Leaf dynamics and protein content of six important fodder trees of the western Himalaya

Leaf production, leaf fall, leaf protein and net primary productivity have been reported for 4-year-old fodder trees in the mid-western Himalaya (altitude 1350 m; rainfall 1100 mm). The species include; Grewia optiva, Robinia pseudoacacia and Celtis australis growing in the open and Quercus glauca, Q. leucotrichophora and Ilex odorata growing in the forest. G. optiva, R. pseudoacacia and C. australis showed two distinct periods of leaf flush, and their leaves were short-lived (180 to 360 days) compared with the remaining three species which retained their leaves for more than two years, and had only one flush period during March–April. The open site species showed much higher leaf production, and the leaves contained a significantly (P < 0.05) higher protein content, compared with the forest group. Net primary productivity of G. optiva, C. australis and R. pseudoacacia was significantly (P < 0.05) higher than the forest species, which may be related to their high leaf production and rapid turnover rate of the leaves. The results are useful for management of these species in agroforestry systems.     

The effects of understory vegetation on P availability in Pinus radiata forest stands: A review

In many second-rotation Pinus radiata forest planta-tions, there has been a steady trend towards wider tree spacing and an increased rate of application of P fertiliser. Under these regimes, the potential for understory growth is expected to in-crease through increased light and greater nutrient resources. Therefore, understory vegetation could become a more signifi-cant component of P cycling in P. radiata forests than under closely-spaced stands. Studies have shown that growth rates and survival of trees is reduced in the presence of understory vegeta-tion due to the competition of understory vegetation with trees. Other studies have suggested that understory vegetation might have beneficial effects on nutrient cycling and conservation within forest stands. This review discusses the significance of understory vegetation in radiata pine forest stands, especially their role in enhancing or reducing P availability to forest trees.     

Carbon dioxide sequestration capability of an unmanaged old-growth broadleaf deciduous forest in a Strict Nature Reserve

The seasonal trend of plant carbon dioxide (CO₂) sequestration is related to the photosynthetic activity, which in turn changes in response to environmental conditions. Great interest has turned to the CO₂ sequestration (CS) potential of temperate forests which play an important role in global carbon (C) cycle contributing to the lowering of atmospheric CO₂ concentration. In such context, the CS of an unmanaged old broad-leaf deciduous forest developing inside a Strict Nature Reserve, and its variations during the year were analyzed considering the monthly variations of leaf area index (LAI) and net photosynthetic rates (N_P). Overall, the total yearly CS of the forest was 141 Mg CO₂ ha^?1 year^?1 with the highest CS value monitored in June (405 Mg CO₂ month^?1) due to the highest LAI (5.0 ± 0.8 m² m^?2) and a high N_P in all the broadleaf species. The first CS decline was observed in August due to the more stressful climatic conditions that constrained N_P rates. Overall, the total CS of the forest reflects the good ecological health of the ecosystem due to its conservative management.     

Differential overstory leaf flushing contributes to the formation of a patchy understory

Understory individuals were found to form patches in a 100-year-old deciduous broad-leaved forest. The closed forest canopy was uniform, and so the light conditions at various locations across the forest floor differed little after the leaf flush of the overstory. To explain the distribution pattern in the understory, a hypothesis was proposed: in spring, the forest floor is divided into patches according to the timing of leaf flush of the overstory individuals, and the light conditions are more favorable for understory plants under the crowns of trees with later-flushing leaves. In the plot, three groups of early, intermediate, and late, were recognized in the overstory concerning the timing of leaf flush. As for the start of leaf flush, a difference of 31.6 days was recognized among tree species, and for the end of leaf flush, a difference of 40.3 days. In the spring of 1998, the relative photosynthetic-photon-flux density under an intensively studiedCastanea crenata tree (late-flushing species) usually showed higher values than that under a similarly studiedAcer mono tree (early-flushing species). Analysis of the spatial-distribution pattern using Morisita’s1δ index revealed that the understory community had an aggregated distribution. In the overstory, the late- and the intermediate-flushing-species groups showed aggregated distributions, while the early-flushing-species group showed random distribution. Spatial correlation between the understory and the overstory was analyzed by using Morisita’sRδ index. The distribution of whole understory community spatially co-occurred with that of the late-flushing-species group of the overstory. In contrast, the understory community was less developed below the members of the early-flushing-species group of the overstory. We consider that the data presented here support our hypothesis, and we suggest that the growth and survival of understory individuals were promoted in the places receiving light for long periods in spring.     

