest Inventory plots. Similarly, the relative recruitment of small size classes of sallow is less than for rowan. Although recruitment is reported to be hampered in wintering areas with high moose or red deer densities, the inventory data, however, dating only back to 1994, do not suggest a general decrease in any of the species. Sallow and rowan saplings show low mortality in moose and deer dominated areas and the species can be characterised as rather resilient to browsing. Of more concern is that browsing can constrain the development of mature rowan and sallow trees locally, with possible consequences for associated epiphytic biodiversity.     

Soil carbon stock assessment in the tropical dry deciduous forest of the Sathanur reserve forest of Eastern Ghats,India

Climate change and carbon mitigation through forest ecosystems are some of the important topics in global perspective. Tropical dry forests are one of the most widely distributed ecosystems in tropics, which remain neglected in research. The soil organic carbon (SOC) stock was quantified on a large scale (30 1-ha plots) in the dry deciduous forest of the Sathanur reserve forest of Eastern Ghats. The SOC stock ranged from 16.92 to 44.65 Mg/ha with a mean value of 28.26 ± 1.35 Mg/ha. SOC exhibited a negative trend with an increase in soil depth. A significant positive correlation was obtained between SOC stocks and vegetation characteristics viz. tree density, shrub basal area, and herb species richness, while a significant negative correlation was observed with bulk density. The variation in SOC stock among the plots obtained in the present study could be due to differences in tree abundance, herb species richness, shrub basal area, soil pH, soil bulk density, soil texture etc. The present study generates a large-scale baseline data of dry deciduous forest SOC stock, which would facilitate SOC stock assessment at the national level as well as to understand its contribution on a global scale.     

Habitat selection by ungulates in relation to predation risk by wolves and humans in the Białowieża Forest, Poland

Predation risk is an important factor influencing the distribution of ungulates and their impact on forest structure. However, simultaneous predation risk by wolves and humans is rarely considered in the analyses of habitat selection by ungulates. We counted ungulate pellets on transects to analyse the influence of wolves and humans on ungulate density distribution in the Białowieża Forest, Poland. We assessed whether (1) forest exploitation influenced ungulate habitat selection, (2) ungulate density was higher in areas without human hunting, (3) ungulates avoided the surroundings of a major road, (4) prey density was higher in the strife zone between home ranges of wolf packs both in the presence and absence of human hunting, (5) ungulates avoided areas selected by wolves, and (6) wolf kill sites were in high prey density areas. Red deer (Cervus elaphus) and wild boar (Sus scrofa) selected unexploited over exploited forests and areas without hunting, whereas roe deer (Capreolus capreolus) preferred exploited forests. Wild boar and European bison (Bison bonasus) avoided the area within 300 m of a major road, whereas we could not detect any avoidance by red or roe deer. Prey density was not higher in wolf strife zones, regardless of human hunting. Ungulates did not avoid areas selected by wolves. Wolves killed red deer in areas with prey density of about 4 red deer/km², regardless of whether the average red deer density in those areas was higher or lower. We conclude that habitat alteration by forest exploitation and hunting by humans influenced the density distribution of ungulates more than predation risk by wolves.     

Boom and bust of a moose population: a call for integrated forest management

There is increasing pressure to manage forests for multiple objectives, including ecosystem services and biodiversity, alongside timber production. However, few forests are currently co-managed for timber and wildlife, despite potential economic and conservation benefits. We present empirical data from a commercial Norway spruce (Picea abies) and Scots pine (Pinus sylvestris) production system in southern Norway in which moose (Alces alces) are an important secondary product. Combining long-term hunting and forestry records, we identified temporal variation in clear-felling over the past five decades, peaking in the 1970s. Herbicide treatment of regenerating stands and a fivefold increase in moose harvest has lead to a reduction in availability of successional forest per moose of >90 % since the 1960s. Field estimates showed that spraying with the herbicide glyphosate reduced forage availability by 60 and 96 % in summer and winter, respectively, 4 years after treatment. It also reduced moose use and habitat selection of young spruce stands compared with unsprayed stands. Together these lines of evidence suggest that forest management led to an increase in moose carrying capacity during the 1970s and a subsequent decline thereafter. This is likely to have contributed to observed reductions in moose population productivity in southern Norway and is counter to sustainable resource management. We therefore call for better integration and long-term planning between forestry and wildlife management to minimise forest damage and the development of large fluctuations in ungulate populations.     

