Mechanical thinning to restore oak savanna promoted predator assemblages of gypsy moth pupae in Michigan

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Establishment and Control of Hay-scented Fern: A Native Invasive Species

Hay-scented fern (Dennstaedtia punctilobula (Michx.) Moore) is a native forest understory species that behaves as an invasive plant under certain conditions. Previous work has shown that both increased understory light intensity following overstory thinning and removal of competing plants by herbivores can lead to accelerated growth of hay-scented fern, allowing it to develop dense, nearly monospecific understories that inhibit tree seedling regeneration. To study the relationship between these two factors, we sampled 28 forest stands thinned at different times and subjected to different levels of browsing by white-tailed deer (Odocoileus virginianus), and concluded that more than 15 years of intensive browsing following thinning was necessary for ferns to form closed understory canopies with densities of >90 fronds/m² and canopy heights of 60–80 cm; thinning alone or intensive browsing alone was not sufficient to cause this level of fern invasion. We applied three treatments to dense fern understories to determine the relative importance of the fern canopy and the dense mat of roots, rhizomes, and dead fronds in the inhibition of tree seedling establishment. Results after two years were: (1) complete removal of the organic mat produced a large germination response of woody and herbaceous species; (2) mixing the organic mat into the mineral soil produced an initial germination response but poor seedling survivorship, as the fern canopy regrew to near pretreatment density; (3) repeatedly clipping the ferns for two years without disturbing the organic mat resulted in a lower germination response than the removal treatment, but rapid growth of seedlings.     

Differences in Spider Community Composition among Adjacent Sites during Initial Stages of Oak Woodland Restoration

Fire suppression has altered the uplands of northern Mississippi (U.S.A.). Once blanketed by open oak woodlands, this region is now experiencing mesophytic tree invasion, canopy closure, reduced oak regeneration, and herbaceous understory loss. In an attempt to reestablish historical conditions, experimental restoration was initiated through thinning and burning treatments. Our study, part of a comprehensive monitoring effort, is the first to examine the impact of oak woodland restoration on the spider community and associated habitat structure. Samples measuring a variety of environmental variables and utilizing an array of spider collecting techniques were taken within four habitats located at the restoration site: fire‐suppressed forest, moderately treated forest, intensely treated forest, and old field. Two main conclusions resulted from this study. (1) Open‐habitat specialists responded positively to increased canopy openness regardless of the availability of herbaceous vegetation. (2) Woodland restoration increased spider diversity, perhaps through the formation of diverse habitat structure and/or by altering species dominance patterns. A rise in open‐habitat specialist diversity was observed as treatment intensity increased, with no compensatory reduction in the diversity of forest specialists. What remains to be seen is whether the continued transition to open woodland habitat will result in losses of forest specialist species. More aggressive overstory tree thinning is currently being administered to encourage the growth of herbaceous grasses and forbs, which will permit future tests of a hypothesized decline in forest specialists.     

An Afforestation System for Restoring Bottomland Hardwood Forests: Biomass Accumulation of Nuttall Oak Seedlings Interplanted Beneath Eastern Cottonwood

Bottomland hardwood forests of the southeastern United States have declined in extent since European settlement. Forest restoration activities over the past decade, however, have driven recent changes in land use through an intensified afforestation effort on former agricultural land. This intense afforestation effort, particularly in the Lower Mississippi Alluvial Valley, has generated a demand for alternative afforestation systems that accommodate various landowner objectives through restoration of sustainable forests. We are currently studying an afforestation system that involves initial establishment of the rapidly growing native species eastern cottonwood (Populus deltoides Bartr. ex Marsh.), followed by enrichment of the plantation understory with Nuttall oak (Quercus nuttallii Palm.). In this article, we examine the growth and biomass accumulation by Nuttall oak seedlings to determine whether this species can be established and whether it will develop beneath the cottonwood overstory. After 3 years of growth beneath cottonwood canopies, Nuttall oak seedlings were similar in height (126 cm), but were 20% smaller in root‐collar diameter than seedlings established in open fields. Seedlings established in the open accumulated more than twice the biomass of seedlings growing beneath a cottonwood canopy. However, the relative distribution of accumulated biomass in seedlings did not differ in the two environments. Ten percent of total seedling biomass was maintained in leaf tissue, 42% was maintained in stem tissue, and 48% was maintained in root tissue on open‐grown seedlings and seedlings established in the understory of cottonwood plantations. Though establishment in the more shaded understory environment reduced Nuttall oak growth, seedling function was not limited enough to induce changes in plant morphology. Our results suggest that an afforestation system involving rapid establishment of forest cover with a quick‐growing plantation species, followed by understory enrichment with species of later succession, may provide an alternative method of forest restoration on bottomland hardwood sites and perhaps other sites degraded by agriculture throughout temperate regions.     

