Factors affecting the occurrence of woody plants in understory of sugi (Cryptomeria japonica D. Don) plantations in a warm-temperate region in Japan

We compared the occurrence of woody plants present in sugi (Cryptomeria japonica D. Don) plantations and evergreen broad-leaved natural forests in a warm-temperate region in order to characterize the understory vegetation of sugi plantations. The influence of stand attributes on species occurrence in the plantations was evaluated using logistic regression analysis. Out of 163 species, 65 plantation-preferring species and 31 natural forest-preferring species were detected. The plantation-preferring species group contained deciduous species (38 of 65 species) that are characteristic of coppice woodland, but all the natural forest-preferring species were evergreen plants. This indicates that the composition of the woody understory in sugi plantations of the region is characterized by the species of open, disturbed habitats. Stand age and the distance from the nearest natural forests were found to significantly affect the occurrence of evergreen shrubs and plants with gravity-dispersed seeds, respectively, but the light environment of the stand had less effect. According to the regression model analyses, long-term strategies such as lengthening the rotation (logging age) or creating patch mosaics of plantations in the matrix of natural forests appeared to be effective in conserving woodland species that are characteristic of evergreen, broad-leaved forests. On the other hand, increasing light penetration through thinning practices is a short-term strategy that is less likely to be effective in promoting these species.     

The influence of canopy patch mosaics on understory plant community composition in boreal mixedwood forest

We assessed the composition of understory vascular plant communities in relation to the mosaic of canopy patch types, and their associated structure and environment, within unmanaged, mature boreal mixedwood forests in western Canada. Within a 30 km² area, we sampled patches of four different canopy types: conifer-dominated, broadleaf-dominated, mixed conifer-broadleaf, and canopy gaps (total n = 98). There were significant differences in understory composition among the four patch types (based on multi-response permutation procedure (MRPP)) and these were mainly due to differences in relative abundances of understory species. The understory communities of conifer patches were characterized by low abundances of shade intolerant species while shade-tolerant and evergreen species were indicators (based on an indicator species analysis (ISA)). Understory communities under gap and broadleaf patches were characterized by higher abundances of grasses and shade intolerant species. Gap, broadleaf, and mixed patches had higher abundances of certain shrub species than did conifer patches. The patch types also differed in terms of their environmental conditions. Conifer patches had drier, cooler soils and the lowest understory light. Broadleaf patches had the warmest soils while understory light during the leaf-off period was similar to that of canopy gaps. Gap patches had the lowest litter cover and PO₄⁻ availability and the highest light. Seven environmental variables (soil moisture, soil temperature, total light during the leaf-off period, cover of coarse and fine downed woody material, and availability of NH₄⁺ and Ca²⁺) were significantly related to understory species composition (in a constrained ordination by means of a distance-based redundancy analysis (db-RDA); 16.5% of variation in understory community data explained). Even within a single patch type, there was substantial environmental variation that was related to understory species composition. Our study suggests that the mosaic of canopy patches within mixedwood forests supports coexistence of both early and late successional understory plant species in mixedwood stands. Maintaining the mixture of canopy patch types within mixedwood stands will be important for conserving the natural patterns of understory plant composition in boreal mixedwood forests.     

Habitat structure influences the spider fauna of short-rotation poplar plantations more than forest age

Plantation forests comprise an important part of the forested areas in European countries. Intensive forestry management and short-rotation cycles of plantation forests reduce habitat diversity and change the composition of invertebrate assemblages, mainly by reducing the number of habitat specialist species. Here, we analysed the effect of vegetation structure, amount of dead organic matter (DOM) and plantation age on spider functional diversity and assemblage structure in short-rotation plantations of native silver poplar in Hungary. Three stages, representing young plantations, 6- to 10-year-old stands (five stands), middle-aged plantations, aged between 23 and 26 years (five stands) and mature, 35- to 37-year-old forests at commercial maturity (five stands) were sampled. Each sample consisted of the data of ten pitfall traps. Traps were installed 5 m from each other in a 2 × 5 grid. Functional diversity was positively related to vegetation structure. High functional diversity indicates a higher number of available niches and potential resources. The abundance of moist habitat species and forest specialist species was positively correlated with DOM. Furthermore, moist habitat species were also related to vegetation structure. The most important environmental parameters affecting spiders were factors attributed to trees (litter and DOM), understory vegetation structure and, to a lesser extent, forest age. Different-aged stands may be similar in terms of species composition of their spider fauna if they comprise the same habitat structural patterns. Our study emphasises that the simple habitat structure of plantation forests has a negative effect on spider communities.     

