Understory Light Availability Research Articles

Understory plants evade shading in a temperate deciduous forest amid climate variability by shifting phenology in synchrony with canopy trees.

Global warming is leading understory and canopy plant communities of temperate deciduous forests to grow leaves earlier in spring and drop them later in autumn. If understory species extend their leafy seasons less than canopy trees, they will intercept less light. We look for mismatched phenological shifts between canopy and understory in 28 years (1995-2022) of weekly data from Trelease Woods, Urbana, IL, USA. The observations cover 31 herb species of contrasting seasonality (for 1995-2017), three sapling species, and the 15 most dominant canopy tree species for all years, combined with solar radiation, temperature and canopy light transmittance data. We estimate how understory phenology, cold temperatures, canopy phenology, and solar radiation have individually limited understory plants' potential light interception over >2 decades. Understory and canopy phenology were the two factors most limiting to understory light availability, but which was more limiting varied greatly among species and among/within seasonality groups; solar radiation ranked third and cold fourth. Understory and canopy phenology shifts usually occurred in the same direction; either both strata were early or both were late, offsetting each other's effects. The four light-limiting factors combined showed significant temporal trends for six understory species, five toward less light interception. Warmer springs were significantly associated with shifts toward more light interception in three sapling species and 19 herb species. Canopy phenology became more limiting in warmer years for all three saplings species and 31 herb species. However, in aggregate, these variables mostly offset one another; only one sapling and seven herb species showed overall significant (and negative) relationships between light interception and spring temperature. The few understory species mismatched with canopy phenology due to changing climate are likely to intercept less light in future warmer years. The few species with data for carbon assimilation show broadly similar patterns to light interception.

Light availability during spring and autumn determines growth and survival of regenerated Korean pine seedlings

ABSTRACT The regeneration of Korean pine (Pinus koraiensis Sieb. et Zucc.) is a core issue for restoration from secondary to primary forests in northeastern China. However, Korean pine exhibits limited recruitment within its own canopy in primary forests, but demonstrates excellent regeneration in secondary forests. Until now, the mechanisms underlying this phenomenon have not been satisfactorily explained. Hence, we conducted studies in both primary and secondary forests to investigate whether light availability during the non-growing season determines the fate of regeneration. In spring and autumn, understory light availability in the secondary forest was significantly better than that in the primary forest. The biomass and non-structural carbohydrate (NSC) concentrations of natural pine seedlings were considerably greater in the secondary forests compared to primary forests. The carbohydrate concentrations increased during spring and autumn and decreased in summer. In the manipulation experiment, the biomass of seedlings was positively correlated with the organ NSCs in spring, while the survival rates were correlated with NSCs in autumn. The light availability during the spring and autumn seasons may have a crucial role in the regeneration of Korean pine seedlings. This finding may support our practical management approach for the transformation of secondary forests into Korean pine forests.

Understory seedlings of Quercus mongolica survive by phenological escape

Understanding understory seedling regeneration mechanisms is important for the sustainable development of temperate primary forests in the context of increasingly intense climate warming events. The poor regeneration of dominant tree species, however, is one of the biggest challenges it faces at the moment. Especially, the regeneration of the shade-intolerant Quercus mongolica seedling is difficult in primary forests, which contrasts with the extreme abundance of understory seedlings in secondary forests. The mechanism behind the interesting phenomenon is still unknown. This study used in-situ monitoring and nursery-controlled experiment to investigate the survival rate, growth performance, as well as nonstructural carbohydrate (NSC) concentrations and pools of various organ tissues of seedlings for two consecutive years, further analyze the understory light availability and simulate the foliage carbon (C) gain in the secondary and primary forest. Results suggested that seedlings in the secondary forest had greater biomass allocation aboveground, height and specific leaf area (SLA) in summer, which allowed the seedling to survive longer in the canopy closure period. High light availability and positive C gain in early spring and late autumn are key factors affecting the growth and survival of understory seedlings in the secondary forest, whereas seedlings in the primary forest had annual negative carbon gain. Through the growing season, the total NSC concentrations of seedlings gradually decreased, whereas those of seedlings in the secondary forest increased significantly in autumn, and were mainly stored in roots for winter consumption and the following year's summer shade period, which was verified by the nursery-controlled experiment that simulated autumn enhanced light availability improved seedling survival rate and NSC pools. In conclusion, our results revealed the survival trade-off strategies of Quercus mongolica seedlings and highlighted the necessity of high light availability during the spring and autumn phenological periods for shade-intolerant tree seedling recruitment.

