Long-term responses of ecosystem components to stand thinning in young lodgepole pine forest: IV. Relative habitat use by mammalian herbivores

Pre-commercial thinning (PCT) is a silvicultural practice that can provide diverse understory and overstory vegetation conditions. We tested the hypothesis that relative habitat use by snowshoe hare (Lepus americanus), mule deer (Odocoileus hemionus), and moose (Alces alces) would increase in response to enhanced abundance of herbs and shrubs, and species diversity and structural diversity of conifers, in heavily thinned (≤1000 stems/ha) stands, at 12–15 years post-thinning. Replicate study areas were located near Penticton, Kamloops, and Prince George in south-central British Columbia, Canada. Each study area had three young pine stands thinned to densities of 500 stems/ha (low), 1000 stems/ha (medium), and 2000 stems/ha (high), with an unthinned young pine and old-growth pine stand for comparison.

Relative habitat use, based on counts of fecal pellets and pellet-groups, was similar among the five treatment stands for hares (P = 0.24), deer (P = 0.23), and moose (P = 0.16). However, low-density stands (500 stems/ha) had ca. 3–20 times as many deer pellet-groups, and ca. 2–4 times as many moose pellet-groups, than other stands. Low-density stands had significantly greater canopy openness, volume of shrubs <2 m, and horizontal hiding cover <1.6 m than other treatments. Relative habitat use by deer and moose was positively related to understory characteristics such as enhanced abundance of forage and security cover. These results support our hypothesis that deer and moose responded positively to enhanced volume of herbs and shrubs as well as to species diversity and structural diversity of conifers and overall vegetation in heavily thinned (≤1000 stems/ha) stands at 12–15 years post-thinning. Our results suggest that ungulate management would be enhanced if greater emphasis was placed on forage enhancement throughout the year, which differs from current management recommendations which tend to focus on winter range and snow-interception cover.     

Influence of conventional and chemical thinning on stand structure and diversity of plant and mammal communities in young lodgepole pine forest

Silvicultural practices that provide a wide variety of vegetative composition and structure (habitats) in young stands should help manage for biological diversity across forested landscapes. This study was designed to test the hypotheses that: (i) abundance and diversity of stand structure attributes (species diversity and structural diversity of herb, shrub and tree layers) and forest floor small mammal communities, and (ii) relative habitat use by large herbivores, will increase from unthinned to conventionally thinned to chemically thinned stands of young lodgepole pine (Pinus contorta) forest. Replicate study areas were located near Summerland, Kelowna and Williams Lake in south-central British Columbia, Canada. Each study area had three treatments: a conventionally thinned, a chemically thinned and an unthinned stand. Pre-commercial thinning was conducted in 1993. Coniferous stand structure and understory vegetation were measured prior to thinning in 1993 and 5 years later in 1998. Small mammal populations were sampled intensively from 1993 to 1998. Relative habitat use by large herbivores was sampled in 1998.

Our results indicate that chemical thinning of young lodgepole pine stands produced an aggregated pattern of crop trees compared with stands subjected to conventional thinning. Diameter growth of crop trees in the chemically thinned stands was similar to that in the conventionally thinned, but also to that in unthinned stands. Although horizontal stratification (aggregates of trees) was enhanced, vertical stratification (structural diversity of vegetation) was less in the chemically than conventionally thinned stands. Abundance and diversity of understory vegetation and small mammal communities were generally unaffected by stand thinning in these particular installations. Relative habitat use by mule deer (Odocoileus hemionus) occurred in a gradient from highest in the conventionally thinned stand to lowest in the unthinned stand. Habitat use by snowshoe hares (Lepus americanus) tended to have the opposite trend. Moose (Alces alces) exhibited no difference in habitat use among stands. Thus, although there were few differences among treatment stands, chemical thinning could be used to develop an aggregated pattern of crop trees in pre-commercially thinned stands to maintain habitat for herbivores such as snowshoe hares and mule deer. Understory plant and forest floor small mammal communities would be maintained in these stands as well.     