Establishment and early growth of introduced indigenous tree species in typical plantations and shrubland in South China

Unsuccessful colonization by indigenous tree species into established plantations has retarded the succession of artificial plantations to more natural, secondary forests in South China. To understand how to improve colonization by seedlings of indigenous species, we determined how performance of indigenous seedlings is affected by seedling species (the shade-intolerant Castanopsis chinensis, the moderately shade-intolerant Michelia chapensis, and the shade-tolerant Psychotria rubra), the site into which the seedlings were transplanted (a mixed-legume plantation, a eucalyptus plantation, a mixed-native plantation, a mixed-conifer plantation, and a shrubland), and site preparation (removal or retention of understory vegetation and litter). Seedling survival and growth were generally increased by removal of understory vegetation and litter. C. chinensis and M. chapensis grew better in the mixed-legume and mixed-conifer plantations, while P. rubra grew better in mixed-native and mixed-conifer plantations. Responses of the transplanted seedlings to environmental factors were species specific. The effects of light on seedling survival and growth were correlated with the shade tolerance of the species. Soil moisture was important; it was positively correlated with survival but negatively correlated with growth of C. chinensis seedlings. Growth of C. chinensis and M. chapensis was positively correlated with soil potassium, while growth of P. rubra was positively correlated with soil organic matter but negatively correlated with soil hydrolyzed nitrogen. These findings suggest that we should select suitable native species under the different plantations before improvement of plantations. Light and soil moisture are most important environmental factors for the selection of species specific. Site preparation and fertilizer are needed during the improvement of those plantations.     

Woody understory plant diversity in pure and mixed native tree plantations at La Selva Biological Station,Costa Rica

Tree plantations can be an important tool for restoration of abandoned pasturelands in the tropics. Plantations can help speed up secondary forest succession by improving soil conditions, attracting seed-dispersal agents, and providing shade necessary for understory growth. In this study, abundance and richness of understory regeneration was measured in three native tree plantations 15–16 years of age at La Selva Biological Station in the Costa Rican Caribbean lowlands. Each plantation contained tree species in pure plots, a mixture of the species, and natural regeneration plots (no trees planted). The greatest abundance of regeneration was found in the understory of pure plots of Jacaranda copaia (Aubl.) D.Don., Vochysia guatemalensis Donn.Sm., Dipteryx panamensis Benth, Vochysia ferruginea Mart., and in two mixed stands, while the lowest was found in the natural regeneration treatments with about half the values as in the plantation stands. There was a significant negative correlation between percent canopy openness and abundance of regeneration in the understory. Two distinctive clusters separated the regeneration treatments from the mixed and pure plantations at a very low Bray–Curtis similarity value. The natural regeneration treatments are separated from mixed and pure plantations in the two-dimensional ordination. The lack of difference between the understory make-up of pure and mixed plantations in abundance, species richness, and seed-dispersal syndromes of understory species suggests that planting mixed stands is not necessarily superior to planting pure stands for promoting understory diversity of woody species. While regeneration of woody species can be faster under pure- or mixed-species plantations than in open pastures, the abundance, richness and species composition depends on each plantation species, or species assemblages in case of the mixtures.     

The response of small mammals to natural and human-altered edges associated with Afromontane forests of South Africa

An increase in edge area reduces the effective size of habitat fragments and thus the area available for habitat-interior specialists. However, it is unclear how edge effects compare at different ecotones in the same system. We investigated the response of a small mammal community associated with Afromontane forests to edge effects at three different habitat transitions: natural forest to grassland (natural edge, structurally different vegetation types), natural forest to mature plantation (human-altered edge, structurally similar vegetation types) and natural forest to harvested plantation (human-altered edge, structurally different vegetation types). We predicted that edge effects should be less severe at natural ecotones and at similarly structured contiguous vegetation types than human-altered ecotones and differently structured contiguous vegetation types, respectively. We found that forest species seemed to avoid all habitat edges in our study area. Surprisingly, natural edges supported a less diverse small mammal community than human-altered forest edges. However, edge effects were observed deeper into native forests surrounded by mature alien plantations (and more so at harvested plantations) than into native forests surrounded by native grasslands. The net effect of mature plantations was therefore to reduce the functional size of the natural forest by creating a larger edge. We suggest that when plantations are established a buffer zone of natural vegetation be left between natural forests and newly established plantations to mitigate the negative effects of plantation forestry.     

