Inherent variation of functional traits in winter and summer leaves of Mediterranean seasonal dimorphic species. Evidence of a 'within leaf cohort' spectrum

The covariation pattern among leaf functional traits involved in resource acquisition has been successfully provided by the leaf economic spectrum (LES). Nevertheless, some aspects such as how the leaf trait variation sources affect LES predictions are still little investigated. Accordingly, the aim of this paper was to test whether leaf trait variations within different leaf cohorts could alter LES. Improving this knowledge can extend the potential of trait-based approaches in simulating future climate effects on ecosystems. A database on leaf morphological and physiological traits from different leaf cohorts of Cistus spp. was built by collecting data from literature. These species are seasonal dimorphic shrubs with two well-defined leaf cohorts during a year: summer leaves (SL) and winter leaves (WL). Traits included: leaf mass area (LMA), leaf thickness (LT), leaf tissue density (LTD), net photosynthetic rate on area (Aa) and mass (Am) base, nitrogen content on area (Na) and mass (Nm) base. The obtained patterns were analysed by standardized major axis regression and then compared with the global spectrum of evergreens and deciduous species. Climatic variable effect on leaf traits was also tested. Winter leaves and SL showed a great inherent variability for all the considered traits. Nevertheless, some relationships differed in terms of slopes or intercepts between SL and WL and between leaf cohorts and the global spectrum of evergreens and deciduous. Moreover, climatic variables differently affected leaf traits in SL and WL. The results show the existence of a 'within leaf cohort' spectrum, providing the first evidence on the role of leaf cohorts as LES source of variation. In fact, WL showed a high return strategy as they tended to maximize, in a short time, resource acquisition with a lower dry mass investment, while SL were characterized by a low return strategy

Individualistic responses of forest herb traits to environmental change

Intraspecific trait variation (ITV; i.e. variability in mean and/or distribution of plant attribute values within species) can occur in response to multiple drivers. Environmental change and land-use legacies could directly alter trait values within species but could also affect them indirectly through changes in vegetation cover. Increasing variability in environmental conditions could lead to more ITV, but responses might differ among species. Disentangling these drivers on ITV is necessary to accurately predict plant community responses to global change. We planted herb communities into forest soils with and without a recent history of agriculture. Soils were collected across temperate European regions, while the 15 selected herb species had different colonizing abilities and affinities to forest habitat. These mesocosms (384) were exposed to two-level full-factorial treatments of warming, nitrogen addition and illumination. We measured plant height and specific leaf area (SLA). For the majority of species, mean plant height increased as vegetation cover increased in response to light addition, warming and agricultural legacy. The coefficient of variation (CV) for height was larger in fast-colonizing species. Mean SLA for vernal species increased with warming, while light addition generally decreased mean SLA for shade-tolerant species. Interactions between treatments were not important predictors. Environmental change treatments influenced ITV, either via increasing vegetation cover or by affecting trait values directly. Species' ITV was individualistic, i.e. species responded to different single resource and condition manipulations that benefited their growth in the short term. These individual responses could be important for altered community organization after a prolonged period

Forest Stand Structure and Primary Production in relation to Ecosystem Development, Disturbance, and Canopy Composition

Temperate forests are complex ecosystems that sequester carbon (C) in biomass. C storage is related to ecosystem-scale forest structure, changing over succession, disturbance, and with community composition. We quantified ecosystem biological and physical structure in two forest chronosequences varying in disturbance intensity, and three late successional functional types to examine how multiple structural expressions relate to ecosystem C cycling. We quantified C cycling as wood net primary production (NPP), ecosystem structure as Simpson’s Index, and physical structure as leaf quantity (LAI) and arrangement (rugosity), examining how wood NPP-structure relates to light distribution and use-efficiency. Relationships between structural attributes of biodiversity, LAI, and rugosity differed. Development of rugosity was conserved regardless of disturbance and composition, suggesting optimization of vegetation arrangement over succession. LAI and rugosity showed significant positive productivity trends over succession, particularly within deciduous broadleaf forests, suggesting these measures of structure contain complementary, not redundant, information related to C cycling

A Comprehensive Emission Inventory of Bbiogenic Volatile Organic Compounds in Europe: Improved Seasonality and Land-cover

