Impact of the invasive shrub <Emphasis Type="Italic">Lonicera maackii</Emphasis> on stand transpiration and ecosystem hydrology in a wetland forest

Invasive shrubs can increase ecosystem transpiration and potentially affect hydrology in forested ecosystems. We examined two adjacent sites in a wetland forest in northern Kentucky, USA. One site contained little Lonicera maackii (Amur honeysuckle), while the other contained considerably more. Using Granier (thermal dissipation) and heat balance probes, transpiration was determined for trees, vines and shrubs at the two sites. Tree and vine transpiration in 2009 was usually 1–2 mm day⁻¹, typical of that seen in humid temperate forests. Additional transpiration from L. maackii was roughly proportional to its basal area, and it totaled 1.0% of tree and vine transpiration from the site with less L. maackii cover and 6.0% from the site with considerable cover. This additional transpiration amounts to roughly 10% of stream flow draining the study area. As L. maackii basal areas at these sites are at the lower end of that seen in other invaded forests in the region, regional impacts on transpiration and hydrology may be larger than those reported here. We expect L. maackii to shorten the lives of ephemeral ponds and streams in wetlands and cause adverse impacts on the organisms, such as amphibians, that require these aquatic environments to complete their life cycle.     

Rainforest expansion reduces understorey plant diversity and density in open forest of eastern Australia

The expansion of rainforest pioneer trees into long‐unburnt open forests has become increasingly widespread across high rainfall regions of Australia. Increasing tree cover can limit resource availability for understorey plant communities and reduce understorey diversity. However, it remains unclear if sclerophyll and rainforest trees differ in their competitive exclusion of understory plant communities, which contain most of the floristic diversity of open forests. Here, we examine dry open forest across contrasting fire histories (burnt and unburnt) and levels of rainforest invasion (sclerophyll or rainforest midstorey) to hindcast changes in understorey plant density, richness and composition. The influence of these treatments and other site variables (midstorey structure, midstorey composition and soil parameters) on understorey plant communities were all examined. This study is the first to demonstrate significantly greater losses of understorey species richness, particularly of dry open‐forest specialists, under an invading rainforest midstorey compared to a typical sclerophyll midstorey. Rainforest pioneers displaced over half of the understorey plant species, and reduced ground cover and density of dry forest specialists by ~90%. Significant understorey declines also occurred with increased sclerophyll midstorey cover following fire exclusion, although losses were typically less than half that of rainforest‐invaded sites over the same period. Understorey declines were closely related to leaf area index and basal area of rainforest and wattle trees, suggesting competitive exclusion through shading and potentially belowground competition for water. Around 20% of displaced species lacked any capacity for population recovery, while transient seed banks or distance‐limited dispersal may hinder recovery for a further 68%. We conclude that rainforest invasion leads to significant declines in understorey plant diversity and cover in open forests. To avoid elimination of local native plant populations in open forests, fires should occur with sufficient frequency to prevent overstorey cover from reaching a level where shade‐intolerant species fail to thrive.     

Present and past old‐growth forests of the Lake Tahoe Basin,Sierra Nevada,US

Abstract. We described 38 relictual old‐growth stands – with data on the mortality, regeneration, floristic richness, fuel load and disease incidence in our study area in the Tahoe Basin of California and Nevada. The stands are within the lower and upper montane zones (1900–2400 m a.s.l.) and they are rare, occupying < 2% of the land in the Basin's watershed. Correlation matrices and ANOVAs of forest types and conifer species with environmental gradients revealed significant relationships with elevation, distance east of the Sierran crest, slope aspect, annual precipitation, date of complete snow melt, litter depth and degree of soil profile development. Pathogens, parasites and wood‐boring insects were present on 23% of living trees; 16% of all trees were dead. We compared these stands to a reconstruction of pre‐contact Basin forests and to ecologically analogous old‐growth forests of Baja California that have never experienced fire suppression management. Currently, overstorey trees (> 180 yr old) in the Basin stands have ca. 33% cover, 54 m².ha^‐1 basal area and 107 individuals.ha^‐1, values very similar to reconstructions of pre‐contact Basin forests and to modern Baja California forests. Understorey trees (60–180 yr old), however, are several times more dense than historic levels and species composition is strongly dominated by A. concolor, regardless of the overstorey composition. The ratio of Pinus: Abies has increased – and the age structure of extant stands predicts that it will continue to increase – from approximately 1:1 in pre‐contact time to 1:7 within the next century. Disease incidence and mortality in Baja forests were lower. Although we quantitatively defined current Basin old‐growth forests – in terms of stand structure – we realize that our definition will differ from that of both past and future old‐growth forests unless management protocols are changed.     