Species richness of the understory woody vegetation in Japanese cedar plantations declines with increasing number of rotations

The possibility of restoring natural broadleaf forests may be decreased by the effects of plantation management, particularly in sites that undergo repeated rotation. We investigated the following two working hypotheses about the effects of repeated plantation of conifers on the natural regeneration of woody saplings in cool-temperate Japanese cedar plantations: (1) that repeated plantation of conifers decreases sapling species richness, and (2) that repeated plantation of conifers changes sapling species compositions. Our result supported the first hypothesis, because species richness was significantly lower in second-rotation plantations than in first-rotation plantations. The second hypothesis was not supported, because no significant or substantial differences in species composition were observed between plantations with different numbers of rotations. However, the abundance of tree (nonshrub) and gravity-dispersed species decreased after the second rotation of large saplings, albeit not those of small saplings, suggesting that response to repeated rotation depended on sapling size. Our results suggest that it is important to consider factors affecting the maintenance of a species in the plantations, such as distance from natural forests and seed sources, to minimize the effects of repeated plantation.     

热带北缘地区种植桉树前后林下植被的变化

分析我国热带北缘地区炼山营造的桉树(Eucalyptus)人工林前后林下植被物种组成与生长的差异,初步探讨桉树与林下植被的关系,为我国热带地区的桉树种植提供参考。造林前的植被包括3个马尾松(Pinus massoniana)林和1个果树林,炼山后全部种植桉树。结果表明:炼山造林2 a后,林下植被得到恢复,其种类组成没有发生改变,而频度的变化则与植物类型有关,频度减小的以灌木为主,频度增大的以草本为主。果树林改造为桉树林后,林下植被出现了一些新的物种,且物种平均高度大幅增加;马尾松林改造为桉树林后,多数物种的平均高度和盖度接近原有群落的水平,而芒萁(Dicranopteris dichotoma)从林下植被层消失,割鸡芒(Hypolytrum latifolium)盖度大幅提高,表明种植桉树对林下植被的影响因物种而异。经过炼山造林,在广东省茂名地区种植桉树对少数物种影响较大,短期内没有降低林下植被的多样性,长期是否存在影响还需要进一步观察。     

Hybrid poplar plantations in a floodplain have balanced impacts on farmland and woodland birds

Hybrid poplar plantations are increasing worldwide and are often accused of impoverishing bird communities. We conducted 124 bird point counts in a landscape where plantations, semi-natural forests and farmland occupied similar surface areas. As expected, birds occurred at higher densities in areas dominated by forests than dominated by plantations, mostly due to the scarcity of late-successional forest birds in plantations. Contrary to expectations, bird communities were the poorest in farmland dominated areas and the most specialised in areas dominated by young plantations. Indeed, many grassland species, including some of conservation concern in Europe, frequently used young plantations. However, plantations have probably depleted grassland bird communities by fragmenting open areas, while playing a limited positive role on forest species. Total length of unpaved roads had a positive effect on community specialisation index, while total length of paved roads and mean forest/plantation patch size played no role. Bird density increased with the development of the understory vegetation in mature poplar plantations. We conclude that poplar should not be planted at the expense of areas of high conservation value; in other cases, they may increase the overall diversity of bird communities. At the plantation scale, we advocate stopping understory removal as soon as possible and maintaining old trees and hedgerows alongside or within plantations to provide suitable breeding habitats for bird species that may be foraging in the plantations.     

A preliminary study on the effects of line and selective thinning on bird communities in Hokkaido, northern Japan

In Japan, selective thinning is a common thinning method, though line thinning receives much attention because of its economic merits. In this study, we examined effects of the two thinning methods on bird communities in Todo fir (Abies sachalinensis) plantations in Hokkaido, Japan. We surveyed bird species in forests under four different management types — unthinned, selectively thinned, line-thinned plantation, and naturally regenerated forest (here after referred to as natural forest) stands — using a line-transect method. We also investigated vegetation structure (canopy tree and understory) of these stands. Bird species richness did not differ between natural forests and plantations, while bird total abundance was greater in plantations than in natural forests. Bird species richness and total abundance were comparable among the three management types for plantations. Abundances of 10 bird species were different among the four management types, and five species were more abundant in line-thinned plantations. However, two species were more abundant in selectively thinned stands than in line-thinned stands, and they frequently appeared in natural forests. There were no distinct differences in vegetation structure among the management types for plantations. Our results suggest that line thinning could be beneficial for some bird species in plantations.     