林分密度对蒙古栎林下草本植物多样性的影响研究

以内蒙古赤峰市阿鲁科尔沁旗沙日温都栎林自然保护区蒙古栎林为主要研究对象,研究不同林分密度对林下草本植物多样性的影响。结果表明,研究区内共有13科、14属、16种草本植物;Simpson指数、Shannon-Wiener指数、Pielou均匀度指数和物种丰富度指数均随着林分密度的增加呈先增大后减小的趋势,当林分密度为750株/hm²时,林下草本植物多样性指数达到最大值;林分密度与Shannon-Wiener多样性指数、Pielou均匀度指数、Simpson多样性指数、物种丰富度指数、树高、胸径均呈极显著负相关,与郁闭度呈极显著正相关,与枝下高不相关。综上所述,最合理的蒙古栎林密度为750株/hm²,该密度下最有利于蒙古栎林及林下草本植物生长发育。     

Shade tolerance and suitability of tree species for planting in rubber plantations

The rapid increase in rubber monoculture in Xishuangbanna has resulted in extensive damage to its local ecosystem. To decrease the negative effects, the concept of the ecological-economic rubber plantation (EERP) system was proposed. The EERP entails intercropping rubber plants with other economically significant plants, which would not only decrease the ecological impact of the rubber plantation, but also maintain and potentially increase its profitability compared with rubber monocultures. In order to select the appropriate species and intercropping pattern in the EERP system, we compared the photosynthetic parameters of five economically important trees (Swietenia mahagoni, Coffea arabica, Mesua ferrea, Myristica yunnanensis and Paramichelia baillonii) under four irradiance levels. The optimal irradiance intensity of C. arabica and M. yunnanensis was approximately 40%, and these species can be planted in the understory of an 8- to 15-year-old rubber forest. Adult S. mahagoni, M. ferrea and P. baillonii grow taller than the rubber trees, and their seedlings also show superior performance in 100% irradiance than in other irradiances. Thus, in an EERP, these valuable trees could be planted adjacent to rubber plantations but on sites that are economically and environmentally marginal for rubber production.     

Correlation of eco-hydrographic benefit and height increment of Robinia pseudoacacia stand with climatic environmental factors in Yellow River Delta Wetland of China

The relationship between eco-hydrographic benefit of forest vegetation and climatic environmental factors is one of the focuses in the research on environmental protection and ecosystem countermeasures in Wetland. The runoff, sediment and soil moisture rate dynamics in Robinia pseudoacacia plantation and its clearcut area were investigated in the natural runoff experiment plots in Yellow River Delta Wetland, Shandong Province, China. The correlation of height increment of R. pseudoacacia with nine climate factors such as light, water, heat, etc. was analyzed by stepwise regression analysis. The results showed that the amounts of runoff and sediment in clearcut area of R. pseudoacacia were 53.9%–150.8% and 172.8%–387.1% higher than that in Robinia pseudoacacia plantation, respectively. The runoff peak value in R. pseudoacacia stand was obviously lower than that in clerarcut area, meantime, the occurrence of runoff peak in R. pseudoacacia stand was 25 min later than in its clerarcut area. The soil moisture rates in R. pseudoacacia stand and its clearcut varied periodically with annual rainfall precipitation in both dry season and humid season. The annual mean soil moisture rate in R. pseudoacacia stand was 23.3%–25.6% higher than that in its clearcut area. Meanwhile, a regression model reflecting the correlation between the height increment of R. pseudoacacia and climatic factors was developed by stepwise regression procedure method. It showed that the light was the most important factor for the height increment of R. pseudoacacia, followed by water and heat factors. Foundation project: Subsidized by the National “11th Five Year” Plan of Science and technology (2006BAD26B06, 2006BAD03A1205) and Ecological Restore Project of Water Resources Ministry of China (2006-2008).     