Converging forest community composition along an edaphic gradient threatens landscape‐level diversity

Aim Plant communities across the temperate zone are changing in response to successional processes and human‐induced disturbances. Here, we assess how upland forest under‐ and overstorey community composition has changed along an edaphic gradient. Location Northern Wisconsin, USA. Methods Forest sites initially sampled in the 1950s were resampled for overstorey composition and diversity, basal area, and understorey composition and diversity. We used clustering methods to identify groups of stands based on overstorey composition, and we used similarity indices, ordination and diversity indices to evaluate changes in species abundance and overall community structure. Results Sites clustered into four overstorey groups along the edaphic gradient: ‘hemlock’ sites dominated by hemlock in 1950, ‘mesic’ sites dominated by northern hardwoods, ‘dry’ sites with a significant pine inclusion in the canopy and diverse ‘dry‐mesic’ sites in the middle. Collectively, forests gained maple, ash and cherry while losing pines, birches and red oaks. The hemlock forest sites gained hardwoods, while the dry‐mesic sites shifted towards a more mesic hardwood composition. Only the driest sites have remained relatively stable in species composition. Main conclusions These trends reflect both ‘mesification’ and homogenization among northern forests. Highly diverse mid‐gradient and mesic hemlock‐dominated stands are transitioning to maple dominance. Fire suppression may be favouring invasions of more mesic plants into historically drier sites, while high deer abundance likely limits hemlock regeneration. If current trends continue, maples will dominate the majority of northern forests, with significant losses of local native species richness and substantial shifts in understorey composition.     

Factors affecting species richness of tree regeneration in mixed‐wood stands of central Maine

Question: Two questions about within‐stand spatial variability are addressed in this paper. How does species richness of tree regeneration respond to small‐scale ecological gradients, and what effect does natural Abies balsamea abundance have on the species richness of other tree regeneration? Location: A long‐term, gap‐silviculture experiment, Acadian mixed‐wood forest, Maine, USA. Methods: Eight stands treated with and without gap harvesting were sampled to capture sub‐stand heterogeneity of understorey tree regeneration concurrently with patterning of local stand conditions. Spatial and non‐spatial models were developed to test the relationships between two response variables [species richness of small (height ≥0.1 m, but <0.75 m) and large (height ≥0.75 m, but <1.4 m) regeneration] and five explanatory variables (depth to water table, percentage canopy transmittance, A. balsamea regeneration density, and overstorey basal area and species richness). Results: Despite high unexplained variance for all models, consistent associations among variables were found. Negative associations were found between: (1) the species richness of small regeneration and A. balsamea regeneration density and (2) the species richness of large regeneration and overstorey basal area. Positive associations were found between: (1) the species richness of small regeneration and both overstorey basal area and species richness and (2) the species richness of small and large regeneration and canopy transmittance. Conclusions: Promoting tree species diversity in Acadian mixed‐wood stands may not be achievable through the use of gap‐harvesting alone if the density of understorey Abies balsamea is not reduced either naturally or through silvicultural intervention.     

Effect of forest canopy composition on soil nutrients and dynamics of the understorey: mixed canopies serve neither vascular nor bryophyte strata

Question: The effect of overstorey composition on above‐ground dynamics of understorey vegetation is poorly understood. This study examines the understorey biomass, production and turnover rates of vascular and non‐vascular plants along a conifer–broadleaf gradient of resource availability and heterogeneity. Location: Canadian boreal forests of northwest Quebec and Ontario. Methods: We sampled mature stands containing various proportions of black spruce (Picea mariana (Mill.) BSP), trembling aspen (Populus tremuloides Michx.) and jack pine (Pinus banksiana Lamb.). Above‐ground biomass of the understorey vegetation was assessed through harvesting; annual growth rates were calculated as the differences between biomass in 2007 and 2008, as estimated by allometric relationships, and turnover rates were estimated as net primary production divided by the biomass in 2007. Results: Higher aspen presence, linked to greater nutrient availability in the forest floor, was generally associated with higher vascular biomass and production in the understorey. This effect was less pronounced in sites of high intrinsic fertility. In contrast, bryophyte biomass was positively associated with conifer abundance, particularly in wet sites of the Quebec study area. Non‐linear responses resulted in total understorey biomass being lower under mixed canopies than under pure aspen or pure conifer canopies. Turnover rates did not differ with overstorey composition. Conclusions: While resource availability is a main driver of understorey productivity, resources as drivers appear to differ with differences in understorey strata components, i.e. vascular versus non‐vascular plants. Resource heterogeneity induced by a mixed canopy had overall negative effects on understorey above‐ground productivity, as this productivity seemed to rely on species adapted to the specific conditions induced by a pure canopy.     

