Forest Herbs Research Articles

Warm Spring Days are Related to Shorter Durations of Reproductive Phenophases for Understory Forest Herbs.

As plants continue to respond to global warming with phenological shifts, our understanding of the importance of short-lived heat events and seasonal weather cues has lagged relative to our understanding of plant responses to broad shifts in mean climate conditions. Here, we explore the importance of warmer-than-average days in driving shifts in phenophase duration for spring-flowering woodland herbs across one growing season. We harnessed the combined power of community science and public gardens, engaging more than 30 volunteers to monitor shifts in phenology (documenting movement from one phenophase to the next) for 198 individual plants of 14 species twice per week for the 2023 growing season (March-October) across five botanic gardens in the midwestern and southeastern US. Gardens included the Holden Arboretum, Kirtland, OH; Dawes Arboretum, Newark, OH; Chicago Botanic Garden, Glencoe, IL; Missouri Botanical Garden, St. Louis, MO; and Huntsville Botanical Garden, Huntsville, AL. We tested: (1) that higher-than-average daily temperatures (deviation from 30-year historical mean daily temperatures for each location) would be related to truncated phenophase durations; and (2) that phenophase durations would vary among species. Our findings support both hypotheses. We documented significant inverse relationships between positive deviations in daily temperature from historic means (e.g., warmer-than-average days) and durations of three reproductive phenophases: "First Bud," "First Ripe Fruit," and "Early Fruiting." Similar (non-significant) trends were noted for several other early-season phenophases. Additionally, significant differences in mean phenophase durations were detected among the different species, although these differences were inconsistent across plant parts (vegetative, flowering, and fruiting). Results underscore the potential sensitivity of understory herb reproductive phenophases to warmer-than-average daily temperatures early in the growing season, contributing to our understanding of phenological responses to short-term heat events and seasonal weather cues.

Cattle exclusion increases encounters of wild herbivores in Neotropical forests

Abstract Ongoing habitat loss and species extinctions require managers to implement and quantify the effectiveness of conservation actions for protecting biodiversity. Fencing, when done properly, is an important management tool for conservation in landscapes where wildlife and domestic animals co‐occur, potentially enhancing habitat use through selective exclusion of domestic species. For instance, the fencing of forest patches in the Neotropics is expected to reduce the degradation of understory vegetation by cattle, releasing these resources for the native community of browsers and fruit consumers. Here, we implemented an ecological experiment using a before‐after control‐impact design to quantify the effect of cattle exclusion on encounter probability of the native community of browsers and fruit consumers, and percent ground cover in multifunctional landscapes of the Colombian Orinoquía. We built 14 km of wildlife‐permeable fences along forest edges in four forest patches (i.e. blocks) containing control and fenced (treatment) sites. We installed 33 camera traps to obtain information about wildlife and cattle encounter probabilities, before and after the fences were constructed. We used Bayesian generalised linear mixed effects models to quantify the effect of fences via the interaction between the time period (before and after the fences were built) and treatment (control or fenced sites). Fencing was effective at reducing encounter probabilities of cattle in the treated sites, and it had a positive impact on relative encounter probabilities of four of seven studied wildlife species (herbivores including the black agouti [dry season only], lowland tapir [dry season only] and spotted paca [both seasons] and an omnivore, the South American coati [rainy season only]). The effect of fencing was negative for the collared peccary but only during the dry season. No statistically significant effect was detected for the white‐lipped peccary or white‐tailed deer. Synthesis and applications: We provide experimental evidence that fences are effective at selectively excluding cattle and increasing encounter rates of wild browsers and fruit consumers in forest patches where these species co‐occur with cattle. Our results highlight an important application of fencing ecology in Neotropical forests, where the implementation of wildlife‐permeable fences is feasible due to smaller body sizes of wildlife compared to domestic animals such as cattle.