Effects of forest structure and microhabitat on the distribution and flowering of a rare understory plant, Actaea elata

Knowing which environmental characteristics constrain forest understory species’ distribution and reproduction can inform conservation decisions about habitat management and locations for reintroduction efforts. Differential plant performance is common in varying environmental conditions, such as levels of canopy closure. Actaea elata (tall bugbane) is a rare, perennial, forest understory herb found from Oregon to British Columbia. Populations throughout this species’ range commonly occur in managed forests. We assessed the importance of environmental variation on the distribution and flowering of A. elata at both microhabitat and site-level scales. At four sites encompassing a variety of forest types, we measured several biotic and abiotic variables at vegetative and reproductive plants, as well as at random points. We measured these same variables at three sites where A. elata has been extirpated. Within occupied sites, reproductive plants were more likely to be located in canopy gaps than were vegetative plants. Both vegetative and reproductive plants were more likely to be found in areas of high herb cover compared to random points. These results indicate that gaps in the canopy may be critical for this species’ ability to reproduce, but survival may be possible in a broader range of forest understory conditions (similar to other co-occurring herb species). Random points at extirpated and occupied sites both tended to be located in areas of lower canopy openness and moisture availability than points in occupied sites where A. elata was present. Extirpated sites were either densely forested or clearcut, while occupied sites contained stands of intermediate density. Circaea alpina, Geranium robertianum, and Lactuca muralis were indicator species for A. elata and normally indicate moist soils, with Circaea alpina being a strong indicator of the presence of flowering A. elata. Taken together, our results suggest that maintenance of canopy gaps is important to promote flowering, and understory species that indicate moist sites, particularly Circaea alpina, can be used as aids to locate new populations of A. elata.     

Photosynthesis and biomass allocation of beech (Fagus crenata) and dwarf-bamboo (Sasa kurilensis) in response to contrasting light regimes in a Japan Sea-Type beech forest

Regeneration of beech (Fagus crenata) forests depends on the formation of canopy gaps. However, in Japan Sea-type beech forests, a dwarf bamboo (Sasa kurilensis) conspicuously occupies sunny gaps. Therefore,F. crenata seedlings must escape the severe interference ofS. kurilensis in the gaps and persist beneath a closed canopy of the beech forest. We hypothesized that the growth ofF. crenata seedlings in the understory would be favored by their being more plastic thanS. kurilensis in photosynthetic and morphological traits, which would support the matter production ofF. crenata seedlings in a wide range of light availabilities. To examine this hypothesis, the photosynthetic-light response of individual leaves and the biomass allocation in aboveground parts (i.e., the culm/foliage ratio) were surveyed at sites with contrasting light availabilities in a Japan Sea-type beech forest in central Japan. InF. crenata, photosynthetic light utilization efficiency at relatively low light was greater, and the dark respiration rate was smaller in the leaves of seedlings (10 cm in height) beneath the closed canopy than in the leaves of saplings at the sunny forest edge. The culm/foliage (C/F) ratio of theF. crenata seedlings at the shady site was small, suggesting effective matter-production beneath the beech canopy. On the other hand,S. kurilensis both in the gap and beneath the beech canopy showed low plasticity in photosynthesis and the culm/foliage ratio. Because the shoot density ofS. kurilensis was smaller beneath the beech canopy than in the gap, the light availability at the bottom of theS. kurilensis layer was greater beneath the beech canopy. These results suggest that the photosynthetic productivity of theF. crenata seedlings would be enough for the seedlings to survive in the understory with a low density ofS. kurilensis shoots beneath the closed beech canopy.     

Distance to flood meadows as a predictor of use of <Emphasis Type="Italic">Nothofagus pumilio</Emphasis> forest by livestock and resulting impact,in Patagonia,Argentina