Characteristics of the Psidium cattleianum invasion of secondary rainforests

Strawberry guava (Psidium cattleianum) is a shade‐tolerant shrub or small tree invader in tropical and subtropical regions and is considered among the world's top 100 worst invasive species. Studies from affected regions report deleterious effects of strawberry guava invasion on native vegetation. Here we examine the life history demographics and environmental determinants of strawberry guava invasions to inform effective weed management in affected rainforest regions. We surveyed the vegetation of 8 mature rainforest and 33 successional sites at various stages of regeneration in the Australian Wet Tropics and found that strawberry guava invasion was largely restricted to successional forests. Strawberry guava exhibited high stem and seedling densities, represented approximately 8% of all individual stems recorded and 20% of all seedlings recorded. The species also had the highest basal area among all the non‐native woody species measured. We compared environmental and community level effects between strawberry guava‐invaded and non‐invaded sites, and modelled how the species basal area and recruitment patterns respond to these effects. Invaded sites differed from non‐invaded sites in several environmental features such as aspect, distance from intact forest blocks, as well as supported higher grass and herb stem densities. Our analysis showed that invasion is currently ongoing in secondary forests, and also that strawberry guava is able to establish and persist under closed canopies. If left unchecked, strawberry guava invasion will have deleterious consequences for native regenerating forest in the Australian Wet Tropics.     

The relative importance of landscape and community features in the invasion of an exotic shrub in a fragmented landscape

Although invasive plants are recognized as a major ecological problem, little is known of the relative importance of plant community characteristics versus landscape context in determining invasibility of communities. We determined the relative importance of community and landscape features of 30 woodlots in influencing the invasion of Lonicera maackii. We sampled woodlots using the point‐quarter method and calculated canopy openness and basal areas and densities of shrub, sapling and tree species, as well as woody species richness. We used aerial photos and ArcView GIS to calculate landscape parameters from the same woodlots using a buffer distance of 1500 m. We used logistic and linear regression analyses to determine the community and landscape factors that best explain L. maackii presence and density. We also tested whether woodlot invasion by L. maackii begins at woodlot edges. Presence of L. maackii was significantly explained only by distance from the nearest town (logistic regression, p=0.017); woodlots nearer town were more likely to be invaded. Among invaded woodlots, density of L. maackii was positively related to the amount of edge in the landscape (partial R²=0.592) and negatively related to total tree basal area (partial R²=0.134), number of native woody species (partial R²=0.054), and sapling shade tolerance index (partial R²=0.054). Lonicera maackii in woodlot interiors were not younger than those on the perimeters, leading us to reject the edge‐first colonization model of invasion. Our findings reveal that landscape structure is of primary importance and community features of secondary importance in the invasion of L. maackii. This shrub is invading from multiple foci (towns) rather than an advancing front. Connectivity in the landscape (i.e. the number of corridors) did not promote invasion. However, edge habitat was important for invasion, probably due to increased propagule pressure. The community features associated with L. maackii invasion may be indicators of past disturbance.     

