Linking dominant Hawaiian tree species to understory development in recovering pastures via impacts on soils and litter

Large areas of tropical forest have been cleared and planted with exotic grass species for use as cattle pasture. These often remain persistent grasslands after grazer removal, which is problematic for restoring native forest communities. It is often hoped that remnant and/or planted trees can jump‐start forest succession; however, there is little mechanistic information on how different canopy species affect community trajectories. To investigate this, I surveyed understory communities, exotic grass biomass, standing litter pools, and soil properties under two dominant canopy trees—Metrosideros polymorpha (‘ōhi‘a) and Acacia koa (koa)—in recovering Hawaiian forests. I then used structural equation models (SEMs) to elucidate direct and indirect effects of trees on native understory. Native understory communities developed under ‘ōhi‘a, which had larger standing litter pools, lower soil nitrogen, and lower exotic grass biomass than koa. This pattern was variable, potentially due to historical site differences and/or distance to intact forest. Koa, in contrast, showed little understory development. Instead, data suggest that increased soil nitrogen under koa leads to high grass biomass that stalls native recruitment. SEMs suggested that indirect effects of trees via litter and soils were as or more important than direct effects for determining native cover. It is suggested that diverse plantings which incorporate species that have high carbon to nitrogen ratios may help ameliorate the negative indirect effects of koa on natural understory regeneration.     

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.     

Understory vegetation response to mechanical mastication and other fuels treatments in a ponderosa pine forest

Questions: What influence does mechanical mastication and other fuel treatments have on: (1) canopy and forest floor response variables that influence understory plant development; (2) initial understory vegetation cover, diversity, and composition; and (3) shrub and non‐native species density in a second‐growth ponderosa pine forest. Location: Challenge Experimental Forest, northern Sierra Nevada, California, USA. Methods: We compared the effects of mastication only, mastication with supplemental treatments (tilling and prescribed fire), hand removal, and a control on initial understory vegetation response using a randomized complete block experimental design. Each block (n=4) contained all five treatments and understory vegetation was surveyed within 0.04‐ha plots for each treatment. Results: While mastication alone and hand removal dramatically reduced the midstory vegetation, these treatments had little effect on understory richness compared with control. Prescribed fire after mastication increased native species richness by 150% (+6.0 species m²) compared with control. However, this also increased non‐native species richness (+0.8 species m²) and shrub seedling density (+24.7 stems m²). Mastication followed by tilling resulted in increased non‐native forb density (+0.7 stems m²). Conclusions: Mechanical mastication and hand removal treatments aided in reducing midstory fuels but did not increase understory plant diversity. The subsequent treatment of prescribed burning not only further reduced fire hazard, but also exposed mineral soil, which likely promoted native plant diversity. Some potential drawbacks to this treatment include an increase of non‐native species and stimulation of shrub seed germination, which could alter ecosystem functions and compromise fire hazard reduction in the long‐term.     

Leaf litter and irrigation can increase seed germination and early seedling survival of the recalcitrant‐seeded tree Beilschmiedia miersii

Although regeneration of recalcitrant‐seeded tree species can be affected by prolonged drought, especially in Mediterranean regions, little is known about the response of such species to varying site conditions. A field experiment was performed to determine the effect of irrigation and leaf litter cover on seed germination and early seedling survival of the vulnerable recalcitrant‐seeded tree Beilschmiedia miersii (Lauraceae). Two levels of irrigation (non‐irrigated and irrigated units) and three levels of leaf litter depth (0, 5 and 12 cm) were applied to 72 groups of 30 seeds along a wet ravine of the Coastal Range of Central Chile, equally distributed across sites with different levels of canopy cover. Seed germination was significantly increased by irrigation only under closed‐canopy cover, and by leaf litter cover (>5 cm) under all canopy conditions. The effect of leaf litter on germination increased with canopy openness, while the effect of irrigation did not show any tendency. Meanwhile, early seedling survival was significantly increased by irrigation under intermediate canopy cover, and by leaf litter (>5 cm) under closed‐canopy cover. As a result of its overall positive effect on germination, leaf litter should be maintained within B. miersii communities, particularly under intermediate to closed‐canopy conditions, where it can also increase early seedling survival, and both seed germination and early seedling survival might be increased through additional water inputs. The presence of leaf litter might help retaining such inputs, prolonging their effect on regeneration of B. miersii communities. We see this as a baseline assessment of regeneration and persistence that needs further testing on species with similar traits, given the expected increase in the frequency and length of dry periods into the future.     