Understory plant diversity assessment of <Emphasis Type="Italic">Eucalyptus</Emphasis> plantations over three vegetation types in Yunnan,China

Biodiversity in managed plantations has become an important issue for long-term sustainability of ecosystems. The environmental effects of plantations comprised of fast-growing introduced trees have been vigorously debated. On one hand, monocultures have been said to exhaust resources, resulting in decreased biodiversity. Conversely, it has been stated that monocultures may favor regeneration of undergrowth plants from surrounding forests, increasing biodiversity. In order to clarify the effects of planting Eucalyptus trees on species composition, diversity, and functional type of understory vegetation in Yunnan province, a field trial was implemented to compare Eucalyptus plantations (EPs) with two other local current vegetation types (secondary evergreen forests (SEs), and abandoned farmlands (AFs)). Each vegetation type was sampled in each of three elevational ranges (low = 1,000–1,400 meters above sea level (masl), medium = 1,400–1,800 masl, and high = 1,800–2,200 masl). Sample sites within each elevational range had similar environmental characteristics (slope, aspect, etc.). Thus, we sampled three vegetation types at each of three sites at each of three elevations for a total of 27 plots. We calculated relative abundance and importance value of species and diversity indexes to evaluate differences among local current vegetation types and elevational ranges, employing multivariate ordination analyses and other methods such as Analyses of Variance (ANOVA) and Indicator Species Analysis. We found that fast growing introduced Eucalyptus plantations led to reduced plant diversity in the study area, and that rare or threatened species were recorded almost exclusively in the SE plots, being essentially absent from the EP and AF plots. The understory plant diversity did not correlate with the altitude gradient significantly. Eucalyptus plantations (EPs) have a simpler community structure than that of either secondary evergreen forests (SEs; similar to natural state) or abandoned farmlands (AFs). No variable significantly explained variation of the understory shrub layer, but soil moisture-holding capacity and overstory coverage were significant in explaining variation of the understory herb layer, suggesting that the study of soil physical properties is necessary for better understanding of their importance in Eucalyptus plantations and other local current vegetation types.     

Do mature pine plantations resemble deciduous natural forests regarding understory plant diversity and canopy structure in historically modified landscapes?

We compared the structure of the arboreal layer and the diversity and species composition of the understory vegetation of three types of mature forest communities: oak (Quercus pyrenaica) and beech (Fagus sylvatica) forests and Scots pine (Pinus sylvestris) plantations. Our main aim was to determine whether differences in these variables existed and were due to the identity of the dominant tree species. We selected four stands or replicates per forest type located geographically close and with relatively similar conditions. We found no differences in the arboreal structure of oak and beech forests, which were characterised by great variability in tree size, while in case of plantations, this variability was lower at both the intra-stand (estimated by the coefficient of variation) and inter-stand (i.e. the four replicates harboured trees of similar sizes) scales. However, the highest variability in the canopy layer of natural forests was not consistently linked to greater understory species richness. Indeed, the lowest plant species richness was found in beech forests, while oak forests harboured the highest value at either the sampling unit (per m²) or stand scales. The greatest negative correlation between plant diversity and the environmental variables measured was found for litter depth, which was the highest in beech forests. The results obtained by the CCA indicated that the four replicates of each forest type clustered together, due to the presence of characteristic species. We concluded that pine plantations did not approach the environmental conditions of native forests, as plantations were characterised by singular understory species composition and low arboreal layer variability, compared to natural woodlands.     

Assessing the relationships between stand development and understory vegetation using a 420-year chronosequence

The large-scale conversion of old forests to tree plantations has made it increasingly important to understand how understory vegetation responds to such landscape changes. For instance, in some forest types a reduction in understory richness and cover is thought to result from the development of canopy closure in plantations, although there is a paucity of empirical data demonstrating this relationship. We used a 420-year forest chronosequence as a case study to assess the relationship between stand age, tree canopy cover and understory vascular plant richness and composition in the Siskiyou Mountains of Oregon. The chronosequence consisted of six young managed (age 7–44) and nine older unmanaged (age 90–427) stands. All stands were similar in underlying geology, slope, elevation, and aspect. We found a non-linear relationship between stand age and richness, in which richness was highest in the youngest stands, reached a low in mid-aged stands (∼55 years), then increased in the oldest stands. We also found that percent tree canopy cover was correlated with total understory cover, richness, diversity, and species composition. In general, young stands were characterized by high shrub and graminoid cover and old stands were characterized by an abundant herb layer. Our work suggests that a major component of our study landscape is currently entering the forest stage (canopy closure) characterized by low levels of vascular plant species richness and cover. We use our results to discuss the potential effects of future forest management on understory plants.     

