An Appraisal of the Classic Forest Succession Paradigm with the Shade Tolerance Index

In this paper we revisit the classic theory of forest succession that relates shade tolerance and species replacement and assess its validity to understand patch-mosaic patterns of forested ecosystems of the USA. We introduce a macroscopic parameter called the “shade tolerance index” and compare it to the classic continuum index in southern Wisconsin forests. We exemplify shade tolerance driven succession in White Pine-Eastern Hemlock forests using computer simulations and analyzing approximated chronosequence data from the USDA FIA forest inventory. We describe this parameter across the last 50 years in the ecoregions of mainland USA, and demonstrate that it does not correlate with the usual macroscopic characteristics of stand age, biomass, basal area, and biodiversity measures. We characterize the dynamics of shade tolerance index using transition matrices and delimit geographical areas based on the relevance of shade tolerance to explain forest succession. We conclude that shade tolerance driven succession is linked to climatic variables and can be considered as a primary driving factor of forest dynamics mostly in central-north and northeastern areas in the USA. Overall, the shade tolerance index constitutes a new quantitative approach that can be used to understand and predict succession of forested ecosystems and biogeographic patterns.     

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.     

Herb layer changes (1954-2000) related to the conversion of coppice-with-standards forest and soil acidification

Question: Did the composition of the herb layer of a deciduous forest on loamy soils sensitive to soil acidification change between 1954 and 2000? How are these change related to the abandonment of traditional coppice-with-standards forest management and increased soil acidification? Location: Central Belgium (Europe). Methods: Twenty semi-permanent phytosociological quadrats from an ancient deciduous forest (Meerdaal forest) were carefully selected out of a total of 70 plots dating from 1954 and were revisited in 2000. Species composition and soil pH H₂O were recorded using an analogous methodology. The studied period coincides with a period of forest conversion from coppice-with-standards towards a high forest structure and with an increase in acidifying and eutrophying deposition. Results: Between 1954 and 2000, species composition of the herb layer changed significantly. Redundancy analysis pointed to increased shade resulting from shifts in cover and species composition of the shrub and tree layer as the main driving force. Soil acidity increased and the majority of plots entered the aluminium buffer range, which potentially affected herb layer composition. Observations at the species level, especially a strong decrease in cover of the vernal species Anemone nemorosa supported this hypothesis. Conclusions: Our results show significant shifts in the forest herb layer in less than five decades. These shifts were related to an alteration in the traditional forest management regime and increased soil acidity. Whereas the effect of a changed management regime can be mitigated, soil acidification is less reversible. Testing the generality of these patterns on more extensive data sets is certainly needed.     

Amazon drought and forest response: Largely reduced forest photosynthesis but slightly increased canopy greenness during the extreme drought of 2015/2016

Amazon droughts have impacted regional ecosystem functioning as well as global carbon cycling. The severe dry‐season droughts in 2005 and 2010, driven by Atlantic sea surface temperature (SST) anomaly, have been widely investigated in terms of drought severity and impacts on ecosystems. Although the influence of Pacific SST anomaly on wet‐season precipitation has been well recognized, it remains uncertain to what extent the droughts driven by Pacific SST anomaly could affect forest greenness and photosynthesis in the Amazon. Here, we examined the monthly and annual dynamics of forest greenness and photosynthetic capacity when Amazon ecosystems experienced an extreme drought in 2015/2016 driven by a strong El Niño event. We found that the drought during August 2015–July 2016 was one of the two most severe meteorological droughts since 1901. Due to the enhanced solar radiation during this drought, overall forest greenness showed a small increase, and 21.6% of forests even greened up (greenness index anomaly ≥1 standard deviation). In contrast, solar‐induced chlorophyll fluorescence (SIF), an indicator of vegetation photosynthetic capacity, showed a significant decrease. Responses of forest greenness and photosynthesis decoupled during this drought, indicating that forest photosynthesis could still be suppressed regardless of the variation in canopy greenness. If future El Niño frequency increases as projected by earth system models, droughts would result in persistent reduction in Amazon forest productivity, substantial changes in tree composition, and considerable carbon emissions from Amazon.     

