Forest Understory Plant Communities Research Articles

Inferring ecological selection from multidimensional community trait distributions along environmental gradients.

Understanding the drivers of community assembly is critical for predicting the future of biodiversity and ecosystem services. Ecological selection ubiquitously shapes communities by selecting for individuals with the most suitable trait combinations. Detecting selection types on key traits across environmental gradients and over time has the potential to reveal the underlying abiotic and biotic drivers of community dynamics. Here, we present a model-based predictive framework to quantify the multidimensional trait distributions of communities (community trait spaces), which we use to identify ecological selection types shaping communities along environmental gradients. We apply the framework to over 3600 boreal forest understory plant communities with results indicating that directional, stabilizing, and divergent selection all modify community trait distributions and that the selection type acting on individual traits may change over time. Our results provide novel and rare empirical evidence for divergent selection within a natural system. Our approach provides a framework for identifying key traits under selection and facilitates the detection of processes underlying community dynamics.

Peeking under the canopy: anomalously short fire-return intervals alter subalpine forest understory plant communities.

Climate change is driving changes in disturbance regimes world-wide. In forests adapted to infrequent, high-severity fires, recent anomalously short fire-return intervals (FRIs) have resulted in greatly reduced postfire tree regeneration. However, effects on understory plant communities remain unexplored. Understory plant communities were sampled in 31 plot pairs across Greater Yellowstone (Wyoming, USA). Each pair included one plot burned at high severity twice in < 30 yr and one plot burned in the same most recent fire but not burned previously for > 125 yr. Understory communities following short-interval fires were also compared with those following the previous long-interval fire. Species capable of growing in drier conditions and in lower vegetation zones became more abundant and regional differences in plant communities declined following short-interval fire. Dissimilarity between plot pairs increased in mesic settings and decreased with time since fire and postfire winter snowfall. Reduced postfire tree density following short-interval fire rather than FRI per se affected the occurrence of most plant species. Anomalously short FRIs altered understory plant communities in space and time, with some indications of community thermophilization and regional homogenization. These and other shifts in understory plant communities may continue with ongoing changes in climate and fire across temperate forests.

Modeled Vegetation Community Trajectories: Effects from Climate Change, Atmospheric Nitrogen Deposition, and Soil Acidification Recovery.

Forest understory plant communities in the United States harbor most of the vegetation diversity of forests and are often sensitive to changes in climate and atmospheric deposition of nitrogen (N). As temperature increases from human-caused climate change and soils recover from long term atmospheric deposition of N and sulfur (S), it is unclear how these important ecosystem components will respond. We used the newly developed US-PROPS model - based on species response functions for over 1,500 species - to evaluate the potential impacts of atmospheric N deposition and climate change on species occurrence probability for a case study in the forested ecosystems of the Great Smoky Mountains National Park (GRSM), an iconic park in the southeastern United States. We evaluated six future scenarios from various combinations of two potential recoveries of soil pH (no change, +0.5 pH units) and three climate futures (no change, +1.5, +3.0 deg C). Species critical loads (CLs) of N deposition and projected responses for each scenario were determined. Critical loads were estimated to be low (< 2 kg N/ha/yr) to protect all species under current and expected future conditions across broad regions of GRSM and these CLs were exceeded at large spatial extents among scenarios. Northern hardwood, yellow pine, and chestnut oak forests were among the most N-sensitive vegetation map classes found within GRSM. Potential future air temperature conditions generally led to decreases in the maximum occurrence probability for species. Therefore, CLs were considered "unattainable" in these situations because the specified level of protection used for CL determination (i.e., maximum occurrence probability under ambient conditions) was not attainable. Although some species showed decreases in maximum occurrence probability with simulated increases in soil pH, most species were favored by increased pH. The importance of our study is rooted in the methodology described here for establishing regional CLs and for evaluating future conditions, which is transferable to other national parks in the U.S. and in Europe where the original PROPS model was developed.

