Linking forest shade tolerance and soil moisture in North America

Increasing occurrence of droughts is a major environmental concern, however its consequences on forested ecosystems are not fully understood at the landscape level. Here we link the forest shade tolerance index to soil moisture in the North America using the U.S. and Quebec forest inventories. We report a significant decrease of shade tolerance index along the hydric–mesic–xeric soil transition in most of the area considered except three subtropical/tropical ecoregions of the Southeastern U.S. We conclude that droughts may alter forest succession, and in particular decrease the role of forest gap dynamics and dominance of the shade-tolerant species in mature forests.     

US forest response to projected climate‐related stress: a tolerance perspective

Although it is widely recognized that climate change will require a major spatial reorganization of forests, our ability to predict exactly how and where forest characteristics and distributions will change has been rather limited. Current efforts to predict future distribution of forested ecosystems as a function of climate include species distribution models (for fine‐scale predictions) and potential vegetation climate envelope models (for coarse‐grained, large‐scale predictions). Here, we develop and apply an intermediate approach wherein we use stand‐level tolerances of environmental stressors to understand forest distributions and vulnerabilities to anticipated climate change. In contrast to other existing models, this approach can be applied at a continental scale while maintaining a direct link to ecologically relevant, climate‐related stressors. We first demonstrate that shade, drought, and waterlogging tolerances of forest stands are strongly correlated with climate and edaphic conditions in the conterminous United States. This discovery allows the development of a tolerance distribution model (TDM), a novel quantitative tool to assess landscape level impacts of climate change. We then focus on evaluating the implications of the drought TDM. Using an ensemble of 17 climate change models to drive this TDM, we estimate that 18% of US ecosystems are vulnerable to drought‐related stress over the coming century. Vulnerable areas include mostly the Midwest United States and Northeast United States, as well as high‐elevation areas of the Rocky Mountains. We also infer stress incurred by shifting climate should create an opening for the establishment of forest types not currently seen in the conterminous United States.     

Modelling secondary succession of neotropical mangroves: Causes and consequences of growth reduction in pioneer species

Mangrove forest structure is the result of interactions between species responses to abiotic stress factors, disturbance, dispersal and competition. The combination of abiotic conditions and disturbance history may determine the growth potential of the species, whereas dispersal, competition and external or biogenic changes in abiotic conditions may tune their succession. Even in forests with only a few species, this set of factors can create multiple species compositions. For example, pure stands of the mangrove Laguncularia are common in newly colonized areas, but they may evolve into mixed stands dominated by Rhizophora or Avicennia. We demonstrate such an evolution with a field study of mangroves established in abandoned rice fields. We use an individual-based simulation model to understand how species-dependent dispersal, growth rate and shade tolerance lead to the observed patterns in horizontal and vertical forest structure. We find that the initially dominating species will be gradually replaced in the canopy. This alteration cannot be explained exclusively by shade tolerance of the succeeding species. Rather, it is essential to assume that the height growth rate of the pioneer species slows down relative to those of the later species. A decrease in nutrient availability may be responsible for these relative changes in the growth rates of different species.     

Secondary succession in disturbed Pinus-Quercus forests in the highlands of Chiapas,Mexico

We present floristic and structural data on serai plant communities (Old-Field, Grassland, Shrubland, and Early Successional, Mid-Successional, and Mature Forest) resulting from the current land use pattern in the Pinus-Quercus forests in the highlands of Chiapas, Mexico. The number of species ranged from 24 in the Old-Field stage up to 100 in the Mature Forest, and the shrub layer was richest in all the forested stands. An almost complete floristic replacement exists between the open and the forested communities. Quercus and Pinus dominate the canopy of old-growth stands, but their recruitment does not occur or is very low in the shade. Vigorous seedlings and saplings of these species are found only in the open serai stages or in forest gaps. Demographic and phytosociological data are used to indicate the successional role of dominant species in the forested stages. Some implications of the regional land use patterns for conservation and succession are discussed.     

Tree species richness promotes productivity in temperate forests through strong complementarity between species

Understanding the link between biodiversity and ecosystem functioning (BEF) is pivotal in the context of global biodiversity loss. Yet, long-term effects have been explored only weakly, especially for forests, and no clear evidence has been found regarding the underlying mechanisms. We explore the long-term relationship between diversity and productivity using a forest succession model. Extensive simulations show that tree species richness promotes productivity in European temperate forests across a large climatic gradient, mostly through strong complementarity between species. We show that this biodiversity effect emerges because increasing species richness promotes higher diversity in shade tolerance and growth ability, which results in forests responding faster to small-scale mortality events. Our study generalises results from short-term experiments in grasslands to forest ecosystems and demonstrates that competition for light alone induces a positive effect of biodiversity on productivity, thus providing a new angle for explaining BEF relationships.     