Planting position and shade enhance native seedling performance in forest restoration for an endangered malagasy plant

The critically endangered tree Schizolaena tampoketsana is confined to a few diminished and degraded forest fragments on the Malagasy highlands. This habitat is vulnerable to loss due to frequent fires in the surrounding grassland that threaten to spread into the forest. One of these fragments is the focus a conservation project and here the managers aim to conserve S. tampoketsana by restoring its forest habitat to its former extent as evidenced by remnant woody plants. To inform this activity the survival and early-stage growth of seedlings of four locally native tree species were compared under contrasting conditions of proximity to the remaining forest and shade. After 12 months, seedlings of three species(Baronia taratana, Eugenia pluricymosa, Uapaca densifolia) survived better and experienced improved growth in height in grassland close to the existing forest rather than distant from it, and two survived better with shade rather than unshaded. A number of mechanisms could explain these results including reduced exposure to desiccating sunlight and winds and better soil and greater water availability close to the forest. The seedlings of one species(Nuxia capitata) survived well under all conditions. This study suggests that reforestation in these dry highlands is most feasible adjacent to remnant forest fragments and in microhabitats that minimize water loss, though young plants of some tree species may be capable of surviving in harsher conditions.     

Leaf structure of Syzygium spp. (Myrtaceae) in relation to site affinity within a tropical rain forest

This study examined four species of Syzygium (S. firmum, S. makul, S. operculatum, S. rubicundum) Myrtaceae, a tree genus that dominates the canopy of rain forests of south‐west Sri Lanka. Syzygium spp. occupy differing habitats with relation to succession and forest topography. We examined differences in leaf morphology and physiology in response to amount of shade, an important environmental variable affecting Syzygium distribution within the forest. To study change in leaf structure and physiology, environmental shelters were constructed simulating forest shade that differed in quality, quantity and duration. Seedlings were exposed to: (i) 0% shade (full sun, FS), red : far red (R : FR) ratio 1.27; (ii) 65% shade (large opening, LO) with direct sunlight similar to the centre of a large canopy opening, R : FR ratio 1.27; (iii) 82% shade (small opening, SO) with direct sunlight similar to the centre of a small canopy opening, R : FR ratio 1.27; (iv) 58% uniform light shade (LS) with a quality similar to the outside edge of a large canopy opening, R : FR ratio 1.05; (v) 85% uniform medium shade (MS) with a quality similar to the inside forest edge of a large canopy opening, R : FR ratio 0.97; (vi) 99% uniform deep shade (DS) similar to that of the forest understorey, R : FR ratio 0.23. The shelters were constructed in a large open area at the field station of the Sinharaja World Heritage site, Sri Lanka. Seedlings of each species were grown for two years in their respective shade treatments before physiological, morphological and anatomical measurements were made on leaves. Variation in leaf structure and physiology between the species was associated with differences in shade‐tolerance and water‐use. All species increased in photosynthesis rates and dimensions in leaf structure (leaf blade and cuticle thickness, stomatal density, thickness of upper and lower epidermis, and thickness of palisade mesophyll) with decrease in shade. In contrast, stomatal conductivity was highest in the DS (99% shade) treatment. Leaves of Syzygium firmum were thickest and largest in area. S. firmum also had highest photosynthesis in the SO (82% shade) treatment. S. firmum was the most shade‐tolerant of all species: it grows well in low shade and its leaf structure suggests it to be the most conservative in water‐use of the Syzygium spp. In the forest S. firmum can persist in the forest shade as established seedlings, but grows best within canopy openings of late‐seral rain forest. Leaves of S. operculatum were thinnest but had highest stomatal densities of the four species. S. operculatum is considered shade‐intolerant, with a leaf structure suggesting it to be prone to desiccation, and by implication susceptible to drought. S. operculatum is found along streams within early seral rain forest habitat, often originating on stream banks after land clearance for cultivation. In the FS (0% shade) treatment, S. rubicundun had highest photosynthesis rates and greatest number of leaves but smallest leaf area of the Syzygium species. S. rubicundum is more shade‐intolerant but more efficient in water‐use than S. operculatum. S. rubicundum is a mid‐seral canopy tree of the midslope stands that are thought to have originated after catastrophic windthrows or swidden cultivation. The leaf physiology and structure of S. makul suggests it to be both moderately shade‐tolerant and conservative in water‐use. It is the most widely distributed Syzygium species across the topography of late‐seral rain forest. We suggest forest disturbance and hydrology are important environmental factors that influence distribution of Syzygium species across the topography. Results from this study contribute to a body of knowledge suggesting that canopy tree species of rain forests in south‐west Sri Lanka have discrete affinities to topography and differences in successional status, and that adaptations in leaf structure and physiology are indicative of such phenomena. © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society, 2003, 141 , 365–377.     