Effects of thinning on the understory light environment of different stands and the photosynthetic performance and growth of the reforestation species Phoebe bournei

Light levels determine regeneration in stands and a key concern is how to regulate the light environment of different stand types to the requirements of the understory. In this study, we selected three stands typical in south China (a Cryptomeria japonica plantation, a Quercus acutissima plantation, and a mixed stand of both) and three thinning intensities to determine the best understory light environment for 3-year-old Phoebe bournei seedlings. The canopy structure, understory light environment, and photosynthesis and growth indicators were assessed following thinning. Thinning improved canopy structure and understory light availability of each stand; species composition was the reason for differences in the understory light environment. Under the same thinning intensity, the mixed stand had the greatest light radiation and most balanced spectral composition. P. bournei photosynthesis and growth were closely related to the light environment; all three stands required heavy thinning to create an effective and sustained understory light environment. In a suitable understory light environment, the efficiency of light interception, absorption, and use by seedlings was enhanced, resulting in a higher carbon assimilation the main limiting factor was stomatal conductance. As a shade-avoidance signal, red/far-red radiation is a critical factor driving changes in photosynthesis and growth of P. bournei seedlings, and a reduction increased light absorption and use capacity and height: diameter ratios. The growth advantage transformed from diameter to height, enabling seedlings to access more light. Our findings suggest that the regeneration of shade-tolerant species such as P. bournei could be enhanced if a targeted approach to thinning based on stand type was adopted.

Restoration of low-intensity fire in Quercus-Pinus mixedwoods following a prolonged period of fire exclusion

Low-intensity surface fire is required to restore and maintain Quercus-Pinus mixedwood composition, structure, and function. However, historical fire exclusion has resulted in altered vegetation-fuel-fire feedbacks in long-unburned mixedwoods. Fire is now being reintroduced to reduce fire-intolerant understory and midstory stem density, consume excessive litter accumulation, eradicate the duff layer, and achieve various other goals; however, the consequences of restoring fire are poorly understood. The goal of this study was to quantify the effects of fire reintroduction on intra-stand Pinus echinata neighborhoods in a mixed Quercus-P. echinata stand following an extended period of fire exclusion. We report results from a 3-year sampling of the effects of one dormant season and one early growing season fire on woody plant dynamics, understory light, fuel conditions, and P. echinata basal duff-ring accumulation. The sapling assemblage was unaffected by one fire, but the second fire resulted in shoot mortality across all taxa. Understory light availability was unchanged after two fires, and litter depth was significantly reduced after the first and second fire. We found that fire seasonality was likely more important than fuel conditions to achieve desired fire effects. Our results indicated that continued fire and midstory treatment will be required to regenerate P. echinata.

Light availability and anthropogenic stress shape plant understory invasions in understory of urban forests: A case study in Shanghai

Light availability and anthropogenic stress shape plant understory invasions in understory of urban forests: A case study in Shanghai

Forest restoration in old pasture areas dominated by <i>Urochloa brizantha</i>

In the present study it was evaluated the density of recruits in an area previously occupied by Urochloa brizantha pasture undergoing forest restoration and in a Semideciduous Forest. Was evaluated the effect of the available light on the biomass of the exotic grass, and investigated associations of recruit density with soil variables and understory light availability. The seed rain in both areas was also evaluated. The biomass of U. brizantha was positively associated with available light and, a negative association was found between recruit density and available light when considering the two areas together. A negative associatio between recruit density and soil pH was found. The results suggest that the low soil fertility did not limited recruitment in the semideciduos forest. The results showed that high resource availability favors the competitivity of U. brizantha, impairing the forest restoration in area previously used as pastures. So, pratices that result in decreasing in light availability would favors the forest restoration in areas previously used as pasture.