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

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

Spatial pattern and process in forest stands within the Virginia piedmont

Abstract. Question: Underlying ecological processes have often been inferred from the analysis of spatial patterns in ecosystems. Using an individual‐based model, we evaluate whether basic assumptions of species’life‐history, drought‐susceptibility, and shade tolerance generate dynamics that replicate patterns between and within forest stands. Location: Virginia piedmont, USA. Method: Model verification examines the transition in forest composition and stand structure between mesic, intermediate and xeric sites. At each site, tree location, diameter, and status were recorded in square plots ranging from 0.25 to 1.0 ha. Model validation examines the simulated spatial pattern of individual trees at scales of 1–25 m within each forest site using a univariate Ripley's K function. Results: 7512 live and dead trees were surveyed across all sites. All sites exhibit a consistent, significant shift in pattern for live trees by size, progressing from a clumped understorey (trees ± 0.1 m in diameter) to a uniform overstorey (trees > 0.25 m). Simulation results reflect not only the general shift in pattern of trees at appropriate scales within sites, but also the general transition in species composition and stand structure between sites. Conclusions: This shift has been observed in other forest ecosystems and interpreted as a result of competition; however, this hypothesis has seldom been evaluated using simulation models. These results support the hypothesis that forest pattern in the Virginia piedmont results from competition involving species’life‐history attributes driven by soil moisture availability between sites and light availability within sites.     

Observed forest trait velocities have not kept pace with hydraulic stress from climate change

The extent to which future climate change will increase forest stress and the amount to which species and forest ecosystems can acclimate or adapt to increased stress is a major unknown. We used high-resolution maps of hydraulic traits representing the diversity in tree drought tolerance across the United States, a hydraulically enabled tree model, and forest inventory observations of demographic shifts to quantify the ability for within-species acclimation and between-species range shifts to mediate climate stress. We found that forests are likely to experience increases in both acute and chronic hydraulic stress with climate change. Based on current species distributions, regional hydraulic trait diversity was sufficient to buffer against increased stress in 88% of forested areas. However, observed trait velocities in 81% of forested areas are not keeping up with the rate required to ameliorate projected future stress without leaf area acclimation.     

Structure of Brazilian Atlantic forests with occurrence of the woolly spider monkey (<Emphasis Type="Italic">Brachyteles hypoxanthus</Emphasis>)

This work examines patches of Atlantic forests in Minas Gerais State where woolly spider monkeys (Brachyteles hypoxanthus) are inhabitants in order to understand the effects of forest structure, diversity, and succession in B. hypoxanthus abundance. This study was carried out in three reserves: Rio Doce State Park (PERD), Mata do Sossego (Sossego), and Serra do Brigadeiro State Park (PESB). Structural studies were carried out by means of point-centered quarter samples in areas used by woolly spider monkeys. Evenness was lower in PERD than in the other two communities. By means of successional functional groups analysis and diametric structure, the PERD forest sample was shown to be in the later, and PESB forest sample in the earlier, stages of secondary succession. We found that tree species evenness and structural traits driven by succession were important factors in determining spider monkey abundance patterns. The PERD forest sample had lower evenness and had lower density of woolly spider monkeys than the Sossego forest sample. The lowest density of these primates was in the PESB forest sample, the earlier successional stage of secondary succession of this study.     

Patterns in potassium dynamics in forest ecosystems

The biotic cycling of potassium (K) in forest systems has been relatively understudied in comparison with nitrogen (N) and phosphorus (P) despite its critical roles in maintaining the nutrition of primary production in forests. We investigated the ecological significance of K in forests from a literature review and data synthesis. We focused on (1) describing patterns of the effects of K availability on aboveground growth and change in foliar tissue of tree species from a variety of forests; and (2) documenting previously unreported relationships between hydrologic losses of K and N in forested watersheds from the Americas. In a review of studies examining tree growth under K manipulations/fertilizations, a high percentage (69% of studies) showed a positive response to increases in K availability in forest soils. In addition, 76% of the tree studies reviewed showed a positive and significant increase in K concentrations in plant tissue after soil K manipulation/fertilization. A meta-analysis on a subset of the reviewed studies was found to provide further evidence that potassium effects tree growth and increased tissue [K] with an effect size of 0.709 for growth and an overall effect size of 0.56. In our review of watershed studies, we observed that concentrations of K typically decreased during growing seasons in streams draining forested areas in the Temperate Zones and were responsive to vegetation disturbance in both temperate and tropical regions. We found a strong relationship (r2 = 0.42-0.99) between concentrations of K and N (another critical plant nutrient) in stream water, suggesting that similar mechanisms of biotic retention may control the flow of these nutrients. Furthermore, K dynamics appear to be unique among the base cations, e.g. calcium, magnesium, and sodium, because the others do not show similar seasonal patterns to K. We suggest that K may be important to the productivity and sustenance of many forests, and its dynamics and ecological significance warrant further study. We suggest that knowledge about the dynamics of this understudied element is imperative for our understanding patterns and processes in forest ecosystems.     