Leaf-cutting ants’ distribution in an elevation gradient in central Argentina

ABSTRACT Leaf-cutting ants are among the most prominent herbivores in neotropical forests and savannas, and Argentina represents the southern limit of their distribution. Despite their importance, few systematic studies analyze the presence and distribution of leaf-cutting ants in Argentina. Our aim was to describe, through an intensive field sampling, the leaf-cutting ant species present in the central Argentine mountains and analyze their distribution along the elevation gradient. We collected ants from 69 nests in 56 out of the 250 plots visited along the study elevation gradient (400–2700 m above sea level) in seven different habitats, from forests to rocky areas. We found six species. The most frequent were Acromyrmex lobicornis and Acromyrmex crassispinus, while Acromyrmex heyeri, Amoimyrmex silvestrii, Amoimyrmex striatus, and Amoimyrmex bruchi were far less frequent. The species distributed at the highest elevation was A. lobicornis (1223 to 2163 m a.s.l.), showing significant differences with the other species. We found leaf-cutting ants in all habitats, with a higher presence in grazing lawns, shrublands, and exotic forests. Our results indicate that leaf-cutting ants are present in most of the mountain range, as the area above their upper limit of distribution represents less than 5% of the total mountain area.

Bumblebees mediate landscape effects on a forest herb's population genetic structure in European agricultural landscapes.

Spatially isolated plant populations in agricultural landscapes exhibit genetic responses not only to habitat fragmentation per se but also to the composition of the landscape matrix between habitat patches. These responses can only be understood by examining how the landscape matrix influences among-habitat movements of pollinators and seed vectors, which act as genetic linkers among populations. We studied the forest herb Polygonatum multiflorum and its associated pollinator and genetic linker, the bumblebee Bombus pascuorum, in three European agricultural landscapes. We aimed to identify which landscape features affect the movement activity of B. pascuorum between forest patches and to assess the relative importance of these features in explaining the forest herb's population genetic structure. We applied microsatellite markers to estimate the movement activity of the bumblebee as well as the population genetic structure of the forest herb. We modelled the movement activity as a function of various landscape metrics. Those metrics found to explain the movement activity best were then used to explain the population genetic structure of the forest herb. The bumblebee movement activity was affected by the cover of maize fields and semi-natural grasslands on a larger spatial scale and by landscape heterogeneity on a smaller spatial scale. For some measures of the forest herb's population genetic structure, that is, allelic richness, observed heterozygosity and the F-value, the combinations of landscape metrics, which explained the linker movement activity best, yielded lower AICc values than 95% of the models including all possible combinations of landscape metrics. Synthesis: The genetic linker, B. pascuorum, mediates landscape effects on the population genetic structure of the forest herb P. multiflorum. Our study indicates, that the movement of the genetic linker among forest patches, and thus the pollen driven gene flow of the herb, depends on the relative value of floral resources in the specific landscape setting. Noteworthy, the population genetic structure of the long-lived, clonal forest herb species correlated with recent land-use types such as maize, which have been existing for not more than a few decades within these landscapes. This underscores the short time in which land-use changes can influence the evolutionary potential of long-lived wild plants.

Soil properties constrain forest understory plant distributions along an elevation gradient.

Projections of spatial biodiversity dynamics under climate warming are often based on models including only climate variables, and when non-climatic factors (e.g. soil) are included, data are often at much coarser spatial resolutions than those experienced by plants. Field studies along elevation gradients permit the gathering of detailed soil data, while still covering a wide climatic gradient. Here, an intensive field survey of four spring forest herbs along an elevation gradient showed that soil properties had substantial impacts on the occurrence/abundance of all species, and that soil effects were more pronounced at higher elevations. For Trillium erectum and Claytonia caroliniana, very infrequent occurrences at high elevation were strongly associated with rare microsites with high pH or nutrients. In a seven-year transplant experiment with T. erectum, we found that individuals grew to much smaller sizes at high than low elevation, suggesting that environmental factors rather than dispersal limitation constrain the species' upper range limit, despite substantial warming in recent decades. Our study demonstrates that soil factors interact strongly with climate to determine plant range limits along climatic gradients. Unsuitable soils for plants at high elevations or latitudes may represent an important constraint on future plant migration and biodiversity change. This article is part of the theme issue 'Ecological novelty and planetary stewardship: biodiversity dynamics in a transforming biosphere'.

