Hydrogeomorphology Influences Swamp Rabbit Habitat Selection in Bottomland Hardwood Forests

In the last century, bottomland hardwood (BLH) forests throughout the Lower Mississippi Alluvial Valley in the United States declined >80% and have been degraded because of habitat loss, fragmentation, and altered hydrology. To better understand how current conditions in BLH forest systems influence wildlife and to better manage land use and vegetation, we characterized winter (Dec–Mar) multi-scale habitat selection of 75 radio-marked swamp rabbits (Sylvilagus aquaticus) based on 850 locations in southern Illinois, USA, during 2010–2016. We investigated habitat selection by fitting resource selection functions with generalized linear mixed models based on Euclidean distances (km) to 8 cover types that described hydrogeomorphic conditions. At the second-order scale of selection (home range selection), swamp rabbits were closer to deciduous forest and low-elevation BLH and farther from agriculture, permanent water, shallow BLH, and woody wetland. At the third-order scale of selection (habitat selection within the home range), swamp rabbits selected areas closer to deciduous forest, low BLH, and shallow BLH, and farther from woody wetlands. For the swamp rabbit in Illinois, a BLH specialist at the northern extent of their range, habitat selection is limited to available terrestrial habitat that provides vegetation for food and hiding cover within linear and flood-prone BLH corridors surrounded by agricultural cover types that are largely unsuitable as habitat. Because hydrologic conditions are spatially and temporally dynamic, wildlife managers should focus on providing diverse habitat conditions across elevations that ensure the continuous availability of terrestrial habitat regardless of water level and flooding extent across the BLH landscape. Further reforestation efforts in BLH ecosystems should target current agricultural land on higher elevations adjacent to characteristically flood-prone forest remnants that escaped agricultural clearing due to frequent flooding. © 2021 The Wildlife Society.     

Temporal variation in the woody understory of an old-growth Fagus-Acer forest and implications for overstory recruitment

Abstract. Changes in woody vegetation were examined over eight years, using a 1.05-ha permanent plot in which the location of every shrub and tree > 1m height was mapped. There was little change in the overstory vegetation, as expected for an old-growth forest. Much greater change occurred in the understory, primarily related to a 40 % increase in density. Differences occurred among species in the under-story, as Acer saccharum and Prunus serotina increased and Fraxinus americana and Fagus grandifolia decreased. Canopy gap dynamics are implicated in differences among species in the establishment and growth of individuals in the understory and their recruitment into the overstory. It is concluded that because understory is temporally variable, overstory recruitment from the understory may take different courses at different times in the same forest.     

Water relations of Quercus palustris: field measurements on an experimentally flooded stand

Summary Field measurements of water relations were used to examine the response of mature Quercus palustris (20 m tall), a flood tolerant tree, to experimental flooding treatments. Flooding treatments included: 1) flooding in the dormant season, 2) short-term flooding in the growing season and 3) long term (<2 years) continuous flooding. Dormant season flooding had no apparent effect on phenology or physiology during the following growing season. Short term flooding in the growing season caused immediate but reversible stomatal closure without significant development of water stresses. Within 10 days after the growing season flooding treatment water relations measurements were not significantly different from pre-treatment measurements or controls. Although no significant differences in water relations were found in growing season measurements on continuously flooded trees, continuous flooding reduced reproductive fitness and caused premature autumn coloration and leafabscission.Unlike upland Quercus species, Quercus palustris does not show evidence of water limitation late in the growing season. Stomatal conductance increased early in the growing season but showed no clear mid- to late-growing season trends. No evidence of mid-day stomatal closure was found throughout the growing season. Stomatal conductance was correlated to both xylem pressure potential and photosynthetically active radiation. Combined osmotic and matric potentials decreased to-2.43 MPa by Julian day 140 and remained constant throughout the growing season. Predawn xylem pressure potentials exceeded-0.4 MPa throughout the growing season.This paper is Scientific Paper SP 6421 of the College of Agriculture Research Center, Washington State University. Work was conducted under Project 0549, ARC-WSU and project 169, 170, 183 of the Missouri Agricultural Experiment Station, University of Missouri. The author acknowledges the support of the Missouri Department of Conservation and the U.S. Fish and Wildlife Service. James Ware and Leigh Fredrickson provided assistance and support without which the project would not have been possible     