When is a ‘forest’ a savanna,and why does it matter?

Savannas are defined based on vegetation structure, the central concept being a discontinuous tree cover in a continuous grass understorey. However, at the high‐rainfall end of the tropical savanna biome, where heavily wooded mesic savannas begin to structurally resemble forests, or where tropical forests are degraded such that they open out to structurally resemble savannas, vegetation structure alone may be inadequate to distinguish mesic savanna from forest. Additional knowledge of the functional differences between these ecosystems which contrast sharply in their evolutionary and ecological history is required. Specifically, we suggest that tropical mesic savannas are predominantly mixed tree–C₄ grass systems defined by fire tolerance and shade intolerance of their species, while forests, from which C₄ grasses are largely absent, have species that are mostly fire intolerant and shade tolerant. Using this framework, we identify a suite of morphological, physiological and life‐history traits that are likely to differ between tropical mesic savanna and forest species. We suggest that these traits can be used to distinguish between these ecosystems and thereby aid their appropriate management and conservation. We also suggest that many areas in South Asia classified as tropical dry forests, but characterized by fire‐resistant tree species in a C₄ grass‐dominated understorey, would be better classified as mesic savannas requiring fire and light to maintain the unique mix of species that characterize them.     

Does Acacia dealbata express shade tolerance in Mediterranean forest ecosystems of South America?

The distribution of Acacia dealbata Link (Fabaceae) in its non-native range is associated with disturbed areas. However, the possibility that it can penetrate the native forest during the invasion process cannot be ruled out. This statement is supported by the fact that this species has been experimentally established successfully under the canopy of native forest. Nonetheless, it is unknown whether A. dealbata can express shade tolerance traits to help increase its invasive potential. We investigated the shade tolerance of A. dealbata under the canopy of two native forests and one non-native for three consecutive years, as well as its early growth and photosynthetic performance at low light intensities (9, 30, and 70 μmol m⁻²sec⁻¹) under controlled conditions. We found many A. dealbata plants surviving and growing under the canopy of native and non-native forests. The number of plants of this invasive species remained almost constant under the canopy of native forests during the years of study. However, the largest number of A. dealbata plants was found under the canopy of non-native forest. In every case, the distribution pattern varied with a highest density of plants in forest edges decreasing progressively toward the inside. Germination and early growth of A. dealbata were slow but successful at three low light intensities tested under controlled conditions. For all tested light regimes, we observed that in this species, most of the energy was dissipated by photochemical processes, in accordance with the high photosynthetic rates that this plant showed, despite the really low light intensities under which it was grown. Our study reveals that A. dealbata expressed shade tolerance traits under the canopy of native and non-native forests. This behavior is supported by the efficient photosynthetic performance that A. dealbata showed at low light intensities. Therefore, these results suggest that Mediterranean forest ecosystems of South America can become progressively invaded by A. dealbata and provide a basis for estimating the possible impacts that this invasive species can cause in these ecosystems in a timescale.     

Effects of Natural and Human-Assisted Regeneration on Landscape Dynamics in a Korean Pine Forest in Northeast China

Improper forest harvesting can potentially degrade forest ecosystem functions and services. Human-assisted regeneration (e.g., planting) is often used to increase the rate of forest recovery and thereby reduce regeneration failure. Seed dispersal is a fundamental ecological process that can also influence spatio-temporal patterns of forest regeneration. In this study, we investigated the relative contribution of planting and seed dispersal on forest regeneration at landscape scales. Because such influences can be further complicated by timber harvest intensity and seed availability within and around harvested area, we also evaluated the effects of those factors on forest landscape dynamics. We used the forest landscape model LANDIS to simulate the dynamics of Korean pine-broadleaf mixed forests in Northeast China. We considered three factors: timber harvest intensity (3 levels), seed dispersal and whether or not planting was used. The results showed that planting was more important in maintaining the abundance of Korean pine (Pinus koraiensis), a climax keystone species in this region, under the high-intensity harvesting option during early succession. In contrast, seed dispersal was more important during late succession. Korean pine can be successfully regenerated through seed dispersal under low and medium harvest intensities. Our results also indicated that effective natural regeneration will require protecting seed-production trees (seed rain). This study results provide a basis for more effectively managing Chinese temperate forests and possibly other similar ecosystems.     