Development of Tree Regeneration in Fern-dominated Forest Understories After Reduction of Deer Browsing

Dennstaedtia punctilobula (hay‐scented fern) can act as a native invasive species in forests in eastern North America where prolonged deer browsing occurs in stands with partially open overstory canopies. Ferns dominate the understory with a 60‐cm tall canopy, with little regeneration of native tree species. It has been hypothesized that, once established, ferns may continue to inhibit tree regeneration after deer browsing has been reduced. To test this hypothesis, we documented the pattern of recovery of the tree seedling understory in plantations of Pinus strobus (white pine) and Pinus resinosa (red pine) on the Quabbin Reservation watershed protection forest in central Massachusetts, where after 40 years of intensive deer browsing the deer herd was rapidly reduced through controlled hunting. Dense fern understories occur on nearly 4,000 ha of the predominantly oak–pine forest. Three years after deer herd reduction, stands with the highest density fern cover (77% of plots with>90% cover) had significantly fewer seedlings at least 30 cm in height, compared with stands with lower fern density, and those seedlings consisted almost entirely of Betula lenta (black birch) and white pine. Height growth analysis showed that black birch and white pine grew above the height of the fern canopy in 3 and 6 years, respectively. In contrast, two common species, Fraxinus americana (white ash) and Quercus rubra (red oak), grew beneath the dense fern cover for 5 years with height growth less than 5 cm/yr after the first year. A study of spring phenology indicated that the ability of black birch to grow through the fern canopy might have been due to its early leaf development in spring before the fern canopy was formed, in contrast to oak and ash with delayed leaf development. Thus, the ferns showed differential interference among species with seedling development after reduction of deer browse.     

Forest community structure and composition following containment treatments for the fungal pathogen oak wilt

In temperate oak forests in Ohio, USA, we examined variability in forest communities within containment treatment sites for oak wilt (Bretziella fagacearum), a fungal pathogen lethal to susceptible oak species. Containment treatments included quarantine lines in soil for limiting belowground fungal spread and sanitation cutting of 1–3 mature black oak (Quercus velutina) trees within oak wilt infection patches. At 28 sites, we compared tree structure and understory plant communities across a gradient of 1- to 6-year-old treatments and reference forest (untreated and without evidence of oak wilt). While oak seedlings were abundant, oak saplings (1–10 cm in diameter) were absent. In contrast, many native understory plant community measures were highest in oak wilt treatments. Plant species richness 100 m^?2 doubled in treatments, regardless of age, compared with reference forest. Plant cover increased with treatment age, with 6-year-old treatments exhibiting 5?×?more cover than reference forest. Non-native plants averaged only a small proportion (<?0.12) of cover across treatments and reference forest. Variability in understory communities was mostly predictable using treatment age, tree canopy cover, and geographic location, as 20 of 25 understory measures had at least 72% of their variance modeled. While oak wilt treatments did not facilitate oak regeneration nor many conservation-priority species of open savanna-woodland habitats, the treatments did diversify and increase cover of native understory communities with minimal invasion of non-native plants.

     

Long‐term and interactive effects of different mammalian consumers on growth,survival, and recruitment of dominant tree species

Throughout the world, numerous tree species are reported to be in decline, either due to increased mortality of established trees or reduced recruitment. The situation appears especially acute for oaks, which are dominant features of many landscapes in the northern hemisphere. Although numerous factors have been hypothesized to explain reductions in tree performance, vertebrate herbivores and granivores may serve as important drivers of these changes. Here, using data from 8‐ and 14‐year‐old exclosure experiments, we evaluated the individual and interactive effects of large and small mammalian herbivores on the performance of three widespread oak species in California—coast live oak (Quercus agrifolia), California black oak (Q. kelloggii), and Oregon white oak (Q. garryana). Although impacts varied somewhat by species and experiment, herbivory by black‐tailed deer (Odocoileus hemionus columbianus) reduced the height and survival of juvenile coast live oaks and altered their architecture, as well as reduced the abundance of black oak seedlings, the richness of woody species and the cover of nonoak woody species. Small mammals (Microtus californicus and Peromyscus maniculatus) had even more widespread effects, reducing the abundance of black oak seedlings and the height and cover of all three oak species. We also detected numerous interactions between small mammals and deer, with one herbivore having positive or negative effects on oak abundance and cover when the other herbivore was either present or absent. For example, deer often had negative effects on seedling abundance only when, or even more so when, small mammals were present. In summary, mammalian consumers play crucial roles in limiting oak recruitment by reducing seedling abundance, maintaining trees in stunted states, and preventing them from reaching sapling stages and becoming reproductive. Interactions between large and small mammals can also alter the intensity and direction of their effects on trees.     