Disparate effects of global‐change drivers on mountain conifer forests: warming‐induced growth enhancement in young trees vs. CO2 fertilization in old trees from wet sites

Theory predicts that the postindustrial rise in the concentration of CO₂ in the atmosphere (c_a) should enhance tree growth either through a direct fertilization effect or indirectly by improving water use efficiency in dry areas. However, this hypothesis has received little support in cold‐limited and subalpine forests where positive growth responses to either rising c_a or warmer temperatures are still under debate. In this study, we address this issue by analyzing an extensive dendrochronological network of high‐elevation Pinus uncinata forests in Spain (28 sites, 544 trees) encompassing the whole biogeographical extent of the species. We determine if the basal area increment (BAI) trends are linked to climate warming and increased c_a by focusing on region‐ and age‐dependent responses. The largest improvement in BAI over the past six centuries occurred during the last 150 years affecting young trees and being driven by recent warming. Indeed, most studied regions and age classes presented BAI patterns mainly controlled by temperature trends, while growing‐season precipitation was only relevant in the driest sites. Growth enhancement was linked to rising c_a in mature (151–300 year‐old trees) and old‐mature trees (301–450 year‐old trees) from the wettest sites only. This finding implies that any potential fertilization effect of elevated c_a on forest growth is contingent on tree features that vary with ontogeny and it depends on site conditions (for instance water availability). Furthermore, we found widespread growth decline in drought‐prone sites probably indicating that the rise in c_a did not compensate for the reduction in water availability. Thus, warming‐triggered drought stress may become a more important direct driver of growth than rising c_a in similar subalpine forests. We argue that broad approaches in biogeographical and temporal terms are required to adequately evaluate any effect of rising c_a on forest growth.     

Post‐fire understorey regeneration in boreal Pinus sylvestris forest sites with different fire histories

Abstract. We studied the characteristics of understorey regeneration on two sites with different fire history in a mature Pinus sylvestris forest in eastern Finland. The study area was a 4‐ha plot, which was divided into two parts based on fire history analysis. In one part the last fire event was a stand‐replacing fire in the early 19^th century, after which the whole stand regenerated, while the other part of the study plot was subsequently burnt by a surface fire in 1906. Understorey P. sylvestris individuals were much more abundant in the area of the 1906 burn compared to the old burn. In both areas the size frequency distribution of living trees was bimodal, with frequency peaks at the < 5 cm and 30–150 cm height classes. In the old burn small understorey trees were mainly associated with microsites created by treefall disturbances while in the 1906 burn most small understorey trees occurred on vegetation‐covered microsites. This indicates that with increasing time since last fire establishment of new understorey trees becomes more restricted by the availability of microsites created by treefall disturbances. In both areas the proportion of vigorous small understorey trees was highest on decayed wood. In the older burn uprooted pits and mounds also had a significant proportion of healthy small understorey trees, while the majority of trees classified as seriously weakened or dying were growing on microhabitats characterized by undisturbed vegetation. Ripley's K‐function analyses showed that spatial distribution of understorey trees was clustered in both areas in all microsite types and clustering at small scales was most pronounced in understorey trees growing in uprooted spots or in association with decayed wood. The bivariate analysis showed a significant repulsion effect between large trees and understorey trees at intermediate spatial scales, indicating that competition had an effect on understorey tree distribution and this effect was more pronounced in the younger burn. The analysis suggests that in Pinus sylvestris forests the abundance, quality and spatial pattern of understorey tree population may vary considerably as a function of disturbance history.     

Initial response of understorey vegetation to fire severity and salvage‐logging in the southern boreal forest of Québec

Abstract. In this study we compared the effects of fire on understorey vegetation in the Québec southern boreal forest with effects of salvage‐logging (clear‐cutting after fire). All 61 400‐m² sampling sites were controlled for overstorey composition (Deciduous, Mixed and Coniferous) and disturbance type, which consisted of three fire impact severity (FIS) classes (Light, Moderate and Extreme) and two harvesting techniques (Stem‐only and Whole‐tree Harvesting). Percent‐cover data of vegetation and post‐disturbance environmental characteristics were recorded in the field during the first two years after fire as well as soil texture. Ordination of fire alone demonstrated that, on Coniferous sites, fire initiates a succession whereby the understorey Coniferous sites approaches that of Deciduous‐Mixed sites, due to the release of the understorey from Sphagnum spp. dominance, this pattern being a function of FIS. On Deciduous‐Mixed stands, increased FIS resulted in a transition from herb to shrub dominance. Ordination of all five disturbance types showed that the impact of salvage‐logging on understorey composition was within the range of fire, but marginalized to the extreme end of the FIS spectrum. Variance partitioning demonstrated that overstorey and soil texture were the most important explanatory variables of fire alone, while disturbance type explained the largest independent fraction of understorey variation when salvage‐logging was introduced. Salvage‐logging also results in significant reductions in understorey abundance, richness and diversity, while indicator species analysis suggests that it favours mesoxerophytic to xeric species. Results are interpreted in light of shade‐tolerance dynamics, forest floor disturbance and soil moisture regimes. Implications for sustainable forest management are discussed.     