COMPARISON OF THE VEGETATION OF THREE LOUISIANA SWAMP SITES WITH DIFFERENT FLOODING REGIMES

Net primary productivity and community structure were studied at three adjacent swamp sites that were all once subject to seasonal flooding. Because of man-related activities one is now permanently flooded, and a second is managed on a summer-fall dry period and a winter-spring flooded period. In the permanently flooded area such flood-intolerant trees as green ash are dying while shrubs and aquatic plants are increasing. Competition among overstory trees is reduced, enhancing growth of existing baldcypress and water tupelo trees. Total areal production, however, is low (887 g/m²/yr) because of fewer trees. Productivity in the managed area is high (1,780 g/m²/yr). The baldcypress-water tupelo community is being replaced by a swamp maple-ash community typical of bottomlands. In the natural swamp forest, baldcypress and water tupelo have remained dominant. Canopy closure has limited understory and aquatic growth but has also increased competition among overstory trees, resulting in a slow increase in basal area. Productivity is about 1,166 g/m²/yr. It is suggested that the flooding regime is an important controlling factor in each of the areas.     

Overstory vegetation and successional trends of Land Between The Lakes,USA

133 stands from the uplands of Land Between The Lakes (LBL) were sampled. Stand data were ordinated to identify clinal patterns in the overstory vegetation. A coeno-cline, accounting for 52% of the species variance, was developed from the first ordination axis. Classification of stands identified nine dominance types: Pinus echinata-Quercus spp., Q. prinus, Q. prinus-Q. alba, Q. stellata-Q. alba, Q. velutina-Q. alba, Q. alba, Q. alba-Q. rubra, Acer saccharum-Q. alba-Fagus grandifolia sadF. grandifolia-A. saccharum. Compositionally-stable (climax) and unstable (successional) stands were analyzed separately. Trajectories of unstable stands include the succession of P. echinata to Quercus-dominated communities on xeric sites and of Quercus types to more mesophytic communities (e.g. A. saccharum and F. grandifolia) onmesic sites. These shifts indicate the nature of the future climax overstory composition at LBL. The first axis DCCA ordination for climax stands accounted for 45% of the vegetation variance. Factor Analysis identified two significant suites of variables related to vegetation: soil chemistry/soil moisture and soil texture. Specific variables related to variation in the climax vegetation include soil pH, calcium and magnesium, effective soil depth, available water capacity, percent rock in the soil, slope position, aspect, elevation and distance to the opposing slope. Modal positions of species along the direct gradient ordination were similar in rank order to those derived from the indirect gradient ordination of all stands. The latter ordination represents the present forest vegetation pattern at LBL. Climax stands showed a shift in modes of Quercus spp. toward the xeric end of the gradient and increases in habitat width of mesophytic species. Following a reduction in disturbance, species appear to adjust to changes in competition and available habitat. That is, the dimensions of realized niches (i.e. habitat widths) change with succession. The climax vegetation at LBL appears to show centrifugal organization.     

Relationships of overstory to understory cover variables in a Ponderosa pine/Gambel oak ecosystem

Many studies have indicated relationships between individual species, but none have related combinations of overstory variables to understory herbaceous vegetation in a Ponderosa pine/Gambel oak ecosystem. Our objective was to determine not only the general relationships between the two sets of variables, but also identify the hyghest contributing variables. We used canonical correlation analysis to relate overstory variables (canopy cover, basal cover and density) to herbaceous vegetation cover variables. Canopy, basal, and ground cover were measured by the line intercept method using a 12.2 m tape as a sample unit. Tree density was measured by the Point-Center-Quarter method. The analysis was made with selected overstory variables and 5 understory herbaceous cover variables. This analysis revealed a significant canonical correlation between the two canonical variables (r=0.69). The analysis showed that among herbaceous cover variables, Oregon grape, Kentucky bluegrass, sedge, and foxtail barley; and among overstory variables, the density and the basal cover of Ponderosa pine indicated the highest positive contribution to the correlation of the two linear combinations while the density and canopy of Gambel oak negatively affected the canonical correlation.     