Chronic browsing by an introduced mammalian herbivore in a tropical island alters species composition and functional traits of forest understory plant communities

AbstractMammalian herbivores have large‐scale impacts on vegetation, altering structure, and species composition, especially in tropical grasslands and savannas. However, there is limited understanding of the potential impacts of mammalian herbivores in tropical wet forests, where they are typically less abundant. We investigated the effects of an introduced mammalian herbivore chital (Axis axis) on vegetation structure, composition and leaf functional traits of tropical evergreen forests of the Andaman Islands, India. Across seven islands, representing a gradient of herbivore densities, increasing chital presence was associated with decreased understory richness, understory density and adult tree richness, but was not related to adult tree density or size class distributions. We also found a significant decrease in community level leaf palatability traits (specific leaf area decreased and leaf thickness increased) with increasing chital habitat use. This community level shift in leaf trait values was better explained by intraspecific variation in leaf traits across islands rather than changes in species composition. In summary, we show persistent long‐term impacts of an introduced mammalian herbivore on understory tropical tree communities although there is little impact on adult tree communities. Our results also show that functional traits of species can be altered in response to novel herbivory, even at herbivore densities where there are no detectable impacts on adult forest structure or composition. Such altered functional traits may potentially alter ecosystem functioning in these forests even without changes in vegetation structure.Abstract in Hindi is available with online material.

Leaf trait variability explains how plant community composition changes under the intense pressure of deer herbivory

AbstractThere is growing evidence indicating that intraspecific trait variation (ITV) plays a prominent role in determining plant community composition. In this study, we investigated the significance of ITV in identifying the processes that shape the development of forest understory plant communities in an area in which excessive herbivory by overabundant deer populations has been an issue. We measured functional traits (height, leaf area, leaf mass per area, leaf dry matter content, and leaf toughness) of 13 forest floor plant species growing in two contrasting sites in which deer were overabundant or had been completely excluded, respectively, and examined the relationships between species dominance and interspecific trait variation and ITV. We detected variations in the trait plasticity of size and morphological features among species, reflecting species‐specific strategies in response to herbivory and consequently observed marked differences in species composition and diversity at the two sites. Notably, small stature and less costly leaves were identified as important traits that enable plants to persist under conditions of intense deer herbivory probably because such plants are less exposed to herbivory and simultaneously optimize carbon assimilation. In contrast, large‐sized plants with costly leaves tended to dominate the plant community in the absence of deer herbivory, as these plants would tend to gain a competitive advantage with respect to the acquisition of space and light. The findings of this study indicate that a focus on ITV represents a promising approach for determining the general ecological patterns of plant communities responding to environmental disturbance.

Plant traits as indicators of recovery of reclaimed wellsites in forested areas: Slow but directional succession trajectory

Trait-based approaches can provide a generalizable mechanistic understanding of complex post-disturbance succession dynamics of plant communities. Much of our knowledge regarding successional trajectories of functional trait composition come from observations of natural disturbances that leave physical and biological legacy on site for self regeneration. We lack, however, understanding of the long-term recovery in severely degraded lands following reclamation through active revegetation. To address this gap, we examined changes in trait composition of forest understory plant communities in reclaimed oil and gas wellsites using chronosequence data (7–48 years since reclamation) to assess recovery towards that of post-harvest and natural post-fire reference forest sites. We used multiple traits associated with resource acquisition (e.g., fast growth) and performance ability metrics (e.g., shade-tolerance) with putative environmental factors to evaluate the trait-environment relationships underlying plant community recovery in reclaimed sites. We found an overall directional change in functional composition with time since reclamation towards that observed in reference sites, but even older reclaimed sites remained significantly different from reference sites. This could be related to differential trajectory patterns among traits where some trait values progressed towards those of reference (long-distance dispersal by wind decreased with time since reclamation) whereas some fast-resource acquisition traits and exotic species showed no change and remained dominant in older reclaimed sites. The strong link between traits and environment suggests a significant influence of time and subsequent developing site conditions (e.g., canopy cover) as well as enduring legacies of wellsite operation/reclamation (e.g., high soil bulk density and exotic species) on functional composition. Knowledge of functional composition trajectories in severely degraded ecosystems, such as wellsites, could improve our understanding of recovery processes and inform more effective reclamation practices by identifying putative underlying environmental factors and specific ecological attributes that may delay successful recovery.