Contrasting responses of forest phenological guilds to complex floodplain change

Abstract Understanding the seasonal dynamics of the forest herb layer and the factors that influence it is essential for predicting how the forest ecosystem, including mutualistic interactions, will respond to global change. However, to date, no research has investigated how understorey phenological guilds respond to major environmental threats and forest interventions. We showed the marked changes in the phenological structure of floodplain forest plant communities over half a century at a multi‐regional scale in Central Europe. Unlike previous studies focusing on individual species' responses to climate change, we examined shifts in species richness, frequency and abundance between phenological guilds within the plant community. We examined the effects of temporal variation in climate, hydrology, soil conditions and canopy structure, and assessed the effects of non‐intervention and intervention management on phenological guild responses. Our results show a significant loss of local species richness and a decline in species frequency in the forest understorey vegetation, associated with species exhibiting summer phenology. In contrast, an increase in abundance and richness of spring flowering species was observed, attributed to high nutrient loads and reduced flooding, respectively. The abundance of spring species increased only in stands where canopy cover increased over time, probably due to the suppression of summer flowering species, allowing spring flowering species to spread on open, bare ground. The community experienced a shift in maximum flowering plant richness from June to May. Despite management interventions such as clearcutting and tree planting, our data show similar temporal trajectories in intervention and non‐intervention forests, indicating that changes in phenological structure are largely independent of recent management activities. Synthesis. We propose that a complex interplay of environmental factors, rather than climate change alone, is shaping shifts towards earlier phenological guilds in floodplain forests. This study highlights the importance of considering a comprehensive view of phenological guild dynamics and the impact of environmental factors on forest plant community structure in the face of global change.

Differences and drivers of leaf stable carbon and nitrogen isotope in herbs under different vegetation types on the eastern Qinghai-Tibet Plateau

The natural abundance of stable carbon and nitrogen isotopes (δ13C and δ15N) in leaves can provide comprehensive information on the physiological and ecological processes of plants and has been widely used in ecological research. However, recent studies on leaf δ13C and δ15N have focused mainly on woody species, few studies have been conducted on herbs in different vegetation types, and their differences and driving factors are still unclear. In this study, we focused on the herbs in subalpine coniferous forests, alpine shrublands, and alpine mea-dows on the eastern Qinghai-Tibet Plateau, and investigated the differences in leaf δ13C and δ15N of herbs and the driving factors. The results showed that there were significant differences in leaf δ13C and δ15N values of herbs among different vegetation types, with the highest δ13C and δ15N values in alpine meadows, followed by alpine shrublands, and the lowest in subalpine coniferous forests. Using variation partitioning analysis, we revealed that differences in leaf δ13C and δ15N of herbs among various vegetation types were driven by both leaf functional traits and climate factors, with the contribution of leaf functional traits being relatively higher than that of climate factors. Hierarchical partitioning results indicated that mean annual temperature (MAT), chlorophyll content index, leaf nitrogen content per unit area (Narea), and leaf mass per area were the main drivers of leaf δ13C variations of herbs across different vegetation types, while the relative importance of Narea and MAT for variation in leaf δ15N of herbs was much higher than those other variables. There was a strong coupling relationship between leaf δ13C and δ15N as indicated by the result of the ordinary least squares regression. Our findings could provide new insights into understanding the key drivers of leaf δ13C and δ15N variations in herbs across different vegetation types.

Declining potential nectar production of the herb layer in temperate forests under global change

Abstract Wild pollinators are crucial for ecosystem functioning and human food production and often rely on floral resources provided by different (semi‐) natural ecosystems for survival. Yet, the role of European forests, and especially the European forest herb layer, as a potential provider of floral resources for pollinators has scarcely been quantified. In this study, we measured the potential nectar production (PNP) of the forest herb layer using resurvey data across 3326 plots in temperate forests in Europe, with an average time interval of 41 years between both surveys in order to assess (i) the importance of the forest herb layer in providing nectar for wild pollinators, (ii) the intra‐annual variation of PNP, (iii) the overall change in PNP between survey periods and (iv) the change in intra‐annual variation of PNP between survey periods. The PNP estimates nectar availability based on the relative cover of different plant species in the forest herb layer. Although PNP overestimates actual nectar production, relative differences amongst plots provide a valid and informative way to analyse differences across time and space. Our results show that the forest herb layer has a large potential for providing nectar for wild pollinator communities, which is greatest in spring, with an average PNP of almost 16 g sugar/m2/year. However, this potential has drastically declined (mean plot‐level decline >24%). Change in light availability, associated with shifts in canopy structure and canopy composition, is the key driver of temporal PNP changes. Synthesis. Our study shows that if management activities are carefully planned to sustain nectar‐producing plant species for wild pollinators, European forest herb layers and European forests as a whole can play key roles in sustaining wild pollinator populations.