Influence of variable retention harvests on forest ecosystems: Plant and mammal responses up to 8 years post-harvest

Green-tree retention systems are an important management component of variable retention harvests in temperate zone coniferous forests. Residual live trees (“legacy trees”) provide mature forest habitat, increase structural diversity, and provide continuity in the regenerating stand. This study was designed to test the hypotheses that, at up to 8 years after harvest, abundance and species diversity of communities of (i) understory plants and (ii) forest-floor small mammals, and (iii) relative habitat use by mule deer (Odocoileus hemionus), will decline with decreasing levels of tree retention. Communities of plants and forest floor small mammals were sampled in replicated clearcut, single seed-tree, group seed-tree, patch cut, and uncut forest sites in mixed Douglas-fir (Pseudotsuga menziesii)—lodgepole pine (Pinus contorta) forest in southern British Columbia, Canada from 2000 to 2003 (5–8 years post-harvest). Habitat use by mule deer was measured during summer and winter periods each year from 1999 to 2003 in these same sites.     

Forage production in natural and afforested grasslands of the Pampas: ecological complementarity and management opportunities

In managed rangelands periods of low primary productivity determine troughs of forage availability, constraining animal production year-round. Although alternative tools to increase forage availability during critical seasons exists, most of them are unaffordable and short-lived in marginal areas. We explore the potential benefits of deciduous tree plantations favoring winter forage productivity by comparing aboveground net primary productivity (ANPP) patterns in herbaceous understory to tree plantations and natural grasslands in the Pampas (Argentina). These temperate subhumid grasslands are characterized by the coexistence of winter species, mainly C3 grasses of the native genera Stipa, Piptochaetium, and Bromus and the exotic genera Lolium and Festuca) and summer species (mainly C4 grasses of the native genera Paspalum, Bothriochloa, and Stenotaphrum) that replace each other throughout the seasons, with domination of the latter. We hypothesize that the natural decoupling of growing seasons between winter deciduous trees and winter grasses could provide the basis for the sustainable promotion of winter forage. We measured ANPP on two 23-year-old Populus deltoides plantations and their understory and compared them with adjacent open grasslands. Afforested stands had 55–75% higher annual ANPP than their non-afforested neighbors, with trees contributing ~70% to total ANPP. Herbaceous canopies beneath plantations achieved about half of the ANPP observed in non-afforested situations with a contrasting seasonal distribution associated with shifts from C4 to C3 grass dominance. Winter ANPP, the most critical source of forage in these grazing systems, was similar or higher in the herbaceous understory of tree plantations to that on their non-afforested counterparts, suggesting that mixed systems involving deciduous trees and understory pastures are a valid and viable option in the region.     

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.     

Mammal movements and foraging at remnant woodlands inside coniferous forest landscapes

Temporary or spatially restricted resources may affect population densities over more or less wide areas in the surrounding landscape or region. They may affect more than one trophic level by facilitation of predation. Areas of influence may differ between species. Such effects of mast-seeding in confined oak–hazel woodlands were examined for a guild of mainly seed-eating mammals and their predators. The mammals were tracked in snow in winter and the foraging of granivorous small mammals was assessed by experimental seed supplies in spring and autumn. Movements and foraging at various locations in and around the woodlands were distinguished from large-scale influences in the surrounding conifer forest landscape. Roe deer moved more abundantly inside the woodlands than in the conifer forest 50 m away and, less clearly, this was also the case for granivorous small mammals. Squirrels were particularly common at the edges while brown hares were somewhat less common at these edges. The edge effects appear therefore species-specific. Brown hares, squirrels, voles and granivorous small mammals were generally less common in the coniferous forest more than 500 m from the woodlands than in the coniferous forest 50 m from the woodlands. The most common generalist predator, the red fox, was as a mean equally common at all locations. However, it demonstrated a spatio-temporal variation in movements related to hare and, to a lesser extent, squirrel occurrence at open woodland sites. In order to sustain several mammalian species in conifer forest landscapes, not only granivorous ones, resource patches of oak and hazel should be retained, regenerated and, if possible, expanded. Such hot spots often need particular management.     