Threshold effects of air pollution and climate change on understory plant communities at forested sites in the eastern United States

Forest understory plant communities in the eastern United States are often diverse and are potentially sensitive to changes in climate and atmospheric inputs of nitrogen caused by air pollution. In recent years, empirical and processed-based mathematical models have been developed to investigate such changes in plant communities. In the study reported here, a robust set of understory vegetation response functions (expressed as version 2 of the Probability of Occurrence of Plant Species model for the United States [US-PROPS v2]) was developed based on observations of forest understory and grassland plant species presence/absence and associated abiotic characteristics derived from spatial datasets. Improvements to the US-PROPS model, relative to version 1, were mostly focused on inclusion of additional input data, development of custom species-level input datasets, and implementation of methods to address uncertainty. We investigated the application of US-PROPS v2 to evaluate the potential impacts of atmospheric nitrogen (N) and sulfur (S) deposition, and climate change on forest ecosystems at three forested sites located in New Hampshire, Virginia, and Tennessee in the eastern United States. Species-level N and S critical loads (CLs) were determined under ambient deposition at all three modeled sites. The lowest species-level CLs of N deposition at each site were between 2 and 11 kg N/ha/yr. Similarly, the lowest CLs of S deposition, based on the predicted soil pH response, were less than 2 kg S/ha/yr among the three sites. Critical load exceedance was found at all three model sites. The New Hampshire site included the largest percentage of species in exceedance. Simulated warming air temperature typically resulted in lower maximum occurrence probability, which contributed to lower CLs of N and S deposition. The US-PROPS v2 model, together with the PROPS-CLF model to derive CL functions, can be used to develop site-specific CLs for understory plants within broad regions of the United States. This study demonstrates that species-level CLs of N and S deposition are spatially variable according to the climate, light availability, and soil characteristics at a given location. Although the species niche models generally performed well in predicting occurrence probability, there remains uncertainty with respect to the accuracy of reported CLs. As such, the specific CLs reported here should be considered as preliminary estimates.

Fire severity and changing composition of forest understory plant communities

AbstractQuestionsGradients of fire severity in dry conifer forests can be associated with variation in understory floristic composition. Recent work in dry conifer forests in California, USA, has suggested that more severely burned stands contain more thermophilic taxa (those associated with warmer and drier conditions), and that forest disturbance may therefore accelerate floristic shifts already underway due to climate change. However, it remains unknown how rapidly thermophilic taxa shifts occur following disturbance, how long such shifts are likely to persist, and how different thermophilic post‐disturbance communities are from pre‐disturbance communities.LocationColorado Front Range, USA.MethodsWe investigated these questions using a unique 15‐year vegetation plot dataset that captures pre‐ and post‐fire understory community composition across a gradient of fire severity in dry conifer forests, classifying taxa using the biogeographic affinity concept.ResultsThermophilization (defined here as a decrease in the ratio of cool‐mesic taxa to warm‐xeric taxa, based on biogeographic affinity of paleobotanical lineages) was observed as early as one year post‐fire for all fire severity classes, but was stronger at sites that burned at higher severity. The ratio of cool‐mesic to warm‐xeric taxa recovered to pre‐fire levels within 10 years in stands that burned at low severity, but not in stands that burned at moderate or high severity. The process of thermophilization after high‐severity fire appears to be driven primarily by the gain of warm‐xeric taxa that were absent before the fire, but losses of cool‐mesic taxa, which did not return during the duration of the study, also played a role.ConclusionsDecreases in canopy cover appear to be a main contributor to understory thermophilization. Fine‐scale heterogeneity in post‐fire forest structure is likely an important driver of floristic diversity, creating the microclimatic variation necessary to maintain floristic refugia for species mal‐adapted to increasingly warm and dry conditions.