大兴安岭森林火烧后不同演替阶段土壤细菌多样性动态

【目的】研究我国最大的林区之一——大兴安岭森林火烧后不同演替阶段土壤细菌多样性动态,为天然林保护工程对于生物多样性的影响增添新的认识。【方法】以空间替代时间的方法分析大兴安岭森林演替对于土壤细菌多样性动态的影响。大兴安岭森林火烧后典型的自然演替序列为火烧迹地(LG-BA)、灌丛(SHR)、白桦林(BP)、白桦落叶松混交林(BP-LG)、落叶松林(LG-OM)。在演替序列的每个典型样地上采集0-10 cm土样,采用Illumina Mi Seq高通量测序技术测定土壤细菌群落组成及其多样性。【结果】细菌操作分类单位(OTU)数量从少到多的顺序为火烧迹地落叶松白桦混交林灌丛落叶松林白桦林。随着森林演替,多样性指数Simpson先增高后降低;Shannon指数先增加后减少再增加;OTU的丰度变化比较平缓,表明物种变化较小。在各演替阶段中,土壤细菌种类主要有变形菌门(Proteobacteria)、酸杆菌门(Acidobacteria)、放线菌门(Actinobacteria)和浮霉菌门(Planctomycetes),4个门的种类含量随演替顺序都呈现先增加后减少的趋势。主成分分析表明不同演替阶段土壤细菌群落存在一定的差异性。冗余分析表明有机质(SOM)、全氮(TN)、全磷(TP)和p H对于土壤细菌群落变化有影响。【结论】随着森林演替,大兴安岭地区土壤细菌种类和生物多样性会发生变化,其变化与土壤化学成分和p H有关。     

Losses of seeds of temperate trees to soil fungi: effects of habitat and host ecology

Evidence suggests that impacts of fungal pathogens on tree recruitment tend to be greater in the forest understory than in openings, and that shade-tolerant trees are less vulnerable than shade-intolerant species. To investigate the role that harmful soil fungi may have in reducing regeneration of temperate trees, we applied fungicide to buried seeds of matched pairs of species differing in their relative shade tolerance and/or successional status (Acer negundo versus Acer saccharum, Prunus virginiana versus Prunus serotina, and Pinus strobus versus Tsuga canadensis), in three habitats that differed in their degree of openness (old field, forest gap, and forest understory). Our results indicated that soil fungi reduced germination of A. negundo, P. virginiana, P. serotina. and T. canadensis, and reduced viability of ungerminated seeds of P. strobus; no significant effects of fungi on seeds of A. saccharum were detected. However, we found seeds were not less likely to survive following burial in forest understory than in gaps. As well, results for only one species pair (A. negundo versus A. saccharum) were consistent with the prediction that shade-intolerant or successional species should be more susceptible to fungal attack than mature forest species. These results contrast with other studies of temperate and especially tropical forest trees.     

Light environment,sapling architecture,and leaf display in six rain forest tree species

Architecture and leaf display were compared in saplings of six rain forest tree species differing in shade tolerance. Saplings were selected along the whole light range encountered in a forest environment. Species differed largely in realized height and crown expansion per unit support biomass, but this could not be related to differences in shade tolerance. The results demonstrate that there exist various solutions to an effective expansion of plant height and crown area. It is argued that choice of the study species and the ontogenetic trajectory regarded determine to a large extent the outcome of interspecific comparisons. No evidence was found that pioneers were characterized by a multilayered and shade tolerants by a monolayered leaf distribution. Yet, sun plants had a similar crown area, a deeper crown, and a higher leaf area index compared to shade plants and their leaves were more evenly distributed along the stem. This suggests that differences in leaf layering are found between plants growing in different light environments, rather than between species differing in shade tolerance.     

Responses of temperate woody seedlings to shade and drought: do trade-offs limit potential niche differentiation?