Influence of tree species on understory vegetation diversity and mechanisms involved—A critical review for temperate and boreal forests

Tree species composition is a primary attribute of forest ecosystems, and is often manipulated by silvicultural practices. Forest management to diversify tree species is now being promoted to favor biodiversity. To assess the soundness of this policy we reviewed and analyzed the literature on the relationship between tree species composition and floristic diversity, including the mechanisms involved therein. Coniferous forests generally provide less diversified vascular understories than broadleaved forests. At the tree species scale, there are not enough reports to draw firm conclusions on the effect of any particular species. Mixing of deciduous and coniferous tree species generally affects understory diversity, but in almost all cases maximum diversity is observed in one of the pure stands, not in mixed stands. Understory vegetation is influenced by overstory composition and structure through modifications of resource availability (light, water and soil nutrients) and other effects, such as physical characteristics of the litter layer. Overstory light transmittance and diverse properties of forest litter are factors that have been most fully studied to date, but other factors such as throughfall water quantity and chemistry may also play a role. While the relative importance of mechanisms that account for the effect of overstory on understory biodiversity has often been discussed, these mechanisms have rarely been the subject of formal experiments. Overall, varying management practices and site attributes make it difficult to generalize results. They combine with the effects of tree species in influencing understory vegetation diversity, but they have been rarely considered. Future research is needed to gain a better understanding of the relationship between overstory and understory diversity and establish general laws.     

Landscape effects in bird assemblages differ between plantations and broadleaved forests in a rural landscape in central Japan

It is important to conserve forest-dependent organisms not only in broadleaved forests but also in plantation forests. We examined how surrounding forest areas affect forest bird assemblages inhabiting conifer plantations and broadleaved forests in a rural landscape in central Japan. Surrounding forest areas were measured separately as plantation area and broadleaved forest area within 200 m of each sampling site. We used hierarchical partitioning to analyze the effects of surrounding forest areas and stand structures (stand height and understory coverage) on the occurrence of four species groups. We especially focused on mature forest users that are most sensitive to loss of broadleaved forests. Occurrence of mature forest users inhabiting plantation sites was positively affected both by plantation area and broadleaved forest area whereas the occurrence of mature forest users inhabiting broadleaved forests was affected by stand height only. These results suggest that surrounding forest areas were more important to mature forest users in plantations than in broadleaved forests. To conserve mature forest users, increasing surrounding forest areas are important in plantations whereas increasing stand heights would be effective in broadleaved forests.     

九龙山多脉青冈-鹅掌楸林演替过程种群生态位

浙江九龙山分布有珍稀濒危落叶树种为主要树种的亚热带常绿落叶阔叶混交林。为探明这些落叶树种在常绿落叶阔叶混交林进展演替过程中生态位变化规律,选择该地区多脉青冈—鹅掌楸林演替过程中4个不同阶段的代表性群落,进展演替过程中,在群落垂直空间上常绿阔叶树不断挤压落叶阔叶树的生存空间,该地区常绿阔叶林顶极群落对珍稀濒危落叶树种是致危生境,不利于生物多样性保护。     

间伐对栓皮栎人工林林下更新的影响

栓皮栎是我国温带落叶阔叶林代表树种之一,是河南省山区典型地带性树种和主要造林树种。间伐作为一个重要的森林经营措施,研究其对林下更新的影响,可为改善栓皮栎人工林林分结构,维持林下物种多样性,制定科学合理的森林经营措施提供支撑。对间伐3 a后栓皮栎人工林林下幼苗更新物种及更新多样性进行调查,结果表明:间伐后林地内幼苗种类和数量显著增加,间伐样地内单物种更新密度和总更新密度均高于未间伐样地,间伐样地内更新苗Shannon-Wiener、Simpson多样性指数、Patrick丰富度指数值、Pielou均匀度指数均高于未间伐样地相应数值,间伐有利于栓皮栎人工林林分结构优化和林分健康。     