Half a Century of Multiple Anthropogenic Stressors has Altered Northern Forest Understory Plant Communities

Boreal forests form the largest and least disturbed forest biome in the northern hemisphere. However, anthropogenic pressure from intensified forest management, eutrophication, and climate change may alter the ecosystem functions of understory vegetation and services boreal forests provide. Swedish forests span long gradients of climate, nitrogen deposition, and management intensity. This makes them ideal to study how the species composition and functions of other, more pristine, boreal forests might change under increased anthropogenic pressure. Moreover, the National Forest Inventory (NFI) has collected systematic data on Swedish forest vegetation since the mid-20th century. We use this data to quantify changes in vegetation types between two periods, 1953-1962 and 2003-2012. The results show changes in forest understory vegetation since the 1950s at scales not previously documented in the boreal biome. The spatial extent of most vegetation types changed significantly. Shade-adapted and nutrient-demanding species (those with high specific leaf area) have become more common at the expense of light-demanding and nutrient-conservative (low specific leaf area) species. The cover of ericaceous dwarf shrubs decreased dramatically. These effects were strongest where anthropogenic impacts were greatest, suggesting links to drivers such as nitrogen deposition and land-use change. These changes may impact ecosystem functions and services via effects on higher trophic levels and faster plant litter decomposition in the expanding vegetation types. This, in turn, may influence nutrient dynamics, and consequently ecosystem productivity and carbon sequestration.

Half a century of multiple anthropogenic stressors has altered northern forest understory plant communities.

Boreal forests form the largest and least disturbed forest biome in the northern hemisphere. However, anthropogenic pressure from intensified forest management, eutrophication, and climate change may alter the ecosystem functions of understory vegetation and services boreal forests provide. Swedish forests span long gradients of climate, nitrogen deposition, and management intensity. This makes them ideal to study how the species composition and functions of other, more pristine, boreal forests might change under increased anthropogenic pressure. Moreover, the National Forest Inventory (NFI) has collected systematic data on Swedish forest vegetation since the mid-20th century. We use this data to quantify changes in vegetation types between two periods, 1953-1962 and 2003-2012. The results show changes in forest understory vegetation since the 1950s at scales not previously documented in the boreal biome. The spatial extent of most vegetation types changed significantly. Shade-adapted and nutrient-demanding species (those with high specific leaf area) have become more common at the expense of light-demanding and nutrient-conservative (low specific leaf area) species. The cover of ericaceous dwarf shrubs decreased dramatically. These effects were strongest where anthropogenic impacts were greatest, suggesting links to drivers such as nitrogen deposition and land-use change. These changes may impact ecosystem functions and services via effects on higher trophic levels and faster plant litter decomposition in the expanding vegetation types. This, in turn, may influence nutrient dynamics, and consequently ecosystem productivity and carbon sequestration.

Plant richness and composition in hardwood forest understories vary along an acidic deposition and soil-chemical gradient in the northeastern United States

AimsA century of atmospheric deposition of sulfur and nitrogen has acidified soils and undermined the health and recruitment of foundational tree species in the northeastern US. However, effects of acidic deposition on the forest understory plant communities of this region are poorly documented. We investigated how forest understory plant species composition and richness varied across gradients of acidic deposition and soil acidity in the Adirondack Mountains of New York State.MethodsWe surveyed understory vegetation and soils in hardwood forests on 20 small watersheds and built models of community composition and richness as functions of soil chemistry, nitrogen and sulfur deposition, and other environmental variables.ResultsCommunity composition varied significantly with gradients of acidic deposition, soil acidity, and base cation availability (63% variance explained). Several species increased with soil acidity while others decreased. Understory plant richness decreased significantly with increasing soil acidity (r = 0.60). The best multivariate regression model to predict richness (p < 0.001, adjusted-R2 = 0.60) reflected positive effects of pH and carbon-to-nitrogen ratio (C:N).ConclusionsThe relationship we found between understory plant communities and a soil-chemical gradient, suggests that soil acidification can reduce diversity and alter the composition of these communities in northern hardwood forests exposed to acidic deposition.