Understory diversity are driven by resource availability rather than resource heterogeneity in subtropical forests

Understory vegetation accounts for most of the plant diversity, particularly in plantation forests, and contributes significantly to energy flows and nutrient cycling. However, we lack a mechanistic understanding of understory diversity associated with resource availability and heterogeneity across forest types over a wide range of stand development stages. We sampled the understory vegetation of Chinese fir plantations and natural secondary forests (4–40 years old) and used primary forests as a reference in subtropical China. The percent covers of all understory plant species were estimated at the shrub layer (height >1.3 m) and herb layer (≤1.3 m). We quantified understory light availability and heterogeneity, substrate diversity and soil nutrients and employed principal component analysis to reduce multiple physical and chemical soil properties to PC1 and PC2, with PC1 positively correlating with soil carbon and total nitrogen, and PC2 with available soil nitrogen and phosphorus. Structural equation modelling (SEM) was employed to test the direct and indirect influences of forest type, stand age, light availability and heterogeneity, substrate diversity and soil nutrients on the diversity of understory species. We found that shrub layer richness was higher in the secondary natural and primary forests than in the plantations, but plantations had higher bryophyte richness. Shrub cover was positively correlated with shrub richness and had a positive indirect effect on shrub richness but a negative indirect effect on herb richness via its effect on soil carbon and nitrogen. Stand age has a positive direct effect on bryophyte richness. There were strong positive correlations among the richness of understory layers and life forms. Our results suggest that physical and chemical soil properties, specifically soil nutrient availability, rather than light availability and heterogeneity or substrate diversity, were the main drivers that affected subtropical forests' understory richness.

Understory plant diversity under variable overstory cover in Mediterranean forests at different spatial scales

Mediterranean forest ecosystems are of high conservation value and therefore increasingly managed to restore and promote their natural biodiversity and complexity. The present study aimed to assess plant species diversity of East-Mediterranean conifer forests as affected by overstory cover manipulation at multiple spatial scales. Understanding this interaction in water-limited forest ecosystems is particularly challenging as overstory-understory relations involve both water and light constrains. The experiment was conducted in a mature (40y) planted Pinus halepensis forest in the Jerusalem Mountains of Israel. Understory plant community structure was monitored for five years in sixteen experimental plots (0.5 ha) representing a range of overstory cover levels (leaf area index-LAI = 0–4 m2m−2) created by tree thinning. Light and water availability in the forest understory were monitored using fisheye camera (irradiance), predawn leaf water potential (PLWP) of dominant species and soil water content (SWC) measurements. Plant diversity was measured at increasing spatial scales (grain sizes, 0.01–400 m2), using hierarchical nested sampling. Irradiance and SWC increased with decreasing LAI while PLWP was not affected. Stand-level understory richness (α diversity) increased linearly with decreasing LAI, indicating a consistent, positive resource availability-diversity relationship. The results pointed towards strong dependency of species richness on light availability as compared to its dependency on water availability that was less definite. Understory species composition varied across LAI levels (β diversity at the forest level) indicating species turnover, mainly between annual herbs benefiting from reduced overstory cover vs. woody species benefiting from higher cover. The relationships between overstory cover and understory richness and composition were rank-invariant across grain sizes. These relationships were stronger at the larger (10, 100, 400 m2) compared to the smaller (0.01, 0.1, 1 m2) grain sizes. The 10-m2 grain-size, corresponding to the crown projection area of individual trees, showed the strongest relationship. Observed patterns of overstory cover–understory richness suggest that resource availability, rather than spatial resource heterogeneity, was the main factor shaping understory diversity. Furthermore, grain-size effects on this relationship support “environmental filtering” rather than “resource density” as the main mechanism through which reduced overstory cover promoted understory diversity.'Synthesis and applications’: Our findings demonstrate the importance of overstory cover as a key factor determining plant diversity in water-limited East-Mediterranean forests. We highlight the important role of spatial heterogeneity of overstory cover at both the stand (intra-plot) and forest (inter-plot) scales and support dynamic variable thinning as a management strategy to enhance forest biodiversity and complexity.