A coenocline of the high-ranked syntaxa of ruderal vegetation

Two sets comprising 1419 and 1350 phytosociological relevés of ruderal vegetation classified into 9 and 7 orders, respectively, from an area in the Podunajská ní zina Lowland, western Slovakia were ordinated using correspondence and detrended correspondence analyses. The paper describes a coenocline of the high-ranked syntaxa contained in the data sets, and discusses some issues of the classification of the syntaxa involved.The Bidentetalia and Potentillo-Polygonetalia should be considered a special category of synanthropic vegetation as habitat moisture (flooding and waterlogging) play the controlling role in the formation of structural and dynamical patterns within these communities. The latter factor complex is responsible for the clear separation of these orders from the other syntaxa included in the ordinations. The coenocline of terrestrial ruderal vegetation units has the following sequence along the CA axis 1: Poo-Polygonetalia, Sisymbrietalia, Eragostrietalia, Onopordetalia, Agropyretalia repentis, Artemisietalia vulgaris and Glechometalia hederaceae.Various complexes of soil factors and anthropogenic disturbance are operational along the concatenation (a series of portions composing the coenocline). The factor complex includes soil compaction, trampling disturbance, nutrient status, soil texture and moisture, and solar irradiation. Step-by-step ordination and interpretation of concatenated portions of the coenocline proved to be useful in the analysis of complex data sets.Abbreviations CA = correspondence analysis - DCA = detrended correspondence analysis - PCA = principal components analysis     

Community Comparison and Determinant Analysis of Understory Vegetation in Six Plantations in South China

Plantations cover large areas in many countries, and the enhancement of plantation biodiversity is an increasingly important ecological concern. Many studies have demonstrated that overstory composition is important because it influences understory regeneration. To compare the understory vegetation and analyze its determinant factors, six typical plantations in South China were investigated: Acacia mangium plantation, Schima superba plantation, Eucalyptus citriodora plantation, E. exserta plantation, mixed‐coniferous plantation, and mixed native species plantation. The results show that native species plantations shaded out more grasses and herbs than exotic species plantations, mixed‐species plantations recruited more understory species than monoculture plantations, the leguminous species plantation had higher soil nitrogen than nonleguminous species plantations, and understory vegetation in the mixed‐coniferous plantation was similar to that of mixed, native broadleaf species plantation. Although light is the crucial environmental factor affecting the understory community and diversity among the 14 measured factors, other environmental variables such as soil nutrients and soil moisture are also important.     

Response to CO2 enrichment of understory vegetation in the shade of forests

Responses of forest ecosystems to increased atmospheric CO₂ concentration have been studied in few free‐air CO₂ enrichment (FACE) experiments during last two decades. Most studies focused principally on the overstory trees with little attention given to understory vegetation. Despite its small contribution to total productivity of an ecosystem, understory vegetation plays an important role in predicting successional dynamics and future plant community composition. Thus, the response of understory vegetation in Pinus taeda plantation at the Duke Forest FACE site after 15–17 years of exposure to elevated CO₂, 6–13 of which with nitrogen (N) amendment, was examined. Aboveground biomass and density of the understory decreased across all treatments with increasing overstory leaf area index (LAI). However, the CO₂ and N treatments had no effect on aboveground biomass, tree density, community composition, and the fraction of shade‐tolerant species. The increases of overstory LAI (~28%) under elevated CO₂ resulted in a reduction of light available to the understory (~18%) sufficient to nullify the expected growth‐enhancing effect of elevated CO₂ on understory vegetation.     