Testing the effect of restoration‐focused silviculture on oak regeneration and groundlayer plant communities in urban–exurban oak woodlands

Throughout their global range, oak‐dominated ecosystems have undergone state changes in stand structure and composition. Land managers face an especially acute challenge in restoring oak ecosystems and promoting oak regeneration in urban–exurban areas, where high‐intensity silvicultural treatments are often not feasible. To investigate low‐intensity management alternatives which could be widely applied in urban–exurban forests, a large‐scale adaptive management experiment was implemented in Lake County, IL, in 2012. Five canopy manipulation treatments of varying intensity, timing, and spatial aggregation were replicated across three study areas and oak seedlings were under‐planted into treatment units following management. Responses of understory light environment, shrub and groundlayer plant communities, and survival and growth of underplanted oak seedlings were evaluated. Understory light availability, canopy openness, total groundlayer plant cover, and groundlayer species diversity all differed among treatments. However, although understory light availability was significantly increased by canopy manipulation, groundlayer communities and oak seedling survival and growth did not differ among treatments. High overall seedling survival rates suggest current conditions are amenable to oak regeneration, but long‐term monitoring will be needed to assess the potential for seedlings to transition to the sapling and canopy layers. Early results demonstrate that canopy‐focused silvicultural treatments can affect the understory light environment and, to some degree, groundlayer plant communities. However, underplanting of oak seedlings paired with subcanopy thinning may be sufficient to restore an oak seedling layer, and (when necessary or preferred) canopy manipulation could potentially be deferred until later in the restoration timeline to promote oak recruitment.     

The regeneration of pioneer tree species under browsing pressure of Sika deer in an evergreen oak forest

The effects of Sika deer (Cervus nippon) browsing on the regeneration of pioneer species were studied in relation to canopy gaps in a warm temperate evergreen oak forest in Kasugayama, Nara City. Four study sites, three in canopy gaps and one under a closed canopy, were selected and each divided into fenced and unfenced plots. Under the closed canopy, seedlings of all the pioneer species died irrespective of browsing pressure. However, in the canopy gap sites, seedlings of the pioneer species could establish and grow well. The seedling survival ratio in the fenced plots in the canopy gaps was >60% 1 year after germination. However, in the unfenced plots, only <20% of the seedlings survived 1 year, with all dying within 3 years after germination. Thus, the regeneration of pioneer trees in this forest was strongly inhibited by deer browsing. Successful regeneration of a pioneer,Zanthoxylum ailanthoides, occurred for several years even after two major wind disturbances during the past 90 years. This may be due to less browsing pressure from the deer.     

Bee community responses to a gradient of oak savanna restoration practices

North American Midwestern oak (Quercus spp.) savannas are rare fire‐dependent ecosystems that can support high levels of biodiversity and are the focus of considerable restoration effort due to widespread fire suppression. Due to the predominance of understory forbs in oak savannas, many of which require insect pollination, restoration practices should be evaluated for their potential impacts on pollinator communities. We evaluated bee community responses during the first 2 years of experimental restoration of fire‐suppressed oak savanna in southern Michigan. We used unmanaged references and two different restoration methods (burning only and burning with thinning) to examine the effects of restoration intensity on the abundance, diversity, and functional groups of bees. We found that thinning and burning rapidly increased bee abundance, richness, and Shannon's diversity, relative to unmanaged references, whereas burn‐only restoration largely failed to do so. Thinning and burning also resulted in a distinct bee community after two seasons, while bee communities in burn‐only restoration plots were similar to those from unmanaged references. Differences in bee diversity and community structure between treatments may be due to the influence of restoration on nesting resources, which is reflected in the differential captures of various nesting guilds. Overall, oak savanna restoration by thinning and burning had positive effects on bee diversity, while burning alone only increased bee abundance. We thus illustrate how restoration strategies that typically target plants have broader‐reaching biodiversity benefits. Although restoring savannas through burning alone may eventually shift bee communities, coupling thinning with burning will influence pollinator communities over the shorter term.     