What potential is there for regeneration of native species from the soil seed bank in Coast Tea Tree‐dominated scrub?

Shrub encroachment generally causes the loss of native species in herbaceous‐dominated communities. The ability of the original ecosystem to return to its pre‐encroachment state (i.e. its ecological resilience) will be partially contingent on the capacity of these species to regenerate from soil‐stored seed. Coast Tea Tree (Leptospermum laevigatum) has formed a dense scrub in many areas previously dominated by grassy woodland, and hence, managers need guidance about the effectiveness of strategies designed to recover the pre‐encroachment vegetation. In this context, we ask: what is the potential of species stored in the soil seed bank to return following Tea Tree removal? A germination experiment was undertaken using soil collected from dense stands of Tea Tree that had been long established. Heat/smoke was applied to soils to simulate the effects of a fire on the soil seed bank, while leaf litter treatments were used to mimic both undisturbed stands and stands where shrubs have been slashed where litter creates a physical barrier to emergence. We found the soil seed bank was dominated by exotic forbs (83% of all germinants) and contained few grasses. Heat and smoke decreased total species density but increased species diversity through the suppression of common exotics. Our data suggest that slashing would result in germination being dominated by exotic flora, but using fire would likely reduce that dominance. However, we conclude that recovery by much of the original flora after site occupation by Coast Tea Tree may be contingent on mechanisms other than soil‐stored seeds.     

Land‐use history,historical connectivity,and land management interact to determine longleaf pine woodland understory richness and composition

Restoration and management activities targeted at recovering biodiversity can lead to unexpected results. In part, this is due to a lack of understanding of how site‐level characteristics, landscape factors, and land‐use history interact with restoration and management practices to determine patterns of diversity. For plants, such factors may be particularly important since plant populations often exhibit lagged responses to habitat loss and degradation. Here, we assess the importance of site‐level, landscape, and historical effects for understory plant species richness and composition across a set of 40 longleaf pine Pinus palustris woodlands undergoing restoration for the federally endangered red‐cockaded woodpecker in the southeastern United States. Land‐use history had an overarching effect on richness and composition. Relative to historically forested sites, sites with agricultural histories (i.e. former pastures or cultivated fields) supported lower species richness and an altered species composition due to fewer upland longleaf pine woodland community members. Landscape effects did not influence the total number of species in either historically forested or post‐agricultural sites; however, understory species composition was affected by historical connectivity, but only for post‐agricultural sites. The influences of management and restoration activities were only apparent once land‐use history was accounted for. Prescribed burning and mechanical overstory thinning were key drivers of understory composition and promoted understory richness in post‐agricultural sites. In historically forested sites these activities had no impact on richness and only prescribed fire influenced composition. Our findings reveal complex interplays between site‐level, landscape, and historical effects, suggest fundamentally different controls over plant communities in longleaf pine woodlands with varying land‐use history, and underscore the importance of considering land‐use history and landscape effects during restoration.     

Canopy thinning,not agricultural history,determines early responses of wild bees to longleaf pine savanna restoration

