Forest herbs show species-specific responses to variation in light regime on sites with contrasting soil acidity: An experiment mimicking forest conversion scenarios

Forest conversion from native deciduous forests to coniferous stands has been performed in many European regions and resulted in dramatic shifts in understorey plant community composition. However, the drivers for changes in specific understorey plant species remained unclear.Here, we experimentally determine the species-specific effects of light availability and chemical soil characteristics, on the vegetative and regenerative performance of five herbaceous forest understorey plants. Topsoil samples from both spruce and deciduous stands at four locations, with two levels of soil acidity, were collected and used in a common garden experiment. Additionally, three different light levels were applied, i.e., ‘light deciduous’, ‘dark deciduous’ (extra light reduction during summer) and ‘evergreen’ (light reduction during winter). In a second experiment we evaluated the germination of two of these species against the acidity and tree species at the site of origin of the soil samples.The light regime affected both the vegetative and regenerative performance of the understorey species: compared to light deciduous, Anemone nemorosa had a significantly lower performance under the evergreen light regime, Convallaria majalis under dark deciduous and Luzula luzuloides and Galium odoratum under both light regimes. The vegetative performance was lower in soil from acid sites for the acid-sensitive species G. odoratum and Primula elatior. Differences between the soils sampled under deciduous or spruce stands had no effect on the vegetative, or the regenerative performance of these species. By contrast, the germination of L. luzuloides and P. elatior was higher in soils sampled in deciduous stands and in neutral sites.Species-specific responses in vegetative and regenerative performance of adult plants to a changed light regime and soil acidification could be a reason for the changed vegetation composition in converted stands. Also lower germination and establishment of forest understorey species in spruce stands could influence the species distribution after conversion.     

The species richness–productivity relationship in the herb layer of European deciduous forests

Aim In contrast to non‐forest vegetation, the species richness–productivity (SR‐P) relationship in forests still remains insufficiently explored. Several studies have focused on the diversity of the tree layer, but the species richness of temperate deciduous forests is mainly determined by their species‐rich herb layer. The factors controlling herb‐layer productivity may differ from those affecting tree layers or open herbaceous vegetation, and thus the SR‐P relationship and its underlying processes may differ. However, the few relevant studies have reported controversial results. Here we explore the SR‐P relationship in the forest herb layer across different areas from oceanic to continental Europe, and put the effect of habitat productivity on species richness into context with other key factors, namely soil pH and light availability. Location North‐western Germany, Czech Republic, Slovakia and southern Urals (Russia). Methods We measured herb‐layer species richness and biomass, soil pH and tree‐layer cover in 156 vegetation plots of 100 m² in deciduous forests. We analysed the SR‐P relationship and the relative importance of environmental variables using regression models for particular areas and separate forest types. Results We found a consistent monotonic increase in the herb‐layer species richness with productivity across all study areas and all forest types. Soil pH and light availability also affected species richness, but their relative importance differed among areas. Main conclusions We suggest that the monotonically increasing SR‐P relationship in the forest herb layer results from the fact that herb‐layer productivity is limited by canopy shading; competition within the herb layer is therefore not strong enough to exclude many species. This differs fundamentally from open herbaceous vegetation, which is not subject to such productivity limits and consequently exhibits a unimodal SR‐P relationship. We present a conceptual model that might explain the differences in the SR‐P relationship between the forest herb layer and open herbaceous vegetation.     

Selection harvest in temperate deciduous forests: impact on herb layer richness and composition

Herb layer richness and composition of four forest types and three different management treatments were investigated in 16 deciduous forest stands of northern Germany. Specifically, we compared the species richness and composition occurring in mature forest stands that were single-tree and group selection harvested to those in unmanaged reference stands. Mean species richness of all herb layer species increased significantly with increasing harvest severity. When analyzing plant groups separately, it became obvious that this overall pattern was not consistent. While a negative relationship was detected between vernal herb richness and harvest severity, group selection harvest significantly increased species richness of summer herbaceous forest species and generalists. Woody species richness was not related to harvest severity. Community composition of the spring aspect was not significantly affected by selection harvest, whereas herb layer species composition in the summer aspect differed significantly among the three harvest treatments. A dominance of highly competitive shrub and generalist species was confined to some parts of the most intensively harvested stands. Overall, our results indicate that the herb layer community was not severely adversely affected by selection harvest at the intensities used in the studied stands. It is suggested that selection harvest systems may be feasible tools with which to conserve local forest vascular plant diversity and at the same time to meet the demand for timber products. However, information about forest history and the implementation of the selection harvest system are basic requirements when interpreting the results of studies on understorey response to selection harvest.     