Diversity in seedling responses to combined shade and drought can drive species niche differences, and thus natural forest and scrub establishment dynamics and diversity. However, inherent trade-offs between shade tolerance and drought tolerance, as hypothesized in the literature, would constrain potential niche differentiation. For thirteen species of European trees and shrubs the impacts were determined of moderate and extreme drought on the growth and survival of first-year seedlings in a given soil volume, in irradiances in the range typical for temperate forest and scrub understories (3% daylight) and large gaps (30% daylight). Pots were watered differentially each second day to equalize soil water content across species and irradiances. Comparisons within and across species supported independent tolerances of shade and drought rather than trade-offs. For all species, drought reduced relative growth rate in dry mass by the same proportion in 3% daylight and in 30% daylight. Consequently, drought generally reduced final dry mass significantly more strongly in 30% daylight than in 3% daylight. Extreme drought led to earlier mortality in 30% daylight than in 3% daylight for nine of the eleven species tested, to earlier mortality in 3% daylight for one species, and to equal mortality rate in both irradiances for one species. For each species, growth-based shade tolerance was quantified as the ratio of absolute growth in low irradiance to that in high irradiance, and growth-based drought tolerance as the ratio of absolute growth in low water supply to that in high water supply. Across species, growth-based shade tolerance varied ≈9-fold, and growth-based drought tolerance ≈2-fold; species’ tolerances correlated respectively with indices of field establishment in shade and drought. Growth-based drought tolerance correlated with survival time in extreme drought. Experimentally quantified shade and drought tolerances varied independently for the 13 species tested, indicating the potential for extensive species niche differentiation in combinations of irradiance and water supply.     

Microclimatic conditions determined by stem density influence leaf anatomy and leaf physiology of beech (Fagus sylvatica L.) growing within stands that naturally regenerate from clear-cutting

Beech forests naturally regenerating from clear-cutting can exhibit different microclimates depending on size of saplings and stem density. When beech trees are young and stem density is low, the level of radiation inside the ecosystem reaching the soil surface is high; consequently, air and soil temperatures rise and the soil water content may decrease. These microclimatic parameters presumably will affect the anatomy, photosynthesis, and carbon metabolism of beech leaves. We studied the morphology and physiology of sun and shade leaves of beech trees differing in age and growing within clear-cut areas with distinct microclimate. Results were compared with those of adult trees in an unmanaged forest. We selected a stand clear-cut in 2001 (14,000 trees ha⁻¹), another clear-cut in 1996 (44,000 trees ha⁻¹) and an unmanaged forest (1,000 trees ha⁻¹). Photosynthetic photon flux density (PPFD) incident on sun leaves, air temperature, soil moisture, and soil temperature within the forests affected water status and carbohydrate storage in all trees. As trees became older, PPFD also influenced pigment composition and Rubisco activity in sun leaves. On the other hand, shade leaves from the oldest trees were the most sensitive to PPFD, air temperature, and soil moisture and temperature inside the forest. Contrariwise, microclimatic parameters slightly affected the physiology of shade leaves of the beech in the stand with the highest light attenuation. Air and soil temperatures were the parameters that most affected the photosynthetic pigments and carbohydrate storage in shade leaves of the youngest trees.     

Tree dynamics and co‐existence in the montane–sub‐alpine ecotone: the role of different light‐induced strategies

Questions: Is light availability the main factor driving forest dynamics in Pyrenean sub‐alpine forests? Do pines and firs differ in growth, mortality and morphological response to low light availability? Can differences in shade tolerance affect predictions of future biome changes in Pyrenean sub‐alpine forests in the absence of thermal limitation? Location: Montane–sub‐alpine ecotones of the Eastern Pyrenees (NE Spain). Methods: We evaluated morphological plasticity, survival and growth response of saplings of Scots pine, mountain pine and silver fir to light availability in a mixed forest ecotone. For each species, we selected 100 living and 50 dead saplings and measured size, crown morphology and light availability. A wood disk at root collar was then removed for every sapling, and models relating growth and mortality to light were obtained. Results: Fir had the lowest mortality rate (<0.1) for any given light condition. Pines had comparable responses to light availability, although in deep shade Scots pine risked higher mortality (0.35) than mountain pine (0.19). Pines and fir developed opposing strategies to light deprivation: fir employed a conservative strategy based on sacrificing height growth, whereas pines enhanced height growth to escape from shade, but at the expense of higher mortality risk. Scots pine showed higher plasticity than mountain pine for all architectural and morphological traits analysed, having higher adaptive capacity to a changing environment. Conclusions: Our results support the prediction of future biome changes in Pyrenean sub‐alpine forests as silver fir and Scots pine may find appropriate conditions for colonizing mountain pine‐dominated stands due to land‐use change‐related forest densification and climate warming‐related temperature increases, respectively.     