Simulating the Long-term Response of Tropical Wet Forests to Fragmentation

In the coming decades, a large fraction of the world's tropical forests will be fragmented into remnants surrounded by secondary vegetation, land-use areas, or roads. It is important to develop integrative tools to monitor the evolution of these fragmented ecosystems. We used the individual-oriented and process-based forest growth simulator FORMIND2.0 to investigate the spatial and temporal effects on standing biomass and functional diversity of various intensities and patterns of fragmentation within a forest landscape. The simulator was calibrated for an old-growth wet forest in French Guiana, South America. We found that the standing biomass of forest remnants was reduced significantly compared to a similar area of nonfragmented forest. When fewer but larger remnants were created rather than many small ones, the total loss in biomass and the increase in the abundance of early-successional species were significantly reduced, confirming that edge effects dominate the functioning of forest remnants. We also performed simulations of secondary succession after the landscape had been abandoned. The simulated recovery time in those secondary forests depends on both the size of cleared area and the spatial pattern of the remnant forests. Received 30 January 2002; accepted 16 July 2002.     

Demographic Drivers of Aboveground Biomass Dynamics During Secondary Succession in Neotropical Dry and Wet Forests

The magnitude of the carbon sink in second-growth forests is expected to vary with successional biomass dynamics resulting from tree growth, recruitment, and mortality, and with the effects of climate on these dynamics. We compare aboveground biomass dynamics of dry and wet Neotropical forests, based on monitoring data gathered over 3–16 years in forests covering the first 25 years of succession. We estimated standing biomass, annual biomass change, and contributions of tree growth, recruitment, and mortality. We also evaluated tree species’ contributions to biomass dynamics. Absolute rates of biomass change were lower in dry forests, 2.3 and 1.9 Mg ha^?1 y^?1, after 5–15 and 15–25 years after abandonment, respectively, than in wet forests, with 4.7 and 6.1 Mg ha^?1 y^?1, in the same age classes. Biomass change was largely driven by tree growth, accounting for at least 48% of biomass change across forest types and age classes. Mortality also contributed strongly to biomass change in wet forests of 5–15 years, whereas its contribution became important later in succession in dry forests. Biomass dynamics tended to be dominated by fewer species in early-successional dry than wet forests, but dominance was strong in both forest types. Overall, our results indicate that biomass dynamics during succession are faster in Neotropical wet than dry forests, with high tree mortality earlier in succession in the wet forests. Long-term monitoring of second-growth tropical forest plots is crucial for improving estimates of annual biomass change, and for enhancing understanding of the underlying mechanisms and demographic drivers.     

海南岛热带低地雨林刀耕火种弃耕地自然恢复过程中的群落构建

研究群落构建机制是群落生态学的一个重要目标, 群落动态过程中的构建规律对于了解群落演替机理有重要的作用。该文以海南岛刀耕火种干扰后自然恢复的10 hm ²热带低地雨林为研究对象, 通过比较不同恢复阶段的次生林(15年、30年和60年)和老龄林在幼苗、幼树和成年树群落的物种组成, 揭示次生演替过程中的群落构建规律。研究结果表明, 老龄林中不同径级群落的物种多样性及不同径级间的物种相似度显著高于各恢复阶段的次生林, 但优势种在群落中的比例低于各恢复阶段的次生林。随着自然恢复过程的进行, 次生林群落物种组成与老龄林的相似性也逐渐增大, 支持演替平衡理论。所有恢复阶段样地中幼苗的个体、物种丰富度和基于多度涵盖估计量(ACE)都低于幼树和成年树群落, 幼苗层物种组成与幼树、成年树也有较大差异, 说明新增到幼苗群落可能是一个难于预测的过程。研究结果说明了确定过程和随机过程共同决定了次生演替的群落构建。     