Response of herbaceous layer species to canopy and soil variables in a central Appalachian hardwood forest ecosystem

Previous work has suggested that excess nitrogen (N) alters the degree to which forest canopy versus soil variables influence forest herb communities. This study tested the hypothesis that excess N would shift this influence on individual herb species from soil N availability to stand structural variables that determine light availability to the forest floor. Two watersheds at the Fernow Experimental Forest, West Virginia, USA were used: WS4 and WS3 as untreated reference and treatment watersheds, respectively. WS3 receives 35 kg N/ha/year via aerial application. Herb cover and composition was measured in seven permanent plots/WS from 1991, currently on-going. In 2011, soil moisture and N availability were measured in each plot, along with several variables of canopy structure. Backwards stepwise regression was used to determine relationships between herb cover/individual species and soil/canopy measurements. Herb cover varied spatially with soil resources on WS4, whereas cover varied spatially with canopy structure on WS3. Although results for total herb layer cover supported this hypothesis, results for individual herb species rejected it. This contrast was especially evident for Rubus allegheniensis (blackberry), a nitrophilic species which increased with increasing soil N on both watersheds, but was not correlated with canopy structure on reference WS4, while being correlated with canopy structure on N-treated WS3. Excess N from atmospheric deposition has been shown to decrease plant biodiversity of impacted forests, especially in its effects on herbaceous layer communities. This work demonstrates that one of the mechanisms of such response is in N-mediated changes in the response of herb communities to soil resources and light availability.

     

Traits of winner and loser species indicate drivers of herb layer changes over two decades in forests of NW Germany

Questions: What are the most likely environmental drivers for compositional herb layer changes as indicated by trait differences between winner and loser species? Location: Weser‐Elbe region (NW Germany). Methods: We resurveyed the herb layer communities of ancient forest patches on base‐rich sites of 175 semi‐permanent plots. Species traits were tested for their ability to discriminate between winner and loser species using logistic regression analyses and deviance partitioning. Results: Of 115 species tested, 31 were identified as winner species and 30 as loser species. Winner species had higher seed longevity, flowered later in the season and more often had an oceanic distribution compared to loser species. Loser species tended to have a higher specific leaf area, were more susceptible to deer browsing and had a performance optimum at higher soil pH compared to winner species. The loser species also represented several ancient forest and threatened species. Deviance partitioning indicated that local drivers (i.e. disturbance due to forest management) were primarily responsible for the species shifts, while regional drivers (i.e. browsing pressure and acidification from atmospheric deposition) and global drivers (i.e. climate warming) had moderate effects. There was no evidence that canopy closure, drainage or eutrophication contributed to herb layer changes. Conclusions: The relative importance of the different drivers as indicated by the winner and loser species differs from that found in previous long‐term studies. Relating species traits to species performance is a valuable tool that provides insight into the environmental drivers that are most likely responsible for herb layer changes.     

Distribution and abundance of trees in floodplain forests of the Wisconsin River: Environmental influences at different scales