Wintering Habitat Evaluation of Red Deer (Cervus elaphus) in Forest Area of Northeastern China

IntroductionTheprofessionalsworkofwildlifemanagementisatargetforpopulationmanagement,butmoreandmoreprofessionalsworkersareinterestedinthemanagementofthewildlifehabitat(Belley,1984).ItisoneofessentiafworksforustoappraisehabitatwiththescientificmanagementontheReddeerpopulation.Soitisusefultoconservation,developmentandutilizationoftheReddeerinananimal'sperspective.Sincethe1970'smanyscientistshavebeenstudyingthehabitatevaluationfordeer.Severalmodelsandmethodssuchasthelineardiagram,thematrixmodel…     

Influence of fire and mountain pine beetle on the dynamics of lodgepole pine stands in British Columbia,Canada

An outbreak of the mountain pine beetle (Dendroctonus ponderosae Hopkins; MPB), currently affecting over 10.1 million hectares of lodgepole pine forests (Pinus contorta Dougl.) in British Columbia, Canada, is the largest in recorded history. We examined the dynamics of even-aged lodgepole pine forests in southern British Columbia, which were undergoing this MPB outbreak. Using dendroecology and forest measurements we reconstructed the stand processes of stand initiation, stand disturbances, tree mortality, and regeneration, and explained the current stand structure and the potential MPB impacts in selected stands. Our results indicate that stand-replacing fires initiated even-aged seral lodgepole pine stands in this region. In the absence of fire in the 20th century, multiple MPB disturbances, which each resulted in partial canopy removal, modified the simple one-layer structure of the fire-origin stands by the initiation of post-MPB disturbance regeneration layers, transforming the stands into complex, multi-aged stands. Despite high overstory mortality due to the current MPB outbreak, regeneration layers, which are likely to survive the current outbreak, will provide important ecological legacies and will contribute to mid-term timber supply.     

Wintering habitat evaluation of Red deer (Cervus elaphus) in forest area of Northeastern China

Six ultimate factors (forage abundance, horizontal sheltering class, distance from human disturbance, snow depth, tree coverage, and edge effect) of Red deer (Cervus elaphus) were investigated in eastern Heilongjiang Province during winter from 1988 to 1991. On the basis of analysis of field data, we evaluated winter habitat conditions of Red deer. The results showed as follows: the wintering habitat quality of Red deer in poplar-birch stands was excellent in the Wanda Mountains, and was moderate in shrub-woods and Korean pine seed stands. According to habitat index (HI) values, the habitat quality in poplar-birch stands (HI=0.8185) was superior to that in shrub-woods (HI=0.4825) and in Korean pine seed stands (HI=0.4385). The wintering habitat quality of Red deer in the Wanda Mountains was superior to that of Dailing Forestry Region. According to the current situations of forestry areas in northeastern China, strengthening habitat management was important for the conservation and management of Red deer population in the next decades. This subject is supported by the Natural Science Foundation of Heilongjiang Province. (Responsible Editor: Zhu Hong)     

Effects of local and regional landscape characteristics on wildlife distribution across managed forests

Understanding the impacts of local and regional landscape characteristics on spatial distributions of wildlife species is vital for achieving ecological and economic sustainability of forested landscapes. This understanding is important because wildlife species such as white-tailed deer (Odocoileus virginianus) have the potential to affect forest dynamics differently across space. Here, we quantify the effects of local and regional landscape characteristics on the spatial distribution of white-tailed deer, produce maps of estimated deer density using these quantified relationships, provide measures of uncertainty for these maps to aid interpretation, and show how this information can be used to guide co-management of deer and forests. Specifically, we use ordinary least squares and Bayesian regression methods to model the spatial distribution of white-tailed deer in northern hardwood stands during the winter in the managed hardwood-conifer forests of the central Upper Peninsula of Michigan, USA. Our results show that deer density is higher nearer lowland conifer stands and in areas where northern hardwood trees have small mean diameter-at-breast-height. Other factors related with deer density include mean northern hardwood basal area (negative relationship), proportion of lowland conifer forest cover (positive relationship), and mean daily snow depth (negative relationship). The modeling methods we present provide a means to identify locations in forest landscapes where wildlife and forest managers may most effectively co-ordinate their actions.     

Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon

Relationships between tree height and crown dimensions and trunk diameter were determined for shade-tolerant species of old-growth forests of western Oregon. The study included both understory and overstory species, deciduous and evergreen angiosperms and evergreen conifers. A comparison of adult understory species with sapling overstory species of similar height showed greater crown width and trunk diameter in the former, whether the comparison is made among conifers or deciduous trees. Conifer saplings had wider crowns than deciduous saplings, but the crown widths of the two groups converged with increase in tree height. Conifer saplings had thicker trunks than deciduous saplings of similar crown width, possibly because of selection for resistance to stem bending under snow loads. The results suggest that understory species have morphologies that increase light interception and persistence in the understory, whereas overstory species allocate their biomass for efficient height growth, thereby attaining the high-light environment of the canopy. The greater crown widths and the additional strength requirements imposed by snow loads on conifer saplings result in less height growth per biomass increment in conifer saplings than in deciduous saplings. However, the convergence in crown width of the two groups at heights greater than 20 m, and the proportionately smaller effect of snow loads on large trees, may result in older conifers equalling or surpassing deciduous trees in biomass allocation to height growth.     

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。     

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.     

Overstory–understory biomass changes over a 35-year period in southcentral Oregon

Forest canopy development is known to influence understory biomass relationships. An 85,268 ha area in southcentral Oregon was examined for changes in overstory canopy closure using 1953 and 1988 aerial and satellite imagery and a geographic information system. A negative exponential curve predicted a loss of approximately half of understory biomass over the 35-year interval. Reductions in understory biomass were most pronounced at higher elevations where growing conditions for conifers and canopy closure changes were most pronounced. The loss of understory biomass was related to declines in mule deer (Odocoileus hemionus) populations in the area.     

The ecology of mixed severity fire regimes in Washington, Oregon, and Northern California

Forests characterized by mixed-severity fires occupy a broad moisture gradient between lower elevation forests typified by low-severity fires and higher elevation forests in which high-severity, stand replacing fires are the norm. Mixed-severity forest types are poorly documented and little understood but likely occupy significant areas in the western United States. By definition, mixed-severity types have high beta diversity at meso-scales, encompassing patches of both high and low severity and gradients in between. Studies of mixed-severity types reveal complex landscapes in which patch sizes follow a power law distribution with many small and few large patches. Forest types characterized by mixed severity can be classified according to the modal proportion of high to low severity patches, which increases from relatively dry to relatively mesic site conditions. Mixed-severity regimes are produced by interactions between top-down forcing by climate and bottom-up shaping by topography and the flammability of vegetation, although specific effects may vary widely across the region, especially the relation between aspect and fire severity. History is important in shaping fire behavior in mixed-severity landscapes, as patterns laid down by previous fires can play a significant role in shaping future fires. Like low-severity forests in the western United States, many dry mixed-severity types experienced significant increases in stand density during the 20th century, threatening forest health and biodiversity, however not all understory development in mixed-severity forests increases the threat of severe wild fires. In general, current landscapes have been homogenized, reducing beta diversity and increasing the probability of large fires and insect outbreaks. Further loss of old, fire tolerant trees is of particular concern, but understory diversity has been reduced as well. High stand densities on relatively dry sites increase water use and therefore susceptibility to drought and insect outbreaks, exacerbating a trend of increasing regional drying. The need to restore beta diversity while protecting habitat for closed-forest specialists such as the northern spotted owl call for landscape-level approaches to ecological restoration.     

Change in historic fire disturbance in a Garry oak (Quercus garryana) meadow and Douglas-fir (Pseudotsuga menziesii) mosaic,University of Victoria,British Columbia,Canada: A possible link with First Nations and Europeans