Simulation of the effects of deer browsing on forest dynamics

I introduced forest dynamics to a deterministic herbivore-vegetation model to examine the effects of vertically stratified forest structure on the dynamics of the herbivore–vegetation system, the resilience and stable states of vegetation, and the interactions between deer populations and vegetation. I constructed a model based on data from field studies performed in Hokkaido, northern Japan. Three phases of state were identified for a given deer density: (1) understory vegetation is maintained with a equilibrium proportion of canopy gaps in the absence of deer; (2) if the equilibrium proportion of canopy gaps is greater than that in the equilibrium state in the absence of deer, the understory vegetation can be maintained; and (3) the understory vegetation cannot be maintained. At the boundary between phases 2 and 3, the herbivore population level had discontinuous effects on vegetation. When the deer density was held below the threshold, forest vegetation had resilience to recover to the equilibrium stable state at the given deer density, although the equilibrium canopy gap ratio and vegetation biomass differed with deer density. However, the forest vegetation–herbivore system could not be maintained in a stable state without artificial deer population management if food limitation was the only mechanism to keep the deer population at low levels. The deer population must be kept below the boundary between phases 1 and 2 to maintain the forest regeneration processes. The level cannot be determined by observing the deer population; careful observation of forest regeneration processes is required.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

Herbivore-induced expansion of generalist species as a driver of homogenization in post-disturbance plant communities

We used a controlled experiment to investigate how disturbance scale (canopy gap area) and herbivory influence post-disturbance plant community dynamics. Twenty canopy gaps were installed in a temperate hemlock-hardwood forest during the winter of 2002–2003: seven small gaps (50–150 m²), seven medium gaps (151–250 m²), and six large gaps (251–450 m²). Within each gap, we established 4–12 sample plots (depending on gap size); 1–3 of which were enclosed with wire mesh white-tailed deer (Odocoileus virginianus) exclosures in 2005. Gaps were revisited and intensively sampled in 2007. After five growing seasons, ground-layer plant communities in non-exclosed plots were more similar compositionally than exclosure plots. Non-exclosed plots in small and medium gaps were more similar to non-exclosed plots in large gaps than they were to exclosed plots in their respective gap size class. Shade-tolerant forbs and trees were less common outside exclosures, while generalist species associated with higher understory light levels and exotics were more prevalent outside exclosures. Our results suggest that even in forests with relatively low deer densities (6.5–9.3 deer km⁻²), white-tailed deer herbivory may influence the developmental trajectory of post-disturbance plant communities and be a mechanism for decreasing β-diversity along environmental gradients.     

Dendroecological study of a subalpine fir (<Emphasis Type="Italic">Abies fargesii</Emphasis>) forest in the Qinling Mountains,China

Dendroecological techniques were used to investigate the stand dynamics and the disturbance history of the subalpine fir forest in the Qinling Mountains of Shaanxi Province, China. The results indicated that 68% of the fir trees experienced 1–2 release events for a total of 10–29 (an average of 15.8) years, and 1–2 suppression events for a total of 10–27 (an average of 13.4) years before they reached canopy. Large number of Abies fargesii and Betula albo-sinensis recruitment coincided temporally with larger increases in the ring-width index from the 1830s to 1880s, suggesting occurrence of a major stand-wide disturbance during this time period. Few seedlings and saplings were found in the forest, and there was a dramatic decline in recruitment after 1890, probably because of the intensive cover of understory umbrella bamboo (Fargesia spathacea). Radial growth analyses indicated frequent canopy opening resulting from small-scale disturbances in the forest. Thus, the subalpine fir forest experienced frequent small-scale disturbances and infrequent large-scale disturbances in its developmental history, and these disturbances coupled with the understory umbrella bamboo might have influenced tree growth and species recruitment.     