Longleaf pine savannas are highly threatened, fire‐maintained ecosystems unique to the southeastern United States. Fire suppression and conversion to agriculture have strongly affected this ecosystem, altering overstory canopies, understory plant communities, and animal populations. Tree thinning to reinstate open canopies can benefit understory plant diversity, but effects on animal communities are less well understood. Moreover, agricultural land‐use legacies can have long‐lasting impacts on plant communities, but their effects on animal communities either alone or through interactions with restoration are unclear. Resolving these impacts is important due to the conservation potential of fire‐suppressed and post‐agricultural longleaf savannas. We evaluated how historical agricultural land use and canopy thinning affect the diversity and abundance of wild bees in longleaf pine savannas. We employed a replicated, large‐scale factorial block experiment in South Carolina, where canopy thinning was applied to longleaf pine savannas that were either post‐agricultural or remnant (no agricultural history). Bees were sampled using elevated bee bowls. In the second growing season after restoration, thinned plots supported a greater bee abundance and bee community richness. Additionally, restored plots had altered wild bee community composition when compared to unthinned plots, indicating that reduction of canopy cover by the thinning treatment best predicted wild bee diversity and composition. Conversely, we found little evidence for differences between sites with or without historical agricultural land use. Some abundant Lasioglossum species were the most sensitive to habitat changes. Our results highlight how restoration practices that reduce canopy cover in fire‐suppressed savannas can have rapid benefits for wild bee communities.     

Biogeochemical and Microbial Legacies of Non‐Native Grasses Can Affect Restoration Success

The restoration of disturbed ecosystems is challenging and often unsuccessful, particularly when non‐native plants are abundant. Ecosystem restoration may be hindered by the effects of non‐native plants on soil biogeochemical characteristics and microbial communities that persist even after plants are removed. To examine the importance of soil legacy effects, we used experimental restorations of Florida shrubland habitat that had been degraded by the introduction of non‐native grasses coupled with either mechanical disturbance or pasture conversion. We removed non‐native grasses and inoculated soils with native microbial communities at each degraded site, then examined how habitat structure, soil nitrogen, soil microbial abundances, and native seed germination responded over two years compared to undisturbed native sites. Grass removal treatments effectively restored some aspects of native habitat structure, including decreased exotic grass cover, increased bare ground, and reduced litter cover. Soil fungal abundance was also somewhat restored by grass removals, but soil algal abundance was unaffected. In addition, grass removal and microbial inoculation improved seed germination rates in degraded sites, but these remained quite low compared to native sites. High soil nitrogen persisted throughout the experiment regardless of treatment. Many treatment effects were site‐specific, however, with legacies in the more degraded vegetation type tending to be more difficult to overcome. These results support the need for context‐dependent restoration approaches and suggest that the degree of soil legacy effects may be a good indicator of restoration potential.     

Effects of longleaf pine planting density and other factors on stand structure and associated wildlife habitat

The primary objective of many longleaf pine (Pinus palustris) restoration programs is to enhance or restore habitat for wildlife dependent on herbaceous plant communities. Because herbaceous cover is inversely related to canopy cover, restoration programs often place restrictions on longleaf pine planting density. However, the influence of planting density on understory plant communities has been inadequately evaluated. Therefore, we initiated a study to examine the relative influences of planting density and other factors on overall understory composition and forage availability for white‐tailed deer (Odocoileus virginianus) and northern bobwhite (Colinus virginianus) in nine longleaf pine stands throughout the Coastal Plain of Alabama during 2017–2018. We found that coverage of herbaceous plants decreased 3.5%, coverage of woody plants decreased 2.4%, and coverage of northern bobwhite forage plants decreased 1.9% for each 1 m²/ha increase in longleaf pine basal area. However, planting density was not a significant predictor of current basal area, nor coverage of any functional group of plants we examined, likely because current longleaf pine density averaged only 46% (range = 30–64%) of seedling planting density. We did not detect an effect of prescribed fire on stand condition or understory plant communities, likely due to variability in fire timing and frequency. Our findings related to planting density were likely a function of low longleaf pine survival, which is not uncommon. Because of this and the inherent variability in growth rates for young longleaf pine stands, restoration programs should consider placing greater emphasis on post‐planting monitoring and management than planting density.     