We investigated late Holocene vegetation and fire changes on southeast Vancouver Island, British Columbia, through high-resolution analysis of pollen, spores and charcoal contained within a forest soil. Located in the Mystic Vale Endowment lands, University of Victoria campus, the site occurs adjacent to Garry oak (Quercus garryana) meadows within a Douglas-fir (Pseudotsuga menziesii) forest regionally influenced by First Nations burning. The age of the sequence is estimated from the occurrence of non-native plantain (Plantago spp.) pollen, between the early 19th century and the present. Prior to the 1850s, the Mystic Vale supported a Douglas-fir forest with a sword fern (Polystichum munitum) understory. Low-intensity fires may have burned either at or near the site, but did not destroy the stand. From the time of European settlement (ca. 1850s to 1872) to the early 20th century, Douglas-fir forests with a sword fern understory persisted but with more openings than in the past. Adjacent forests were cleared leading to successional stands of red alder (Alnus rubra). The site may have supported more grasses near or after 1872 because of adjacent cultivation and the introduction of livestock. Local establishment of Garry oaks at the margin of a field adjacent to the deposition site may have occurred at this time. The strong charcoal signal may be attributable to clearing and local fires of European settlers. Exotic weedy species such as plantain arrived during this time. From about the mid 20th century to the present a closed canopy Douglas-fir forest returned. Pre-European charcoal concentrations were significantly greater than modern-day charcoal concentrations. Peak charcoal concentrations coincided with the onset of European settlement in the area, a period in which land-clearing is thought to have been aided by the use of fire. This study suggests that the observed changes in charcoal concentrations in pre- and post-European contact time may be linked with a switch from aboriginal to European ignition sources. The study provides a historical impetus to consider partially restoring parts of southeast Vancouver Island by re-introducing fire via the application of controlled burns.     

A COMPARISON OF RAINFALL IN DIFFERENT WOODLANDS

At Bedgebury the rainfall in thirteen forest plots planted withdifferent tree species has been compared with that in the open.In all these forest plots the trees have closed canopy within20 years of planting and the canopies differ considerably intheir structure. From 6 to 93 per cent. of the gross rainfallmay be retained on the tree canopies and lost to the soil. Theproportion of precipitation intercepted is greatest in lightrains and least in heavy rains. During light showers considerablymore moisture is retained on the conifer than on the hardwoodcanopies. In the heavier showers rainwater flows down the stems,but stem flow represents a small proportion of the total waterreaching the forest floor. The amount of water which reachesthe ground along the trunks varies with the species and thecharacter of branching. Some raindrops penetrate directly throughthe canopy, but many are intercepted by the canopies and coalesceto fall as large drops on the forest floor. These large waterdropstend to fall continuously on the same spot so that in the plantationsthe water is distributed unevenly. In the winter months, whenthe deciduous species have shed their leaves, snow penetratesmore easily through their canopies than those of the evergreenconifers.     

Effect of snow cover on the distribution of non-migratory sika deer on the top of Mt. Tanzawa,Japan

On the basis of 268 direct sika deer observations and 212 ratio locations of three individuals, the influence of snow on deer distribution was studied. During the period of snow absence or its shallow cover, the deer did not show any preferences towards slope aspect. However, when the snow coverage became deeper, the deer used westerly exposures (having shallower snow cover) more intensively. Later on, when the snow started to melt on the easterly slope aspects deer mostly used those exposures due to relatively abundant food resources.     

Influence of fire intensity on structure and composition of jack pine stands in the boreal forest of Quebec: Live trees,understory vegetation and dead wood dynamics

North American jack pine (Pinus banksiana Lamb.) stands are generally characterized by an even-aged structure resulting from high intensity fires (HIF). However, non-lethal fires of moderate intensity (MIF), which leave behind surviving trees, have also been reported. The objectives of this study were two-fold: (1) assess the concurrent dynamics of live trees, understory vegetation and different types of coarse woody debris (CWD) during succession after HIF; and (2) document how MIF affects stand structure component dynamics compared to HIF. Stands affected by both HIF and MIF were selected. Tree characteristics and age structure, understory biomass, and CWD volume were assessed. Our results suggest that the structural succession of jack pine stands following HIF comprises three stages: young stands (<48 years), premature and mature stands (58–100 years) and old stands (>118 years). Canopy openness and jack pine density significantly decreased with time since HIF, while black spruce density and CWD volume significantly increased. The highest structural diversity was measured in the premature and mature stands. Compared to HIF, MIF increased mean jack pine basal area, decreased average stand density, delayed the replacement of jack pine by black spruce replacement in the canopy, decreased CWD volume, and significantly increased bryophytes mass. MIF increased the diversity of live trees and generally decreased CWD structural diversity. The study confirms the diversity of natural disturbance magnitude and successional processes thereby initiated. Thereafter, it appeared to be relevant for adjustment of disturbance emulating forest-management systems.