Distribution and conservation status of Camellia longzhouensis (Theaceae), a critically endangered plant species endemic to southern China

Camellia longzhouensis (Theaceae) is an endemic evergreen shrub or small tree with a distribution restricted to South China. It is listed as Grade-II in the National Key Protected Wild Plants List. In this study, we surveyed its distribution, examined its population structure, identified factors affecting its survival, and reassessed its extinction risk. We found that C. longzhouensis was only distributed in the Nonggang National Nature Reserve and the surrounding area. Its individuals only grew under the secondary forest canopy in the karst mountain. A total of 58 individuals of C. longzhouensis in three sub-populations were found. Soil fertility and understory light availability were the main habitat factors influencing the survival of C. longzhouensis. Anthropogenic disturbances and reproductive obstacles have caused a low seed-setting rate, poor seedling survival, and a lack of adult plants of C. longzhouensis in the natural habitat. The population structure of C. longzhouensis is spindle-shaped, indicating poor natural regeneration and inhibited seedling recruitment. Cleistanthus petelotii had a significant positive interspecific interaction with C. longzhouensis in the community. Based on the information obtained here and IUCN criterion C2ai, we recommend that C. longzhouensis be categorized in the International Union for Conservation of Nature (IUCN) Red List as Critically Endangered. We also developed a comprehensive protection strategy, consisting of in situ conservation, ex situ conservation, reintroduction, and commercial utilization. This strategy can be readily applied to protect other endangered plants with economic value in karst poor regions.

Initial plant community responses to hardwood control treatments in restoration of remnant longleaf pine (Pinus palustris) woodlands

Changes in land use over the past century have contributed to substantial losses of longleaf pine (Pinus palustris) woodlands in the southeastern USA and replacement with higher density, mixed pine and hardwood stands that suppress understory development and limit application of prescribed fire. To increase understanding of limiting factors and identify potential approaches for restoring longleaf pine woodlands, we studied initial availability of light and soil water and 2-year (2018–2019) plant community responses after controlling overstory hardwoods in five remnant longleaf pine woodlands having no evidence of previous agriculture at the Savannah River Site near Aiken, SC, USA. Seven hardwood control treatments and a non-treated check were compared in a randomized compete block experiment: cutting, cutting + shredding of logging residues, stem injection with imazapyr herbicide, cutting + basal spray with imazapyr herbicide, cutting + basal spray with triclopyr herbicide, cutting + directed foliar spray with a mixture of glyphosate and imazapyr herbicides, and cutting + broadcast foliar spray with the same herbicide mixture. In the year prior to hardwood cutting (2016), understory light availability averaged 23% of full sun. Throughout the year of treatment installation (2017), average soil water content (SWC) was below permanent wilting point (5.5% SWC) where overstory hardwoods were retained; whereas, it was above wilting point (7.5%) where they had been cut. Thus, combined effects of shade and root competition from overstory hardwoods probably limited cover of herbaceous species in the non-treated check. In the 2 years following treatment, hardwood survival averaged > 99%, 62%, 42% and < 1% for the non-treated check, cutting, shredding, and herbicide treatments, respectively. Relative to the non-treated check, herbaceous species richness was increased by shredding, stem injection, basal spray, or directed foliar treatments; herbaceous cover was increased by basal spray or directed foliar treatments; and woody cover was decreased by directed or broadcast foliar treatments. The directed foliar spray was the most effective treatment for achieving many of the desired understory characteristics of a longleaf pine woodland, including a diverse understory dominated by herbaceous vegetation capable of supporting periodic prescribed fires. Although the imazapyr basal spray treatment increased cover of remnant woodland indicator species, none of the treatments increased richness of this species group, suggesting that restoration of native species composition will be expedited by enrichment seedings or plantings.