Limited effects of population age on the genetic structure of spatially isolated forest herb populations in temperate Europe.

Due to multiple land-cover changes, forest herb populations residing in forest patches embedded in agricultural landscapes display different ages and, thus, experience differences in genetic exchange, mutation accumulation and genetic drift. The extent of divergence in present-day population genetic structure among these populations of different ages remains unclear, considering their diverse breeding systems and associated pollinators. Answering this question is essential to understand these species' persistence, maintenance of evolutionary potential and adaptability to changing environments. We applied a multi-landscape setup to compare the genetic structure of forest herb populations across forest patches of different ages (18-338 years). We studied the impact on three common slow-colonizer herb species with distinct breeding systems and associated pollinators: Polygonatum multiflorum (outcrossing, long-distance pollinators), Anemone nemorosa (outcrossing, short-distance pollinators) and Oxalis acetosella (mixed breeding). We aimed to assess if in general older populations displayed higher genetic diversity and lower differentiation than younger ones. We also anticipated that P. multiflorum would show the smallest while O. acetosella the largest difference, between old and young populations. We found that older populations had a higher observed heterozygosity (H o) but a similar level of allelic richness (A r) and expected heterozygosity (H e) as younger populations, except for A. nemorosa, which exhibited higher A r and H e in younger populations. As populations aged, their pairwise genetic differentiation measured by D PS decreased independent of species identity while the other two genetic differentiation measures showed either comparable levels between old and young populations (G" ST) or inconsistency among three species (cGD). The age difference of the two populations did not explain their genetic differentiation. Synthesis: We found restricted evidence that forest herb populations with different ages differ in their genetic structure, indicating that populations of different ages can reach a similar genetic structure within decades and thus persist in the long term after habitat disturbance. Despite their distinct breeding systems and associated pollinators, the three studied species exhibited partly similar genetic patterns, suggesting that their common characteristics, such as being slow colonizers or their ability to propagate vegetatively, are important in determining their long-term response to land-cover change.

High-resolution data are necessary to understand the effects of climate on plant population dynamics of a forest herb.

Climate is assumed to strongly influence species distribution and abundance. Although the performance of many organisms is influenced by the climate in their immediate proximity, the climate data used to model their distributions often have a coarse spatial resolution. This is problematic because the local climate experienced by individuals might deviate substantially from the regional average. This problem is likely to be particularly important for sessile organisms like plants and in environments where small-scale variation in climate is large. To quantify the effect of local temperature on vital rates and population growth rates, we used temperature values measured at the local scale (in situ logger measures) and integral projection models with demographic data from 37 populations of the forest herb Lathyrus vernus across a wide latitudinal gradient in Sweden. To assess how the spatial resolution of temperature data influences assessments of climate effects, we compared effects from models using local data with models using regionally aggregated temperature data at several spatial resolutions (≥1 km). Using local temperature data, we found that spring frost reduced the asymptotic population growth rate in the first of two annual transitions and influenced survival in both transitions. Only one of the four regional estimates showed a similar negative effect of spring frost on population growth rate. Our results for a perennial forest herb show that analyses using regionally aggregated data often fail to identify the effects of climate on population dynamics. This emphasizes the importance of using organism-relevant estimates of climate when examining effects on individual performance and population dynamics, as well as when modeling species distributions. For sessile organisms that experience the environment over small spatial scales, this will require climate data at high spatial resolutions.