Questions: 1. How do physiography, flooding regime, landscape pattern, land‐cover history, and local soil conditions influence the presence, community structure and abundance of overstorey trees? 2. Can broad‐scale factors explain variation in the floodplain forest community, or are locally measured soil conditions necessary? Location: Floodplain of the lower 370 km of the Wisconsin River, Wisconsin, USA. Methods: Floodplain forest was sampled in 10 m × 20 m plots [n= 405) during summers of 1999 and 2000 in six 12‐ to 15‐km reaches. Results: Species observed most frequently were Fraxinus pennsylvanica, Acer saccharinum and Ulmus americana. Physiography (e.g. geographic province) and indicators of flooding regime (e.g. relative elevation and distance from main channel) were consistently important in predicting occurrence, community composition, and abundance of trees. Correspondence analysis revealed that flood‐tolerant and intolerant species segregated along the primary axis, and late‐successional species segregated from flood‐tolerant species along the secondary axis. Current landscape configuration only influenced species presence or abundance in forests that developed during recent decades. Land‐cover history was important for tree species presence and for the abundance of late‐successional species. Comparison of statistical models developed with and without soils data suggested that broad‐scale factors such as geographic province generally performed well. Conclusions: Physiography and indicators of flood regime are particularly useful for explaining floodplain forest structure and composition in floodplains with a relatively high proportion of natural cover types.     

Multiple environmental changes drive forest floor vegetation in a temperate mountain forest

Human‐induced changes of the environment and their possible impacts on temperate forest understory plant communities have been examined in many studies. However, the relative contribution of individual environmental factors to these changes in the herb layer is still unclear. In this study, we used vegetation survey data covering a time period of 21 years and collected from 143 permanent plots in the Northern Limestone Alps, Austria. Data on soil chemistry (49 plots), light condition (51 plots), soil temperature and moisture (four and six plots), disturbance (all plots), climate (one station in a clearing area), and airborne sulfur (S) and nitrogen (N) deposition (two forest stands) were available for analyses. We used these data together with plot mean Ellenberg indicator values in a path analysis to attribute their relative contributions to observed vegetation changes. Our analysis reveals a strong directional shift of the forest understory plant community. We found strong evidence for a recovery of the ground‐layer vegetation from acidification as response to decreased S deposition. We did not observe a community response to atmospheric N deposition, but we found a response to altered climatic conditions (thermophilization and drying). The path analysis revealed that changes in the light regime, which were related to small‐scale disturbances, had most influence on herb layer community shifts. Thermophilization and drying were identified as drivers of understory community changes independent of disturbance events.     

Vegetation changes in forests of the Krkonoše Mts. over a period of air pollution stress (1980–1995)

Species composition of the understory of spruce (Picea abies (L.) Karsten) and beech (Fagus sylvatica L.) forests subjected to intense air pollution stress in the Krkonoe Mts., Czech Republic, showed directional changes over the last 15 years. The changes were documented with repeated observations of 29 permanent plots, initiated in 1980 and analysed with constrained ordination methods (Canonical Correspondence Analysis). Ground layer changes were mainly associated with the loss of canopy foliage from air pollution stress. The increased amount of the light at ground level resulted in increased herb cover. The initially dominant species (e.g., Calamagrostis villosa (Chaix) J. F. Gmelin, Deschampsia flexuosa (L.) P. B., Vaccinium myrtillus L., Athyrium distentifolium Tausch) were those that increased in cover. Moss cover, and moss and herb richness, declined over the course of the study. Thus, the changes in tree canopy are accompanied with changes in the forest understory, which in turn can affect the dynamics of these forests.     

How forest edge-center transitions in the herb layer interact with beech dominance versus tree diversity

Aims Forest fragmentation and the associated augmentation of forest edge zones are increasing worldwide. Forest edges are characterized by altered plant species richness and community composition. As the tree layer and its species composition has been shown to influence herb layer composition, changes in tree species composition or richness may weaken or strengthen edge effects in forest ecosystems. We studied effects of the edge–center transition, tree species composition and their potential interaction on the understory vegetation in the Hainich National Park, Germany's largest connected deciduous forest, allowing to cover large edge–center transects.Methods We established 12 transects in an area of 75 km 2 of continuous forest, 6 beech-dominated and 6 in multispecies forest stands. Each transect reached from the forest edge up to 500 m into the forest interior. Vegetation relevés were conducted in regular, logarithmic distances along each transect.Important findings Herb species richness was influenced by an interaction of edge effects and tree diversity level. With increasing distance from the forest edge, herb species richness remained constant in multispecies forest stands but rapidly decreased in beech-dominated forest stands. Further, herb richness was higher in the interior of multispecies forest stands. Percent forest specialists increased and percent generalists decreased with distance from the edge and this contrasting pattern was much more pronounced in beech-dominated transects. By using structural equation modeling, we identified litter depth mediated by tree species composition as the most important driver of herb layer plant species richness.     