The Spatial-Temporal Heterogeneity of Understory Light Availability in a Temperate Forest of North China

The spatial-temporal variation of understory light availability has important influences on species diversity and community assembly. However, the distribution characteristics and influencing factors of understory light availability have not been fully elucidated, especially in temperate deciduous, broad-leaved forests. In this study, the understory light availability was monitored monthly (May–October) in a temperate deciduous, broad-leaved forest in Henan Province, China. Differences in the light availability among different months and habitat types were statistically analyzed using Kruskal–Wallis method, respectively. Partial least squares path modeling (PLS-PM) was used to explore the direct and/or indirect effects of stand structure, dominant species and topographic factors on the light environment. Results showed that there were differences in light environments among the four habitat types and during the studied six months. The PLS-PM results showed that the stand structure and the dominant species were negatively correlated with the light environment, and the path coefficient values were −0.089 (P = 0.042) and −0.130 (P = 0.004), respectively. Our result indicated that the understory light availability exhibit a distinct spatial and temporal heterogeneity in temperate deciduous, broad-leaved forest of north China. The characteristics of woody plant community, especially the abundance of one of the dominant plant species, were the important factors affecting the understory light availability.

Characterizing Canopy Openness in Open Forests: Spherical Densiometer and Canopy Photography Are Equivalent but Less Sensitive than Direct Measurements of Solar Radiation

AbstractIn forest ecosystems, canopy openness affects understory light availability, plant growth, and tree species recruitment, thus shaping future forest composition, structure, and functional diversity. Foresters must correctly and quickly measure canopy openness to meet their management objectives. To help guide the selection of an appropriate method for measuring canopy openness, we compared three common techniques that vary in cost, complexity, and time required for measurements and data processing: smartphone-based hemispherical photography, spherical densiometer measurements, and direct measurements of solar radiation (using AccuPAR ceptometer). We measured canopy openness using these three methods on 28 permanent forest health monitoring plots in pine-oak forests of the Central Pine Barrens of Long Island in New York State. By analysis of variance and regression analyses, we found the three methods (particularly densiometer and hemispherical photographs) yielded broadly equivalent and strongly positively correlated descriptions of canopy openness. The direct measurements of solar radiation seemed to have a greater potential to detect subtle variation in forest understory light. Forest managers may sufficiently characterize canopy openness using quick and cheap methods (e.g., spherical densiometers) and avoid larger costs of devices for direct light measurements (e.g., ceptometers) and the larger data-processing times of hemispherical photography.

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.

Early indications of success rehabilitating an underperforming teak (Tectona grandis) plantation in Panama through enrichment planting

Enrichment planting can be an effective strategy for increasing the ecological and economic value of timber plantations, but success depends on appropriate matching of under- and overstory species and site conditions. This case study in the Panama Canal Watershed explores the viability of enrichment planting for rehabilitating underperforming teak (Tectonia grandis) plantations, which are common in the area. Two high-value timber species native to the neotropics, Dalbergia retusa and Terminalia amazonia, were underplanted in an established teak plantation at the Agua Salud research site in the province of Colon, Panama. Seedling survival, basal diameter, height, total biomass and relative growth rate were assessed for the two years following planting. In contrast with the widespread belief that teak is a poor nurse tree, both species achieved promising early growth with low mortality in plantation understory conditions. Neither understory light availability nor combined above- and below-ground effects of crowding pressure from teak strongly predicted growth of either species. D. retusa, thought to be more shade-tolerant, performed equally across a range of intermediate light levels whereas T. amazonia, thought to be more heliotropic, performed best at the highest light levels, though light relationships explained little variation in seedling growth. These early findings support the suitability of either species for use in enrichment plantings in established, underperforming teak plantations in the Panama Canal Watershed. Longer-term research is needed to evaluate the potential of enrichment planting to increase profitability and ecosystem services such as carbon sequestration and water resource management in these plantations.