Using space‐for‐time substitution and time sequence approaches in invasion ecology

1. Invasion biologists use two main approaches to evaluate the effects of non‐native species (NNS) on diversity of native species (DNS), namely space‐for‐time and time approaches. These approaches have pitfalls related to lack of controls: the former lacks pre‐invasion data, while the latter often lacks data from non‐invaded sites. 2. We propose a framework that combines space‐for‐time and time approaches and which should result in more focused mechanistic hypotheses and experiments to test the causes of invasibility and the effects of NNS on DNS. We illustrate the usefulness of our framework using two case studies: one with the submersed macrophyte, Hydrilla verticillata, in reservoir and the other with the fish, Geophagus proximus, in a large river–floodplain system. 3. Hydrilla verticillata invaded sites with DNS similar to that found in non‐invaded sites, indicating that biotic and/or abiotic factors did not influence invasion success; however, DNS increased over time in invaded sites compared with non‐invaded sites, suggesting that H. verticillata facilitated natives. In contrast, G. proximus invaded sites with higher DNS than non‐invaded sites, suggesting that biotic and/or abiotic factors favouring natives were important for invasion success, but DNS increased in invaded and non‐invaded sites over time, indicating that an independent factor contributed to DNS increases. 4. Conclusions from both studies would have been inaccurate or incomplete if the space‐for‐time and time approaches had not been used in combination as proposed in our framework.     

Foliar production and decomposition rates in urban forests invaded by the exotic invasive shrub, <Emphasis Type="Italic">Lonicera maackii</Emphasis>

Exotic invasive shrubs can form dense monocultures in forest understories, which can have cascading effects on ecosystem structure and function. Amur honeysuckle, an exotic shrub that forms dense canopies in eastern forests, has the potential to alter plant community structure and ecosystem functions, such as primary production and decomposition. The goal of this study was to examine foliar productivity and leaf litter decomposition in forests invaded by Amur honeysuckle (Lonicera maackii) and to determine the extent to which the presence of this dominant exotic species may alter ecosystem function in these forests. We found that forests invaded by Amur honeysuckle had 16 times greater honeysuckle foliar biomass and 1.5 times lower total foliar biomass than forests of equivalent tree basal area, but having few honeysuckle shrubs. This suggests that productivity of native tree and shrub species may be reduced where honeysuckle density is high. Additionally, honeysuckle litter decayed four times faster and released nitrogen more rapidly than sugar maple litter, and sugar maple litter decayed 19% faster in forests invaded by Amur honeysuckle. These findings suggest that forests invaded by Amur honeysuckle may exhibit lower rates of organic matter accrual and less nitrogen retention in the forest floor. Since honeysuckle leaves develop in early spring before those of other shrubs or trees in the area, the rapid release of nitrogen from honeysuckle litter that we measured in early spring is timed to benefit this invasive species. The temporally coincident phenologies of nitrogen release during decomposition with the foliar growth needs of this shrub indicates that a potential positive feedback loop may exist between these processes that promotes continued growth and dominance of honeysuckle shrubs in these forested systems.     

Pine savanna overstorey influences on ground‐cover biodiversity

Question: Does the overstorey of pine savannas influence plant species biodiversity in the ground cover? Location: Camp Whispering Pines (30°41’N; 90°29’W), eastern Louisiana (USA). Methods: We used ecologically sensitive restoration logging to remove patches of Pinus palustris (longleaf pine) in a second‐growth loess plain Pinus palustris savanna managed using frequent lightning season fires. Five years later, we measured numbers of vascular plant species and transmitted light in replicated 100‐m² plots. Treatments involved three different overstorey conditions: no overstorey for 5 years, no overstorey for several decades, and overstorey pines present for decades. Results: Both recent and long‐term openings contained, on average, about 100 vascular plant species per 100 m², 20% more than in similar‐sized areas beneath overstorey trees. Responses varied with life form; more herbaceous species occurred in recent and older overstorey openings than beneath overstorey trees. Total numbers of all species and of less abundant forb species were positively and linearly related to light transmitted to ground level. Those species responding to openings in the overstorey and positively associated with increased transmitted light levels were monocarpic and shortlived perennial forb and grass species with a seed bank in the soil. In addition, community structure, as reflected in species composition and abundances, appeared to vary with canopy condition. Conclusions: Restoration involving ecologically sensitive removal of patches of overstorey pines in frequently burned pine savannas should benefit the ground cover and increase plant species biodiversity as a result of increased abundance of seed bank species.     