基于MapInfo的塞罕坝地区植被景观空间结构分析

运用景观生态学原理,借助M apInfo技术,选取斑块密度、边缘密度、景观多样性、景观优势度等指标,分析塞罕坝地区不同地貌下景观类型的空间分布和空间格局特征,深入探讨了森林景观的林分类型、起源和分布特点,结果显示塞罕坝山地的斑块密度、边缘密度、破碎度,分别都高于沙地和曼甸,但山地的景观多样性低于曼甸和沙地。森林景观以落叶针叶林面积最大,为38 306.7 hm2,其次是阔叶林和常绿针叶林,分别为16 470.5hm2,9 455.5hm2,灌木林和混交林面积最小。曼甸以落叶针叶林景观优势度最高,且多为落叶松人工纯林;山地以落叶针叶林和阔叶林景观优势度较高,其中,阔叶林多为天然次生林,树种以桦树为主;沙地以落叶针叶林和常绿针叶林优势度较高,其常绿针叶林多为人工樟子松纯林。     

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.     

Variation in post-wildfire regeneration of boreal mixedwood forests: underlying factors and implications for natural disturbance-based management

To test the direct regeneration hypothesis and support natural disturbance-based forest management we characterized the structure and composition of boreal mixedwood forests regenerating after large wildfires and examined the influence of pre-fire stand composition and post-fire competing vegetation. In stands which had been deciduous (Populus sp.)-dominated, conifer (white spruce)-dominated, or mixed pre-fire we measured regeneration stocking (presence in 10 m² plots), density and height 10–20 years post-burn in five wildfires in Alberta, Canada. Most plots regenerated to the deciduous or mixed stocking types; plots in the older fire and in stands that were pure conifer pre-fire had higher amounts of conifer regeneration. Surprisingly, regeneration in pre-fire ‘pure’ white spruce stands was most often to pine, although these had not been recorded in the pre-fire inventory. Pre-fire deciduous stands were the most resilient in that poplar species dominated their post-fire regeneration in terms of stocking, density and height. These stands also had the highest diversity of regenerating tree species and the most unstocked plots. High grass cover negatively affected regeneration density of both deciduous and conifer trees. Our results demonstrate the natural occurrence of regeneration gaps, pre- to post-fire changes in forest composition, and high variation in post-fire regeneration composition. These should be taken into consideration when developing goals for post-harvest regeneration mimicking natural disturbance.     

Spatial heterogeneity of winter forest resources used by deer

Monitoring red deer (Cervus elaphus) and roe deer (Capreolus capreolus) populations is a key aspect of wildlife management in Europe, so the utilization of resources must be considered. However, the spatial distribution of resources used has often been overlooked. We studied spatial heterogeneity of winter forest resources used by deer and applied an indirect method to evaluate the browsing pressure of these cervids in a forest of the Cévennes National Park (Lozère, France). We analyzed the pattern of browsing by principal component analysis (PCA) and graphical display. We demonstrated that heather (Calluna vulgaris), bilberry (Vaccinium sp.) and beech (Fagus sylvatica), which figured in more than 50% of the browsing observed, were unevenly consumed in the study area. Moreover, the deer consumed more beech outside high beech forest. Our methodology provides a helpful tool to monitor browse–ungulate interaction by taking into account the availability of key browse resources.     

基于MaxEnt模型的白水河国家级保护区有蹄类保护动物生境适宜性评价

通过MaxEnt模型对白水河国家级自然保护区4种有蹄类保护物种进行生境适宜性评价,研究结果如下:(1)20次重复后斑羚、鬣羚、林麝、羚牛的训练集AUC值分别为0.962、0.947、0.924和0.945;(2)斑羚、鬣羚、林麝、羚牛得适宜栖息地分别占保护区总面积的14%、27%、25%、24%,次适宜栖息地占保护区总面积的27%、19%、35%、34%,不适宜栖息地占保护区总面积59%、54%、40%、42%;(3)对斑羚贡献最大的3个环境因子是坡向、坡度和落叶灌丛,对鬣羚贡献最大的3个环境因子是竹林、栽培竹林和落叶灌丛,对林麝贡献最大的3个环境因子是坡向、落叶灌丛和竹林,对羚牛贡献最大的3个环境因子是坡向、竹林以及落叶阔叶林。     

Species proportions by area in mixtures of Scots pine (<Emphasis Type="Italic">Pinus sylvestris</Emphasis> L.) and European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.)