Post-fire recovery of ant communities in Submediterranean Pinus nigra forests

This study analyzes the variations in the structure and composition of ant communities in burned Pinus nigra forests in central Catalonia (NE Spain). Pinus nigra forests do not recover after fire, changing to shrublands and oak coppices. For this reason, we suggest that ant communities of burned P. nigra forests will change after fire, because the post‐fire scenario, in particular with the increase of open areas, is different to the unburned one, and more favourable for some species than for others. In four locations previously occupied by P. nigra forests where different fires occurred 1, 5, 13 and 19 yr before the sampling, we sampled the structure and composition of ant communities with pitfall traps, tree traps and net sweeping in unburned plots and in plots affected by canopy and understory fire. The results obtained suggest that canopy and understory fire had little effect on the structure of ant communities. Thus, many variables concerning ant communities were not modified either by fire type (understory or canopy fire) or by time since fire. However, a number of particular species were affected, either positively or negatively, by canopy fire: three species characteristic of forest habitats decreased after fire, while eight species characteristic of open habitats increased in areas affected by canopy fire, especially in the first few years after fire. These differences in ant community composition between burned and unburned plots imply that the maximum richness is achieved when there is a mixture of unburned forests and areas burned with canopy fire. Moreover, as canopy cover in P. nigra forests burned with canopy fire is not completed in the period of time studied, the presence of the species that are characteristic of burned areas remains along the chronosequence studied, while the species that disappear after fire do not recover in the period of time considered. Overall, the results obtained indicate that there is a persistent replacement of ant species in burned P. nigra forests, as is also the case with vegetation.     

Improving restoration success through microsite selection: an example with planting sagebrush seedlings after wildfire

Post‐fire restoration of foundation plant species, particularly non‐sprouting shrubs, is critically needed in arid and semi‐arid rangeland, but is hampered by low success. Expensive and labor‐intensive methods, including planting seedlings, can improve restoration success. Prioritizing where these more intensive methods are applied may improve restoration efficiency. Shrubs in arid and semi‐arid environments can create resource islands under their canopies that may remain after fire. Seedlings planted post‐fire in former canopy and between canopies (interspace) may have different survival and growth. We compared planting Wyoming big sagebrush (Artemisia tridentata Nutt. ssp. wyomingensis Beetle & Young) seedlings post‐fire in former sagebrush canopy and interspace microsites at five locations. Four growing seasons after planting, seedling survival was 46 and 7% in canopy and interspace microsites, respectively. Sagebrush cover was 5.8 times greater in canopy compared to interspace microsites. Sagebrush survival and cover were likely greater because of less competition from herbaceous vegetation as well as benefiting from resource island effects in canopy microsites. Initially, post‐fire abundance of exotic annual grasses was less in canopy microsites, but by the third year post‐fire it was substantially greater in canopy microsites, indicating that resource availability to seedlings was greater, at least initially, in canopy microsites. These results suggest microsites with greater likelihood of success should be identified and then utilized to improve restoration success and efficiency. This is important as the need for restoration greatly exceeds resources available for restoration.     

Positive Effects of Non-Native Grasses on the Growth of a Native Annual in a Southern California Ecosystem

Fire disturbance is considered a major factor in the promotion of non-native plant species. Non-native grasses are adapted to fire and can alter environmental conditions and reduce resource availability in native coastal sage scrub and chaparral communities of southern California. In these communities persistence of non-native grasses following fire can inhibit establishment and growth of woody species. This may allow certain native herbaceous species to colonize and persist beneath gaps in the canopy. A field manipulative experiment with control, litter, and bare ground treatments was used to examine the impact of non-native grasses on growth and establishment of a native herbaceous species, Cryptantha muricata. C. muricata seedling survival, growth, and reproduction were greatest in the control treatment where non-native grasses were present. C. muricata plants growing in the presence of non-native grasses produced more than twice the number of flowers and more than twice the reproductive biomass of plants growing in the treatments where non-native grasses were removed. Total biomass and number of fruits were also greater in the plants growing in the presence of non-native grasses. Total biomass and reproductive biomass was also greater in late germinants than early germinants growing in the presence of non-native grasses. This study suggests a potential positive effect of non-native grasses on the performance of a particular native annual in a southern California ecosystem.     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

Multivariate path analysis of the relationships between seedling regeneration and environmental factors beneath a dwarf bamboo understory