Seedling browsing by livestock has been indicated as major threat for forest sustainability use. Nothofagus pumilio forests are part of the livestock raising system in Patagonia, but because of the sparse understory cover, livestock graze mainly on flood meadows within the forest matrix. The complexity of the environment under study (forests and flood meadows) means that an adaptive predictor is needed to evaluate the intensity of resource use by livestock in order to assess its effect on the forest. Distance to flood meadows was evaluated as a predictor of the use intensity of the forest by livestock and its effect on the understory. The study was conducted at three sites in Chubut Province, Patagonia–Argentina. We established transects in the forest 320 m long, starting at the edge flood meadow-forest. In these transects, the livestock presence indicators (soil compaction, density of feces and trails) and composition of the understory were evaluated. Generalized Linear Model for repeated measures for longitudinal data were used. The indicators showed that distance is efficient for estimating forest use intensity by livestock. The understory varied with distance, the cover of exotic herbaceous was higher near the flood meadow. Far from the flood meadow, the cover was entirely composited of native species. The results support the use of distance to flood meadows as a tool for decision making in livestock and forest management in N. pumilio forests, and for further research on livestock effect on the forest.     

Studies on the diversity of soil animals in Taishan Mountain

Taishan Mountain has diverse habitats and abundant groups of soil animals. Five habitats,Platycladus orientalis forest,Pinus densiflora forest,Robinia pseudoacacia forest,Pinus tablaerormis forest and Grassland, were selected and the diversity of soil animals in different habitats in Mt. Taishan were investigated and studied in 1997–1999. Totally 52 groups of soil animals were found, belonging separately to 7 Phyla, 11 Classes, 25 Orders and 28 suborders or families. There exists remarkable difference in groups, quantity and diversity of soil animals among the five habitats. Grassland habitat was the highest andPinus densiflora forest was the lowest in groups, quantities, Density-group index (DG) and Diversity index of comparing many communities (DIC) of soil animals. Shannon-Wiener’s diversity index (H′) for soil animals of different habitats was in order ofPlatycladus orientalis forest>Pinus densiflora forest>Pinus tablaerormis forest>Grassland>Robinia pseudoacacia forest, Pielou’s equality index (J′) wasPlatycladus orientalis forest>Pinus densiflora forest>Grassland>Pinus tablaerormis forest>Robinia pseudoacacia forest, and Simposn’s dominant index (C) was Grassland>Robinia pseudoacacia forest>Pinus tablaerormis forest>Pinus densiflora forest>Platycladus orientalis forest. The H index and J index of macro-soil animal all were larger than that of mid-small soil animal. Clustering analysis for five soil animal communities and ordination analysis for community structure by multidimensional scaling (MDS) were made and completely identical results were obtained. Soil animal communities of five habitats were divided into 3 groups. Biography: Y{upuan} Xing-zhong (1963-), male, associate professor of East China Normal University, Shanghal 200062, China Responsible editor: Chai Ruihai     

Light-related competitive effects of overstory trees on the understory conifer saplings in a subalpine forest

To clarify the mechanism by which overstory trees shade understory saplings, we investigated the relationships among light conditions of the saplings (measured as indirect site factor; ISF and direct site factor; DSF), the calculated competition effects of overstory trees on the saplings (W), and relative height growth rate of the saplings (RHGR). We calculated several W values in order to find a W value which can express the light conditions as appropriately as possible, and the results indicated that W explained only 21.9%–24.7% of the total variance of light conditions in the cases where W gave the best fit. In this study, W was calculated based on the basal areas of overstory trees. However, it is known that canopy structure also affects the light regimes in the forest understory, and this might yield the possible errors even within W representing the shading effects most adequately. Therefore, although W significantly represents the shading effect from overstory trees, a great proportion of the variance remained without being explained by W. RHGR was negatively correlated with W, and the W value which had the most adequate explanation of the shading effect also showed the best negative correlation with RHGR. This provides the evidence that the competitive effect of overstory trees on sapling growth is mediated by the shading effect, indicating that competition for light clearly exists within this forest. Such competition for light may closely relate to the well-known phenomenon of gap regeneration in subalpine forests in central Japan.     