Habitat fragmentation affects climate adaptation in a forest herb

Abstract Climate change and the resulting increased drought frequencies pose considerable threats to forest herb populations, particularly where additional environmental challenges jeopardize responses to selection. Specifically, habitat fragmentation may hamper climate adaptation by altering the distribution of adaptive genetic variation and may also induce evolutionary changes in mating systems. To assess how habitat fragmentation disrupts climate adaptation, we conducted a common garden experiment with Primula elatior offspring originating from 24 populations sampled along a latitudinal gradient with varying climate and landscape characteristics. We then quantified a range of vegetative, regulatory and reproductive traits under distinct soil moisture regimes to evaluate imprints of local adaptation and phenotypic plasticity. Additionally, we conducted a more extensive field campaign in 60 populations along the same latitudinal gradient to evaluate the potential evolutionary breakdown of reciprocal herkogamy. For large, connected populations, our results demonstrated an evolutionary shift from a strategy in southern populations that seems aligned with drought avoidance—where plants minimize their exposure to dry conditions and optimize photosynthesis—to a drought tolerance strategy in northern populations, where plants are adapted to function despite water scarcity. However, habitat fragmentation disrupted climate clines and the adaptive responses to drought stress in key traits related to growth, biomass allocation and water regulation. Additionally, our findings indicate the onset of evolutionary breakdown in reciprocal herkogamy and divergence in other key flower traits. The disruption of climate clines, drought responses and adaptations in mating systems contributed to a substantially diminished flowering investment across the distribution range, with the most pronounced effects observed in southern fragmented populations. Synthesis. We present novel empirical evidence of how habitat fragmentation disrupts climate adaptation and drought tolerance in a wide range of traits along the range of the forest herb Primula elatior. These findings emphasize the need to account for habitat fragmentation while designing effective conservation strategies in order to preserve and restore resilient meta‐populations of forest herbs amidst ongoing global changes.

More losses than gains? Distribution models predict species-specific shifts in climatic suitability for European beech forest herbs under climate change

IntroductionHerbaceous plant species constitute an essential element of the flora of European beech (Fagus sylvatica) forests. There is increasing evidence that rapidly changing climate is likely to modify the spatial distribution of plant species. However, we lack understanding of the impact that climate change might have on beech forest herbs across the European continent. We investigated the possible effects of predicted increasing rates of global warming and altered precipitation regimes on 71 forest herbs closely associated with beech forests, but with varying biogeographic and climatic niche attributes.MethodsBy using a total of 394,502 occurrence records and an ensemble of species distribution models (SDMs), we quantified the potential current distribution and future (2061-2080) range shifts in climatic suitability (expressed as occurrence probability, OP) according to two climate change scenarios (moderate SSP2-4.5 and severe SSP5-8.5).ResultsOverall, precipitation of the warmest quarter and temperature seasonality were the most influential predictors in shaping current distribution patterns. For SSP5-8.5 scenario, all studied species experienced significant reductions (52.9% on average) in the total size of highly suitable areas (OP >0.75). However, the magnitude and directions of changes in the climatic suitability were highly species-specific; few species might even increase OP in the future, particularly in case of SSP2-4.5 scenario. The SDMs revealed the most substantial decline of climatic suitability at the trailing edges in southern Europe. We found that climatic suitability is predicted to show unidirectional northward shift and to move toward higher elevations. The gain/loss ratio was generally higher for narrow-ranged species compared to widespread taxa.DiscussionOur findings are contextualized with regards to potential confounding factors (dispersal limitation, microclimatic buffering) that may mitigate or accelerate climate change impacts. Given the low long-distance migration ability, many beech forest herbs are unlikely to track the velocity with which macroclimatic isotherms are moving toward higher latitudes, making this species group particularly vulnerable to climate change.