ORDINATION AND CLASSIFICATION OF WESTERN OAK FORESTS IN OKLAHOMA

Ordination and classification techniques were used to analyze patterns of forest vegetation, species diversity, and soil type in the Wichita Mountains Wildlife Refuge of southwestern Oklahoma. Cluster analysis based on tree species produced three general community types: 1) Quercus stellata-Q. marilandica forests; 2) Q. stellata forests; and 3) mesophytic forests. A polar ordination produced a gradient of vegetation that corresponded to a moisture gradient. Many high diversity forests were located on loamy drainageway soils or north facing slopes. Tree species diversity (H') was inversely related to the importance of Quercus stellata. Cluster analysis based on species composition of the tree seedlings produced four general community types: 1) Q. marilandica type; 2) Q. marilandica-Q. stellata-Juniperus virginiana type; 3) Ulmus americana-Celtis reticulata-Bumelia lanuginosa type; and 4) Acer saccharum type. The third seedling type occurred almost exclusively on loamy drainageway soils. There was no relationship between stand location on the first axis of the tree ordination and the first axis of the seedling ordination suggesting that trees and seedlings respond differently along the moisture gradient.     

Ordination of simulated complex forest succession: A new test of ordination methods

A model of a 1/12th ha forest stand, FORET, generated 10 000 years of simulated species succession. Approximately the first third of these results were analyzed by principal component analysis as if they were collected field data to give the trajectory of the community particle in a collapsed species space. The ordination axis orientation was performed on a dispersion matrix and correlation matrix between species. In both cases, however, the eigen vectors were applied to the data matrix which had not been transformed to unit species variance. This facilitated comparison of species dispersion and correlation structure; it emerged they were very different. Correlation structure gave large weights to understory species while dispersion emphasized the dominant overstory species. This implies a decomposition of simulated stand behavior into overstory and understory, even though such decomposition was not formally built into the model. This decomposition would seem to pertain to real vegetation. Principal component analysis was able to express insightful differences between data structure with and without the unit variance transformation implicit in the correlation matrix. This flexibility of the ordination method proved valuable in uncovering unsuspected ordering principles in the model. Complex simulated data allow the ordination technique to demonstrate its capacity to generate new hypotheses, which hypotheses can then be simply validated by a return to the structure of the model but with the hindsight of the analysis. The generation of new hypotheses is not possible if the simulation is of a simple coenocline; on the other hand, ordination of test field data does not allow the simple validation of new hypotheses, for in the field there is not a defined algorithm to which the researcher can return.     

Deer Browsing,Forest Edge Effects,and Vegetation Dynamics Following Bottomland Forest Restoration

Vegetation composition and forest stand development are frequently mediated by browsing herbivores. These relationships have received little attention in a forest restoration context even though White‐tailed deer (Odocoileus virginianus) is likely to influence these agriculture, forest, and restored ecosystem mosaic landscapes. Tree species composition, herbaceous vegetation, and deer browsing patterns were assessed 5 and 7 years following bottomland hardwood forest restoration on a 526‐ha site in the Cache River watershed in southern Illinois, United States. Light‐seeded tree species (Fraxinus pennsylvanica, Acer negundo, Liquidambar styraciflua, and Platanus occidentalis) of volunteer origin dominated the woody vegetation component, with especially high stocking and density near existing forest cover and potential seed sources. At more distant locations, presumably planted Quercus spp. were more likely to dominate and were the only tree species found in 15% of plots in year 7. Quercus stocking increased over the course of the study, constituting 7% of trees during year 7. Deer herbivory was associated with reduced stem height and disproportionately impacted seedlings of Quercus palustris and Celtis spp. Our results suggest that deer browsing influences forest stand composition and density as a function of distance from the nearest forest edge. Herbaceous vegetation had little impact on early stand development. Continued spread of the exotic and invasive Lonicera japonica and potential mortality of F. pennsylvanica due to an anticipated Emerald ash borer (Agrilus planipennis) epidemic, combined with low stand density and delayed canopy closure, may result in persistent overstory gaps and compromise long‐term restoration success.     

Contributions of Understory and/or Overstory Vegetations to Soil Microbial PLFA and Nematode Diversities in Eucalyptus Monocultures