Impacts of understory species removal and/or addition on soil respiration in a mixed forest plantation with native species in southern China

Although the removal or addition of understory vegetation has been an important forest management practice in forest plantations, the effects of this management practice on soil respiration are unclear. The overall objective of this study was to measure and model soil respiration and its components in a mixed forest plantation with native species in south China and to assess the effects of understory species management on soil respiration and on the contribution of root respiration (R_r) to total soil respiration (R_s). An experiment was conducted in a plantation containing a mixture of 30 native tree species and in which understory plants had been removed or replaced by Cassia alata Linn. The four treatments were the control (Control), C. alata addition (CA), understory removal (UR) and understory removal with C. alata addition (UR + CA). Trenched subplots were used to quantify R_r by comparing R_s outside the 1-m² trenched subplots (plants and roots present) and inside the trenched subplots (plants and roots absent) in each treatment. Annual soil respiration were modeled using the values measured for R_s, soil temperature and soil moisture. Our results indicate that understory removal reduced R_s rate and soil moisture but increased soil temperature. Regression models revealed that soil temperature was the main factor and soil moisture was secondary. Understory manipulations and trenching increased the temperature sensitivity of R_s. Annual R_s for the Control, CA, UR and UR + CA treatments averaged 594, 718, 557 and 608 g C m⁻² yr⁻¹, respectively. UR decreased annual R_s by 6%, but CA increased R_s by about 21%. Our results also indicate that management of understory species increased the contribution of R_r to R_s.     

Five-year effects of cutting and herbicide treatments on control of <Emphasis Type="Italic">Rhododendron flavum</Emphasis> Don., and macronutrient pools in eastern beech (<Emphasis Type="Italic">Fagus orientalis</Emphasis> Lipsky) forests of Turkey

This experiment was carried out in a mature (>100-year-old) eastern beech forest in the Eastern Black Sea Region (BSR) of Turkey. The effects of experimental practices to control the woody plant Rhododendron flavum Don., including grubbing, manual cutting, and foliar and cut-stump spraying with imazapyr and triclopyr, were assessed on understory and forest floor biomass and nutrients, as well as soil bulk density and nutrient content. The cut plots had 5, 15, and 38 times greater competing vegetation biomass than did the plots treated with triclopyr, grubbing, and imazapyr, respectively. Because of the increased biomass, the amount of understory N, P, Ca, K, S, and Mg content on the cut plots was also substantially higher when compared to the grubbed, imazapyr-treated, and triclopyr-treated plots. Triclopyr and imazapyr applications brought about two- and threefold greater dead organic matter on the forest floor than did cutting and grubbing, respectively. As a result, forest floor macronutrient contents on the imazapyr- and triclopyr-treated plots were significantly greater than on the cut plots. Five years after treatments, soil bulk density at the 10–20 cm depth on the imazapyr-sprayed plots was about 14% higher than that on other treatments. Herbicides, especially imazapyr and grubbing were effective in controlling rhododendron. Macronutrients stored in the rhododendron were released to the forest floor after vegetation was controlled with herbicides.     

Stand structure and the regeneration ofChamaecyparis pisifera (Sieb. et Zucc.) Endl. on sites with different soil development in an old-growth coniferous forest, central Japan

Stand structure and the regeneration ofChamaecyparis pisifera on sites with and without well-developed soil in an old-growth coniferous forest, Akasawa Forest Reserve, central Japan, were investigated, along with their relationships to other important coniferous species. Of stems ofC. pisifera≥5 cm diameter at breast height, stems with intermediate size were absent in stands on the developed-soil site, while those with smaller size increased in stands on the undeveloped-soil site, which supported a higher density ofC. pisifera. In the stands without recent tree-fall of canopy stems on the developed-soil site, canopy stems ofC. pisifera were distributed uniformly and the understory stems, patchily. On the other hand, both canopy and understory stems ofC. pisifera in stands on the undeveloped-soil site were distributed patchily. In the understory,C. pisifera occurred as clonal patches formed by vegetative reproduction on various types of microsites including bare rocks. However, the clonal patches did not occur in a stand on the developed-soil site with dense saplings of an evergreen conifer,Thujopsis dolabrata, which can expand vegetatively with a well developed soil layer. Gap formation may induceC. pisifera to develop tree-form stems from shrubby clonal patches. On developed-soil sites,C. pisifera stands that survive a long disturbance-free period on this site need catastrophic disturbance for their regeneration, which will eliminateT. dolabrata in the understory and create bare soil for colonization ofC. pisifera from seeds. On undeveloped-soil sites,C. pisifera replaces itself continuously by effective vegetative reproduction. Ground instability and the wet condition of this site may promote the replacement.     