Effects of light availability on leaf attributes and seedling growth of four tree species in tropical dry forest

BackgroundIn tropical dry forests, variation in understory light availability due to season and canopy tree density could be a governing factor in establishment and growth of tree seedlings. Species with varying life history traits are expected to respond differentially to such heterogeneity. We investigated the response of seedlings of four tree species in a tropical day forest in relation to spatiotemporal variability of light. We attempt to explore the role of leaf attributes in explaining intra- and inter-specific variations in relative growth rate. Four study sites, each with three contrasting canopy conditions, were selected along a soil moisture gradient. Seedlings of four tree species (viz., Acacia catechu, Bridelia retusa, Lagerstroemia parviflora, and Shorea robusta), varying in life history traits, were monitored for seasonal variations in growth traits across canopy condition and sites for 2 years.ResultsWe observed a larger variation in leaf attributes for pioneer species. A. catechu showed highest mean values for leaf dry matter content, leaf nitrogen concentration, leaf phosphorus concentration, net stomatal conductance, net photosynthetic rate, and relative growth rate in high light conditions. S. robusta and B. retusa demonstrated highest mean values for all the leaf attributes (except leaf dry matter content) in low light conditions. However, intermediate values for leaf attributes were observed in L. parviflora which preferred moderate light conditions.ConclusionsSeasonal variations in light availability at the forest floor appear to play an important role in the establishment and growth of tree seedlings in seasonal dry forests. Leaf attributes can be used to explain intra- and inter-specific variation in response to light availability. Leaf attributes in combinations can be used to predict relative growth rate of tree species in tropical dry environment, which apart from soil moisture is also determined by light availability due to seasonal changes and canopy tree density.

Understory Floristic Diversity in Poplar Plantations in the Dongting Lake Wetlands, China

Large-scale poplar plantations have evoked great concern over their adverse effects on floristic diversity in understory vegetation. The aim of this study was to investigate the effects of tree density on the composition of understory vegetation in poplar (Populus deltoides) plantations in the Dongting Lake wetlands, China. Stand characteristics and soil properties had significant effects on understory plant distribution. Stand density was positively correlated with number of species, Shannon diversity index, and proportions of mesophytes and partial-shade plants but was negatively correlated with the proportions of hygrophytes and heliophytes. Understory light availability was negatively correlated with number of species, Shannon diversity index, and the proportions of mesophytes and partial-shade plants and was positively correlated with the proportions of hygrophytes and heliophytes. Soil water content was negatively correlated with species richness and the proportion of mesophytes and was positively correlated with the proportions of hygrophytes and heliophytes. Compared to low-density stands, high-density stands had a higher number of species and higher proportion of annual/biennial plants, mesophytes, and xerophytes and had lower proportions of perennial plants and hygrophytes during the one-year monitoring period. Therefore, poplar plantations should be managed at lower densities in the Dongting Lake wetlands to conserve the understory flora.

How can the shade intolerant Korean pine survive under dense deciduous canopy?

The regeneration of Korean pine (Pinus koraiensis Sieb. et Zucc) is a core issue for restoration from secondary to primary forests. There is a lack of Korean pine regeneration in primary forests, compared to an abundance of regeneration in secondary forests. However, the mechanisms behind this mysterious phenomenon have not been studied. This study investigated seasonal understory light availability in addition to carbon (C) gain of understory Korean pine, nonstructural carbohydrate (NSC) storage of Korean pine in secondary forests, and the growth and survival of seedlings under manipulated light regimes. Understory light intensities in early spring and late autumn in secondary forests were significantly higher than in primary forests. The C gain of understory Korean pine occurred mostly in early spring and late autumn, whereas it was negative in the growing season (summer). Total NSC concentrations of Korean pine saplings increased between growing seasons, and decreased during the growing season. In particular, the NSC was dominated by a progressive increase in starch in early spring and by a peak in soluble sugar in late autumn. In this study, the biomass, NSC concentration and pool sizes, as well as survival rates, were positively correlated with light availability, revealing that the species is shade intolerant. The accumulated survival rate, examined in the autumn, was less correlated with biomass, but positively correlated with the carbohydrate storage in autumn. High light availability and positive C gain in early spring and late autumn are key factors affecting the growth and survival of understory seedlings. These two periods in secondary forests are crucial for the survival of seedlings by ensuring sufficient energy to live through the long-lasting dark summer. In primary forests, however, light availability was relatively stable across all seasons, especially in the leafless period, and insufficient through-canopy radiation, along with dense lateral closure, caused seedlings mortality.