Ecological interactions between aboveground and belowground biodiversity have received many attentions in the recent decades. Although soil biodiversity declined with the decrease of plant diversity, many previous studies found plant species identities were more important than plant diversity in controlling soil biodiversity. This study focused on the responses of soil biodiversity to the altering of plant functional groups, namely overstory and understory vegetations, rather than plant diversity gradient. We conducted an experiment by removing overstory and/or understory vegetation to compare their effects on soil microbial phospholipid fatty acid (PLFA) and nematode diversities in eucalyptus monocultures. Our results indicated that both overstory and understory vegetations could affect soil microbial PLFA and nematode diversities, which manifested as the decrease in Shannon–Wiener diversity index (H′) and Pielou evenness index (J) and the increase in Simpson dominance index (λ) after vegetation removal. Soil microclimate change explained part of variance of soil biodiversity indices. Both overstory and understory vegetations positively correlated with soil microbial PLFA and nematode diversities. In addition, the alteration of soil biodiversity might be due to a mixing effect of bottom-up control and soil microclimate change after vegetation removal in the studied plantations. Given the studied ecosystem is common in humid subtropical and tropical region of the world, our findings might have great potential to extrapolate to large scales and could be conducive to ecosystem management and service.     

Overstory Community Composition and Elevated Atmospheric CO2 and O3 Modify Understory Biomass Production and Nitrogen Acquisition

Elevated atmospheric CO₂ and O₃ have the potential to affect the primary productivity of the forest overstory, but little attention has been given to potential responses of understory vegetation. Our objective was to document the effects of elevated atmospheric CO₂ and O₃ on understory species composition and biomass and to quantify nitrogen (N) acquisition by the understory vegetation. The research took place at the aspen free-air CO₂ and O₃ enrichment (FACE) experiment, which has four treatments (control, elevated CO₂, elevated O₃, and elevated CO₂+O₃) and three tree communities: aspen, aspen/birch, and aspen/maple. In June 2003, each FACE ring was uniformly labeled with 15N applied as NH₄Cl. Understory biomass was harvested in June of 2004 for productivity, N, and 15N measurements, and photosynthetically active radiation (PAR) was measured below the canopy. The understory was divided into five species groups, which dominate in this young aggrading forest: Taraxacum officinale (dandelion), Solidago sp. (goldenrod), Trifolium repens and T. pretense (clover), various species from the Poaceae family (grass), and composited minor components (CMC). Understory species composition, total and individual species biomass, N content, and 15N recovery showed overstory community effects, but the direct effects of treatments was masked by the high variability of these data. Total understory biomass increased with increasing light, and thus was greatest under the open canopy of the aspen/maple community, as well as the more open canopy of the elevated O₃ treatments. Species were different from one another in terms of 15N recovery, with virtually no 15N recovered in clover and the greatest amount recovered in dandelion. Thus, understory species composition and biomass appear to be driven by the structure of the overstory community, which is determined by the tree species present and their response to the treatments. However, N acquisition by the understory does not appear to be affected by either the overstory community or the treatments at this point.     

Variation in radial growth responses to drought among species, site, and canopy strata

 Radial growth responses to drought were examined in the tree-ring records of six species growing within two locations of differing land-use history and soil moisture characteristics, and in overstory and understory canopy positions in northern Virginia. Tree species experienced differential ring-width reductions during or immediately following four severe drought periods occurring from 1930 to 1965 and were influenced by climatic variables including annual and summer temperatures, annual precipitation, and annual Palmer Drought Severity Index. Relative growth comparisons averaged across species before and after drought years indicated that understory trees on dry-mesic sites grew 11% faster after drought compared to pre-drought rates while mesic site trees in both canopy positions grew approximately 4% slower. Superposed epoch analysis indicated that Liriodendron tulipifera growing on mesic sites experienced greater ring-width reductions associated with drought than co-occurring, more drought-tolerant Quercus alba and Q. velutina. On dry-mesic sites, L. tulipifera also experienced greatly reduced growth as a result of drought but exhibited significant growth increases following individual drought events. Quercus alba was the only species that exhibited a consistent, significant ring-width decrease associated with all droughts on dry-mesic sites. In contrast, Pinus virginiana was least impacted by drought on dry-mesic sites but was much more impacted by drought on mesic sites, indicating a drought×site interaction for this species. Overstory Carya glabra and Q. alba experienced larger growth decreases during drought on dry-mesic versus mesic sites. Understory tree growth reductions did not differ between site types but were often significantly larger than overstory responses of the same species on mesic sites. Following drought, most trees exhibited growth reductions lasting 2–3 years, although several species experienced reductions lasting up to 6 years. The results of this study suggest that tree rings represent an important long-term proxy for leaf-level ecophysiological measurements of growth responses to drought periods. Received: 31 July 1996 / Accepted: 16 April 1997     