Passive Restoration of Atlantic Forest Following Pinus taeda Harvesting in Southern Brazil

The understory of exotic tree plantations can have non‐negligible native species richness. Ecological restoration of these sites may include the harvest of trees, depending on the tradeoff between timber income and harvest impacts on biodiversity. This study aimed to investigate how a site can recover from harvest disturbance, by comparing the regeneration of woody species in the understory of two types of 37‐year‐old Pinus taeda plantation (P1 and P2, high and low relative density of pine seedlings in the understory, respectively), with stands that were similar to P2 but subjected to harvest and then abandoned for 15 years (R sites). Secondary forests (SF) were used as references. We sampled three different sites for each stand condition; soil chemical properties, estimations of litter mass, and canopy cover were measured. P1 had low species diversity, and P2 and R had 50 and 46% of SF richness, respectively. The R site contained few pine saplings and was floristically similar to P2; this indicated that 15 years was sufficient for the recovery of plant diversity to near pre‐harvesting levels. Soil fertility was highest in SF and lowest in P1. Thus old plantations of P. taeda with low relative density of pine juveniles can be cost‐effective starting points for restoration. Despite the destructive effects of pine harvest, recovery of native species can occur rapidly. In situations in which clearcutting of pine stands is not planned or possible, modest thinning of P. taeda adults and/or intensive thinning of juveniles could expedite restoration.     

Woodland canopy structure and the light response of juvenile Quercus lobata (Fagaceae)

I investigated competition for light between canopy plants and juvenile valley oaks (Quercus lobata Nee) in a mixed-broadleaf woodland of California's northern Coast Ranges. Canopy effects on understory light supply were separated among the overlying adult valley oak, the adult's woody understory, and neighboring trees and shrubs through a series of light sampling surveys and measurements of the number, size, and spatial distribution of neighboring plants. Light supply in the understory was primarily influenced by neighboring plants, with no detectable effect of the overlying adult valley oak. Light supply in the understory averaged 25% full sun due to a high frequency of canopy gaps and a typically open understory. Seedling response to understory light supply was investigated in an experimental sunfleck gradient (10%, 19%, and 100% full sun). Between 10% and 100% full sun, seedling growth increased by 90% and the shoot:∗∗∗root ratio changed from 1.561 to 0.607. Shade seedlings were also taller and produced fewer, larger, and thinner leaves than seedlings grown in full sun. A field survey of the spatial distribution and crown morphology of saplings and young adults found 1) the distance between young valley oaks and neighboring overstory trees to increase with neighbor size, and 2) crowns of the young oaks to be skewed away from neighbors. Although shading by the canopy was only moderate, canopy effects on understory light supply may restrict juvenile recruitment of valley oak in this woodland.     

Experimental Manipulation of Restoration Barriers in Abandoned Eucalypt Plantations

Expansion of the nature conservation estate in northeastern New South Wales, Australia, has captured weed‐infested timber plantations amid a mosaic of high conservation value lands. We adopted a state‐and‐transition approach to test the hypothesis that restoration barriers restrict the natural regeneration of native species in Eucalyptus grandis plantations infested by Lantana camara in Bongil Bongil National Park, New South Wales. Plantation tree thinning and weed control were applied in factorial combination at three sites (totaling to 4.5 ha). Topsoil chemistry responses to these interventions were attributable to the “ash bed” effect, with temporary increases in topsoil pH_W and nitrate, particularly where canopy reduction was greatest. Other soil changes were minor, indicating that thinning and burning did not risk soil degradation. Plant species richness and functional group representation in the regenerating understorey were improved by the interventions. Regeneration of native potential canopy trees, understorey trees, shrubs and woody climbers, and perennial forbs all increased with canopy retention. Grass cover dominated the regeneration where canopy cover was less than 50%. In the absence of weed control, the cover of introduced shrubs increased with reduction in canopy cover, as did the rate of understorey regeneration generally. These responses indicate that thinning and weed control can reinstate succession, leading to structurally and compositionally diverse forest. Given the abundance of native woody regeneration under retained canopy, the lantana understorey was more important in inhibiting native regeneration. The experimental approach will promote efficient use of resources across the remaining 200 ha of low conservation value plantations in this national park.     