Unraveling Shade Tolerance and Plasticity of Semi-Evergreen Oaks: Insights From Maritime Forest Live Oak Restoration.

Quercus spp. (oaks) are generally intermediate in shade tolerance, yet there is large variation within the genus in shade tolerance and plasticity in response to varying resource availability. Ecophysiological knowledge specific to semi-evergreen Quercus spp. from subtropical maritime forests is lacking relative to temperate deciduous oaks. We studied the influence of light availability and plant competition on leaf physiology and performance of semi-evergreen Quercus virginiana on a barrier island along the US southern Atlantic coast. Seedlings were underplanted in pine (Pinus taeda) plantation stands with varying overstory density (clear-cut, heavy thin, light thin, and non-thinned; creating a gradient of understory light availability) and vegetation (no competition removal or herbaceous competition removal) treatments. After 2 years, seedling survival was higher with increasing light availability (clear-cut = heavy thin > light thin > non-thinned). Seedling growth (i.e., diameter, height, and crown width) increased similarly with increasing thinning intensity, while vegetation control was mainly beneficial to seedling growth in clear-cuts. These responses were partially explained by foliar nitrogen and leaf trait measurements, which followed the same pattern. Q. virginiana seedlings demonstrated high plasticity in their ability to acclimate to varying resource availability, as indicated by light response curves, specific leaf area, stomatal density, stomatal pore index, and maximum theoretical stomatal conductance. Light compensation and saturation points illustrated seedling capacity to increase net CO2 assimilation with increased light availability. Leaves on trees in the high light environment had the highest net CO2 assimilation, stomatal density, stomatal pore index, maximum theoretical stomatal conductance, and lowest specific leaf area. Although we demonstrated the relative shade tolerance of Q. virginiana in lower light environments (i.e., heavy and light thin plots), this semi-evergreen species shows high plasticity in capacity to respond to varying resource availability, similar to other Quercus spp. from mesic and Mediterranean environments.

Similar Impacts of Alien and Native Tree Species on Understory Light Availability in a Temperate Forest

Research Highlights: We evaluated influence of alien and native trees and shrubs on stand leaf area index to basal area ratio, indicating that both groups provide similar amounts of foliage. Background and Objectives: Foliage traits determine tree species effect on understory light availability. Direct comparisons of understory light availability due to different foliage traits of tree species are conducted less often at the stand level. We hypothesized that light availability is driven by canopy leaf area, and alien species contribution to canopy foliage will be similar to native species due to analogous patterns of biomass allocation in tree species. Materials and Methods: We studied forests dominated by alien and native tree species in Wielkopolski National Park (Western Poland). We measured light availability using the LAI-2200 canopy analyzer (Li-Cor Inc., Lincoln, NE, USA) and we calculated leaf area index (LAI) in 170 stands using published models of foliage biomass and data on specific leaf area. Results: Our study confirmed an impact of LAI on light availability in the understory layer. Analyzing the proportion of contribution to stand LAI and basal area (BA) we found that most alien species did not differ in LAI to BA ratio from native species. The exception was Prunus serotina Ehrh., with a LAI to BA ratio higher than all native and alien trees. However, the highest LAI to BA ratios we found were for the alien shrub Cotoneaster lucidus Schltdl. and native shrubs of fertile broadleaved forests. Conclusions: Our study showed that alien species contribution to shading the understory is comparable to native species, with the exception of P. serotina due to its dominance in the higher shrub canopy strata where it exhibits different patterns of biomass allocation than native trees. Our study explained that invasive tree species impact on light availability in forest ecosystems is mainly mediated by the increased quantity of foliage, not by more effective LAI to BA ratio.