Plant Recruitment in Early Development Stages on Rehabilitated Quarries in Hong Kong

To understand vegetation development during the ecological succession of rehabilitated quarries, floristic composition and structure were evaluated at different restoration phases on three quarries in Hong Kong that were planted with exotic woody species over the course of 2–14 years. A total of 113 species, of which 82 were woody species, were recorded. Exotic species dominated the overstory, and species number, richness, and diversity increased with age. Some light‐demanding early successional species were becoming dominant in the overstory vegetation at the older phases of revegetation. These species could be potential candidates for early enrichment planting with Acacia spp. Common secondary forest species occurred naturally in the understory vegetation, and were more abundant and dominant after 10 years of ecological development. The most successful restorations were on scree slopes using leguminous Acacia spp. as nurse species.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

The Effects of Intermittent Flooding on Seedlings of Three Forest Species

Under greenhouse conditions, seedlings of three forest species, baldcypress (Taxodium distichum), nuttall oak (Quercus nuttallii), and swamp chestnut oak (Quercus michauxii) were subjected to an intermittent flooding and subsequent physiological and growth responses to such conditions were evaluated. Baldcypress showed no significant reductions in stomatal conductance (g _s) or net photosynthetic rate (P _N) in response to flood pulses. In nuttall oak seedlings g _s and P _N were significantly decreased during periods of inundation, but recovered rapidly following drainage. In contrast, in swamp chestnut oak g _s was reduced by 71.8 % while P _N was reduced by 57.2 % compared to controls. Baldcypress displayed no significant changes in total mass while oak species had significantly lower leaf and total mass compared to their respective controls. Thus baldcypress and nuttall oak showed superior performance under frequent intermittent flooding regimes due to several factors including the ability for rapid recovery of gas exchange soon after soil was drained. In contrast, swamp chestnut oak seedlings failed to resume gas exchange functions after the removal of flooding.     

Effect of fire intensity on understory composition and diversity in aKalmia-dominated oak forest, New England, USA

This study investigates the understory dynamics of two mixed-oak stands following fire of varying intensity. Composition and diversity of woody and herbaceous species in the understory were measured in two stands 7–8 years after a prescribed burn. On both sites, unburned areas, low-intensity fire areas, and areas where the overstory had been severely damaged were measured. Patterns of species presence and absence following fire were consistent with an initial-floristics model. Most species increased in density and frequency following fire; onlyAralia nudicaulis andQuercus alba showed statistically significant decreases in density. Overall, 29 species increased in density following fire, while 8 declined; 29 species increased in frequency, while 6 declined. However, diversity and equitability measures were depressed on the moderately burned sites, due to rapid regrowth ofKalmia latifolia. Ordination using binary discriminant analysis suggested species responded individualistically to both burning and site variation.     

Response of overstory and understory vegetation 37 years after prescribed burning in an aspen-dominated forest in northern Minnesota,USA – A case study

Many studies have examined short-term changes in understory vegetation following prescribed burning. However, knowledge concerning longer term effects on both forest understory and overstory vegetation is lacking. This investigation was initiated to examine changes in understory (herbaceous and shrub) and overstory species composition almost four decades after logging and prescribed burning at the Pike Bay Experimental Forest in Minnesota. The experiment was established in 1964 with a randomized block design with four treatments: control (c); burned in spring 1967 (S0); burned in spring 1967 + repeat burn spring 1969 (S2); and burned in spring 1967 + repeat burn fall 1970 (F4). Overstory and understory species diversity indices and richness varied within and among treatments but were not strongly or consistently affected by the treatments. Multivariate analyses (multi-response block permutation procedures and non-metric multidimensional scaling) reveal some lingering effects of burning intensity and seasonal variation as well as some compositional differentiation among treatments, but only in the herb layer. In this environment, the effects of two repeated burnings (fire) have essentially disappeared for overstory and understory species diversity and community composition and have failed to convert an aspen-dominated stand to a coniferous stand (an original goal of the study).