Trait–micro‐environment relationships of forest herb communities across Europe

AbstractAimThe microclimate and light conditions on the forest floor are strongly modified by tree canopies. Therefore, we need to better consider the micro‐environment when quantifying trait–environment relationships for forest understorey plants. Here, we quantify relationships between micro‐environmental conditions and plant functional traits at the community level, including intraspecific trait variation, and their relationship with microclimate air temperature, light and soil properties.LocationDeciduous temperate forests across Europe.Time period2018.Major taxa studiedHerbaceous vegetation.MethodsWe sampled 225 plots across 15 regions along four complementary gradients capturing both macro‐ and microclimatic conditions including latitude, elevation, forest management and distance to forest edges. We related the community‐weighted mean of five plant functional traits (plant height, specific leaf area [SLA], plant carbon [C], plant nitrogen [N] and plant C:N ratio) across 150 vascular plant species to variation in local microclimate air temperature, light and soil properties. We tested the effect of accounting for intraspecific variation in trait–environment relationships and performed variation partitioning to identify major drivers of trait variation.ResultsMicroclimate temperature, light availability and soil properties were all important predictors of community‐weighted mean functional traits. When light availability and variation in temperature were higher, the herb community often consisted of taller plants with a higher C:N ratio. In more productive environments (e.g. with high soil nitrogen availability), the community was dominated by individuals with resource‐acquisitive traits: high SLA and N but low C:N. Including intraspecific trait variation increased the strength of the trait–micro‐environment relationship, and increased the importance of light availability.Main conclusionsThe trait–environment relationships were much stronger when the micro‐environment and intraspecific trait variation were considered. By locally steering light availability and temperature, forest managers can potentially impact the functional signature of the forest herb‐layer community.

Life forms affect beta-diversity patterns of larch forests in China

Beta-diversity reflects the spatial changes in community species composition which helps to understand how communities are assembled and biodiversity is formed and maintained. Larch (Larix) forests, which are coniferous forests widely distributed in the mountainous and plateau areas in North and Southwest China, are critical for maintaining the environmental conditions and species diversity. Few studies of larch forests have examined the beta-diversity and its constituent components (species turnover and nestedness-resultant components). Here, we used 483 larch forest plots to determine the total beta-diversity and its components in different life forms (i.e., tree, shrub, and herb) of larch forests in China and to evaluate the main drivers that underlie this beta-diversity. We found that total beta-diversity of larch forests was mainly dependent on the species turnover component. In all life forms, total beta-diversity and the species turnover component increased with increasing geographic, elevational, current climatic, and paleoclimatic distances. In contrast, the nestedness-resultant component decreased across these same distances. Geographic and environmental factors explained 20%–25% of total beta-diversity, 18%–27% of species turnover component, and 4%–16% of nestedness-resultant component. Larch forest types significantly affected total beta-diversity and species turnover component. Taken together, our results indicate that life forms affect beta-diversity patterns of larch forests in China, and that beta-diversity is driven by both niche differentiation and dispersal limitation. Our findings help to greatly understand the mechanisms of community assemblies of larch forests in China.

Biomass carbon sink stability of conifer and broadleaf boreal forests: differently associated with plant diversity and mycorrhizal symbionts?