Diversity of naturally-regenerated native woody species in forest plantations in the Ethiopian highlands

Woody species diversity and ground layervegetation cover were studied in plantations ofPinus patula, Cupressus lusitanica,Grevillea robusta, and Juniperusprocera, and in surrounding natural forests inWondo Genet, Ethiopia. In the understory of theplantations, a total of 53 naturallyregenerated tree and shrub species belonging to31 families were recorded. Important indigenoustimber species were also represented. Treesaccounted for 72% of all naturally-regeneratedwoody plant species. On the average, 78% ofthe woody species found in plantations werealso found in the natural forest. There was nosignificant variation in understory woodyspecies richness among plantations. C.lusitanica plantations had a significantlyhigher abundance of woody species than those ofP. patula, but the difference wasnot significant in relation to J. procera,G. robusta and the adjacent natural forest.The herbaceous ground cover percentage in G. robusta and P. patula stands wasconsiderably higher than in C. lusitanicaand J. procera stands. Relative toplantations of other species, plantations ofthe native tree species J. procera hadhigher understory species richness, mediumwoody species abundance and low ground cover.Although there was a marked variation inoverstory plantation density, its effect onunderstory species richness and abundance wasnot significant. The density ofnaturally-regenerated woody species inplantations was over three times the usualplanting density in Ethiopia, indicating a highpotential of forest plantations for restoringthe natural forest ecosystems on degraded landsat a comparatively low cost.     

Inhibition of the regeneration of Japanese black pine (<Emphasis Type="Italic">Pinus thunbergii</Emphasis>) by black locust (<Emphasis Type="Italic">Robinia pseudoacacia</Emphasis>) in coastal sand dunes

Pine forests are declining because of pine wilt disease and Robinia pseudoacacia, a nitrogen-fixing species, is dominating coastal forests along the Sea of Japan. We examined the effects of R. pseudoacacia on the regeneration of a native pine species, Pinus thunbergii. Two 200 m² plots were set up at the border of a P. thunbergii and R. pseudoacacia-dominated area in a coastal forest. We conducted monthly censuses of emergence, distribution and survival of pine seedlings in the plots from May 2003 to December 2004. Light intensity and soil properties were also measured to analyze the relationships between the survival of pine seedlings and environmental conditions using the Mantel test and the structural equation model. Pinus thunbergii seedlings emerged in spring–early summer and in late autumn. Survival of pine seedlings in the R. pseudoacacia-dominated subplots was less than half that in the pine-dominated subplots. Survival of pine seedlings emerging in May 2003 was significantly reduced by the lower light intensity and higher soil nitrogen in R. pseudoacacia-dominated subplots. The tendency was the same for seedlings emerging from April to May 2004. We concluded that R. pseudoacacia reduced the intensity of light during the growing season and increased the nitrogen content of soil, which resulted in inhibition of the natural regeneration of P. thunbergii.     

Non-additive Effects of Leaf Litter Mixtures from Robinia pseudoacacia and Ten Tree Species on Soil Properties

ABSTRACT

Litter decomposition plays a critical role in both nutrient cycling and the interspecific relationships between tree species. In this study, leaf litter from Robinia pseudoacacia was mixed with litter from 10 other species, this mixture was then ground and incubated in soil. The soil indexes (i.e., quantity of microbes, activities of enzymes, and chemical properties) were analyzed to study the effects of litter decomposition on soil properties and the interaction between the litters. If mixed separately with Larix principis-rupprechtii or Betula platyphylla, R. pseudoacacia litter exerted synergistic effects on the activities of most enzymes. The mixed litters of R. pseudoacacia and Pinus sylvestris var. mongolica or Ulmus pumila exerted synergistic effects on the soil available P. With regard to the soil properties as a whole, the mixed litters of R. pseudoacacia and B. platyphylla, Pinus tabulaeformis, P. sylvestris var. mongolica, or L. principis-rupprechtii exerted synergistic effects on soil during their decomposition. However, the mixed litters of R. pseudoacacia and Populus simonii, Quercus liaotungensis, U. pumila, or Caragana microphylla did not. This indicated that R. pseudoacacia mixed with the litter of other needle-leaf tree species benefits the development and regeneration of soil and thus can be used for sustainable forest management.     

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.