Biogenic volatile organic compounds (BVOC) emitted from vegetation are important for the formation of secondary pollutants such as ozone and secondary organic aerosols (SOA) in the atmosphere. Therefore, BVOC emission are an important input for air quality models. To model these emissions with high spatial resolution, the accuracy of the underlying vegetation inventory is crucial. We present a BVOC emission model that accommodates different vegetation inventories and uses satellite-based measurements of greenness instead of pre-defined vegetation periods. This approach to seasonality implicitly treats effects caused by water or nutrient availability, altitude and latitude on a plant stand. Additionally, we test the influence of proposed seasonal variability in enzyme activity on BVOC emissions. In its present setup, the emission model calculates hourly emissions of isoprene, monoterpenes, sesquiterpenes and the oxygenated volatile organic compounds (OVOC) methanol, formaldehyde, formic acid, ethanol, acetaldehyde, acetone and acetic acid. In this study, emissions based on three different vegetation inventories are compared with each other and diurnal and seasonal variations in Europe are investigated for the year 2006. Two of these vegetation inventories require information on tree-cover as an input. We compare three different land-cover inventories (USGS GLCC, GLC2000 and Globcover 2.2) with respect to tree-cover. The often-used USGS GLCC land-cover inventory leads to a severe reduction of BVOC emissions due to a potential miss-attribution of broad-leaved trees and reduced tree-cover compared to the two other land-cover inventories. To account for uncertainties in the land-cover classification, we introduce land-cover correction factors for each relevant land-use category to adjust the tree-cover. The results are very sensitive to these factors within the plausible range. For June 2006, total monthly BVOC emissions decreased up to −27% with minimal and increased up to +71% with maximal factors, while in January 2006, the changes in monthly BVOC emissions were −54 and +56% with minimal and maximal factors, respectively. The new seasonality approach leads to a reduction in the annual emissions compared with non-adjusted data. The strongest reduction occurs in OVOC (up to −32 %), the weakest in isoprene (as little as −19 %). If also enzyme seasonality is taken into account, however, isoprene reacts with the steepest decrease of annual emissions, which are reduced by −44% to −49 %, annual emissions of monoterpenes reduce between −30 and −35 %. The sensitivity of the model to changes in temperature depends on the climatic zone but not on the vegetation inventory. The sensitivity is higher for temperature increases of 3K (+31% to +64 %) than decreases by the same amount (−20 to −35 %). The climatic zones “Cold except summer” and “arid” are most sensitive to temperature changes in January for isoprene and monoterpenes, respectively, while in June, “polar” is most sensitive to temperature for both isoprene and monoterpenes. Our model predicts the oxygenated volatile organic compounds to be the most abundant fraction of the annual European emissions (3571–5328 Gg yr−1), followed by monoterpenes (2964–4124 Gg yr−1), isoprene (1450–2650 Gg yr−1) and sesquiterpenes (150–257 Gg yr−1). We find regions with high isoprene emissions (most notably the Iberian Peninsula), but overall, oxygenated VOC dominate with 43–45% (depending on the vegetation inventory) contribution to the total annual BVOC emissions in Europe. Isoprene contributes between 18–21 %, monoterpenes 33–36% and sesquiterpenes contribute 1–2 %.We compare the concentrations of biogenic species simulated by an air quality model with measurements of isoprene and monoterpenes in Hohenpeissenberg (Germany) for both summer and winter. The agreement between observed and modelled concentrations is better in summer than in winter. This can partly be explained with the difficulty to model weather conditions in winter accurately, but also with the increased anthropogenic influence on the concentrations of BVOC compounds in winter. Our results suggest that land-cover inventories used to derive tree-cover must be chosen with care. Also, uncertainties in the classification of land-cover pixels must be taken into account and remain high. This problem must be addressed together with the remote sensing community. Our new approach using a greenness index for addressing seasonality of vegetation can be implemented easily in existing models. The importance of OVOC for air quality should be more deeply addressed by future studies, especially in smog chambers. Also, the fate of BVOC from the dominant region of the Iberian Peninsula should be studied more in detail

Optimization of leaf morphology in relation to leaf water status: A theory.

The leaf economic traits such as leaf area, maximum carbon assimilation rate, and venation are all correlated and related to water availability. Furthermore, leaves are often broad and large in humid areas and narrower in arid/semiarid and hot and cold areas. We use optimization theory to explain these patterns. We have created a constrained optimization leaf model linking leaf shape to vein structure that is integrated into coupled transpiration and carbon assimilation processes. The model maximizes net leaf carbon gain (NPPleaf) over the loss of xylem water potential. Modeled relations between leaf traits are consistent with empirically observed patterns. As the results of the leaf shape-venation relation, our model further predicts that a broadleaf has overall higher NPPleaf compared to a narrowleaf. In addition, a broadleaf has a lower stomatal resistance compared to a narrowleaf under the same level of constraint. With the same leaf area, a broadleaf will have, on average, larger conduits and lower total leaf xylem resistance and thus be more efficient in water transportation but less resistant to cavitation. By linking venation structure to leaf shape and using water potential as the constraint, our model provides a physical explanation for the general pattern of the covariance of leaf traits through the safety-efficiency trade-off of leaf hydraulic design

Forest Net Primary Production Resistance Across a Gradient of Moderate Disturbance