Seedling emergence and establishment are fragile processes that determine the direction and structure of forest succession and regeneration. However, seedling emergence and establishment are easily affected by biotic and abiotic (environmental) factors. A dense and expanding understory of dwarf bamboo is one such important factor that can seriously hinder the seedling regeneration. We conducted a field experiment to investigate the emergence and establishment of canopy tree seedlings under artificially controlled densities of dwarf bamboo. We found that understory dwarf bamboo obstructed seedling emergence but reduced the death of seedlings. Although understory dwarf bamboo reduced the median retention time of seedlings, dense bamboo increased the mean survival time of seedlings. Our results suggest that understory dwarf bamboo has multiple selectivities for tree seedling emergence and establishment: high‐density dwarf bamboo was beneficial to evergreen species but lower‐density of bamboo was conducive to the survival of deciduous species, it means the dwarf bamboo potentially alters successional trajectories of forest communities. Path analysis revealed that the most important factors affecting tree seedling emergence and death were the abundance of seeds in the seed bank and the density of emerged seedlings, and that the soil temperature promoted seedling emergence but increased seedling death, the thickness of litter limited seedling emergence, and the leaf area index of the bamboo canopy limited seedling death. The present study suggests that dwarf bamboo can directly alter the microenvironment, significantly reducing light levels and soil temperature but increasing the thickness of litter and soil humus, thereby indirectly impacting the regeneration of tree seedlings. Our results indicate that various factors affected seedling emergence, and there were complex indirect relationships among these factors. In general, biological factors had a stronger influence on tree seedling regeneration than environmental factors.     

Understory Vegetation Dynamics of North American Boreal Forests

Understory vegetation is the most diverse and least understood component of North American boreal forests. Understory communities are important as they act as drivers of overstory succession and nutrient cycling. The objective of this review was to examine how understory vegetation abundance, composition, and diversity change with stand development after a major stand replacing disturbance. Understory vegetation abundance and diversity increase rapidly after fire, in response to abundant resources and an influx of disturbance adapted species. The highest diversity occurs within the first 40 years following fire, and declines indefinitely thereafter as a result of decreasing productivity and increased dominance of a small number of late successional feather mosses and woody plant species. Vascular plant and bryophyte/lichen communities undergo very different successional changes. Vascular plant communities are dynamic and change more dramatically with time after fire, whereas bryophyte and lichen communities are much slower to establish and change over time. Considerable variations in these processes exist depending on canopy composition, site condition, regional climate, and frequently occurring non-stand-replacing disturbances. Forest management practices represent a unique disturbance process and can result in different understory vegetation communities from those observed for natural processes, with potential implications for overstory succession and long-term productivity. Because of the importance of understory vegetation on nutrient cycling and overstory composition, post-harvest treatments emulating stand-replacing fire are required to maintain understory diversity, composition, and promote stand productivity in boreal forests.     

Seedling Ecology and Restoration of Blackbrush (Coleogyne ramosissima) in the Mojave Desert,United States

Increases in fire frequency are disrupting many ecological communities not historically subjected to fire. In the southwestern United States, the blackbrush (Coleogyne ramosissima) community is among the most threatened, often replaced by invasive annual grasses after fire. This long‐lived shrub is vulnerable because it recruits sporadically, partially due to mast seeding and the absence of a seed bank. The goal of this study was to evaluate if shrub restoration can be enhanced by identifying and ameliorating recruitment limitations. Specifically, we tested the effect of encapsulating seeds in predation‐deterring “seed balls.” We also tested the effects of nurse plants and mammalian exclusion cages on seedling emergence, growth, and survivorship. These experiments were conducted in a full‐factorial design across three sites differing in elevation. Over 2 years, 13% of all planted seeds emerged and the effect of seed balls was overwhelmingly negative because of low emergence. Nurse plants had overall positive effects at Low Elevation, but negative effects at Mid‐ and High Elevation. Emergence and survival were highest in caged plots everywhere, and effect sizes increased with elevation. Interactions between the cage and the nurse plant treatments indicated that nurse plants tended to attract mammalian predators, lowering emergence and seedling survivorship, particularly at higher elevations. Findings conform to the stress‐gradient hypothesis in that interactions among seedlings and mature plants shifted from facilitation to competition as environmental stress decreased with increasing elevation, suggesting that they are transferable to ecologically similar communities elsewhere. Knowledge of site‐specific recruitment limitations can help minimize ineffective restoration efforts.     

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.     