Patterns and factors in early-stage vegetation recovery at abandoned plantation clearcut sites in Oita, Japan: possible indicators for evaluating vegetation status

The abandonment of sites after clearcutting plantations is increasing rapidly in Japan. Although a few stand-level studies have been conducted, a broad-scale study is needed to understand the general tendency of vegetation recovery and to obtain information for the management of such extensively distributed abandoned clearcut sites. This study is a province-level study that aims to interpret the patterns and factors in early-stage vegetation recovery at abandoned plantation clearcut sites. We also discuss the potential indicators that may determine whether a certain site recovers towards evergreen broad-leaved forest consisting of Castanopsis cuspidata and Quercus glauca or towards deciduous broad-leaved forest comprising Q. serrata and Q. acutissima. A cluster analysis revealed that five types of vegetation (evergreen trees, evergreen trees–deciduous pioneer shrubs, pioneer species, deciduous trees, and deciduous shrubs) were observed from an early stage of succession. The first two vegetation types were considered to recover successfully to the target vegetation of evergreen broad-leaved forest, but the remaining three vegetation types did not show any tendency to recover to the target vegetations. According to the results of the detrended correspondence analysis, a smaller size of the abandoned site with adjacent evergreen broad-leaved forest and a longer time after clearcutting were identified as factors that enhance the invasion of evergreen tree species; these factors are also useful to predict the future successional tendency. In conjunction with these factors, the existence of regeneration sources (i.e., stumps) that allow sprouting appeared to be an important indicator for predicting the early-stage vegetation recovery.     

Understory diversity and composition after planting of teak and mahogany in Yogyakarta,Indonesia

ABSTRACT

Forest rehabilitation is when a desired tree species is planted in degraded forests or lands. Rehabilitation by planting a single tree species is a common way to restore exploited forests to maintain ecological processes. We compared woody and herbaceous understory vegetation between forests rehabilitated by mahogany (N = 12) or teak (N = 12) planted from 1941 until 2003 in Yogyakarta, Indonesia. Understory vegetation of these areas was compared with that of three native forests. Species richness, species diversity, density of plants and proportion of native plants did not differ between the rehabilitated areas and the native forest. Recently rehabilitated areas were different from the native forests while 41–74 yr after rehabilitation, characteristics of understory vegetation approached those of native forest. We described species composition using ordination, and found it to differ between areas rehabilitated with teak and with mahogany and, particularly, between the rehabilitated areas and the native forests. Time since rehabilitation and tree species planted were important for the species composition of understory vegetation. We conclude that the selection of species for rehabilitation and letting rehabilitated areas mature are important for understory development and species diversity.     

Understory plant communities of boreal mixedwood forests in western Canada: Natural patterns and response to variable-retention harvesting

We examined patterns of variation in richness, diversity, and composition of understory vascular plant communities in mixedwood boreal forests of varying composition (broadleaf, mixedwood, conifer) in Alberta, Canada, before and for 2 years following variable-retention harvesting (clearcut, 20 and 75% dispersed green tree retention, control). Broadleaf-dominated forests differed from mixedwood or conifer-dominated forests in that they had greater canopy cover, litter depth, soil nitrogen, warmer soils, as well as greater shrub cover, herb and shrub richness and diversity (plot scale). In contrast, conifer, and to a lesser extent mixedwood, forest had greater β diversity than broadleaf forest. Overall, mixedwood and conifer forests were similar to one another, both differed from broadleaf forest. Several species were found to be significant indicators of broadleaf forest but most of these also occurred in the other forest types. Understory composition was related to canopy composition and edaphic conditions. Variable-retention harvesting had little effect on understory cover, richness, or diversity but resulted in reduced richness and β diversity at a larger scale. The clearcut and 20% treatments affected composition in all forest types. Early successional species and those common in disturbed sites were indicators of harvesting while evergreen, shade-tolerant understory herbs were indicators of the control forest and 75% retention harvest. We conclude that it is important to maintain a range of variation in canopy composition of mixedwood forests in order to conserve the associated understory communities. The presence of conifers in these forests has a particularly important influence on understory communities. The threshold for a lifeboat effect of variable-retention harvesting is between 20 and 75% retention. Examination of richness and β diversity at a variety of scales can provide interesting information on effects of harvesting on spatial reorganization and homogenization of understory plant communities.