Forest biomass carbon stability is crucial in achieving carbon neutrality in the high-latitude northern hemisphere, and identifying the differences among forest types and decoupling their associations with plant traits and geoclimatic conditions is the basis for precise forest management. We conducted a large-scale field survey in state-owned forest areas in northeastern China, covering a total of 280,000 km2 forest area, 1275 arbor plots (30m × 30m), 5285 shrub plots (5m × 5m), and 7076 herb plots (1m × 1m). We hypothesized that the conifer and broadleaf forest differences in biomass carbon (C) storage and stability (environmental stability to climatic changes-ES and recalcitrant stability to be decomposed-RS) are associated with mycorrhizal abundance (EcM: ectomycorrhizal, AM: arbuscular mycorrhizal, NM-AM: non-mycorrhizal or arbuscular mycorrhizal), taxon diversity traits (richness, Simpson, Shannon-Wiener, and evenness), and structural differences (diameter, height, and density) in the arbor, shrub, and herb layers. Our results showed that (1) conifer forests had 13.1Mg/ha higher C stocks and 30.9% higher RS, but 8.6% lower ES than broadleaf forests (p < 0.05). Trees in conifer forests had 1.5m taller and 2.4cm thicker trees, but 15% less tree density than those in broadleaf forests. Herbs in conifer forests were 14% shorter and 57% denser than in broadleaf forests. (2) The abundance of EcM-symbiont trees in conifer forests was 15% higher than in broadleaf forests, while their EcM-symbiont shrubs and AM-symbiont herbs were 5-6% lower (p < 0.05). Broadleaf forests had 7% higher tree richness and 19% higher herb richness but 9% lower shrub richness than conifer forests (p < 0.05). Tree and herb evenness was 5-6% higher in conifer forests (p < 0.05). (3) Variations of biomass C sink traits could be explained more by plant diversity in conifer forests (7%) than in broadleaf forests (3.4%). Mycorrhizal symbionts could explain more in broadleaf forests (9.7%) than conifer forests (6.7%). In conifer forests, fewer EcM trees (higher AM trees) and AM herbs, higher tree richness were accompanied by higher biomass C storage and ES. Broadleaf forests underwent similar changes, characterized by an elevation in both RS and ES. (4) Our research emphasized that variations in carbon sequestration between conifer and broadleaf forests could be attributed to mycorrhizal symbionts and species diversity besides tree size-related structural differences. Our findings support the precise management of boreal forests to achieve carbon neutrality based on leaf blade types, plant diversity, and mycorrhizal symbionts.

Light more than warming impacts understory tree seedling growth in a temperate deciduous forest

Understanding the distinct impacts of temperature and light on seedling growth is crucial for predicting forest regeneration trajectories under future climate change and forest disturbance. This is because temperature and light can change independently or together, influencing the competitive status of tree seedlings and forest herbs. However, most prior studies tend to explore temperature and light effects either separately or in combination, lacking comprehensive full-factorial designs. Here, we utilized two large-scale full-factorial experiments to explore warming and light effects on tree seedlings growing in mesocosms with herbaceous plants. We found that light increased seedling height, diameter, and biomass, while warming alone had no significant effects. Moreover, we observed that there was an interaction effect between light and warming, where warming increased seedling height, diameter, and biomass under illumination. Understory herbaceous plant cover alone did not affect seedling height, diameter, and biomass, but it decreased seedling biomass when mesocosms were warmed or illuminated. This highlights the importance of considering the indirect negative effects induced by the interaction between forest opening and understory herbaceous plants. By disentangling the effects of increased temperature and light availability on understory seedling height, diameter, and biomass, our results contribute valuable knowledge for future forest management. It is imperative to carefully control the size of the gaps artificially created to facilitate the understory regeneration. Small gaps are recommended considering that the herbaceous plants may interact with both warming and light to negatively affect seedlings.

Mycoheterotrophic plants as indicators of post-agricultural forest regeneration: abundance of Hypopitys monotropa and Monotropa uniflora in post-agricultural forests changes through time

Herbaceous layers in second-growth forests are shaped by past land use. Disturbances such as agriculture may impact populations of mycoheterotrophs, non-photosynthetic mycorrhizal plants that obtain carbon from fungal networks by altering mycorrhizal communities or removing trees they derive carbon from. I tested the hypotheses that two mycoheterotrophic forest herbs increase in abundance during succession and become most common in older forests as plant communities reassemble through time. Distributions of Hypopitys monotropa and Monotropa uniflora were sampled in Athens County, Ohio, USA. I surveyed populations in a 40-site post-agricultural forest chronosequence with five upland and five valley sites in each of four age classes: 40–60, 61–80, 81–100, and &gt;130 years since canopy closure. Aspect and elevation were measured to assess environmental influence. Both H. monotropa and M. uniflora were most common in older stands with EM tree-rich canopy composition and west- or south-facing aspects, indicating influence of historical, biotic, and edaphic factors. Hypopitys was exclusive to forests &gt;80 years old, while M. uniflora was present in younger stands. Abundance of both species was also significantly predicted by Fagaceae basal area. Because EM trees were also most abundant in south- and west-facing uplands, environmental influence appears to be mediated through canopy composition.