A long‐term record of Nothofagus dominance in the southern Andes,Chile

Abstract The general model of regeneration dynamics in Nothofagus forests of southern South America could have value in community ecology if predictive relationships between disturbance history, functional traits and site attributes could be identified. Examined here is the proposal that on favourable sites shade‐intolerant Nothofagus are likely not to survive in competition with shade‐tolerant, broad‐leaved evergreen taxa of temperate rain forests, and persistence, thus, is dependent on periodic coarse‐scale disturbance. Comparison of stand dynamics of three old‐growth Nothofagus forests at different elevations in the southern Andes, Chile where deciduous Nothofagus alpina dominates the upper canopy, and examination of the life history trade‐offs of this variation were made. Stem density of all stems ≥5.0 cm d.b.h. was 233–303 stems per hectare, and basal area was 123.9–171.0 m²ha^?1. Maximum lifespan of N. alpina was found to be greater than ca 640 years, exceeding all previously reported ages for this species in the region. Forests had a stable canopy composition for this long‐term, but some appeared to lack effective regeneration of N. alpina in recent years. Regeneration of N. alpina was generally greater in disturbed stands and higher elevation than in undisturbed stands and at lower elevation. Recruitment emerged to be strongly affected by competitive over‐ and understorey associates. There was a gradient of increasing dependence of N. alpina on disturbance towards the more productive end of the environment gradients, and hence less dependence of N. alpina on disturbance for its regeneration towards higher elevation. The study confirms that changes in forest composition may be explained by processes occurring in accordance with the predictions of the existing model of Nothofagus regeneration dynamics, providing stronger evidence specifically directed at mid‐tolerant N. alpina, and by factoring out regeneration dynamics on favourable sites. Thus, for N. alpina, trait differences probably contribute to the competitive advantage over its associates in productive habitats, and may be linked to small‐to‐intermediate‐sized disturbances which inevitably occur as older trees die, enabling N. alpina to persist in forests and therefore maintain species coexistence for the long‐term.     

Distinct seasonal dynamics of responses to elevated CO2 in two understorey grass species differing in shade‐tolerance

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Community‐level changes in Australian subalpine vegetation following invasion by the non‐native shrub Cytisus scoparius

Question: What are the changes associated with the recent invasion by the non‐native legume, Cytisus scoparius? Location: Subalpine vegetation (1500 m a.s.l.) in Australia. Methods: We used multivariate techniques and regression analyses to assess vegetation and environmental changes across six study sites. Vegetation and environmental variables were investigated at three different stages of invasion: (1) recent invasion (8–10 yr), (2) mature invasion (15–16 yr) and (3) long‐term invasion (25 yr). Results: Substantial changes in floristic composition and species richness were evident after 15 yr and these changes became more pronounced after 25 yr. Changes due to invasion were associated with a dramatic loss of native species or a reduction in their abundance. No ‘new species’ were evident under invaded stands. Forbs were most affected by the establishment of C. scoparius, although all growth forms responded negatively. Dense canopy shading and an increasingly dense, homogeneous litter layer in the understorey as a result of C. scoparius were strong environmental drivers of vegetation change. Greenhouse studies confirmed the importance of these processes on the germination and growth of two native species. Conclusions: This study highlights the potential for C. scoparius to alter both vegetation and environmental processes in the subalpine region.     