Regional Patterns of Floristic Diversity and Composition in Forests Invaded by Garlic Mustard (Alliaria petiolata)

The impacts of invasive species on native plant communities are often studied on small spatial scales but may vary across regionally heterogeneous landscapes. Comparisons of vegetation across several similar sites with and without an invasive species present can be logistically challenging but highly informative to both scientists and land managers. We examined regional geographic variation in the diversity and composition of 8 replicate northeastern forest-understory plant communities invaded by the non-native species Alliaria petiolata (Garlic Mustard). Despite variation in underlying soil conditions and horizon development, several native species and their associated functional groups were either negatively or positively associated with Garlic Mustard invasion at the regional scale, and soil moisture and pH were higher in invaded plots across all sites. Most tree species were less common at invaded sites, but high tree-seedling abundances at some sites led to regionally higher seedling abundance in the presence of Garlic Mustard. Our study highlights the importance of species-specific responses, as well as site-specific soil conditions, for better understanding potential impacts of invasion.

Multiple environmental changes drive forest floor vegetation in a temperate mountain forest.

Human‐induced changes of the environment and their possible impacts on temperate forest understory plant communities have been examined in many studies. However, the relative contribution of individual environmental factors to these changes in the herb layer is still unclear. In this study, we used vegetation survey data covering a time period of 21 years and collected from 143 permanent plots in the Northern Limestone Alps, Austria. Data on soil chemistry (49 plots), light condition (51 plots), soil temperature and moisture (four and six plots), disturbance (all plots), climate (one station in a clearing area), and airborne sulfur (S) and nitrogen (N) deposition (two forest stands) were available for analyses. We used these data together with plot mean Ellenberg indicator values in a path analysis to attribute their relative contributions to observed vegetation changes. Our analysis reveals a strong directional shift of the forest understory plant community. We found strong evidence for a recovery of the ground‐layer vegetation from acidification as response to decreased S deposition. We did not observe a community response to atmospheric N deposition, but we found a response to altered climatic conditions (thermophilization and drying). The path analysis revealed that changes in the light regime, which were related to small‐scale disturbances, had most influence on herb layer community shifts. Thermophilization and drying were identified as drivers of understory community changes independent of disturbance events.

Stand- and plot-level changes in a boreal forest understory community following wildfire

Background: Boreal forest understory plant communities are known to be resilient to fire – the species composition of stands after a fire is quite similar to the pre-fire composition. However, we know little about recovery of individual plants within particular locations in forest stands (i.e. plot-level changes) since we usually do not have pre-fire data for plots.Aims: We wanted to determine whether species recruited into the same or different locations in a Pinus banksiana stand that experienced a severe wildfire.Methods: We used pre-existing permanent plots to evaluate the cover of understory after an unplanned wildfire.Results: Across the entire stand nine of 47 species showed a significant change in cover. The largest change in a plant functional group was in the mosses, with all species present before fire being eliminated. There was no change in species diversity or total cover. At the plot level, species composition showed a much greater change. An average of 47% of the species present in a plot before the fire were absent in the same plot after the fire, and the total species turnover in plots was 88% of the species present before the fire. The plots showed a similar shift in species composition.Conclusions: These results confirm that boreal forest communities show a high degree of resilience to fire, but within a forest stand species will be found in different locations following fire, potentially exposing them to a different set of biotic and abiotic conditions in these new locations.

Rapid Increases in forest understory diversity and productivity following a mountain pine beetle (Dendroctonus ponderosae) outbreak in pine forests.

The current unprecedented outbreak of mountain pine beetle (Dendroctonus ponderosae) in lodgepole pine (Pinus contorta) forests of western Canada has resulted in a landscape consisting of a mosaic of forest stands at different stages of mortality. Within forest stands, understory communities are the reservoir of the majority of plant species diversity and influence the composition of future forests in response to disturbance. Although changes to stand composition following beetle outbreaks are well documented, information on immediate responses of forest understory plant communities is limited. The objective of this study was to examine the effects of D. ponderosae-induced tree mortality on initial changes in diversity and productivity of understory plant communities. We established a total of 110 1-m2 plots across eleven mature lodgepole pine forests to measure changes in understory diversity and productivity as a function of tree mortality and below ground resource availability across multiple years. Overall, understory community diversity and productivity increased across the gradient of increased tree mortality. Richness of herbaceous perennials increased with tree mortality as well as soil moisture and nutrient levels. In contrast, the diversity of woody perennials did not change across the gradient of tree mortality. Understory vegetation, namely herbaceous perennials, showed an immediate response to improved growing conditions caused by increases in tree mortality. How this increased pulse in understory richness and productivity affects future forest trajectories in a novel system is unknown.