Disturbance of the herbaceous layer after invasion of an eutrophic temperate forest by wild boar

Disturbances of the soil and the tree canopy are crucial factors determining the diversity, composition and biomass of the herbaceous layer in forests. This study presents a detailed account of ground vegetation in permanent plots surveyed before and after invasion of wild boar (Sus scrofa) to a temperate deciduous broadleaf forest. Specifically, we aimed to quantify the effect of wild boar rooting on cover, richness and composition of spring ephemerals, summer green herbs and saplings of woody species in relation to tree canopy cover. Rooting frequency in sample plots increased from 0% in 2010 to 61% in 2013. In heavily rooted plots, the mean cover of spring ephemeral geophytes (mainly Anemone nemorosa, A. ranunculoides and Ranunculus ficaria) decreased from 75% to 39% between 2010 and 2013. Species richness of summer green herbs generally increased between 2010 and 2013 and was additionally positively affected by heavy rooting and low canopy cover. Rooting also caused heterogenization of the herbaceous layer and amplified ongoing compositional changes induced by changing light conditions. Frequency and richness of spring ephemeral and woody species remained unchanged. We conclude that overall species richness of the herbaceous layer may increase in the short‐term as a result of increased plant recruitment and seed dispersal. However, wild boar rooting can greatly reduce the ground cover of spring ephemerals in eutrophic broadleaf forests, thereby threatening their important ecological function. To avoid long‐term losses of characteristic spring flora elements, local population control of wild boar is necessary to reduce abundance and frequency of soil rooting.     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.     

Influence of ungulates on the vegetation composition and diversity of mixed deciduous and coniferous mountain forest in Austria

The often highly elevated stocks of ungulates (red and roe deer and chamois) in the Alps shape the composition of the woody vegetation. The aim of this study was to investigate the influence of ungulates on the mixed deciduous and coniferous mountain forest in the district of Reutte, which boasts the highest density of ungulates in Tyrol (Austria), with a special focus on the effect of browsing by ungulates on plant diversity of the herb layer, different shrub layers. and the tree layer. Our results showed that within the fenced ungulate exclosures, (1) the composition of trees shifted towards fir (Abies alba) and various deciduous trees, whereas outside the fences, spruce became the dominant species; (2) the cover of dwarf shrubs and upper and lower shrub layers (1.3–5.0 and 0.5–1.3 m, respectively) increased significantly; (3) the cover of grasses decreased significantly and (4) the diversity decreased as an increase in the diversity of the tree and shrub layer was overcompensated by a significant decrease in the diversity of the undergrowth vegetation. Browsing by ungulates benefited grass species in the understory and altered the relative abundance of tree species in the lower layer which could, over time, result in compositional shifts in the canopy.     

桂西地区南亚热带落叶栎林群落组成及物种多样性研究北大核心CSCD

落叶栎林是桂西地区南亚热带的典型次生林,也是该区域落叶阔叶林的重要群系组。该研究采用典型样方法对桂西地区落叶栎林群落进行调查,分析了该区域落叶栎林群落的物种组成、区系成分、物种多样性特征及其与地形因子的关系,为桂西地区南亚热带植物多样性保护与恢复提供依据。结果显示:(1)研究区落叶栎林群落维管束植物共计269种,隶属80科178属。(2)种子植物区系以热带成分为主,同时表现出一定程度的温带过渡性质。(3)聚类分析表明,调查的落叶栎林群落可分为云南波罗栎林、栓皮栎林、白栎林、麻栎林4种林分类型,其中以白栎林群落的物种多样性最高,且灌木层的物种多样性显著高于乔木层和草本层。(4)RDA分析显示,落叶栎林群落不同层次物种多样性的差异受地形因子的影响,多样性指标与经度、纬度、海拔之间具有明显的相关性(P<0.05),其中乔木层物种多样性主要与经度、纬度呈显著的相关性,灌木层物种多样性与纬度、海拔相关,草本层物种多样性与经度、纬度、海拔之间均有相关性。     