Scots pine (Pinus sylvestris L.) and European beech (Fagus sylvatica L.) dominate many of the European forest stands. Also, mixtures of European beech and Scots pine more or less occur over all European countries, but have been scarcely investigated. The area occupied by each species is of high relevance, especially for growth evaluation and comparison of different species in mixed and monospecific stands. Thus, we studied different methods to describe species proportions and their definition as proportion by area. 25 triplets consisting of mixed and monospecific stands were established across Europe ranging from Lithuania to Spain in northern to southern direction and from Bulgaria to Belgium in eastern to western direction. On stand level, the conclusive method for estimating the species proportion as a fraction of the stand area relates the observed density (tree number or basal area) to its potential. This stand-level estimation makes use of the potential from comparable neighboring monospecific stands or from maximum density lines derived from other data, e.g. forest inventories or permanent observations plots. At tree level, the fraction of the stand area occupied by a species can be derived from the proportions of their crown projection area or of their leaf area. The estimates of the potentials obtained from neighboring monospecific stands, especially in older stands, were poorer than those from the maximum density line depending on the Martonne aridity index. Therefore, the stand-level method in combination with the Martonne aridity index for potential densities can be highly recommended. The species’ proportions estimated with this method are best approximated by the proportions of the species’ leaf areas. In forest practice, the most commonly applied method is an ocular estimation of the proportions by crown projection area. Even though the proportions of pine were calculated here by measuring crown projection areas in the field, we found this method to underestimate the proportion by 25% compared to the stand-level approach.     

Deer Browsing and Impact on Forest Development

Abstract

Browsing by large herbivores can cause serious costs in forest production. The most important costs related to browsing damage are insufficient stand regeneration, volume losses and timber quality impairment. In Sweden, moose (Alces alces)and roe deer (Capreolus capreolus) are the most abundant deer species and therefore cause the majority of browsing damage. Balancing costs and benefits related to deer density is difficult since few data about the final losses in timber volume or quality impairment is available. Simulations of browsing and the impact on stand growth suggest that a minor or moderate browsing intensity has little effect on final harvest volume. The reason is that trees seem to recover some of the lost growth as they grow out of reach of the animals. Balancing between different uses of the forestry resource is complicated by our limited capability to predict the impact of different management regimes.     

Effects of deer on the post-fire recovery of a Mediterranean plant community in Central Portugal

We studied the effect of dense populations of red and fallow deer on a burned Mediterranean plant community in Portugal. We sampled a set of 12 open and 12 fenced plots 2, 3, 4, and 8 years after fire occurrence. Each plot was sub-sampled using a quadrat grid to estimate the relative plant frequency (proportion of quadrats with plants). The effect of deer on temporal trends of overall plant frequency and diversity, and on the abundance of the four most common woody species, was modeled using Generalized Linear Mixed Models. The effect of deer on plant assemblage composition was assessed by use of Redundancy Analysis. In open plots the relative plant frequency increased along the study period from 0.59 to 0.85, whereas in fenced plots there was a significantly faster recovery from 0.35 to 0.96. Cistus salvifolius and Rubus ulmifolius were significantly favored by fencing but a similar effect was not observed for Erica scoparia and Ulex jussiaei. The Redundancy Analysis showed a divergent floristic composition of the two sets of plots. The effect of deer was significant for the Evenness index but not for the Shannon and the Richness indices. Our results show that a high deer density may significantly delay the post-fire recovery of a Mediterranean community of woody plants and cause shifts in the direction of plant succession.     

Ungulate browsing on conifers during summer as revealed by DNA

Young trees are particularly vulnerable to browsing by ungulates, since most, or all, of their canopies are within browsing height. In tree species with strong apical dominance, browsing may also affect the structural development of the tree, causing impaired timber quality. Across Sweden, ungulate communities comprise multiple coexisting species, making it a challenge to determine which herbivore species have caused the browsing damages. However, correct identification of the culprits is pivotal for the appropriate management actions to be undertaken. In central Sweden, forest plantation owners recently reported a perceived increase in browsing damages on conifers, primarily Scots pine (Pinus sylvestris). Concurrent with this increase in browsing damages, they noticed an increase in the population of red deer in the area, leading them to suspect that the red deer (Cervus elaphus) were the primary browsers responsible for the damages. Browsed twig samples were sent to us so that we could identify the responsible browsing species using DNA. We discovered that 73.7% of the damages were actually attributable to moose (Alces alces) with 24.7% caused by red deer and 1.5% by fallow deer (Cervus dama). DNA diagnostics can thus yield important information regarding multispecies assemblages enabling managers to correctly identify the relative contributions of different species to browsing damages.     

Stand Structure in Pine,Spruce and Deciduous Forests: A Comparison Between Finnish and Russian Karelia

This study investigated the stand structure in pine, spruce and deciduous forests in the border district of Finland and Russia. A total of 46 mature forest stands was selected as pairs, the members of each pair being as similar as possible with respect to their forest site type, age, moisture and topography. The stands were then compared between the two countries by means of basal areas and number of stems. The proportions of dominating tree species were 2-12% lower, and correspondingly the proportions of secondary tree species higher, in Russian forests. The density of the forest stock was also higher in each forest type in Russia. The forests in the two countries differed most radically in terms of the abundance of dead trees. The amount was two to four times higher in Russian deciduous and spruce forests, and in pine forests the difference was 10-fold. The stand structures indicated that Russian coniferous stands, in particular, were more heterogeneous than intensively managed pine and spruce stands in Finland.