Growth decline and divergent tree ring isotopic composition (δ13C and δ18O) contradict predictions of CO2 stimulation in high altitudinal forests

Human‐induced changes in atmospheric composition are expected to affect primary productivity across terrestrial biomes. Recent changes in productivity have been observed in many forest ecosystems, but low‐latitude upper tree line forests remain to be investigated. Here, we use dendrochronological methods and isotopic analysis to examine changes in productivity, and their physiological basis, in Abies religiosa (Ar) and Pinus hartwegii (Ph) trees growing in high‐elevation forests of central Mexico. Six sites were selected across a longitudinal transect (Transverse Volcanic Axis), from the Pacific Ocean toward the Gulf of Mexico, where mature dominant trees were sampled at altitudes ranging from 3200 to 4000 m asl. A total of 60 Ar and 84 Ph trees were analyzed to describe changes in growth (annual‐resolution) and isotopic composition (decadal‐resolution) since the early 1900s. Our results show an initial widespread increase in basal area increment (BAI) during the first half of the past century. However, BAI has decreased significantly since the 1950s with accentuated decline after the 1980s in both species and across sites. We found a consistent reduction in atmosphere to wood ¹³C discrimination, resulting from increasing water use efficiency (20–60%), coinciding with rising atmospheric CO₂. Changes in ¹³C discrimination were not followed, however, by shifts in tree ring δ¹⁸O, indicating site‐ and species‐specific differences in water source or uptake strategy. Our results indicate that CO₂ stimulation has not been enough to counteract warming‐induced drought stress, but other stressors, such as progressive nutrient limitation, could also have contributed to growth decline. Future studies should explore the distinct role of resource limitation (water vs. nutrients) in modulating the response of high‐elevation ecosystems to atmospheric change.     

Restoration of three forest herbs in the Liliaceae family by manipulating deer herbivory and overstorey and understorey vegetation

Background: Research on herbaceous vegetation restoration in forests characterised by overstorey tree harvests, excessive deer herbivory, and a dominant fern understorey is lacking. Most of the plant diversity found in Eastern hardwood forests in the United States is found in the herbaceous understorey layer. Loss of forest herbaceous species is an indicator of declining forest conditions.

Aims: The combined effects of deer herbivory, competitive understorey vegetation removal, and overstorey tree removal on the abundance and reproductive capacity of three understorey herbs in the Liliaceae family were evaluated.

Methods: A split-plot randomised block design was used with three replicates. Treatments included three harvest intensities, fenced/unfenced, herbicide/no herbicide-treated, prescribed burn/no prescribed burn, and all combinations. A generalised linear model was used to compare treatment effects over 8 years.

Results: Both fruit production and cover increased significantly in fenced areas for all three species. There was a significant 6-year recovery period for cover of the three species in response to herbicide. There was a significant 4-year recovery period of fire-treated plots for fruit production of the three species. The most intensively cut, fenced, and herbicide-treated plots had the greatest increases in sapling and Rubus spp. cover. Cover and fruit production of the three herbs were significantly greatest in the moderate-cut treatment.

Conclusions: Restoration of these three liliaceous species is most likely to occur in Eastern deciduous forests and similar forests using a combined fenced and moderate-cut treatment.     

Population dynamics of four understorey shrub species during a 7‐yr period in a primary beech forest

Abstract. Shrubs persist in the understorey layer of forests throughout their lives, while tall trees remain there only during the juvenile stage and then grow into the canopy layer. Thus demographic parameters (recruitment‐, mortality‐, and growth‐rates) of shrub species are expected to differ from those of tall tree species. We investigated aspects of the demography of four dominant deciduous‐shrub species (Viburnum furcatum, Lindera umbellata var. membranacea, Magnolia salicifolia, and Hydrangea paniculata) in Fagus crenata forests at the beginning and at the end of a 7‐yr period in a 1‐ha permanent plot. For each species, the number of stems changed little (within ± 10%) during the study period, while total basal area increased markedly (11.7–33.8%), because (1) new stems continuously recruited by vegetative growth replaced the substantial number of dead stems, and (2) vegetative stems grew vigorously, probably due to nutrient support from parents. The results indicate that these four understorey shrub species have high ability to maintain their population size in the shaded forest understorey. While in each species more than half of the dead stems were standing dead, a substantial proportion of the dead stems (9.0–38.5%) showed signs of mechanical damage, such as stem breakage and suppression by fallen branches or trees. Snow pressure that resulted in decumbent stems was also an important mortality agent for V. furcatum (20.7%) and L. umbellata var. membranacea (5.6%). Probability of damage was constant across all DBH‐classes for all study species. In each species, newly recruited stems and dead stems were spatially aggregated, largely due to habits of vegetative growth and mechanical damage, respectively. This study revealed that several demographic traits, resulting from recruitment by vegetative growth and death by mechanical damage, were shrub‐species specific and markedly different from those of tall tree species.