浙江天童常绿阔叶林地被层的研究

 对天童常绿阔叶林地被层植物种类组成及群落学作用进行了初步研究。研究表明：组成地被层计有维管植物44科78属114种。蕨类植物优势度最大,里白占绝对优势。39种草木植物,以世界分布科和地面芽植物为主,常绿种类占84.6％,未见亚洲和北美洲间断分布的地理成分。乔灌木幼苗51种,80%的苗量集中于4～6个种中,优势种突出。藤本植物24种,草质藤本占25％,落叶种类42％。地面苔藓植物(22种)与枯枝落叶构成镶嵌分布,发育较成熟的群落中苔藓种类多,盖度大。里白及其枯死物形成层层交错重叠的地被层,阻碍凋落物进入地表,减弱光照,使近地表光照长期低于幼苗光补偿点,抑制了乔灌木优势种的更新。凋落物阻碍种子下移至半分解层,栲树种子下移率为17％至22.3％,冬季种子的动物损耗率平均为98.2％,凋落物中种子的平均腐烂率为28.2％,表层保存下来的栲树种子因缺水萌发速度慢且成活率低,下层萌发速度虽快,但腐烂率高,栲树天然更新幼苗主要来源于草窗。     

临安次生灌丛植物多样性对林火烈度空间异质性的响应

以同一过火3a后临安市太阳镇天然次生灌丛为对象,采用样地调查法按不同林火烈度设置火干扰样地进行植物群落调查,以检验林分内部的林火烈度异质性是否与局部的植物多样性变化相关。结果表明:研究区共有高等植物83种,分属于38科67属,群落区系组成以亚热带科属为主,表现出常绿阔叶林已退化过渡到位于演替早期阶段的落叶次生灌丛群落的性质;低林火烈度对灌木层的树种组成有影响,但不明显;中烈度林火对灌木层的物种组成影响较大;低、中林火烈度下草本层的物种组成变化都很明显;灌木层的物种数和多样性指数都表现出低烈度火未火烧中烈度火的趋势;草本层的物种数、多样性指数和均匀度指数表现出中烈度火低烈度火未火烧的趋势;草本层的物种组成和多样性受林火烈度的影响较灌木层更大。研究表明次生灌丛群落过火区内部林火烈度异质性在初期会引起植物多样性的响应差异;低烈度火干扰可以增加次生灌丛生物多样性、促进群落更新;中烈度火干扰下木本植物物种多样性丧失较大,而草本植物多样性显著增加,不利于群落的正向演替。     

川西亚高山针叶林土壤动物群落对模拟林下植物丧失的响应

受研究手段的限制,有关森林物种组成对土壤动物群落影响的研究少有报道。采用人工除灌和除草的林地控制实验方法,研究了亚高山人工林灌草层关键物种丧失对土壤动物群落结构的影响。结果表明:(1)灌木层去除后土壤动物密度极显著低于对照(P0.01);线虫纲(Nematoda)优势度持续增加,处理15个月时极显著高于对照(P0.01);土壤动物群落多样性指数极显著低于对照(P0.01);腐食性功能团类群数及个体数百分比有所下降。(2)草本层去除后土壤动物密度显著低于对照(P0.05);线虫纲优势度持续增加,处理15个月时显著高于对照(P0.05);土壤动物群落多样性指数极显著低于对照(P0.01);腐食性功能团类群数略低于对照,个体数百分比无显著差异。综上所述,林下灌草层去除,尤其是灌木层去除,导致土壤动物群落个体数量、多样性指数降低,优势类群格局、腐食性功能团构成发生变化,从而在一定程度上影响到森林生态系统的物质循环功能。     

Leaf litter fall and soil acidity during half a century of secondary succession in a temperate deciduous forest

Vegetation, leaf litter fall and soil pH were sampled repeatedly within semipermanent plots in a South-Swedish deciduous forest, 1935–1983. Leaf litter fall was summarized in a litter quality index. Vegetation types were differentiated along similar gradients in soil pH and leaf litter quality. The greatest shifts in dominance among field layer species were found in those plots where the quality of the leaf litter had improved. These plots also showed a halt in the general tendency towards a decreasing pH in the top soil.     