Effects of Light Gaps and Litter Removal on the Seedling Performance of Six African Timber Species1

Valuable timber tree species frequently show poor regeneration after selective logging in tropical forests. Small size of logging gaps, lack of soil disturbance, and limited seed availability have each been blamed for observed regeneration failures. We investigated seed germination and seedling performance using a split‐plot factorial design involving light availability and litter removal for six Central African timber tree species, hypothesizing that canopy gaps and litter removal would improve seedling establishment, and that less shade‐tolerant species would show stronger responses to both factors. Contrary to our expectations, significantly more germinants established on intact litter than on exposed mineral soil 3 mo after seeding. After 18 mo, seedling survival, height and diameter growth, leaf area, and rooting depth were all much higher in gap plots than in the understory for all species, with the exception of Gilbertiodendron dewevrei, a highly shade‐tolerant species whose survival was higher in the understory. Leaf production was negatively influenced by litter removal in the least shade‐tolerant species, Nauclea diderrichii, with weak or positive effects in other species. G. dewevrei, while displaying a low‐light threshold for growth, exhibited a surprisingly high growth response to increasing light comparable to more shade‐intolerant species, a response that may help explain its local competitive dominance in the region. Due to the rapid closure of small gaps, we suggest that shade‐intolerant species such as N. diderrichii, Khaya anthotheca, and Entandrophragma utile might benefit from more intensive silvicultural practices that create larger canopy gaps.     

Altering Light Availability to Restore Invaded Forest: The Predictive Role of Plant Traits

Many studies have demonstrated that reduced light availability, which can be manipulated at local scales by planting or seeding canopy species, can curtail the growth of invasive species and promote the growth of native species. Species differences in functional traits, such as light use and stress tolerance, may be used to determine how native and invasive species will respond to these resource manipulations. We altered light availability in a mesic Hawaiian forest understory and found that low light levels reduced the biomass and growth of two invasive grasses (Pennisetum clandestinum and Ehrharta stipoides) relative to two native shrubs (Pipturus albidus and Coprosma rhynchocarpa) and two native canopy species (Metrosideros polymorpha and Acacia koa). Native species generally displayed traits associated with shade tolerance (high quantum yield, chlorophyll content, and leaf mass per area), whereas the two invasive grasses displayed traits associated with shade intolerance (high photosynthetic rate and growth rate). Several key traits pertaining to light acquisition and shade tolerance (quantum yield, chlorophyll content, and leaf mass per area) predicted seedling survival in low‐light treatments. Our data suggest that differences in light use among native and invasive species can help to determine the utility of resource manipulation as a restoration tool and, more specifically, to predict which native species will be optimal for restoration efforts that manipulate light availability.     

Competition and facilitation structure plant communities under nurse tree canopies in extremely stressful environments

Nurse plant facilitation in stressful environments can produce an environment with relatively low stress under its canopy. These nurse plants may produce the conditions promoting intense competition between coexisting species under the canopy, and canopies may establish stress gradients, where stress increases toward the edge of the canopy. Competition and facilitation on these stress gradients may control species distributions in the communities under canopies. We tested the following predictions: (1) interactions between understory species shift from competition to facilitation in habitats experiencing increasing stress from the center to the edge of canopy of a nurse plant, and (2) species distributions in understory communities are controlled by competitive interactions at the center of canopy, and facilitation at the edge of the canopy. We tested these predictions using a neighbor removal experiment under nurse trees growing in arid environments. Established individuals of each of four of the most common herbaceous species in the understory were used in the experiment. Two species were more frequent in the center of the canopy, and two species were more frequent at the edge of the canopy. Established individuals of each species were subjected to neighbor removal or control treatments in both canopy center and edge habitats. We found a shift from competitive to facilitative interactions from the center to the edge of the canopy. The shift in the effect of neighbors on the target species can help to explain species distributions in these canopies. Canopy‐dominant species only perform well in the presence of neighbors in the edge microhabitat. Competition from canopy‐dominant species can also limit the performance of edge‐dominant species in the canopy microhabitat. The shift from competition to facilitation under nurse plant canopies can structure the understory communities in extremely stressful environments.