庐山森林景观格局变化的长期动态模拟

在以植被格局为基础的森林景观动态分析中,可通过森林演替推断景观格局的动态变化以及相应的景观生态过程。运用空间直观景观模型LANDIS,以庐山风景区为案例地,模拟森林植被在未来300 a的自然演替动态,在此基础上选取斑块面积比、聚集度、分维数、多样性指数和均匀度指数等景观格局指数,分析森林景观格局随森林演替的动态变化。结果表明：（1）阔叶林树种的绝对优势地位保证其斑块面积比呈现持续增长的稳定趋势,森林植被将朝着地带性常绿阔叶林方向演替;（2）景观聚集度特征方面,阔叶林树种在前150 a缓慢增长,而后150 a保持相对稳定,杉木林一直保持平稳,毛竹林在整个模拟阶段一直在不断下降直至演替结束;（3）各优势树种植被斑块的分维数都保持在1-1.1之间,说明各景观斑块的边缘相对较规则且变化较小;（4）景观多样性指数呈现出先上升后缓慢下降的趋势,而均匀度指数则呈现出先下降后上升再缓慢下降的变化态势。景观格局指数的变化特征与植被向顶极群落演替的趋势相吻合,该模拟结果可运用到庐山森林景观的管理实践中。从长远来看,应该继续实行严格的封山育林政策。     

Determinants of nocturnal Lepidopteran diversity and community structure in a conifer-dominated forest

Prediction of community response of fauna to anthropogenic or environmental disturbance requires knowledge of faunal distribution and abundance as well as an understanding of the mechanisms underlying community organization. We investigated linkages between Lepidoptera, one of the most influential insect taxa in forested ecosystems, and vegetation in riparian areas of conifer-dominated forests of western Oregon, USA. Using model selection techniques, we found that canopy cover explained variability in patterns of moth species dominance and diversity better than all other factors investigated, whereas elevation best explained patterns of moth species richness. Using canonical correspondence analysis, we determined that gradients in shrub species richness, elevation, and understory species richness accounted for the most variation in regional moth community structure. Results suggest that reductions of riparian forest canopy are likely to have the greatest impact on the variety and relative abundance of moths. Due to the relative rarity and patchy distribution of the majority of individual species, we predict that lower intensity vegetation manipulations distributed across larger spatial scales are likely to be less damaging to regional moth biodiversity than more intensive alterations at smaller scales. Finally, if global warming trends continue, upward elevational shifts in moth species distributions will make forested regions at higher elevation worthy of concerted protection.     

A shade tolerance index for common understory species of northeastern North America

Since Baker’s [Baker, F.S., 1949. A revised tolerance table. J. For. 47, 179–181] classic contribution, shade tolerance indices have not been much modified for North American plant species. While many common tree and shrub species are included in the shade tolerance index, much less is known about this characteristic for the abundant and rich understory vascular and nonvascular plant layers. The classification of the shade tolerance is widely used to compare relative growth and survival among plant species under closed canopies and is also fundamental to an understanding of stand development following small and large scale disturbances. Although qualitative, it is frequently used both in research and management implications. Here we provide a significant revision to Baker’s shade tolerance table to include the most common forest understory plant species found in northeastern North America forests. Our index is based on: (1) the compilation of the opinions of five experts, (2) a comparison with Ellenberg’s index from Europe as well as, (3) information from current literature. For most of the 347 plant species investigated, a consistent and robust shade tolerance index, ranging between 1 (very tolerant) and 9 (very intolerant), was found. Here we present revised shade indices for 71 tree and shrub species, 185 herbaceous species, and 91 bryophyte and lichen species.     

The limited role of canopy gaps in the successional dynamics of a mature mixed Quercus forest remnant

Question: Are canopy gap dynamics responsible for driving the structural and compositional changes that have occurred over a 26‐year period in a mature Quercus forest remnant? Location: Dobbs Natural Area, an unlogged 3.6‐ha forest preserve in west‐central Indiana, USA. Methods: We analyzed mapped permanent plot data for a site that illustrates a trend common in Quercus‐dominated forests in eastern North America, where recruitment of new stems is dominated by mesophytic, shade‐tolerant species such as Acer saccharum, rather than Quercus. We developed a GIS database from stand census measurements taken in 1974 and 2000, employing it to conduct tree‐by‐tree comparisons that allow direct determination of ingrowth, mortality and survivorship, and to relate the spatial patterns of subcanopy dynamics to canopy gap occurrence. Results: The re‐census shows modest changes in canopy composition, but much greater turnover in the subcanopy. Nearly half of all individuals originally present died; much of this mortality resulted from a major decline in subcanopy Ulmus americana. While overall density remained fairly constant, the subcanopy experienced substantial ingrowth of shade‐tolerant Acer saccharum, Fagus grandifolia, and Tilia americana. Canopy gaps, although forming at rates in the upper range of regional averages, did not significantly benefit subcanopy populations of Quercus spp. or most other taxa with limited shade tolerance. Conclusions: Canopy gaps play a minor role in driving the recent demographic trends of this stand. The spatial and temporal scales of light availability in gaps do not support regeneration of most shade‐intolerant species. Compositional change parallels a historical shift in light regimes.