White‐tailed deer (Odocoileus virginianus) positively affect the growth of mature northern red oak (Quercus rubra) trees

Understanding and predicting the effects of deer (Cervidae) on forest ecosystems present significant challenges in ecosystem ecology. Deer herbivory can cause large changes in the biomass and species composition of forest understory plant communities, including early life‐cycle trees (i.e., seedlings and saplings). Such changes can impact juvenile to adult transitions and the future age structure and species composition of mature forests. Changes to understory vegetation also impact flow of energy and nutrients in forest ecosystems. Studies examining the influence of deer on mature trees, however, are rare and rely on extrapolating effects from early life cycle stages of trees. We tested the hypothesis that the absence of deer would result in an increase in the growth rate of mature trees by examining the impact of white‐tailed deer (Odocoileus virginianus) on mature canopy trees. We examined incremental growth in individuals of Quercus rubra, an important component of temperate deciduous forests in North America, inside and outside 16‐year deer exclosures in eastern U.S. deciduous forests. We found that adult trees inside exclosures grew less than those directly exposed to deer. Our findings highlight the indirect effects of white‐tailed deer on the growth of adult individuals of Q. rubra in a way opposite of what would be expected from previous studies based on immature or understory tree populations. We suggest the increased growth of adult trees in the presence of deer may be explained by increased nutrient inputs through deer fecal and urine deposits and the alteration of the competitive environment belowground through the reduction of understory vegetation by browsing. Underscoring the ecological and demographic importance of adult trees in forest ecosystems, results from this study suggest the direct and indirect effects of deer on mature trees should not be overlooked.

Floristic Diversity in Managed Forests: Demography and Physiology of Understory Plants Following Disturbance in Southern New England Forests

Current interest in the conservation of biodiversity is generating a need for forest management and silvicultural techniques designed to maintain the integrity of ecosystems while satisfying society's need for timber resources. The conservation of forest understory plant communities should be a major emphasis of this effort as they contain the majority of plant diversity in most U.S. forests and play a significant role in many ecosystem functions. This article reviews the literature regarding plant responses to disturbance—most importantly changes in light environments—and applies that information to forest management. A comparison of developmental plasticity and rapid acclimation as response pathways is used to discuss plant level responses. At the landscape level, diversity models, silviculture treatments, and site characteristics are used to discuss changes in understory community composition following disturbance. Results of ongoing research on the effects of forest management on floristic patterns for southern New England forests are summarized.

Leaf longevity of Oxalis acetosella (Oxalidaceae) in the Catskill Mountains, New York, USA

Leaf habit correlates with multiple physiological traits. Understanding ecophysiology is therefore dependent on knowledge of leaf habit. A variety of leaf habits exists within forest understory plant communities. Oxalis acetosella is one such understory plant and has long been considered a wintergreen, meaning that it keeps a set of leaves for one full year, replacing them with a new set during spring. To assess the leaf habit of O. acetosella and place it into a classification scheme of leaf habits, leaves of four populations of O. acetosella were repeatedly censused for two years in a northern hardwood forest of the Catskill Mountains, New York, USA. New leaves developed and old leaves senesced throughout the year, yielding a continual replacement of leaves and a summer peak in leaf number. Leaves that developed in the fall and winter had longer maximum life spans than leaves that developed during the summer. The name "seasonalgreen" is suggested to describe the continual development, senescence, and presence of leaves and annual peak in leaf number within O. acetosella. The functional significance of this leaf habit in this species and the possibility of its presence in other species deserve further study.