The impact of grazing pressure in clearfelled,burned and undisturbed eucalypt forest

An area of dry eucalypt forest on dolerite in southeastern Tasmania with comparable sites that were a) unlogged, b) clearfelled, c) clearfelled and slash-burned, was monitored over a 24 month period. Detailed records were made of plant species composition and cover on floristically similar fenced and unfenced permanent plots in two major vegetation types. Mammal faecal pellets were cleared from each of the unfenced plots and subsequently counted and attributed to species of origin. For the six combinations of treatment and vegetation type pellet deposition rate was highly correlated with differences in the rate of change of species cover between fenced and unfenced plots and the lifeform composition of the vegetation. The greatest changes were observed in the clearfelled, burned area, while there was relatively little change in the undisturbed forest. Grass and herb species were generally disadvantaged by mammal exclusion while shrubs and graminoids generally grew faster in the fenced plots.Nomenclature of plant species follows Curtis & Morris (1975).This project was supported financially by the Forest Ecology Research Fund, Tasmania.     

Distribution and effects of tree leaf litter on vegetation composition and biomass in a forest-grassland ecotone

Aims After abandonment of grasslands, secondary succession leads to the invasion by woody species. This process begins with the accumulation of tree litter in the forest–grassland ecotone. Our objectives were to determine the relationships between litter amounts and vegetation composition and cover along natural forest–grassland ecotones and to experimentally study the initial effects of tree litter accumulation on grassland vegetation and on microsite conditions.Methods We established 11 transects varying from 12 to 15 m in length in different forest–grassland ecotones in the Lahn-Dill highlands, Germany, and measured the mass and cover of tree litter and the cover and composition of vegetation at five sequential positions along each transect by using 1 m 2 plots with five replications. In a field experiment, we established plots subjected to different litter amounts (0, 200 and 600g m ?2) and evaluated changes in grassland vegetation, soil temperature and soil nutrient availability below the litter layer.Important findings Tree litter amounts decrease from 650 to 65g m ?2 across the forest–grassland ecotone. Vegetation changed from shrubs and annual species (adapted to more stressful conditions) in the forests edge to grasses, rosettes and hemirosette species (with higher competitive abilities) in the grassland. These anthropogenic forest–grassland ecotones showed abrupt edges, and the two adjacent ecosystems were characterized by different species pools and functional groups. In the field experiment, the presence of a litter layer reduced vegetation biomass and cover; the species richness was only reduced in the treatment with high litter (600g m ?2). Additionally, adding litter on top of vegetation also reduced thermal amplitude and the number of frost days, while increasing the availability of some nutrients, such as nitrogen and aluminium, the latter being an indicator of soil acidification. Adding a tree litter layer of 600g m ?2 in grassland areas had strong effects on the composition and diversity of grassland vegetation by reducing the cover of several key grassland species. In, or near, forest edges, litter accumulation rapidly changes established vegetation, microsite conditions and soil nutrients.     

Effects of grazing and cultivation on forest plant communities in Mount Elgon National Park, Uganda

Plant communities in the montane forest of Mount Elgon National Park were studied in order to assess the impact of grazing and cultivation on species composition. Present and former land uses, tree, shrub and herb species, soil properties and the percentage cover and height of trees, shrubs and herbs were determined in 40 plots. An indirect ordination of these plots showed that species composition was primarily determined by successional stage and agricultural disturbance. In forest plots (ordinated separately) where the widest range of former and current grazing intensities had occurred, evidence of grazing history, soil phosphorus and vegetation height correlated negatively with the strongest ordination axis. Least grazed forest plots had fewer tree seedlings and saplings than more intensively grazed plots. This may be due to the increase in Mimulopsis alpina (Acanthaceae) in less grazed forest where tree regeneration might otherwise be more advanced. Tree seedlings and saplings were uncommon in the forest, rarely exceeding 30cm in height and there was no tree understorey. Although grazing is important for preserving species diversity in Mount Elgon National Park through the maintenance of species-rich grasslands, long-term effects on montane forest communities must be considered in future park management.