Effects of introduced ungulates on forest understory communities in northern Patagonia are modified by timing and severity of stand mortality

Natural disturbances such as fires, windstorms, floods, and herbivory often act on plant communities, affecting their structure and the abundance and composition of their species. Most research has focused on the effects of single disturbances on plant communities whereas the synergistic effects of several disturbances have received less attention. In this study, we evaluated how timing and severity of tree mortality modified plant use by introduced deer and early post-mortality successional trajectories in northern Patagonian conifer forests. We sampled understory composition and deer use in Austrocedrus chilensis (ciprés de la cordillera) forest stands undergoing varying timing and severity of forest mortality as reconstructed using dendroecological techniques. In addition, we evaluated the effect of fallen logs on plant composition and deer use of plants by monitoring areas of massive dieback where fallen logs had been removed for fire hazard reduction, and nearby control areas not subjected to such removal. Stepwise regression analyses showed that history and severity of tree mortality strongly influence plant composition and deer use of plants. For deer use (with pellet counts and browsing index as response variables), results showed a positive relationship with degree of stand mortality and a negative relationship with cover of fallen logs. Similarly, cover of unpalatable shrub species was explained by canopy mortality history, whereas cover of palatable shrub species was positively associated with severity of canopy mortality. In areas where fallen logs had been removed, pellet counts were six times higher than those in control areas. Though total shrub species cover was similar between log removal and control areas, proportion of unpalatable shrubs increased in areas where fallen logs had been removed. In conclusion, deer use of plants was strongly limited by tall fallen logs, allowing palatable species to establish and grow. Fallen log removal accelerated deer entrance and changed understory composition toward more browse-resistant and unpalatable species. These results underscore the importance of considering the dynamics (timing, severity, and extent) of fallen woody debris influencing understory herbivory and post-disturbance succession. In addition, experimental results underpin the importance of maintaining snags and large woody debris in disturbed landscapes where salvage logging is a routine procedure.     

Restoring Perennial Warm‐Season Grasses as a Means of Reversing Mesophication of Oak Woodlands in Northern Mississippi

Fire suppression has removed an important ecological force previously responsible for shaping many plant communities throughout the world. Upland areas of north‐central Mississippi that have been protected from fire are now closed‐canopy forests including species known to be uncommon as bearing/witness trees in upland portions of the landscape (historically off‐site species) and sparse ground cover vegetation. Anecdotal evidence suggests that warm‐season grasses were prevalent in the understory of these communities, which could have provided more consistent fuel. We corroborate the historic presence of these grasses by looking at their natural co‐occurrence with oak regeneration (a requisite of self‐replacing stands of oaks found historically). Restoration of these communities has typically focused on burning and off‐site tree thinning. Utilizing a restoration experiment implementing these treatments, we found significantly reduced understory leaf litter in treatment areas. To test which variables associated with restoration treatments were most important for the survival of these grasses, we measured the effect of leaf litter removal and its interaction with environmental conditions on the survival of transplanted shoots. Survival of little bluestem increased with decreasing canopy density and decreasing leaf litter. Leaf‐litter removal did not increase survival, nor did it interact with either pre‐treatment leaf litter depth or canopy density. These results show that little bluestem benefits from conditions expected historically: increased light and possibly fire.     

Forest plant community as a driver of soil biodiversity: experimental evidence from collembolan assemblages through large‐scale and long‐term removal of oak canopy trees Quercus petraea

Plant–soil interactions are increasingly recognized to play a major role in terrestrial ecosystems functioning. However, few studies to date have focused on slow dynamic ecosystems such as forests. As they are vertically stratified by multiple vegetation strata, canopy tree removal by thinning operations could alter forest plant community through tree canopy opening. Very little is known about cascading effects on soil biodiversity. We conducted a large‐scale, multi‐site assessment of collembolan assemblage response to long‐term canopy tree removal in sessile oak Quercus petraea temperate forests. A total of 33 experimental plots were studied covering a large gradient of canopy tree basal area, stand age and local abiotic contexts. Collembolan abundance strongly declined with canopy tree removal in early forest successional stage and this was mediated by negative effect of understory plant community composition changes, i.e. shift from moss and forb to tree seedling, fern, shrub and grass species. Negative effect of this composition shift on collembolan species richness was largely offset by positive effect of the increase in understory plant species richness. This gives support to both the plant mass‐ratio and functional diversity hypotheses. Collembolan functional groups had contrasting response patterns, which were mediated by different ecological factors. Epedaphic (r‐strategist) abundance and species richness increased with canopy tree removal in relation with the increase in understory plant species richness. In contrast, euedaphic (K‐strategist) abundance and species richness declined with canopy tree removal in early forest successional stage in relation with changes in understory plant community composition and species richness, as well as microclimatic conditions. Overall, our study provides experimental evidence that forest plant community can be a strong driver of collembolan assemblages. It also emphasizes the role of trees as foundation species of forest ecosystems that can shape soil biodiversity through their regulation of understory plant community and ecosystem abiotic conditions.