Differential effect of shade,water and soil type on emergence and early survival of three dominant species of the Atacama Desert

Understanding the regeneration niche of species may allow us to gain insight into how communities are structured. In deserts, the regeneration niche is usually related to spaces beneath shrubs where shade cast by shrubs creates microenvironments that benefit seedlings and where even small amounts of rain may favour germination and establishment. Shade and water may also interact with different types of soils. However, species may have different requirements for germination and seedling survival. We could expect that shrub species with different drought tolerances exhibit different responses to the combination of these factors. We ask if responses of dominant species of the Atacama Desert to abiotic factors (shade, water and soil type) are related to their drought tolerance, a topic not exhaustively explored in shrubs growing in true deserts. We conducted two factorial experiments. The first one was designed to evaluate how shade (microhabitat) in combination with water may affect germination (emergence) and early survival. In the second experiment, we assessed the influence of shade in relation to soil type. Each species responded distinctively to the three variables under study, but in general, their emergence responses were more influenced by water (more water, greater emergence) than by microhabitat or soil type. Survival was influenced both by microhabitat and by water and was higher under shade and abundant water. Soil type affected only one of our species in terms of emergence. Species responses in general depended on their tolerance to stress. In one species, there was indication of a seed–seedling conflict. Our results show similar species responses to environmental constraints but also more or less unique responses that are related to their tolerance to drought and which may ultimately permit species coexistence. We found that shade may not be important for germination but may be crucial for survival in dry years.     

Hydraulics and life history of tropical dry forest tree species: coordination of species' drought and shade tolerance

Plant hydraulic architecture has been studied extensively, yet we know little about how hydraulic properties relate to species' life history strategies, such as drought and shade tolerance. The prevailing theories seem contradictory. We measured the sapwood (K(s) ) and leaf (K(l) ) hydraulic conductivities of 40 coexisting tree species in a Bolivian dry forest, and examined associations with functional stem and leaf traits and indices of species' drought (dry-season leaf water potential) and shade (juvenile crown exposure) tolerance. Hydraulic properties varied across species and between life-history groups (pioneers vs shade-tolerant, and deciduous vs evergreen species). In addition to the expected negative correlation of K(l) with drought tolerance, we found a strong, negative correlation between K(l) and species' shade tolerance. Across species, K(s) and K(l) were negatively correlated with wood density and positively with maximum vessel length. Consequently, drought and shade tolerance scaled similarly with hydraulic properties, wood density and leaf dry matter content. We found that deciduous species also had traits conferring efficient water transport relative to evergreen species. Hydraulic properties varied across species, corresponding to the classical trade-off between hydraulic efficiency and safety, which for these dry forest trees resulted in coordinated drought and shade tolerance across species rather than the frequently hypothesized trade-off.     

Effects of overstory and understory vegetation on the understory light environment in mixed boreal forests

Abstract. The percentage of above-canopy Photosynthetic Photon Flux Density (%PPFD) was measured at 0, 50 and 100 cm above the forest floor and above the main understory vegetation in stands of (1) pure Betula papyrifera (White birch), (2) pure Populus tremuloides (Trembling aspen), (3) mixed broad-leaf-conifer, (4) shade-tolerant conifer and (5) pure Pinus banksiana (Jack pine) occurring on both clay and till soil types. %PPFD was measured instantaneously under overcast sky conditions (nine locations within each of 29 stands) and continuously for a full day under clear sky conditions (five locations within each of eight stands). The percentage cover of the understory layer was estimated at the same locations as light measurements. Mean %PPFD varied from 2% at the forest floor under Populus forests to 15% above the understory vegetation cover under Betula forests. Percent PPFD above the understory vegetation cover was significantly higher under shade intolerant tree species such as Populus, Betula and Pinus than under shade tolerant conifers. No significant differences were found in %PPFD above the understory vegetation cover under similar tree species between clay and till soil types. The coefficient of variation in %PPFD measured in the nine locations within each stand was significantly lower under deciduous dominated forests (mean of 19%) than under coniferous dominated forests (mean of 40%). %PPFD measured at the forest floor was positively correlated with %PPFD measured above the understory vegetation and negatively correlated with cumulative total percent cover of the understory vegetation (R² = 0.852). The proportion of sunflecks above 250 and 500 mmol m^–2 s^–1 was much lower and %PPFD in shade much higher under Populus and Betula forests than under the other forests. Differences in the mean, variability and nature of the light environment found among forest and soil types are discussed in relation to their possible influences on tree succession.     