Canopy gaps are less susceptible to disturbance-related and invasive herbs than clear-cuts: Temporal changes in the understorey after experimental silvicultural treatments

Forest management has a major impact on the understorey vegetation, with the intensity and type of the applied silvicultural treatments driving variable vegetation responses. We compared understorey variables across different experimental silvicultural treatments in a temperate oak-hornbeam forest in Central Hungary. Five treatment types were used in six replicates representing rotation and selection silvicultural systems: control (C), clear-cutting (CC), gap-cutting (G), preparation cutting (P), and retention tree group (R). The response of several understorey variables was investigated to the treatments in the first six years after their implementation in 2014.We assessed how understorey variables change in response to different forestry treatments, how these responses vary with time, and how game exclusion affects them. We then evaluated how well the treatments can preserve the forest character of the vegetation.We found a large temporal variability in the understorey variables over the study period. In all cases, the interventions led to an initial increase in species richness, followed by a decline later, where the regeneration layer started to close. The regeneration layer grew most intensively in G and CC. At the end of the study, R had the highest average species number, comprising a heterogeneous group of perennial forb species. The interventions all resulted in a rapid increase in total herb layer cover, mainly in favour of graminoid and perennial species. The extent of cover increase depended primarily on the amount of additional light received (CC > G > P > R > C). Turnover and beta diversity values also decreased in a similar order. The effect of game exclusion was especially pronounced in the case of the CC and G, where game browsing significantly slowed the regeneration outside the fences. The most significant changes in almost all variables were in the CC. It had the highest number of indicator species, many of them annual, disturbance-related, and invasive. G preserved the forest character of the vegetation better and proved to be less susceptible to the mass appearance of disturbance-related and invasive herbaceous species.Increasing the share of continuous cover forestry methods is crucial to preserve the forest herb layer. Rotation forestry with large cutting areas is not recommended or should be kept at low landscape rates, as these areas are highly exposed to disturbance-related and invasive species. Leaving retention tree groups can be key to the survival of numerous forest plant species.

Forest management drives evolution of understorey herbs

Forest management has a strong impact on the forest structure and subsequently on the biotic and abiotic forest understorey environment. Forest understorey herbs can thus be expected to exhibit an evolutionary response to management-induced environmental variation (provided sufficient time for adaptation), but this has been little tested to date.Here we use one common garden, to test for genetically based variation in phenotypic traits in populations of forest herbs sampled along a forest management intensity gradient, covering protected areas, selection forests and age-class forests. Five different herbaceous species were sampled from 70 to 100 populations in three regions in Germany and were tested for genetically based variation in flowering start, proportion of flowering ramets, and plant height. Additionally, we performed structural equation modelling to study how forest management drives trait differentiation via its effects on the microenvironment.Our results show the studied forest understorey herbs varied genetically in the measured functional and phenological traits among the sampled populations. Forest management likely affected the traits in various directions and strengths depending on the species, either directly through variation in forest structural attributes or indirectly through changes in the microclimatic environment on the forest floor.In sum, we show that forest management can have evolutionary consequences for forest understorey plants. In an applied context, diverse forest management actions within landscapes are recommended as it creates heterogeneity that selects for different plant traits and thus helps conserving genetic diversity.

Spring temperature drives phenotypic selection on plasticity of flowering time.

In seasonal environments, a high responsiveness of development to increasing temperatures in spring can infer benefits in terms of a longer growing season, but also costs in terms of an increased risk of facing unfavourable weather conditions. Still, we know little about how climatic conditions influence the optimal plastic response. Using 22 years of field observations for the perennial forest herb Lathyrus vernus, we assessed phenotypic selection on among-individual variation in reaction norms of flowering time to spring temperature, and examined if among-year variation in selection on plasticity was associated with spring temperature conditions. We found significant among-individual variation in mean flowering time and flowering time plasticity, and that plants that flowered earlier also had a more plastic flowering time. Selection favoured individuals with an earlier mean flowering time and a lower thermal plasticity of flowering time. Less plastic individuals were more strongly favoured in colder springs, indicating that spring temperature influenced optimal flowering time plasticity. Our results show how selection on plasticity can be linked to climatic conditions, and illustrate how we can understand and predict evolutionary responses of organisms to changing environmental conditions.