The global carbon (C) balance is vulnerable to disturbances that alter terrestrial C uptake and loss. Moderate disturbances that kill or defoliate only a subset of canopy trees such as insect defoliation, drought, and age-related senescence are increasing in extent and frequency; yet, little is known about the effect of moderate disturbance on forest production and the mechanisms sustaining or supporting the recovery of the C cycle across a range of moderate disturbance severities. We used a broad plot-scale gradient of upper canopy tree mortality within a large manipulation of forest disturbance to: 1) quantify how aboveground wood net primary production (ANPPw) responds to a range of moderate disturbance severities and; 2) identify the primary mechanisms supporting ANPPw resistance or resilience following moderate disturbance. We found that ANPPw was highly resistant to moderate disturbance, with production levels sustained following the senescence of 9 to \u3e 60 % of the upper canopy tree basal area. As upper canopy gap fraction increased with rising disturbance severity, greater light availability to the subcanopy enhanced leaf-level C uptake and the growth of this formerly light-limited canopy stratum, compensating for upper canopy production losses. As a result, whole-ecosystem production efficiency (ANPPw/LAI) increased at high levels of disturbance severity and leaf area loss. These findings provide a mechanistic explanation for sustained ANPPw across the disturbance gradient, in which the physiological and growth enhancement of undisturbed vegetation was proportional to the level of disturbance severity. Our results have important ecological and management implications, showing that moderate disturbances may minimally alter ecosystem functions such as C storage

Divergent drivers of leaf trait variation within species, among species, and among functional groups.

Understanding variation in leaf functional traits-including rates of photosynthesis and respiration and concentrations of nitrogen and phosphorus-is a fundamental challenge in plant ecophysiology. When expressed per unit leaf area, these traits typically increase with leaf mass per area (LMA) within species but are roughly independent of LMA across the global flora. LMA is determined by mass components with different biological functions, including photosynthetic mass that largely determines metabolic rates and contains most nitrogen and phosphorus, and structural mass that affects toughness and leaf lifespan (LL). A possible explanation for the contrasting trait relationships is that most LMA variation within species is associated with variation in photosynthetic mass, whereas most LMA variation across the global flora is associated with variation in structural mass. This hypothesis leads to the predictions that (i) gas exchange rates and nutrient concentrations per unit leaf area should increase strongly with LMA across species assemblages with low LL variance but should increase weakly with LMA across species assemblages with high LL variance and that (ii) controlling for LL variation should increase the strength of the above LMA relationships. We present analyses of intra- and interspecific trait variation from three tropical forest sites and interspecific analyses within functional groups in a global dataset that are consistent with the above predictions. Our analysis suggests that the qualitatively different trait relationships exhibited by different leaf assemblages can be understood by considering the degree to which photosynthetic and structural mass components contribute to LMA variation in a given assemblage

ENVIRONMENTAL VARIABLES ASSOCIATED WITH INVASIVE GLOSSY BUCKTHORN (FRANGULA ALNUS MILL.) AND INDIRECT CONTROL STRATEGIES FOR FOREST MANAGERS

Glossy buckthorn (Frangula alnus Mill.) is one of the most prominent non-native invasive plant species affecting New England forests. It quickly invades a forest and can create a dense understory effectively altering the species composition and dynamics of that forest. To gain a better understanding of the environmental variables associated with glossy buckthorn density we sampled forests across New Hampshire with varying degrees of buckthorn invasion. The effect on tree regeneration was analyzed with measurements of height and abundance of glossy buckthorn and native regeneration. Glossy buckthorn was found to be at its highest densities in disturbed softwood forests that were historically old fields, specifically eastern white pine (Pinus strobus L.), with a thin organic layer and low herbaceous cover on drained loam and clay soils. The data show there is direct competition between glossy buckthorn and forest tree regeneration, although no relationship with regeneration shade tolerance was found. This information was used to create a prescription risk tree to aid forest managers in assessing the risk of buckthorn invasion and inhibition of tree regeneration associated with harvesting and suggests how to adapt their silvicultural prescriptions

Plant and soil microbe responses to light, warming and nitrogen addition in a temperate forest

1. Temperate forests across Europe and eastern North America have become denser since the 1950s due to less intensive forest management and global environmental changes such as nitrogen deposition and climate warming. Denser tree canopies result in lower light availability at the forest floor. This shade may buffer the effects of nitrogen deposition and climate warming on understorey plant communities. 2. We conducted an innovative in situ field experiment to study the responses of co-occurring soil microbial and understorey plant communities to nitrogen addition, enhanced light availability and experimental warming in a full-factorial design. 3. We determined the effects of multiple environmental drivers and their interactions on the soil microbial and understorey plant communities, and assessed to what extent the soil microbial and understorey plant communities covary. 4. High light led to lower biomass of the soil microbes (analysed by phospholipid fatty acids), but the soil microbial structure, i.e. the ratio of fungal biomass to bacterial biomass, was not affected by light availability. The composition of the soil bacterial community (analysed by high-throughput sequencing) was affected by both light availability and warming (and their interaction), but not by nitrogen addition. Yet, the number of unique operational taxonomic units was higher in plots with nitrogen addition, and there were significant interactive effects of light and nitrogen addition. Light availability also determined the composition of the plant community; no effects of nitrogen addition and warming were observed. The soil bacterial and plant communities were co-structured, and light availability explained a large part of the variance of this co-structure. 5. We provide robust evidence for the key role of light in affecting both the soil microbial and plant communities in forest understoreys. Our results advocate for more multifactor global change experiments that investigate the mechanism underlying the (in) direct effects of light on the plant-soil continuum in forests