Seasonal and long-term soil water regime in West African tropical forest

Abstract. Variation of soil matric potential of a Wet Evergreen and a Moist Semi-deciduous West African forest were compared. The two forest types differed strongly in their soil water regime. Wet Evergreen forest experienced matric potentials below ?100 kPa only occasionally, while in Moist Semi-deciduous forest matric potentials were less than ?2.5 MPa for periods of several weeks or more each season. A water balance equation was used to simulate the soil water regime at both sites and predict severity and length of the dry period. The predictions showed good agreement with the field measurements of soil water potential over a 2-yr period. The methodology was used to estimate the occurrence and severity of droughts over longer periods at the two sites. The balance calculations suggest that droughts occur occasionally in the Wet Evergreen forest under study. The potential impact of droughts on species distribution and vegetation disturbance in tropical forests is discussed.     

The shrub Rhodomyrtus tomentosa acts as a nurse plant for seedlings differing in shade tolerance in degraded land of South China

Questions: What are the nurse effects of Rhodomyrtus tomentosa in degraded land of South China? Are canopy or soil factors responsible for the main nurse effect? Do facilitative effects increase with the shade tolerance of the target species? Location: Degraded shrubland in South China. Methods: Seedlings of three native climax woody species (Schima superba, Michelia macclurei, Castanopsis fissa) that differ in shade tolerance were subjected to four treatments by transplantation: (1) under the canopy of R. tomentosa shrubs; (2) in open interspaces without vegetation cover (control); (3) under the canopy of R. tomentosa from which canopies had been removed; and (4) in open interspaces without vegetation but covered by branches and leaves of R. tomentosa. Results: At low soil nutrient levels, increased canopy shade, soil porosity and soil moisture under the canopy of R. tomentosa increased seedling survival of the climax tree species S. superba, C. fissa and M. macclurei, and shoot height of S. superba. The nurse effect (a form of facilitation) of R. tomentosa depended more on canopy shade than on soil amelioration. The magnitude of facilitation or nurse effect was positively correlated with shade tolerance of the target species. Conclusions: Use of nurse plants in restoration differs from traditional reforestation (clearing and/or burning to reduce interspecific competition between target tree species and non‐target species) because it focuses on positive interactions between nurse plants and target plants that increase establishment of target species and reduce time required for restoration. Because nurse effects of R. tomentosa shrubs tended to be larger on target species with greater shade tolerance, shade‐tolerant plants are suggested as target species to accelerate restoration.     

Drought legacies are dependent on water table depth,wood anatomy and drought timing across the eastern US

Severe droughts can impart long‐lasting legacies on forest ecosystems through lagged effects that hinder tree recovery and suppress whole‐forest carbon uptake. However, the local climatic and edaphic factors that interact to affect drought legacies in temperate forests remain unknown. Here, we pair a dataset of 143 tree ring chronologies across the mesic forests of the eastern US with historical climate and local soil properties. We found legacy effects to be widespread, the magnitude of which increased markedly in diffuse porous species, sites with deep water tables, and in response to late‐season droughts (August–September). Using an ensemble of downscaled climate projections, we additionally show that our sites are projected to drastically increase in water deficit and drought frequency by the end of the century, potentially increasing the size of legacy effects by up to 65% and acting as a significant process shaping forest composition, carbon uptake and mortality.     

Recovery of soil productivity with forest succession on abandoned agricultural land

Much of the primary forest in the eastern United States that was converted to farmland between 1600 and 1900 has reverted back to second growth forest as a result of agriculture abandonment. This reversion back to forest gives soil productivity a chance to recover, though the rates of recovery are not well understood. Understanding the legacy effects of past disturbances like agriculture can provide important insights to support ecological restoration efforts on disturbed soils. Our goal with this study was to further understand the effects of forest development on soil productivity after agriculture abandonment. We used a chronosequence approach to examine soil properties over a 60‐year temporal scale of forest development on abandoned agricultural lands in Saratoga and Rensselaer Counties in New York, U.S.A. We measured soil properties within this chronosequence to test the hypothesis that there would be measurable recoveries of soil physical properties and fertility over time. We observed rapid recovery of physical properties (lower bulk density and higher macroporosity) of surface soils within 5–10 years after agricultural abandonment. However, we found a legacy effect of agricultural compaction still evident in subsoils, with soil strength measurements indicating that past agricultural practices still limited root growth 55–60 years after abandonment. Soil percent organic matter and mineralizable nitrogen (N) both increased with forest development, but biomass accumulation may be slowed by limited root growth in the subsoil due to high strength. We recommend assessing subsoil physical properties when developing ecological restoration plans for agricultural lands.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.